ACTA

AMAZONICA http://dx.doi.org/10.1590/1809-4392201800961

ORIGINAL ARTICLE

Relevance of wood anatomy and size of Amazonian trees

in the determination and allometry of sapwood area
Luiza Maria Teophilo APARECIDO¹,3*, Joaquim dos SANTOS¹, Niro HIGUCHI¹, Norbert KUNERT¹,²,4
¹ Instituto Nacional de Pesquisas da Amazônia − INPA, Coordenação de Dinâmica Ambiental − CODAM, Av. Andre Araújo 2936, CEP 69060-001, Manaus, Amazonas, Brazil
² Max-Planck-Institut für Biogeochemie, Abteilung Biogeochemische Prozesse, Hans-Knöll Str. 10, 07745 Jena, Germany
3
Present address: School of Life Sciences, Arizona State University, Tempe, AZ, 85281, USA4 Present address: Conservation Ecology Center, Smithsonian Conservation
Biology Institute, Front Royal, VA, 22630, USA
* Corresponding author: [email protected]

ABSTRACT
Hydrological processes in forest stands are mainly influenced by tree species composition and morpho-physiological
characteristics. Few studies on anatomical patterns that govern plant hydraulics were conducted in tropical forest ecosystems.
Thus, we used dye immersion to analyze sapwood area patterns of 34 trees belonging to 26 species from a terra firme forest
in the central Brazilian Amazon. The sapwood area was related with wood anatomy and tree size parameters (diameter-at-
breast-height - DBH, total height and estimated whole-tree volume). Exponential allometric equations were used to model
sapwood area using the biometrical variables measured. Sapwood area traits (cross-section non-uniformity and heartwood
visibility) varied significantly among and within species even though all were classified as diffuse porous. DBH was strongly
and non-linearly correlated with sapwood area (R2 = 0.46, P < 0.001), while no correlation was observed with vessel-lumen
diameter (P = 0.94) and frequency (P = 0.58). Sapwood area and shape were also affected by the occurrence of vessel obstruction
(i.e., tyloses), hollow stems and diseases. Our results suggest that sapwood area patterns and correlated variables are driven
by intrinsic species characteristics, microclimate and ecological succession within the stand. We believe that individual tree
sapwood characteristics have strong implications over water use, hydrological stand upsaling and biomass quantification. These
characteristics should be taken into account (e.g., through a multi-point sampling approach) when estimating forest stand
transpiration in a highly biodiverse ecosystem.
KEYWORDS: ecohydrology; active xylem; tyloses; vessels; Brazilian Amazon

Relevância da anatomia da madeira e dimensões de árvores amazônicas

na determinação e alometria do alburno
RESUMO
Processos hidrológicos de povoamentos florestais são predominantemente influenciados pela composição de espécies arbóreas
e suas características morfo-fisiológicas. No entanto, existem poucos estudos sobre os padrões anatômicos que determinam o
sistema hidráulico de plantas em ecossistemas tropicais. Por isso, nosso objetivo foi o de analisar os padrões da área do xilema
ativo em 34 árvores de 26 espécies de uma floresta de terra firme na Amazônia central por meio de imersão em solução de
corante. A área do xilema ativo foi relacionada a características autoecológicas das espécies, anatomia da madeira e parâmetros
de crescimento (diametro à altura do peito - DAP, altura total e volume total). Equações alométricas exponenciais foram
utilizadas para ajustar a área do xilema às variáveis medidas. Características do alburno (área transversal não-uniforme e
visibilidade do cerne) variaram significativamente entre e dentro de espécies, apesar de que todas as espécies apresentaram
vasos difusos. DAP foi fortemente e não-linearmente correlacionado à área do alburno (R2 = 0,46; P < 0,001), enquanto
diâmetro (P = 0,94) e frequência (P = 0.58) de vasos não apresentaram nenhum grau de relacionamento. O tamanho e forma
do alburno foram afetados pela ocorrência de obstrução de poros (tilose) e troncos ocos. Estes padrões sugerem que a área do
xilema é influenciada por características intrínsicas de cada espécie, microclima e estágio sucessional dentro do povoamento.
Nossos resultados implicam que características individuais de árvores podem fortemente influenciar o transporte de água e,
consequentemente, os processos hidrológicos e a quantificação de biomassa do povoamento. Essas caracteristicas deveriam ser
consideradas (por exemplo, por meio da coleta de amostras da área do xilema ativo ao longo da área transversal) ao estimar-se
a transpiração de uma floresta altamente biodiversa.
PALAVRAS-CHAVE: ecohidrologia; xilema ativo; tilose; vasos; Amazônia brasileira

