Ryoichi Doi and Senaratne L. Ranamukhaarachchi

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1 SILVA FENNICA Silv Fennic 43(5) reserch rticles ISSN The Finnish Society of Forest Science The Finnish Forest Reserch Institute Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis Plnttion on Degrded Lnd in Skert, Thilnd Ryoichi Doi nd Senrtne L. Rnmukhrchchi Doi, R. & Rnmukhrchchi, S.L Community-level physiologicl profiling in monitoring rehilittive effects of Acci uriculiformis plnttion on degrded lnd in Skert, Thilnd. Silv Fennic 43(5): This study ws conducted to investigte the rehilittive effects of plnting Acci uriculiformis trees on degrded lnd y oserving vritions in soil cteril community profiles provided y community-level physiologicl profiling. Soil cteril nd physicochemicl comprisons etween n originl evergreen forest nd the Acci plnttion plot, estlished on n re severely degrded s result of deforesttion, showed tht most soil chrcteristics were rehilitted 18 to 19 yers fter the plnttion of Acci ccording to single vriles, Shnnon nd Simpson diversity indices sed on the community-level physiologicl profiles, principl component nlysis nd redundncy nlysis. However, more strict sttisticl comprison, discriminnt nlysis, completely discriminted etween the Acci plnttion nd the evergreen forest soils when the community-level physiologicl profiles were compred. Thus, the Acci plnttion soil ws shown to still e in the process to full recovery. Here, we discuss the relevnce of plnting A. uriculiformis in lnd rehilittion schemes in svnn regions. Keywords Acci uriculiformis, community-level physiologicl profile, lnd degrdtion nd rehilittion, multivrite nlysis, soil cteril community Addresses AFE Building, School of Environment, Resources nd Development, Asin Institute of Technology, Pthumthni 12120, Thilnd E-mil roird@eiou.pt Received 4 August 2008 Revised 25 My 2009 Accepted 3 Septemer 2009 Aville t 739

2 Silv Fennic 43(5), 2009 reserch rticles 1 Introduction Deforesttion hs emerged s chllenge to socio-economic development in Thilnd (The tenth ntionl 2006) s in mny other countries. In the pst four decdes, forest res in Thilnd s whole re estimted to hve declined y 50% or more (Fisher nd Hirsch 2008). In the tropics, deforesttion often leds to lnd degrdtion nd results in degrded soils under tropicl climtic conditions (Eden nd Prry 1996). Deforesttion is seen s mjor cuse of incresingly severe prolems of drought nd flood (Krirpnond nd Atkinson 1998). Since the lte 1980s, the Thi government hs een tking mesures to rehilitte the degrded lnds. Reforesttion is one of the rehilittion mesures, nd trees hve een plnted on the degrded lnds under susidy scheme of the government (Shrp nd Nkgoshi 2006). Becuse the ntive tree species re prone to fil to survive due to the degrded soil conditions, in the reforesttion strtegy, exotic plnt species re often introduced to rehilitte the degrded lnds with hrsh soil conditions (Ashton et l. 2001). However, the strtegy is often criticized ecuse the introduced exotic species my result in iologicl deserts (Wuethrich 2007). Another criticl opinion is tht the exotic tree species my escpe to djcent res, thretening ntive species (Hrtley 2002). When considering these constrints, we question whether introducing n exotic tree species is truly rehilittive. There hs een some evidence of the rehilittive effects of reforesttion. The oveground diversity of plnt species drws ttention reltively well. Surveys of the distriution of plnt species following reforesttion hve een used to demonstrte the success or filure of the reforesttion in terms of recovery of the originl ecosystems (Kmo et l. 2002). In ddition to the oveground plnt community structure, soil qulity hs lso een used s n indictor of rehilittive effects of reforesttion (Young 1997). On the other hnd, soil iotic chnges re not well documented in investigtions of the rehilittive effects of reforesttion, while the chnges hve een reported s responses to lnd degrdtion cused y deforesttion. When soil microes function well, the soil my support trees nd other plnts in the ecosystem (Bere et l. 1995). Soil cteri, s prt of the soil microil community, contriute to plnt growth y minerl soluiliztion (Derylo nd Skorupsk 1992), nitrogen fixtion (Alrecht et l. 1981) nd production of plnt growth hormones (Nieto nd Frnkenerg 1989). Therefore, n indictor of rehilittion is the recovery of soil microes nd their functions, though they hve seldom een monitored in investigtions of the rehilittive effects of reforesttion. Rehilittive effects of reforesttion re shown y compring soils of the rehilitted re with reforesttion nd the djcent degrded lnd without reforesttion. To compre the soils, we hve used community-level physiologicl profiling of soil cteril community s method for multivrite profiling of the soils (Grlnd nd Mills 1991). This method revels the functionl potentil of the soil cteril community through oservtion of the utiliztion pttern of different cron sources. In this study, we profiled soils over lnd degrdtion-rehilittion grdient in Skert, Thilnd (Doi nd Rnmukhrchchi 2007). The originl vegettion type of Skert is dry evergreen forest. But humn ctivities, minly slsh-nd-urn cultivtion, hve deforested some prts of Skert. Some res were cultivted nd ndoned fter intensive exploittion of the soil resources, cusing the ground to ecome re. In some spces, the re ground ws rehilitted y plnting Acci uriculiformis trees (Ski et l. 2009). Bre ground ws the most degrded vegettion type, nd the rehilitted Acci plnttion plots nd the dry evergreen forest represented lnd degrdtionrehilittion grdient. The min ojective of this study ws to investigte the rehilittive effects of the Acci plnttion y profiling the grdient in wet-to-dry sesonl chnge. 2 Mterils nd Methods 2.1 Site Description This study ws conducted in Skert Environmentl Reserch Sttion, Thilnd. The detils were descried elsewhere (Doi nd Rnmukhrchchi 2007, Doi nd Skuri 2004). The nnul precipittion is 1260 mm nd the verge 740

