10.07.2015 Views

Plant Species Diversity in Tropical Planted Forests and Implication ...

Plant Species Diversity in Tropical Planted Forests and Implication ...

Plant Species Diversity in Tropical Planted Forests and Implication ...

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

BNM/RH/CIR 010-9Consumer <strong>and</strong> MarketConduct DepartmentMarket Value of Motor VehiclesPage1/21. INTRODUCTION1.1 The current market practice of <strong>in</strong>sur<strong>in</strong>g motor vehicles without reference to anyvehicle valuation database has not only given rise to over-<strong>in</strong>surance or under<strong>in</strong>suranceby consumers but also disputes on market value of vehicle at the po<strong>in</strong>tof loss. The subjectivity <strong>in</strong> determ<strong>in</strong><strong>in</strong>g the market value of vehicle couldpotentially lead to abusive practices to the detriment of consumers.1.2 With the availability of <strong>in</strong>dustry motor vehicle valuation databases, <strong>in</strong>surers /takaful operators are now able to determ<strong>in</strong>e the market value of most of themotor vehicles at the po<strong>in</strong>t of sale as well as at the po<strong>in</strong>t of loss.1.3 The objective of the circular is to ensure consumers are properly advised on theappropriate market value of motor vehicle to address over-<strong>in</strong>surance or under<strong>in</strong>surance.In addition, consumers will also be assured of objective determ<strong>in</strong>ationof market value of vehicle at the po<strong>in</strong>t of loss.2. APPLICABILITY2.1 The circular is applicable to general <strong>in</strong>surers licensed under the Insurance Act1996 <strong>and</strong> takaful operators registered under the Takaful Act 1984, <strong>and</strong> their<strong>in</strong>termediaries.3. LEGAL PROVISIONS3.1 The circular is issued pursuant to section 201 of the Insurance Act 1996 <strong>and</strong>section 69 of the Takaful Act 1984.4. EFFECTIVE DATE4.1 The circular is effective from 1 August 2011.


<strong>Plant</strong> <strong>Species</strong> <strong>Diversity</strong> <strong>in</strong> <strong>Tropical</strong> <strong>Plant</strong>ed <strong>Forests</strong>Methods of studyA sample plot was set up <strong>in</strong> each of the 6 forest st<strong>and</strong>s<strong>in</strong> May 1998 (12 years after the plantation of the st<strong>and</strong>s) <strong>and</strong><strong>in</strong> the natural forest (Table 1). All the trees <strong>in</strong> the plantedforests <strong>and</strong> trees with a DBH (diameter at breast height) of4.5 cm or more <strong>in</strong> the natural forest were measured for DBH<strong>and</strong> height <strong>in</strong> May 1998 <strong>and</strong> November 1998. Sample treeswere cut down <strong>in</strong> the planted forests, <strong>and</strong> tree dimensions<strong>and</strong> weight of the leaves, branches <strong>and</strong> stems were measured<strong>in</strong> November 1998 to determ<strong>in</strong>e the biomass of the st<strong>and</strong>s.To assess the understorey floral composition, we recordedthe names of all the vascular plants established <strong>in</strong> all the 7plots. We also identified vascular plant species outside thesample plots with<strong>in</strong> each st<strong>and</strong> to determ<strong>in</strong>e the suitability ofthe size of each sample plot to estimate the species richness<strong>in</strong> each st<strong>and</strong>. To measure the biomass of understorey plants<strong>in</strong> the planted forests, we set up 12 quadrats, 1 m 2 <strong>in</strong> size,outside of the sample plot <strong>in</strong> each st<strong>and</strong>. All the plantsgrow<strong>in</strong>g <strong>in</strong> the quadrat were identified <strong>and</strong> cut, <strong>and</strong> theaboveground dry weight was determ<strong>in</strong>ed separately for thetrees, shrubs, herbs <strong>and</strong> v<strong>in</strong>es.In the I. cyl<strong>in</strong>drica <strong>and</strong> S. spontaneum grassl<strong>and</strong>s,sample plots, 100 <strong>and</strong> 200 m 2 <strong>in</strong> size, respectively, were setup <strong>in</strong> November 1998. All the vascular plants were identified<strong>in</strong>side <strong>and</strong> outside of the sample plots. To estimate thebiomass of the grassl<strong>and</strong>s, we set up 6 <strong>and</strong> 8 