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
of Swietenia macrophylla King and Cedrela odorata L.
was investigated by dendroecological methods in a
primary forest near Aripuan¼, Mato Grosso, Brazil
(1009'S, 5926'W). The annual period of cambial cell
division and its intra-annual variation were determined by
dendrometer measurements of 30 trees of each species.
Tree-ring width chronologies for Swietenia and Cedrela
were developed from cross-dated increment curves of 33
out of 47 Swietenia and 51 out of 64 Cedrela trees.
Simple correlations were computed between the radial
growth increment and monthly precipitation for the period
1890–2000. In Swietenia, cambium activity occurred
throughout almost the whole year, but in Cedrela it was
restricted to the rainy period from September of the
previous year to June of the current year. Tree-rings were
formed annually in the juvenile and adult wood of
Cedrela, while in Swietenia the annual formation of tree-
rings was restricted to the adult wood. Consequently the
age of the Swietenia trees could be dated by the tree-rings
in good approximation, while age dating of the Cedrela
trees was exact. Correlation analyses revealed a signifi-
cant relationship between the precipitation at the begin-
ning and at the end of the growth season and the width of
the increment zones in the adult xylem of Swietenia. In
contrast, the width of the growth increment in the xylem
of Cedrela was significantly correlated with the precip-
itation in March and May of the previous growth period.
O. Dnisch · J. Bauch
Institute of Wood Biology, University of Hamburg,
Leuschnerstrasse 91, 21031 Hamburg, Germany
O. Dnisch ())
Department of Soil Science, Federal University of Parana,
Rua dos Funcionarios 1540, 80035-50 Curitiba, PR, Brazil
e-mail: oliver_duenisch@gmx.de
Tel.: +55-41-3505789
Fax: +55-41-3534144
V. R. Montia
EMBRAPA Amazonia Ocidental, Rodovia AM 010, km 29,
69048–970 Manaus, AM, Brazil
Keywords Wood formation · Cambial activity ·
Increment zones · Tree-ring analysis · Dendroecology
Introduction
Earlier investigations of the influence of exogenous
factors on the wood anatomy of trees (e.g. Denne and
Dodd 1981; Vysotskaya and Vaganov 1989; Yasue et al.
2000) showed that dendrochronological methods are
useful for historical, ecological, and climatological stud-
ies (Fritts 1976; Schweingruber 1988). Microscopical
(Barnett 1992; Catesson 1994) and physiological inves-
tigations (Kozlowski et al. 1991), indicate that inter- and
intra-annual variation in wood formation of many species
is predominantly correlated with climatic factors, namely
temperature (Denne 1971; Briffa et al. 1990, 1995) and
precipitation (Hughes et al. 1994). Consequently, tree-
ring analysis for dendroecological applications is most
successful for sites with a strong variation of meteoro-
logical parameters (Kienast et al. 1987). Most tree-ring
chronologies were, therefore, established especially for
tree species growing in climates with a strong seasonality
(Fritts 1976).
Due to the lack of strong seasonality in the humid
tropics, the suitability of tropical trees for dendrochro-
nology has been questioned for many years, even though
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-
icity in some tropical areas. Studies on Tectona grandis,
growing under the Asian monsoon climate, showed that
wood formation is correlated with the seasonal course of
precipitation. This allowed the elaboration of tree-ring
chronologies of T. grandis, with applications on histor-
ical, climatological, and ecological studies (Berlage 1931;
Pumijumnong et al. 1995; Priya and Bhat 1999). Annual
patterns of wood formation of tropical trees also has been
reported for inundation forests of the Amazon basin
(Worbes 1988, 1989), swamp forests in the Brazilian
Atlantic Rain Forest (Callado et al. 2001) as well as for

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
the xylem of Swietenia was restricted to adult trees. This
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
species.
This study was designed to investigate the possibility
of establishment of tree-ring chronologies (width of
increment zones) of S. macrophylla and C. odorata
growing in the southern part of the Amazon basin. We
determined the growth period of the trees by dendrometer
measurements and established a master chronology for
each species by dendrochronological methods in order to
determine the age of these species growing in primary
forests. The relationship between monthly precipitation
and the intra-annual growth increment of the trees was
also investigated.
