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Different Kinds of Mangrove Forests Provide Different Goods and Services Author(s): Katherine C. Ewel, Robert R.

Twilley, Jin Eong Ong Reviewed work(s): Source: Global Ecology and Biogeography Letters, Vol. 7, No. 1, Biodiversity and Function of Mangrove Ecosystems (Jan., 1998), pp. 83-94 Published by: Blackwell Publishing Stable URL: . Accessed: 27/01/2012 10:56
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Global Ecology Biogeography and Letters (I1998)7, 83-94

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kinds mangrove of forests different Different provide goods and services

KATHERINE C. EWEL*, ROBERT R. TWILLEYt and JIN EONG ONGt USDA ForestService, PacificSouthwest ResearchStation,1151 Punchbowl Rm. 323, HonoluluHI 96813 USA (e-mail: St., LA 70504 U.S.A. of of Louisiana,Lafayette tfDepartment Biology,University Southwestern Sains Malaysia, 11800 Penang,Malaysia and tCentreMarine Coastal Studies,Universiti Abstract. The goods and servicesthat mangrove forests provideto societyare widelyunderstoodbut maybe too generally statedto serveas useful guidelines in decision-making. Understandingthe differences between fringe, riverine, basin forests and mayhelp to the and to determine bestuse of focustheseguidelines a particularforest.Fringe mangrovesare important nutrient concentrationsassociated with sediment serveas nutrient sinksforboth trapping. Basin forests enhanced ecosystem natural and anthropogenically sources of wood processes and are oftenimportant products.Exploitationof a forestfor one particular othergoods reasonmaymakeitincapableofproviding and services. flood protection, Key words. Mangrove forests, nutrients,wastewater recycling,timber harvesting, shrimp ponds.

primarily shoreline for protection. Riverine forests,

to of whichare likely be themostproductive thethree are to typesof forests, particularly important animal and plant productivity, perhaps because of high

are Mangroveforests widelyrecognizedas providing to a widevariety goods and services people,including of of protection from floods,provision a variety plant of sediment and nutrient and animal products, trapping, uptake ahd transformation (FAO, 1994). Destruction of these forestscontinues,however,in spite of this understanding theirimportance.Although these of wetlands abundantalongmany are protected, shorelines but around the world, the life-sustaining stillpoorly can provide, suchas support documented benefits they to offshore fisheries, likelyto be diminishing. are The lack of a direct,easily observed relationship a and thebenefits provides between mangrove forest it (and sometimesthe lack of sufficient research to document it) may be one reason for continued exploitation, often and loss,ofthesewetlands. Another reason may be the generalitythat cloaks many discussionsof the importanceof these wetlands.In fact,not all mangroveforestsprovide all the goods * Corresponding author.
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and servicesattributed them.There are significant to in of differences thecharacteristics mangrove habitats, not only betweencontinents and regionsbut within individual standsof mangroves well.Using a simple as rationaleforclassifying givenmangrovestand may a in its value assistland-usemanagers determining likely to societyand subsequently using it more wisely. in The purpose of this paper is to develop a simple of functionalclassification mangroveforestsand to identify whichgoods and servicesare likelyto derive fromwhichkindsof forests.


Efforts understandmangroveforestsfocused for to many years on the significanceto tree species in distributionsof spatial differences soil water characteristics (Macnae, 1968), short-term differences in propaguledispersal and survival (Rabinowitz,1978; Smith, 1987), competition among species (Clarke & Hannon, 1971), and geomorphologicalhistory or characterization of estuaries (Thom, Wright & 83