quadrats,respectively, 1 m 2 <strong>in</strong> size, <strong>in</strong> the sample plots. All the vascularplants <strong>in</strong> each quadrat were identified <strong>and</strong> cut, <strong>and</strong>the dry weight of the trees, shrubs, herbs <strong>and</strong> v<strong>in</strong>es wasmeasured.Relative light <strong>in</strong>tensity was measured with M<strong>in</strong>oltadigital photometers <strong>in</strong> the forest sample plots <strong>in</strong> November1998.Fig. 1. Relationship between leaf biomass <strong>and</strong> relative light<strong>in</strong>tensityResultsSt<strong>and</strong> descriptionThe growth of the st<strong>and</strong>s varied considerably amongthe planted forests. The aboveground biomass rangedfrom 28 to 196 t ha –1 , be<strong>in</strong>g larger <strong>in</strong> the exotic speciesst<strong>and</strong>s than <strong>in</strong> the <strong>in</strong>digenous species st<strong>and</strong>s. The largestbiomass was recorded <strong>in</strong> the A. mangium st<strong>and</strong>, which ischaracterized by a high biomass production <strong>in</strong> tropicalplantations 10–12, 26 . The smallest biomass was recorded <strong>in</strong>the P. macrocarpus st<strong>and</strong>. This small biomass was associatedwith the widest tree spac<strong>in</strong>g <strong>and</strong> a smaller biomass<strong>in</strong>crement than that of the other species. The leaf biomassshowed a narrower range among the st<strong>and</strong>s than theaboveground biomass, be<strong>in</strong>g 2.7 to 5.9 t ha –1 , except forthe P. macrocarpus st<strong>and</strong>, whose canopy was not completelyclosed. The leaf biomass <strong>in</strong> these st<strong>and</strong>s wasalmost the same as that <strong>in</strong> various fast-grow<strong>in</strong>g plantations<strong>in</strong> the tropics 10–12,26,30 , which have a generally lowleaf biomass among the various forests 29 .The relative light <strong>in</strong>tensity varied among the studyst<strong>and</strong>s from 2.6 to 27.6% under closed canopy <strong>and</strong>amounted to 30.6% <strong>in</strong> the P. macrocarpus st<strong>and</strong>. Leafbiomass showed a significant relation to the relative light<strong>in</strong>tensity (Fig. 1). The relative light <strong>in</strong>tensity <strong>in</strong> tropicalforests with a larger leaf biomass (7.4 to 9.1 t ha –1 ) thanthat <strong>in</strong> our study st<strong>and</strong>s ranged from 0.5 to 1.9% 16,20 .Thus, leaf biomass was considered to be an appropriate<strong>in</strong>dicator of light <strong>in</strong>tensity under the canopy among differentforests.Understorey species richnessFig. 2 shows the understorey species richness <strong>in</strong>each sample plot. The number of understorey plant species<strong>in</strong> the sample plots reflected well the composition <strong>in</strong>the planted forests, natural forest <strong>and</strong> grassl<strong>and</strong>s with aclose positive correlation (R = 0.93, p < 0.01). The numberof understorey plant species <strong>in</strong> the planted forests waslarger than that <strong>in</strong> the grassl<strong>and</strong>s <strong>and</strong> smaller than that <strong>in</strong>the natural forest, account<strong>in</strong>g for 31 to 52% of the understoreyspecies <strong>in</strong> the natural forest.The number of understorey plant species <strong>in</strong> the sampleplots <strong>in</strong> the planted forests ranged from 35 to 59; therewas no difference between the exotic <strong>and</strong> <strong>in</strong>digenousst<strong>and</strong>s. The S. spontaneum grassl<strong>and</strong> exhibited a slightlygreater species richness than the I. cyl<strong>in</strong>drica grassl<strong>and</strong>.Trees <strong>and</strong> v<strong>in</strong>es comprised a large proportion (29 to40% <strong>and</strong> 34 to 40%, respectively) of the total plant species.The number of understorey tree species found <strong>in</strong> theplanted forests was 1.5 to 2.8 times as large as that <strong>in</strong> theS. spontaneum grassl<strong>and</strong> <strong>and</strong> 4.0 to 7.3 times as large as113