Materials and methods
Study site and experimental trees
The study site is located in the indigenous forest reserve “Rio
Branco” approximately 50 km west of the city of Aripuan¼, Mato
Grosso, Brazil (1009'S, 5926'W; 190 m above sea-level). The soil
is of a xanthic ferralsol type (FAO 1990). The mean air temperature
is 22.9C, and the annual precipitation is approximately 3,000 mm
(Fig. 1; Lisboa et al. 1976). Whereas the mean monthly temperature
and the relative humidity of the air show low intra-annual variation,
the precipitation is distributed unequally over the year. From May
until October on the study site the precipitation is significantly
reduced compared to the other months of the year (Fig. 1). The
primary forest of the study site has been slightly disturbed by recent
logging activities. The actual density of old growth (BHD >0.4 m)
Swietenia macrophylla King and Cedrela odorata L. in the forest
reserve (area: 800 km2) is 178 and 243 trees per km2, respectively.
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
increment zones, stem discs were sampled directly after felling of
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
logging companies. Discs were sampled of all trees at the bottom of
the stem and at one to three locations in the upper parts of the stem.
The stem discs for tree-ring analyses were sampled in an area of
7.2 km2 (four logging plots) for Swietenia and of 5.3 km2 (three
logging plots) for Cedrela. The Swietenia and Cedrela trees
245
Fig. 1 Mean monthly air temperature (C) and precipitation (mm)
of the study site Aripuan¼, Mato Grosso (1009'S, 5926'W). n=7
selected for dendrometer measurements were distributed over an
area of 0.7 km2 and 1.0 km2, respectively.
Dendrometer measurements
Strain gauges (accuracy 0.1 mm; UP, Osnabrck, Germany) were
installed in October 1998 at breast height (1.3 m) of 30 Swietenia
(BHD: 0.27–0.61 m; tree height: ca. 17–31 m) and 30 Cedrela
(BHD: 0.38–0.67 m; tree height: ca. 24–29 m) trees. The increase in
stem circumference was measured weekly from October 1998 to
November 2001. To determine whether the cambium was dormant
or active during periods without an increase in stem circumference,
samples of the cambial zone were collected with a mini-increment
borer (Ø 1.4 mm; Bucker et al. 1998; Sack 1998). The samples
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.
Wood anatomy and width of increment zones
Wood anatomy and the width of increment zones were studied on
polished discs along the north, south, west, and east radius. A total
of 131 discs from 47 Swietenia trees and 137 discs from 64 Cedrela
trees were analyzed. Annual growth increments in the adult xylem
of Swietenia are marked by terminal parenchyma bands. Annual
growth increments in the xylem of Cedrela are indicated by
alternating fibre and vessel bands embedded in paratracheal
parenchyma (Dnisch et al. 2002a). In preparation for the
increment measurements, the markers of each increment zone
were identified by light microscopy of the discs or in microtome
cross-sections of the xylem blocks (Reichert, Austria; section
thickness approximately 20 m). The width of the increment zones
was obtained by means of a measuring ocular lens (accuracy
0.1 mm).
Development of increment chronologies and study
of the response to precipitation
In order to study the relationship between the monthly precipitation
and the width of the increment zones of the Swietenia and Cedrela
trees, ring width chronologies were developed. The increment
curves were visually cross-dated within and between trees accord-
ing to Fritts (1976) using percentage of parallel run and correlation
246

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

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
for S. macrophylla and C. odorata
The increment curves of 33 out of 47 Swietenia trees and
of 51 out of 64 Cedrela trees were included in the
increment chronologies (Table 2, Fig. 3). Due to the fact
that growth increments in the juvenile wood of Swietenia
(Dnisch et al. 2002a) are not suitable for the determi-
nation of the tree age, while increment zones in the adult
wood are suitable for cross-dating and age counting, the
age of the sampled Swietenia trees could only be dated as
a good approximation, while exact dating of the age of the
Cedrela trees was possible from all tree-ring records. The
age of the Swietenia trees varied between approximately
90 and 170 years, while the age of the sampled Cedrela
trees varied between 73 and 157 years. The mean annual
growth increment of the Swietenia and Cedrela trees was
3.49 and 2.95 mm, respectively.
