ENNRJ-2015!01!01format01-12_Effects of Climate Variability on Monthly Growth of Aglaia Odoratissima...

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Effects of Climate Variability on MonthlyGrowth of Aglaia odoratissima and

Hydnocarpus ilicifolia at the Sakaerat

Environmental Research Station (SERS),

Northeastern Thailand

ARTICLEJUNE 2015

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Environment and Natural Resources J. Vol 13, No.1, January-June 2015:1-12 1

*Corresponding author:

E-mail: [email protected]

Effects of Climate Variability on Monthly Growth ofAglaia odoratissimaandHydnocarpusilicifoliaat the Sakaerat Environmental Research

Station (SERS), Northeastern Thailand

Kritsadapan Palakit1*,Khwanchai Duangsathaporn2, Somkid Siripatanadilok3,

Pichit Lumyai2

1Faculty of Environment and Natural Resource Studies, Mahidol University,Nakhon Pathom, Thailand.

2Laboratory of Tropical Dendrochronology, Department of Forest Management, Faculty of Forestry,Kasetsart University, Thailand

3Department of Forest Management, Faculty of Forestry, Kasetsart University, Thailand

Abstract

The research objective was to investigate effects of climate variability on monthly growth ofAglaia odoratissimaandHydnocarpus ilicifoliawhich are the dominant tree species in lower canopies ofdry evergreen forest and generally found at the Sakaerat Environmental Research Station (SERS),northeastern Thailand. For one year of the investigation, monthly data of tree leaf phenology, inside bark

diameter (IBD) and outside bark diameter (OBD) increments were examined. These data were related tosoil moisture content and climatic data of monthly rainfall, temperature and relative humidity. The resultsshowed that leaf phenology ofA. odoratissimaandH. ilicifoliaillustrated leaf maturation throughout theyear, while young leaves were abundant in the rainy season and leaf abscission was rarely foundthroughout the year. The IBD increments of these species on transverse surfaces could be detectedthroughout the year and the most rapid increments were detected in the rainy season, while OBDincrements of both species shrank in the dry season and swelled in the rainy season. Using path analysis(PA), climate variability was found to be significantly related to leaf phenology of A. odoratissimaandIBD increments ofH. ilicifolia; and it was also significantly related to OBD increments of both species.

Keywords:Leaf phenology / Outside bark diameter/ Path analysis /Inside bark diameter

1. IntroductionForests are extremely responsive to

climate change and variability; it has been shownby observations from past research, experimentalstudies, and simulation models (Andreu et al.,2007; Nitschke and Innes, 2008; Polgar andPrimack, 2011). Tree-ring analysis (calleddendrochronology) is widely and successfullyused to explain long-term climatic effects of treegrowth especially in the temperate and borealregions (Garcia-Suarez et al., 2009; Trouet et al.,2013). The dendrochronologists attempted toextend their research to the tropical andsubtropical regions due to increasing demand for abetter understanding of global climate dynamicsassociated with the gap of the paleoclimatic

information in the tropics (Buckley et al., 1995;Pumijumnong et al., 1995; DArrigo et al., 1997;Pumijumnong and Wanyaphet, 2006;Pumijumnong and Eckstein, 2011). However, thestudy of climate-growth response using thetechnique of tree-ring analysis is still limited andthe question of which tree species form annualrings and respond to climate are not yet solved.Additionally, the one criterion of tree selection forclimate-growth studies is focused on dominanttrees with non-suppression by the nearest trees(Fritts, 1976). This traditional procedure in tree-ring analysis is suitable for distinct annual ringspecies with the upper canopy but is not

appropriate for indistinct and non-distinct annualring species with the lower canopy. Therefore, in

this paper, we investigated monthly growth of twoevergreen species within distinct annual ringformation, namely Aglaia odoratissima andHydnocarpus ilicifolia, in order to understand theeffects of climate variability on their growth in thelower canopy. The study can be applied to confirmthe principle of dendrochronology in tree selectionfor climate-growth relationship and climaticreconstruction analysis. Additionally, in forestmanagement, the study illustrates the benefit ofthe emergent and upper forest canopy in localclimatic stabilization which is suitable for treespecies in the lower canopy growing throughoutthe year.