K. Kamo et al.Fig. 2. <strong>Species</strong> richness of understorey plants <strong>in</strong> the study plotsOw<strong>in</strong>g to differences <strong>in</strong> plot sizes, the numbers of plant species <strong>in</strong> the plots except for A. auriculiformis <strong>and</strong> E.camaldulensis were adjusted with a species-area curve for each plot to the plot size of planted forests (575 m 2 ) .that <strong>in</strong> the I. cyl<strong>in</strong>drica grassl<strong>and</strong>, or 26 to 48% of that <strong>in</strong>the natural forest.No seedl<strong>in</strong>gs of the plantation species had becomeestablished <strong>in</strong> the planted forests, except for Leucaenaleucocephala.Similarity <strong>in</strong>dexesSimilarity <strong>in</strong>dexes (Nomura-Simpson’s coefficient 14 )for all the understorey vascular plant species between thenatural forest, the planted forests <strong>and</strong> the 2 grassl<strong>and</strong>s areshown <strong>in</strong> Table 2. The degree of similarity was notappreciably high between the plots. The degree of similaritywas also not very high among the tree species (Fig.3).The mean similarity <strong>in</strong>dex of the understorey treespecies was calculated for all the tree species found <strong>in</strong> theplanted forests, <strong>and</strong> large tree species (DBH ≥ 10 cm) <strong>and</strong>small tree species (DBH < 10 cm) <strong>in</strong> the sample plot ofthe natural forest (Fig. 3). The similarity <strong>in</strong>dex for largetree species <strong>in</strong> the natural forest was significantly higherFig. 3. Mean similarity <strong>in</strong>dex between the planted forestsfor understorey tree speciesTable 2. Similarity <strong>in</strong>dex between the study plots for all the plant speciesSite Aa Am Ec Dc Xk Pm Ic SsAm 0.54Ec 0.58 0.41Dc 0.44 0.58 0.48Xk 0.55 0.49 0.44 0.59Pm 0.46 0.51 0.45 0.57 0.52Ic 0.71 0.54 0.63 0.67 0.58 0.50Ss 0.53 0.56 0.35 0.65 0.53 0.47 0.38DEF 0.42 0.45 0.42 0.57 0.50 0.46 0.54 0.56Aa: A. auriculiformis st<strong>and</strong>, Am: A. mangium st<strong>and</strong>, Ec: E. camaldulensis st<strong>and</strong>, Dc: D. coch<strong>in</strong>ch<strong>in</strong>ensis st<strong>and</strong>, Xk: X. kerrii st<strong>and</strong>,Pm: P. macrocarpus st<strong>and</strong>, Ic: I. cyl<strong>in</strong>drica grassl<strong>and</strong>, Ss: S. spontaneum grassl<strong>and</strong>, DEF: dry evergreen forest.114 JARQ 36 (2) 2002


<strong>Plant</strong> <strong>Species</strong> <strong>Diversity</strong> <strong>in</strong> <strong>Tropical</strong> <strong>Plant</strong>ed <strong>Forests</strong>Fig. 4. Biomass of understorey plant species <strong>in</strong> the study plotsthan that for the small tree species <strong>in</strong> the natural forest<strong>and</strong> for all the tree species with<strong>in</strong> the planted forests.Understorey biomassThe biomass of the understorey plant species <strong>in</strong> theplanted forests was smaller than that of the grassl<strong>and</strong>s(Fig. 4). No I. cyl<strong>in</strong>drica or S. spontaneum grasses grew<strong>in</strong> the sample plots of the planted forests, although a fewwere found outside the sample plots. Among the plantedforests, the total biomass of the understorey plant specieswas largest <strong>in</strong> the P. macrocarpus st<strong>and</strong>. However, alarge proportion of biomass was occupied by shrubs <strong>and</strong>herbs, presumably because of the higher relative light<strong>in</strong>tensity (30.6%) <strong>in</strong> this st<strong>and</strong> than <strong>in</strong> the other st<strong>and</strong>s.Yet even <strong>in</strong> this light environment, I. cyl<strong>in</strong>drica <strong>and</strong> S.spontaneum did not become established <strong>in</strong> the sampleplots, <strong>in</strong>dicat<strong>in</strong>g the strong shade <strong>in</strong>tolerance of these 2grasses.St<strong>and</strong> parameters <strong>and</strong> understorey plant species richness<strong>and</strong> biomassThe correlations between the st<strong>and</strong> parameters, <strong>and</strong>the species richness <strong>and</strong> the biomass of the understorey<strong>and</strong> tree species among the planted forests, are shown <strong>in</strong>Table 3. None of the st<strong>and</strong> parameters showed a significantcorrelation with the species richness of the understoreyplant species. On the other h<strong>and</strong>, the tree densityexhibited a significant negative correlation with the biomassof all the understorey plant species. A significantpositive correlation was found between the biomass ofthe understorey tree species <strong>and</strong> the