The parallel run between the standardized increment
curves included in the two chronologies (Fig. 3) was
higher than 90%, but the correlation of the standardized
tree-ring records of the Swietenia and Cedrela trees,
respectively was not significant, indicating somewhat low
The monthly climatic records of Manaus show similar
tendencies to those near Aripuan¼, while the annual
precipitation of Aripuan¼ is significantly higher than the
annual precipitation of Manaus. The monthly precipita-
tion from January until June and from September until
December at the Aripuan¼ site exceeds the precipitation
in Manaus, while the precipitation is lower in July and
August at the Aripuan¼ site than in Manaus (Table 1).
This shows that at the study site near Aripuan¼ the annual
course of precipitation shows a more seasonal pattern than
in Manaus.
Simple correlations between the tree-ring chronologies
and the monthly precipitation records reveals a significant
influence of the precipitation in January, May, November,
and December of the current growth season on the width
of the increment zones in Swietenia trees (Fig. 4a). The
lowest correlation coefficients were found for the rela-
tionship between the precipitation during the period of
dormancy and the annual increment of Swietenia. No
significant relationship was found between the precipita-
tion of the current growth period as well as the
precipitation during the period of dormancy and the
standardized growth increment of Cedrela. However, the
248
Fig. 4 Correlation coefficient of third degree polynomial regres-
sions between the standardized monthly precipitation of Manaus,
Amazonia (0308'S, 5952'W) and the standardized growth incre-
ment of a Swietenia macrophylla King and b Cedrela odorata L.
grown near Aripuan¼, Mato Grosso. Regressions are calculated for
width of the growth increment in the xylem of Cedrela
was significantly correlated with the precipitation at the
end of the previous growth period (March–May; Fig. 4b).
Discussion
This dendroecological study showed that the cambial
activity of old-growth Swietenia and Cedrela grown on
the Aripuan¼ study site in the Amazon follows an annual
course. This result corresponds to findings in other studies
on the cambial growth dynamics of Swietenia and
Cedrela (Coster 1927, 1928; Fujii et al. 1998; Worbes
1999; Dnisch et al. 2002a). Nevertheless it still has to be
questioned, whether growth increments in the xylem of
tropical Meliaceae are formed annually as a rule (cf.
Worbes 1999). Bauch and Dnisch (2000) reported that
the formation of increment zones in the xylem of the
neotropical Meliaceae species Carapa guinanensis often
did not correspond to the annual increment of the trees. In
addition, shorter periods of dormancies, the formation of
non-annual increment zones, and false rings are reported
from young Swietenia trees growing in plantations near
Manaus, where the precipitation is distributed more
equally over the year (Dnisch et al. 2002a). This
indicates that the annual pattern of cambial growth of
tropical tree species has to be analysed separately on each
site before master chronologies can be established.
ngart et al.
Dormant phases of the cambium in Swietenia and
Cedrela trees became apparent during the moderately dry
period of the year, while cambial cell divisions were
restricted to the more humid months. From these findings
it might be concluded that the period of cambial growth of
both species is determined by the water supply. On the
other hand during the drier period day-length is near its
ll 1985; Sauter 2000). Investigations on
minimum, which makes it impossible to evaluate the
significance of water supply and day-length to the
the period from 1890 to 2000. n=99 (missing meteorological data:
13 years). Significant correlations (P<0.05) are indicated by shaded
columns. Precipitation data were standardized by calculating the
residues of the monthly precipitation of each year from the average
long-term monthly precipitation (%)
cambial activity of the trees independently (Borchert
1999; Borchert et al. 2002).
The increment chronology developed for Swietenia
showed a higher sensitivity compared to the increment
chronology developed for Cedrela. This was due to the
higher variation of the absolute increment in the adult
wood of the cross-dated increment curves of the Swietenia
trees compared to the Cedrela trees and the higher
autocorrelation within the Cedrela chronology compared
to the Swietenia chronology. Ecophysiological investiga-
tions on the light, water, and nutrient demand for the net
photosynthesis and the biomass production of Swietenia
and Cedrela showed that Swietenia has a lower ecological
amplitude and responds more sensitively to unfavourable
site conditions than Cedrela (Dnisch et al. 2002b). This
indicates that the higher sensitivity of the Swietenia
chronology might be due to the variability of micro site
conditions on the study area. In agreement Pennington et
al. (1981) reported that the natural distribution of
Swietenia is restricted to open sites with more fertile
soils and a good water supply, and a growing season of at
least 8 months per year.