CITE AS: Aparecido, L.M.T.; Santos, J. dos; Higuchi, N.; Kunert, N. 2018. Relevance of wood anatomy and size of Amazonian trees in the
determination and allometry of sapwood area. Acta Amazonica 49: 1-10.

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AMAZONICA APARECIDO et al. Sapwood area allometry of Aazonian trees

INTRODUCTION modelling (Enquist 2002; Čermák et al. 2004). Thus, to

avoid errors that might lead to over or underestimation of
The Amazonian forest harbors the most biodiverse plant
evapotranspiration and carbon storage, it is important to assess
communities of the world (ter Steege et al. 2013) and maintains
the drivers that might influence the development of sapwood
a significant role in global water recycling and carbon storage
in tropical trees (e.g., wood anatomy and ecological traits), and
within its mostly old, pristine forests (Shuttleworth 1988;
which variables are the most reliable for the indirect estimation
Fauset et al. 2015). These older stands have developed
of sapwood area. Hence, we determined the sapwood area of
very complex vertical canopy structures composed of large
34 trees (26 species) from an old-growth terra firme forest in
trees with vast canopies that shade less robust midstory
the central Brazilian Amazon. Our objective was to determine
strata and an even sparser understory canopy (Amaral et al.
whether wood anatomical variables (vessel-lumen diameter,
2000). These multi-layered, heterogeneous stands influence
frequency and vessel element length) and tree size variables
the microclimatic conditions within the canopy, which
(diameter at breast height, total height and volume) are
will directly affect the development of plant individuals
directly related to xylem development. We used macroscopic
(Kammesheidt 2000; Dietz et al. 2007). The increase in light
and microscopic wood descriptions and allometric equation
intensity, lower relative humidity and higher vapor pressure
modelling for a heterogeneous group of Amazonian tree
deficit resulting from natural and anthropogenic forest gaps
species. We hypothesized that sapwood area allometry is not
can be detrimental for late successional trees and favor the
universal (Meinzer et al. 2001) and will vary as a function of
expansion of early successional species (Kammesheidt 2000).
ecological successional stage, independently of size and species.
Not only will species differ, but reminiscent trees are required
to develop certain physiological strategies to cope with these
changes, such as increasing sapwood area to avoid cavitation- MATERIAL AND METHODS
embolism (Brodersen and McElrone 2013). Study site and tree sampling
Sapwood area plays an important role when estimating The study was conducted in an old growth, lowland forest
whole-tree and stand transpiration (Cermak and Nadezhdina located approximately 60 km northeast of the city of Manaus,
1998). The use of sapwood area as an upscaling parameter has Amazonas state, Brazil, in the Experimental Station for
been widely studied and applied in the field of ecohydrology, Tropical Silviculture ZF-2 (02°37’31.8”S; 60°08’24.7”W)
specifically when coupled with sap flux measurements of the Instituto Nacional de Pesquisas da Amazônia (INPA).
(Meinzer et al. 2004; Horna et al. 2011; Aparecido et al. 2016; The non-inundating, flat terrain is covered with a mature
Kunert et al. 2017; Moore et al. 2017). However, sapwood area and species-rich evergreen terra firme forest. The climate
determination is difficult when using destructive methods, is characterized as hot and humid with an average annual
such as dye-immersed or thermo-imaged fallen tree samples temperature of 25.8°C, relative air humidity between 84 and
(Granier et al. 1994), or non-destructive methods, such as 90% throughout the year, and 2547 mm of annual rainfall
dyed increment cores (Gebauer et al. 2008) or electromagnetic (Kunert et al. 2017).
imaging equipment (Bieker and Rust 2010). The major Thirty-four sample trees were selected at random between
issue underlying non-destructive methods is the clarity of December 2012 and May 2013 close to a vicinal road
sapwood delimitation (e.g., heartwood color distinction; and up to 100 m inside the forest. The tree selection was
Granier et al. 1994; Giothiomi and Dougal 2002) and its based on logistics and tree size criteria (ability to transport
uniformity throughout the cross-section, especially when few and handle log samples). The trees belonged to 26 species
measurement points are taken and extrapolated to all other (Supplementary Material, Table S1), identified by means of
directions (Gebauer et al. 2008). their wood anatomy characteristics and, whenever necessary,
With secondary growth, trees naturally tend to a gradual botanical samples (Aparecido et al. 2015). None of the trees
deactivation of old xylem tissues, although at different rates had buttressed stems reaching above a meter from the ground.
and magnitude (Stewart 1966), due to regulatory processes Log sections of 0.40 m in length were extracted at breast height
that maintain the sapwood area at an optimum metabolic level (1.30 m) for sapwood area determination and wood samples
(i.e., water use and growth; Bamber 1975). However, xylem for macro and microscopic analyses. Total height (including
deactivation can be accelerated through vessel embolism- the crown) was measured in the field after the tree was felled.
cavitation, when trees are fully exposed to high amounts After logs were extracted, the diameter at breast height (DBH)
of radiation, vapor pressure and/or soil moisture deficit was obtained by averaging two measurements from the tree’s
(Machado and Tyree 1994; Schultz and Matthews 1997). cross-section (widest and narrowest diameters, as done when
There are very few studies that describe in detail the external using tree calipers). Volume was estimated using an equation
and internal wood characteristics of tropical tree species and developed for the same area [Volume (m³) = a·DBHb·Htc;
the different growth rates that might affect sapwood area where, a = 3.29; b = 2.16; c = 4.20] (Fernandes et al. 1983).
formation and over or underestimation of its allometric We chose an equation that uses total height and DBH as