3 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis Fig. 1. The vegettion types of the Skert Environmentl Reserch Sttion nd the smpling points. DDF nd DEF indicte dry deciduous forest nd dry evergreen forest, respectively. temperture is 26 C. The climte is clssified s svnn (Köppen 1931). The soil is originlly n Orthic Acrisol, ccording to the FAO/UNESCO scheme (FAO/UNESCO 1979). We compred soils of dry evergreen forest (the originl vegettion), Acci plnttion nd re ground (the most degrded vegettion). These vegettion types represent lnd degrdtionrehilittion grdient. The vegettion types were rndomly distriuted. Thus, the vegettion mosic ws regrded s completely rndomized design (Fig. 1, Doi nd Skuri 2004). The numers of replictions were 7, 7 nd 6 for dry evergreen forest, Acci plnttion nd re ground, respectively. All the smpling points were on slight slopes (less thn 10 ). The dry evergreen forest is primrily dominted y Hope ferre nd Shore spp. tht form the upper storey 20 to 40 m ove ground. A typicl dry evergreen forest contins more thn 1000 trees (trunk dimeter t rest height > 5 cm) h 1, the totl sl re t 1.3 m height exceeds 30 m 2 h 1 nd the ove ground iomss is over 200 tons h 1 (Knzki et l. 1995). The Acci uriculiformis plnttion plots re scttered in the re (Fig. 1). The Acci plots were estlished in 1986 nd 1987 in the prts tht used to e sujected to slsh-nd-urn shifting cultivtion (Keonium et l. 1976). In these spces, the originl vegettion hd een removed nd the iomss hd een urned. The clered lnd hd een cultivted for few yers, nd then ndoned when the soil qulity deteriorted to the extent tht could not support crop production. Some of the ndoned prts of Skert hd een converted to plnttion plots of Acci mngium, Euclyptus cmldulensis nd other tree species. A. uriculiformis ws one of the introduced tree species. The re ground soil hs een intensively deprived of nutrients nd hs lost conditions seen in forest soils. At these smpling points, recovery of vegettive cover did not occur since the hrsh conditions for plnts mke the re ground to remin so. Morphologicl fetures of re ground cn still e seen t some points in the site. For typicl re ground, the A horizon cn not e recognized. The uppermost horizon is reddish rown (Doi nd Rnmukhrchchi 2007) rich in grvel, nd hs few roots nd other plnt orgns/ deris. The oundry etween the uppermost horizon nd the deeper horizon is not cler, while the horizon deeper thn 50 cm is ple ornge. 2.2 Soil Smpling nd Physicochemicl Profiling Soils were smpled on Sep 25 nd 26, Nov 25 nd 26 nd Dec 24 nd 25, 2005 (Fig. 2). The 741

4 Silv Fennic 43(5), 2009 reserch rticles Fig. 2. Chnges in temperture nd precipittion during the reserch period. smpling ws done within 26 hours, in which the site hd negligile precipittion (< 1 mm). At ech smpling point, hundred-ml core smplers, 5 cm in dimeter, were inserted from the surfce to depth of 5.1 cm. A circle, 10 m in dimeter ws estlished, nd 8 soil cores were rndomly tken within the circle. In ddition, two other cores were rndomly tken in the circle for soil moisture nd ulk density mesurements. The 8 soil cores were immeditely plced into single plstic g, mixed nd rought to the lortory. For community-level physiologicl profiling, the moist soil ws immeditely pssed through 2 mm sieve nd ws rought to the lortory within 12 h. For soil physicochemicl profiling, the soil smples were ir-dried, pssed through 2 mm sieve then nlyzed. Physicochemicl profiling of soils were performed s previously descried (Doi nd Rnmukhrchchi 2007). 2.3 Community-level Physiologicl Profiling The cteril community in ech composite smple ws profiled with three Biolog EcoPltes s previously descried (Doi 2005). Five grms of ech soil smple were suspended in 45 ml of sterilized 0.85% (w/v) NCl nd reciproclly shken t room temperture for 30 min t 120 rpm. The suspension ws centrifuged t 1000 g for 5 min, decnted, nd the pellet ws re-suspended in 45 ml of the NCl solution. Then, the centrifugtion nd suspending ws repeted twice. The soil suspension ws left still for minute, nd 10 ml of the uppermost section ws diluted 40-fold with the NCl solution. This suspension ws used to inoculte Biolog EcoPlte t rte of 0.1 ml/well. Oxidtion of ech cron source ws mesured y quntifying purple color development s result of the reduction of tetrzolium violet. The pltes were incuted t 26 C in the drk nd sornce t 405 nm ws red using microplte reder (Perlong DNM-9602G, Nnjing, PR Chin) t 4 to 12 h intervls for 7 dys. During the incution, the pltes were wrpped in plstic film to void desicction. Vlues for the ove mentioned three pseudo-replictes were verged, nd used for the following sttisticl nlyses. 2.4 Dt Anlyses The following nlyses were performed using the sttisticl softwre, SPSS (SPSS Inc.). Repeted mesures nlysis of vrince for ech of the soil physicochemicl chrcteristics ws performed. As the post-hoc test, Dunnett T3 t-test 742

