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Downloaded from Brill.Com09/29/2021 09:58:02PM Via Free Access 102 IAWA Journal, Vol IAWA Journal, Vol. 23 (2), 2002: 101–119 FORMATION OF INCREMENT ZONES AND INTRAANNUAL GROWTH DYNAMICS IN THE XYLEM OF SWIETENIA MACROPHYLLA, CARAPA GUIANENSIS, AND CEDRELA ODORATA (MELIACEAE)* by Oliver Dünisch1, 2, Josef Bauch1 & Luadir Gasparotto2 SUMMARY The pattern of growth increment zones, the cambial growth dynamics and the structural variation in wood formation of Swietenia macrophylla King, Carapa guianensis Aubl., and Cedrela odorata L. (Meliaceae) were in- vestigated in order to understand the relationship of site conditions and sustainable growth in Central Amazonian plantations. Trees were avail- able from 8-, 17-, 23-, and 57-year-old plantations, and from primary for- ests in Manaus (Amazônia), Santarem (Pará), and Aripuanã (Mato Grosso). The wood anatomical structure and the annual increments of 61 Swietenia, 94 Carapa, and 89 Cedrela trees were studied for different tree heights. The curves of annual increments were cross-dated and tested for synchro- nisation. The cambial growth dynamics of up to 52 trees per species were dated by means of dendrometer measurements, monthly labelling by pin- markers, and extracted cambium samples investigated using a microscope. The intraannual course of the growth and structural variation was com- pared with the water supply of the soil and insect attacks (Hypsipyla grandella (Zeller) Lep.). In Swietenia and Carapa parenchyma and vessel bands as well as bands of resin canals were observed. Within the xylem of Cedrela, alternating bands of fibres and vessels surrounded by paratracheal parenchyma were found; bands of resin canals were only occasional. In the juvenile wood of Swietenia and Carapa no synchronization of the increment curves was possible, whereas the increment curves obtained in the juvenile wood of Cedrela showed parallel run in growth. The increment curves obtained in adult wood of Swietenia and Cedrela indicate an annual formation of in- crement zones, whereas the number of increment zones in the xylem of Carapa was approximately 50% higher than the tree age (years) indicat- ing that the growth increments of Carapa also were not annual during the adult phase of growth. *) Dedicated to Prof. Dr. R.W. Kennedy on occasion of his 70th birthday. 1) Institute of Wood Biology, University of Hamburg, Leuschnerstr. 91, D-21031 Hamburg, Germany. 2) EMBRAPA Amazonia Ocidental, AM 010, km 29, 69048-970 Manaus, AM, Brazil. Corresponding address: Prof. Dr. Josef Bauch, Dr. Oliver Dünisch, Institute of Wood Biology, University of Hamburg, Leuschnerstrasse 91, D-21031 Hamburg, Germany [[email protected]; [email protected]]. Downloaded from Brill.com09/29/2021 09:58:02PM via free access 102 IAWA Journal, Vol. 23 (2), 2002 The study of the intraannual growth dynamics of the trees showed that the formation of parenchyma bands in Swietenia is induced by dry periods before a cambial dormancy. The formation of parenchyma bands of Carapa was induced by extremely dry and extremely wet periods before a cambial dormancy, whereas fibre bands in Cedrela were induced by dry periods before a cambial dormancy and the formation of vessel bands embedded in paratracheal parenchyma was induced by wet periods after a cambial dormancy. In addition, insect attack (Hypsipyla grandella) induced lo- cally restricted formation of parenchyma bands and bands of resin canals in Swietenia, Carapa and Cedrela. Key words: Wood formation, increment zones, intraannual growth dynam- ics, water supply, Swietenia macrophylla King, Carapa guianensis Aubl., Cedrela odorata L., Meliaceae. INTRODUCTION In the Central Amazon the demand for wood for the local market and for export is satisfied exclusively from primary forests, which leads to strong exploitation, espe- cially of high quality species such as those of the Meliaceae (comp. Loureiro et al. 1979; Dahms 1989; Mayhew & Newton 1998; Wagenführ 2000). In Brazil, logging of the mahogany species Swietenia macrophylla King is prohibited. Resources of Carapa guianensis Aubl. and Cedrela odorata L. are also seriously declining. In ad- dition, logging in primary forest is often associated with serious negative effects on the ecosystem and a degradation of the area (Lamprecht 1986; Brünig1996). In order to counteract these negative effects special attention is given to cultivating native tree species for high quality timber (Lamprecht 1986; Whitmore 1995; Brünig 1996; Bauch et al. 1999). Particularly on the “terra firme” of the Amazon basin mixed plantations are promising for sustainable growth and the production of fruits and wood (Sanchez 1976; Lieberei & Gasparotto 1998; Dünisch 2001). Successful establishment of plantations needs information on the relationship of site conditions and growth characteristics of the planted trees. Information on growth dynamics and the influence of exogenous impact on the wood formation of most of the native timber tree species of the Amazon is still limited (Baas & Vetter 1989; Jacoby 1989). Investigations on the periodicity of cambial growth and