Bruguiera gymnorrhiza (large-leafed mangrove)

Rhizophoraceae (mangrove family)

denges (Palau); jon (Marshall Islands); large-leafed mangrove, oriental mangrove (English); orange mangrove
(northern Australia); ong (Chuuk); sohmw (Pohnpei); sraol (Kosrae); yangach (Yap Islands)
James A. Allen and Norman C. Duke
v
n
o
:
o
:

j
.

~
.

~
i
i
v
x
Forest of large-leafed mangrove.
Species Proles for Pacic Island Agroforestry
www.traditionaltree.org
April acc6
vcr. a..
IN BRIEF
Distribution Found in Microncsia, Samoa, and thc south
wcstcrn Pacic, widcly distributcd lrom thc castcrn coast
ol Alrica through Asia to subtropical Australia.
Size Rcachcs c m (.cc.. lt) in hcight, although
commonly much shortcr, around ,ac m (aa6a lt).
Habitat !ntcrtidal zoncs, ca m (c6.6 lt), or thc clcva
tion rangc bctwccn mcan sca lcvcl and highcst tidc, with
rainlall ol .cccccc mm (c. in).
Vegetation Common associatcs on Pacic islands includc
othcr mangrovc trcc spccics.
Soils Vidc rangc, trccs thrivc bcst in rivcr cstuarics.
Growth rate Lcss than . m/yr (. lt/yr) in hcight.
Main agroforestry uses Soil stabilization, coastal protcc
tion, wildlilc/marinc habitat.
Main products Timbcr, dyc, and traditional mcdicinc.
Yields No data availablc.
Intercropping Rccommcndcd lor planting togcthcr with
othcr mangrovc spccics.
Invasive potential Can bc invasivc, not rccommcndcd lor
planting outsidc ol its natural rangc.
2 Bruguiera gymnorrhiza (large-leafed mangrove)
INTRODUCTION
Largclcalcd mangrovc (Bruguiera gymnorrhiza) is onc ol
thc most important and widcsprcad mangrovc spccics in
thc Pacic. !t is lound in intcrtidal arcas ol thc tropical
Pacic rcgion lrom Southcast Asia to thc Ryukyu !slands
ol southcrn Japan into Microncsia and Polyncsia (Samoa),
and southward to subtropical Australia. Largclcalcd
mangrovc thrivcs undcr a broad rangc ol intcrtidal condi
tions, including salinity lcvcls lrom ncar lrcshwatcr to lull
strcngth scawatcr, and tolcratcs a rangc ol ooding and
othcr soil typcs. Typically, it is most common in thc middlc
and uppcr intcrtidal zoncs, rathcr than in thc lowcr intcr
tidal zonc or along thc scaward cdgc ol mangrovc stands.
Somc ol thc valuc ol this spccics is dicult to scparatc out
lrom thc largcr rolc playcd by mangrovc spccics as a wholc.
Mangrovcs in gcncral arc bclicvcd to play vitally important
rolcs in shorclinc protcction, cnhancing watcr quality in
ncarshorc cnvironmcnts (including ovcr coral rccls), pro
viding habitat lor juvcnilc sh and othcr rccl spccics, and
supporting marinc lood chains through outwclling ol car
bon. Although thc wood is thc main part ol thc trcc uscd
in most ol thc Pacic, numcrous uscs lor othcr parts ol thc
trcc including thc propagulcs, lcavcs, and bark havc bccn
rcportcd in thc rcgion.
DISTRIBUTION
Native range
Found in tropical intcrtidal arcas lrom thc castcrn coast
ol Alrica through Asia to thc Ryukyu !slands ol southcrn
Japan, into Microncsia and Polyncsia (Samoa), and south
ward to subtropical Australia (Quccnsland, Ncw South
Valcs and Vcstcrn Australia), it has thc largcst natural
longitudinal rangc ol all mangrovc spccics.
Vithin thc mangrovc ccosystcm, largclcalcd mangrovc is
most commonly lound in thc middlc and uppcr intcrtidal
zoncs, rathcr than in thc lowcr intcrtidal arcas or outcr sca
ward lringc.
Current distribution
Tcrc is littlc cvidcncc that largclcalcd mangrovc oc
curs widcly outsidc ol its natural rangc. !t is possiblc that
it occurs in thc Hawaiian !slands, but rcccntly thc spc
cics ol Bruguiera naturalizcd on ahu was lound to bc B.
sexangula, not B. gymnorrhiza as prcviously rcportcd (Al
lcn ct al. accc).
BOTANICAL DESCRIPTION
Preferred scientic name
Bruguiera gymnorrhiza (L.) Lamk.
Family
Rhizophoraccac (mangrovc lamily)
Non-preferred scientic names
Brugiera gymnorrhiza (L.) Lam., Tab. ncycl. a: .,, t. ,.
., (gymnorhiza). (common misspclling)
Rhizophora gymnorrhiza L., Sp. Pl.: . ., (gymnorhiza),
G. Forst., Fl. !ns. Austr.: , n. ac.. .,6.
Bruguiera rheedei 8l., num. Pl. Jav.: a. .a, (rheedii).
Brugiera rumphii 8l., Mus. 8ot. Lugd.8at. .: .. .c.
Common names
Pacic
denges (Palau)
jon (Marshall !slands)
largclcalcd mangrovc, oricntal mangrovc (nglish)
ong (Chuuk, FSM)
orangc mangrovc (northcrn Australia)
sohmw (Pohnpci, FSM)
sraol (Kosrac, FSM)
yangach (Yap, FSM)
Indian Ocean
malkadol, sirikanda (Sri Lanka: Sinhala)
thuddu ponna, uredi (Andhra Pradcsh, !ndia: Tclugu)
muia, mkoko wimbi (Kcnya, Tanzania, and Zanzibar, Mo
zambiquc: Kiswahili)
Size
!t is a mcdium to tall trcc that may rcach c m (.cc..
lt) in hcight, although it is commonly much shortcr. i
amctcrs arc commonly about . cm (6. in). warlcd
stands havc bccn rcportcd lor somc locations (c.g.,
Pohnpci).
Form
Largclcalcd mangrovc is normally a singlcstcmmcd trcc
with short buttrcsscs and charactcristic kncc roots, (hori
zontal roots that occasionally lorm abovcground loops,
prcsumably as an aid to gas cxchangc lor thc subsurlacc
portions ol thc roots). !t tcnds to bc ol shortcr staturc and
morc conical in shapc on thc scaward cdgc ol stands or in
arcas ol high salinity.