5 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis ws performed to exmine the significnt differences etween mens. Community-level physiologicl profiles of the soils were nlyzed to construct dt sets for using the informtion. The verge well color development pproch (AWCD, Grlnd nd Mills 1991) ws employed s the first choice. AWCD t prticulr time ws clculted using the following eqution. Averge well color development (AWCD) = Σ(ABSit ABSct ) /31 (1) where ABSit is the sornce t 405 nm for the i-th cron source t the time (t) nd ABSct is the sornce for the control well, including no cron sources, t the time (t). In this study, vlues of AWCD for the re ground soil smples t the convergence stge were oviously poor, then comprle to tht for the forest soil smples t the exponentil stge. This mde it impossile to use profiles of the forest soils t their convergence stges ecuse the color development for the re ground soils did not rech to the color intensity tht the forest soil smples mnifested. Therefore, the kinetic pproch proposed y Lindstrom et l. (1998) ws pplied. Vlues of AWCD in the color development were used to determine the symptote (K) sttisticlly. The vlue of prmeter K ws regrded s the symptote tht the vlue of AWCD finlly reches. The 0.5 K time point ws regrded s the exponentil stge (Lindstrom et l. 1998) nd the 0.95 K time point s the convergence stge. This pproch provided 0.5 K nd 0.95 K dt sets. In ddition to these dt sets, nother dt set, clled re dt set, ws constructed (Hckett nd Griffiths 1997). Then, multivrite profile of the soil smple ws otined. A rtio-trnsformtion ws employed, i.e., ech oservtion ws divided y the sum of ll the oservtions for the smple nd used for sttisticl nlyses. To compre the dt sets in discriminting mong the soils, discriminnt nlysis ws performed. The 31 vlues for ech soil smple were used for discriminnt nlysis of the communitylevel physiologicl profiles. Rw soil physicochemicl dt were used for discriminnt nlysis of soil physicochemicl profiles. The puttingindependents-together method ws chosen. Principl component nlysis ws performed to extrct principl components from the dt set on physicochemicl profiles or the communitylevel physiologicl profiles. Then, the principl component scores were used for descriing the lnd degrdtion-rehilittion grdient. It hd een demonstrted tht the intensity of the lnd degrdtion in Skert ws solely descried y the first principl component derived from the physicochemicl dt set (Doi nd Skuri 2004). Hence, multiple regression nlysis ws performed to otin regression model for descriing the lnd degrdtion-rehilittion grdient sed on the dt set on the communitylevel physiologicl profiles. Scores on the first principl component derived from the physicochemicl dt set were exmined if the vrition hs significnt liner reltionships with scores on principl components extrcted from the dt set on the community-level physiologicl profiles. The multiple regression nlyses involved only the significnt principl components hving eigenvlues of 1 or greter (Kiser 1960). The stepwise method t the defult criteri (p = 0.05 for inclusion nd 0.10 for removl) ws chosen. We clculted iodiversity indices sed on the cron source utiliztion pttern y the soil cteril community (Stddon et l. 1997). Shnnon diversity, Shnnon evenness nd Simpson indices were clculted. The Shnnon diversity nd evenness indices indicte diversity nd evenness of distriution pttern of the community, while the Simpson index indictes dominnce y prticulr portion of the community (Stddon et l. 1997). Repeted mesures nlysis of vrince ws performed for nlyztion, determintion nd comprison of sources of vritions of the diversity indices. Dunnett T3 t-test ws chosen s the post-hoc t-test. CANOCO for Windows 4.02 nd CnoDrw 3.10 (Microcomputer Power, NY) were used for redundncy nlysis of the reltionships mong the soil physicochemicl vriles, the soil cteril community profiles nd the vlues of Shnnon diversity index (ter Brk nd Šmiluer 1998). The smpling dtes were used s the co-vrile to tke the temporl chnges into ccount. 743