84 K. C. Ewel, R. Twilley Jin R. and EongOng

Coleman, 1975). Some, studies have de-emphasized tree species zonation because of the large numberof exceptions simplepatterns to (e.g. West, 1977). High and low intertidal as zones are likelyto differ a result in of gradients frequency inundation, porewater of soil salinity, and soil waterlogging,that are usually interrelated often and difficult predict theabsence to in of information about both regional hydrologyand animal activity. No consistentpatternor cause of mangrove zonation has yet to be derived (Smith, 1992). Two systems thatmaybe particularly usefulto land managers havebeenproposedforclassifying mangrove forests.Six different kinds of mangroveforestsare in where commonly distinguished theNeotropics, many are underlain a carbonatelimestone base: overwash, by fringe, riverine, basin, scrub,and hammock(Lugo & Snedaker,1974, Cintr6n, Lugo & Martinez,1986). A moregeneralsystem been proposedforOld World has mangroves,which are more likely to develop on accreting sedimentsdeposited by rivers and tides three (Woodroffe,1992). This systemdistinguishes based on dominant riverextremes, physical processes: and interiormangrove dominated, tide-dominated, Intermediate kindsof forests, the forests. including six can this New Worldtypes, be locatedwithin framework (Fig. la). In this paper,we adopt a hybridof thesesystems, of the combining familiarity Lugo & Snedaker'sterms withthe flexibility Woodroffe's of We to system. refer rivertide-dominated as mangroves fringe mangroves, dominated mangroves as riverinemangroves,and interior mangroves basinmangroves as (Fig. Ib). Basin mangrovesare likelyto contain the most variation within region, a low zones including and highintertidal far as wellas smallforests inland,fedonlyoccasionally by storm tides. Others have used the same system for already morelocal descriptions Florida:Odum (e.g. & Mclvor, 1990). We suggest thatthisframework can be used to define extremes the amongmangrove forests withinany regionwithouthaving to determine how fringe mangroveforests (for instance)in one part of the world are related to fringe mangroveforestsin another. The three extremesare easily described. Fringe mangrovesreceivethe bruntof the tides,which are often seawater. full-strength Prop roots,buttresses, and are pneumatophores commonamongtreesin thispart of a forest.Riverinemangrovesare floodedby river wateras well as by tides,so thatsalinity moderate. is Trees in this zone are likelyto be among the most productivein a forest(Twilley,Lugo & Pattersoncover large Zucca, 1986). Basin mangrovesgenerally areas behindfringe and riverine mangroves, and only occasionallydo tides inundatean entirebasin forest. Soil salinitymay be very high at higherelevations whereevapotranspiration causes salts to accumulate. In small forests that are frequently flooded,or where is rainfall highand/or groundwater flowis substantial, a basin mangroveforest can be of moderateor even low salinity (e.g. Cintr6n al., 1978; Semeniuk,1983; et Ewel et al., 1998a). Clearly,theremay be substantial differencesin hydrology, nutrient cycling, and productivity between these three types of forests (Twilley,1995). Productivity measured by (generally alone in these forests)is closely related to litterfall water turnover, with riverine>fringe >basin (Pool, Lugo & Snedaker,1975), because higherturnover is likelyto mean: (i) an increasedsupply of nutrients, of and silts, clays;(ii) lessaccumulation toxicsubstances in porewater;and (iii) greateraeration of the soil matrix. Distinct boundaries between the three hydroin geomorphic types mangrove of forest thesamestand to Some characteristics each, of maybe difficult define. such as inundationtimeand porewatersalinity, may vary fromregionto region,and thereis likelyto be no singlemeasurethat enables one kind of forestto be unambiguously assigned to one zone or another. these broadgroupings should Nevertheless, recognizing assist in formulating generalizationsthat may be particularly helpful in establishing appropriate management policies(e.g. Bacon, 1994).


In this paper, we consider a range of goods and services provided by mangrove forests (Table 1). Some, such as sediment trapping, nutrientprofood and habitatfor animals, cessing,and providing are essentialfor preserving ecosystemintegrity and regional biodiversity. Others are not obtained without considerable human intervention,such as harvesting mangrove trees.The magnitude and quality of each of these goods and services are likely to vary among the threehydrogeomorphic zones, and no zone can be designated as 'most' or 'least' overall. important

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Hammock Scrub Riverine

rngly ouIeling

Fri Frne

bidirectiona flux



Flooding Frequency from Tides

Flooding Frequency from River


of Salinity Floodwaters


tide-dominated, interior), and (river-dominated, forests typesof mangrove among threefunctional Fig. 1. (a) The relationships 1992). (b) describedfor the Neotropics(Woodroffe, dominantphysicalprocesses(in italics),and six typesof mangroveforests withinany given region typesof mangroveswamps that can be distinguished among threefunctional Proposed relationship (original).

Sediment trapping
Entiremangrove forests trapand retainsediments can in generated theuplandsby virtueof their positionin thelandscape(Lynchet al., 1989;Parkinson, DeLaune & White, 1994). Riverine forestsare likely to be in particularly important this respect,because river waters a usually sediment carry heavier load thanocean tides. Sedimentsdeposited in fringeforestscan be riverine origin,however, in having been recirculated within nearshore the waters (Wolanski, Mazda & Ridd,

1992). Basin swampsalso trapsediments, the receiving finest particles thatare carriedpast riverine fringe and forests by floods and tides. They may also trap sediments depositedby runoff fromuplands along the landwardedge of the swamp. intactavails society Simply leavingmangrove forests of the serviceof sediment because removal trapping, of the forest, particularly along the banks of fringe and riverine mangroveforests, opens up vulnerable soils to erosion and offshore sedimentdeposition. Mangroveforests also be exploited thisservice can for

