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

harrisonii
(AtlanticEast Pacic red mangrove)
Rhizophoraceae (mangrove family)
American mangrove (English, Australia); red mangrove (USA); mangle rojo (Central and Latin America, Pacic and
Caribbean coasts); tiri wai (Fiji); togo (Samoa)
Norman C. Duke and James A. Allen
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AtlanticEast Pacic red mangrove Rhizophora mangle growing
along an estuarine shoreline near Braganza, Amazonian Brazil.
Species Proles for Pacic Island Agroforestry
www.traditionaltree.org
April acc6
vcr. a..
IN BRIEF
Distribution Nativc to Amcrican wcst and
cast coasts and Alrican wcst coast. nc spccics,
Rhizophora mangle, was introduccd to thc ccn
tral Pacic, including Hawaii and thc Socicty
!slands. Closcly allicd with !ndoVcst Pacic
stilt mangrovcs whosc rangcs naturally ovcrlap
AP mangrovcs only in thc southcrn Pacic.
Size Can rcach cc m (.cc.6c lt) in hcight,
although commonly attains 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 Commonly associatcd with othcr
mangrovc spccics.
Soils Adaptcd to a vcry widc rangc ol soils but
thrivcs bcst in nc mud scdimcnts ol down
strcam rivcr cstuarics.
Growth rate Grows lcss than . m/yr (. lt/yr)
in hcight.
Main agroforestry uses Soil stabilization,
coastal protcction, wildlilc/marinc habitat.
Main products Timbcr, luclwood, charcoal,
dycs, and traditional mcdicinc.
Yields Timbcr volumc was cstimatcd at .cc
.c m