6 Silv Fennic 43(5), 2009 reserch rticles 3 Results The climtic conditions re summrized in Fig. 2. The site hd hevy precipittion in Septemer nd Octoer, ut received less wter in Novemer nd Decemer, resulting in wet-to-dry trnsition. Most soil physicochemicl vriles reflected the degrdtive/rehilittive effects significntly (Tle 1, p < 0.05). The lnd degrdtion ws explined y high vlues of ulk density nd exchngele cidity (Al, H), nd the low vlues of moisture content, ph, orgnic mtter content, sic ction contents (K, C, Mg), nd ville phosphorus. The vlues for the Acci plnttion soil were not significntly different from those for the evergreen forest soil, except for the ulk density vlue in Septemer. The re ground soil ws shown to e significntly poorer thn the other soils in some soil physicochemicl vriles. Acci plnttion soil showed rehilittive effects of the plnttion on the degrded soil. The effects of the wet-to-dry sesonl chnge were pronounced s chnges in soil moisture content, ulk density, nd ph. Fig. 3 shows community-level physiologicl profiles of the soils. Some significnt differences were found etween the vegettion types. The re ground soil ws significntly different from the others, ut there were fewer cses in which the mens for the Acci plnttion nd the evergreen forest were differentited t significnce level of p = These results gin indicte tht the soil of the Acci plnttion plot ws progressively recovering the originl community-level physiologicl profile. To extend the use of the dt, multivrite nlyses were performed. Tle 2 indictes the results of principl component nlysis. Regrding vrition in scores on the first principl component extrcted from the overll physicochemicl dt set, vegettion type ws significnt source (p = 0.001), ut the smpling time (p = 0.908) nd the interction (p = 0.798) were not. Thus, the first principl component ws confirmed to e mesure of the intensity of lnd degrdtion in Skert (Doi nd Skuri 2004). The cteril dt sets provided lrger numers of significnt principl components (eigenvlue > 1) thn the physicochemicl dt set. For the overll dt sets on the community-level physiologicl profiles, repeted mesures nlyses of vrince were performed to determine the principl components explining the sesonl chnge. When smpling time ws exmined s source of the vrition, the principl components hving the smllest p vlues were the fifth (0.5 K), the eighth (0.95 K), nd the third (Are). Thus, sesonlity ws found to e minor source of the vrition. Fig. 4 summrizes the regression models tht descrie the lnd degrdtion-rehilittion grdient. These models show tht the first cteril principl components hd significnt liner reltionships with the lnd degrdtion-rehilittion grdient. The soil smple groups of the evergreen forest nd the Acci plnttion overlp while the re ground smples re seprted in the digrms. It ws indicted tht the soil of the Acci plnttion plot ws recovering the originl physiologicl functions. The discriminnt nlyses of the cteril dt sets properly clssified ll the soil smple groups in ll three smpling times nd resulted in no misclssifiction. On the other hnd, discriminnt nlysis of the physicochemicl dt sets resulted in two cses of misclssifiction: n Acci smple ws misclssified s n evergreen forest smple in Septemer nd Novemer, respectively. Hence, the cteril dt sets enled discrimintion mong the soils t lest s well s the physicochemicl dt set. Anlysis of the iodiversity indices reveled significnt sources of the vrition (Tle 3). Smpling time nd vegettion type were significnt (p < 0.003) for ll the dt sets nd ll the indices. No significnt interctions etween these sources of vrition were recognized (p > 0.212); consistent vrition ptterns mong the smpling times or the vegettion types were seen. For exmple, the diversity indices tended to e greter in Septemer nd smller in Decemer. The cteril community in the evergreen forest soil tended to hve the highest functionl diversity, nd tht in the re ground soil hd the poorest. In most comprisons, the diversity indices of the soil cteril community of the evergreen forest were not significntly different from those of the Acci plot. Therefore, it ws gin shown tht the soil of the Acci plot ws incresing in similrity to tht of the evergreen forest during the rehilittion period. This ws indicted y recovering vlues of verge well color development s well. 744

7 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis Tle 1. Soil physicochemicl chrcteristics nd results of repeted mesures nlysis of vrince. Physicochemicl chrcteristics Seprtion of mens for the vegettion types t ech smpling time Significnce of sources of vrition (p vlue) Septemer Novemer Decemer BG* Acci DEF BG Acci DEF BG Acci DEF Vegettion Smpling V S type (V) time (S) Moisture content (%) Bulk density (kg L 1 ) ph Orgnic mtter (%) Bry II P (mg kg 1 dry soil) Exch K (m eq kg 1 dry soil) Exch C (m eq kg 1 dry soil) Exch Mg (m eq kg 1 dry soil) Exch N (m eq kg 1 dry soil) CEC** (m eq kg 1 dry soil) Exch Al (m eq kg 1 dry soil) Exch H (m eq kg 1 dry soil) * BG, Acci, nd DEF in the men seprtion columns indicte re ground, Acci plnttion nd dry evergreen forest, respectively, for ech month nd ech soil physicochemicl chrcteristic. ** CEC, ction exchnge cpcity. The mens indexed y the sme letter do not differ significntly t p = 0.05 ccording to the Dunnett T3 t-test. 745

8 Silv Fennic 43(5), 2009 reserch rticles Tle 2. Structures of the dt sets. Smpling time Dt set Sttistic Principl components Septemer 0.5 K Eigenvlue 9.1* Vrition explined (%) K Eigenvlue Vrition explined (%) Are Eigenvlue Vrition explined (%) Physicochemicl Eigenvlue Vrition explined (%) Novemer 0.5 K Eigenvlue Vrition explined (%) K Eigenvlue Vrition explined (%) Are Eigenvlue Vrition explined (%) Physicochemicl Eigenvlue Vrition explined (%) Decemer 0.5 K Eigenvlue Vrition explined (%) K Eigenvlue Vrition explined (%) Are Eigenvlue Vrition explined (%) Physicochemicl Eigenvlue Vrition explined (%) Septemer, 0.5 K Eigenvlue Novemer, Vrition explined (%) nd 0.95 K Eigenvlue Decemer Vrition explined (%) Are Eigenvlue Vrition explined (%) Physicochemicl Eigenvlue Vrition explined (%) * The underlined figures indicte tht the principl component is significnt ccording to the Keiser s criterion (eigenvlue 1, Keiser 1960). Fig. 3. Community-level physiologicl profiles of the soil cteril communities of the re ground (solid r), the Acci plnttion (open r), nd the evergreen forest (gry r) in ech month nd for ech dt set. The mens indexed y different letters differ significntly t p = 0.05 ccording to the Dunnett T3 t-test. The mens without indexing letters indicte no significnt differences mong the vegettion types. The error r indictes the stndrd devition. (n = 6, re ground; n = 7, Acci plnttion nd evergreen forest) 746