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independent variables to evaluate to which extent sapwood

area may be influenced by variables that have different degrees
of dependence to stand density and quality (Machado and
Figueiredo Filho 2006).
Sapwood area determination
Sapwood area was determined through immersion of the
harvested tree logs (Chavarro-Rincon 2009) in an aqueous dye
solution (0.1% indigo carmine) (Andrade et al. 1998; Meinzer
et al. 2001) immediately after harvest. Logs were left immersed
for up to four hours until blue staining was fully apparent on
the top cross-section surface, indicating the active conductive
area. Most of the sections showed signs of staining within a
couple of hours after immersion. Samples that did not show
clear sapwood staining after that time were excluded from
the dataset. Sapwood area (As) was measured using five radii
marked along the cross-section starting from the pith (Nutto
et al. 2012). The first radius was the longest (i.e., the reference
radius – Rn* of Figure 1a). Each radius had two measurements:
total radius [distance from the pith to the bark base (R)]
and heartwood (or inactive area) radius [distance from the
pith to the base of the dye-stained area (r)] (Figure 1a). The
difference between R and r resulted in the sapwood depth
(Supplementary Material, Table S1). Through each radius we
estimated total cross-section area and heartwood cross-section
area and subtracted these variables to achieve total sapwood
area of each tree 𝑅𝑅̅ 𝑟𝑟̅ (Aparecido et al. 2015).
Measuring five different radii guaranteed that variations
within an irregularly shaped cross-section and non-continuous
conductive sapwood area were properly accounted for (Figure
1b). To test this hypothesis, we also estimated sapwood area
using only two radii measurements (longest and shortest)
and compared them with the estimates obtained through the
5-radii method (Supplementary Material, Table S2).
Figure 1. Indigo carmine stained sapwood area of a few log sections. A. Protium
Wood anatomy characterization heptaphyllum sapwood area demarcation and illustration of the 5-radii (Rn)
measurements in a non-uniform cross-section. Rn* indicates the larger, reference
The tree samples were characterized through macroscopic
1

