Trees (2003) 17:244–250

DOI 10.1007/s00468-002-0230-2

ORIGINAL ARTICLE

Oliver D
nisch · Valdinez Ribeiro Montia ·

Josef Bauch

Dendroecological investigations on Swietenia macrophylla King

and Cedrela odorata L. (Meliaceae) in the central Amazon

Received: 16 March 2002 / Accepted: 31 October 2002 / Published online: 23 January 2003

Springer-Verlag 2003

Abstract The width of the increment zones in the xylem Keywords Wood formation · Cambial activity ·
of Swietenia macrophylla King and Cedrela odorata L. Increment zones · Tree-ring analysis · Dendroecology
was investigated by dendroecological methods in a
primary forest near Aripuan¼, Mato Grosso, Brazil
(1009'S, 5926'W). The annual period of cambial cell Introduction
division and its intra-annual variation were determined by
dendrometer measurements of 30 trees of each species. Earlier investigations of the influence of exogenous
Tree-ring width chronologies for Swietenia and Cedrela factors on the wood anatomy of trees (e.g. Denne and
were developed from cross-dated increment curves of 33 Dodd 1981; Vysotskaya and Vaganov 1989; Yasue et al.
out of 47 Swietenia and 51 out of 64 Cedrela trees. 2000) showed that dendrochronological methods are
Simple correlations were computed between the radial useful for historical, ecological, and climatological stud-
growth increment and monthly precipitation for the period ies (Fritts 1976; Schweingruber 1988). Microscopical
1890–2000. In Swietenia, cambium activity occurred (Barnett 1992; Catesson 1994) and physiological inves-
throughout almost the whole year, but in Cedrela it was tigations (Kozlowski et al. 1991), indicate that inter- and
restricted to the rainy period from September of the intra-annual variation in wood formation of many species
previous year to June of the current year. Tree-rings were is predominantly correlated with climatic factors, namely
formed annually in the juvenile and adult wood of temperature (Denne 1971; Briffa et al. 1990, 1995) and
Cedrela, while in Swietenia the annual formation of tree- precipitation (Hughes et al. 1994). Consequently, tree-
rings was restricted to the adult wood. Consequently the ring analysis for dendroecological applications is most
age of the Swietenia trees could be dated by the tree-rings successful for sites with a strong variation of meteoro-
in good approximation, while age dating of the Cedrela logical parameters (Kienast et al. 1987). Most tree-ring
trees was exact. Correlation analyses revealed a signifi- chronologies were, therefore, established especially for
cant relationship between the precipitation at the begin- tree species growing in climates with a strong seasonality
ning and at the end of the growth season and the width of (Fritts 1976).
the increment zones in the adult xylem of Swietenia. In Due to the lack of strong seasonality in the humid
contrast, the width of the growth increment in the xylem tropics, the suitability of tropical trees for dendrochro-
of Cedrela was significantly correlated with the precip- nology has been questioned for many years, even though
itation in March and May of the previous growth period. the pioneering investigations by Coster (1927, 1928)
indicated a regular pattern in wood formation of some
tropical tree species related to a distinct rainfall period-
O. Dnisch · J. Bauch icity in some tropical areas. Studies on Tectona grandis,
Institute of Wood Biology, University of Hamburg, growing under the Asian monsoon climate, showed that
Leuschnerstrasse 91, 21031 Hamburg, Germany wood formation is correlated with the seasonal course of
O. Dnisch ()) precipitation. This allowed the elaboration of tree-ring
Department of Soil Science, Federal University of Parana, chronologies of T. grandis, with applications on histor-
Rua dos Funcionarios 1540, 80035-50 Curitiba, PR, Brazil ical, climatological, and ecological studies (Berlage 1931;
e-mail: [email protected] Pumijumnong et al. 1995; Priya and Bhat 1999). Annual
Tel.: +55-41-3505789
Fax: +55-41-3534144
patterns of wood formation of tropical trees also has been
reported for inundation forests of the Amazon basin
V. R. Montia (Worbes 1988, 1989), swamp forests in the Brazilian
EMBRAPA Amazonia Ocidental, Rodovia AM 010, km 29, Atlantic Rain Forest (Callado et al. 2001) as well as for
69048–970 Manaus, AM, Brazil
 245