9 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis Reltive cron source utiliztion=asornce vlue for the i-th cron source/σ[sornce vlue for cron source] Pyruvic cid methy ester 0.5 K Septemer 0.5 K Novemer 0.5 K Decemer Tween 40 Tween 80 Alf-Cyclodextrin Glycogen D-Celloiose 0.95 K Septemer 0.95 K Novemer Are Septemer Are Novemer Are Decemer Alf-D-Lctose Bet-Methyl-D-glucoside D-Xylose i-erythritol D-Mnnitol N-Acetyl-D-glucosmine D-Glucosminic cid Glucose-1-phosphte D,L-Alf-glycerol phosphte D-Glctonic cid gmm-lctone D-Glcturonic cid 2-Hydroxy enzoic cid 4-Hydroxy enzoic cid Gmm-Hydroxyutyric cid Itconic cid Alf-Ketoutyric cid D-Mlic cid L-Arginine 0.95 K Decemer c L-Asprgine L-Phenyllnine L-Serine L-Threonine Glycyl-L-glutmic cid Phenylethylmine Putrescine 747

10 Silv Fennic 43(5), 2009 reserch rticles Tle 3. Vlues of the iodiversity indices for the soil cteril communities of the vegettion types nd the descriptive sttistics t ech smpling time sed on community-level physiologicl profiling. Dt set Diversity index or Septemer Novemer Decemer Significnce of lnd degrdtion/ descriptive sttistic rehilittion (vegettion type, V) nd smpling time (S) s sources of vrition (p vlue) Bre Acci Evergreen Bre Acci Evergreen Bre Acci Evergreen V S V S ground plnttion forest ground plnttion forest ground plnttion forest 0.5 K Shnnon A * A A A B B A AB AB Shnnon evenness A AB A A B B A A AB / Simpson 20.0 A 20.7 A 21.5 A 18.3 A 19.7 B 20.4 B 17.7 A 19.4 AB 19.9 AB Averge well color A A A A A A c A A A development (AWCD) Numer of cron 28.2 A 27.7 A 29.1 A 25.8 A 25.6 A 26.1 B 28.5 A 26.6 A 28.0 A sources with AWCD > K Shnnon A A A A A A A A A Shnnon evenness A A A A A A A A A / Simpson 27.8 A 28.9 A 29.4 A 25.9 A 28.8 A 29.4 A 24.7 A 27.3 A 28.5 A Averge well color A A A A A A c A A A development (AWCD) Numer of cron 30.7 A 30.9 A 30.7 A 30.0 A 30.9 A 30.9 A 30.5 A 30.4 A 30.9 A sources with AWCD > 0 Are Shnnon A A A A B B A B B Shnnon evenness A A A A B AB A B B / Simpson 26.2 A 29.4 A 30.0 A 26.0 c A 28.6 B 29.2 B 22.8 A 27.2 B 27.9 C Are (Asornce hour) 3470 A 5121 A 5464 A 2915 A 4021 B 4292 B 3161 A 4779 A 5150 A Numer of cron 30.2 A 31.0 A 31.0 A 30.0 A 30.9 A 30.9 A 30.0 A 30.9 A 30.7 A sources with > 0 AWCD * For ech month nd ech diversity index, the mens indexed y the sme smll letter do not differ significntly mong the vegettion types t p = 0.05 ccording to the Dunnett T3 t-test. For ech index nd ech vegettion type, the mens indexed y the sme cpitl letter do not differ significntly mong the smpling times t p = 0.05 ccording to the t-test. 748

11 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis Fig. 4. Regression models descriing the lnd degrdtion-rehilittion grdient shown s the vritions of score on the first principl components derived from the physicochemicl dt sets. For ech month, the solid dimond, the open circle nd the gry tringle indicte re ground, Acci plnttion nd evergreen forest, respectively. The formule long the horizontl xes were provided y the multiple regression nlysis. PCi indictes the score on the i-th principl component for the soil smple. The p vlue following the R vlue indictes the significnce of the regression model. For the overll dt sets, the dimond, the circle nd the tringle indicte re ground, Acci plnttion nd evergreen forest, respectively. The solid, the gry nd the open symols indicte the soil smples tken in Septemer, Novemer nd Decemer, respectively. By direct grdient nlyses, the first nd second xes were extrcted to explore the reltionships etween the vlues of the Shnnon diversity index nd soil environmentl grdients. The first xes extrcted y redundncy nlysis hd eigenvlues of to (Fig. 5). The sptil vrition in soil moisture content ws the most significnt soil environmentl grdient relted to the Shnnon diversity index nd soil cteril community profiles. Fig. 5 shows similrity etween the cteril communities in the Acci plnttion nd the evergreen forest soils. Agin, the re ground soil cteril community ws clerly different from the others. In ddition to moisture content, ulk density, ph nd/or orgnic mtter content were significnt soil environmentl grdients relted to the vrition in the communitylevel physiologicl profiles. In other words, the community-level physiologicl profile of the re ground soil, hving the lowest functionl diversity in Skert, ppered when the soil ecme drier, more cidic, hevier, nd/or more rren ecuse of humn ctivities. 4 Discussion Besides its high dptility in degrded svnn res (Bdejo 1998), A. uriculiformis is known for its nitrogen-fixtion property (Sprent nd Prsons 2000), enriching mcrofunl composition 749