radius and rn the inner radii (equivalent to heartwood area). B. Eschweilera

(wood characteristics for species identification) and wachenheimii (bottom log section) and Cecropia sciadophylla (two log sections
microscopic wood description techniques (histological on top). This figure is in color in the electronic version.
slides and maceration for vessel diameter and frequency
measurements). The macroscopic description followed the lumen-diameter (µm) and frequency (vessels per mm2), and
technical standards recommended by the Pan-American
to record tylose occurrence. Slides were mounted with 18
Technical Standards Commission (COPANT 1974). Dry
wood samples were taken from each tree section and were later μm thick wood sections (American Optical 860 microtome,
shaped into 1.5 × 2.0 × 3.0 cm blocks. Wood structures and AO Scientific Division, Buffalo, NY) previously dehydrated
characteristics such as heartwood color, smell, texture, type in an ethyl series (50, 70, and 100%; Johansen 1940) and
of bark, and parenchyma description were used to determine stained with safranin-fucsin for tissue observations. Vessel
and validate tree species identification. Characteristics
diameter and frequency were measured using an Olympus
observed during harvesting (e.g., exudates, smell, disease,
hollow stem) were also considered for tree identification and Tokyo projector at 100× resolution (TandD VR-71, T&D
as complementary wood characteristics. Corporation, Shimadachi Matsumoto City, Japan) by
Histological microscope slides were prepared for the randomly choosing 25 vessels per tree sample for each
measurement of tangential vessel dimensions, specifically for measurement type (Carlquist 1988).

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Vessel element length (µm) required a chemical maceration and 147% (DR% in Supplementary Material, Table S2).
process to dissociate the vessel elements from fiber and Consequently, cross-sections with highly variable radii lengths
parenchyma (Johansen 1940; Franklin 1945). Small wood resulted in drastically different sapwood area estimates when
slivers were submerged and stored in sealed beakers for considering randomly only one radius. For example, sapwood
48 hours in a maceration solution consisting of 1:4:5 of area was 49% smaller when using the shortest radius of Vantanea
hydrogen peroxide, distilled water and acetic acid. After micrantha (Figure 1b; Supplementary Material, Table S2), with
washing with distilled water, the samples were loosened and an overall average of 34 ± 11% smaller than the reference radius.
dissociated, and then submerged in safranin-fucsin dye. The Wood anatomy analyses showed interspecific differences
vessel elements were then set on microscope slides and 25 in the presence of visible heartwood in the sapwood area
vessel elements were randomly measured using a ruler and the (65% with complete or partial visibility), exudates (26% had
same Olympus Tokyo projector at 100× resolution. All wood some type of exudate), tyloses (50% had partial or complete
anatomy variables were quantified and categorized according apparent vessel obstruction; Figure 2), hollow stem or disease
to COPANT (1974) technical standard classifications for (38%), and/or sapwood area that did not reach the heartwood
wood structure characteristics. This classification allowed a
broader view of the wood property diversity of the sampled
trees (Supplementary Material, Figure S1). Wood samples were
registered and stored in the xylarium of INPA.
Data analyses
The correlations of tree size and wood anatomy variables with
sapwood area were analyzed using Pearson’s correlation coefficient
(r) and linear and exponential regressions to determine variable
relationship and line-curve trend and to develop a practical
allometric equation to estimate sapwood area in a non-destructive
manner for this region and forest type (Supplementary Material,
Table S2). The primary models used were an exponential power
model (Y = aXb) and an exponential decay model (Y = ae(-bX)),
where Y is sapwood area, a is the normalization constant, X is
one of the biometrical or wood anatomy variables and b is an
allometric scale exponent) (Wullscheleger et al. 2001; Meinzer et
al. 2005; Gebauer et al. 2008). Model selection was based on the
coefficient of determination (R2), F-values and P-values (α = 0.05,
0.01 and 0.001) obtained from each model fit. Statistical analyses
were performed with R version 2.6.2 (R Development Core Team
2008) and SYSTAT 12 (SYSTAT Software, Inc.; Richmond, CA).