some “terra firme” sites (non-inundated sites; Mariaux

1967; Dtienne 1989; Breitspecher and Bethel 1990;
Worbes 1999). In view of the urgent need for the
preservation of tropical rainforests and its sustainable
timber production, we need sound information on the
periodicity of cambial growth of tropical timber trees,
especially those growing on “terra firme” sites with a
seasonal distribution of the rainfall through the year
(Brnig 1996). In a recent study, the cambium activity of
two important timber species, Swietenia macrophylla
(true mahogany) and Cedrela odorata (cedar), was
studied in plantations and in natural forests on the “terra
firme” near Manaus (state of Amazonas) and in the
northern part of the state of Mato Grosso, Brazil (Dnisch
et al. 2002a). It turned out that the formation of increment
zones in the juvenile and adult xylem of Cedrela was
annual while an annual formation of increment zones in
Fig. 1 Mean monthly air temperature (C) and precipitation (mm)
the xylem of Swietenia was restricted to adult trees. This of the study site Aripuan¼, Mato Grosso (1009'S, 5926'W). n=7
led to the conclusion that exact dating of tree-ring records
of these two species by dendrochronological methods
might be possible at least for the adult wood of these selected for dendrometer measurements were distributed over an
species. area of 0.7 km2 and 1.0 km2, respectively.
This study was designed to investigate the possibility
of establishment of tree-ring chronologies (width of Dendrometer measurements
increment zones) of S. macrophylla and C. odorata
growing in the southern part of the Amazon basin. We Strain gauges (accuracy 0.1 mm; UP, Osnabrck, Germany) were
installed in October 1998 at breast height (1.3 m) of 30 Swietenia
determined the growth period of the trees by dendrometer (BHD: 0.27–0.61 m; tree height: ca. 17–31 m) and 30 Cedrela
measurements and established a master chronology for (BHD: 0.38–0.67 m; tree height: ca. 24–29 m) trees. The increase in
each species by dendrochronological methods in order to stem circumference was measured weekly from October 1998 to
determine the age of these species growing in primary November 2001. To determine whether the cambium was dormant
forests. The relationship between monthly precipitation or active during periods without an increase in stem circumference,
samples of the cambial zone were collected with a mini-increment
and the intra-annual growth increment of the trees was borer (Ø 1.4 mm; Bucker et al. 1998; Sack 1998). The samples
also investigated. were fixed with 70% alcohol and embedded in polyethylene glycol.
Cross-sections were cut either by hand or a LKB Historange 2218-
020 microtome. The slides were stained with safranin and astrablue
and studied under the light microscope.
Materials and methods
Study site and experimental trees Wood anatomy and width of increment zones
The study site is located in the indigenous forest reserve “Rio Wood anatomy and the width of increment zones were studied on
Branco” approximately 50 km west of the city of Aripuan¼, Mato polished discs along the north, south, west, and east radius. A total
Grosso, Brazil (1009'S, 5926'W; 190 m above sea-level). The soil of 131 discs from 47 Swietenia trees and 137 discs from 64 Cedrela
is of a xanthic ferralsol type (FAO 1990). The mean air temperature trees were analyzed. Annual growth increments in the adult xylem
is 22.9C, and the annual precipitation is approximately 3,000 mm of Swietenia are marked by terminal parenchyma bands. Annual
(Fig. 1; Lisboa et al. 1976). Whereas the mean monthly temperature growth increments in the xylem of Cedrela are indicated by
and the relative humidity of the air show low intra-annual variation, alternating fibre and vessel bands embedded in paratracheal
the precipitation is distributed unequally over the year. From May parenchyma (Dnisch et al. 2002a). In preparation for the
until October on the study site the precipitation is significantly increment measurements, the markers of each increment zone
reduced compared to the other months of the year (Fig. 1). The were identified by light microscopy of the discs or in microtome
primary forest of the study site has been slightly disturbed by recent cross-sections of the xylem blocks (Reichert, Austria; section
logging activities. The actual density of old growth (BHD >0.4 m) thickness approximately 20 m). The width of the increment zones
Swietenia macrophylla King and Cedrela odorata L. in the forest was obtained by means of a measuring ocular lens (accuracy
reserve (area: 800 km2) is 178 and 243 trees per km2, respectively. 0.1 mm).
Thirty dominant trees each of both S. macrophylla and C.
odorata were used to determine cambium growth periodicity by
dendrometer measurements. For the analysis of the width of the Development of increment chronologies and study
increment zones, stem discs were sampled directly after felling of of the response to precipitation
47 Swietenia (BHD: 0.43–1.07 m; tree height: 24–41 m) and 64
Cedrela trees (BHD: 0.41–0.78 m; tree height: 19–36 m) by In order to study the relationship between the monthly precipitation
logging companies. Discs were sampled of all trees at the bottom of and the width of the increment zones of the Swietenia and Cedrela
the stem and at one to three locations in the upper parts of the stem. trees, ring width chronologies were developed. The increment
The stem discs for tree-ring analyses were sampled in an area of curves were visually cross-dated within and between trees accord-
7.2 km2 (four logging plots) for Swietenia and of 5.3 km2 (three ing to Fritts (1976) using percentage of parallel run and correlation
logging plots) for Cedrela. The Swietenia and Cedrela trees
246