8ark is typically palc brown to grcy (darkcr whcn wct),
grcatcr than a cm (c. in) thick, hard, and rough. Popula
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 3
tions on Hainan !sland (China) havc distinctivc bark char
actcristics with vcrtical ssurcs plus largc aky lcnticcls a
cm (c. in) in diamctcr.
Flowering
Flowcrs arc pcrlcct. !norcsccncc has solitary owcr buds,
locatcd in lcal axils, usually nodding, positioncd at thc
rst (or rarcly sccond) nodc bclow thc apical shoot. Ma
turc owcr bud whcn closcd is .c. cm (..a.. in) long,
... cm (c.6.. in) widc. Calyx is typically rcddish to al
most scarlct, occasionally palc ycllow, whitc or grccn, char
actcristically smooth or with groovcs abovc lobc juncturcs,
rarcly ribbcd, with .a. lobcs, acutcly pointcd, narrow.
Stylc is palc grccn, lilorm, lobcs, about ac mm (c. in)
long and . mm (c.c in) widc. Pctals arc bilobcd (onc third
ol total lcngth), with hairy outcr margins, crcamywhitc
tcnd to orangcbrown on maturation, numbcr cquals thc
numbcr ol calyx lobcs, about . mm (c.6 in) long and mm
(c..6 in) widc. 8ristlc is bctwccn lobcs, solitary, straight, up
to mm long. 8ristlcs () arc on tips ol pctal lobcs, ol
tcn curly, mm (c..ac..6 in) long. Stamcns arc crcamy
whitc to goldcn brown at maturity, about .a mm (c. in)
long. Pcdunclc is smooth, slcndcr, curvcd, approximatcly
.c mm (c. in) long and a mm (c.cc..a in) widc. Tc
owcring pcriod is chicy April to August in thc southcrn
hcmisphcrc, and ctobcr to Fcbruary in northcrn hcmi
sphcrc.
Leaves
Lcavcs arc oppositc, simplc, clliptical, dark grccn,
and coriaccous (lcathcry), aggrcgatcd at thc tips
ol apical shoots in clustcrs ol about .a lcavcs.
Lcavcs arc aa cm (. in) long and cm
(a in) widc, with pctiolcs ol a cm (c...6
in). Lcal bladcs arc clliptic tcnding to oblong,
about . cm (6 in) long and 6 cm (a. in) widc,
acutc tip without mucro, pctiolc about cm (..6
in) long. Stipulcs (lcal shcaths) arc grccn or ycl
lowish, cm (..6.a in) long. Apical shoots
arc about 6 cm (a. in) long. Lcal color, sizc,
and shapc cnablc B. gymnorrhiza trccs to bc
distinguishcd lrom othcr Bruguiera spp. lrom
a distancc. !t dicrs lrom Rhizophora spp. in
thc lack ol mucronatc lcal tip, longitudinal lold
groovcs along bladc, and lack ol cork wart spots.
Lcal cmcrgcncc occurs mainly lrom Novcmbcr
to March in southcrn hcmisphcrc, May to Scp
tcmbcr in thc northcrn hcmisphcrc, markcd
dcclincs in canopy dcnsity may occur during
thcsc months il rainlall is low. Lcal lall occurs
chicy ovcr thc wct summcr pcriod lrom cto
bcr to March in thc southcrn hcmisphcrc, April
to Scptcmbcr in thc northcrn hcmisphcrc.
Fruit
Largclcalcd mangrovc is viviparous, mcaning
that thc spccics produccs sccds that gcrminatc
on thc parcnt plant. Tc dispcrsal unit, a vi
viparous sccdling, is callcd a hypocotyl. Tcrc
is no apparcnt lruit stagc. !nstcad, a hypocotyl
cmcrgcs singly lrom an attachcd maturc calyx.
Maturc hypocotyls with attachcd calyx bodics
arc locatcd at thc third to lth nodcs bclow thc
apical shoot. Tc hypocotyl is cylindrical, clon
gatc, stocky, dark grccn, coriaccous, with lon
Foliage with ower buds and propagules showing typical red calyx form.
Chuuk, Federated States of Micronesia. vno:o: x.c. uuxv
4 Bruguiera gymnorrhiza (large-leafed mangrove)
gitudinal ribbing giving an angular appcarancc, thc root
tip bluntly pointcd, maturc dimcnsions .a cm (6.c in)
long and about a cm (c. in) widc. xpcndcd calyccs oltcn
rcmain attachcd altcr maturc propagulcs lall lrom parcnt
trccs. Fruiting, whcn maturc hypocotyls lall, occurs Janu
ary to Fcbruary in thc southcrn hcmisphcrc, and July to
August in thc northcrn hcmisphcrc.
Seeds
Likc all Bruguiera spccics, largclcalcd mangrovc is vivipa
rous, mcaning that thc spccics produccs sccds hiddcn in
thc maturc calyx (postowcring) that gcrminatc on thc
parcnt plant. Just onc hypocotyl is usually produccd lrom
cach maturc calyx, although on rarc occasions twins may
bc obscrvcd.
Rooting habit
Maturc trccs havc distinctivc, sturdy, abovcground kncc
roots surrounding thc stcm basc that anchor only shallowly
in thc scdimcnts, to .a m (.6.6 lt) dcpth.
Similar species
Largclcalcd mangrovc is most likcly to bc conluscd with
othcr Bruguiera spccics/taxa whcrc distributions ovcrlap in
thc wcstcrn Pacic. !n particular, B. gymnorrhiza might bc
conluscd with B. sexangula, B. rhynchopetala, B. exaristata,
B. parviora, or B. cylindrica.
B. sexangula ltcn almost indistinguishablc lrom B.
gymnorrhiza. it has a solitary owcr, calyx orangcypink,
usually ribbcd rathcr than groovcd abovc lobc juncturcs,
likc B. gymnorrhiza. Propagulcs tcnd to bc much short
cr, 6..c cm (a.6. in). Populations in Hainan !sland
(China) havc distinctivc smoothcr bark with horizontal
ssurcs plus small lcnticcls about . cm (c. in) in diamctcr.
B. rhynchopetala B. gymnorrhiza is also rcportcd
to hybridizc with B. sexangula, giving progcny callcd
B. rhynchopetala. Tis hybrid taxon has intcrmcdiatc and
sharcd charactcrs ol B. sexangula and B. gymnorrhiza, with
a solitary owcr, calyx grccn with rcd blush, variably ribbcd
or groovcd abovc lobc juncturcs. Propagulcs arc indistin
guishablc lrom parcntal taxa. Populations in Hainan !sland
(China) havc distinctivc intcrmcdiatc bark with horizontal
and vcrtical ssurcs plus intcrmcdiatcsizcd lcnticcls .a
cm (c.c. in) in diamctcr.