/ha/yr (.cca.cc lt

/ac/yr).
Intercropping Rccommcndcd lor planting to
gcthcr with othcr mangrovc spccics.
Invasive potential Tcsc plants arc rcady col
onizcrs ol ncw mud banks, making thcm op
portunistically invasivc with a high potcntial to
invadc alicn cnvironmcnts, gcncrally not rcc
ommcndcd lor planting outsidc ol thcir natural
rangc.
2 Atlanticast Pacic Rhizophora spccics
INTRODUCTION
Atlanticast Pacic rcd mangrovcs (AP Rhizophora
spccics) arc thc most important and dominant mangrovc
spccics ol tropical coastal arcas ol thc Atlantic ccan, thc
Amcrican Pacic coast, and scvcral islands in thc south
wcstcrn Pacic ccan. Tis spccics group is onc ol two
that makc up thc gcnus Rhizophora and consists ol thrcc
spccics (two bcing closcly allicd) and onc hybrid: R. mangle,
R. samoensis, R. racemosa, and R. harrisonii, rcspcctivcly.
Rcd mangrovcs, notably R. mangle, havc also bccn intro
duccd into ncw sitcs in thc !ndoVcst Pacic (!VP) rc
gion during thc past ccntury. !n thc Hawaiian and Socicty
!slands, no mangrovcs wcrc prcscnt until introductions
wcrc madc in thc .acs, sincc which timc thcir prcscncc has
bccomc quitc noticcablc. !n thc Hawaiian !slands, man
grovcs havc rcportcdly ovcrgrown channcls, rcduccd tidal
ows, and ovcrgrown archcological sitcs. Rcd mangrovcs
thrivc undcr a rangc ol intcrtidal wctland conditions, in
cluding high salinity lcvcls lrom grcatcr than lull strcngth
scawatcr to lrcshwatcr (Cintron ct al. .,), and thcy tolcr
atc a rangc ol ooding, soil typcs, and othcr physical sitc
lactors. Typically, thcy arc most common in thc middlc to
low intcrtidal zonc abovc mcan sca lcvcl, cxtcnding oltcn
along thc scaward margin ol mangrovc stands.
Many ol thc valucs ol rcd mangrovcs arc dicult to scpa
ratc lrom thc largcr rolc playcd by mangrovc plants. As
such, thc rolcs ol particular spccics ol Rhizophora arc oltcn
not distinguishcd lrom othcr mcmbcrs ol thc gcnus, in
cluding thc !VP stilt mangrovcs. 8ccausc Rhizophora spc
cics dominatc most tropical mangrovcs worldwidc thcy arc
gcncrally bclicvcd to play a vital rolc lor mangrovc ccosys
tcms including shorclinc protcction, cnhanccmcnt ol watcr
quality in ncarshorc cnvironmcnts (plus coral rccl arcas),
and support ol cstuarinc and marinc lood chains.
Rcd mangrovcs arc gcncrally considcrcd nonnativc to
thc !VP, not withstanding thc disjoint but natural occur
rcncc ol R. samoensis in Ncw Calcdonia, Ncw Hcbridcs,
Fiji, Tonga, and Samoa. Howcvcr, whcrc introductions ol
R. mangle havc bccn madc, thc uscs ol such plants nccd
to bc wcighcd carclully against thcir cccts as potcntially
invasivc spccics. !n Hawaii, lor cxamplc, scvcral important
ncgativc cccts havc bccn documcntcd, including rcduc
tion in thc habitat quality lor cndangcrcd watcr birds such
as thc Hawaiian stilt (Himantopus mexicanus knudensi),
colonization ol habitats to thc dctrimcnt ol nativc spccics
(c.g., in anchialinc pools), ovcrgrowing nativc Hawaiian
archacological sitcs, and causing localizcd drainagc prob
lcms by rcducing thc ow through tidal crccks or drainagc
channcls.
DISTRIBUTION
Native range
Atlanticast Pacic rcd mangrovcs arc nativc to tropical
rcgions along thc Amcrican cast and wcst coasts to thc
Alrican wcst coast, as wcll as in isolatcd occurrcnccs in thc
southwcstcrn Pacic islands.
Rhizophora mangle is thc spccics that most charactcrizcs
AP Rhizophora. !t occurs naturally and dominatcs tropi
cal tidal arcas along both sidcs ol thc Atlantic. Tc closcly
rclatcd and almost idcntical sibling spccics, R. samoen-
sis (=R. mangle sensu lato Tomlinson .6, -R. mangle var.
samoensis Hochr.), is nativc to thc Amcrican wcst coast and
islands in thc southwcstcrn Pacic, notably Ncw Calcdonia,
Ncw Hcbridcs, Fiji, Tonga, and Samoa. Tc taxonomic sta
tus ol thcsc sibling taxa is arguablc bascd on morphological
data alonc. Tc status ol obscrvcd dicrcnccs in likc lorms
R. mangle and R. samoensis, howcvcr, will only bc rcsolvcd
in gcnctic studics and sclcctcd brccding programs.
Rhizophora racemosa is lcss common than R. mangle and
R. samoensis but occurs as a distinct coinhabitant ol man
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
grovc stands in wcttcr arcas and largcr catchmcnt cstuarics
ol thc Atlantic. Tc spccics lavors rivcrinc cstuarics, and
it is rcstrictcd in thc AP to a lcw stands along thc Pa
cic coast ol thc Amcricas and docs not occur in thc !VP.
At lcast onc othcr possiblc spccics, R. harrisonii, occurs
across thc samc rangc. Givcn this and its intcrmcdiatc
charactcrs, thc taxon is considcrcd thc putativc hybrid ol
R. mangle and R. racemosa. Tc distribution ol R. racemosa
and thc putativc hybrid R. harrisonii appcars rcstrictcd
mostly to cquatorial cstuarics ol largcr rivcr systcms with
morc continuous lrcshwatcr ows.
l grcat intcrcst also is thc natural prcscncc ol rcd man
grovc in thc southwcstcrn Pacic islands. Also occurring
in this rcgion is anothcr hybrid taxon, R. selala (Salvoza)
Tomlinson, whosc putativc parcnts includc R. samoensis
(-R. mangle var. samoensis) and R. stylosa (-R. mucronata
var. stylosa). Tis hybrid taxon is spccial bccausc R. stylosa
is a dominant mcmbcr ol thc !VP stilt mangrovcs. Tc oc
currcncc ol this hybrid mcans that thcrc appcars to bc vcry
littlc gcnctic scparation bctwccn thcsc dcning and most
divcrgcnt ol Rhizophora spccics.
Current distribution
8ccausc rcd mangrovcs arc rccognizcd as valuablc timbcr
produccrs, bcnccial to shorclinc stabilization and shcr
ics, it is lcasiblc that thcir dispcrsal wcstward in thc Pa
cic may havc bccn assistcd by aboriginal travclcrs in thc
past. Howcvcr, thcrc is no cvidcncc lor this happcning, so
currcntly thcrc is no agrccd cxplanation lor thc prcscncc
ol R. samoensis in thc southwcstcrn Pacic. !n thc abscncc
ol human intcrvcntion, and acccpting that longdistancc
dispcrsal was not possiblc, onc cxplanation is islandhop
ping by natural dispcrsion across an ancicnt archipclago
ol scamounts that crosscd thc southcastcrn Pacic during
carly lormation ol thc Pacic Platc in thc Latc Crctaccous
pcriod (ukc .a, .). Any migration would havc bccn
wcstward, with distanccs bctwccn thc putativc ancicnt is
lands largc cnough to rcstrict all mangrovc spccics cxccpt
thc dispcrsalspccialist rcd mangrovcs. !n support ol this
thcory, thcrc arc tantalizing similaritics in crossoccan
linkagcs lor scvcral shallowwatcr rccl sh.
Rcd mangrovcs cxtcnd notably bcyond thcir nativc rangc.
Rhizophora mangle has bccn introduccd to thc Hawaiian
and Socicty !slands lrom Atlantic populations in Florida.
!n cach casc, loundcr populations havc incrcascd and cx
pandcd dramatically, cspccially in thc Hawaiian !slands.
Plants introduccd in thc carly .ccs to Molokai and ahu
now cxtcnd to most islands ol thc group, and thc cxpccta
World distribution of red mangroves, the AEP Rhizophora species. Given the apparent hybrid status of R. harrisonii, its dis-
tribution is likely where the distributions R. racemosa and R. mangle/R. samoensis overlap. Coastlines marked in red indicate the