2. Methodology2.1 The Study SiteThe study site was located at the dry

evergreen forest of the Sakaerat EnvironmentalResearch Station (SERS) in Udomsap Sub-district,Wang Nam Khiao District, Nakhon RatchasimaProvince, northeastern Thailand (Figure 1). TheSERS occupies an area of 80 km on the edge ofthe Khorat Plateau, at longitude 101 51 E andlatitude 14 30 N. It rises from 250 m above meansea level to 762 m at the top of its highestmountain. The dry evergreen forest has a denseand three-storey canopy which is dominated bythe upper canopy species ranging from 23-38 m

high such as Hopea ferrea, Lagerstroemiaduppereana, Irvingia oliveri and Shorea


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2 Kritsadapan Palakit et al./ Research Article: 1-12

henryana. The majority of the lower canopyspecies ranging from 16-22 m high includeWalsura trichostenon, Memecylon caeruleum,Hydnocarpus ilicifolia, and Aglaia spp.The lowest layer is 4-14 m high tree species suchas Memecylon ovatum, Ixora ebarbata, and

Randia wittii (Thompson and Landsberg, 1975;Lamotte et al., 1998).

The climatic condition at SERS istropical with no occurrence of frost. The winter iscool and dry, while the summer is hot and humid.

Figure 1: The study site at Sakaerat Environmental Research Station (SERS)

Mean maximum temperature is 35 C, and meanminimum temperature is 16 C. The wet seasongenerally occurs from May to mid-October, withrainfall peaks in May and September. The averageannual precipitation is 1,200 mm. Climatic dataduring the investigated periods derived from

SERS are shown in Figure 2. Gray band in Figure2a indicated dry period in November 2009-February 2010.

2.2 MethodsTwo dominant species of the lower

canopy, namely A. odoratissima (code AO01-AO06) and H. ilicifolia (code HI01-HI06) wereselected. Six trees of each species which were;healthy, located on good drained area, and hadsymmetrical crowns and straight trunks werestudied. Each tree location was marked usingGlobal Positioning System (GPS). Leaf phenologyincluding leaf flushing (LF), mature light greenleaves (MLL), mature dark green leaves (MDL)and leaf abscission (LA),was routinely

investigated once a month for a year in September

2009 till September 2010 using visual estimateswith binoculars (OBrien et al., 2008; Vitasse etal., 2009). The observer scored 0 for a bare crown,1 for > 020 percent, 2 for > 2040 percent, 3 for> 40-60 percent, 4 for > 60-80 percent and 5 for >80 percent of the crown cover occupied by each

class. Outside and inside bark diameterincrements were other periodic growthinvestigations. Outside bark diameter increment(OBD) at breast height (130 cm) was recordedusing a modified manual band dendrometer with a0.1 mm accurate vernier scale (Figure 3a). Themeasurements of OBD were taken for a year;coincided with the studies of leaf phenology.Inside bark diameter increments (IBD) werestudied using the techniques of cambial woundedmarking (Figure 3b). Once a month, cambialzones of the selected trees were injured in order tomark and investigate the boundaries of monthlygrowth zone and cell differentiation. A Year Later

when the markings were completed,


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Environment and Natural Resources J. Vol 13, No 1, January-June 2015:1-12 3

Figure 2: Meteorological data in August 2009 - August 2010; Rainfall and temperature data(a); Relative humdity and temperature data (b)

all marked points were extracted and woundedblocks were carefully polished using severalgrades of abrasive papers to obtain clean andsmooth wood surfaces until marked wounds wereprominently visible. The distances from thebaseline (first marked point) to each monthlymarked point was measured and calculated formonthly and total wood increments(Siripatanadilok, 1983) using the Velmexmeasuring system to the nearest 0.001 mm

accuracy. Leaf phenological data and monthlywood increments of OBD and IBD were thencorrelated with monthly climatic data of rainfall,temperature, relative humidity and soil moisturecontents using the multiple linear regressiontechnique, namely path analysis (PA), to explainthe effect of climate variability on tree growth inthe lower canopy with multi-collinearity removal(Hu and Bentler, 1999; O'Brien, 2007).

Figure 3: Manual band dendrometer installation (a) and cambial wounded marking (b)


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3. Results

During the investigation period of oneyear, leaf phenology of 6 mature Aglaiaodoratissimashowed leaf flushing (LF) in Augustto November. Mature light green leaf (MLL) wascommonly found associated with LF and abundantin April and May, which was the beginning of therainy season. Mature dark green leaf (MDL) wasabundantly found in every month, except in Apriland May which MLL abundantly found instead ofother leaf pheno-phases. Leaf abscission (LA) wascommonly found in dry period of December toMarch (Figure 4a). In case of 6 mature (Figure4a).

In case of 6 mature Hydnocarpus ilicifolia, LFwas not found during the investigated period. Thesmall amounts of MLL were commonly found inMay to August, while MDL were abundantlyfound in every month, especially in rainy seasonas similar as A. odoratissima. Small amounts ofLA were commonly found throughout theinvestigation periods (Figure 4b).