RESULTS
Sapwood area description
Sapwood area varied between 32 and 951 cm², and the ratio
between sapwood area and cross-section varied between 10%
and 100% (Supplementary Material, Table S1; Figure S1).
Sapwood area depth varied substantially between the measured
radii along the cross-section of most trees. Longest and shortest
radii differed in up to 37% in some trees. Consequently, the
diameter estimated by five radii resulted in only nine trees with
± 1% difference from the diameters estimated by two radii
measurements (Supplementary Material, Table S2). The most
extreme differences between 5-radii and 2-radii estimates were
88% (12% underestimated) and 108% (8% overestimated), Figure 2. Macroscopic view of diffuse vessels in 10× resolution from nine tree
species. A. Cecropia sciadophylla; B. Virola calophylla; C. Protium heptaphyllum; D.
whith an average of 99 ± 12% (DR_Avg% in Supplementary Ocotea neesiana; E. Eschweilera wachenheimii; F. Gustavia speciosa; G. Byrsonima
Material, Table S1). When considering only one of the radii crispa; H. Pouteria caimito; I. Micrandropsis scleroxylon. Panels A, E and I show vessels
to estimate DBH, our estimate differences ranged from 67% obstructed by tyloses. This figure is in color in the electronic version.

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(47%) (Supplementary Material, Table S3). The only characters when above 30 cm, which led to a weaker fit (Figure 3a;
common to all species were diffuse porosity, which is a common Supplementary Material, Table S4). Outliers were included
feature of Amazonian tree species (Ferreira et al. 2004; Marques while performing the regression analyses to maintain a
2008; Moutinho 2008), and the predominant occurrence of representative sample, since ~ 90% of trees in the study area are
solitary vessels (68% of samples), followed by groupings of vessel in DBH classes < 30 cm (Higuchi et al. 2012). Additionally,
multiples of 2 (20%) (Supplementary Material, Figure S1). excluding outliers did not result in a significantly better
model fit for any of the variables analyzed in this study. Total
Allometric relationships height had a slightly weaker fit when compared to DBH, due
DBH had a positive and strong relationship with sapwood to greater variability among trees (Figure 3a-b; R² = 0.33;
area (R2 = 0.46, r = 0.68; P < 0.001). DBH varied widely r = 0.54; P < 0.001). The relationship of tree volume with

Figure 3. Non-linear relationships between sapwood area and tree size and wood anatomy variables. Only significant (P < 0.05) curves represented. A. Diameter
at breast height (DBH, cm); B. Total height (Ht; m); C. Volume estimated from DBH and Ht (Vol, m3); D. Vessel diameter (µm); E. Vessel element length (µm); F. Vessel
frequency (vessels per mm2).