distribution of all mangroves. i:~cv: x.c. uuxv
4 Atlanticast Pacic Rhizophora spccics
tion is that thcy will sprcad lurthcr. Accordingly, R. mangle
is trcatcd as an invasivc spccics in thcsc islands.
Similar introductions ol R. mangle clscwhcrc arc appar
cntly lcss invasivc, possibly bccausc othcr mangrovcs
that alrcady occupy thc habitat rcducc cstablishmcnt op
portunitics lor any ncw introductions. !n thc Townsvillc,
Australia, arca, local authoritics wcrc not prcparcd to takc
thc risk, howcvcr, so whcn a small introduccd stand ol R.
mangle was discovcrcd in thc uppcr part ol Ross Rivcr, it
was dcstroycd.
BOTANICAL DESCRIPTION
Preferred scientic names
Rhizophora mangle L.
Rhizophora samoensis (Hochr.) Salvoza (-R. mangle var.
samoensis Hochr.)
Rhizophora racemosa G.F.V. Mcycr
Rhizophora harrisonii Lccchman (-R. mangle R.
racemosa)
Family
Rhizophoraccac (mangrovc lamily)
Common names
Atlanticast Pacic rcd mangrovcs, Amcrican mangrovc
(nglish)
rcd mangrovc (USA)
mangle rojo (Ccntral and Latin Amcrica, Pacic and
Caribbcan coasts)
tiri wai (Fiji)
togo (Samoa)
Size
Atlanticast Pacic rcd mangrovcs arc mcdium to tall
trccs. Tcy may rcach cc m (.cc.6c lt) in hcight, al
though thcy arc commonly much shortcr, around m
(.6a6 lt). Stcm diamctcrs arc about . cm (6. in)
takcn just abovc thc highcst prop root. Tis mcthod ol
mcasurcmcnt dicrs lundamcntally lrom thc standard di
amctcr at brcast hcight (dbh) uscd lor most lorcst survcys,
as diamctcr hcight abovc thc substratc varics lrom c.,
m (..a lt) (considcr thc trcc in thc photo on thc rst
pagc).
Form
Rcd mangrovcs arc oltcn multistcmmcd rambling to co
lumnar trccs with distinct abovcground prop roots. Tcy
tcnd to bc ol shortcr staturc and morc sprcading in shapc
on thc scaward cdgc ol stands or in sitcs ol highcr salinity.
Tallcr, singlcstcmmcd trccs arc oltcn lound just bchind
thc scaward cdgc ol stands downstrcam in major rivcr cs
tuarics. Multistcmmcd trccs occur in lrontal arcas but arc
morc common in uppcr intcrtidal rcgions. Prop roots arc
sturdy cvcn whcn rclativcly thin.
Flowering
Flowcrs arc pcrlcct, containing both malc and lcmalc parts.
!norcsccnccs havc lcw to many joints with .,a,,choto
mous branching and onc to many buds possiblc pcr in
orcsccncc. pcn owcrs arc locatcd within or bclow lcal
axils at lcal nodcs bclow thc apical shoot, dcpcnding on
spccics. For R. mangle, R. harrisonii, and R. racemosa,
maturc buds and owcrs arc locatcd at .a, , and ,
nodcs down lrom thc apical shoot, rcspcctivcly. Tc ca
lyx is typically waxy ycllow to crcamy whitc and grccn at
maturity, with lour lobcs. 8uds clongatc to ovatc, grccn
whcn immaturc to lightcr colors as thcy maturc, smooth,
dimcnsions .a cm (c.c. in) long, ~c. cm (c.a in) widc.
Pctals, usually lour, arc lanccolatc to lincar, crcamy whitc,
with woolly to sparscly hairy margins, ~.a mm (c. in) long
and ~ mm (c..6 in) widc. Stamcns numbcr cight, palc
ycllow, to goldcn brown at maturity. Stylc is palc grccn, tc
rctc, lilorm, c. mm (c.cac..6 in) abovc ovary basc,
.. mm (c.c6c..a in) widc, dichotomous tip, palc ycl
low. 8racts and bractcolcs arc distinct. Pcdunclc is cm
long (..a..6 in), ~c. cm (c.c. in) widc. Flowcring pcriod
is chicy (but not cxclusivcly) Augustcccmbcr in thc
southcrn hcmisphcrc and FcbruaryJunc in thc northcrn
hcmisphcrc.
Leaves
Lcavcs arc oppositc, simplc, bright grccn, obovatc, lcathcry,
margins rcvolutc, gcncrally curvcd surlacc, obtusc blunt
apcx with a minutc lip loldcd undcr. Cork wart spots occur
on most spccics lcal undcrsurlaccs, scattcrcd cvcnly, not
raiscd, rcddish brown (may bc distinguishcd lrom inlcc
tions and wounds by thcir unilorm covcr). Notc that non
spottcd lcavcs arc lound on an unusual lorm ol R. racemosa
in northcrn 8razil. Lcal cmcrgcncc occurs chicy during
NovcmbcrFcbruary in thc southcrn hcmisphcrc and May
August in thc north. Lcal lall occurs chicy ovcr thc wct
summcr pcriod, ctobcrFcbruary in thc southcrn hcmi
sphcrc and AprilAugust in thc northcrn hcmisphcrc.
Fruit
Fruits, whcn maturc, arc pcarlikc, clongatc, waist constric
tion, smooth brown surlacc, calyx lobcs clongatc sprcading
(whcn thc hypocotyl is rcady to cmcrgc). For R. mangle,
R. harrisonii, and R. racemosa maturc lruit locatcd in lcal
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 5
axils , , (rarc) and .c.. nodcs down lrom apical shoot,
rcspcctivcly.
Seeds/hypocotyls
Likc all Rhizophora spccics, rcd mangrovcs arc viviparous,
mcaning that thcy producc sccds hiddcn in thc maturc
lruit, and thcsc gcrminatc whilc thc lruit is still attachcd to
thc parcnt plant. Tc dispcrsal 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
widcr, distal tip pointcd. Rhizophora mangle and R. samo-
ensis both havc distinct brown distal portions ol othcr
wisc grccn, rclativcly short hypocotyls (scc photo). For R.
mangle, R. harrisonii, and R. racemosa, maturc hypocotyls
arc locatcd in lcal axils , ~ (rarcly obscrvcd), and ..
nodcs down lrom thc apical shoot, rcspcctivcly.
Fruiting, whcn maturc hypocotyls lall, occurs chicy (but
not cxclusivcly) in NovcmbcrJanuary in thc southcrn
hcmisphcrc and MayJuly in thc northcrn hcmisphcrc.
Bark
8ark is smooth and rcdbrown in scaward and cxposcd lo
cations (rocky and sandy substratcs), to grayssurcd with
smooth, rcdbrown prop roots in shcltcrcd locations (mud
substratcs). Tcrc is total covcragc ol grayssurcd bark in
somc localitics.
Top left: Te blunt apex of leaves is characteristic of all AEP red mangroves, as in this R. samoensis from Fiji. Top right: Leafy ro-
sette of Rhizophora mangle (similar to R. samoensis) showing open owers with elongate, reexed calyx lobes. Bottom left: Open
ower of Rhizophora racemosa showing at, slightly hairy petals and sti-erect, non-reexed calyx lobes. Bottom right: Mature
fruit of Rhizophora mangle (similar to R. samoensis) showing its elongate pear-shape, prior to emergence of the hypocotyl. Note
the persistent style at the distil tip of the fruiting body. Atlantic Panama. vno:os: x.c. uuxv
6 Atlanticast Pacic Rhizophora spccics
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 m) dcpth. Tis
conlorms to thc anoxic conditions commonly obscrvcd in
mangrovc scdimcnts.
Similar species
Rcd mangrovcs arc distinguishcd lrom !VP stilt man
grovcs principally by thc lcal apcx: rcd mangrovcs havc
a blunt lcal apcx and lack thc spikcd, mucronatc lcal tip
prcscnt in stilt mangrovcs.
Rhizophora mangle and R. samoensis, with mostly c in
orcsccncc joints, arc distinguishcd lrom R. racemosa and
R. harrisonii, which havc inorcsccncc joints.
Rhizophora racemosa is distinguishcd lrom R. harrisonii
by thc nodc position ol maturc buds and owcrs in lcal
axils, down lrom thc apical shoot, bcing , nodcs and
rcspcctivcly. Tc possiblc hybrid charactcr is shown in
R. harrisonii, whcrc it has charactcrs intcrmcdiatc bc
twccn R. racemosa and R. mangle.
Rhizophora mangle and R. samoensis, thc sibling spccics, arc
distinguishcd by minutc/abscnt bracts on pcdiclcs at thc