12 Silv Fennic 43(5), 2009 reserch rticles Fig. 5. Redundncy nlysis ordintion digrms indicting (1) scores on the ordintion xes for the soil smples (top); nd (2) chnges in functionl diversity of soil cteril community in the ordintion plne (ottom). For the top figures, the solid dimond, the open circle nd the gry tringle indicte re ground, Acci plnttion nd evergreen forest, respectively. For the ottom figures, the solid nd the dotted contour lines indicte interpoltion nd extrpoltion, respectively. (Moukou-Kimts et l. 1998), low llelopthic effects (Bernhrd-Reverst 1999), nd pumping nutrients from the susoil (Kng 1993). Such rehilittive effects were lso seen in our study s the ove results. The Acci plnttion soil hd lrgely recovered the originl conditions nd cteril functions seen in the evergreen forest soil. At the sme time, in the study period in 2005, the A. uriculiformis plnttion soil seemed to still e recovering the originl chrcteristics of the evergreen forest. This ws especilly indicted y the results of discriminnt nlysis which still clerly discriminted etween the cteril community profiles of the evergreen forest nd the Acci plnttion soils. A possile explntion is tht slight physicochemicl soil chnges (p > 0.10) my significntly ffect soil iotic vriles, such s soil enzyme ctivity (Jh et l. 1992) nd/or soil cteril community structure (Doi nd Skuri 2003). Hence, the oserved significnt differences etween the soil cteril communities my e results of the mplifiction of physicochemicl differences etween the soils of the Acci plot nd the evergreen forest. Soil physicochemicl chnges hve een oserved reltively more often in ssocition with lnd rehilittion involving reforesttion. Recently, some surveys hve reported recovery of single soil iotic vriles following reforesttion (Doi nd Rnmukhrchchi 2009; Ren et l. 2007; Lee et l. 2006). In comprison with oservtion of single iotic vriles, multivrite profiling of soil cteril communities my offer more chnces to find differences in soil qulity. In Skert, protecting the plnttion plot from fire seems to e importnt for succession (Shunlu nd Dhnmnond 1995). The A. uriculiformis plnttion soil ws concluded to hve 750

13 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis lredy een rehilitted eyond the soil of the dry deciduous forest (Fig. 1) distured y humn ctivities, minly frequent running fire (Doi nd Rnmukhrchchi 2007). In North Americn forests, fire cn ccelerte succession if the dominnt species re urnt nd then opportunities for growth re given to suppressed ntive tree species (Arms nd Scott 1989). On the other hnd, in cse study in Jpn, fire set ck succession of secondry forest dominted y Jpnese red pine trees to n erly stge, while pine wilt enhnced succession y promoting the growth of previously suppressed ok trees (Fujihr 1996). In Skert, strong solr rdition, thus higher temperture, in ddition to soil dryness (Fig. 5), ws indicted to stress the ntive evergreen tree species (Fig. 5, Skuri et l. 1998; Stott 1984). Recovery of moister soil nd cooler microsite conditions y cnopy development would e required for succession (Ren et l. 2007) while fire would destroy the chnces for the ntive tree species, s well s the iotic life in the soil, to recover. Once A. uriculiformis plnttion estlishes cnopy, self-thinning fvors succession (Ashton et l. 2001). In 1998, mong 6 tree species introduced to the Skert Environmentl Reserch Sttion, the A. uriculiformis plnttion recorded the second highest species richness of plnts (51), following the Dlergi cochinchinensis plnttion, which scored species richness of 59, when the evergreen forest hd 114 species (Kmo et l. 2002). We oserved mny tree species in the A. uriculiformis plots on the smpling dys in This perceivle progress in succession is thought to contriute to the reltive proximity of the A. uriculiformis plnttion soil to the evergreen forest soil. Incresing plnt community diversity cn help enrich soil fertility (El-Kelwy nd Ksiksi 2005) nd estlish the soil ecologicl structure (Bere et l. 1995) formed y plnt roots nd root exudtes (Grlnd 1996) on which soil nimls nd microes rely. These processes involved in succession were likely recovering the soil functions oserved in this study. On the other hnd, we re sure tht the Acci plnttion plots will require severl more yers to fully recover the originl soil qulity so there re no recognizle differences from the evergreen forest soil y discriminnt nlysis (Young 1997). Yemefck et l. (2005) recognized incomplete soil fertility recovery in comprle rehilittion period of 15 yers fter shifting cultivtion in Cmeroon. In the southern Yuctn peninsul of Mexico, under similr climtic conditions, yers ws estimted for recovery of totl oveground iomss following shifting cultivtion, sed on the most optimistic estimte (Chzdon 2003). Thus, the A. uriculiformis plnttion soil nd the ecosystem were very likely to e in succession towrd the climx. Therefore, even when we see similrity in multivrite profiles etween soils of reforesttion plots nd originl forest, we need to keep in mind criteri such s discriminnt nlysis nd sttistics (Doi nd Rnmukhrchchi 2007). The fitness of introduced exotic tree species is site-specific (Ehrenfeld 2003). In the region, A. uriculiformis plnttion looks suitle to invest for the environment, thus, in turn the society nd the economy. In Skert, the fitness of A. uriculiformis ws higher thn Euclyptus cmldulensis nd, nd thinning in the A. uriculiformis plot encourged growth of ntive evergreen forest species, Hope odort (Ski et l. 2009). A similr cse of ccelerted succession y thinning in n A. uriculiformis stnd in Vietnm ws reported y McNmr et l. (2006). In cses when the lor force for forest mngement is limited (Nwir et l. 2007), self-thinning is n option. Therefore, Acci monoculture ppels s n lterntive for rehilittion of degrded lnds in svnn regions (Lugo 1997). Acknowledgements All the stff memers of the Skert Environmentl Reserch Sttion ssisted this reserch ctivity. The uthors grtefully cknowledge their relentless support. References Arms, M.D. & Scott, M.L Disturnce-medited ccelerted succession in two Michign forest types. Forest Science 35(1): Alrecht, S.L., Okon, Y., Lonnquist, J. & Burton, 751