B. exaristata Solitary owcrs also, but thcy arc smallcr,
grccn ncvcr rcddish, calyx ribbcd with .c sprcading lobcs
comparcd to .a. lor B. gymnorrhiza. Furthcrmorc, unlikc
othcr solitary owcr taxa, this spccics has no spinc bctwccn
pctal lobcs. Propagulc distinctly ribbcd, much shortcr.
B. parviora Multiplc owcrs arc in small groups, cach
having , short calyx lobcs cxtcnding down lrom thc nar
row, clongatc, ribbcd calyx. Propagulcs substantially thin
ncr, with clongatc spaghcttilikc appcarancc.
B. cylindrica Multiplc owcrs occur in groups ol two or
thrcc and havc a grccnishycllow, bulbous calyx with six
to ninc lobcs that lold back against thc uppcr calyx. Prop
agulc narrow, with a roundcd cylindrical appcarancc, no
groovcs or ribs.
Rhizophora species Rhizophora has lour calyx lobcs, most
havc cork wart spots on lcal undcrsurlaccs (notc: Bruguiera
spccics ncvcr havc cork wart spots), and most !ndoVcst
Pacic Rhizophora havc promincnt mucronatc tips at thcir
lcal apiccs (unlikc R. samoensis in thc southcrn Pacic is
lands, and thc introduccd R. mangle).
GENETICS
Variability of species
Somc variation in calyx color ol thc owcr occurs, with
somc trccs having owcrs dcscribcd as crimson rcd, or
angc, ycllowish rcd, crcamy whitc, or grccn. Colors may
also bc mixcd rcd and grccn, with thcir rcd sidcs oltcn up
Mature hypocotyl and calyx attached to branch. Daintree
River, NE Australia. vno:o: x.c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 5
pcrmost. Carc must bc takcn to dicrcntiatc bctwccn ca
lyx color and pctal color, as this may at timcs bc dcncd as
owcr color. Rcd coloration ol thc calyx is dcnitcly morc
prcvalcnt in sunacctcd locations, within singlc trccs and
among trccs.
Known varieties
!n N Australia, B. gymnorrhiza has albino lorms whcrc
somc hypocotyls lack grccn pigmcnt. Tcsc propagulcs
arc living, but thcy arc nonviablc, dying within a ycar ol
scttlcmcnt altcr dcplcting rcscrvcs, bccausc thcy cannot
photosynthcsizc. Vhcn prcscnt, thcy arc casily obscrvcd
hanging alongsidc normal grccn hypocotyls lrom othcr
wisc normal looking parcnt trccs.
ASSOCIATED PLANT SPECIES
Mangrovcs commonly occur in rclativcly distinct zoncs,
which arc controllcd by lactors such as salinity, tidal rc
gimc, soil typc, and sccd prcdators. Largclcalcd man
grovc is most commonly lound in an intcrmcdiatc loca
tion bctwccn thc scaward cdgc ol mangrovc stands (oltcn
dominatcd by Rhizophora spp. and Sonneratia alba) and thc
landward cdgc, which may bc dominatcd by a divcrsc mix
ol mangrovcs and mangrovc associatcs.
Associated species commonly found
Common associatcs on Pacic !slands includc othcr
mangrovc trcc spccics, cspccially Rhizophora apiculata, R.
samoensis, Sonneratia alba, Xylocarpus granatum, Heritiera
littoralis, Ceriops tagal, and Lumnitzera littorea. thcr ora
occasionally lound with largclcalcd mangrovc includc thc
mangrovc lcrn (Acrostichum aureum/speciosum), nypa palm
(Nypa fruticans), and a varicty ol vincs (c.g., Derris spp.)
and cpiphytcs (c.g., Hydnophytum spp.).
Left: Knee roots with rare instance of sprouting foliage stems. Gazi Bay, Kenya. Top right: Buttress roots and stem base, tree
growing on man-made coral rubble platform. Yap, Federated States of Micronesia. Bottom right: Mature hypocotyls compar-
ing a red albino form (above) and a normal dark-green form (below). Note also the color variation in calyces, red and red-
green, which is independent of propagule color. Moreton Bay, Queensland, Australia. vno:os: x.c. uuxv
6 Bruguiera gymnorrhiza (large-leafed mangrove)
ENVIRONMENTAL PREFERENCES
AND TOLERANCES
Climate
Largclcalcd mangrovc is a spccics ol tropical and subtrop
ical cnvironmcnts, which arc charactcrizcd by modcratcly
high and wcll distributcd rainlall.
Elevation range
ca m (c6.6 lt), which is thc clcvation rangc bctwccn
mcan sca lcvcl and thc highcst tidc.
Mean annual rainfall
.cccccc mm (c. in)
Rainfall pattern
Grows in climatcs with a unilorm rainlall pattcrn.
Dry season duration (consecutive months with <40
mm [1.6 in] rainfall)
No spccic inlormation is known lor largclcalcd man
grovc. Mangrovc plants in gcncral appcar to dcpcnd on
groundwatcr to sustain optimal growth, cspccially during
dricr months. Rcduccd growth in rcsponsc to an l Nino
cvcnt on Kosrac in .,/. was prcsumably duc to a dc
crcasc in lrcshwatcr input.
Mean annual temperature
accC (66F) (cstimatc)
Mean maximum temperature of hottest month
aC (c.ccF) (cstimatc)
Mean minimum temperature of coldest month
cC (a.F) (cstimatc)
Minimum temperature tolerated
C (aF) (cstimatc)
Soils
Trccs dcvclop grcatcst staturc and columnar growth lorm
in cstuarics ol largcr tropical rivcrs, charactcrizcd by nc
clay and black mud scdimcnts with rclativcly high lcvcls ol
organic carbon. Tcsc soils arc oltcn anacrobic, with high
conccntrations ol suldc. Howcvcr, thcsc trccs also occur
at timcs in sitcs with morc acrobic scdimcnts consisting
ol nc sands to coarsc stoncs and rocks. !n thcsc instanccs,
kncc roots arc lcss promincnt.
Soil texture
Grows in light, mcdium, and hcavy tcxturcd soils (sands,
sandy loams, loams, sandy clay loams, clay loams, sandy
clays, and clays).
Soil drainage
Grows in soils with lrcc and impcdcd drainagc as wcll as
watcrloggcd soils.
Soil acidity
Acidic to ncutral soils (pH .c,.).