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86 K. C Ewel, R. Twilley Jin R. and EongOng

forests providing in goods and services.1=most important of typesof mangrove Table 1. Relativeimportance different Role Trap sediments and Processnutrients organicmatter to waters Providea source of detritus nearshore and Serveas a sinkfornutrients carbon C, N P Improvewaterquality Providefood and habitatforanimals Provideaesthetically pleasingenvironments Protectshorelines Provideplantproducts Riverine 1 1 2 1 2 1 1 2 2 Basin 2 3 1 3 1 3 3 3 1 Fringe 3 2 3 2 3 2 2 1 3

when excess sedimentsgeneratedby anthropogenic activities such as road construction upland forest and clearingare prevented fromwashingout to offshore seagrass beds and coral reefs. Riverine forestsare mostimportant thisrespect, basin forests in but often performthis servicealso, as well as narrow fringe foreststhat directly abut against uplands (Nixon et to al., 1984). The abilityof mangroveforests receive sediments limited, is however, because treesare killed when lenticelson pneumatophores, prop roots, and young stemsare buried.The second largestcause of mangrove loss on the Pacific island of Pohnpei, FederatedStatesof Micronesia, overa 10-year period, was road construction, to both clearingthe road due trees bed and the death of adjacent and downstream fromexcess sedimentation (W. Raynor,The Nature Conservancy, pers. comm.). of a The prospect sea levelrisebrings newdimension It to the importance sediment of trapping. is possible of that thoughtful to manipulation sediment delivery a mangrove forest could ensurecontinued existence of the forest even in the face of rising tides. An understandingof the balance between sediment accretion and compaction as well as of potential vegetation and hydrodynamic changes due to associated climate change would also be required, however.

Ong, 1990; Wattayakorn, Wolanski& Kjerfve,1990). Measurementsof the net flux of different formsof nutrients and organic matterare required in order to understandthe role and relativeimportanceof mangrove forests biogeochemistry productivity to and of coastal waters(Boto & Wellington, 1988; RiveraMonroy et al., 1995). As observedfor salt marshes, into theremay be a net flux of inorganicnutrients mangrovesfromcoastal waters and a net exportof organic nutrientsassociated with particulate and dissolvedorganicmatter export(Twilleyet al., 1997). Even though there are still few studies of these between processes,highlighting important differences fringe, riverine, and basin mangrovesmay transcend muchof thiscomplexity.

matter Organic export

Estimatesof carbon exportto offshore watersrange across two ordersof magnitude;the average rate is about 210gC m-2 yr-1,with greatestexport values comingfromfringe mangroves (Table 2). In Rookery Bay, Florida, for instance,fringemangroveforests exporttwice as much organicmatterper hectareas basin forests(but taking the area of each type of in wetlandinto account results equal loading ratesof between of detritus them;Twilley, 1982). The presence crabs thatconsumelitter (28-79% of annual leaf fall; Robertson, 1986, 1988; Robertson & Daniel, 1989; Twilley et al., 1997) may increase the tightness of nutrient cycles in the mangroveforestand decrease the amount of outwelling, particularly frominterior and especiallyin Old World portionsof basin forests sites, where crabs are more common (Jones, 1984). High productivity relatively and shortresidence times

Processing of organic matter and nutrients

The exchanges of nutrientsand detritusbetween mangroves and near-shore watersin areas of different geomorphology and hydrodynamics are widely but complex and difficult accepted as important to quantify (Boto & Bunt, 1981; Twilley, 1985; Gong &

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et (afterTwilley al., 1992). Table 2. Exportof organiccarbon frommangroveforests Swamp type Export (gC m-2 yr-1) 2 64 91 110 186 292 340 401 420 210 Location Reference

Fringe Basin Basin Swamp* Fringe Fringe Swamp* Fringe Swamp* Average

Hong Kong Florida, USA Florida, USA New Zealand Florida, USA Florida, USA Australia Hinchinbrook, PuertoRica, USA Australia Hinchinbrook,

Lee (1989) Twilley(1985) Lugo & Snedaker(1974) (1985) Woodroffe Heald (1969) Odum & Heald (1972) Robertson(1986) Golley,Odum & Wilson(1962) Boto & Bunt (1981)

* Because fluxmeasurements both fringe the weremade at themouthof a tidalcreek,exchangerepresents combinedexportfrom the swamp. and basin mangroves and therefore entire

of litter in riverine and fringemangroves, both of make associatedwithhigher frequency inundation, exceptwherebasin zones themparticularly important, are muchlarger.