calyx basc in R. mangle, whilc R. samoensis has bracts twicc
as widc as thc pcdiclc. As notcd, R. mangle and R. samoen-
sis appcar vcry closcly rclatcd and arc likcly to bc thc samc
spccics. iscriminating bctwccn thcm rcliably is not pos
siblc in many instanccs without dctailcd cxamination ol
kcy morphological and gcnctic charactcristics.
GENETICS
Variability of species
Atlanticast Pacic rcd mangrovcs arc thosc Rhizophora
spccics that occur naturally along thc cast and wcst coasts
ol thc Amcricas, as wcll as along thc wcst Alrican coast
and thosc island populations in thc southwcstcrn Pacic.
!n this group thcrc appcar to bc lour rclativcly distinct taxa,
although at timcs thcir morphological and taxonomic dil
lcrcnccs arc qucstionablc. Tc unccrtainty is chicy bascd
on: .) thc prcscncc ol onc intcrmcdiatc individual that is
rccognizcd as a distinct hybrid, namcly R. harrisonii, and
a) thc occurrcncc ol two sibling spccics, R. mangle and R.
samoensis, which may, on closcr cxamination, bc shown to
bc thc samc spccics.
Tc rclationship ol R. mangle and R. samoensis is pcrhaps
thc most contcntious. Tcy appcar vcry closcly rclatcd, as
thcy arc distinguishcd by only a lcw morphological charac
tcrs, as wcll as thcir distinct gcographic rangcs. Tc sibling
Mature hypocotyls of Rhizophora mangle (similar to R. samo-
ensis), showing distinctive brown distal end, Hawaii. vno:o:
x.c. uuxv
Expended fruits of Rhizophora samoensis (left) and R. stylosa
(right), Fiji. vno:o: x.c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 7
spccics scparatc naturally across thc Amcrican land barricr,
with R. mangle in thc Atlantic and R. samoensis in thc Pa
cic. Tc situation bccamc morc complcx during thc .ccs
whcn R. mangle (mostly lrom around Atlantic Florida)
was introduccd to thc Pacic in scvcral isolatcd instanccs,
notably to Hawaii, thc Socicty !slands, and northcastcrn
Australia. Tc Australian introduction was cradicatcd a
lcw ycars ago, but in both othcr locations thc sprcad ol
R. mangle has procccdcd unchcckcd, and thc spccics now
dominatcs most intcrtidal wctland arcas ol cach location.
!n Hawaii changcs in shorclinc ccology havc bccn dra
matic, cspccially bccausc no mangrovcs grcw in this iso
latcd location prcviously. A similar situation is cxpcctcd in
thc Socicty !slands, but no dctailcd rcports arc availablc.
Tcsc introductions highlight thc uncxplaincd disconti
nuity ol R. samoensis in thc southwcstcrn Pacic islands.
Tis is thc only cxamplc whcrc AP and !VP Rhizophora
spccics naturally cocxist. Curiously, thc human introduc
tions ol R. mangle dcmonstratc that island habitats with
out mangrovcs cxistcd bctwccn Samoa and South Amcrica,
and that thcsc arc suitablc lor mangrovc colonization. Tc
qucstion raiscd by this natural discontinuity rcmains unrc
solvcd. Tis obscrvation cmphasizcs thc prolound limita
tions ol longdistancc dispcrsal lor this mangrovc group ol
othcrwisc longdistancc dispcrsal spccialists.
Rccognizing thc morphological dicrcnccs bctwccn R.
mangle and R. samoensis is considcrcd usclul in distinguish
ing bctwccn introduccd populations ol R. mangle in thc
Hawaiian and Socicty !slands, as comparcd with natural
populations ol R. samoensis whcrc thcy occur clscwhcrc in
thc !ndoVcst Pacic.
8y contrast, R. racemosa is rcadily distinguishcd lrom R.
mangle and R. samoensis. Tis spccics appcars morc com
mon in cstuarics inucnccd by largcr and morc continu
ous lrcshwatcr ows in cquatorial rcgions ol thrcc distinct
rcgions: thc cast Pacic (Costa Rica to cuador), wcst At
lantic (\cnczucla to 8razil), and cast Atlantic (wcstcrn Al
rica). Tcrc arc no rcports ol dicrcnccs within this taxon
bctwccn thcsc arcas, but rcccntly (N.C. ukc, pcrs. obs.),
two lorms ol R. racemosa wcrc lound to cocxist in north
crn 8razil. nc ts thc typc dcscription in cvcry rcspcct
including thc prcscncc ol cork wart spots on lcal undcr
surlaccs, whilc thc othcr lorm lackcd thcsc spots. Tis was
particularly notablc sincc all rcd mangrovcs until now havc
bccn rcportcd to posscss such spots on thcir lcal undcr
surlaccs. Tis charactcr is prcscnt in !VP stilt mangrovcs
and in a similar way dcncs thc two lorms ol R. apiculata
occurring in Australasia and thc northwcstcrn Pacic.
Known varieties and hybrids
Rhizophora harrisonii is thc apparcnt hybrid ol R. mangle
and R. racemosa bascd on its intcrmcdiatc and sharcd mor
phological charactcristics. Furthcr invcstigations arc nccd
cd to adcquatcly dcscribc Rhizophora taxa and thcir dis
tributions throughout thc AP rcgion. Currcnt cvidcncc
indicatcs thc situation may bc morc complcx, with anothcr
possiblc hybrid whcrc R. racemosa apparcntly occurs on thc
castcrn Pacic coast ol thc Amcricas with R. samoensis, thc
allicd partncr to R. mangle.
Furthcrmorc, as notcd abovc, two lorms ol R. racemosa
wcrc obscrvcd in 8razil. Tc samc rcccnt invcstigation
also discovcrcd two intcrmcdiatc lorms, rccognizcd as po
tcntial hybrids bctwccn thc two R. racemosa lorms and R.
mangle. Kcy qucstions arisc lrom thcsc ncw obscrvations
including: what is thc distribution ol thc two lorms ol R.
racemosa, and how do thcsc comparc with populations in
wcstcrn Alrica: Tcrc arc clcarly morc qucstions than an
swcrs conccrning gcnctic variation in rcd mangrovcs.
Abcrrant individuals arc rcportcd to producc chlorophyll
dccicnt propagulcs, callcd albinos. Tis is particularly no
tablc and common in R. mangle stands throughout thc Ca
ribbcan arca to northcrn 8razil. Ycllow or rcd propagulcs
can bc obscrvcd hanging in acctcd trccs alongsidc normal
grccn propagulcs. Tcir rclativc numbcrs on an individual
trcc arc bclicvcd to quantily thc amount ol outcrossing
that occurs among ncighboring normal trccs. thcr trccs
havc also bccn obscrvcd with varicgatcd loliagc.
Culturally important related species
All Rhizophora spccics arc closcly similar in trcc lorm, and
cultural groups in thc Pacic rcgion may not always dis
tinguish bctwccn thcm. thcr mangrovc gcncra, such as
Bruguiera, arc considcrcd closc in lorm and valuc also, and
thcsc togcthcr arc oltcn uscd in similar ways by indigcnous
pcoplcs.
ASSOCIATED PLANT SPECIES
AP rcd mangrovcs can bc lound growing with othcr man
grovc spccics, prclcrably within intcrtidal wctland condi
tions abovc mcan sca lcvcl clcvations in tropical rcgions. R.
mangle and R. samoensis grow in marinc salinc conditions,
whilc R. racemosa lorms and hybrid grow in proximity to a
rcgular lrcshwatcr sourcc. R. racemosa arc commonly lound
in lowcr to middlc tidal rcachcs ol rivcrs and strcams with
rcgular, lrcqucnt lrcshwatcr runo, whilc R. mangle and R.
samoensis occur in coastal cmbaymcnts and coral islands. !n
Fiji, howcvcr, R. samoensis occurs togcthcr with R. stylosa (a
stilt mangrovc). !n this casc, R. samoensis occurs morc lrc
8 Atlanticast Pacic Rhizophora spccics
qucntly in upstrcam locations ol thc lrcshwatcr dominatcd
strcams, whilc R. stylosa rcmains thc marinc spccialist, prc
lcrring downstrcam and cxposcd coastal locations.
