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

lamarckii
(IndoWest Pacic 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, ngochango-
chara (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)
Norman C. Duke
v
n
o
:
o
:

x
.

c
.

u
u
x
v
Rhizophora mucronata trees showing stilt roots,
Chuuk, Federated States of Micronesia.
Species Proles for Pacic Island Agroforestry
www.traditionaltree.org
April acc6
vcr. a..
IN BRIEF
Distribution Nativc to tropical and subtropi
cal coastal arcas lrom thc Alrican cast coast,
throughout Asia to Australia and to most is
lands ol thc castcrn Pacic ccan. Closcly al
licd with Atlanticast Pacic rcd mangrovcs
whosc rangcs naturally ovcrlap only in a small
numbcr ol southcrn Pacic islands.
Size Can rcach cc m (.cc.c lt) in hcight,
although commonly rcachcs m (.6a6 lt).
Habitat !nhabits thc intcrtidal wctland zonc,
c6 m (cac lt) clcvation bctwccn mcan sca
lcvcl and highcst tidcs, with variablc rainlall.
Vegetation Mangrovc communitics.
Soils Adaptcd to a widc rangc but thrivcs bcst
in nc mud scdimcnts ol downstrcam rivcr cs
tuarics.
Growth rate Grows . m/yr (. lt/yr) in
hcight.
Main agroforestry uses Soil stabilization,
coastal protcction, wildlilc/marinc habitat lor
marinc launa.
Main products Timbcr, luclwood, charcoal,
dycs, and traditional mcdicinc.
Yields !n Malaysia, thc cycar rotation har
vcstcd yicld ol grccn wood has bccn about .6
a mt/ha (6.. t/ac).
Intercropping Rccommcndcd lor planting to
gcthcr with othcr mangrovc spccics.
Invasive potential Has potcntial to invadc ncw
cnvironmcnts, not rccommcndcd lor planting
outsidc ol thc natural rangc. Tcsc plants arc
rcady colonizcrs ol ncw mud banks, making
thcm opportunistically invasivc.
2 !ndoVcst Pacic Rhizophora spccics
INTRODUCTION
!ndoVcst Pacic stilt mangrovcs (!VP Rhizophora spc
cics) arc widcsprcad throughout most tropical coastal arcas
ol thc wcstcrn Pacic rcgion to cast Alrica. Tis group is
onc ol two that makc up thc gcnus Rhizophora and consists
ol thrcc spccics (two bcing closcly allicd), R. mucronata, R.
stylosa, and R. apiculata, and two hybrids, R. lamarckii
and R. annamalai.
Stilt mangrovcs thrivc undcr a rangc ol intcrtidal condi
tions, including a rangc ol salinity lcvcls lrom ncar lrcsh
watcr to lull strcngth scawatcr. Tcy tolcratc a rangc ol
ooding rcgimcs, soil typcs, and othcr physical sitc lactors.
Typically, thcsc mangrovcs arc common in thc midintcr
tidal zonc, and particularly along thc scaward margin ol
tropical mangrovc stands.
Rhizophora arc considcrcd thc most important ol all man
grovc gcncra across thc Pacic tropical rcgion. Howcvcr,
thc bcncts providcd by stilt mangrovcs arc dicult to
scparatc lrom thc largcr rolc ol mangrovcs and mangrovc
ccosystcms in gcncral. Mangrovcs arc known to play a vital
rolc in shorclinc protcction, cnhanccmcnt ol watcr quality
in ncarshorc cnvironmcnts (including coral rccls), and in
supporting cstuarinc and marinc lood chains. !n most parts
ol thc Pacic, trccs arc harvcstcd lor rcwood, so thc trunk
is thc main part ol thc trcc considcrcd lor dircct usc.
DISTRIBUTION
Native range
!ndoVcst Pacic stilt mangrovcs occur widcly through
out thc wcstcrn Pacic. Spccically, thcy occur in tropical
and subtropical, intcrtidal wctlands lrom thc cast coast ol
Alrica, through Asia to Microncsia cxtcnding cast in thc
northwcstcrn Pacic through thc Fcdcratcd Statcs ol Mi
croncsia to thc Marshall !slands, and south to northcrn
Australia cxtcnding cast in thc southcrn Pacic as lar as
Samoa. istributions appcar continuous, but scparatc spc
cics havc dicrcnt prclcrrcd locations whcrc thcy domi
natc:
R. mucronata occurs mostly in arcas subjcct to rcgu
lar lrcshwatcr ows (at lcast in thc castcrn part ol its
rangc)
R. stylosa occurs in marinc situations oltcn prclcrring
morc cxposcd oshorc sitcs
R. apiculata is oltcn lound midlowcr cstuary in largcr
rivcrinc cstuarics and cmbaymcnts
thc hybrid R. lamarckii is lound downstrcam in mid
dlc to highcr intcrtidal locations.

Rhizophora mucronata and R. stylosa arc sibling spccics (i.c.,

possibly R. stylosa -R. mucronata var. stylosa (Gri.) Sal
voza), and togcthcr thcy charactcrizc most stands ol !VP
stilt mangrovcs. Tc rangc ol R. mucronata is thc widcst ol
all !VP spccics, cxtcnding lrom cast Alrica, whcrc it oc
curs as thc solc Rhizophora, to thc wcstcrn Pacic, whcrc it
ovcrlaps with all othcr !VP spccics. !n contrast, R. stylosa
cxtcnds cxclusivcly cast, notably into thc southwcstcrn Pa
cic. Tc rangc ol R. apiculata also cxtcnds lurthcr cast in
thc south Pacic than R. mucronata, but othcrwisc its rangc
mostly ts within thc rangcs ol othcr !VP Rhizophora. Tc
status ol obscrvcd dicrcnccs in similar lorms, R. mucro-
nata and R. stylosa, will only bc rcsolvcd in gcnctic studics
and sclcctcd brccding programs.
Tc two hybrid taxa prcsumably havc distributions in kccp
ing with thcir hybrid status. As such, thcy arc locatcd whcr
cvcr thc rangcs ol thcir putativc parcnts ovcrlap. Conrmcd
rccords ol hybrid distributions show R. lamarckii to bc
widcsprcad in thc wcstcrn Pacic, whilc R. annamalai
was lound in !ndia and Sri Lanka along thc wcstcrn limits
ol R. apiculata. Howcvcr, considcrablc conlusion is cxpcct
cd in thc cld distribution rccords, bccausc thcsc hybrids
WHAT IS A MANGROVE?
Mangrovcs lorm a uniquc and dominant ccosystcm
compriscd ol intcrtidal marinc plants, mostly trccs,
prcdominantly bordcring margins ol tropical coast
lincs around thc world. Tcsc halophytic (salt tolcr
ant) plants thrivc in salinc conditions and daily inun
dation bctwccn mcan sca lcvcl and highcst astronomi
cal tidcs, and thcy providc vital structurc as habitat
and lood lor similarly adaptcd rcsidcnt and transicnt
launa. Mangrovc plants cxchangc gascs lrom cxposcd
roots using spccial lcnticcls, whilc ooding tidcs allow
uptakc ol rivcrbornc nutricnts and lrcqucnt dispcrsal
by thcir buoyant propagulcs. Tc ccological limits dc
ncd by thc diurnal tidal rangc cxplain thc sctting and
why just ,c spccics around thc world arc considcrcd
to bc mangrovcs (Tomlinson .6, ukc ct al. .),
comparcd with adjaccnt rainlorcsts that may havc
hundrcds ol trcc spccics pcr hcctarc. Spccializcd mor
phological and physiological charactcristics largcly
dcnc and charactcrizc mangrovc plants, such as but
trcss trunks and roots providing support in solt scdi
mcnts, abovcground roots allowing vital gas cxchangc
in anacrobic scdimcnts, and physiological adaptations
lor cxcluding or cxpclling salt. Fcwcr than aa plant
lamilics havc dcvclopcd such csscntial attributcs, rcp
rcscnting indcpcndcnt instanccs ol cocvolution ovcr
millions ol ycars to lorm todays mangrovc habitats.
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 3
arc dicult to distinguish bascd on morphological char
actcrs alonc.
l grcat intcrcst also in thc southwcstcrn Pacic islands
is thc only occurrcncc ol anothcr hybrid taxon, R. selala
(Salvoza) Tomlinson, dcrivcd lrom putativc parcnts R.
samoensis (-R. mangle:) and R. stylosa (-R. mucronata var.
stylosa:). Tis hybrid taxon is spccial bccausc R. samoen-
sis is a kcy mcmbcr ol thc AP rcd mangrovc Rhizophora
spccics. Tc occurrcncc ol this hybrid mcans thcrc appcars
to bc vcry littlc gcnctic scparation bctwccn thcsc dcning
and most divcrgcnt ol Rhizophora spccics.
Current distribution
Tcrc arc no rcports ol any stilt mangrovcs occurring out
sidc thcir natural rangc. Howcvcr, thcsc spccics arc rccog
nizcd as valuablc timbcr produccrs, so it is possiblc that
thcir dispcrsal cast in thc Pacic and clscwhcrc may havc
bccn assistcd by indigcnous pcoplcs in thc past.
BOTANICAL DESCRIPTION
Preferred scientic names
Rhizophora mucronata Lamk.
Rhizophora stylosa Gri. (- R. mucronata var. stylosa (Gri.)
Salvoza)
Rhizophora apiculata 8l.
Rhizophora lamarckii Montr. (- R. stylosa R. apiculata)
Rhizophora annamalai K. Kathircsan. (- R. mucronata R.
apiculata)
Family
Rhizophoraccac (mangrovc lamily)
Common names
aak, akelel, akapah (Pohnpci, FSM)
abat (Ccram, !ndoncsia)
ailave, kailau (Aru, !ndoncsia)
bakau (Malay Pcninsula, Singaporc, 8ornco, Sumatra)
bakauan, bakau, bakhaw (Philippincs)
bako ( Java, !ndoncsia)
bangka (Achch, !ndoncsia)
chia, cia (Chuuk, FSM)
!ndoVcst Pacic stilt mangrovcs (nglish)
kahrak, suhkasrik, sakasrik (Kosrac, FSM)
koriki (aru and Kiwai, PNG)
lolaro, belukap (Sulawcsi, !ndoncsia)
mangle hembra (Guam: Chamorro)
ngochangochara, petu rogha (Marovo, Solomon !slands)
prop root mangrovcs (northcastcrn Australia)
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. i:~cv: x.c. uuxv
4 !ndoVcst Pacic Rhizophora spccics
raway, roway, ravey (Yap !slands, FSM)
rcd, tallstiltcd mangrovc (Quccnsland)
spottcd lcavcd rcd mangrovc (Vcstcrn Australia)
tebechel, bngaol, agpat, apgal (Palau)
tiri tambua (Fiji)
totoa (Motu, PNG)
Size
!ndoVcst Pacic stilt mangrovcs arc mcdium to tall trccs
that may rcach cc m (.cc.c lt) in hcight, although
thcy arc commonly much shortcr, around m (.6a6 lt).