The correlation analyses of the relationship between
precipitation and the width of the increment zones in the
xylem of Swietenia and Cedrela showed that the different
availability of water between the years caused at least a
portion of the variance of the annual increment found in
both species (Worbes 1997; Gnter 2001; Sch
2002). The analyses revealed only significant correlations
between the precipitation at the end of the previous
growth season and the width of the increment zones in the
xylem of Cedrela. This could indicate that in Cedrela the
water supply influences especially the formation of food
reserves, which are mobilized at the beginning of the next
growth period (H
the assimilate balance of Cedrela showed that growth
during the first 2 months after the dormant period
predominantly results from the mobilization of food

reserves (Dnisch and Puls 2002). In contrast to these
findings, significant correlations were found between the
precipitation at the beginning and at the end of the growth
season and the width of increment zones in the xylem of
Swietenia. In view of a strong influence of the water
supply on the period of cambial growth in Swietenia this
finding indicates that the positive effect of high precip-
itation from November until February and in May on the
width of the increment zones in Swietenia is especially
due to a longer period of cambial cell divisions. This
effect has been observed in other species (G
ttsche-Khn Carapa guianensis Aubl. in Central Amazonia. IAWA J
1988; Antonova and Stasova 1993, 1997; Dnisch et al.
1996; Borchert et al. 2002). Moreover, the intrinsic
regulation of cambial growth has also to be taken into
account (Savidge 2000).
Due to the lack of long-term meteorological observa-
tions close to our Aripuan¼ study site, meteorological data
from Manaus were used for the correlation analysis.
However, no old-growth of Swietenia and Cedrela is
available near Manaus to enable a correlation to be
undertaken of the tree growth of both species with
meteorological parameters recorded far from the study
site (Pennington et al. 1981). Although the meteorological
data from Aripuan¼ are strongly correlated with the
meteorological data from Manaus, it might be that the
influence of precipitation on the width of the increment
zones in the xylem of the Swietenia and Cedrela trees is
slightly overestimated, because the precipitation of the
relevant months of the study site is higher than on the
Manaus site (Table 1). In addition, the cambial growth of
Swietenia and Cedrela leads to the expectation of a high
sensitivity towards precipitation due to their root system
in the upper soil (Noldt et al. 2001), while many other
tropical tree species withdraw a high amount of water
from deep soil water storages (Nepstad et al. 1994).
From this study, it was concluded that Swietenia
macrophylla and Cedrela odorata develop distinct annual
increment zones in the xylem, which are suitable for the
development of increment chronologies. With regard to
the urgent need of information on the autecology of
tropical timber trees of the Amazon, dendroecological
approaches offer the chance to study the relationship of
exogenous input and wood formation of Swietenia and
Cedrela by time series analyses.
Acknowledgements We thank the Federal Ministry for Education
and Research (BMBF) and the DLR, Bonn, Germany and the
CNPq/IBAMA, Brasilia, Brazil for financial support within the
German-Brazilian cooperation program SHIFT (Studies on Human
Impacts on Forests and Floodplains in the Tropics). The compre-
hensive cooperation of Dr. L. Gasparotto, EMBRAPA, Manaus, is
very much acknowledged. We thank the Municipio Aripuan¼ for
providing the experimental trees and are also grateful to G.R.
Montia and I.R. Montia for joining in the experiments. We wish
to thank M. Mller, T. Schwarz and R. Rebello for technical
assistance. The critical reading and improvement of the manuscript
by Prof. R. Borchert is especially appreciated.
249
References
Antonova GF, Stasova VV (1993) Effects of environmental factors
on wood formation of Scots pine stems. Trees 7: 214–219
Antonova GF, Stasova VV (1997) Effects of environmental factors
on wood formation in larch (Larix sibirica Ldb.) stems. Trees
11:462–468
Barnett JR (1992) Reactivation of the cambium in Aesculus
hippocastanum L.: a transmission electron microscope study.