Rcd mangrovcs oltcn cxtcnd up thc tidal prolc to thc
tcrrcstrial lringc in arcas ol highcr rainlall. !n lowcrrain
lall arcas, rcd mangrovcs occur on cstuarinc margins just
abovc mcan sca lcvcl. 8cst dcvclopmcnt has bccn obscrvcd
in lowcr tidal rcachcs and on solt, nc, muddy scdimcnt
substratcs.
Associated species commonly found
!n Atlantic and cast Pacic populations, rcd mangrovcs
occur commonly with Avicennia germinans and Laguncu-
laria racemosa. !n thc southwcstcrn Pacic, R. samoensis is
associatcd with !VP mangrovcs, including R. stylosa and
Bruguiera gymnorrhiza prcdominantly. !n othcr locations,
rcd mangrovcs arc associatcd with othcr mangrovc spc
cics. Tc typcs ol associatcd spccics arc notably variablc
whcrc this spccics has bccn introduccd to thc !VP rcgion.
!n Hawaii, whilc thc spccics lrcqucntly occurs in ncarly
monospccic stands, it docs somctimcs occur alongsidc a
mangrovc introduccd lrom thc !VP (Bruguiera sexangula)
and spccics that typically occur in lowcrsalinity cnviron
mcnts, such as Hibiscus tiliaceus.
ENVIRONMENTAL PREFERENCES
AND TOLERANCES
Climate
Atlanticast Pacic rcd mangrovcs thrivc in tropical and
subtropical cnvironmcnts charactcrizcd by modcratcly
high and wcll distributcd rainlall. Mangrovc plants appcar
to dcpcnd on groundwatcr to sustain optimal growth, cs
pccially during dricr months. !n dricr locations, such as
thc 8aja Pcninsula on thc northcrn Pacic coast ol Mcxico,
thc stuntcd but dcnsc thickcts ol R. samoensis (=R. man-
gle:) attcst to thc grcat adaptability ol rcd 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
Tcy grow in climatcs with summcr or unilorm rainlall
pattcrns.
Dry season duration (consecutive months with <40
mm [1.6 in] rainfall)
8ccausc ol acccss to groundwatcr, rcd mangrovcs grow in
rcgions with up to months ol drought. Across a widc va
ricty ol climatic rcgions, mangrovc covcr cxpands and con
tracts through timc. Tis has bccn cvidcnt in corrclations
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 (c.ccF) (cstimatc)
Mean minimum temperature of coldest month
cC (c.F)
Minimum temperature tolerated
cC (aF) (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. Tcy arc also anacrobic with high conccntra
tions ol suldc. Trccs also grow wcll in sitcs with acrobic
scdimcnts consisting ol nc sands to coarsc stoncs and
rocks, and coral ramparts.
Soil texture
Plants grow in light, mcdium, and hcavy tcxturc soils
(sands, sandy loams, loams, and sandy clay loams, sandy
clays, clay loams, and clays).
Soil drainage
Tc trccs grow in soils with lrcc and unimpcdcd drainagc,
as wcll as watcrloggcd soils.
Soil acidity
pH 6.
Special soil tolerances
Plants grow bcst in salinc soils but survivc wcll in lrcsh wa
tcr. Tc optimal salinity rangc is rcportcd to bc lrom to
a6 ppt (parts pcr thousand), comparcd with approximatcly
6 ppt lor scawatcr.
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 9
Tolerances
Drought
Rcd mangrovcs can tolcratc drought pcriods wcll, although
trccs havc apparcntly bccn killcd by drought in somc sitcs
on Molokai, Hawaii whcrc groundwatcr sourccs appcar
to bc limitcd. rought conditions prcsumably causc soil
salinity to incrcasc in cxccss ol tolcrablc limits lor thcsc
plants.
Full sun
Tcy grow bcst in lull sun.
Shade
Tcy appcar to havc low tolcrancc ol shadc. Howcvcr, rc
ccnt cvidcncc shows this morc likcly duc to wccvils that
inlcst and kill coolcr, shadcd sccdlings (8rook acc., Sousa
ct al. acc).
Fire
Rcd mangrovcs havc no tolcrancc ol rc in closc proxim
ity.
Frost
Tolcrancc ol sublrcczing tcmpcraturcs is low to nonc.
Waterlogging
Rcd mangrovcs arc tolcrant ol daily tidal ooding up to
dcpths ol up to .. m ( lt). Vhilc tolcrating pcrmancntly
saturatcd soils, thcy arc intolcrant ol drying soils.
Salt spray
Tc trccs arc highly tolcrant ol salt spray.
Wind
Rcd mangrovcs arc typically lound in scaward arcas subjcct
to wind and salt spray but largcly protcctcd lrom wavcs.
Planting in such highly windpronc locations is rccom
mcndcd, but only whcrc rcquircd and within thc trccs na
tivc rangc.
Waves
!n gcncral, cxposcd, wavcpronc coastlincs arc inhospitablc
to mangrovcs. !n arcas whcrc wavc action is inlrcqucnt, rcd
mangrovcs arc bclicvcd to providc signicant bucring ol
coastal arcas during storm and tsunami surgc cvcnts (ah
douhGucbas acc).
Abilities
Self-prune
Rcd mangrovcs scllprunc wcll in dcnsc stands, but thcy
commonly maintain lowcr branchcs in morc opcngrow
ing locations.
Coppice
Tc trccs havc notably poor coppicc ability. Gcncrally, il
c or morc ol thc lcavcs arc rcmovcd lrom a trcc, it will
dic.
GROWTH AND DEVELOPMENT
Growth ratcs vary with agc. Tcy gcncrally grow lcss than
MANGROVES AND CLIMATE
CHANGE
Atlanticast Pacic rcd mangrovcs, likc othcr man
grovc 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 dcci
sivcly to shilts in kcy lactors, likc tcmpcraturc, rainlall,
and sca lcvcl, as cach spccics has dcncd rangcs ol tol
crancc lor cach lactor. For instancc, bccausc mangrovcs
arc charactcristically rcstrictcd to clcvations bctwccn
mcan sca lcvcl and highcst tidcs, as sca lcvcl riscs thcir
communitics must movc upland to survivc. Sincc man
grovcs havc narrow optimal tcmpcraturc rangcs, ris
ing tcmpcraturcs 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
rccstablishmcnt, and thc availability ol suitablc ncw
spacc. Clcarly, such changcs havc occurrcd throughout
history, so thc distribution ol mangrovcs today rcprc
scnts 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 Atlanticast Pacic Rhizophora spccics
. m/yr (. lt/yr) but can cxcccd this in lavorablc circum
stanccs. Hcight growth is rapid shortly altcr cstablishmcnt
whilc lood rcscrvcs arc takcn up lrom thc hypocotyls ol
cstablishcd sccdlings. Growth ratc slows whcn trccs ap
proach a sitcspccic maximum canopy hcight. Vhcn ncar
maturity and maximum hcight, trccs broadcn thcir cano
pics and incrcasc stcm diamctcr rathcr than grow tallcr.
Flowering and fruiting
Flowcring and lruiting pcriods ol rcd mangrovcs arc dis
tinctly scasonal. Pcak phcnocvcnts arc cxpcctcd to shilt
into latcr months 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 maturc hypocotyls. Tc duration dcpcnds on spc
cics, with thc longcst bcing R. racemosa with an cxpcctcd
pcriod ol around c months lor cach rcproductivc cyclc.
Yields/growth rates
Growth ratcs vary with spccics, spatial position in thc stand,
compctition, vigor, and agc. For northcrn Pacic sitcs in
Costa Rica, Jimcncz (in Chong .) rcportcd annual di
amctcr growth incrcmcnts lor R. mangle and R. racemosa
trccs ol c..c.. cm (c.c6c.c, in) lor diamctcr sizc
classcs lcss than .c cm ( in), and c.cc.., cm (c.cc.c,
in) lor diamctcr sizc classcs grcatcr than .c cm.
Chong (.) providcs inlormation on potcntial yiclds
lrom Playa Garza Pilot Arca in an invcstigation ol thc lca
sibility ol managing mangrovc lorcsts thc TcrrabaSicrpc
Forcst Rcscrvc on thc southcrn (wcttcr) Pacic coast ol
Costa Rica. Tc cstimatcd mcan annual incrcmcnt (MA!)
was 6 m