Stcm diamctcrs arc oltcn about . cm (6. in) takcn
just abovc thc highcst prop root. Tis mcasurc dicrs lun
damcntally lrom thc standard diamctcr at brcast hcight
(dbh) uscd lor most lorcst survcys, as diamctcr hcight
abovc thc substratc varics lrom c. to , m (..6a lt) (con
sidcr thc trcc in thc photo on thc rst pagc).
Form
Stilt mangrovcs arc rambling to columnar trccs with distinct
abovcground prop roots. Trccs tcnd to bc ol shortcr staturc
and morc sprcading in shapc on thc scaward cdgc ol stands
or in arcas ol highcr salinity. Tallcr, singlcstcmmcd trccs
arc lound most oltcn just bchind thc watcrs cdgc ol stands
midstrcam in major rivcrinc cstuarics. Multistcmmcd
trccs arc common in morc arid or marginal habitats.
Flowering
Flowcrs arc pcrlcct. !norcsccnccs havc lcw to many
joints with .,achotomous branching and onc to many
buds possiblc pcr inorcsccncc. pcn owcrs arc locatcd
within or bclow lcal axils at lcal nodcs bclow thc apical
shoot, dcpcnding on spccics. For R. mucronata, R. stylosa,
R. annamalai. R. lamarckii, and R. apiculata maturc
buds and owcrs arc locatcd at ., ., , 6, and 6..
nodcs down lrom apical shoot, rcspcctivcly. Calyccs arc
typically palc ycllow at maturity with lobcs, rarcly . 8uds
arc obovatc, grccn whcn immaturc to palc ycllowish grccn
as thcy maturc, dimcnsions .a cm (c.c. in) long and ~.
cm (c. in) widc. Pctals, usually , arc lanccolatc to lincar,
crcamy whitc, woolly to hairlcss, ~.c mm (c. in) long and
~a mm (c.c in) widc. Stamcns numbcr ,.a, palc ycllow.
Stylc is palc grccn, tcrctc, c.6 mm (c.cac.a in) abovc
basc, dichotomous tip. 8racts and bractcolcs arc variablc
dcpcnding on spccics. Maturc bud bracts arc slcndcr grccn
(R. mucronata, R. stylosa), swollcn grccn (R. annamalai,
R. lamarckii) and swollcn corky grccn (R. apiculata). Pc
dunclcs arc ., cm (c.a. in) long, and ~ mm (c..a in)
widc. Flowcring pcriod is chicy during Augustcccm
bcr in thc southcrn hcmisphcrc, and during FcbruaryJunc
in thc northcrn hcmisphcrc.
Leaves
Lcavcs arc oppositc, simplc, light or dark grccn, obovatc,
lcathcry, margins rcvolutc, bluntly acutc apcx with a dis
tinct mucronatc tip, ., mm (c.cc. in) long. Uppcr lcal
surlacc is smooth, shiny. Cork wart spots occur on undcr
surlacc, scattcrcd cvcnly, not raiscd, prcscnt in most spccics
in most locations, but abscnt only lrom R. apiculata and
R. lamarckii in southcrn Ncw Guinca and Australia. Ma
turc lcal dimcnsions arc 6. cm (a.,. in) long, and .c
cm (..a. in) widc. Pctiolc . mm (c.cc..6 in) long.
Lcal cmcrgcncc is mostly around NovcmbcrFcbruary in
thc southcrn hcmisphcrc and MayAugust in thc northcrn
hcmisphcrc. Lcal lall occurs mostly during thc wct summcr
pcriod lrom ctobcr to Fcbruary in thc southcrn hcmi
sphcrc and AprilAugust in thc northcrn hcmisphcrc.
Fruit
Fruits, whcn maturc, arc pcarshapcd, clongatc, waist
constriction, smooth brown surlacc, calyx lobcs clongatc
sprcading (whcn hypocotyl rcady to cmcrgc). For R. mu-
cronata, R. stylosa, R. annamalai, R. lamarckii, and R.
apiculata maturc lruit locatcd in lcal axils , ,, (rarc),
, (rarc), and nodcs down lrom apical shoot, rcspcctivcly.
Seeds/hypocotyls
Likc all Rhizophora spccics, stilt mangrovcs arc viviparous,
mcaning that thc trccs producc sccds hiddcn in thc maturc
lruit, and thcsc gcrminatc on thc parcnt trcc. Tc dispcr
sal unit, a viviparous sccdling, is callcd a hypocotyl. nc
hypocotyl is usually produccd lrom cach lruit, although on
rarc occasions twins may bc obscrvcd.
Hypocotyls arc narrowly cylindrical, clongatc, grccn,
smooth with irrcgular small brown lcnticcls, distal hall is
slightly widcr, distal tip is pointcd in most taxa, but round
cd to blunt lor R. apiculata. For R. mucronata, R. stylosa,
R. annamalai, R. lamarckii, and R. apiculata maturc
hypocotyls arc locatcd in lcal axils .c, , nonc,
(rarcly obscrvcd), and . nodcs down lrom thc apical
shoot, rcspcctivcly. Hypocotyl dimcnsions arc variablc and
not consistcntly spccicsspccic, .c cm (6. in) long,
.a cm (c.c. in) at thc widcst point, and c... cm
(c.ac.6 in) widc at thc collar, thc lruiting structurc that
cnvclops thc plumulc (cmbryonic lcavcs).
Fruiting, whcn maturc hypocotyls lall, occurs chicy (but
not cxclusivcly) lrom Novcmbcr to January in thc southcrn
hcmisphcrc and May to July in thc northcrn hcmisphcrc.
Bark
Tc bark is gray to dark gray and hcavily ssurcd, occa
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 5
sionally rcdbrown and smooth. Prop roots arc sturdy cvcn
whcn rclativcly thin.
Rooting habit
Maturc trccs havc distinctivc, sturdy, abovcground prop
roots surrounding thc stcm basc that anchor only shal
lowly in thc scdimcnts to .a m (.6.6 lt) dcpth. Tis
conlorms to thc oxygcn dccicnt (anoxic) conditions com
monly mcasurcd in mangrovc scdimcnts.
Similar species
Stilt mangrovcs arc distinguishcd lrom Atlanticast Pa
cic rcd mangrovc spccics principally by thc spikcd, mu
cronatc tip at thc lcal apcx ol stilt mangrovcs that is abscnt
in rcd mangrovcs.
Rhizophora mucronata and R. stylosa havc slcndcr (i.c., lcngth
much grcatcr than thc width) bracts at thc basc ol maturc
buds as distinguishcd lrom R. apiculata, R. lamarckii, and
R. annamalai that havc bracts almost as widc, or widcr
than thc lcngth.
Rhizophora apiculata is distinguishcd lrom R. lamarckii
and R. annamalai plus othcr !VP spccics by swollcn
(widcr than long), corky brown bracts, onc inorcsccncc
joint, and nodc position ol maturc buds and owcrs in lcal
axils at 6.. nodcs down lrom apical shoot, wcll bclow
lcavcs in thc lcaly shoot.
Hybrid charactcristics arc shown in R. lamarckii whcrc
Top left: Te distinct mucronate (abrupt, pointed) tip at the apex of leaves is characteristic of all IWP stilt Rhizophora. Te
coloration of the mucronate tip shown in this image of R. stylosa is variably red or green. Northeastern Australia. Top right:
Inorescence of Rhizophora mucronata (similar to R. stylosa) showing open owers with distinctive woolly petals and slender
smooth bract beneath the calyx. Bottom left: Inorescence of Rhizophora apiculata showing open owers with distinctive hair-
less petals, non-reexed 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. vno:os: x.c. uuxv
6 !ndoVcst Pacic Rhizophora spccics
it has kcy charactcrs intcrmcdiatc bctwccn R. apiculata
and R. stylosa. Similarly, hybrid charactcristics arc shown
in R. annamalai, which has kcy charactcrs intcrmcdiatc
bctwccn R. apiculata and R. mucronata. Hybrids can oltcn
bc largcr and tallcr than ncighboring parcnt trccs.
Rhizophora lamarckii is distinguishcd lrom R. annamalai
by stylc lcngth .., mm (c.c, in).
Rhizophora mucronata and R. stylosa, thc sibling spccics, arc
distinguishcd by short stylcs (a mm |c.c in| long) in R.
mucronata, whilc R. stylosa has long stylcs (~a mm |c.c in|
long). As notcd, R. mucronata and R. stylosa appcar closcly
rclatcd and arc possibly thc samc spccics. !l gcnctic stud
ics show thcm to bc thc samc, thcn R. stylosa would bc
rcnamcd R. mucronata var. stylosa. iscriminating bctwccn
thcm rcliably is not possiblc in many instanccs without
dctailcd cxamination ol kcy morphological and gcnctic
charactcristics. !ndividuals with intcrmcdiatc lcngth stylcs
occur in scvcral locations.
GENETICS
Variability of species
!ndoVcst Pacic stilt mangrovcs arc thosc Rhizophora
spccics that occur naturally lrom cast Alrica to thc wcst
crn Pacic ccan (scc thc map). Tis group compriscs
vc rclativcly distinct taxa, although at timcs thcir mor
phological and taxonomic dicrcnccs appcar qucstionablc.
Tc unccrtainty is chicy bascd on: .) thc prcscncc ol two
intcrmcdiatc individuals that arc rccognizcd as distinct
hybrids, namcly R. lamarckii and R. annamalai, and a)
thc occurrcncc ol two sibling spccics, R. mucronata and R.
stylosa, which may, on closcr cxamination, bc shown to bc
thc samc spccics.
Tc rclationship ol R. mucronata and R. stylosa is pcrhaps
thc most contcntious. Tcy appcar closcly rclatcd bccausc
thcy arc distinguishcd only by onc kcy charactcr, namcly
stylc lcngth. Tc sibling spccics arc largcly not scparatcd
gcographically, with a signicant portion ol thc rangc ol
cach ovcrlapping thc othcr. Howcvcr, thcir rangcs do in
cludc arcas ol cxclusivity in thc wcst and cast, rcspcctivcly.