Fig. 2 Relative monthly growth increment (% of the growth triangles Relative monthly growth increment of the tree with the
increment of a single growth period) of a Swietenia macrophylla earliest commencement of cambial growth after a dormant period.
King and b Cedrela odorata L. grown near Aripuan¼, Mato Grosso. Clear triangles Relative monthly growth increment of the tree with
Solid line, Mean (€SD) of 30 trees studied during three growth the latest finish of cambial growth before a dormant period
periods (1998–2001) by dendrometer measurements (n=90). Black

Table 1 Linear regression and correlation coefficient (R2) for the Results
relationship of the monthly precipitation (mm) of the Aripuan¼
study site, Mato Grosso (1009'S, 5926'W) and the monthly
precipitation (mm) of Manaus, Amaznia (0308'S, 5952'W). n=7 Intra-annual cambial growth dynamics of S. macrophylla
and C. odorata
Period Linear regression R2
January 0.7825x+127.18 0.87 The cambial growth dynamics of Swietenia and Cedrela
February 0.6769x+163.93 0.76 showed one period of dormancy per year (Fig. 2a, b).
March 0.7866x+120.70 0.73 Active cambium was found in Swietenia from late August
April 1.1335x+41.04 0.96 until the beginning of August of the following year at the
May 1.2303x+1.29 0.88
June 1.3552x+21.81 0.91 utmost (Fig. 2a), while cambial activity in Cedrela was
July 1.2515x 24.81 0.60 found from mid-September until late May of the subse-
August 0.5664
55.77 0.98 quent year at the utmost (Fig. 2b). In comparison, the
September 0.5056x+7.73 0.59 beginning and the end of cambial growth was more
October 1.1828x+54.36 0.83 variable in Swietenia than in Cedrela. Cambial reactiva-
November 0.6281x+103.29 0.76
December 0.9290x+49.46 0.57 tion after a period of dormancy was found in Swietenia
trees between 18 August and 20 September , while the
end of cambial growth was dated between 26 April and 4
analyses (linear correlation). The synchronized increment curves August indicating a considerable variation between the 30
were standardized by fitting logarithmic regression lines. The investigated trees. Cambial reactivation after a period of
individual standardized increment curves were then averaged to dormancy was found in the Cedrela trees between 11
establish the corresponding chronologies for Swietenia and Cedrela September and 28 September. The end of cambial growth
trees. of the Cedrela trees was dated between 20 May and 15
Simple correlations (third degree regression curves) were
computed between the chronologies and monthly total precipitation June. Compared to Swietenia, the periods of growth and
from January of the previous growth season until August of the dormancy of the Cedrela trees were rather uniform.
current growth season. Meteorological data for “Aripuan¼” are only When calculating the mean monthly growth increment
available for 1974–2001, while long-term meteorological data of the 30 trees each, it became obvious that Swietenia has
(since 1866 with missing values) for the Central Amazon are only
available from meteorological stations in Manaus (0308'S, the highest monthly increment in the middle of the growth
5952'W; approximately 50 m above sea-level). The increment season (January to March; Fig. 2a), while the highest
chronologies of Swietenia and Cedrela were correlated with the monthly growth increment of Cedrela was found at the
precipitation data from Manaus for the years 1890–2000 (13 years beginning of the growth season (September until January;
were excluded due to the lack of meteorological data). To prove the
suitability of the precipitation records from Manaus for the Fig. 2b). The seasonal course of cambial growth in
analyses, the linear correlation between the precipitation in Manaus Cedrela followed a more regular pattern than in Swiete-
and "Aripuan¼" was calculated (Table 1). nia, which was visible in the lower variability of the
relative monthly growth increment between trees and
between the three growth seasons (1998–2001) in Cedrela
compared to Swietenia (Fig. 2a, b).
 247
Fig. 3. a Absolute (1) and
standardized (2) chronologies of
growth increments of (heavy
line) Swietenia macrophylla
King and (light line) Cedrela
odorata L. grown in a primary
forest near Aripuan¼, Mato
Grosso. b The number of trees
included in the chronology is
shown below and reveals the
year of the beginning of the
formation of annual increments
zones in the xylem of the indi-
vidual trees. Each increment
curve was established by fitting
logarithmic regression lines