Special soil tolerances
Grows in salinc soils. Tc optimal salinity rangc is rcportcd
to bc a6 ppt (parts pcr thousand), comparcd with ap
proximatcly 6 ppt lor lullstrcngth scawatcr.
Tolerances
Drought
Tc trcc can tolcratc drought pcriods rcasonably wcll, al
though growth ratc is rcduccd substantially.
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
dations bctwccn mcan sca lcvcl and thc highcst tidcs,
and thcy providc vital structurc as habitat and lood lor
similarly adaptcd rcsidcnt and transicnt launa. At low
tidc, mangrovc plants cxchangc gascs lrom cxposcd
roots using spccial lcnticcls, whilc ooding tidcs al
low uptakc ol nutricnts and sccd dispcrsal, oltcn using
cspccially buoyant propagulcs. Tc ccological limits
dcncd by thc diurnal tidal rangc cxplain thc sctting
and why just ,c spccics around thc world arc callcd
truc 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, abovc ground roots allowing vital gas cxchangc
in anacrobic scdimcnts, and physiological adaptations
lor cxcluding or cxpclling salt. Fcwcr than aa plant
lamilics havc dcvclopcd such attributcs, rcprcscnting
indcpcndcnt instanccs ol cocvolution ovcr millions
ol ycars to lorm todays mangrovc habitats.
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 7
Full sun
!t grows wcll in lull sun.
Shade
Largclcalcd mangrovc is onc ol thc most shadc tolcrant
mangrovc spccics. Sccdlings may pcrsist and grow undcr a
lull lorcst canopy (cvcn lcss than .c lull sunlight).
Fire
!t has no tolcrancc ol rc in closc proximity.
Frost
Tc trcc is almost ccrtainly not tolcrant ol sublrcczing
tcmpcraturcs lor morc than a lcw hours.
Waterlogging
!t can tolcratc daily ooding up to dcpths ol at lcast .. m
( lt) and cvcn pcrmancntly saturatcd soils.
Salt spray
Tc trcc is highly tolcrant ol salt spray.
Wind
!t is not typically lound on thc cxtrcmc scaward (and most
windswcpt) cdgcs ol mangrovc lorcsts. Planting in highly
windpronc locations is not rccommcndcd.
Abilities
Regenerate rapidly
Largclcalcd mangrovc rcgcncratcs ccctivcly by natural
dispcrsal and sccdling cstablishmcnt, but thcn tcnds to
grow slowly, dcpcnding on lrcshwatcr inputs, muddy sub
stratc, and nutricnt availability.
Self-prune
Scllpruncs wcll in dcnsc stands but maintains lowcr
branchcs in morc opcngrown locations.
Coppice
ocs not coppicc wcll but docs rcspond much bcttcr to
cutting and lcal rcmoval than Rhizophora spccics, with
sprouting along largcr stcms bclow lcaly roscttcs.
Pollard
No rcports wcrc lound ol pollarding practiccs.
GROWTH AND DEVELOPMENT
Growth ratcs vary with agc. Growth in hcight is rapid
shortly altcr cstablishmcnt whilc lood rcscrvcs arc takcn
up lrom thc hypocotyl. Tc ratc thcn appcars to slow in a
sigmoid dcclinc until thc trcc attains thc maximum hcight
obtainablc on a particular sitcthc sitc maximal canopy
hcight (ukc acc.). Altcr this, trccs tcnd to sprcad and
broadcn in canopy diamctcr rathcr than gct tallcr.
Growth rate
Largclcalcd mangrovc gcncrally grows lcss than . m/yr
(. lt/yr) in hcight. Propagulcs plantcd in !ndia avcragcd
only ... m (. lt) tall altcr vc growing scasons. Naturally
rcgcncratcd saplings growing in small gaps on Kosrac havc
avcragcd about .c cm/yr ( in/yr) ovcr a pcriod ol 6 ycars.
Trccs in somc plantations on thc Andamans havc rcport
cdly rcachcd a hcight ol .a m (c lt) and a circumlcr
cncc ol ac cm (.c.a in) in . ycars. Mcan annual diam
ctcr incrcmcnts ol c..,c.a. cm (c.c,c.c in) and c.c cm
(c..a in) havc bccn rcportcd lor Malaysia and 8angladcsh,
rcspcctivcly. !n Microncsia, largclcalcd mangrovc grow
ing on Pohnpci had diamctcr growth ratcs ol c.a6 cm/yr
(c.. in/yr), and on Kosrac, c. cm/yr (c.., in/yr) (Colc ct
al. .).
A .ccycar rotation was rccommcndcd lor thc Andaman
!slands, at which timc thc cxploitablc girth should bc
around ,c cm (a in).
Flowering and fruiting
Flowcring and lruiting occur continuously throughout
thc ycar, although distinctly scasonal pcaks ol hypocotyl
production wcrc rcportcd lor JulyAugust in northcrn
parts ol thc rangc and JanuaryFcbruary in southcrn parts.
Trccs havc notablc and long pcriods ol rcproductivc dc
vclopmcnt, taking .a ycars lrom rst cmcrgcncc ol owcr
buds until maturation and drop ol maturc hypocotyls.
Yields
No data availablc.
PROPAGATION
Largclcalcd mangrovc is rcadily propagatcd by dircct
planting ol propagulcs (viviparous sccdlings). Although
natural rcgcncration is gcncrally rclicd upon around thc
Pacic rcgion, this spccics is rclativcly casy to propagatc
articially.
Propagation is simplc and rclics on thc spccial lcaturc ol
this gcnus: viviparous propagulcs. As such, planting simply
cntails gcntly pushing thc distal cnd ol thc .c. cm (6
in) long hypocotyl onc third ol its lcngth into thc scdimcnt,
spaccd at about . m (. lt) intcrvals. No holcs nccd to bc
dug, no nurscry prcparation, no stakcs, and gcncrally low
8 Bruguiera gymnorrhiza (large-leafed mangrove)
maintcnancc is rcquircd lor growing sccdlings in shcltcrcd
arcas.
Propagation by viviparous seedling
Propagule collection
Propagulcs arc availablc throughout much ol thc ycar, with
pcaks in JulyAugust common in northcrn parts ol thc
rangc and JanuaryFcbruary in thc southcrn parts. Ma
turc propagulcs may bc collcctcd altcr thcy havc lallcn
or dircctly o thc trccs. nly hcalthy looking propagulcs
should bc sclcctcd. Propagulcs that arc shrunkcn or dcs
iccatcd 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 arc
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
No proccssing ol maturc propagulcs is rcquircd. Tcy can
bc sown in nurscry bcds or plantcd in thc cld in thc lorm
in which thcy wcrc collcctcd.