improvement Waterquality

The service of nutrientprocessinghas often been Mangrove exploitedfor water quality improvement. forests, like wetlands in general around the world, receive untreatedwastewaterof often inadvertently Nutrient sink Basin mangrove forests bothhumanand animalorigin. nutrients, usefulfortransforming may be particularly Basin mangroveforestsmay rank lower in organic particularly N, and immobilizing microbes and but frequency, matter exportbecause oflowerflooding chemicalssuch as pesticides (Clough,Boto & Attiwill, they may have higherrates of organic matterand removednitrate from effectively 1983). Denitrification nutrientaccumulation. Soil redox potential is a measureof thesoil environment a heavy wastewaterload in a Puerto Rican basin particularly important mangroveforest,for instance (Corredor & Morell, of characteristics nutrientcycles for determining of 1994). However,the full implications disposal of because of its controloveroxidationstatesof P, N, S, wastewater into mangrove forests are seldom It Mn, Fe, and manyotherelements. shouldbe lowest humanhealth, effluent should To considered. safeguard wherehydroperiods longest, are althoughthepresence in be retained an area freefromcontactwithhumans of pneumatophores and prop roots (McKee, i.e. or commonly eaten fishand shellfish, as farback Mendelssohn Hester,1988)and theabsenceofstrong & in a basin forest as possible. Although increased in ratesbetweenthe zones differences waterturnover was once consideredas productivity an ecosystem of this.Basin mangrove (Ewel et al., 1998a) maymitigate of the a usefulby-product this kind of enrichment, forests morelikelyto serveas a sinkforinorganic are changes in a habitat that can follow such dramatic in N because of the likelihoodof denitrification an but anaerobicenvironment, in factbothdenitrification changes in ecosystem function,both on-site and harmful are downstream, now regardedas potentially rates in mangroveswamps are and nitrogen fixation (Ewel, 1997). Nevertheless,when the expense of slow (e.g. Rivera-Monroy & Twilley, 1996). constructing a wastewater treatment plant is can be significant, even in a fringe Denitrification considered,wetlands are still commonlyselected as however mangroveforest, under sewage enrichment, receiving areas foreffluent. & of (Corredor Morell,1994).Formation H2S and CH4 Retrofittingmangrove forests for wastewater should be particularly common in basin mangrove treatment preferable destruction, wastewater is to and forests,but oxygen transportvia prop roots (e.g. and McKee etal., 1988)mayslowtheformation release as disposalmayevenbe considered a tool in mangrove of thesegases. treatedwastewater restoration. Directingsecondarily
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and EongOng R. 88 K. C Ewe!, R. Twilley Jin

through a complex of coastal ecosystems on a Caribbean island resulted in both 'mangrove rejuvenation'and increaseduse by water birds and care (Bacon & Morgan,1996). Although otherwildlife must stillbe taken to meet acceptable public health the wetland'modelused widely standards, 'constructed for now in the USA (Ewel, 1997) may be appropriate coastal wetlandsin manytropicalcountries. portions of basin mangroves(Robertson & Blaber, data indicate that shrimpobtain 1992). Conflicting carbon fromplankton and possibly epiphyticalgae ratherthan frommangroves(Primavera,1996) and that mangrove-derived carbon fuels bacteria trophic foundin higher thatis subsequently production et levels(Cifuentes al., 1996).It is possiblethattheroles in productivity detritus insituprimary and ofmangrove highertrophiclevelsvaryamong different supporting typesof estuary. once grew The use of land wheremangroveforests to cultureshrimp(prawns) has a long traditionin farming expandedmore and shrimp Asia. Mariculture into Ecuador, where,by the late 1980s,there recently were more shrimpproduced than anywhereelse in the world (Twilley,Bodero & Robadue, 1993). The of intensity use ranges fromtrappingshrimpwithin untilharvest, forests holdingthemthere and mangrove where systems to usuallyfora lunarcycle, moreintense as mangrovetreescoexistwithpond and shrimp, in the traditional'tambak' of Indonesia. Much more intensiveculture techniques have been introduced recently, primarily in basin mangroves, where mangrovetreesare totallyclearedand ponds are dug to a depth of 1-2 m. Much of this startedin the of (Chanos initially theculture milkfish for Philippines, dependedon thenatural chanosForsskal.).The system of fertility the ponds,whichdecreasedin productivity and the and very quickly, required clearing construction of aquaculture.Most ofnewponds in a system shifting in had of themangroves thePhilippines been degraded culture Fabricius)took whenshrimp (Penaeusmonodon forests hold. By theearly1990s,50% of themangrove in Thailand (Spalding,Blasco & Field, 1997) and 21% in of the mangroves Ecuador (CLIRSEN, 1992) had of ponds. beenlost,mostly due to construction shrimp 24% of shrimpponds in Overall, approximately Thailand have been abandoned because of diseases (which have also affected ponds in India, the Philippines,Taiwan, and Indonesia) (Stevenson & Burbridge, 1997). Also, many mangrove soils, in particularly the Old World tropics,are potentially acid sulphate soils, and sulphuric acid will form waterpH to around 3) if the soil is oxidized (reducing beginsafterthe ponds are (as whenwatercirculation built). This can be overcome by heavy limingand extensiveflushing (which is not only expensivebut reduces water quality in the neighbouring estuary). are beingusedbecausemangrove Mangroveforests still land is grosslyundervaluedand oftenbelongsto the need to establish real state.Thereis thusan urgent the