Table 2 Dendrochronological properties of the master chronolo- common signals within the trees (Table 2). The high
gies for Swietenia macrophylla and Cedrela odorata on the standard deviation of both chronologies indicates a high
Aripuan¼ study site, Mato Grosso (1009'S, 5926'W)
sensitivity of both chronologies. Although parallel run
Master chronology Swietenia Cedrela between the standardized Swietenia and Cedrela
chronologies became apparent in some growth periods,
No. of trees investigated 47 64
No. of trees included 33 51 the statistical data did not give any evidence for
Tree age (years) 90–170 (approx.) 73–157 synchronicity between the two chronologies (parallel
Mean annual radius 3.49€1.82 2.95€0.80 run: 67%, linear correlation: 0.0016).
increment (mm)
Standardised master chronology Swietenia Cedrela
Correlation between trees 0.19 0.24
SD 0.27 0.18 The relationship between precipitation and the width
First order autocorrelation 0.045 0.067 of the increment zones in the xylem of S. macrophylla
and C. odorata

Establishment of a master chronology The monthly climatic records of Manaus show similar
for S. macrophylla and C. odorata tendencies to those near Aripuan¼, while the annual
precipitation of Aripuan¼ is significantly higher than the
The increment curves of 33 out of 47 Swietenia trees and annual precipitation of Manaus. The monthly precipita-
of 51 out of 64 Cedrela trees were included in the tion from January until June and from September until
increment chronologies (Table 2, Fig. 3). Due to the fact December at the Aripuan¼ site exceeds the precipitation
that growth increments in the juvenile wood of Swietenia in Manaus, while the precipitation is lower in July and
(Dnisch et al. 2002a) are not suitable for the determi- August at the Aripuan¼ site than in Manaus (Table 1).
nation of the tree age, while increment zones in the adult This shows that at the study site near Aripuan¼ the annual
wood are suitable for cross-dating and age counting, the course of precipitation shows a more seasonal pattern than
age of the sampled Swietenia trees could only be dated as in Manaus.
a good approximation, while exact dating of the age of the Simple correlations between the tree-ring chronologies
Cedrela trees was possible from all tree-ring records. The and the monthly precipitation records reveals a significant
age of the Swietenia trees varied between approximately influence of the precipitation in January, May, November,
90 and 170 years, while the age of the sampled Cedrela and December of the current growth season on the width
trees varied between 73 and 157 years. The mean annual of the increment zones in Swietenia trees (Fig. 4a). The
growth increment of the Swietenia and Cedrela trees was lowest correlation coefficients were found for the rela-
3.49 and 2.95 mm, respectively. tionship between the precipitation during the period of
The parallel run between the standardized increment dormancy and the annual increment of Swietenia. No
curves included in the two chronologies (Fig. 3) was significant relationship was found between the precipita-
higher than 90%, but the correlation of the standardized tion of the current growth period as well as the
tree-ring records of the Swietenia and Cedrela trees, precipitation during the period of dormancy and the
respectively was not significant, indicating somewhat low standardized growth increment of Cedrela. However, the
248