Ann Bot 70:169–177
Bauch J, Dnisch O (2000) Comparison of growth dynamics and
wood characteristics of plantation-grown and primary forest
21:321–333
Bucker E, Bues CT, Vogel M (1998) Radial growth dynamics of
spruce (Picea abies) measured by micro-cores. IAWA J 19:301-
309
Berlage HP (1931) Over het verband tusschen de dikte der
jaarringen van djatiboomen (Tectona grandis L.f.) en den
regenval op Java. Tectona 24:939–953
Borchert R (1999) Climatic periodicity, phenology, and cambium
activity in tropical dry forest trees. IAWA J 20:239–247
Borchert R, Rivera G, Hagnauer W (2002) Modification of
vegetative phenology in a tropical semideciduous forest by
abnormal drought and rain. Biotropica 34:381–393
Breitspecher A., Bethel JS (1990) Stem-growth periodicity of trees
in a tropical wet forest of Costa Rica. Ecology 71:1156-1164
Briffa KR, Bartholin TS, Eckstein D, Jones PD, Karln W,
Schweingruber FH, Zetterberg P (1990) A 1400-year tree-ring
record of summer temperatures in Fennoscandia. Nature
346:434–439
Briffa KR, Jones PD, Schweingruber FH, Shiyatov SG, Cook ER
(1995) Unusual twentieth-century summer warmth in a 1000-
year temperature record from Siberia. Nature 376:156–159
Brnig EF (1996) Conservation and management of tropical
rainforests. An integrated approach to sustainability. CAB
International, Wallingford
Calladao CH, da Silva Neto SJ, Scarano FR, Costa CG (2001)
Periodicity of growth rings in some flood-prone trees of the
Atlantic Rain Forest in Rio de Janeiro, Brazil. Trees 15:492–
497
Catesson AM (1994) Cambial ultrastructure and biochemistry:
changes in relation to vascular tissue differentiation and the
seasonal cycle. Int J Plant Sci 155:251–261
Coster C (1927) Zur Anatomie und Physiologie der Zuwachszonen
und Jahresringbildung in den Tropen I. Ann Jard Buitenzorg
37:49-161
Coster C (1928) Zur Anatomie und Physiologie der Zuwachszonen
und Jahresringbildung in den Tropen II. Ann Jard Buitenzorg
38:1-114
Denne MP (1971) Temperature and tracheid development in Pinus
sylvestris seedlings. J Exp Bot 22:362–370
Denne MP, Dodd RS (1981) The environmental control of xylem
differentiation. In: Barnett JR (ed) Xylem cell development.
Castle House, Kent, pp 236–255
Dtienne P (1989) Appearance and periodicity of growth rings in
some tropical woods. lAWA J 10:123-132
Dnisch O, Puls J (2002) Photosynthesis and assimilate allocation
of three plantation-grown Meliaceae of the Amazon. Tree
Physiol 22 (in press)
Dnisch O, Bauch J, Puls, Mller M (1996) Biological and
chemical wood properties of long-term polluted spruce (Picea
abies [L.] Karst.) at high-altitude stands of the Erzgebirge.
Holzforschung 50:497–506
Dnisch O, Bauch J, Gasparotto L (2002a) Cambial growth
dynamics and formation of increment zones in the xylem of
Swietenia macrophylla King., Carapa guianensis Aubl., and
Cedrela odorata L. (Meliaceae). IAWA J 23:101–119
Dnisch O, Azevedo CP, Gasparotto L, Montia GR, da Silva GJ,
Schwarz T (2002b) Light, water, and nutrient demand for the
growth of three high quality timber species (Meliaceae) of the
Amazon. J Appl Bot 76:29–40
250
FAO-UNESCO (1990) Soil map of the world, Revised Legend.
Food and Agriculture Organization of the United Nations,
Rome
Fritts HC (1976) Tree rings and climate. Academic Press, London
Fujii T, Marsoem SN, Fujiwara T (1998) Annual growth rings in
mahogany (Swietenia macrophylla) growing in Java. IAWA J
19:449–450
G
ttsche-Khn H (1988) Bildung und Eigenschaften des Holzes
von Fichten (Picea abies [L.] Karst.] aus Waldschadensge-
bieten. Mitt Bundesforschanst Forst Holzwirtsch 157
Gnter S (2001) kologie und Verjngung von Mahagoni (Swi-
etenia macrophylla King) in Naturwldern Boliviens. Ph.D.
thesis, University of G
ttingen. Cuvillier, G
ttingen Growth Regul 20:52–77
H
ll W (1985) Seasonal fluctuation of reserve materials in the
ngart J, Piedade MTF, Ludwigshausen S, Horna V, Worbes M
trunkwood of spruce (Picea abies (L.) Karst.). J Plant Physiol
117:355–362
Hughes MK, Wu X, Shao X, Garfin GM (1994) A preliminary
reconstruction of rainfall in north-central China since A.D.