/ha/yr (6 lt

/ac/yr) with somc stands as high as

... m

/ha/yr (.,acc lt

/ac/yr).
Reaction to competition
Rapid carly growth ol rcd mangrovc sccdlings in lull sun
light cnsurcs thcir succcss and dominancc in prclcrrcd cs
tuarinc and intcrtidal conditions. Ncwly cstablishcd sccd
lings grow bcst in closc proximity with thcir samc spccics
cohort. Tis aords thcm maximal protcction lrom physi
cal damagc by drilt logs and crosivc wavcs. Sincc compcti
tion is high bctwccn ncighboring sccdlings, slowcr plants
dic and dccomposc quickly, lcaving lastcr compctitors thc
bcnct ol not only thc spacc thcy occupicd but also thcir
nutricnts.
PROPAGATION
Although natural rcgcncration is gcncrally rclicd upon
lrom Pacic to Atlantic rcgions, thcsc spccics arc rclativc
ly 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 dis
tal cnd ol thc ac6c cm (a in) long hypocotyl onc third
ol its lcngth into thc scdimcnt, spaccd at about ... m
(. lt) intcrvals. No holcs nccd bc dug, and ncithcr nurs
cry prcparation nor stakcs arc nccdcd. Low maintcnancc is
gcncrally sucicnt lor maximizing sccdling cstablishmcnt
succcss in shcltcrcd arcas. Howcvcr, substantial protcction
is rcquircd in morc cxposcd coastal locations during thc
rst dccadc ol cstablishmcnt. Such protcction mcthods
may includc cncascmcnt ol individual 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. Propagulc maturation occurs latcr in rclativcly
highcr latitudc locations north or south bcyond thc tropic
zonc (c.g., Julyctobcr in Hawaii).
Maturc propagulcs may bc collcctcd lrom thc ground altcr
thcy havc lallcn or pickcd dircctly lrom 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 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
Proccssing ol maturc propagulcs is not rcquircd lor rcd
mangrovcs, although damagcd and inscctinlcstcd indi
viduals should bc rcmovcd. Tc lollowing prctrcatmcnt
scction givcs 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
burlap bags and kccping thcm out ol dircct sunlight. Prc
trcatmcnt is considcrcd csscntial il considcring such stor
agc (scc bclow). !t is likcly that propagulcs can bc kcpt
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 11
longcr, but storagc bcyond a wccks is not rccommcndcd,
and longtcrm storagc is not lcasiblc.
Propagule pretreatment
Prctrcatmcnt ol propagulcs is gcncrally considcrcd unncc
cssary. Howcvcr, a study ol scolytid bcctlc larvac (Cocco-
trypes fallax) inlcctions on Rhizophora propagulcs in Aus
tralia lound at lcast . wcrc hcavily inlcstcd (8rook acc.).
!nlcstcd propagulcs wcrc cstablishcd undcr canopyshadcd
arcas. Tc study wcnt lurthcr to nd that prctrcatmcnt in
a cC (.aaF) watcr bath lor minutcs killcd thc bcctlcs
and rcmovcd thc risk ol cstablishmcnt losscs duc to borcr
damagc. Hcat trcatmcnt might bc casily achicvcd by lcav
ing thc collcctcd propagulcs in thc sun lor a short pcriod
(a lcw hours) bclorc planting.
Growing area
Rcd 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
Sccdlings arc rcady lor outplanting at thc sixlcal (thrcc
nodc) stagc il grown up in nurscry conditions, which can
takc up to 6 months to achicvc. ldcr sccdlings up to c.
m (.. lt) tall havc also bccn succcsslully transplantcd, but
this is not rccommcndcd. ircct planting ol largc numbcrs
ol propagulcs is rcstrictcd by thcir pcak scasonal availabil
ity, as propagulcs do not kccp lor cxtcndcd pcriods.
Guidelines for outplanting
utplanting anytimc ol thc ycar is suitablc. Propagulcs or
nurscrygrown sccdlings usually havc cxccllcnt survival in
sitcs corrcctly sclcctcd and, il appropriatc, protcctcd lrom
disturbancc. Survival ratcs ol c or grcatcr arc rcasonablc
in such circumstanccs.
n thc othcr hand, survival may bc zcro on sitcs cxposcd
to cxccssivc wavc action, on sitcs with inappropriatc hy
drologic or salinity rcgimcs, or (rarcly) subjcctcd to distur
bancc by grazing animals (c.g., goats, shccp, cattlc, 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, rcd mangrovcs posc lcw signicant disadvan
tagcs whcn plantcd within thcir nativc rangc. Tcy arc not
cspccially susccptiblc to pcsts or pathogcns and havc not
bccn rcportcd to host major pcsts or pathogcns ol impor
tant crop spccics.
Potential for invasiveness
Tc vcry succcsslul introduction and rapid sprcad ol rcd
mangrovcs in thc Hawaiian !slands clcarly dcmonstratcs
thc potcntial lor invasivcncss ol mangrovcs in arcas whcrc
suitablc habitat is availablc. Tc plants arc rcadily opportu
nistic duc to thcir rclativcly widc tolcrancc lor salinity and
Mature hypocotyl of Rhizophora racemosa showing distinc-
tive, slender, smooth propagule, Brazil. vno:o: u. :vniic
12 Atlanticast Pacic Rhizophora spccics
soil conditions. Also, thcir oating propagulcs arc sprcad
widcly by occan currcnts ovcr grcat distanccs.
Tc spccics R. mangle has unlortunatcly now takcn on thc
public status ol invasivc wccd and pcst spccics in Hawaii.
To allcviatc public conccrn, cradication corts havc bccn
carricd out in scvcral locations on ahu and Hawaii. !t
is not clcar about thc succcss or ccctivcncss ol this cam
paign, as it appcars to havc bccn bascd on subjcctivc inlor
mation and no monitoring.
Pests and diseases
Susccptibility to pcsts and discascs is bclicvcd to bc low,
with thc cxccption ol inscct borcrs and crabs that prcy on
propagulcs. For introduccd stands in thc Hawaiian islands,
damagc to propagulcs and lcavcs is notably lowcr than
within thc spccics nativc rangc, and productivity (as cx
prcsscd in littcr lall) is highcr.
Host to crop pests/pathogens
No rcports arc lound ol rcd mangrovcs scrving as hosts lor
known major crop pcsts or pathogcns. !n Hawaii, howcvcr,
R. mangle stands havc scrvcd as idcal sitcs lor thc nonna
tivc cattlc cgrct (Bubulcus ibis) and also somctimcs harbor
signicant populations ol rats.
AGROFORESTRY/ENVIRONMENTAL
PRACTICES
Mulch/organic matter
Mulch in Rhizophora lorcsts is hiddcn lrom vicw. !l it wcrc
not lor thc small mangrovc crabs, lallcn 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
Rcd mangrovc lorcsts stabilizc soils with thcir nctwork
ol sturdy ovcrlapping prop roots, which dampcn watcr
movcmcnt and promotc scdimcntation in arcas that might
othcrwisc bc crodcd. A major rcason that rcd mangrovc
was introduccd into Hawaii was to stabilizc mud ats that
wcrc cxpanding as a rcsult ol sugarcanc production and
rcsultant crosion. Rhizophora mangle has provcn quitc cl
lcctivc lor this purposc and has bccn shown to improvc
watcr clarity in ncarshorc cnvironmcnts, prcsumably duc
to its rolc in scdimcnt trapping and stabilization.
Fence posts
Rcd mangrovc stcms makc good posts, as thc wood is gcn
crally hard and rcsistant to inscct borcrs.
Windbreaks
Rcd mangrovc lorcsts providc a windbrcak along coastal
margins gcncrally, and placcs to scck sanctuary during ty
phoons and hurricancs.
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. Rcd mangrovc timbcr is vcry usclul lor small con
struction and lor lucl lor cooking. Convcrting thc wood
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
Rcd mangrovc is a known sourcc ol nativc animal loods.
Scvcral obscrvations dcmonstratc thc divcrsity and quan
tity, and it is thought to bc cxtrcmcly important in man
grovc ccosystcms. Numcrous insccts, crabs, and mollusks
grazc on grccn lcavcs in thc lorcst canopy and on lallcn
lcavcs on thc lorcst oor. Vhcrc prcscnt, 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
mammals (probably nonc in introduccd Hawaiian stands)
appcar to usc rcd mangrovc as a major lood sourcc.