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sapwood area was only slightly stronger than using only DBH found that heliophylle (or pioneer) trees (i.e., plants with
as a predictive variable (R² = 0.48, r = 0.70; P < 0.001). rapid growth under high light intensities) transpire more
Vessel-lumen diameter did not vary widely among the than co-existing late successional trees, specially closer to the
sampled tree species. Although average vessel diameter of forest edge (Horna et al. 2011; Kunert et al. 2015; Kunert
all trees ranged between 59 and 238 μm, 51% of the trees et al. 2017). To compensate water loss, pioneer species may
had diameters of 100 - 200 μm (Figure 3d; Supplementary require a larger sapwood area in order to transport larger
Material, Figure S1), with an average of 127.6 ± 49.2 μm volumes of water. For instance, one of our sampled species,
(see Supplementary Material, Table S1 for individual tree Cecropia sciadophylla, is a common pioneer tree in Amazonian
averages). Vessel-lumen diameter had the worst fit among the forests, and its sapwood area occupied > 80% of the cross-
variables tested and, thus, had no influence on sapwood area section in all three sampled specimens. In contrast, in Ocotea
variation (R2 = 0.001; r = 0.03). Similarly to diameter, vessel neesiana, a late successional tree (Paula et al. 2004), sapwood
frequency also varied little among trees. A third (34%) of area occupied only 30% of the cross-section, which suggests
the trees had few vessels per mm² (Figure 3f; Supplementary that inherent successional stage might be one of the factors
Material, Figure S1b), and there was no correlation between that influence the size of the active xylem area and, possibly,
vessel frequency and sapwood area (R2 = 0.01; r = -0.10). other physiological adaptations, such as leaf shedding.
Within-tree average vessel frequency varied between 2.5 and The weaker relationship between total height and sapwood
29.2 vessels per mm², with an overall average of 9.1 ± 6.5 area, when compared to DBH, may be attributed to vertical
vessels per mm² (see Supplementary Material, Table S1 for tree growth dynamics not being related to horizontal tree
individual tree averages). Contrary to all other variables, vessel growth (in diameter), or to the ability of generating new
frequency decreased with sapwood area increase (Figure 3f; vessels in its cross-section (Brodersen and McElrone 2013).
Supplementary Material, Table S1). Average vessel element Although a weaker predictor than DBH, total height was
length varied between 148 and 1177 μm, with an average significantly correlated with sapwood area, which suggests that
of 555 ± 202 μm. Most vessel element lengths (54%) were sapwood growth may be related to tree height to some extent,
shorter than 300 μm (Supplementary Material, Figure i.e., enabling vertical growth expansion and maintenance
S1). Unexpectedly, vessel element length had the strongest (Kostner et al. 2002; Gebauer et al. 2008). Even though
relationship with sapwood area (R² = 0.65, r = 0.69; P < 0.001) an increase in photosynthetic activity due to greater solar
(Figure 3e; Supplementary Material, Table S1). exposure may lead to tree growth (Andrade et al. 1998), it
does not necessarily mean that total height will increase in
DISCUSSION the same manner as sapwood area. Thus, if tall trees are not
assimilating the amount of carbon required to maintain their
Tree size
size/biomass, they may reduce sapwood area to avoid water
Our results showed that vessel element length was the strongest loss. Tall, younger trees are likely to have more exposure to
predictor of sapwood area among the tested variables, followed solar radiation, which will increase their water use. In this
by volume and DBH. Of the three variables, DBH is the easiest way, the combination of short and tall young trees in a forest
to measure, and, hence, more practical to estimate sapwood stand will result in a high variability of sapwood area and,
area. Even though DBH was not as strongly correlated to consequently, in a weaker model fit. Taller, more exposed,
sapwood area as expected, other studies have considered DBH trees also have a greater disadvantage during the dry season,
as the best variable for sapwood area estimation (Gebauer et al. when high vapor pressure deficits (VPD > 1 kPa) may result
2008; Moore et al. 2010; Horna et al. 2011). The assumption in significant loss of hydraulic conductivity due to cavitation
of a strong relationship between DBH and sapwood area is and embolism (Tyree and Sperry 1989; Gebauer et al. 2008).
based on the hypothesis that water consumption increases Therefore, some trees can maintain a smaller sapwood area
with tree size (Meinzer et al. 2005). To allocate the necessary to avoid embolism and cavitation (Cochard and Tyree 1990;
amount of carbon for biomass production (e.g., increase in Christman et al. 2012), specially those with medium to large-
trunk, branch and leaf biomass), trees need to maximize their sized vessels (Tyree et al. 1994).
water absorption through the increase of their sapwood area
(Gebauer et al. 2008). Sapwood area
The weaker than expected model fit of the relationship The sapwood areas determined in this study (32 − 951 cm2)
between sapwood area and DBH may have been due to were similar in range to those observed in other Neotropical
a wide range of intrinsic differences within each tree. For lowland and montane forests, with extremes ranging between
example, species belonging to different successional stages 20 and 4000 cm2 (Granier et al. 1996; Anhuf et al. 1999;
transpire at different rates and, consequently, alter the size of Motzer et al. 2005; Parolin et al. 2008; Aparecido et al.
their sapwood area according to their water use (Horna et al. 2016; Kunert et al. 2017; Moore et al. 2017). The significant
2011; Kunert et al. 2017). Other studies in tropical regions heterogeneity in sapwood area patterns in our samples suggests