Outside bark diameter (OBD) at breastheight (130 cm) of all selected A. odoratissimaand H. ilicifolia were monthly measured at thesame time as leaf phenological investigation.

Figure 4: Leaf phenology ofA. odoratissima (a) and H. ilicifolia (b)

Not only direct values of OBD, but

standardized values of OBD increments were alsocalculated using z-score which was the value ofthe element (X) minus by the population mean (X)and divided by the standard deviation (SD). If a z-score was positive or negative, it could interpretthat the direct value was above (greater than) orbelow (less than) the mean, respectively. The z-score formula was shown as below.

z =

(1)

The cumulative OBD and standardizedz-score profile of sixA. odoratissimafrom August

2009 - September 2010 are shown in Figure 5aand 5b. During the investigated periods, the

vernier scales of all installed band dendrometers

were not changed for 5 months in September 2009to January 2010,except A. odoratissima codedAO01 which OBD mildly increased by 0.07 mm.

The OBD of A. odoratissima rapidlyshrank in February 2010 and gently increased inMarch - April 2010, unchanged in May-June2010, and gently increased in July-September2010. The cumulative diameters of all A.odoratissima were averaged, and the averageincrement was 0.024 cm/year.

Based on the cumulative OBD of six H.ilicifolia, as shown in Figure 6a and 6b, bothdirect value and standardized increment dataexhibited dormant growth from September 2009


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Figure 5:Outside (OBD) and inside bark diameter (IBD) increments ofA. odoratissima

till January 2010 as similar as A. odoratissima.The OBD rapidly in May 2010 and gentlyincreased until September 2010. However, thecumulative OBD of H. ilicifolia at the end of theinvestigation period was lower than the baseline

data recorded in August 2009 of 0.045 cm.Inside bark diameter (IBD) increment of

A. odoratissima increased in August 2009 tillSeptember 2010; it averaged 1.56 mm (Figure 5c).The dormant period where monthly woodincrement did not change was found to be inSeptember 2009 after the cambial markings.However, it was difficult to identify this period asa stop in growth or dormant period of A.odoratissimadue to the continuous occurrences ofwood increment in September 2010.

Wood increment stopped in September2009 and was re-stimulated in October 2009,averaging 0.03 mm (1.34% of total wood

increment) and continuously increased to 0.13 mm(8.46%) in both of November and December

2009. Wood increment gradually declined andfluctuated from 0.03 till 0.09 mm in January tillJune 2010. Total wood increment in these sixmonths was 0.39 mm (24.89% of overallincrement). In July till September 2010, wood

increment rapidly increased to 0.87 mm (55.97%).Although wood increment could be detected inevery month during the investigated periods, thepatterns of wood increments in all trees were notsimilar. It might be assumed that wood incrementsof A. odoratissimawere not dependent on factorsaffecting tree growth, especially climatic factors.Wood increments continuously increased asshown in Figure 5c and 5d, which showed thefluctuation of directly measured and relativemonthly wood increments of A. odoratissimaduring investigation periods.

During September 2009 till theSeptember 2010, the OBD increment of H.

ilicifolia decreased (Figure 6a and 6b), while theIBD increment increased to 1.03 mm (Figure 6c


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and 6d). After cambial marking at the beginningof September 2009, monthly IBD did not changefor 3 months in September November 2009,except a tree coded HI05 whose wood incrementin November 2009 was 0.06 mm (4.31% fromtotal growth of HI105). Unchanged woodincrement in these 3 months seemed to be thedocument period of H. ilicifolia at SERS. Theaverage of wood increments of H. ilicifolia trees,which started in December 2009 and continuously

grew until February 2010, was 0.07 mm (5.59% oftotal growth). In April to June 2010, woodincrement was 0.06 mm (5.29%). It could bestated that, after dormant periods, wood incrementin December 2009 till June 2010 was very slow. InJuly to September 2020, wood growth rapidlyincreased to 0.89 mm (88.49%) and the rate of

wood increment continuously increased. Figure 6cand 6d illustrate the variations of direct andrelative monthly wood increments of H. ilicifoliaduring investigation periods.

In order to calculate effects of climatevariability on tree growth, climatic (rainfall,Tmax, Tmean, Tmin, RH) and soil moisture (SM)variables were converted to 2 components ofclimatic data based on the criteria of Eigen values1.

Based on varimax rotation of principlecomponent analysis (Kaiser, 1958), the rotatedcomponent matrix of 2 non-significantlycorrelated components was observed. The firstcomponent was related to relative humidity,rainfall and soil moisture contents in all soildepths and was re-named as Moisture.