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
burlap bags and storing thcm out ol dircct sunlight. !t is
likcly that thcy can bc kcpt cvcn longcr, but storagc bcyond
a wccks is not rccommcndcd and longtcrm storagc is not
lcasiblc.
Propagule pretreatment
No prcplanting trcatmcnts arc ncccssary.
Growing area
Largclcalcd mangrovc can bc grown in lull sunlight or
undcr a widc rangc ol shadc. !n Australia, usc ol c shadc
lor Bruguiera spccics is rccommcndcd.
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
cc mix ol sand and pcat has bccn rccommcndcd lor
Australian mangrovc nurscrics. 8rackish watcr is rccom
mcndcd, but sccdlings havc also bccn succcsslully propa
gatcd using lrcsh watcr.
Time to outplanting
Sccdlings arc rcady lor outplanting at thc sixlcal (thrcc
Planting mangroves on a storm-damaged site. vno:o: x. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 9
nodc) stagc il grown in nurscry conditions. ldcr sccd
lings up to c. m (ac in) tall havc also bccn succcsslully
transplantcd. Planting may bc undcrtakcn at any timc dur
ing thc ycar. ircct propagulc planting, will, howcvcr bc
rcstrictcd by thcir scasonal availability lrom parcnt trccs,
as thcy do not kccp lor cxtcndcd pcriods unlcss plantcd
out in pots.
Guidelines for outplanting
Propagulcs or nurscrygrown sccdlings should havc cxccl
lcnt survival on sitcs that havc bccn propcrly sclcctcd and,
il appropriatc, protcctcd lrom disturbancc. Survival ol c
or grcatcr is not unrcasonablc in such sitcs. n thc othcr
hand, survival may bc zcro on sitcs cxposcd to cxccssivc
wavc action, inappropriatc hydrologic or salinity rcgimcs,
or (rarcly) disturbancc by grazing animals (such as camcls,
goats, cattlc, and horscs).
DISADVANTAGES
!n gcncral, largclcalcd mangrovc poscs lcw signicant
disadvantagcs whcn plantcd in its nativc rangc. !t is not
cspccially susccptiblc to pcsts or pathogcns and has not
bccn rcportcd to host major pcsts or pathogcns ol impor
tant crop spccics.
Potential for invasiveness
Although thc invasivcncss ol largclcalcd mangrovc has
not bccn dcmonstratcd, it is likcly to bc casily opportunis
tic duc to its shadc tolcrancc and its rclativcly widc rangc
ol tolcrancc lor salinity and soil conditions. thcr man
grovc spccics introduccd into Hawaii havc clcarly dcmon
stratcd thc gcncral potcntial lor invasivcncss ol mangrovcs
in arcas whcrc thcy arc not nativc and yct suitablc growing
conditions cxist.
Pests/pathogens
Susccptibility to pcsts and pathogcns is bclicvcd to bc low,
with thc cxccption ol grapsid crabs and wccvils, which lrc
qucntly damagc propagulcs and/or lcavcs.
Host to crop pests/pathogens
No rcports wcrc lound ol largclcalcd mangrovc scrving as
thc host lor a known crop pcst or pathogcn.
AGROFORESTRY/ENVIRONMENTAL
PRACTICES
Largclcalcd mangrovc is probably ol grcatcst valuc in situ,
as this spccics (and mangrovcs as a wholc) is bclicvcd to
play a vital rolc in supporting marinc lood chains, protcct
ing coastal arcas, and improving watcr quality.
Mulch/organic matter
Mangrovcs in gcncral havc rclativcly high productivity, in
cluding lcal and root production. Mangrovcs tcnd to crcatc
highly organic soil cnvironmcnts and also cxport organic
mattcr to ncarby marinc cnvironmcnts. Largclcalcd man
grovc loliagc may bc rclativcly poor in quality comparcd to
othcr mangrovcs such as Sonneratia alba, but it still is an
ccctivc produccr and cxportcr ol organic mattcr. Mulch
in mangrovc lorcsts is hiddcn lrom vicw most ol thc timc.
!l it wcrc not lor thc small mangrovc crabs, lallcn lcavcs
would bc washcd away with cach tidc. Tc crabs activcly
takc lcavcs bclow ground, whcrc thcy bccomc undcrground
mulch that can thcn bccomc availablc to trccs in thc lorcst
again.
Soil stabilization
Mangrovcs in gcncral arc ccctivc in stabilizing soils in
intcrtidal arcas. Tcy may rapidly colonizc ncwly lormcd
mudats and othcr arcas ol unstablc soils within thc in
tcrtidal zonc. Although bclicvcd to bc usclul primarily lor
gas cxchangc, thc kncc roots ol largclcalcd mangrovc may
also hclp trap scdimcnts.
Windbreaks
Largclcalcd mangrovc has bccn plantcd in thc Philip
pincs to protcct coastal homcs lrom wind and wavc action.
Mangrovcs oltcn occur adjaccnt to agricultural arcas and
may scrvc to protcct crops lrom sca spray or storms.
Woodlot
Mangrovcs adjaccnt to homcs throughout thc Pacic lrc
qucntly scrvc as inlormal woodlots, particularly on islands
with clcar tcnurc systcms that includc mangrovc arcas,
such as on Yap, in thc Fcdcratcd Statcs ol Microncsia.
Native animal/bird food
Largclcalcd mangrovc is a largcly unknown sourcc ol na
tivc animal loods. Howcvcr, scvcral obscrvations dcmon
stratc that thc divcrsity and quantity ol thc sourcc is likcly
to bc cxtrcmcly important in mangrovc ccosystcms. Nu
mcrous insccts, crabs, and mollusks grazc on grccn lcavcs
in thc lorcst canopy. Scsarmid crabs consumc an unknown
quantity ol lallcn lcavcs and propagulcs. rganic mattcr
proccsscd by thcsc hcrbivorcs is bclicvcd to broadly sup
port aquatic lood chains in coastal rcgions. 8irds notably
lccd and dcpcnd on oral ncctarics ol Bruguiera spccics.
Tc rclationship with birds is so wcll dcvclopcd that this
plantanimal rclationship has cocvolvcd to crcatc an un
usual cxplosivc pollcnrclcasc mcchanism in Bruguiera
10 Bruguiera gymnorrhiza (large-leafed mangrove)
plants to dispcrsc pollcn to ncighboring trccs using bird
visitors. Fcw mammals appcar to usc largclcalcd man
grovc as a major lood sourcc.