Tcrc arc also indications that morphological and ccologi
cal charactcristics vary lor cach taxon across thcir widc
rangcs. For instancc, R. mucronata in cast Alrica has thc
samc cxposcd watcr cdgc habit as R. stylosa in northcrn
Australia. 8y contrast, R. mucronata in Australia and Asia
occurs in upstrcam locations ol lrcshwatcr dominatcd cstu
arics. Tcrc arc also subtlc but distinct dicrcnccs in mor
phological charactcristics in R. mucronata lor bud shapc,
inorcsccncc structurc, and lcal dimcnsions that may bc
Mature hypocotyls of Rhizophora stylosa (similar to R. mucro-
nata). Great Sandy Straits, NE Australia. vno:o: x.c. uuxv
Left to right: Mature ower buds of Rhizophora apiculata,
R. lamarckii, R. stylosa, and R. mucronata. Tese samples
were collected in NE Australia, where the four species grow
side by side in several estuaries. vno:o: x.c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 7
rclatcd to dicrcnccs in habitat. !t is clcar that
R. stylosa is thc taxon bcst adaptcd to marinc
cxposcd locations whcrcvcr it occurs.
!t is ol grcat intcrcst to lully cvaluatc thc rcla
tivc and prccisc distributions ol R. mucronata
and R. stylosa, bccausc thcsc arc likcly to cxplain
thc way spccics havc naturally cvolvcd and dis
pcrscd. !n somc island groups to thc north ol
Australia, lor instancc, thcrc is a curious pattcrn
in scparatc sympatric occurrcnccs whcrc thcsc
spccics arc casily distinguishcd in somc popu
lations (Philippincs, Sulawcsi) whilc in othcrs
(Fcdcratcd Statcs ol Microncsia, Palau) thcir
stylc lcngths ovcrlap, making it impossiblc to
always tcll thcm apart using this charactcr. Tis
may bc cxplaincd by a progrcssivc lcngthcning
ol thc stylc in R. stylosa, corrcsponding possibly
with its morc rcccnt introduction in locations
with longcr stylcs. !l this is corrcct, R. stylosa
has sprcad lrom Australia and Fcdcratcd Statcs
ol Microncsia both wcstward and castward,
lor cxamplc, in Sulawcsi and Fiji whcrc stylc
lcngths arc thc longcst rccordcd. Tis idca con
curs with thc spccializcd marinc habit ol this
spccics.
8y contrast, R. apiculata is rcadily distinguishcd
lrom R. mucronata and R. stylosa. Tis spccics
appcars morc prcvalcnt in cstuarics inucnccd
by largcr and morc continuous lrcshwatcr ows,
and it is lound in a continuous distributional
rangc, lrom !ndia to thc wcstcrn Pacic and northcrn
Australia. Across this rangc thcrc arc also two lorms ol R.
apiculata, and thcir occurrcnccs do not ovcrlap. nc lorm,
lound north and wcst lrom thc northcrn Ncw Guinca
coast, ts thc typc dcscription in cvcry rcspcct including
thc prcscncc ol cork wart spots on lcal undcrsurlaccs. Tc
othcr lorm lacking thcsc spots occurs south and cast lrom
thc southcrn Ncw Guinca coast. Tis charactcr was thought
to bc uniquc to R. apiculata until rcccntly, whcn trccs ol R.
racemosa (an AP rcd mangrovc) without spots wcrc dis
covcrcd in 8razil. Tis charactcr hclps dcnc two lorms ol
R. apiculata in Australasia and thc wcstcrn Pacic.
Known varieties and hybrids
Rhizophora mucronata and R. stylosa arc both known to
hybridizc with R. apiculata, giving risc to morphologi
cally distinct and gcographically widcsprcad hybrid lorms,
R. annamalai and R. lamarckii, rcspcctivcly. Tcir hy
brid status was initially bascd on intcrmcdiatc morpho
logical charactcrs but has sincc bccn conrmcd in gcnctic
invcstigations.
As discusscd abovc, R. mucronata and R. stylosa arc sibling
spccics with lcw diagnostic charactcrs to scparatc thcm
and a corrcsponding lack ol gcnctic scparation. Tis clcarly
makcs it dicult to scparatc rcspcctivc hybrid lorms with
R. apiculata, although this may bc donc by also using thc
stylc lcngth charactcr. Tc situation is morc complcx, how
cvcr, whcn it must bc considcrcd lurthcr that thcrc arc thc
two lorms ol R. apiculata. Tcsc cach havc hybrids with
R. stylosa bascd on occurrcnccs north and south ol Ncw
Guinca.
!VP stilt mangrovcs arc also charactcrizcd by a numbcr ol
notablc gcnctic, morphological, and growth variants. Ab
crrant traits arc obscrvcd in a small numbcr ol individuals
throughout thc rangc. Pcrhaps thc most consistcnt abcr
rant trait, howcvcr, is albino hypocotyls (i.c., thosc lacking
any grccn pigmcnt, lcaving thcm rcd or ycllow) obscrvcd in
Australia, Fcdcratcd Statcs ol Microncsia, Palau, and othcr
placcs. Ycllow or rcd propagulcs can bc obscrvcd hanging
in acctcd trccs alongsidc normal grccn propagulcs ol all
!VP spccics. Tcir rclativc numbcr on an individual trcc is
thought to quantily thc amount ol outcrossing that occurs
with ncighboring normal trccs. n rarc occasions, othcr
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. i:~cvs: x. c. uuxv
8 !ndoVcst Pacic Rhizophora spccics
trccs havc also bccn obscrvcd with varicgatcd
loliagc.
Culturally important related species
All Rhizophora spccics arc closcly similar in trcc
lorm, and dicrcnt cultural groups in thc Pa
cic rcgion may not always distinguish bctwccn
thcm. thcr mangrovc gcncra, likc Bruguiera,
arc considcrcd closc in lorm and valuc also, and
thcsc arc oltcn uscd in similar ways by indig
cnous pcoplcs.
ASSOCIATED PLANT SPECIES
!VP stilt mangrovcs arc naturally associatcd
with a numbcr ol othcr mangrovc and non
mangrovc spccics across thcir widc rangc ol
tidal and cstuarinc locations. Planting with
othcr spccics is highly rccommcndcd, par
ticularly tcrrcstrial and bcach strand associatcs
growing naturally abovc mcan sca lcvcl clcva
tions. Tc idca is to cstablish and achicvc wholc
olbank stabilization as soon as possiblc. Sincc
stilt mangrovcs also occupy a rangc ol substratc
typcs, including coral, rocks, gravcl, sand, and mud, it is
important to choosc associatcd spccics bcst suitcd to thc
particular local conditions.
Associated species commonly found
Tc associatcd spccics vary with gcographic location, lati
tudc, soil typc, cstuarinc upstrcam location, and tidal posi
tion.
!n lrontal stands, !VP stilt mangrovcs may bc associatcd
with Sonneratia alba and Avicennia marina downstrcam.
Vhcrc downstrcam stands ol stilt mangrovcs arc backcd
by sand ridgcs and bcach margins, thc associatcd spccics
includc Calophyllum spp., Tespesia spp., Casuarina spp.,
Barringtonia asiatica, and Cocos nucifera.
Upstrcam in rivcrinc cstuarics ol tropical coastlincs ol
largcr islands and contincntal margins, associatcd spccics
includc Sonneratia caseolaris, S. lanceolata, and Aegiceras cor-
niculatum. !n such upstrcam locations, stilt mangrovcs arc
commonly backcd by Bruguiera and othcr highcr intcrtidal
mangrovc spccics, as wcll as associatcd upland spccics in
cluding Barringtonia racemosa and Hibiscus tiliaceus.
ENVIRONMENTAL PREFERENCES
AND TOLERANCES
Climate
!VP stilt mangrovcs thrivc in tropical and subtropical
cnvironmcnts charactcrizcd by modcratcly high and wcll
distributcd rainlall. Howcvcr, in dricr locations, likc in
northcrn arcas ol Vcstcrn Australia, thc stuntcd but dcnsc
thickcts ol R. stylosa attcst to thc grcat adaptability ol stilt
mangrovcs to a widc rangc ol climatic typcs.
Elevation range
c6 m (cac lt), in rclcrcncc to mcan sca lcvcl.
Mean annual rainfall
Tcsc mangrovcs grow in all rainlall conditions. Tcir cx
tcnt, lorm, and biomass rccct thc dicrcnt rainlall condi
tions.
Rainfall pattern
Grows in climatcs with summcr or unilorm rainlall pat
tcrns.
Dry season duration (consecutive months with <40
mm [1.6 in] rainfall)
Mangrovc plants appcar to dcpcnd on groundwatcr to sus
tain optimal growth, cspccially during dricr months. Across
a widc varicty ol climatic rcgions, mangrovc covcr cxpands
Foliage of Rhizophora mucronata showing a tree with a rare yellow albino
form of hypocotyls among the normal green hypocotyls. Johnstone River,
NE Australia. vno:o: x. c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 9
and contracts through timc. Tis has bccn cvidcnt in cor
rclations bctwccn l Nino cvcnts and rcduccd growth as
possiblc causcs ol somc damagc to mangrovcs, prcsumably
duc to dccrcascs in lrcshwatcr availability.
Mean annual temperature
accC (66F) (cstimatc)
Mean maximum temperature of hottest month
aC (,.ccF) (cstimatc)
Mean minimum temperature of coldest month
..C (6F)
Minimum temperature tolerated
.cC (cF) (cstimatc)
Soils
Trccs dcvclop grcatcst staturc and columnar growth lorm
in cstuarics ol largcr tropical rivcrs, charactcrizcd by nc
clay, black mud scdimcnts with rclativcly high loads ol or
ganic carbon, and anacrobic soils with high conccntrations
ol suldc. Trccs also grow wcll in sitcs with acrobic scdi
mcnts consisting ol nc sands to coarsc stoncs and rocks,
as wcll as coral ramparts.
Soil texture
Plants grow bcst in light, mcdium, and hcavy tcxturc soils
(sands, sandy loams, loams, sandy clay loams, sandy clays,
clay loams, and clays).
Soil drainage
Plants grow bcst in soils with lrcc and unimpcdcd drain
agc, as wcll as watcrloggcd soils.
Soil acidity
pH 6.
Special soil tolerances
Plants grow bcst in salinc soils but can survivc wcll in lrcsh
watcr. Tc optimal salinity rangc is rcportcd to bc a6 ppt
(parts pcr thousand), comparcd to approximatcly 6 ppt
lor scawatcr.