parameters related tothe st<strong>and</strong> growth performance (mean height, basal area,Table 3. Correlation analysis between various st<strong>and</strong> parameters of planted forests <strong>and</strong> species richness<strong>and</strong> biomass of understorey plant species <strong>and</strong> tree speciesDependent variables(Category of plant)Treedensity(No. ha –1 )Meanheight(m)St<strong>and</strong> parametersBasalarea(m 2 ha –1 )Leafbiomass(t ha –1 )Abovegroundbiomass(t ha –1 )Relativelight<strong>in</strong>tensity(%)Understorey species richnessTree species ns ns ns ns ns nsAll the plant species ns ns ns ns ns nsUnderstorey biomassTree species ns positive* positive* ns positive* nsAll the plant species negative** ns ns ns ns nsns: Not significant, * Significant at 5 % level, ** Significant at 1% level.115


K. Kamo et al.aboveground biomass), but there was no significant correlationbetween the biomass of all the understorey plantspecies or understorey tree species <strong>and</strong> leaf biomass orrelative light <strong>in</strong>tensity.Discussion<strong>Plant</strong>ations of s<strong>in</strong>gle tree species are usually set upfor reafforestation <strong>in</strong> the tropics. However, s<strong>in</strong>gle speciesplantations have often been criticized for be<strong>in</strong>g associatedwith a low level of diversity <strong>in</strong> the ecosystems. Theresults of this study <strong>in</strong>dicate that many plant species,<strong>in</strong>clud<strong>in</strong>g tree seedl<strong>in</strong>gs, became established <strong>in</strong> theplanted forests with s<strong>in</strong>gle tree species <strong>in</strong> the neighborhoodof a natural forest. It is noteworthy that relativelymany plant species became established <strong>in</strong> the E. camaldulensisst<strong>and</strong>, although eucalypts are often considered tohave very few plant species on their forest floor, which isa cause for concern <strong>in</strong> eucalypt plantations 4 .In Sakaerat, before the plantations were established,fire occurred frequently <strong>in</strong> the I. cyl<strong>in</strong>drica <strong>and</strong> S. spontaneumgrassl<strong>and</strong>s, ma<strong>in</strong>ly <strong>in</strong> the dry season, hamper<strong>in</strong>g the<strong>in</strong>itiation of secondary succession to natural forest. Afterthe forest st<strong>and</strong>s were established, the fire frequencydecreased markedly (personal communication, Tha<strong>in</strong>gam).We found that I. cyl<strong>in</strong>drica <strong>and</strong> S. spontaneum hadalmost disappeared under the canopy of the planted forests,ow<strong>in</strong>g to their strong shade <strong>in</strong>tolerance. Thus, thedisappearance of the 2 grasses with<strong>in</strong> the st<strong>and</strong>s was amajor cause of the reduction of fires after st<strong>and</strong> establishment.In the reversion of I. cyl<strong>in</strong>drica <strong>and</strong> S. spontaneumgrassl<strong>and</strong>s to natural forest, the planted forests at theresearch site appeared to play a significant role <strong>in</strong> accelerat<strong>in</strong>gsecondary succession. Fire had not been recorded<strong>in</strong> the S. spontaneum grassl<strong>and</strong> at the site for more than10 years, while the I. cyl<strong>in</strong>drica grassl<strong>and</strong> had often beenburned. The tree species richness was slightly higher <strong>in</strong>the S. spontaneum grassl<strong>and</strong> than <strong>in</strong> the I. cyl<strong>in</strong>dricagrassl<strong>and</strong> (Fig. 2). In some parts of the grassl<strong>and</strong> whereS. spontaneum had died, tree species began to <strong>in</strong>vade.These observations imply that prevent<strong>in</strong>g fire should <strong>in</strong>itiatesecondary succession. The planted forests conta<strong>in</strong>edmore plant species, <strong>in</strong>clud<strong>in</strong>g tree seedl<strong>in</strong>gs, than the S.spontaneum grassl<strong>and</strong>, which <strong>in</strong> turn conta<strong>in</strong>ed more speciesthan the I. cyl<strong>in</strong>drica grassl<strong>and</strong> (Fig. 2). These factsimply that the plantations at the research site facilitatedthe establishment of various plant species, <strong>and</strong> consequentlypromoted secondary succession.One of the major factors that affect the abundantcolonization of tree species <strong>in</strong> a planted forest is the proximityto a natural forest, i.e. seed source 18, 24 . The plantedforests we studied were located near a natural forest witha high species diversity (Kamo et al., unpublished data).This might be one of the factors that contributed to theabundance of tree species with<strong>in</strong> the planted forests.Analysis of similarity <strong>in</strong>dexes (Table 2) showed thatthe species composition differed between the studyst<strong>and</strong>s to more or less extent <strong>and</strong> that there were relativelyfew common species. However, among the largetree species (canopy tree species) <strong>in</strong> the natural forest thatbecame established <strong>in</strong> the planted forests, common speciestended to prevail (Fig. 3). This shows that the seedl<strong>in</strong>gsof large tree species <strong>in</strong> a natural forest can becomewidely established <strong>in</strong> plantations. The larger <strong>and</strong> mostcommonly established tree species <strong>in</strong>cluded Xerospermum<strong>in</strong>termedium, Melia azedarach, Hopea ferrea, Vitexp<strong>in</strong>nata <strong>and</strong> Shorea henryana, a major canopy dom<strong>in</strong>ant<strong>in</strong> the natural forest <strong>in</strong> Sakaerat. In the process of naturalregeneration, animals play an important role <strong>in</strong> seed dispersal<strong>in</strong> the tropics 8,33 . In this connection, birds mightperch on the overstorey trees 6,33 of the planted forests <strong>and</strong>disperse seeds.Many plant species, <strong>in</strong>clud<strong>in</strong>g tree seedl<strong>in</strong>gs,became established with<strong>in</strong> the planted forests (Fig. 2),while almost no I. cyl<strong>in</strong>drica or S. spontaneum grew (Fig.4). The relative light <strong>in</strong>tensity <strong>in</strong> the planted forestsranged from around 3 to 30%. The small leaf biomass ofthe st<strong>and</strong>s caused this low degree of shade (Fig. 1). However,such a light environment was sufficient to allowseedl<strong>in</strong>g establishment while shad<strong>in</strong>g out the 2 grasses.Watt 32 has recognized 4 characteristic phases afterdisturbance <strong>in</strong> a plant community: regenerative phase,build<strong>in</strong>g phase, mature phase <strong>and</strong> degenerative phase. Inthe build<strong>in</strong>g phase, or the stem exclusion stage <strong>in</strong> Oliver<strong>and</strong> Larson’s st<strong>and</strong> development model 21 , the colonizationof understorey species was markedly <strong>in</strong>hibited by the<strong>in</strong>tense competition associated with vigorous growth ofthe st<strong>and</strong>. Thereafter, understorey species became graduallyestablished 5 . In vigorously grow<strong>in</strong>g evergreen forestswith a large leaf biomass, the forest floor often lacksundergrowth 5,27 for a relatively long period of time aftercanopy closure. The relative light <strong>in</strong>tensity then isextremely low, often less than 1%. St<strong>and</strong> leaf biomass<strong>in</strong>creased as the st<strong>and</strong> aged <strong>and</strong> reached a maximumvalue at the time of vigorous growth after canopy closure,<strong>and</strong> thereafter decreased to reach a constant level 15 . Theperiod of large leaf biomass would correspond to that ofvigorous growth of the st<strong>and</strong> 15 , i.e. the build<strong>in</strong>g or stemexclusion stage. The <strong>in</strong>tervention of the build<strong>in</strong>g or stemexclusion stage dur<strong>in</strong>g st<strong>and</strong> development would causethe <strong>in</strong>itiation of the much-delayed natural regenerationwith<strong>in</strong> these st<strong>and</strong>s. Many late successional tree specieshad become established at the late stage of forest devel-116 JARQ 36 (2) 2002