14 Silv Fennic 43(5), 2009 reserch rticles R.H Nitrogen fixtion y corn-azospirillum ssocition in temperte climte. Crop Science 21(2): Ashton, M.S., Guntilleke, C.V.S., Singhkumr, B.M.P. & Guntilleke, I.A.U.N Restortion pthwys for rin forest in southwest Sri Lnk: review of concepts nd models. Forest Ecology nd Mngement 154(3): Bdejo, M.A Agroecologicl restortion of svnn ecosystems. Ecologicl Engineering 10(2): Bere, M.H., Colemn, D.C., Crossley, D.A. Jr., Hendrix, P.F. & Odum, E.P A hierrchicl pproch to evluting the significnce of soil iodivresity to iogeochemicl cycling. Plnt nd Soil 170(1) : Bernhrd-Reverst, F The leching of Euclyptus hyrids nd Acci uriculiformis lef litter: lortory experiments on erly decomposition nd ecologicl implictions in Congolese tree plnttions. Applied Soil Ecology 12(3): Csid, L.E. Jr., Klein, D.A. & Sntoro, T Soil dehydrogense ctivity. Soil Science 98(6): Chzdon, R.L Tropicl forest recovery: legcies of humn impct nd nturl disturnces. Perspectives in Plnt Ecology, Evolution nd Systemtics 6(1 2): Derylo, M. & Skorupsk, A Rhizoil siderophore s n iron source for clover. Physiologi Plntrum 85(1): Doi, R Effects of ir-drying soils on discriminting the soils with community-level physiologicl profiling. Journl of Generl nd Applied Microiology, 51(6): & Rnmukhrchchi, S.L Soil color designtion using Adoe PhotoshopTM in estimting soil fertility restortion y Acci uriculiformis plnttion on degrded lnd. Current Science 92(11): & Rnmukhrchchi, S.L Soil dehydrogense in lnd degrdtion-rehilittion grdient: oservtions from svnn site with wet/ dry sesonl cycle. Revist de Biologi Tropicl 57(1 2): & Skuri, K Soil environmentl fctors relting to diversity of culturle soil cteril communities in the Skert Environmentl Reserch Sttion, Thilnd. Tropics 12(3): & Skuri, K Principl components derived from soil physico-chemicl dt explined lnd degrdtion grdient, nd suggested the pplicility of new indexes for estimtion of soil productivity in the Skert Environmentl Reserch Sttion, Thilnd. Interntionl Journl of Sustinle Development nd World Ecology 11(3): Eden, M.J. & Prry, J.T Lnd degrdtion in the tropics: environmentl nd policy issues. Glol Development & the Environment Series. Pinter, London, UK. Ehrenfeld, J.G Effects of exotic plnt invsions on soil nutrient cycling processes. Ecosystems 6(6): El-Kelwy, A. & Ksiksi, T Artificil forests s conservtion sites for the ntive flor of the UAE. Forest Ecology nd Mngement 213(1): FAO/UNESCO Soil mp of the world. IX, Southest Asi. UNESCO, Pris. Fisher, R. & Hirsch, P Poverty nd grrin-forest interctions in Thilnd. Geogrphicl Reserch 46(1): Fujihr, M Development of secondry pine forests fter pine wilt disese in western Jpn. Journl of Vegettion Science 7(5): Grlnd, J.L Anlyticl pproches to the chrcteriztion of smples of microil communities using ptterns of potentil C source utiliztion. Soil Biology nd Biochemistry 28(2): & Mills, A.L Clssifiction nd chrcteriztion of heterotrophic microil communities on the sis of ptterns of community-level sole-cronsource utiliztion. Applied nd Environmentl Microiology 57(8): Hckett, C.A. & Griffiths, B.S Sttisticl nlysis of the time-course of Biolog sustrte utiliztion. Journl of Microiologicl Methods 30(1): Hrtley, M.J Rtionle nd methods for conserving iodiversity in plnttion forests. Forest Ecology nd Mngement 155(1 3): Ji, B., Zhou, G, Wng, F., Wng, Y., Yun, W. & Zhou, L Prtitioning root nd microil contriutions to soil respirtion in Leymus chinensis popultions. Soil Biology nd Biochemistry 38(4): Keonim, P., Khoort, P., Sunthornsn, W., Issreey, M., Cherdchun, C. & Buchum, W A study of illegl deforesttion in the reserved forest re 752