Tolerances
Drought
Stilt mangrovcs usually grow bcst in wcttcr conditions.
Howcvcr, R. stylosa and wcstcrn rangc R. mucronata can
tolcratc drought pcriods wcll.
Full sun
Tcy usually grow bcst in lull sun.
Shade
Stilt mangrovcs arc usually considcrcd to havc a vcry low
tolcrancc ol shadc. Howcvcr, in castcrn Australia at its
southcrn latitudinal limit, R. stylosa trccs commonly grow
undcr a closcd canopy ol Avicennia marina. Tis may bc
MANGROVES AND CLIMATE
CHANGE
!ndoVcst Pacic stilt mangrovcs, likc othcr mangrovc
spccics, arc acctcd by climatc changc. Tc uniquc
physiological charactcristics ol cach spccics dcnc its
capacity lor survival in thc lacc ol changc. Mangrovcs
arc cxpcctcd to rcspond rapidly and dccisivcly to shilts
in kcy lactors, likc tcmpcraturc, rainlall, and sca lcvcl,
as cach spccics has dcncd rangcs ol tolcrancc lor cach
lactor. For instancc, bccausc mangrovcs arc charactcr
istically rcstrictcd to clcvations bctwccn mcan sca lcvcl
and highcst tidcs, as sca lcvcl riscs thcir communitics
must movc upland to survivc. Sincc mangrovcs havc
narrow optimal tcmpcraturc rangcs, rising tcmpcra
turcs will causc thcir distributions to shilt north or
south to arcas whcrc tcmpcraturc conditions arc most
suitablc, and thcy will dic o in arcas whcrc thcy arc
not suitcd. l coursc, thcir succcss in making thcsc
shilts dcpcnds on thcir succcsslul dispcrsal and rccs
tablishmcnt, and thc availability ol suitablc ncw spacc.
Clcarly, such changcs havc occurrcd throughout his
tory, so thc distribution ol mangrovcs today rcprcscnts
thc survivors ol all past changcs.
Kcy indicators ol changc can bc idcnticd and mappcd
as incrcmcntal shilts and rcsponscs ol mangrovc com
munitics. Tcsc might bc obscrvcd as shilts in vcgcta
tion, lor cxamplc: .) in thc total tidal wctland habitat
zonc, as cxpcctcd with changcs in sca lcvcl, and a) in
thc salt marshmangrovc ccotonc, as cxpcctcd with
changcs in longcrtcrm rainlall pattcrns as this accts
moisturc strcss in salinc cnvironmcnts. !n both cascs,
thc rcsponsc zoncs will lollow clcvation contours.
Changcs along contours can bc quanticd lrom long
tcrm spatial asscssmcnts ovcr dccadc and ccntury
long timc pcriods, dcpcnding on thc ratcs ol changc.
Knowlcdgc ol thcsc changcs and thcir causcs allows
bcttcr prcdiction ol luturc changc.
10 !ndoVcst Pacic Rhizophora spccics
duc to thc abscncc ol scolytid bcctlc larvac that
usually inlcct shadcd propagulcsccdlings in
thc tropics (8rook acc.). !l this is thc casc, it
implics Rhizophora sccdlings might bc morc
shadc tolcrant than originally thought (Sousa
ct al. acc). Tis is supportcd lurthcr by thc
obscrvation that Bruguiera spccics, thc appar
cnt shadc spccialists ol tropical arcas, had lcw
scolytid inlcctions.
Fire
Tc trccs havc no tolcrancc ol rc in closc prox
imity.
Frost
Tcy havc low tolcrancc ol sublrcczing tcm
pcraturcs.
Waterlogging
Stilt mangrovc trccs arc tolcrant ol daily tidal
ooding up to dcpths ol a. m ( lt). Vhilc
tolcrating pcrmancntly saturatcd soils, thcy arc
intolcrant ol drying soils.
Salt spray
Tcy arc highly tolcrant ol salt spray.
Wind
Somc spccics ol stilt mangrovc trccs arc typi
cally lound on thc cxposcd watcrs cdgc ol largc
rivcrinc cstuarics.
Abilities
Self-prune
Stilt mangrovcs scllprunc wcll in dcnsc stands, but thcy
commonly maintain lowcr branchcs in morc opcn loca
tions.
Coppice
Tc trccs havc notably poor coppicc ability. Gcncrally, il
grcatcr than c or morc ol thc lcavcs arc rcmovcd lrom
a trcc, thcy will dic. Tcrc appcars to bc a variant ol R.
apiculata in thc Philippincs and in somc islands ol thc
Fcdcratcd Statcs ol Microncsia (Pohnpci and Kosrac) that
is uscd by local pcoplc lor sustainablc rcwood collcction.
Tcsc trccs rcgrow lrom rcgular cutting, which kccps thcm
stuntcd.
GROWTH AND DEVELOPMENT
Growth ratcs vary with agc. Tc trcc gcncrally grows lcss
than . m/yr in hcight but can cxcccd this in lavorablc cir
cumstanccs. Hcight growth is rapid shortly altcr cstablish
mcnt whilc lood rcscrvcs arc takcn up lrom thc hypocotyls
ol cstablishcd propagulcs. Tc growth ratcs appcar to slow
whcn trccs approach a sitc maximal canopy hcight, dcpcn
dant on spccic sitc/location conditions. Vhcn ncar ma
turity, trccs tcnd to sprcad and broadcn thcir canopy and
stcm diamctcr rathcr than grow tallcr.
Flowering and fruiting
Flowcring and lruiting pcriods ol stilt mangrovcs arc dis
tinctly scasonal. Furthcrmorc, pcak phcnocvcnts shilt
latcr with coolcr tcmpcraturcs and highcr latitudcs. Trccs
havc notablc and rclativcly long pcriods ol rcproductivc
dcvclopmcnt, taking .c months lrom rst cmcrgcncc
ol owcr bud primordia until maturation and drop ol ma
turc hypocotyls. Tc duration dcpcnds on spccics, with thc
Trunk of Rhizophora mucronata showing distinctive stilt roots. Kenya, E
Africa. vno:o: x. c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 11
longcst bcing Rhizophora apiculata, with around c months
lor cach rcproductivc cyclc.
Yields/growth rates
Growth ratcs vary with spccics, spatial position in thc
stand, compctition, vigor, and agc. !n thc Matang Man
grovc Forcst Rcscrvc in Malaysia, Putz and Chan (.6)
rcportcd that diamctcr growth ratcs ol R. apiculata trccs
wcrc c.ac.a cm (c.cc... in) lor diamctcr sizc classcs
lrom .c to 6c cm (a in).
Vatson (.a) cstimatcd that undcr Malaysian conditions
mcan annual incrcmcnt (MA!) ol stilt mangrovcs culmi
natcd at around .c.6 m

/ha/yr (.a lt

/ac/yr) at c ycars.
Reaction to competition
Rapid carly growth ol sccdlings ol stilt mangrovcs in lull
sunlight cnsurcs thcir succcss and dominancc in prclcrrcd
cstuarinc and intcrtidal conditions. Ncwly cstablishcd
sccdlings grow bcst in closc proximity with thcir samc
spccics cohort. Tis aords thcm maximal protcction lrom
physical damagc by drilt logs and crosivc wavcs. Sincc
compctition is high bctwccn ncighboring sccdlings, slowcr
plants dic and dccomposc quickly, lcaving lastcr compcti
tors thc bcnct ol not only thc spacc thcy occupicd but
also thcir nutricnts.
PROPAGATION
!VP stilt mangrovcs arc rcadily propagatcd by dircct
planting ol thcir propagulcs. Although natural
rcgcncration is gcncrally rclicd upon around thc
Pacic rcgion, thcsc spccics arc rclativcly casy to
propagatc. Propagation is simplc and rclics on
thc spccial lcaturc ol thc gcnus in having largc
viviparous propagulcs. Planting simply cntails
gcntly pushing thc distal cnd ol thc acc cm
(. in) long hypocotyl onc third ol its lcngth
into thc scdimcnt, spaccd at about ... m (.
lt) intcrvals. No holcs nccd bc dug, and ncithcr
nurscry prcparation nor stakcs arc nccdcd. Low
maintcnancc is gcncrally rcquircd lor maximiz
ing sccdling cstablishmcnt succcss in shcltcrcd
arcas. Howcvcr, substantial protcction is rc
quircd in morc cxposcd coastal locations during
thc rst dccadc ol cstablishmcnt. Such protcc
tion mcthods may includc cncascmcnt ol indi
vidual sccdlings in P\C piping (Rilcy and Kcnt
.), or installation ol tcmporary structurcs to
dampcn wavc action and rcducc dcbris drilt
across rcstoration sitcs as obscrvcd in \ictnam and China
(Ficld .6).
Propagule collection
Propagulcs may bc availablc throughout thc ycar, but pcak
production occurs around JulyAugust in northcrn parts
ol thc rangc and around JanuaryFcbruary in thc southcrn
hcmisphcrc. Maturc propagulcs may bc collcctcd altcr thcy
havc lallcn or bccn pickcd dircctly o trccs. nly hcalthy
looking propagulcs should bc sclcctcd. Propagulcs that
arc shrunkcn or dcsiccatcd in appcarancc or that cxhibit
signicant physical damagc should bc rcjcctcd. Although
propagulcs with only minor borcr damagc may survivc and
grow, sclcction ol propagulcs with no signs ol borcr or crab
damagc is strongly prclcrrcd. Propagulcs that alrcady havc
somc root or lcal dcvclopmcnt can bc uscd in most cascs
but should not bc storcd lor long.
Propagule processing
Proccssing ol maturc propagulcs is not rcquircd lor stilt
mangrovcs, although damagcd and inscct inlcstcd individ
uals should bc rcmovcd. Also scc thc lollowing prctrcat
mcnt scction lor possiblc additional stcps. Propagulcs can
bc sown in nurscry bcds, or prclcrably plantcd dircctly in
thc cld soon altcr collcction.
Propagule storage
Propagulcs can bc kcpt viablc lor at lcast 6, days by stor
ing thcm in brackish watcr or by wrapping thcm in wct
Left: Mature hypocotyl and fruits of Rhizophora mucronata showing dis-
tinctive collar on attached propagule and expended fruit. vno:o: x. c. uuxv
Right: A sectioned wilting seedling of Rhizophora stylosa shows the extent
of scolytid infection marked by brown frass. vno:os: n. :. nvoox
12 !ndoVcst Pacic Rhizophora spccics
burlap bags and kccping thcm out ol dircct
sunlight. Prctrcatmcnt is considcrcd csscntial il
considcring such storagc (scc bclow). !t is likcly
that propagulcs can bc kcpt longcr, but storagc
bcyond a wccks is not rccommcndcd, and long
tcrm storagc is not lcasiblc.