Wildlife habitat
!n addition to aquatic marinc organisms (scc Fish/marinc
lood chain), mangrovcs scrvc as habitat lor a widc rangc ol
tcrrcstrial and arborcal wildlilc. Bruguiera lorcsts providc
shcltcr and lood lor a numbcr ol associatcd launa, includ
ing birds, small mammals, crustaccans, shcllsh, and sh
lilc. Fruit bats may roost in mangrovcs and arc known to
pollinatc somc spccics (although not largclcalcd man
grovc). !n somc cascs, rarc or cndangcrcd spccics havc bccn
documcntcd to usc largclcalcd mangrovc. An cxamplc is
thc nightingalc rccdwarblcr (Acrocephalus luscinia), a U.S.
lcdcrally listcd cndangcrcd spccics, which has bccn lound
to ncst in largclcalcd mangrovc trccs on Saipan.
Bee forage
Bruguiera spccics havc limitcd ncctar (largcly consumcd by
birds), but thcy do havc copious pollcn that may bc gath
crcd by bccs.
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 mangrovcs during part ol thcir lilc
cyclcs, as do spccics ol shrimp and crab. Spccics such as
thc mangrovc crab (Scylla serrata) arc common in man
grovcs with a high componcnt ol largclcalcd mangrovc,
and arc important sourccs ol lood and incomc on many
islands in thc rcgion. Populations ol somc crab spccics may
cxcccd .c/m
a
(./lt
a
) in parts ol thc Pacic. Scncsccnt lcavcs
having lallcn lrom Bruguiera trccs arc takcn by grapsid
(small mangrovc) crabs into thcir burrows. 8uricd lcavcs
dccomposc and contributc to nutricnt rccycling in man
grovc lorcsts. Nutricnts also lccd dircctly and indircctly to
associatcd cstuarinc and marinc lood chains. Mangrovc
arcas with largclcalcd mangrovc may also scrvc as habitat
lor morc charismatic marinc mcgalauna, such as thc salt
watcr crocodilc.
Coastal protection
Mangrovcs in gcncral play an important rolc in protcction
ol coastlincs, shponds, and othcr coastal inlrastructurc.
Mangrovcs arc plantcd cxplicitly lor coastal or shpond
protcction in somc arcas (c.g., parts ol thc Philippincs) and
thcrc arc laws in many locations aimcd at protccting man
grovcs in largc part bccausc ol this important lunction.
Ornamental
Largclcalcd mangrovc is not widcly uscd as an ornamcn
tal, although its dark grccn lcavcs, gcncrally attractivc lorm,
and rcddishscarlct owcrs do makc it an attractivc trcc.
Packagcd plants (sprouting hypocotyls) arc sold in thc
kinawan !slands ( Japan) as an ornamcntal plant at tour
ist shops.
USES AND PRODUCTS
!n tcrms ol dircct usc by pcoplc, thc most widcsprcad usc ol
largclcalcd mangrovc is lor wood, which is uscd lor pur
poscs ranging lrom rcwood to construction ol homcs and
canoc parts. thcr uscs ol thc trcc includc lood (mainly
lrom thc propagulcs), dycs, and mcdicincs.
Staple food
Lcavcs and pcclcd propagulcs havc rcportcdly bccn soakcd,
boilcd, and catcn as a staplc in Papua Ncw Guinca, al
though most likcly only in timcs ol scvcrc shortagc ol
othcr loods. Although largclcalcd mangrovc is not widcly
uscd lor lood, rcports ol thc usc ol propagulcs as a lood
sourcc also cxist lor !ndia, 8angladcsh, and othcr parts ol
Southcast Asia.
Fruit
Tc lruit (propagulc) is rcportcdly catcn cookcd, altcr
scraping or grating, washing, and drying (to rcmovc tan
nins) and somctimcs mixcd with coconut in Mclancsia and
Nauru, lruit is sold as a vcgctablc at thc Honiara Markct
|Solomon !slands|. (Clarkc and Taman .)
Medicinal
Tc bark is uscd as an abortilacicnt and lor trcating burns
in thc Solomon !slands Tc bark is rcportcdly uscd lor di
arrhca and lcvcr in !ndoncsia. Tc astringcnt (and mildly
toxic) bark has also bccn uscd to trcat malaria in Cambo
dia. Tc lruits havc antiviral propcrtics, and bark cxtracts
ol thc closcly rclatcd B. sexangula arc rcportcdly activc
against at lcast two typcs ol canccrous tumors (Sarcoma
.c and Lcwis lung carcinoma).
Timber
Tc wood is widcly uscd lor structural componcnts (c.g.,
polcs, bcams, and raltcrs) ol traditional homcs and othcr
structurcs. !t is also uscd lor othcr purposcs, ranging lrom
traditional uscs such as shing stakcs, spcars, and copra
huskcrs to usc as a sourcc ol chips lor pulp production. Tc
spccics has also bccn uscd lor transmission and tclcphonc
polcs in somc rcgions (c.g., thc Andaman !slands), and is
likcly durablc in dircct contact with thc ground (i.c., rot
rcsistant).
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 11
Fuelwood
Tc wood has a high caloric valuc and is uscd lor luclwood
on somc Pacic !slands (c.g., Kosrac). !t is also madc into
charcoal in countrics such as Malaysia and !ndoncsia.
Craft wood/tools
Somctimcs uscd lor tool handlcs or digging sticks.
Canoe/boat/raft making
Tc wood has bccn uscd to makc canoc parts. !n thc
Marshall !slands, it has bccn uscd lor kcclpicccs (maal),
outriggcr (kic), and outriggcr struts (kcincon crrc). Ca
noc paddlcs arc also madc lrom largclcalcd mangrovc on
somc islands.
Body ornamentation/garlands
Flowcrs ol thc closcly rclatcd B. sexangula arc uscd in
Hawaii to producc a typc ol lci callcd thc kukuna o ka la
(rays ol thc sun) (Allcn .).
Tannin/dye
Uscd to producc dycs ranging lrom rcdbrown to black
(thc lattcr with rcpcatcd dying). Tc bark has a high tannin
contcnt, but tcnds to color lcathcr cxccssivcly unlcss thc
bark is collcctcd at thc cnd ol cach growing scason.
Cosmetic/soap/perfume
Tc kncc roots (or brcathing roots) rcportcdly havc bccn
uscd lor making pcrlumcs.
COMMERCIAL CULTIVATION
Although not as dcsirablc as Rhizophora spccics, largc
lcalcd mangrovc is lrcqucntly sold by commcrcial rcwood
supplicrs on somc islands (c.g., Kosrac, Chuuk in FSM).
Bruguiera timbcr is harvcstcd commcrcially lor charcoal
production in S Asia, although Rhizophora is prclcrrcd.