Wildlife habitat
!n addition to aquatic marinc organisms (scc Fish/marinc
lood chain), rcd mangrovcs scrvc as habitat lor a widc
rangc ol tcrrcstrial and arborcal wildlilc. !n various loca
tions throughout thc rcgion, thcsc mangrovc lorcsts pro
vidc shcltcr and lood lor a numbcr ol associatcd launa, in
cluding birds, small mammals, shcllsh, and othcr marinc
lilc.
Bee forage
Rhizophora spccics havc no ncctar, but thcy do producc
copious pollcn that is usually distributcd by wind. !ntcr
cstingly, onc rcason statcd lor thc introduction ol rcd man
grovc to Hawaii was as a pasturc plant lor bccs (Cookc
..,). Clcarly, it wasnt thc bcst choicc lor this purposc!
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 rcd mangrovcs during part ol thcir
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 13
lilc cyclcs, as do spccics ol shrimp and crabs. As mcntioncd
abovc, scncsccnt lcavcs that havc lallcn lrom Rhizophora
trccs arc takcn by grapsid (small mangrovc) crabs into thcir
burrows. !n Hawaiian populations, thcrc appcars to bc cx
ccssivc lcal accumulation in somc locations, suggcsting
that normal associatcd launa and othcr dccompositional
biota arc lacking.
Coastal protection
Rcd mangrovc lorcsts and mangrovcs in gcncral play an
important rolc in protcction ol coastlincs, shponds, and
othcr coastal inlrastructurc. Rcd mangrovcs arc plantcd lor
coastal or shpond protcction in somc arcas (c.g., in Tcr
rabaSicrpc Forcst Rcscrvc, Costa Rica), and thcrc arc laws
in many locations aimcd at protccting mangrovcs in largc
part bccausc ol this important lunction.
USES AND PRODUCTS
Rcd mangrovcs arc probably ol grcatcst valuc lor thcir cn
vironmcntal bcncts, bccausc thcy (and mangrovcs gcncr
ally) arc bclicvcd to play a vital rolc in supporting marinc
lood chains, protccting coastal arcas, and improving watcr
quality.
!n tcrms ol dircct bcncts to pcoplc, thc most widcsprcad
usc ol rcd mangrovcs is lor wood uscd lor a rangc ol pur
poscs lrom cooking lucl to construction ol homcs and ca
noc parts. thcr uscs ol thc trcc includc dycs, mcdicincs,
and tannin lor tanning lcathcr. !t sccms likcly that rcd
mangrovcs may havc scvcral othcr uscs (c.g., as cattlc lccd)
that to datc havc not bccn lully cxplorcd.
Staple food
Lcavcs and hypocotyls arc cdiblc but not widcly uscd lor
lood.
Medicinal
Rcd mangrovc bark has rcportcdly bccn uscd to trcat an
gina, 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 cl
lcctivc thc trcatmcnts havc bccn in thcsc cascs.
Animal fodder
nc rcport (Morton .6) concludcd that rcd mangrovc
lcavcs might scrvc as a valuablc sourcc ol cattlc lccd, but
this potcntial has yct to bc rcalizcd.
Timber
Tc wood ol rcd mangrovcs is widcly uscd lor structural
componcnts ol traditional homcs (c.g., polcs, bcams, oor
ing, wallcladding, raltcrs) and othcr componcnts includ
ing undcrground minc supports, lcncing, cabinct works,
tool handlcs, and boat anchors. Tc wood is also uscd lor
othcr purposcs, ranging lrom traditional uscs such as sh
ing stakcs, spcars, and copra huskcrs to usc as a sourcc ol
chips lor pulp production.
Tc hcartwood ol R. mangle is light rcd to dark rcd or
rcddish brown or purplish, with unilorm or morc or lcss
stripcd grain. Tc sapwood is ycllowish, grayish, or pink
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 Atlanticast Pacic Rhizophora spccics
ish. Tc wood tcxturc is nc to mcdium, grain
straight to irrcgular, lowlustcr, without dis
tinctivc odor or tastc. Tc spccic gravity (ovcn
dry wcight/grccn volumc) is c..
Fuelwood
Rcd mangrovc wood is uscd lor luclwood in
many placcs along thc Amcrican Pacic coast
(c.g., Panama, Costa Rica, Nicaragua). Tc
wood is also madc into charcoal in many Ccn
tral Amcrican countrics, including Panama and
Costa Rica.
Canoe/boat/raft making
Tc wood has bccn uscd to makc canoc parts.
Tannin/dye
Tc bark and hypocotyls arc uscd to tan lcathcr
and to producc dycs ranging lrom rcdbrown to
black (thc lattcr with rcpcatcd dying). Tannin
contcnt ol thc bark is high lor most Rhizophora
spccics.
COMMERCIAL CULTIVATION
Rcd mangrovc timbcr is harvcstcd commcr
cially lor charcoal production through much ol
its rangc, including Ccntral and Latin Amcrica.
Tc caloric valuc ol thc timbcr is signicantly
cnhanccd by convcrting it to charcoal. Tis is
donc with all local Rhizophora spccics, as wcll
as Pelliciera and Laguncularia.
Rehabilitation and replanting
Projccts to rcplant and rchabilitatc mangrovc
lorcsts havc bccn conductcd whcrc thcy havc
bccn damagcd in a signicant way. For cxamplc,
in 8ahia las Minas, Atlantic Panama, a largc
oil spill killcd around c ha (.a ac) ol man
grovcs in .6 (ukc ct al. .,), and Rcncria
Panama lcad a projcct to rcplant thc damagcd
arcas. 8ccausc thc soils wcrc oilcd it was dccidcd to plant
R. mangle sccdlings using clcan tcrrcstrial scdimcnts. To
tal plantings cvcntually amountcd to in cxccss ol .cc,ccc
sccdlings. A subscqucnt invcstigation (ukc .6) lound
natural rccruitmcnt was c timcs grcatcr than plantcd
numbcrs, and natural sccdlings grcw cqually as wcll as, or
bcttcr than, plantcd sccdlings. !t was ol grcat importancc
that naturally rccruitcd sitcs rccovcrcd morc quickly than
plantcd sitcs, possibly bccausc ol sitc damagc during plant
ing. Furthcrmorc, damagc to cxposcd and damagcd man
grovc arcas incrcascd altcr 6 ycars whcn dcad standing
timbcr dramatically dctcrioratcd and was movcd by wavc
action with cach tidc and storm.
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
multiplc stcmmcd and/or shortcr trccs. Vidcr spacing and
thcrclorc sprcading trccs may bc dcsircd lor coastal protcc
tion projccts but not lor timbcr production. !n thc abscncc
Top: Charcoal treatment plant established with the DANIDA Mangrove
Project in Pacic Costa Rica, supported by IUCN and CATIE. Bottom:
Typical earthen kiln used to convert stacked red mangrove poles into char-
coal for local heating needs, Pacic Panama. vno:os: x.c. uuxv
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org) 15
ol signicant natural mortality, timbcr plantations should
bc thinncd to spacing ol a. to . m (.. lt) bctwccn trccs
as thc stand dcvclops and bccomcs crowdcd.
Management objectives and design consider-
ations
Somc publishcd guidclincs lor mangrovc silviculturc cxist
and arc rclcrcnccd bclow, but spccic guidclincs on thin
ning, lcrtilizing, ctc., arc lacking.
Growing in polycultures
Rcd mangrovcs naturally occur in mixcdspccics stands,
and cach spccics has its own ccological and cconomic val
ucs. !t is also important to plant associatcd bucr arcas,
particularly along thc shorclinc whcrc mangrovcs grow
bcttcr adjaccnt to banks stabilizcd by shorclinc upland
plants. Togcthcr thcy will complcmcnt and cnhancc thc
richncss and stability ol thc plantcd cnvironmcnt.
Estimated yield
For thc southcrn (wcttcr) Pacic coast ol Costa Rica,
Chong (.), in an FAsponsorcd projcct, providcd in
lormation on potcntial yiclds lrom thc ac ha (c ac) Pla
ya Garza Pilot Arca in an invcstigation ol thc lcasibility ol
managing around acc ha (.a,cc ac) ol mangrovc lorcsts
in thc surrounding TcrrabaSicrpc Forcst Rcscrvc. !t was
rcportcd that yiclds rcsulting lrom uncontrollcd lclling ol
.cc ha/yr (ac ac/yr) could bc achicvcd by lclling just .c
ha/yr (a ac/yr) il conductcd morc systcmatically.
Chong cstimatcd that cach hcctarc ol man
grovc in this rcscrvc could producc USs6.
(sac/ac) annually. A dctailcd managcmcnt
plan was proposcd in thc rcport. Tc avcragc
stand volumc lor rcd mangrovcs was .6 m