Propagule pretreatment
Prctrcatmcnt ol propagulcs is gcncrally con
sidcrcd unncccssary. Howcvcr, a study ol sco
lytid bcctlc larvac (Coccotrypes fallax) inlcctions
on Rhizophora propagulcs in Australia lound
at lcast . that wcrc hcavily inlcstcd (8rook
acc.). !nlcstcd propagulcs wcrc cstablishcd un
dcr canopyshadcd arcas. Tc study wcnt lur
thcr, nding that prctrcatmcnt in a cC (.aaF)
watcr bath lor minutcs killcd thc bcctlcs and
rcmovcd thc risk ol cstablishmcnt loscs duc to
borcr damagc. Hcat trcatmcnt might bc casily
achicvcd by lcaving thc collcctcd propagulcs in
thc sun lor a short pcriod (a lcw hours) bclorc
planting.
Growing area
Stilt mangrovcs should bc grown in lull sunlight.
Seedling establishment
Lcavcs and roots may bcgin to dcvclop within a wcck or
two ol sowing.
Media
Although a widc varicty ol soil mcdia arc acccptablc, a mix
ol sand and pcat in cqual parts has bccn rccommcndcd lor
mangrovc nurscrics.
Time to outplanting
Anytimc during thc ycar is suitablc. Sccdlings arc rcady
lor outplanting at thc 6lcal (- nodc) stagc il grown up
in nurscry conditions. ldcr sccdlings up to c. m (ac in)
tall havc also bccn succcsslully transplantcd, but this is not
considcrcd bcnccial.
ircct planting ol largc numbcrs ol propagulcs is rcstrict
cd by thcir pcak scasonal availability, as propagulcs do not
kccp lor cxtcndcd pcriods unlcss plantcd out in nurscry
conditions.
Guidelines for outplanting
Propagulcs or nurscrygrown sccdlings usually havc cxccl
lcnt survival in sitcs corrcctly sclcctcd and, il appropriatc,
protcctcd lrom disturbancc. Survival ratcs ol c or grcat
cr arc not unrcasonablc in such circumstanccs.
n thc othcr hand, survival may bc zcro on sitcs cxposcd
to cxccssivc wavc action, on sitcs with inappropriatc hydro
logic or salinity rcgimcs, or (rarcly) disturbancc by grazing
animals (c.g., goats, shccp, cattlc, and horscs). A mcthod ol
cncasing propagulcs in P\C pipc has bccn uscd in Florida
and thc Caribbcan to protcct sccdlings in placcs with a
high likclihood ol disturbancc.
DISADVANTAGES
!n gcncral, thc planting ol stilt mangrovcs poscs lcw signil
icant disadvantagcs whcn plantcd within thcir nativc rang
cs. Tcy arc not cspccially susccptiblc to pcsts or pathogcns
and thcy havc not bccn rcportcd to host major pcsts or
pathogcns ol important crop spccics.
Potential for invasiveness
Although thc invasivcncss ol stilt mangrovcs has not bccn
dcmonstratcd, thcy arc likcly to bc opportunistic duc to
thcir rclativcly widc tolcrancc lor salinity and soil condi
tions. Tc sprcad ol thc closcly allicd AP rcd mangrovc
spccics R. mangle whcn introduccd into Hawaii has clcarly
dcmonstratcd thc potcntial lor invasivcncss ol thcsc man
grovcs. Although thcy wcrc not nativc, suitablc growing
conditions cxistcd, dcmonstrating that thc global distribu
tion ol Rhizophora spp. is currcntly limitcd by thcir natural
New planting on seaward edge of R. apiculata at Ao Khung Krabaen Man-
grove Forestry Center, Som Lek, Tailand. vno:o: c. vivvi:cn
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 13
dispcrsal rangc. Tis also mcans gcographically isolatcd
populations arc vulncrablc to introductions ol gcnotypcs
lrom clscwhcrc.
Susceptibility to pests/pathogens
Susccptibility to pcsts and discascs is bclicvcd to bc low,
with thc cxccption ol inscct borcrs and crabs that lccd on
propagulcs.
Host to crop pests/pathogens
No rcports wcrc lound ol stilt mangrovcs scrv
ing as hosts lor known major crop pcsts or
pathogcns. Tc scolytid bcctlcs arc spccic to
thc gcnus Rhizophora.
AGROFORESTRY/
ENVIRONMENTAL
PRACTICES
Mulch/organic matter
Mulch in Rhizophora lorcsts is hiddcn lrom vicw.
!l it wcrc not lor thc small mangrovc crabs, lall
cn lcavcs would bc washcd away with cach tidc.
Tc crabs activcly takc lcavcs into undcrground
burrows and chambcrs. Tc rcsulting mulch is
rapidly colonizcd by bactcria and consumcd by
othcr burrowing launa to rclcasc nutricnts that appcar to
lurthcr cnhancc thc lorcst.
Soil stabilization
Stilt mangrovc lorcsts stabilizc soils with thcir nctwork
ol sturdy ovcrlapping prop roots dampcning watcr movc
mcnt and promoting scdimcntation in arcas that might
othcrwisc bc crodcd.
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. vno:o: x. c. uuxv
8cncts includc, in no particular ordcr (adaptcd lrom
Tomlinson .6):
visual amcnity and shorclinc bcautication
nutricnt uptakc, xation, trapping, and turnovcr
habitat usc by launa
mcsoclimatc, whcrc lorcsts modcratc cvapotranspira
tion to crcatc a spccializcd nichc climatc
nurscry habitat lor young launa, whcrc mangrovcs
providc a sourcc ol lood and physical protcction lrom
prcdation
sanctuary nichc lor maturc launa, including migratory
birds and sh, whcrc mangrovcs providc protcction
and a lood rcsourcc
primary production bascd on photosynthcsis, giving
risc to lorcst growth and lorcst products, notably tim
bcr
sccondary production, including microbial and launal
production, as wcll as grazcrs, and via dccomposition
shcry products, including both cstuarinc and coastal
shorclinc protcction, bascd on gcncral mangrovc trcc
and root structurc, as wcll as cdgc trccs, which rcducc
crosion and providc stand protcction lrom wavcs and
watcr movcmcnt
carbon scqucstration and a sink whcrc carbon is bound
within living plant biomass
scdimcnt trapping, bascd on mangrovcs bcing a dcp
ositional sitc lor both watcr and airbornc scdimcnts,
which in turn rcduccs turbidity ol coastal watcrs.
BENEFITS OF MANGROVE TIDAL WETLAND
14 !ndoVcst Pacic Rhizophora spccics
Fence posts
Stilt mangrovc stcms makc good posts sincc thcy arc gcn
crally hard wood and rcsistant to inscct borcrs.
Windbreaks
Rhizophora lorcsts providc a windbrcak along coastal mar
gins, gcncrally, and as placcs to scck sanctuary during ty
phoons and cycloncs. Planting in highly windpronc loca
tions is not rccommcndcd unlcss thc location has somc
history, or rcasonablc cxpcctation, that mangrovc survival
is likcly.
Woodlot
Mangrovcs adjaccnt to pcoplcs homcs throughout thc Pa
cic lrcqucntly scrvc as inlormal woodlots, particularly on
islands with clcar tcnurc systcms that includc mangrovc
arcas. Stilt mangrovc wood is vcry usclul lor small con
struction, and lor cooking lucl. Convcrting it to charcoal
can lurthcr cnhancc thc timbcrs caloric valuc. Tis is
donc commcrcially in S Asia and Ccntral Amcrica with
various Rhizophora spccics.
Native animal/bird food
!VP stilt mangrovcs arc largcly an unknown sourcc ol
nativc animal loods. Howcvcr, scvcral obscrvations dcm
onstratc that thc divcrsity and quantity ol thc sourcc is
thought to bc cxtrcmcly important in mangrovc ccosys
tcms. Numcrous insccts, crabs, and mollusks grazc on
grccn lcavcs in thc lorcst canopy. Scsarmid crabs consumc
a largc quantity ol lallcn lcavcs and propagulcs. rganic
mattcr proccsscd by thcsc hcrbivorcs is bclicvcd to broadly
support aquatic lood chains in coastal rcgions. Fcw mam
mals appcar to usc stilt mangrovcs as a major lood sourcc,
although nativc rats oltcn chcw into thc wood in scarch ol
boring inscct larvac.
Wildlife habitat
!n addition to aquatic marinc organisms (scc Fish/marinc
lood chain), stilt mangrovcs scrvc as habitat lor a widc
rangc ol tcrrcstrial and arborcal wildlilc. !n various loca
tions throughout thc rcgion, thcsc lorcsts providc shcltcr
and lood lor a numbcr ol associatcd launa, including birds,
lruit bats, small mammals, shcllsh, and othcr marinc lilc.
Bee forage
Rhizophora spccics havc no ncctar, but thcy do producc co
pious pollcn that is usually distributcd by wind.
Fish/marine food chain
Mangrovcs in gcncral arc bclicvcd to play a vitally impor
tant rolc in protccting and supporting marinc lood chains.
Many sh spccics usc stilt mangrovcs during part ol thcir
lilc cyclcs, as do spccics ol shrimp and crabs. Spccics such
as thc mangrovc mud crab (Scylla serrata) arc common in
!VP mangrovcs, and thcsc arc important sourccs ol lood
and incomc on many islands in thc rcgion. Populations
ol somc smallcr spccics may cxcccd .c crabs/m
a
(. crab/
lt
a
) in parts ol thc Pacic. Scncsccnt lcavcs having lallcn
lrom Rhizophora trccs arc takcn by grapsid (small man
grovc) crabs into thcir burrows. 8uricd lcavcs dccomposc
and contributc to nutricnt rccycling in mangrovc lorcsts.
Nutricnts also lccd dircctly and indircctly to associatcd cs
tuarinc and marinc lood chains.
Coastal protection
Stilt mangrovc lorcsts, and mangrovcs in gcncral, play an
important rolc in protcction ol coastlincs, shponds, and
othcr coastal inlrastructurc. Stilt mangrovcs arc plantcd
lor coastal or shpond protcction in somc arcas (c.g., in
Kalibo 8akhawan co Park, thc Philippincs) and thcrc arc
laws in many locations aimcd at protccting mangrovcs in
largc part bccausc ol this important lunction.