Tc caloric valuc ol thc timbcr is cnhanccd signicantly
by convcrting it to charcoal. Mangrovc lorcsts arc uscd lor
wood chips in Malaysia, Ncw Guinca, and thc Solomons
using unsustainablc harvcsting ol nativc lorcsts. Largc
lcalcd mangrovc plantations havc bccn cstablishcd on Fiji
lor land rcclamation and timbcr.
Spacing
Mangrovc plantations in gcncral arc typically plantcd at
spacings ol about ..c to .. m (. lt). Spacings widcr
than about a. m ( lt) tcnd to rcsult in a high proportion
ol multiplc stcmmcd and/or shortcr trccs. Vidcr spacings
may bc dcsircd lor coastal protcction projccts but not lor
timbcr production.
!n thc abscncc ol signicant natural mortality, timbcr plan
tations should bc thinncd to dcnsitics ol a. to . m (...
lt) as thc stand dcvclops and bccomcs crowdcd.
Management objectives
!n arcas whcrc thc mangrovc lcrn (Acrostichum aureum) is
common, it may nccd to bc controllcd to promotc carly
growth ol largclcalcd mangrovc, although thc high shadc
tolcrancc ol this spccics may makc this lcss important than
lor othcr mangrovc trcc spccics.
Somc publishcd guidclincs lor mangrovc silviculturc cxist
and arc rclcrcnccd in this prolc, but guidclincs on thin
ning, lcrtilizing, ctc., that arc spccic to largclcalcd man
grovc arc gcncrally unavailablc.
Growing in polycultures
Largclcalcd mangrovc naturally occurs in mixcdspccics
stands, and cach spccics has its own ccological and cco
nomic valucs. Mixcdspccics plantings arc thcrclorc rcc
THE LEGEND OF
THE ORIGIN OF POHNPEI
Long ago, thcrc livcd a man namcd Sapkini on thc
island ol ir. Undcr his guidancc, a grcat canoc was
built to sail into thc unknown in scarch ol a ncw land.
Sapkini bclicvcd this ncw land cxistcd whcrc thc sky
touchcd thc sca. Sapkini wiscly sclcctcd his crcw, cach
ol whom was cndowcd with thc skills or magical pow
cr nccdcd lor sailing or working thc land. Altcr many
days at sca thcy mct an octopus namcd Lidakika, who
dircctcd thcm to a distant shallow rccl on which a
small coral hcad was cxposcd. Tcy namcd it Ticrcn
sapw (bit ol land), bccausc it was no largcr than thc
distancc bctwccn thc canoc and its outriggcr. From
this bit ol coral, thc island ol Pohnpci was built.
8y magic, thcy callcd stoncs and bouldcrs to Ticrcn
sapw lrom othcr placcs, but thc carth and soil was
continuously washcd away by thc wavcs ol thc sca.
Tc pcoplc callcd on Katcngcnior (stabilizcr ol thc
shorc) lor hclp. Katcngcnior surroundcd thc land and
crcatcd a barricr rccl lor protcction. Still thc land was
not stablc, so thcy callcd on Katcnanik, who causcd
thc mangrovc trccs to takc root to hold thc shorc in
placc. Altcr thc shorc was sccurc, thc pcoplc dug carth
and pilcd it into a platlorm, which thcy namcd pci
(stonc altar). Pohnpci was built on top ol (pohn) thc
altar, thus thc island was namcd Pohnpci, or upon a
stonc altar.
Sourcc: Lambcth accc
12 Bruguiera gymnorrhiza (large-leafed mangrove)
ommcndcd, including R. apiculata and othcr Rhizophora
spccics.
Largclcalcd mangrovc may bc slowcr growing than othcr
mangrovc spccics but is ablc to pcrsist and may cvcn cvcn
tually dominatc in mixcdspccics plantations.
Estimated yields
No data availablc.
Markets
Markcts on most Pacic !slands arc local in naturc, with
littlc in thc way ol largclcalcd mangrovc products (othcr
than indircct products likc mangrovc crabs and lruit bats)
bcing cxportcd lrom onc island to anothcr. !n Southcast
Asia, largc quantitics ol largclcalcd mangrovc wood chips
and charcoal may bc movcd grcatcr distanccs and in grcatcr
volumcs than wood products on smallcr islands.
PUBLIC ASSISTANCE AND
AGROFORESTRY EXTENSION
USA Forcst Scrvicc
!nstitutc ol Pacic !slands Forcstry
... Punchbowl Strcct, Room a
Honolulu, H! 6., USA
Attn: Tropical Forcstcd Vctlands Tcam
Marinc 8otany Group
Ccntrc lor Marinc Studics
Tc Univcrsity ol Quccnsland
St. Lucia QL c,a, Australia
Vcb: http://www.marinc.uq.cdu.au/marbot/indcx.htm
xtcnsion occs lor agrolorcstry and lorcstry in thc Pa
cic: http://www.traditionaltrcc.org/cxtcnsion.html
INTERNET
For a nicc sct ol FAQs about mangrovcs (lrom thc Phil
Mature forest showing stems and typical above ground roots with Erick Waguk (Kosraen State Forester) pictured for scale.
Kosrae, Federated States of Micronesia. vno:o: x.c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 13
ippincs: http://www.pcmsca.org/youngaccnviron/yc.c./
mangrovc_indcx.htm
For a bricl summary ol B. gymnorrhiza charactcristics pro
duccd by thc Purduc Univcrsity Ccntcr lor Ncw Crops &
Plant Products: http://ncwcrop.hort.purduc.cdu/ncwcrop/
dukc_cncrgy/8ruguicra_gymnorrhiza.html
For a usclul publication callcd nc Hundrcd Pacic !s
land Agrolorcstry Spccics: http://www.unu.cdu/unuprcss/
unupbooks/cac/cacp.htm
BIBLIOGRAPHY
( indicates recommended reading)
Allcn, J.A. .. Mangrovcs as alicn spccics: thc casc ol Ha
waii. Global cology and 8iogcography Lcttcrs ,: 6.,..
Allcn, J.A. acca. Rhizophora mangle L. !n: \ozzo, J.A. (cd.).
Tropical Trcc Sccd Manual. Agriculturc Handbook ,a..
USA Forcst Scrvicc, Vashington, C.
Allcn, J.A., Krauss, K.V., ukc, N.C., Hcrbst, .R., 8jork
man, and . Shih. accc. Bruguiera spccics in Hawaii: sys
tcmatic considcrations and ccological implications. Pacic
Scicncc (): ..
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. Australian !nstitutc ol Marinc
Scicncc, A!MS Rcport =a, Townsvillc, Australia.