<strong>Plant</strong> <strong>Species</strong> <strong>Diversity</strong> <strong>in</strong> <strong>Tropical</strong> <strong>Plant</strong>ed <strong>Forests</strong>opment <strong>in</strong> Cryptomeria japonica st<strong>and</strong>s, suggest<strong>in</strong>g that a60-year rotation would not be sufficient to develop a highspecies diversity <strong>in</strong> the st<strong>and</strong>s 9 . In a sitka spruce—westernhemlock forest <strong>in</strong> Alaska, dur<strong>in</strong>g the first 70 to 80years of 100-year rotations, the forest floor accumulatedlittle understorey 2 . The leaf biomass of these evergreenforests 29 would be much larger than that of the st<strong>and</strong>s westudied. In contrast, the seedl<strong>in</strong>gs of the tree species <strong>in</strong>our study st<strong>and</strong>s would have begun to colonize the forestfloor at an early stage of st<strong>and</strong> development, s<strong>in</strong>ce the 12-year-old plantations already had many seedl<strong>in</strong>gs. Thismight be ascribed to the small leaf biomass <strong>in</strong> the st<strong>and</strong>s.The leaf biomass rema<strong>in</strong>ed low even dur<strong>in</strong>g the period oflargest biomass production at an early stage of forestdevelopment <strong>in</strong> a tropical st<strong>and</strong> of fast-grow<strong>in</strong>g tree specieswith a small leaf biomass 12 . The peak of leaf biomasstended to be less conspicuous, rema<strong>in</strong><strong>in</strong>g low <strong>in</strong> st<strong>and</strong>swith a small leaf biomass dur<strong>in</strong>g forest development 28 .This temporal leaf dynamics may obscure the build<strong>in</strong>g orstem exclusion stage <strong>in</strong> the st<strong>and</strong>, with the small leaf biomassallow<strong>in</strong>g seedl<strong>in</strong>g establishment early <strong>in</strong> the forestdevelopment <strong>and</strong> promot<strong>in</strong>g an earlier build-up of treeseedl<strong>in</strong>gs <strong>in</strong> these st<strong>and</strong>s than <strong>in</strong> the st<strong>and</strong>s with a largerleaf biomass. Thus, fast-grow<strong>in</strong>g tree species with asmall leaf biomass would act as a better catalyst than treespecies with a large leaf biomass.St<strong>and</strong>s of both exotic <strong>and</strong> <strong>in</strong>digenous species conta<strong>in</strong>edapproximately the same number of plant species<strong>and</strong> tree species (Fig. 2). This suggests that both types ofst<strong>and</strong>s may allow similar plant species establishment <strong>in</strong>the understorey, although their st<strong>and</strong> biomass <strong>and</strong> growthperformance differed. On the other h<strong>and</strong>, exotic speciesst<strong>and</strong>s were superior to <strong>in</strong>digenous species st<strong>and</strong>s for theaccumulation of biomass of the understorey tree species(Fig. 4). The biomass of the understorey tree speciesshowed a positive correlation with the st<strong>and</strong> parametersrelated to biomass <strong>and</strong> growth performance of the st<strong>and</strong>s(Table 3). This <strong>in</strong>dicates that the faster-grow<strong>in</strong>g tree speciesare likely to accumulate a larger biomass of understoreytree species. The growth of the understorey is<strong>in</strong>fluenced by various factors, <strong>in</strong>clud<strong>in</strong>g competition forlight <strong>and</strong> nutrients, pattern of tree regeneration, soil <strong>and</strong>microclimatic effects <strong>and</strong> past st<strong>and</strong> conditions (disturbance)2 . It rema<strong>in</strong>s to be determ<strong>in</strong>ed which factors weremost important <strong>in</strong> our study st<strong>and</strong>s. However, one explanationassociated with the better growth of the st<strong>and</strong>s thatfavored the accumulation of understorey might be earliercanopy closure <strong>in</strong> the st<strong>and</strong>s of the faster-grow<strong>in</strong>g treespecies. Canopy closure would shade out the 2 grasses<strong>and</strong> v<strong>in</strong>es <strong>and</strong> herbs (Fig. 4), allow<strong>in</strong>g the establishment<strong>and</strong> growth of seedl<strong>in</strong>gs on the forest floor. Among theplanted forests, the largest understorey biomass accumulatorwas found to be A. mangium. This species was alsoreported to be a promis<strong>in</strong>g tree species <strong>in</strong> degraded grassl<strong>and</strong>s<strong>in</strong> Indonesia 17 . A. mangium displayed a very lowsprout<strong>in</strong>g capacity <strong>and</strong> a relatively narrow crown <strong>in</strong>Sakaerat (Kamo et al., unpublished data). These traits arelikely to free regenerat<strong>in</strong>g seedl<strong>in</strong>gs from competition<strong>and</strong> allow them to escape with lighter damage after thest<strong>and</strong> is th<strong>in</strong>ned or cut dur<strong>in</strong>g