Ornamental
No signicant ornamcntal tradc is known. Howcvcr, as an
Rhizophora stylosa sprouted propagules for sale in Iriomote,
Japan. vno:o: x. c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 15
apparcntly isolatcd cxamplc, a small numbcr ol plants arc
sold as souvcnirs to dcparting tourists lrom !riomotc and
othcr southcrn islands ol Japan. Sproutcd hypocotyl sccd
lings arc uscd to makc bonsai lorcst gardcns.
USES AND PRODUCTS
!VP stilt mangrovcs arc probably ol grcatcst valuc lor thcir
cnvironmcntal bcncts, bccausc thcy (and mangrovcs gcn
crally) arc bclicvcd to play a vital rolc in sup
porting marinc lood chains, protccting coastal
arcas, and improving watcr quality.
!n tcrms ol dircct bcncts to pcoplc, thc most
widcsprcad usc ol stilt mangrovcs is lor wood
lor a rangc ol purposcs lrom cooking lucl to
construction ol homcs and canoc parts. thcr
uscs ol thc stilt mangrovcs includc tannin and
dycs. A lcss laudcd bcnct has bccn dcrivcd
lrom somc ccotourism vcnturcs. Tc dual bcnc
t in this practicc comcs lrom both a modcratc
cconomic rcturn combincd with a substantial
longcrtcrm cducational valuc in raising cnvi
ronmcntal awarcncss in thc community.
Staple food
Lcavcs and hypocotyls arc cdiblc but not widcly
uscd lor lood.
Medicinal
Stilt mangrovc bark has rcportcdly bccn uscd
to trcat angina, boils, and lungal inlcctions. Tc
lcavcs and bark havc bccn uscd as an antiscptic
and to trcat diarrhca, dyscntcry, lcvcr, malaria,
and lcprosy, although it is not clcar how ccc
tivc thc trcatmcnts havc bccn in cach ol thcsc
cascs.
Timber
Tc wood ol stilt mangrovcs is widcly uscd
lor structural componcnts (c.g., polcs, bcams,
ooring, wallcladding, and raltcrs) ol tradi
tional homcs and othcr structurcs likc undcr
ground minc supports, lcncing, cabinct works,
tool handlcs, and boat anchors. Tc wood is also
uscd lor othcr purposcs, ranging lrom tradi
tional uscs such as shing stakcs, spcars, and co
prahuskcrs to usc as a sourcc ol chips lor pulp
production. Howcvcr, pulp lrom Rhizophora
spccics docs not havc good strcngth propcrtics,
so thcsc spccics arc not oltcn sought altcr lor
this purposc (Pcrcival and Vomcrslcy .,).
Fuelwood
Stilt mangrovc wood is uscd lor luclwood on many Pacic
islands (c.g., Kosrac and Chuuk). Tc wood is also madc
into charcoal in countrics such as thc Philippincs, !ndonc
sia, and Malaysia (ng ct al. .c, .).
Canoe/boat/raft making
Tc wood has bccn uscd to makc canoc parts.
Top: Mangrove tourism in Hainan, China, complete with Rhizophora
motifs on the jetty handrail panels. vno:o: x. c. uuxv Bottom: Boardwalk
and signage for visitors (R. mucronata on left and R. apiculata on right), Ao
Khung Krabaen Mangrove Forestry Center, Som Lek, Tailand. vno:o: c.
vivvi:cn
16 !ndoVcst Pacic Rhizophora spccics
Tannin/dye
Tc bark and hypocotyls arc uscd to producc dycs ranging
lrom rcdbrown to black (thc lattcr with rcpcatcd dying).
Rhizophora spccics arc thc bcst produccrs ol tannins ol all
thc mangrovcs with aca ol bark wcight (Pcrcival and
Vomcrslcy .,). Choppcd bark is trcatcd by continuous
countcrcurrcnt cxtraction with rst cold thcn warm wa
tcr producing an cxtract callcd cutch. Tc cutch can bc
cxportcd lor proccssing and nal tannin cxtraction. Tan
nins wcrc uscd cxtcnsivcly lor prcscrving cotton ropcs and
nctting. Mangrovc tannins can bc uscd to prcparc tannin
lormaldchydc adhcsivcs. Tcsc adhcsivcs arc rccognizcd
lor thcir high moisturc rcsistancc and watcrprool gradcs in
plywood and particlcboard production. Rhizophora tannins
arc also known to impart a rcd color to nishcd lcathcr
tanning, but this is considcrcd undcsirablc sincc blcnding
with othcr tannin cxtracts is nccdcd.
Ecotourism
Tc mangrovc cnvironmcnt is attractivc and intcrcsting
to both local pcoplc and to visiting tourists. !n China, lor
cxamplc, lacilitics havc bccn providcd to assist and rcgu
latc this lorm ol ccotourism, including a dcdicatcd hotcl
and rcstaurants. Simplc boardwalks in othcr placcs (c.g.,
Australia, Tailand) arc rclativcly low kcy but thc intcn
tion is thc samc, to providc public acccss lor thosc wanting
to scc and lcarn about this lascinating cnvironmcnt. Tc
boardwalks arc thcrc to makc thc mangrovc cxpcricncc as
cortlcss as possiblc whilc protccting thc mangrovcs lrom
dircct damagc. Valks arc oltcn also accompanicd with in
lormativc and cducational signagc.
COMMERCIAL CULTIVATION
Stilt mangrovc timbcr is harvcstcd commcrcially lor char
coal production in S Asia. Tc caloric valuc ol thc tim
bcr is signicantly cnhanccd by convcrting it to charcoal.
Tis is donc with various Rhizophora spccics. Charcoal
production has bccn conductcd in a sustainablc way on
thc Malay Pcninsula lor ovcr .cc ycars using silvicultural
practiccs dcvclopcd lor Rhizophora spccics, cspccially R.
apiculata (ng ct al. .c, .). !n addition, lorcsts in Ma
laysia, Ncw Guinca, and thc Solomon !slands havc bccn
clcarlcllcd lor wood chips using unsustainablc harvcsting
practiccs.
Spacing
Mangrovc plantations in gcncral arc typically plantcd at
spacings ol about ..c.. m (. lt). Spacing widcr than
about a. m ( lt) tcnds to rcsult in a high proportion ol
multiplcstcmmcd and/or shortcr trccs. Vidcr spacing
and thc rcsulting sprcading trccs may bc dcsircd lor coastal
protcction projccts, but not lor timbcr production. !n thc
abscncc ol signicant natural mortality timbcr plantations
should bc thinncd to spacing ol a. to . m (.. lt) bc
twccn trccs as thc stand dcvclops and bccomcs crowdcd.
Management objectives
!n arcas whcrc thc mangrovc lcrn (Acrostichum speciosum)
is common, it may nccd to bc controllcd to promotc carly
growth ol stilt mangrovcs. Somc publishcd guidclincs lor
mangrovc silviculturc cxist and arc rclcrcnccd bclow, but
spccic guidclincs on thinning, lcrtilizing, ctc., arc cur
rcntly unavailablc.
Growing in polycultures
Stilt mangrovcs naturally occur in mixcdspccics stands,
and cach spccics has its own ccological and cconomic val
ucs. Mixcdspccics plantings arc rccommcndcd togcthcr
with largclcal mangrovc, Bruguiera gymnorrhiza. !t is also
Eleven-year-old plantation of Rhizophora apiculata in the
ha ( ac) Bakhawan Eco Park at Kalibo, northern Panay,
Philippines. vno:o: x. c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 17
important to plant associatcd bucr arcas, cspccially along
thc shorclinc whcrc mangrovcs grow bcttcr adjaccnt to
banks stabilizcd by shorclinc upland plants. Suitablc up
land trcc gcncra includc Calophyllum, Casuarina, Hibiscus,
Tespesia, and Barringtonia. Vithin thc mangrovc lorcst,
natural rccruitmcnt will oltcn bring additional mangrovcs
such as Bruguiera spccics, but thcsc might bc plantcd as
wcll. Togcthcr thcy will complcmcnt and cnhancc thc
richncss and stability ol thc plantcd cnvironmcnt.
Estimated yield
Sncdakcr and 8rown (.a) statc that mangrovc lorcst prod
ucts in 8angladcsh havc an annual valuc ol USs6,ccc,ccc.
ng (.a) rcports that thc Matang Rcscrvc in Malaysia
with ,ccc ha (6,ccc ac) ol productivc, almost monospc
cic, lorcsts ol R. apiculata, thc valuc ol mangrovc wood
alonc is closc to USs,ccc,ccc pcr ycar.
Tc Matang Mangrovc Forcst Rcscrvc in Malaysia has
bccn managcd lor timbcr production ol chicy Rhizophora
apiculata sincc thc bcginning ol thc ccntury and is rcputcd
ly thc bcst managcd mangrovc lorcst in thc world (Khoon
and ng ., ng ct al. .). Tc avcragc stand volumc
lor thcsc stilt mangrovcs was . m

/ha (a.c lt

/ac). Stand
volumcs up to aa6 m

/ha (ac lt

/ac) havc bccn rcportcd

lrom Tailand.
Tc prcscnt managcmcnt plan lor thc Matang Rcscrvc
is a cycar rotation pcriod with two thinnings, at . and
ac ycars. Tc cycar rotation harvcstcd yiclds havc bccn
around .6a mt/ha (6.. t/ac) ol grccn wood. How
cvcr, thcrc has bccn a dcclinc in yicld lrom a mt/ha (.
t/ac) lrom virgin stands to thc sccondgcncration yiclds ol
. mt/ha (,c t/ac) in .6,6 to an cvcn lowcr .6 mt/ha
(6. t/ac) in .,c,,. 8ccausc thc standing biomass ol thc
trccs did not incrcasc lrom a ycars (. mt/ha |6 t/ac|)
to a ycars (. mt/ha |6 t/ac|), it was suggcstcd that a
rotation ol a ycars bc uscd instcad ol thc prcvious rccom
mcndation ol c ycars.
Markets
Markcts on most Pacic islands arc local in naturc, with
littlc in thc way ol stilt mangrovc products (othcr than in
dircct products likc mangrovc crabs and lruit bats) bcing
cxportcd lrom onc island to anothcr. Tc cxccption is prob
ably rcwood and charcoal, which is availablc lrom local
markcts in most placcs. !n Southcast Asia, largc quantitics
ol stilt mangrovc wood chips and charcoal may bc movcd
grcatcr distanccs and in grcatcr volumcs than wood prod
ucts on smallcr islands.