Animal habitat
Mangrove forestssupport animal populations both areas. Some of these and in offshore within forest the life animalsspendonlypartof their cyclein mangrove forests,either during a particular stage of their et maturationor as migrants(Yainez-Arancibia al., 1988; Yainez-Arancibia,Lara-Dominguez & Day, 1993). Where mangrovesare a dominantsource of carbon, they are importantto estuarineconsumers (Rodelli et al., 1984). Crabs are among the fauna of mangroveforests characteristic invertebrate in important humanfood chains. thatare particularly on likely be highest to Densitiesof crabs are especially with mudbanks adjacent to mangroves, unvegetated theirdensities mangroves; along fringe lowernumbers however, perhaps areoften inside low forests, mangrove because of low food quality as a result of high of 1992). concentrations tannins (Alongi& Sasekumar, There, they are more common at high intertidal locationsthanat lowerzones,perhapsbecause feeding timesare longer(Mclvor & Smith,1995). as Some speciesof crabs processdetritus, described earlier,and some feed on propagules (and thereby forest structure; Smith,1987). They are affect directly in processes ecosystem clearlyimportant maintaining in mangroveforests, and many provide food, both and to mangrove directly indirectly, people. Culturing crabs (specifically Scylla serrata (Forsk.)) has been proposed, but this would have to be done without waterflowsin orderto the or clearing forest impeding maintainsuitablehabitat.Crab productionmightbe food supply,such as with increasedby augmenting kitchen scraps, but because mangrove crabs are territorial can be cannibalistic J.SmithIII, US and (T. BiologicalResourcesDivision,pers. GeologicalSurvey, comm.), this is not likely to be sustainable on a commercial scale. Juvenileshrimp (e.g. penaeid prawns) are also fauna. importantcomponents of the invertebrate Shrimp are more common in fringeand riverine inundated mangrovesas well as in more frequently

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value of mangroveland not only in economic terms but also in termsof its ecological and sociological value. There is no clear explanationforthe highdensities in offish thathavebeenreported and aroundmangrove forests, although they are often associated with prop roots forests, wherewell-developed Rhizophora may provide safetyfrom predation (Robertson & Blaber, 1992). Habitat availabilityfor juvenile and in and riverine adult fishis likelyto be greater fringe as forests, well as in those portionsof basin forests wheretidal channelsprovideaccess. forests. also vertebrates use mangrove Many higher Few amphibiansare found in mangroves(although, for example, ranid frogsoccur in mangrovesin SE reptiles, including but Asia and thePhilippines), several crocodiles,snakes, and lizards,use mangroveforests & (Macnae, 1968; Hutchings Recher,1983). Some sea rootsand leaves and appear turtles feedon mangrove areas (Odum, to use mangroveestuariesas nursery Mclvor & Smith,1982). for Large numbersof birdsuse mangroves feeding in and/or nesting manypartsoftheworld.For example, but fromFlorida mangroves, 181 speciesare reported alone. on veryfeware heavilydependent theseforests Those most closelyassociated withmangroveforests in common feedprimarily thecanopyand areespecially in fringe riverine and forests (Odum etal., 1982).Fringe and riverine mangrovesare especiallyimportantto birds(Ogden, 1994). More than200 species migrating of birds have also been reportedfrom Australian including someendemicspecies(e.g. forests, mangrove Butorides striatus themangrove (L.)) (Hutchings heron, & Recher,1983). most Many mammals also use mangroveforests, or of them in addition to other terrestrial aquatic There are a fewendemicmammalspecies ecosystems. rat myoides e.g. in-mangroves, a crab-eating (Xeromys & Thomas) in Australia(Hutchings Recher,1983),the in cristatus (Raffles)) Malaysia, leafmonkey (Presbytis and theproboscismonkey (Wurmb.)) (Nasalis larvatus in Borneo(Macnae, 1968). Mangrovesare particularly to important a fewlargemammals,such as largecats Pantheratigris Linnaeus, (e.g. the Royal Bengal tiger, in theSundarbans, India and Bangladesh),as between wellas a variety deerand otters of (Odum et al., 1982; Dugan, 1993; FAO, 1994). along stately particularly Boardwalksand canoe trails, mangrove riverine corridorsand the edges of fringe for forests, providean opportunity obtainingincome from otherwiseundeveloped land as well as for and values of educatinglaypeople on the functions wetlands. mangroves, Healthy stands of fringeand riverine In are however, not alwaysappreciated. the USA, the thatpermits passed legislation StateofFloridarecently to homeowners prunemangrovesalong the shoreline view,in spiteof thefactthat to allow an unobstructed mangle one of the most common species,Rhizophora after cutting to the L., does notretain ability regenerate (Gill & Tomlinson,1969; Snedakeret al., 1992). The 'Ding' Darling National Wildlife Refuge, also in Florida, cuts 'windows' in the stands of mangroves that grow up along dikes to allow easier viewingof alligatorsand wading birdsthat wildlife, particularly use the ditcheson the otherside.

Protection fromfloods
surges, from of Protection humaninfrastructure storm touted tidalwaves,and floodsis one ofthemostwidely services providedby wetlands.This is because (1) the of boundariesof a wetlandindicatetheextent normal flooding and thereforethe zone where human shouldcease, and (2) wetlandvegetation development decreases the rate at which water passes over land, as force of floodwaters they slowingthe destructive approach the uplands. Like many other ecosystem it in services, spiteof the generalrecognition receives, (or the value of shoreline protection flood protection in general)providedby a wetlandlies in the cost that societydoes not have to pay untilthat serviceis lost (Goulder & Kennedy,1997). Mangroves develop only along low-energyor protectedcoasts, where sedimentsare retained and mangrove seedlings can become established. Only eventssuch as major stormsurges wherehighenergy forests likely and tidalwavesare episodicare mangrove In in human structures. to be important protecting are suchplaces,fringe forests believedto be mangrove but hypothesis important, thisis a difficult particularly to prove, and anecdotal evidence is not always convincing (Clough, 1993). Perhaps the most widely cited observationis Fosberg's (1971) suggestion that the loss of more than 100,000lives in Bangladeshin and tidalwave might have 1970 following hurricane a been reducedhad large areas of mangroves not been replaced by rice paddies. Other anecdotes are