the process of st<strong>and</strong> conversion.These species characteristics would also make A.mangium a superior catalyst.Leav<strong>in</strong>g st<strong>and</strong>s untended after establishment mayfacilitate the establishment of <strong>in</strong>digenous tree specieswith<strong>in</strong> planted forests 23 . This was the case <strong>in</strong> Sakaerat.However, v<strong>in</strong>es should still be cut: we observed abundantv<strong>in</strong>es (Figs. 2, 4) that were damag<strong>in</strong>g the st<strong>and</strong>s.Traditionally, foresters tend to consider tree plantationsas a renewable resource for produc<strong>in</strong>g timber <strong>and</strong>cellulose. Us<strong>in</strong>g a plantation to catalyze the naturalregeneration of tree species is a very different conceptfrom the traditional approach. However, the 2 practicesshould not be alternative but complementary for creat<strong>in</strong>ga forest ecosystem with high biological diversity. Variousunderstorey tree species aris<strong>in</strong>g <strong>in</strong> st<strong>and</strong>s with s<strong>in</strong>gletree species could be used to achieve high biologicaldiversity at the l<strong>and</strong>scape level by creat<strong>in</strong>g various typesof forests—such as human-<strong>in</strong>duced natural forest, mixed<strong>in</strong>digenous st<strong>and</strong>s <strong>and</strong> mixed exotic <strong>and</strong> <strong>in</strong>digenousst<strong>and</strong>s—through various silvicultural treatments, whilest<strong>and</strong>s with s<strong>in</strong>gle tree species could be managed for traditionaltimber <strong>and</strong> cellulose production at the same time.High biological diversity at the l<strong>and</strong>scape level couldbr<strong>in</strong>g about many benefits from forests, <strong>in</strong>clud<strong>in</strong>g woodproduction, water <strong>and</strong> environmental conservation, carbonstock<strong>in</strong>g, education <strong>and</strong> scenic recreation. Toachieve this objective, it is essential to reta<strong>in</strong> some naturalforests <strong>in</strong> the reafforestation areas, avoid<strong>in</strong>g largescaleclear-cutt<strong>in</strong>g.References1. Aide, T. M. et al. (1995): Forest recovery <strong>in</strong> ab<strong>and</strong>onedtropical pastures <strong>in</strong> Puerto Rico. For. Ecol. Manage., 77,77–86.2. Alaback, P. B. (1982): Dynamics of understory biomass<strong>in</strong> sitka spruce — western hemlock forests of southeastAlaska. Ecology, 63, 1932–1948.3. Bruenig, E. F. (1996): Conservation <strong>and</strong> management oftropical ra<strong>in</strong>forests; an <strong>in</strong>tegrated approach to susta<strong>in</strong>ability.CABI Publish<strong>in</strong>g, Wall<strong>in</strong>gford, Oxford, pp.339.4. Calder, I. R., Hall, R. L. & Adlard, P. G. (eds.). (1992):Growth <strong>and</strong> water use of forest plantations. John Wiley& Sons, Chichester, pp.381.5. Cousens, J. (1974): An Introduction to Woodl<strong>and</strong> Ecology.Oliver <strong>and</strong> Boyd, Ed<strong>in</strong>burgh, pp.151.117


K. Kamo et al.6. Guevara, S. et al. (1986): The role of remnant forest trees<strong>in</strong> tropical secondary succession. Vegetatio, 66, 77–84.7. Haggar, J., Wightman, K. & Fisher, R. (1997): The potentialof plantations to foster woody regeneration with<strong>in</strong> adeforested l<strong>and</strong>scape <strong>in</strong> lowl<strong>and</strong> Costa Rica. For. Ecol.Manage., 99, 55–64.8. Howe, H. F. & Smallwood, J. (1982): Ecology of seeddispersal. Annu. Rev. Ecol. Syst., 13, 201–228.9. Ito, S. et al.(1998): <strong>Species</strong> richness of sugi (Cryptomeriajaponica D. Don) plantations <strong>in</strong> relation to their st<strong>and</strong>age, area <strong>and</strong> spatial arrangement. In Proceed<strong>in</strong>gs of VIIInternational Congress of Ecology, 203.10. Kamo, K., Ishizuka, M. & Ohsumi, K. (1989): Growthanalysis of fast-grow<strong>in</strong>g species <strong>and</strong> Benguet p<strong>in</strong>e st<strong>and</strong>.Nettai Nouken Shuho (Bull. Trop. Agric. Res. Cent.), 65,65–79 [In Japanese].11. Kamo, K. (1990): Productivity of fast-grow<strong>in</strong>g species <strong>in</strong>Thail<strong>and</strong>. Trop. For., 19, 26–34 [In Japanese].12. Kawahara, T., Kanazawa, Y. & Sakurai, S. (1981): Biomass<strong>and</strong> net production of man-made forests <strong>in</strong> the Philipp<strong>in</strong>es.J. Jpn. For. Soc., 63, 320–327.13. Keenan, P. et al.