15 Doi nd Rnmukhrchch Community-level Physiologicl Profiling in Monitoring Rehilittive Effects of Acci uriculiformis t the Skert Environmentl Reserch Sttion. Environmentl nd Ecologicl Reserch Deprtment, Applied Scientific Reserch Corportion of Thilnd, Bngkok. 53 p. Kiser, H.F The ppliction of electronic computers to fctor nlysis. Eductionl nd Psychologicl Mesurement 20(1): Kmo, K., Vchrngkur, T., Tiynon, S., Viriyunch C., Nimpil, S. & Dongsrisen, B Plnt species diversity in tropicl plnted forests nd impliction for restortion of forest ecosystems in Skert, Northestern Thilnd. JARQ 36(2): Kng, B.T Sustinle groforestry systems for the tropics: concepts nd exmples. IITA Reserch Guide 26. IITA, Idn. Knzki, M., Yod, K. & Dhnmnond, K Mosic structure nd tree growth pttern in monodomoionnt tropicl sesonl evergreen forest in Thilnd. In: Box, E. O., Peet, R. K., Msuzw, T., Ymd, I., Fujiwr, K., & Mycosk, P.F. (eds). Vegettion science in forestry. Kluwer Pulishers, Netherlnds. p Köppen, W Grundriss der Klimkunde, Wlter de Gruyter, Berlin. Krirpnond, N. & Atkinson, A Wtershed mngement in Thilnd: concepts, prolems nd implementtion. Regulted Rivers: Reserch nd Mngement 14(6): Lee, Y. K., Lee, D. K., Woo, S. Y., Prk, P. S., Jng, Y. H. & Arhm, E.R.G. Effect of Acci plnttions on net photosynthesis, tree species composition soil enzyme ctivities, nd microclimte on Mt. Mkiling. Photosynthetic 44(2): Lindstrom, J.E., Brry, R.P. & Brddock, J.F Microil community nlysis: kinetic pproch to constructing potentil C source utiliztion ptterns. Soil Biology nd Biochemistry 30(2): Lugo, A.E The pprent prdox of reestlishing species richness on degrded lnds with tree monoculture. Forest Ecology nd Mngement 99(1 2): Moukou-Kimts, I.M.C., Bernhrd-Reverst, F. & Loumeto, J.J Chnge in soil mcrofun nd vegettion when fst-growing trees re plnted on svnn soils. Forest Ecology nd Mngement 110(1 3): McNmr, S., Tinh, D.G., Eriskine, P.D., Lm, D., Ytes, D. & Brown, S Rehilitting degrded forest lnd in centrl Vietnm with mixed ntive species plntings. Forest Ecology nd Mngement 233(2 3): Nwir, A.A., Murniti & Rumoko, L Forest rehilittion in Indonesi. Center for Interntionl Forestry Reserch. SMK Grfk Des Puter, Jkrt, Indonesi. Nieto, K.F. & Frnkenerg, W.T Biosynthesis of cytokinins in soil. Soil Science Society of Americ Journl 53(3): Ren, H., Li, Z., Shen, W., Yu, Z., Peng, S., Lio, C., Ding, M. & Wu, J Chnges in iodiversity nd ecosystem function during the restortion of tropicl forest in south Chin. Science in Chin Series C: Life Sciences 50(2): Shunlu, P. & Dhnmnond, P Structure nd dynmics of dry dipterocrp forest, Skert, northestern Thilnd. In: Box, E.O., Peet, R.K., Msuzw, T., Ymd, I., Fujiwr, K. & Mycosk, P.F. (eds.). Vegettion science in forestry. Kluwer Pulishers, Netherlnds. p Ski, A., Visrtn, T., Vchrngkur, T., Thingm, R., Tnk, N., Ishizuk, M. & Nkmur, S Effect of species nd spcing of fst-growing nurse trees on growth of n indigenous tree, Hope odort Rox., in northest Thilnd. Forest Ecology nd Mngement 257(2): Skuri, K., Tnk, S., Ishizuk, S. & Knzki, M Differences in soil properties of dry evergreen nd dry deciduous forests in the Skert Environmentl Reserch Sttion. Tropics 8(1 2): Shrp, A & Nkgoshi, N Rehilittion of degrded forests in Thilnd: policy nd prctice. Lndscpe nd Ecologicl Engineering 2(2): Sprent, J.I. & Prsons, R Nitrogen fixtion in legume nd non-legume trees. Field Crops Reserch 65(2 3): Stddon, W.J., Duchesne, L.C. & Trevors, J.T Microil divesity nd community structure of postdistrunce forest soils s determined y solecron-source utiliztion ptterns. Microil Ecology 34(2): Stott, P The svnn forests of minlnd southest Asi: n ecologicl survey. Progress in Physicl Geogrphy 8(3): ter Brk, C. J. F. & Šmiluer, P Cnoco reference mnul nd user s guide to Cnoco for windows: softwre for cnonicl community ordintion 753

16 Silv Fennic 43(5), 2009 reserch rticles (version 4). Microcomputer Power, Ithc, NY. The tenth ntionl economic nd socil development pln ( ) Ntionl Economic nd Socil Development Bord of Thilnd, Office of the Prime Minister, Bngkok. Wuethrich, B Reconstructing Brzil s Atlntic rinforest. Science 315(5815): Yemefck, M., Rossiter, D.G. & Njomgng, R Multi-scle chrcteriztion of soil vriility within n griculturl lndscpe mosic system in southern Cmeroon. Geoderm 125(1 2): Young, A Agroforestry for soil mngement. 2nd ed. CAB Interntionl nd ICRAF, New York Totl of 52 references 754

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