Figure 6: Outside (OBD) and inside bark diameter (IBD) increments of H. ilicifolia


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Figure 7: Grouping of SERS climatic and soil moisture variables into two components of Temperatureand Moisture using principle components analysis

The second factor was related to Tmax,Tmin and Tmean and was re-named as Tempe-

rature. These 2 components (Figure 7) were then

used as independent factors in regression analysisand could explain all selected climatic data(90.83%).

As show in Figure 8, using path analysiswith statistically significant p


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Figure 8: Path diagrams: factors affecting tree growth of A. odoratissima and e1, e2 and e3 indicatederrors in predicting the values of mature dark green leaf, leaf abscission and standardizedoutside bark diameter increment, respectively

Figure 9: Path diagrams: factors affecting tree growth of H. ilicifoliaand e1 and e2 indicated errors inpredicting the values of relative monthly wood increment, and standardized outside barkdiameter increment, respectively.


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4. DiscussionBased on leaf phenology,A. odoratissima

and H. ilicifolia produced abundant leaf crownsthroughout the year, especially in the rainy season(Gutierrez-Soto, 2008). Young leaves wereabundantly found in rainy season and leafabscission rarely found throughout the year. Xiaoet al. (2006) suggested that leaf pheno-phases ofevergreen trees in the tropics were not related tothe seasonality of rainfall. Instead, the fluctuationof leaf phenology may be forced by solar radiationavailability. Williams et al. (2008) studied leafphenology of A. odorata and A. spectabilisgrowing in western Thailand and also found theevergreen and leaf flushed characteristics at thebeginning of the rainy season.

The increments of OBD in A.odoratissima and H. ilicifolia continuouslydeclined from the installed period in the secondhalf of the rainy season to the end of the dry

season (September 2009 - February 2010). In therainy season, OBD rapidly increased from March2010 until the end of the investigated period inSeptember 2010. A. odoratissimaand H. ilicifoliashowed fast increment at the beginning of therainy season but illustrated the unchanged OBD atthe end of the rainy season and the dry season.The extreme shrinkage of OBD occurred for onemonth in February before rainfall in March. TheOBD showed extreme shrinkage in Februarybefore diameter increment gently increased inMarch. Therefore, the suitable period of banddendrometer installation for tree growthmeasurement at SERS should be during the

transition of dry and wet periods at the end ofFebruary. Pelissier and Pascal (2000) stated thatOBD variations originated from hydrostatic forceinducing stem flexibility which depended on thedegree of turgidity or water stress of the tree. Itwas similar to the study of radial stem variationsof several tree species relating water statusincluding precipitation and vapor pressure deficit(Bruning and Burchardt, 2005; Lisi et al., 2008;Oberhuber and Gruber, 2010; Volland-Voigt et al.,2011).

As dendrometer bands were installedover bark in order to continue the measurement oftree growth, changes in phloem differentiation andbark width relating to total OBD increment wasimportant to understand. Several authorssuggested that the swelling and the shrinkage instems mostly occurred in bark, phloem and thezone of cambium, while variation in xylem zonewas insignificantly related (Brough et al., 1986;Daudet et al., 2005; Dobbs and Scott, 1971; Molzand Klepper, 1973; Zweifel et al., 2000). TheOBD variation in Eucalyptus nitens,approximately 40%, had been attributed toshrinkage of phloem and bark (Downes et al.,1999). This information must be taken in toconsideration, when analyzing and interpreting theOBD data

Monthly wood increments of A.

odoratissima and H. ilicifolia derived from

cambial marking studies were found throughoutthe year as similar as leaf phenology. Growth ratesof these species rapidly increased and maximizedin the rainy season. Several authors claimed thatlower rainfall during the winter strongly reducedwood increment, while wet condition during therainy season induced wood increment in both ofbroad leaf and long leaf trees (Lisi et al., 2008;Marcati et al., 2006 and 2008; Singh andVenugopal, 2011). There were few investigationson wood formation in evergreen tree species suchas Sass et al. (1995) who found a continuousprocess in wood formation of Dryobalanopssumatrensisand Shorea leprosula, which was notrelated to seasonality in rainfall and phenology.

Based on the occurrences and abundancesof leaf phenology, monthly wood increments ofthese evergreen species was not significantlyrelated to the occurrences of their leaves, resultssimilar to those of the study of OBrien et al.