(1997): Restoration of plant biodiversitybeneath tropical tree plantations <strong>in</strong> northern Australia.For. Ecol. Manage., 99, 117–131.14. Kimoto, S. & Takeda, H. (1989): Introduction of communityecology. Kyoritsu, Tokyo, pp. 198 [In Japanese].15. Kira, T. & Shidei, T. (1967): Primary production <strong>and</strong>turnover of organic matter <strong>in</strong> different forest ecosystemsof the Western Pacific. Jpn. J. Ecol., 17, 79–87.16. Kira, K. (1978): Community architecture <strong>and</strong> organicmatter dynamics <strong>in</strong> tropical lowl<strong>and</strong> ra<strong>in</strong> forests of SoutheastAsia with special reference to Pasoh forest, WestMalaysia. In <strong>Tropical</strong> Trees as Liv<strong>in</strong>g Systems. eds.Toml<strong>in</strong>son, P. B. & Zimmermann, M. H., CambridgeUniversity Press, Cambridge. pp. 657.17. Kuusipalo, J. et al. (1995): Restoration of natural vegetation<strong>in</strong> degraded Imperata cyl<strong>in</strong>drica grassl<strong>and</strong>; understorydevelopment <strong>in</strong> forest plantations. J. Veg. Sci., 6,205–210.18. Lamb, D. et al.(1997): Rejo<strong>in</strong><strong>in</strong>g habitat remnants; restor<strong>in</strong>gdegraded ra<strong>in</strong>forest l<strong>and</strong>s. In <strong>Tropical</strong> Forest Remnants.eds. Laurance, W. F. & Bierregaard, R, O., TheUniversity of Chicago Press, Chicago, 336–385.19. Lugo, A.E., Parrotta, J.A. & Brown, S. (1993): Loss ofspecies caused by tropical deforestation <strong>and</strong> their recoverythrough management. Ambio, 22, 106–109.20. Ogawa, H. et al.(1965): Comparative ecological studieson three ma<strong>in</strong> types of forest vegetation <strong>in</strong> Thail<strong>and</strong>. II<strong>Plant</strong> biomass. Nat. Life Southeast Asia, 4, 49–80.21. Oliver, C. D. & Larson, B. C. (1990): Forest St<strong>and</strong>Dynamics. McGraw-Hill, Inc., New York, pp.467.22. Parrotta, J. A. (1995): The <strong>in</strong>fluence of overstory compositionon understory colonization by native species <strong>in</strong>plantations on a degraded tropical site. J. Veg. Sci., 6,627–636.23. Parrotta, J. A., Turnbull, J. W. & Jones, N. (1997): Catalyz<strong>in</strong>gnative forest regeneration on degraded tropicall<strong>and</strong>s. For. Ecol. Manage., 99, 1–7.24. Parrotta, J. A. et al.(1997): Development of floristicdiversity <strong>in</strong> 10-year-old restoration forests on a bauxitem<strong>in</strong>ed site <strong>in</strong> Amazonia. For. Ecol. Manage, 99, 21–42.25. Rundel, P. W. & Boonpragob, K. (1995): Dry forest ecosystemsof Thail<strong>and</strong>. In Seasonally Dry <strong>Tropical</strong> <strong>Forests</strong>.eds. Bullock, S., Mooney, H. & Med<strong>in</strong>a, E., CambridgeUniversity Press, Cambridge, 93–123.26. Sato, A., Kanazawa, Y. & Kamo, K. (1989): St<strong>and</strong> structure<strong>and</strong> growth analysis of giant ipil-ipil (Leucaena leucocephala(Lam.) de Wit). Nettai Nouken Shuho (Bull.Trop. Agric. Res. Cent.), 65, 80–93 [In Japanese].27. Shidei, T. et al. (1974): H<strong>in</strong>oki St<strong>and</strong>s: Ecology <strong>and</strong> NaturalRegeneration. Chikyusha, Tokyo, pp. 375 [In Japanese].28. Sukachev, V. & Dylis, N. (1964): Fundamentals of ForestBiogeocoenology. Oliver & Boyd, Ed<strong>in</strong>burgh, pp.672.29. Tadaki, Y. (1977): Leaf biomass. In Primary Productivityof Japanese <strong>Forests</strong>. JIBP Synth. eds. Shidei, T. & Kira,T., 16, 39–44.30. Thoranisorn, S., Sahunalu, P. & Yoda, K. (1991): Litterfall<strong>and</strong> productivity of Eucalyptus camaldulensis <strong>in</strong>Thail<strong>and</strong>. J. Trop. Ecol., 7, 275–279.31. Yoshioka, J. (1986): Trial classification of forest soil forafforestation <strong>in</strong> Sakaerat <strong>in</strong> Thail<strong>and</strong>. Trop. For., 7, 21–28 [In Japanese].32. Watt, A. S. (1947): Pattern <strong>and</strong> process <strong>in</strong> the plant community.J. Ecol., 35, 1–22.33. Wunderle, J. M. (1997): The role of animal seed dispersal<strong>in</strong> accelerat<strong>in</strong>g native forest regeneration on degradedtropical l<strong>and</strong>s. For. Ecol. Manage., 99, 223–235.118 JARQ 36 (2) 2002

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!