the formation of incre- ment zones of tropical trees revealed different patterns of structural variation in wood formation (e.g. Coster 1927, 1928; Mariaux 1969; Worbes 1988, 1989, 1999; Détienne 1989). Dendroecological studies showed that on the one hand the structural variation in wood formation of tropical trees could be annual and correlated with exogenous input such as water supply and inundation (Worbes 1988, 1999; Pumijumnong et al. 1995). On the other hand wood formation of tropical trees often was not annual and not correlated with exogenous input (Jacoby 1989; Breitspecher & Bethel 1990). With regard to exogenous input the seasonal variation of the water supply is of main importance in the Central Amazon (Dünisch et al. 1999b). The water supply is significant for the turgor of differentiating xylem cells (Larson 1969; Dünisch & Bauch Downloaded from Brill.com09/29/2021 09:58:02PM via free access Dünisch, Bauch & Gasparotto — Formation of increment zones 103 Fig. 1. Study sites for sample collection of 8-, 17-, 23-, and 57-year-old (%) plantation grown and ($) primary forest grown (N) Swietenia macrophylla (Sw), Carapa guianensis (Ca), and Cedrela odorata (Ce) near Manaus-Amazônas, Santarem-Pará, and Aripuanã-Mato Grosso. 1994), the biosynthesis of carbohydrates (Langenfeld-Heyser 1987), and the trans- port of mineral elements (Kramer 1985; Kozlowski et al. 1991). The growth of Meliaceae is often affected by the attack of the insect Hypsipyla grandella (Zeller) Lep., which causes severe damage of the primary meristems of the trees (Whitmore 1976; Newton et al. 1993). According to the concept for the com- partmentalization of wounds proposed by Shigo (1984), wounding of the primary meristem caused by Hypsipyla grandella might also influence wood formation at the cambium. The main objective of this study is to elucidate the relationship of exogenous input and the intraannual cambial growth dynamics of Swietenia macrophylla, Carapa guianensis and Cedrela odorata (Meliaceae) with regard to sustainable growth. MATERIAL AND METHODS Study sites and tree selection For this study we used 61 trees of Swietenia macrophylla King, 94 trees of Carapa guianensis Aubl. and 89 trees of Cedrela odorata L. grown in plantations of known age, and in primary forests (Fig. 1). Samples were collected in the region of Manaus, Downloaded from Brill.com09/29/2021 09:58:02PM via free access 104 IAWA Journal, Vol. 23 (2), 2002 Amazônia (03° 08' S, 59° 52' W), the region of Santarem, Pará (02° 52' S, 54° 45' W), and the region of Aripuanã, Mato Grosso (10° 09' S, 59° 26' W). Manaus region: Samples of 8-year-old plantation grown Swietenia, Carapa, and Cedrela, as well as three primary forest grown Carapa trees were collected on the research station of the EMBRAPA Amazônia Ocidental, 24 km from Manaus. The area is located at approximately 50 m above sea-level with an annual precipitation of about 2,500 mm (min. 110 mm (August), max. 295 mm (February)), a mean air tem- perature of 26.4 °C, and a mean air humidity of 87%. The soil is a poor ferralitic soil (FAO 1990) with a low cation exchange capacity. This study site has been used for interdisciplinary research projects within the Brazilian-German cooperation program “SHIFT” since 1992 (Lieberei & Gasparotto 1998; Bauch et al. 1999). In addition, two 17-year-old Carapa trees were collected from a plantation of the National Re- search Institute of Amazonas (INPA) located 45 km north of Manaus with corre- sponding climatic and soil conditions (Bauch & Dünisch 2000). Santarem region: Samples of eight 57-year-old plantation grown Swietenia, Carapa, and Cedrela were collected from the research station of the EMBRAPA Amazônia Oriental in Belterra, 35 km south of the city of Santarem. In addition, samples of three primary forest grown Carapa trees were collected at the “Floresta Nacional de Tapajos” located approximately 55 km south of Santarem. Climatic and soil condi- tions correspond to the Manaus region, but soil analyses (Dünisch 2001) showed a higher K and Mg content of the soil in the Santarem region compared to the Manaus region (Schroth et al. 2000). Aripuanã region: In the neighbourhood of the city of Aripuanã xylem samples were collected from two 23-year-old plantation grown Swietenia trees planted by small land owners. In addition, eight primary forest grown Swietenia and Cedrela trees were collected from the “Reserva Rio Branco” 50 km west of the city (comp. Loureiro & Lisboa 1979). The experimental area is located approximately 190 m above sea-level with an annual precipitation of approximately 3,000 mm (wettest months February and March, driest months September and October) and a mean tem- perature of 24.9°C. Soil analyses (Dünisch 2001) showed a higher soil fertility of this site compared to the Manaus and the Santarem sites (Lisboa et al. 1976). Microscopical characteristics and width of increment zones Selected microscopical characteristics and the width of increment zones of all trees were analyzed on discs or samples collected with an increment borer in the four car- dinal directions.
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