/ha
(ac lt

/ac), with a total mangrovc lorcst stand

volumc around ac m

/ha (ccc lt

/ac). sti
matcd yiclds ol timbcr volumc undcr bark wcrc
.cc.c m

/ha/yr (.cca.cc lt

/ac/yr) lor rcd

mangrovcs, and lor thc codominant mangrovc
Pelliciera, yiclds arc cstimatcd at 6c m

/ha/yr
(6c lt

/ac/yr).
Markets
Markcts lor rcd mangrovc products arc local
in naturc, with littlc availablc cxccpt lor rc
wood and charcoal in most placcs, othcr than
indircct products such as mangrovc crabs and
lruit bats. !n Ccntral Amcrica, largc quanti
tics ol rcd mangrovc wood chips and charcoal
may bc movcd grcatcr distanccs and in grcatcr
volumcs than othcr wood products. !n Panama thcrc is a
strong tradc in rcd mangrovc polcs around m (.c lt) in
lcngth (around .c. ycars old) lor construction ol bohio,
thc locally popular outdoor rccrcation shcltcrs lor barbc
cucs and partics.
INTERPLANTING
Somc intcrplanting systcms includc:
Example Bahia las Minas oil spill (ukc .6,
ukc ct al. .,)
Location
8ahia las Minas, Panama Atlantic coast
Description
Planting was undcrtakcn in . and .. Around c ha
(.cc ac) wcrc plantcd with Rhizophora mangle, sponsorcd
and implcmcntcd by Rcncria Panama.
Crop/tree interactions
Not applicablc.
Spacing/density of species
Spacing was about .. m ( lt).
Planting of red mangroves in an attempt to replace around ha of tree
loss following a major oil spill in Bahia las Minas, Atlantic Panama. vno:o:
x.c. uuxv
16 Atlanticast Pacic Rhizophora spccics
Example Terraba-Sierpe Forest Reserve
(Chong .)
Location
Costa Rica, TcrrabaSicrpc.
Description
Planncd projcct. Tis projcct rcachcd thc planning stagcs
only and was sponsorcd by FA with !UCN and CAT!.
Tc spccics to bc plantcd includcd R. racemosa, R. mangle,
and Pelliciera rhizophorae.
Crop/tree interactions
Not applicablc, as thcrc wcrc no crops.
Spacing/density of species
Spacing was planncd to bc .. m ( lt) lor all 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~
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plantnctwork.org~.
18 Atlanticast Pacic Rhizophora spccics
Species Proles for Pacic Island Agroforestry (www.traditionaltree.org)
Rhizophora mangle, R. samoensis, R. racemosa, R. harrisonii
(AtlanticEast Pacic red mangroves)
Authors: Norman C. ukc
.
and Jamcs A. Allcn
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~.
a. Paul Smiths Collcgc, P 8ox a6, Paul Smiths, Ncw York .a,c, USA, Vcb: http://www.paulsmiths.cdu
Acknowledgments: Tc authors and publishcr thank Tom Colc, Farid ahdouhGucbas, alc vans, Kathy wcl, Aricl Lugo, and
ianc Ragonc lor thcir input. A photo contribution by Ull Mchlig is grcatly apprcciatcd. Norm ukc 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.
Recommended citation: ukc, N.C., and J.A. Allcn. acc6. Rhizophora mangle, R. samoensis, R. racemosa, R. harrisonii (Atlantic
ast Pacic rcd 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.