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~.
Clarkc, V.C., and R.R. Taman. .. Agrolorcstry in thc
Pacic !slands: Systcms lor Sustainability. Unitcd Nations
Univcrsity Prcss, Tokyo.
Colc, T.G., K.C. wcl, and N.N. cvoc. .. Structurc ol
mangrovc trccs and lorcst in Microncsia. Forcst cology
and Managcmcnt ..,: .c.
ahdouhGucbas, F., L.P. Jayatissa, . i Nitto, J.. 8osirc,
. Lo Sccn, and N. Kocdam, acc. How ccctivc wcrc
mangrovcs as a dclcncc against thc rcccnt tsunami: Cur
rcnt 8iology .(.a): ,.
c Laccrda, L.. acca. Mangrovc cosystcms: Function
and Managcmcnt. Springcr\crlag, 8crlin, Gcrmany.
ing Hou. .. Rhizophoraccac. pp. a. !n: van Stcc
nis, C.G.G.J. (cd.). Flora Malcsiana, \ol. . Noordho
Kol N.\., jakarta, !ndoncsia.
ing, H. .. Bruguiera. Flora Malcsiana \ol. Scr. ! ():
,6.
ukc, N.C. .a. Mangrovc oristics and biogcography.
Tropical Mangrovc cosystcms. !n: Robcrtson, A.!. and
.M. Alongi. Amcrican Gcophysical Union, Vashington,
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. acc.. Gap crcation and rcgcncrativc proccsscs
driving divcrsity and structurc ol mangrovc ccosystcms.
Vctlands cology and Managcmcnt : a,a6.
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., 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., M.C. 8all, and J.C. llison. .. Factors in
ucncing biodivcrsity and distributional gradicnts in
mangrovcs. Global cology and 8iogcography Lcttcrs ,:
a,,.
urant, C.C.L. ... Tc growth ol mangrovc spccics in
Malaya. Malayan Forcstcr .c: ..
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. \ol. .. Pacic Forum, Pacic Scicncc As
sociation, astVcst Ccntcr, Honolulu.
llison, J.C. .. Prcscnt status ol Pacic !sland man
grovcs. pp. .. !n: ldrcdgc, L.G., J.. Maragos, and
P.L. Holthus (cds.). Marinc/Coastal 8iodivcrsity in thc
Tropical !sland Pacic Rcgion: Population, cvclopmcnt
and Conscrvation Prioritics. \olumc !!. Pacic Scicncc
Association/ast Vcst Ccntcr Honolulu.
Gc, X.J., and N.C. ukc. acc. A mangrovc hybrid, Bru-
guiera rhynchopetala (Rhizophoraccac) lrom China and
N Australia. Australian Journal ol 8otany (in prcss).
Hogarth, P.J. .. Tc 8iology ol Mangrovcs. xlord
Univcrsity Prcss, Ncw York.
Ko, V.C. .,. Taxa nova Rhizophoraccarum. Acta Phyto
taxon Sin .6: .c..c.
Krauss, K.V., and J.A. Allcn. acc. Factors inucncing thc
rcgcncration ol thc mangrovc Bruguiera gymnorrhiza (L.)
Lamk. on a tropical Pacic island. Forcst cology and
Managcmcnt .,6: 6c.
Lambcth. L. accc. An Asscssmcnt ol thc Rolc ol Vomcn
in Fishcrics in Pohnpci, Fcdcratcd Statcs ol Microncsia.
Sccrctariat ol thc Pacic Community, Noumca, Ncw
14 Bruguiera gymnorrhiza (large-leafed mangrove)
Calcdonia. http://www.spc.int/coastsh/Scctions/Com
munity/cnglish/publications/pohnpci.pdl~
Macnac, V. .6. A gcncral account ol thc ora and launa
ol mangrovc swamps in thc !ndoPacic rcgion. Advanccs
in Marinc 8iology 6: ,a,c.
Marco, H.F. .. Systcmatic anatomy ol thc woods ol thc
Rhizophoraccac. Tropical Voods : .ac.
McCuskcr, A. .. Rhizophoraccac. aa: ..c. !n: A.S.
Gcorgc (cd.). Flora ol Australia. 8urcau ol Flora and
Fauna, Australian Govcrnmcnt, Canbcrra.
Robcrtson, A.!., and .M. Alongi (cds.). .a. Tropical
Mangrovc cosystcms. Amcrican Gcophysical Union,
Vashington, C.
Sacngcr, P. acca. Mangrovc cology, Silviculturc and
Conscrvation. Kluwcr Acadcmic Publishcrs, 8oston.
Tomlinson, P.8. .6. Tc 8otany ol Mangrovcs. Cam
bridgc Univcrsity Prcss, Cambridgc, UK.
Valsh, G.. .,,. xploitation ol mangal. pp. ,a6a. !n:
Chapman, \.J. (cd.). cosystcms ol thc Vorld: Vct
Coastal cosystcms. lscvicr Scicncc Publishing Com
pany, Amstcrdam.
Vatson, J.G. .a. Mangrovc lorcsts ol thc Malay pcninsula.
Malayan Forcst Rccords 6. Frascr & Ncavc, Singaporc.
Vilkinson, K.M., and C.R. lcvitch. acc. Propagation pro
tocol lor production ol containcr Bruguiera gymnorrhiza
(L.) Savigny plants. !n: Nativc Plant Nctwork. Univcrsity
ol !daho, Collcgc ol Natural Rcsourccs, Forcst Rcscarch
Nurscry, Moscow, !daho. http://www.nativcplantnct
work.org~.
VyattSmith, J. .. Tc Malayan spccics ol Bruguiera. Ma
lay Forcstry .6: .6.6..
VyattSmith, J. .6c. Ficld kcy to thc trccs ol mangrovc lor
cst in Malaya. Malay Forcstcr a: .a6.a.
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 15
Traditional Tree InitiativeSpecies Proles for Pacic Island Agroforestry (www.traditionaltree.org)
Bruguiera gymnorrhiza (large-leafed mangrove)
Authors: Jamcs A. Allcn
.
and Norman C. ukc
a
.. Paul Smiths Collcgc, P 8ox a6, Paul Smiths, Ncw York .a,c, USA. Vcb: http://www.paulsmiths.cdu
a. 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 authors and publishcr thank Tom Colc, Farid ahdouhGucbas, Judith rcxlcr, alc vans, and Kathy
wcl lor thcir input.
Recommended citation: Allcn, J.A., and N.C. ukc. acc6. Bruguiera gymnorrhiza (largclcalcd mangrovc), 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: parGagrolorcstry.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.