Aestheticallypleasing environment
stands of mangroveforestsare Large, undisturbed attractive tourists(Hamilton & Snedaker, 1984). to

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Erosion along a coastal road behind accumulating. mangroveson St. John,US VirginIslands, aftera in hurricane fall 1995 was least wherethe mangroves (T.J. Smith III, pers. comm.). Loss of were thickest of on following diversion a river thePacific mangroves island of Kosrae, FederatedStatesof Micronesia,has with increased wearon thecoastalroad beencorrelated by and nearbystructures stormtides(K.C. Ewel and pers. obs.). Instancesin which storm R. R. Twilley, have affected on vegetation thelandwardsideof surges unscathed remained whilemangrove mangrove forests (e.g. Steers, 1977) do not prove that the disturbed had the further destruction area would have suffered mangrovesnot been there.In the absence of more it data, however, seemsapparentthatzoning definitive to limit, that a regulations protect wetland itslandward of should as upperlimit flooding, recognized thenormal enable it in turnto protecthomes,roads, and other infrastructure flooddamage. from Carolinesrecognize PacificIslandersin the western drooping the importance of the characteristically forests for mangrove along theedge of fringe branches breakingthe force of stormtides (K.C. Ewel, pers. obs.). The roots of mangrovetrees in both fringe role in binding forests and riverine play an important and soilstightly Scoffin, 1970).Bothfringe (e.g. surface important are particularly mangroves therefore riverine not shorelinesfromeroding,thereby for preventing shoreline protectionbut protecting only affording offshore seagrassbeds and coral reefsfromsediment depositionas well. from mayassistin protection Basinmangrove forests watervelocity episodicfloodsas well,bothby reducing and by adding flood storagecapacity behind fringe The amount of forestnecessary and riverine forests. in for adequate protection a particulararea depends of on the geomorphology an individualshorelineas and magnitudeof possible well as on the frequency storms. Plant products An impressive of diversity plantproductsis harvested from mangrove trees, including tannins, honey, medicinal and (Hamilton& Snedaker, products, thatch impact 1984). Some of thesecan be obtainedwithlittle on the forest;harvesting for firewoodand timber effect. probablyhas the greatest Net primary as of productivity mangroveforests, to measuredby litterfall, oftenhighrelative upland is at forests thesamelatitude 1993), (Saenger& Snedaker, fromsite to but tree growthrates vary considerably withlatitude(Twilleyet al., decreasing site,generally appear to be mostproductive 1996). Mangroveforests where thereis no distinctdry season, and biomass can exceed 250 Mg/ha (Ong et al., 1979), reaching 350-400Mg/hain protectedareas of Malaysia (Putz of & Chan, 1986). The best growth mangrovetreesis where forests, believedto occur in riverine generally (e.g. Lacerda et floodsdepositsediments periodically is al., 1993; Hussain, 1995),but harvesting oftenmost common in basin forestswhere large monospecific flooding maymake extraction standsand less frequent moreeconomical. have different timber speciesof mangrove Different physical properties(FAO, 1994) but there is no on information how variations within and among thataffect forces speciesmaybe relatedto thephysical Some species,likeAvicennia, kindsof forests. different are widespreadecologically;they are very soft and whichis also have littlecommercialuse. Rhizophora, environments often and of foundin a variety different in extensive, stands, has many almost monospecific commercial genus. exploited usesand is themostwidely The wood is veryhard and dense but tends to split but is very readily.It is not suitable for furniture and charcoal and is ideal for oftenused forfirewood conversionto rayon. It is also oftenused for poles resistantto rot under and, because it is extremely material. suitableas piling anaerobicconditions, very is A number less widelyspreadgeneralikeXylocarpus of in and Heritieraare foundprimarily basin or riverine grade mangroveforests;they are excellentfurniture timberbut are seldom exploited for that purpose tendto occurin low numbers. Xylocarpus because they in is used extensively Micronesiaforcarving. In most of the tropics, mangrove timber has been used mainly as fuelwood (either traditionally as to or directly firewood afterconversion charcoal), as fishing stakes, and as buildingmaterials(pilings, poles, and timberfor buildings and boats). Here, mangroveuse is not a matterof choice but ratherof and of survival, intensity use dependson ease of access and populationpressures. in Commercial productionis common primarily for are largestandsofRhizophora harvested Asia,where pilings),charcoal and, more recently, poles (including intorayon).Wherethenet woodchips(forconversion of productivity mangrovesis high, it is possible to timber a sustainable harvest on basis. A simpleruleof thumb clearfelling around smallpatchesand thinning 15 years and again around 20 years was developed