Fig. 4 Correlation coefficient of third degree polynomial regres- the period from 1890 to 2000. n=99 (missing meteorological data:
sions between the standardized monthly precipitation of Manaus, 13 years). Significant correlations (P<0.05) are indicated by shaded
Amazonia (0308'S, 5952'W) and the standardized growth incre- columns. Precipitation data were standardized by calculating the
ment of a Swietenia macrophylla King and b Cedrela odorata L. residues of the monthly precipitation of each year from the average
grown near Aripuan¼, Mato Grosso. Regressions are calculated for long-term monthly precipitation (%)

width of the growth increment in the xylem of Cedrela cambial activity of the trees independently (Borchert
was significantly correlated with the precipitation at the 1999; Borchert et al. 2002).
end of the previous growth period (March–May; Fig. 4b). The increment chronology developed for Swietenia
showed a higher sensitivity compared to the increment
chronology developed for Cedrela. This was due to the
Discussion higher variation of the absolute increment in the adult
wood of the cross-dated increment curves of the Swietenia
This dendroecological study showed that the cambial trees compared to the Cedrela trees and the higher
activity of old-growth Swietenia and Cedrela grown on autocorrelation within the Cedrela chronology compared
the Aripuan¼ study site in the Amazon follows an annual to the Swietenia chronology. Ecophysiological investiga-
course. This result corresponds to findings in other studies tions on the light, water, and nutrient demand for the net
on the cambial growth dynamics of Swietenia and photosynthesis and the biomass production of Swietenia
Cedrela (Coster 1927, 1928; Fujii et al. 1998; Worbes and Cedrela showed that Swietenia has a lower ecological
1999; Dnisch et al. 2002a). Nevertheless it still has to be amplitude and responds more sensitively to unfavourable
questioned, whether growth increments in the xylem of site conditions than Cedrela (Dnisch et al. 2002b). This
tropical Meliaceae are formed annually as a rule (cf. indicates that the higher sensitivity of the Swietenia
Worbes 1999). Bauch and Dnisch (2000) reported that chronology might be due to the variability of micro site
the formation of increment zones in the xylem of the conditions on the study area. In agreement Pennington et
neotropical Meliaceae species Carapa guinanensis often al. (1981) reported that the natural distribution of
did not correspond to the annual increment of the trees. In Swietenia is restricted to open sites with more fertile
addition, shorter periods of dormancies, the formation of soils and a good water supply, and a growing season of at
non-annual increment zones, and false rings are reported least 8 months per year.
from young Swietenia trees growing in plantations near The correlation analyses of the relationship between
Manaus, where the precipitation is distributed more precipitation and the width of the increment zones in the
equally over the year (Dnisch et al. 2002a). This xylem of Swietenia and Cedrela showed that the different
indicates that the annual pattern of cambial growth of availability of water between the years caused at least a
tropical tree species has to be analysed separately on each portion of the variance of the annual increment found in
site before master chronologies can be established. both species (Worbes 1997; Gnter 2001; Sch
ngart et al.
Dormant phases of the cambium in Swietenia and 2002). The analyses revealed only significant correlations
Cedrela trees became apparent during the moderately dry between the precipitation at the end of the previous
period of the year, while cambial cell divisions were growth season and the width of the increment zones in the
restricted to the more humid months. From these findings xylem of Cedrela. This could indicate that in Cedrela the
it might be concluded that the period of cambial growth of water supply influences especially the formation of food
both species is determined by the water supply. On the reserves, which are mobilized at the beginning of the next
other hand during the drier period day-length is near its growth period (H
ll 1985; Sauter 2000). Investigations on
minimum, which makes it impossible to evaluate the the assimilate balance of Cedrela showed that growth
significance of water supply and day-length to the during the first 2 months after the dormant period
predominantly results from the mobilization of food
 249

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