INTERPLANTING/FARM
APPLICATIONS
Somc intcrplanting systcms includc:
Example Bakhawan Eco Park (Primavcra ct al.
acc)
Location
Aklan, Ncw 8uswang, and Kalibo, Philippincs
Description
Planting was undcrtakcn in . and .. Spccics plant
cd includcd R. apiculata and R. mucronata on a total arca
plantcd ol 6 ha (.6 ac). Sponsors and implcmcntcrs wcrc
NR, PACAP AusA!, USVAG, and KASAMA.
Crop/tree interactions
Not known.
Spacing/density of species
Spacing was about .. m ( lt) lor both spccics.
Example (Primavcra ct al. acc)
Location
Guimaras and Nucva \alcncia, Philippincs
Description
Tc planting was undcrtakcn in .. Spccics plantcd in
cludcd R. apiculata and R. mucronata on a total arca ol
. ha (6 ac). Funding sourcc and implcmcntcrs wcrc
NRCP.
Crop/tree interactions
Not known.
Spacing/density of species
Spacing was around .. m ( lt) lor both spccics.
Example (Primavcra ct al. acc)
Location
!loilo and Carlcs, Philippincs
Description
Tc planting was donc in acc.. Spccics plantcd includcd
R. apiculata and R. mucronata on a total arca ol c ha
(.cc ac). Funding sourcc and implcmcntcrs wcrc NR
Forcstry Scctor Program: MACA8ATAARM Fcdcratcd
Pcoplcs rganisation.
18 !ndoVcst Pacic Rhizophora spccics
Crop/tree interactions
Not known.
Spacing/density of species
Spacing was around .. m ( lt) lor both spccics.
PUBLIC ASSISTANCE
Ccntrc lor Marinc Studics, Marinc 8otany Group
Tc Univcrsity ol Quccnsland
St. Lucia QL c,a
Australia
Vcb: http://www.marinc.uq.cdu.au/marbot/indcx.htm~.
BIBLIOGRAPHY
( indicates recommended reading)
8andaranayakc, V.M. .. Survcy ol mangrovc plants
lrom Northcrn Australia lor phytochcmical constitucnts
and U\absorbing compounds. Currcnt Topics in Phyto
chcmistry .: 6,.
8andaranayakc, V.M. .. Traditional and mcdicinal uscs
ol mangrovcs. Mangrovcs and Salt Marshcs a: ...
8andaranayakc, V.M. .. conomic, traditional and mc
dicinal uscs ol mangrovcs. A!MS Rcport a. Australian
!nstitutc ol Marinc Scicncc, Townsvillc, Australia.
8oonnitcc, V. .,. Tc Rhizophoras ol Tailand. 8iotrop.
Spcc. Publ. .c: a..
8osirc, J.., J. Kazungu, N. Kocdam, and F. ahdouhGuc
bas. acc. Prcdation on propagulcs rcgulatcs rcgcncration
in a highdcnsity rclorcstcd mangrovc plantation. Marinc
cology Progrcss Scrics a: ...
8rook, 8.M. acc.. Tc ccct ol Coccotrypes fallax (Colcop
tcra, Scolytidac) on thc rccruitmcnt ol Rhizophora stylosa
(Family Rhizophoraccac) in North Quccnsland man
grovcs. Ph Tcsis. School ol 8iological Scicnccs, Jamcs
Cook Univcrsity, Townsvillc, Quccnsland.
Chan, H.T. .. Mangrovc lorcst managcmcnt in Malaysia.
!TT Tropical Forcst Updatc (a): .a.
Clarkc, A., and L. Johns. acca. Mangrovc nurscrics: Con
struction, propagation and planting. Fish Habitat Guidc
lincs FHG cc. cpartmcnt ol Primary !ndustrics,
Quccnsland Fishcrics Scrvicc. http://www.dpi.qld.gov.
au/shwcb/.cca.html~.
Clough, 8.F., and K. Scott. .. Allomctric rclationship lor
cstimating abovcground biomass in six mangrovc spccics.
Forcst col. & Manag. a,(a): ..,.a.
Cuong, H.\.\. .6. Rhizophoraccac. Nat. Mus. Nat. Hist.
: ...,.
ahdouhGucbas, F., S. Hcttiarachchi, . Lo Sccn, .
8atclaan, S. Sooriyarachchi, L.P. Jayatissa, and N. Kocdam.
acc. Transitions in ancicnt inland lrcshwatcr rcsourcc
managcmcnt in Sri Lanka acct biota and human popu
lations in and around coastal lagoons. Currcnt 8iology
.(6): ,6.
ahdouhGucbas, F., L.P. Jayatissa, .. Nitto, J.. 8o
sirc, .L. Sccn, and N. Kocdam. acc. How ccctivc wcrc
mangrovcs as a dclcncc against thc rcccnt tsunami: Cur
rcnt Scicncc .: ,.
ahdouhGucbas, F., C. Mathcngc, J.G. Kairo, and N.
Kocdam. accc. Utilization ol mangrovc wood products
around Mida Crcck (Kcnya) amongst subsistcncc and
commcrcial uscrs. conomic 8otany (): .a,.
ahdouhGucbas, F., M. \crncirt, J.F. Tack, and N. Kocdam.
.,. Food prclcrcnccs ol Neosarmatium meinerti dc Man
(ccapoda : Scsarminac) and its possiblc ccct on thc rc
gcncration ol mangrovcs. Hydrobiologia ,: .
ahdouhGucbas, F., M. \crncirt, J.F. Tack, . \an Spcy
brocck, and N. Kocdam. .. Propagulc prcdators in Kc
nyan mangrovcs and thcir possiblc ccct on rcgcncration.
Marinc and Frcshwatcr Rcscarch (): c.
c Laccrda, L.. acca. Mangrovc cosystcms: Function
and Managcmcnt. Springcr\crlag, 8crlin, Gcrmany.
ing, H. .6c. A rcvicw ol thc gcnus Rhizophorawith
spccial rclcrcncc to thc Pacic spccics. 8lumca .c(a):
6a6.
ukc, J.A. n.d. Rhizophora mucronata Lam. !n: Handbook
ol ncrgy Crops. http://www.hort.purduc.cdu/ncwcrop/
dukc_cncrgy/Rhizophora_manglc.html~.
ukc, N.C. .a. Mangrovc oristics and biogcography. !n:
Robcrtson, A.!. and .M. Alongi (cds.). Tropical Man
grovc cosystcms. Amcrican Gcophysical Union, Vash
ington, C.
ukc, N.C. .. Tc . survcy ol Rhizophora spccics in
Microncsia. Rcport to thc USA Forcst Scrvicc. Marinc
8otany Group, 8otany cpartmcnt, Tc Univcrsity ol
Quccnsland, 8risbanc, Australia.
ukc, N.C., and J.S. 8unt. .,. Tc gcnus Rhizophora
(Rhizophoraccac) in northcastcrn Australia. Australian
Journal ol 8otany a,: 6,6,.
ukc, N.C., and Z.S. Pinzn. .a. Aging Rhizophora
sccdlings lrom lcal scar nodcs: a tcchniquc lor studying
rccruitmcnt and growth in mangrovc lorcsts. 8iotropica
a(aa): .,.6.
ukc, N.C., M.C. 8all, and J.C. llison. .. Factors in
ucncing biodivcrsity and distributional gradicnts in
mangrovcs. Global cology and 8iogcography Lcttcrs ,:
a,,.
ukc, N.C., J. S. 8unt, and V. T. Villiams. .. bscrva
tions on thc oral and vcgctativc phcnologics ol north
castcrn Australian mangrovcs. Australian Journal ol 8ot
any a: ,.
ukc, N.C., .Y.Y. Lo, and M. Sun. acca. Global distribu
tion and gcnctic discontinuitics ol mangrovcscmcrging
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 19
pattcrns in thc cvolution ol Rhizophora. Trccs Structurc
and Function .6: 6,.
urant, C.C.L. ... Tc growth ol mangrovc spccics in
Malaya. Malay. For. .c: ..
llison, J.C. .. Status ol mangrovcs in thc South Pacic.
Asian Vctland Ncws ,: .
llison, J.C. .. Systcmatics and distributions ol Pacic
!sland mangrovcs. !n: Maragos, J.., M.N.A. Pctcrson,
L.G. ldrcdgc, J.. 8ardach, and H.F. Takcuchi (cds.),
Marinc and Coastal 8iodivcrsity in thc Tropical !sland
Pacic Rcgion, vol. .. Pacic Scicncc Association, ast
Vcst Ccntcr, Honolulu.
llison, J.C. .. Prcscnt status ol Pacic !sland mangrovcs.
Marinc/Coastal 8iodivcrsity \ol. !!. pp .. !n: ldrcdgc,
L.G., J.. Maragos, and P.L. Holthus (cds.). Tc Tropi
cal !sland Pacic Rcgion: Population, cvclopmcnt and
Conscrvation Prioritics. Pacic Scicncc Association, ast
Vcst Ccntcr, Honolulu.
Ficld, C.. (cd.). .6. Rcstoration ol mangrovc ccosystcms.
!ntcrnational Socicty lor Mangrovc cosystcms !SM,
kinawa, Japan, and thc !ntcrnational Tropical Timbcr
rganisation (!TT).
Fosbcrg, F.R. .,. Phytogcography ol Microncsian man
grovcs. !n: Valsh, G., S. Sncdakcr, and H. Tcas. (cds.). pp.
aa. Procccdings ol thc !ntcrnational Symposium on
8iological Managcmcnt ol Mangrovcs. !nstitutc ol Food
and Agricultural Scicncc, Gaincsvillc, Florida.
Gong, V.K., J.. ng, C.H. Vong, and G. hanarajan.
.c. Productivity ol Mangrovc Trccs and its Signicancc
in a Managcd Mangrovc cosystcm in Malaysia. Man
grovc nvironmcntal Rcscarch and Managcmcnt, Uni
vcrsity ol Malaya, Kuala Lumpur and UNSC.
Guppy, H.8. .c6. bscrvations ol a Naturalist in thc Pa
cic bctwccn .6 and .. \ol. !!. Plant dispcrsal. Mac
millan and Co., London.
Hogarth, P.J. .. Tc 8iology ol Mangrovcs. xlord
Univcrsity Prcss, Ncw York.
Kairo, J.G., F. ahdouhGucbas, J. 8osirc, and N. Kocdam.
acc.. Rcstoration and managcmcnt ol mangrovc sys
tcmsA lcsson lor and lrom thc ast Alrican rcgion.
South Alrican Journal ol 8otany 6,: .
Kathircsan, K. .. Rhizophora annamalayana: a ncw spccics
ol mangrovcs. nvironmcnt and cology .(.): aca..