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in in theearlypartof thiscentury Malaysia by British foresters (Watson, 1928). The classic success case is in the Matang Mangrove Forest in Malaysia where Rhizophora apiculata BL. has been harvested sustainably (annual coupe of about 1000ha) sincethe earlypartof thiscentury (Ong, 1995). Here thetimber is used as poles forpilingas wellas fortheproduction of charcoal (for export). Yet, in the same country, pristine mangroves(e.g. the mangroveforestof the Rajang Delta in Sarawak) have been badly degraded as a resultof harvesting woodchips(annual coupe for of about 600 ha). This operation startedin the late timber after about 25 1960sand ran out of mangrove was years.A minimum girthsystemof management practiced but appears to have failed. A similar State of Sabah was operation in the neighbouring closed down earlier. is The mangrove woodchipindustry now veryactive in some of the best mangrove forests Kalimantan, (in it Sumatraand IrianJaya)in Indonesia,where operates almost unnoticedin remote,sparselypopulated, or unpopulated areas, where some of the world's best pristine mangrove forests (Rhizophora-dominated) last generally for have been targeted. These operations about 25 yearsand thenare moved to anothersite.It is not known if the Indonesian operations can be sustainedas theyare presently run, but a sustainedhas not been implemented yield management system elsewhere (Ong, 1995). parts of a forestmay help in formulating different supply policiesthatenable thecontinued management a of essentialgoods and services.For instance, basin forestfromwhichcrabs are harvestedshould not be and timberharvest used for wastewatertreatment, in and or shouldbe restricted evenprohibited riverine whichare more subjectto erosionthan fringe forests, basin forests. All three kinds of forests can be in pleasingand are important production aesthetically althoughin very processing, of animals and nutrient different ways. Exploitinga mangroveforestfor one product or Timber can reduceitsability provideothers. to service and ponds waterqualityimprovement, shrimp harvest, in wheretheyare are all concentrated basin forests, likely to reduce a forest'spotentialfor creatingan change the nature aesthetically pleasingenvironment, of its nutrient cycles, and alter habitat for animal it withregional planning should However, populations. goods and whereconflicting be possible to determine zones within services could be obtainedfromdifferent for the same forest, perhapsas mitigation preserving a similarforestin an area where habitat value or For instance, some recreation benefits may be higher. of the concernabout unregulated growthof shrimp ponds centresaround the loss of water quality in effluent Distributing coastal ecosystems. neighbouring forests basinmangrove pondsintonearby from shrimp would not onlylimit impactson waterquality negative in coastal ecosystems (Robertson& Phillips,1995),but it could also increase mangrovetree growthrates, in of intensity harvesting morepristine reducing thereby stands. all We do not yet understand the consequencesof For instance,does to disturbances mangroveforests. removal of large volumes of timber from basin compaction dead as forests lead to sediment mangrove roots decay,in turnleadingto lowerredox levelsand changesin habitatvalue forboth plantsand animals? Lower redox but no apparentsubsequentchangesin in speciescomposition small basin-zonegaps in highforests have beenrecorded rainfall (Ewel et al., 1998b), and these effects mightbe accentuatedin more arid release Lower redoxvaluescould also increase regi-ons. of N,, H2S, and perhapsCH4 to theatmosphere the to extent thattheyaffect regionalor even global climate. However,we cannot yet defineacceptablelimitsthat can be used to formulatemanagementpolicies in different partsof theworld. the that Acknowledging understanding diversity and exists within forests shouldprovide broader a mangrove



can be attributed The usefulness mangroveforests of and the diversity to both the diversity among forests of goods and services thattheysupply.A synopsisof the relativeimportance fringe, of and basin riverine, mangrove forestsindicates that all three may be consideredcriticalin at least one respect(Table 1). Riverine forests,with their nutrientinflows and moderatesalinitiesare important interfaces between themoreexpansive basin forests thefresh and and salt waterinflows. Basin forests, the otherhand, with on theirmore restricted water flows,are oftenthe sites of greatest human activity. Fringemangroveforests, to water movement consistently and subject thegreatest barrier the for highsalinity, critical a protective are as rest of the forest and, occasionally, for human as structures well. Understandingthe importanceand best use of