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that certain stem anatomical traits may affect the results of is the case of heartwood wounds that develop after cambium
allometric modeling and determination of sapwood area. For injuries (Bamber 1975). This can result in significant errors
example, the fact that in 47% of our sampled trees the active in hydrological measurements, such as sap flux, when probes
xylem did not reach the inferred and visible heartwood area are inserted in partially deactivated sapwood (Clearwater et al.
suggests that sapwood area can be overestimated by some 1999; McDowell et al. 2002; Gebauer et al. 2008).
methodologies, such as the determination of sapwood area
solely on heartwood coloration (Granier et al. 1996; Gartner
Wood anatomy
2002; Giothiomi and Dougal 2002). Heartwood coloration Our results indicated clearly that the anatomical characters
may therefore not be suitable as a sapwood estimation method analyzed were not responsible for sapwood area growth or
for most Amazonian species. Additionally, most of the trees deactivation. Unlike sapwood area, the number and tangential
sampled did not have a uniformly, round-shaped stem, which dimensions of vessels are not altered due to seasonal variations
would also over or underestimate sapwood area based on a or ecological traits (Sterck et al. 2008). These variables can
single core extraction. be altered only through certain genetic mutations or as an
adaptation to local microclimates at different altitudes and/or
Another possible source of sapwood area (and transpiration)
latitudes (Fisher et al. 2007). Since our samples were collected
overestimation was the unexpected presence of hollow
in a small area on a flat terrain, the occurrence of genetic or
stems and tyloses, which are rarely acknowledged in other
geographical variation is unlikely. Likewise, Noshiro and Baas
ecohydrological studies (James et al. 2003). Hollow stems
(2000) concluded that insufficient distance between forest
were found in 20% of our trees, but can affect 50-75% of
stands explained the lack of significant variation in vessel
trees and account for up to 83% of the stem’s cross section
diameter and frequency in relation to DBH and sapwood
in other tropical sites (Bakshi 1960; Heineman et al. 2015).
area between stands.
Other studies in the Brazilian Amazon found hollow stems
in 30-40% of sampled trees (Apolinario and Martius 2004; The surprisingly strong relationship observed between
Eleuterio 2011). Hollow stem incidence is higher in larger vessel element length and sapwood area may be attributed to
trees (DBH > 50 cm) (Heineman et al. 2015), which may the local microclimate (Baas 1982). Through paleoecology,
explain why hollow trees were relatively less frequent in our Baas (1982) associated smaller vessel element length to
sample than in the aforementioned studies. Hollow stems accentuated seasonality of the local climate (e.g., elevated
can lead to 7% less biomass in a forest stand, affecting carbon precipitation rates or rigorous drought periods). Considering
stock quantification (Heineman et al. 2015). Hence, without that the Amazon region does not have a strong dry season, and
proper sapwood determination methods, there is increased temperatures and precipitation are predominantly constant
uncertainty as to what degree factors such as heart rot or over the years, we can assume that interspecific variation in
termite and ant colonies have affected the heartwood. Since vessel element lengths is due to different water use strategies
hollow trunks are not exclusive to a certain tree species or and the effect of microclimatic variation within a forest stand,
age, and are typically associated with tree size, habitat and such as the occurrence of forest gaps (Kunert et al. 2015),
soil conditions (Heineman et al. 2015), it is only possible to which may also directly affect sapwood area size (Copenhaver
non-destructively predict whether a certain tree cross-section and Tinker 2014). Although being an evolutionary insightful
is whole by employing specialized equipment, such as laser parameter, vessel element length is extremely difficult to
scanners, x-rays or electromagnetic resonance (Nikolova et measure, for it requires tree harvesting and laborious and
al. 2009; Bieker and Rust 2010). However, these methods expensive laboratory analyses, which makes it unpractical as
are expensive, laborious and practically inviable to use in the a variable to estimate sapwood area.
tropics (due to humidity, and the number and size of samples
needed). Increment borers may provide a less laborious and CONCLUSIONS
little-invasive alternative to assess trunk hollowness, yet, as Our study sheds light on the importance of considering wood
previously mentioned, non-uniformity of tree stems may lead anatomy traits and tree size variations when estimating sapwood
to errors if only one core is extracted. area in a heterogeneous tropical forest stand. The frequent
Tyloses have been reported to occur in some specific occurrence of tyloses and hollow stems can potentially affect
tree families, such as Lecythidaceae (Moutinho 2008) and the accuracy of allometric modeling and the estimation of
Salicaceae (Mota et al. 2017). Considering that half the trees ecohydrological parameters. Neglect in accounting for hollow
from this study presented some type of vessel obstruction, we stems may lead to significant overestimation of aboveground
can assume that tyloses are a major contributor to sapwood tree biomass in the Amazon, while not accounting for the
area deactivation or partial obstruction of water and nutrient occurrence of tyloses (specially in Lecythidaceae) may lead
pathways in the active xylem of Amazonian trees (Bamber to overestimation of stand transpiration. Even though xylem
1975; Kitin et al. 2010). These deactivated portions may not vessels are important components of sapwood area, the
be homogeneous and can occur in any portion of the stem, as variables that mainly affected this parameter were related to