Khoon, G. V., and J.. ng. .. Tc usc ol dcmographic
studics in mangrovc silviculturc. Hydrobiologia a: a
a6..
Landon, F.H. .. Mangrovc volumc tablcs. Malayan For.
..(): ..,.ac.
LaRuc, C.., and T.J. Muzik. ... ocs thc mangrovc rc
ally plant its sccdlings: Scicncc ..: 66.66a.
Lo, ., N.C. ukc, and M. Sun. acca. Phylogcnctic cvalu
ation ol Rhizophora taxa lor conscrvation managcmcnt.
Socicty lor Conscrvation 8iology, Unitcd Kingdom.
Marco, H.F. .. Systcmatic anatomy ol thc woods ol thc
Rhizophoraccac. Trop. Voods : .ac.
McCuskcr, A. .. Rhizophoraccac. !n: Gcorgc, A.S. pp.
..c. Flora ol Australia aa. 8urcau ol Flora and Fauna,
Australian Govcrnmcnt, Canbcrra, Australia.
Morton, J.F. .6. Can thc rcd mangrovc providc lood, lccd
and lcrtilizcr: conomic 8otany .: ...a.
Noakcs, .S.P. .. Mcthods ol incrcasing growth and ob
taining natural rcgcncration ol thc mangrovc typc in Ma
laysia. Malay. For. .: ac.
ng, J.. .a. Mangrovcs and aquaculturc in Malaysia.
Ambio ..(): aaa,.
ng, J.., G.V. Khoon, and 8.F. Clough. .. Structurc
and productivity ol a acycarold stand ol Rhizophora
apiculata 8l. mangrovc lorcst. Journal ol 8iogcography aa:
.,a.
Panapitukkul, N., C.M. uartc, U. Tampanya, P. Khc
owvongsri, N. Srichai, . GccrtzHanscn, J. Tcrrados, and
S. 8oromthanarath. .. Mangrovc colonization: man
grovc progrcssion ovcr Pak Phanang (S Tailand) mud
at. stuarinc, Coastal and Shcll Scicncc ,: .6..
Pcrcival, M., and J.S. Vomcrslcy. .,. Floristics and cology
ol thc Mangrovc \cgctation ol Papua and Ncw Guinca.
cpartmcnt ol Forcsts, ivision ol 8otany, Lac, Papua
Ncw Guinca.
PohTaySoon and P.8.L. Srivastava. .. Crop composi
tion and dcnsity altcr thinnings and bclorc nal lclling
in thc Matang Mangrovc Rcscrvc, Pcrak, Malaysia. Pcr
tanika (.): .c.
Primavcra, J.H., R.8. Sadaba, M.J.H.L. Lcbata, and J.P.
Altamirano. acc. Handbook ol Mangrovcs in thc Phil
ippincsPanay. SAFC Aquaculturc cpartmcnt,
!loilo, Panay, Philippincs.
Pullc, A. .... Rhizophoraccac. Nova Guinca. Rcsultat
dc Lcxpcdition scicntiquc nccrlandaisc a la Nouvcllc
Guincc cn .c, ct .c. H.A. Lorcntz. , ac Partic, 8ota
niquc: 6,.
Putz, F.., and H.T. Chan. .6. Trcc growth, dynamics
and productivity in a maturc mangrovc lorcst in Malaysia.
Forcst cology and Managcmcnt .,: a..ac.
Rao, T.A., P.\. Surcsh, and A.N. Shcric. .6. Multiplc vi
viparity in a lcw taxa ol mangrovcs. Currcnt Scicncc :
aa6..
Rilcy, R.V., and C.P. Salgado Kcnt. .. Rilcy cncascd
mcthodology: principlcs and proccsscs ol mangrovc habi
tat crcation and rcstoration. Mangrovcs and Salt Marshcs
: ac,a..
Robcrtson, A.!., and .M. Alongi (cds.). .a. Tropi
cal Mangrovc cosystcms. Amcrican Gcophysical Union,
Vashington, .C.
Sacngcr, P. acca. Mangrovc cology, Silviculturc and
Conscrvation. Kluwcr Acadcmic Publishcrs, 8oston.
20 !ndoVcst Pacic Rhizophora spccics
Salvoza, F.M. .6. Rhizophora. Univ. Philip. Nat. Appl. Sc.
8ull. (): .,a,.
Smith, A.C. ... Family 6. Rhizophoracacac. Flora
\iticnsis Nova: A Ncw Flora ol Fiji, vol. a. Pacic Tropi
cal 8otanical Gardcn, Lwai, Kauai, Hawaii.
Sncdakcr, S.C., and M.S. 8rown. .a. Primary produc
tivity ol mangrovcs. !n: Mitsui, A. and C.C. 8lack (cds.).
CRC Handbook ol 8iosolar Rcsourccs. CRC Prcss !nc.,
8oca Raton, Florida.
Sousa, V.P., S.P. Quck, and 8.J. Mitchcll. acc. Rcgcncra
tion ol Rhizophora mangle in a Caribbcan mangrovc lor
cst: intcracting cccts ol canopy disturbancc and a stcm
boring bcctlc. ccologia .,: 6.
Stccnis, C.C.\. .. !ntroduction to Rhizophoraccac. Flora
Mal. (.): a.
Taylor, F.J. .6. Mangrovcs in lrcshwatcr. 8lumca .: a,.
a,a.
Taman, R.R. .. nc hundrcd Pacic island agrolorcstry
spccics. !n: Clarkc, V.C. and R.R. Taman. Agrolorcstry
in thc Pacic !slands: Systcms lor Sustainability. Unitcd
Nations Univcrsity Prcss, Tokyo. http://www.unu.cdu/
unuprcss/unupbooks/cac/cacp.htm~.
Tomlinson, P.8. .6. Tc 8otany ol Mangrovcs. Cam
bridgc Univcrsity Prcss, Cambridgc, UK.
Tomlinson, P.8., R.8. Primack, and J.S. 8unt. .,. Prc
liminary obscrvations on oral biology in mangrovc
Rhizophoraccac. 8iotropica ..(): a6a,,.
Tsuda, S., and M. Ajima. .. A prcliminary study ol rc
sprouting ability ol somc mangrovc spccics altcr cutting.
Tropics (): aa.aa.
Valsh, G.. .,,. xploitation ol mangal. !n: Chapman, \.J.
(cd.). cosystcms ol thc Vorld: Vct Coastal cosystcms.
lscvicr Scicncc Publishing Company, Amstcrdam.
Valtcrs, 8.8. acc. Pattcrns ol local wood usc and cutting
ol Philippinc mangrovc lorcsts. conomic 8otany (.):
66,6.
Vatson, J.G. .a. Mangrovc lorcsts ol thc Malay pcninsula.
Malayan Forcst Rccords 6. Frascr & Ncavc, Singaporc.
Vcinstock, J.A. .. Rhizophora mangrovc agrolorcstry.
conomic 8otany (a): a.ca..
VyattSmith, J. .6c. Ficld kcy to thc trccs ol mangrovc lor
cst in Malaya. Malay. For. a: .a6.a.
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 21
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org)
Rhizophora apiculata, R. mucronata, R. stylosa, R. annamalai, R. lamarckii
(IndoWest Pacic stilt mangroves)
Authors: Norman C. ukc, Marinc 8otany Group, Ccntrc lor Marinc Studics, Tc Univcrsity ol Quccnsland, St. Lucia QL c,a,
Australia, http://www.marinc.uq.cdu.au/marbot/indcx.htm~.
Acknowledgments: Tc author and publishcr thank Tom Colc, Farid ahdouhGucbas, alc vans, Aricl Lugo, and ianc Ragonc
lor thcir input. Photo contributions by 8cth 8rook and Craig lcvitch arc grcatly apprcciatcd. Tc author acknowlcdgcs a host ol
pcoplc who havc ovcr thc ycars supportcd his journcy into Rhizophora and who sharc thc passion lor this group ol plants. Collabora
tors and lricnds includc: Jim Allcn, Uta 8crgcr, John 8unt, Molly Crawlord, Farid ahdouhGucbas, tto alhaus, Kathy wcl,
Candy Fcllcr, XucJun Gc, Gina Holguin, L.P. Jayatissa, Kandasamy Kathircsan, Kcn Krauss, ugcnia Lo, Ull Mchlig, Jaimc Polania,
Jurgcnnc Primavcra, Utc Stcinickc, Mci Sun, and Nick Vilson.
Rccommcndcd citation: ukc, N.C. acc6. Rhizophora apiculata, R. mucronata, R. stylosa, R. annamalai, R. lamarckii (!ndoVcst
Pacic stilt mangrovcs), vcr. a... !n: lcvitch, C.R. (cd.). Spccics Prolcs lor Pacic !sland Agrolorcstry. Pcrmancnt Agriculturc
Rcsourccs (PAR), Hlualoa, Hawaii. http://www.traditionaltrcc.org~.
Sponsors: Publication was madc possiblc by gcncrous support ol thc Unitcd Statcs cpartmcnt ol Agriculturc Vcstcrn Rcgion Sus
tainablc Agriculturc Rcscarch and ducation (USAVSAR) Program, SPC/GTZ PacicGcrman Rcgional Forcstry Projcct,
USA Natural Rcsourccs Conscrvation Scrvicc (USA NRCS), Statc ol Hawaii cpartmcnt ol Land & Natural Rcsourccs ivi
sion ol Forcstry & Vildlilc, and thc USA Forcst Scrvicc Forcst Lands nhanccmcnt Program. Tis matcrial is bascd upon work
supportcd by thc Coopcrativc Statc Rcscarch, ducation, and xtcnsion Scrvicc, U.S. cpartmcnt ol Agriculturc, and Agricultural
xpcrimcnt Station, Utah Statc Univcrsity, undcr Coopcrativc Agrccmcnt acca,cc.c.a,.
Series editor: Craig R. lcvitch
Publisher: Pcrmancnt Agriculturc Rcsourccs (PAR), P 8ox a, Hlualoa, Hawaii 6,a, USA, Tcl: caa,, Fax: ca
.a, mail: parCagrolorcstry.nct, Vcb: http://www.agrolorcstry.nct~. Tis institution is an cqual opportunity providcr.
Reproduction: Copics ol this publication can bc downloadcd lrom http://www.traditionaltrcc.org~. Tis publication may bc rcpro
duccd lor noncommcrcial cducational purposcs only, with crcdit givcn to thc sourcc. acc6 Pcrmancnt Agriculturc Rcsourccs. All
rights rcscrvcd.