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92 K. C. Ewe/, R. Twilley in EongOng R. and

of perspective judgingall theimplications theuses for of this important formof wetland.Aggregating this diversityaround simplifying concepts such as the distinctions amongfringe, riverine, basin zones in and mangroveforestsmay assist local natural resource managersin restricting uses of thesewetlandsto the are to kindofforest where they mostlikely be tolerated and even sustained. Boundaries among these zones may be difficult define, to and particular zones in one area maynot be directly comparableto the same zone in anotherarea. A conservative approach,untilmore definitive guidelines be obtained,may be to define can Best Management Practices that limit potentially destructive management practices, such as harvesting, 50 withinthe first or 100m (or some multipleof the normal tidal range) of a fringeor riverineforest. Being able to restrict development means of easily by understoodguidelines may be the first step not only toward reducingthe loss of mangroveforestsin a regionbut also to reducingthe rate of loss of their goods and services. elementalvariationsof carbon and nitrogen a in mangroveestuary.EstuarineCoast Shelf Sci. 43, 781-800. G. Cintr6n, Lugo, A.E., Pool, D.J. & Morris, (1978) G., Mangroves arid environments PuertoRico and of in adjacent islands. Biotropica, 110-121. 10, Cintr6n, Lugo,A.E.:& Martinez, (1986) Structural G., R. The andfunctional properties mangrove of forest. botany e and natural history Panama: La botanica historia of naturalde Panama (ed. by W.G. D'Arcy,and M.D. Correa),pp. 53-66. Monogr.Syst.Bot. 10, Missouri Bot. Gardens, Louis MO, USA. St. Clarke, L.D. & Hannon, N.J.(1971)Themangrove swamp and salt marshcommunities the Sydneydistrict. of of J. IV. The significance speciesinteraction. Ecol. 59, 535-553. de CLIRSEN (1992) Estudiomultitemporal manglares, de camaroneras areas salinas,mediante y informaci6n sensores remotos. Centrode levantamientos integrados naturales sensores remotos. Memoria de recursos por Tecnica. Quito,Ecuador. Clough,B.F. (1993) The statusand value of mangrove in forests Indonesia, Malaysiaand Thailand: Summary. The economic and environmental values of mangrove and state in Southforests their present ofconservationthe eastAsialPacific Region byB. F. Clough)pp. 1-10. (ed. ITTO/ISME/JIAM Project PD71/89 Rev. 1. ACKNOWLEDGMENTS Clough, B.F., Boto, K.G. & Attiwill,P.M. (1983) Tasksfor Mangrovesand sewage: a re-evaluation. Dr 8 science (ed. byH. J.Teas),pp. 151-161. vegetation We thankJamesA. Allen,Judy Drexler,and three Z. W. Junk, Hague. The reviewsof the for thoughtful anonymousreviewers J.E. Corredor, & Morell,J.M.(1994) Nitrate depuration manuscript. in sediments. of secondary sewageeffluents mangrove Estuaries, 295-300. 17, Dugan,P. (ed) (1993) Wetlands danger, in 187pp.Oxford REFERENCES New York. University Press, Ewel,K.C. (1997)Water quality improvement wetlands. by Island services byG. Daily),pp. 329-344. Ecosystem (ed. Alongi, D.M. & Sasekumar, A. (1992) Benthic DC. Press, Washington communities. Tropicalmangrove ecosystems:coastal and Ewel, K.C., Bourgeois, J.A., Cole, T.G. & Zheng, S. estuarine studies 41 (ed by A.I. Robertson D.M. & in characteristics and (1998a) Variation environmental Alongi) pp. 137-171.AmericanGeophysical Union, in in vegetation high-rainfall mangrove swamps Kosrae, DC. Washington Micronesia. GlobalEcol. Biogeogr. Letts, 49-56. 7, Bacon,P.R. (1994)Template evaluation impacts for of of J. Ewel, K.C., Zheng S., Pinz6n,Z. & Bourgeois, A. sea levelriseon Caribbean coastalwetlands. Ecol. Eng. (1998b)Environmental effects canopygap formation of 3, 171-186. in high-rainfall in mangrove swamps. Biotropica, press. Bacon,P.R. & Morgan, A.C. (1996) Integrating wetland 319 FAO (1994) Mangrove forest management guidelines, restoration wastewater with in treatment a Caribbean pp. Rome. V coastal environment(abstract). INTECOL's Fosberg, F.R. (1971) Mangroves versus tidalwaves.Biol. International Wetlands Conference, Perth, Western Conserv. 38-39. 4, Australia. P.B. on Boto,K.G. & Bunt, (1981)Tidalexport particulate Gill,A.M. & Tomlinson, (1969)Studies thegrowth J.S. of of redmangrove matter from northern a (Rhizophora mangle I. Habit and L.). organic Australian mangrove system. Estuarine Coast Shelf Sci. 13,247-255. general morphology. Biotropica, 1-9. 1, Boto,K.G. & Wellington, (1988) Seasonalvariations Golley,F.B., Odum, H.T. & Wilson,R.F. (1962) The J.T. in concentrations fluxes dissolved and of structure and metabolism a Puerto Rican red organic and of in inorganic materials a tropical,tidally-dominated, mangrove in forest May.Ecology, 9-19. 43, Mar. mangrove waterway. Ecol. Progr.Ser. 50, 151-160. W.K. & Ong,J.E.(1990)Plantbiomass nutrient and Gong, Cifuentes, L.A., Coffin, R.B., Solarzano,L., Cardenas, flux in a managed mangroveforestin Malaysia. W., Espinosa,J. & Twilley, R.R. (1996) Isotopicand Estuarine CoastShelf Sci. 31, 519-530.
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