1600 from tree ring density and width. Q Res 42:88–99
Kienast F, Schweingruber FH, Brker OU, Schr E (1987) Tree-
ring studies on conifers along ecological gradients and the
potential of single-year analyses. Can J For Res 17:683–696
Kozlowski T, Kramer PJ, Pallardy SG (1991) The physiological
ecology of woody plants. Academic Press, San Diego
Pumijumnong N, Eckstein D, Sass U (1995) Tree-ring research on
Tectona grandis in Northern Thailand. lAWA J 16:385-392
Sack M (1998) Charakterisierung der Holzbildung und des
Zuwachses von Swietenia macrophylla King und Carapa
guianensis Aubl. aus der Familie der Meliaceae unter Planta-
genbedingungen in Zentralamazonien. Master thesis, Hamburg
University
Sauter J (2000) Photosynthate allocation of the vascular cambium:
facts and problems. In: Cell and molecular biology of wood
formation. Savidge RA, Barnett JR, Napier R (eds) BIOS
Scientific, Oxford, pp 71–84
Savidge RA (2000) Intrinsic regulation of cambial growth. J Plant
Sch
(2002) Phenology and stem-growth periodicity of tree species
in Amazonian floodplain forests. J Trop Ecol 18 (in press)
Schweingruber FH (1988) Tree rings. Basics and applications of
dendrochronology. Kluwer Academic, Dordrecht
Vysotskaya LG, Vaganov EA (1989) Components of the variability
of radial cell size in conifers. IAWA Bull 10:417–427
Worbes M (1988) Variety in structure of annual growth zones in
Tabebuia barbata (E. Mey) Sandw., Bignoniaceae, a tropical
tree species from Central Amazonian inundation forests.
Dendrochronologia 6:71-89
Lisboa PLB, Prance GT, Lisboa RCL (1976) Contribui
es ao Worbes M (1989) Growth rings, increment and age of trees in
projeto Aripuan¼. Acta Amazonica 6 Suppl
Mariaux A (1967) Les cernes dans les bois tropicaux africains,
nature et periodicit. Rev Bois For Trop 113:3–14
Nepstad DC, Carvalho CR, Davidson EA, Jipp PH, Lefebvre PA,
Negreiros GH, Trumbore SE, Vieira S (1994) The role of deep
roots in the hydrological and carbon cycles of Amazonian
forests and pastures. Nature 372:666–669
Noldt G, Bauch J, Koch G, Schmitt U (2001) Fine roots of Carapa
guianensis Aubl. and Swietenia macrophylla King: Cell struc-
ture and adaptation to the dry season in Central Amazonia. J
Appl Bot 75:152–158
Pennington TD, Styles BT, Taylor DAH (1981) Flora Neotropica,
Monograph 28. Academic Press, New York
Priya PB, Bhat KM (1999). Influence of rainfall, irrigation and age
on the growth periodicity and wood structure in Teak (Tectona
grandis). IAWA J 20:181–192
inundation forests, savannas and a mountain forest in the
Neotropics. IAWA J 10:109-122
Worbes M (1997) The forest ecosystem of the floodplains. In: Junk
WJ (ed) The Amazonian floodplains: ecology of a pulsing
system. Ecological Studies 126. Springer, Berlin Heidelberg
New York, pp 223–265
Worbes M (1999) Annual growth rings, rainfall dependent growth
and long-term growth patterns of tropical trees from the Forest
Reserve Caparo in Venezuela. J Ecol 87:391–403
Yasue K, Funada R, Kobayashi O, Ohtani J (2000) The effects of
tracheid dimensions on variations in maximum density of Picea
glehnii and relationships to climatic factors. Trees 14:223–229