(2008).All climatic data did not showed the

significant correlation with the monthly woodincrement of A. odoratissima but moisture wassignificantly related to wood increment of H.ilicifolia. Lamotte et al. (1998) and Marod et al.(2003) explained that H. ilicifolia were co-dominant tree species and A. odoratissima werethe lower layers of crown covers and weresuppressed by other trees. Microclimatic factorsvaried vertically within the forest stand andinduced plant water status changing vertically.Response to increased air temperature, higher leaftemperature in the upper canopy caused a steeper

gradient of vapor pressure from the leaf to the airgreater than the observation from the shadedunderstory and ground vegetation (Thompson andHinckley, 1977). Therefore, the fluctuation ofclimate could directly affect these dominant andco-dominant tree species and explained theinsignificant correlation or the lower effects to thesuppressed or understory trees. However, Borchert(1999) explained that seasonal rainfall drove leafphenology, cambial growth and wood incrementin deciduous trees, but were progressivelyuncoupled in evergreen trees. Lisi et al. (2008)also found trunk increment dynamics of many treespices in a seasonal semi-deciduous forest ofBrazil corresponded to seasonal changes inprecipitation.

Increments of A. odoratissima and H.ilicifolia OBD did not show the positiverelationship with IBD increments. Depending onthe stem shrink and swelling, increased ordecreased temperature, respectively, inducedshrinkage or swelling of OBD and moistureshowed direct and positive effects on OBDincrements. Several studies explained OBDincrements in rainy or wet season related to newcell formation and development, while swellingand shrinkage in dormant period probablydepended on the water content in bark, rather thanchanges in wood increment (Bruning and

Burchardt, 2005; Bruning et al., 2008;


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Deslauriers et al., 2003; Gruber et al., 2009; Jeziket al., 2007). However, to avoid the effect of barkswelling and shrinkage, several studiesrecommended the cambial wounding technique toinvestigate the seasonal rhythms of xylem growthinstead of dendrometer bands (Kuroda andKiyono, 1997; Makinen et al., 2008).

The study of IBD increments for one yearwas design to measure exact wood incrementresponding to monthly and seasonal climatevariability. It could not refer to the long-termassociation of tree growth and climate change. Themeasurement of OBD increment using a modifiedmanual band dendrometer is appropriated to studylong-term periodic growth of indistinct and absentannual ring species which errors from barkcharacteristics are acceptable (Keeland andSharitz, 1993; Schongart et al., 2002).

5. ConclusionsTwo species of the evergreen trees,

including Aglaia odoratissima and Hydnocarpusilicifolia, were selected to study effects of monthlyclimate variability on monthly tree growth for ayear. For leaf phenological studies, these selectedtrees illustrated leaf maturation throughout theyear, while young leaves were abundantly foundin rainy season, and leaf abscission rarely foundthroughout the year. The increments of outsidebark diameters (OBD) of A. odoratissima and H.ilicifolia continuously decreased from theinstallation period in the second half of rainyseason to the end of dry season, and rapidlyincreased at the beginning of the rainy season until

the end of the investigation d period; the insidebark diameter (IBD) increments of both speciesshowed the dormancy period in the dry season.

The climatic factors of moisture andtemperature components derived from thePrinciple Component Analysis (PCA) showedboth direct and indirect effects on monthly woodincrements. The increased moisture directlyinduced IBD increments of H. ilicifolia, but thesetwo climatic components did not show asignificant relationship with IBD of A.odoratissima.Using the technique of path analysis(PA), the fluctuation of leaf phenology of thesetwo species was not significantly related to theincrements of IBD. The OBD increments of H.ilicifoliawere directly and positively related to theincrement of IBD, whileA. odoratisimaOBD wasnot significantly correlated with IBD. Dependingon the shrink and swelling of OBD increments,A.odoratissima OBD was directly related tomoisture and temperature components, and H.ilicifolia OBD showed both direct and indirectrelationships to the fluctuation of all the climaticcomponents.

For further studies, A. odoratissima andH. ilicifoliagrowing in the different sites shall bestudied to confirm the finding of this research assimilar as the study of long-term periodic growthfor climate change investigation. Several

researches suggested that photoperiod involved

photosynthesis, tree growth, and phenologicalchanges (Basler and Korner, 2014; Jackson,2009). Its relationship with tree growth shall be in-depth studies in the next step. Other tree species inthe lower canopy shall also be applied to analyzeclimate-growth relationship and the weatherstation shall be installed to collect the localclimatic data in each selected forest stand.

6. AcknowledgementsThe project was partly funded by a grant

from the Center for Advanced Studies in TropicalNatural Resources (CASTNR) under the NationalResearch University-Kasetsart University (NRU-KU).

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