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AMAZONICA APARECIDO et al. Sapwood area allometry of Aazonian trees

tree size, ecological succession and location inside the stand canopy conditions in a Costa Rican premontane tropical forest.
(canopy exposure). Our results suggest that sapwood area Hydrological Processes, 30: 5000-5011.
allometry may not be constant in Amazonian forests, but vary Apolinario, F.E.; Martius, C. 2004. Ecological role of termites )
with tree diameter, species autoecology and stand dynamics. Insecta, Isoptera) in tree trunks in central Amazonian rain forests.
Future studies aiming at sapwood area estimation in the Forest Ecology and Management, 194: 23-28.
Amazon should take into account these aspects when sampling Baas, P. 1982. Systematic, phylogenetic, and ecological wood
highly heterogeneous stands. Our findings also can improve anatomy—history and perspectives. In: P, Baas. (Ed.), New
sapwood area determination through non-destructive multi- perspectives in wood anatomy. Forestry Sciences. Springer
point sampling, especially when dealing with non-uniform Science & Business Media, Dordrecht, p.23-58.
tree stems. Differential adaptability to microclimates will Bakshi, B.K. 1960. Heart rot of Sal in India. Phytopathological News,
affect water use rates of trees, and consequently their sapwood 6: 45-46.
area size. Our study provides useful insights for future studies Bamber, R.K. 1975. Heartwood, its function and formation. Wood
on Amazonian tree water use and carbon stock estimation. Science and Technology, 10: 1-8.
Bieker, D.; Rust, S. 2010. Non-destructive estimation of sapwood
and heartwood width in Scots Pine. Silva Fenica, 44: 267-273.
ACKNOWLEDGMENTS
Brodersen, C.R.; McElrone, A.J. 2013. Maintenance of xylem
We would like to thank CAPES (Coordenação de Network Transport Capacity: A Review of Embolism Repair in
Aperfeicoamento de Pessoal de Nível Superior) for financing Vascular Plants. Frontiers in Plant Science, 4: 1-11.
the first author’s master’s degree, CNPq (Conselho Nacional de
Carlquist, S. 1988. Comparative wood anatomy: Systematic, ecological,
Desenvolvimento Científico e Tecnológico) for their program and evolutionary aspects of dicotyledon wood. Wood Science. 1st.
INCT- Madeiras da Amazônia, that provides financial support ed. Springer Verlag, Berlin, 448p.
for the Forest Management Laboratory and ZF-2 Reserve Čermák, J.; Kučera, J.; Nadezhdina, N. 2004. Sap flow measurements
infra-structure, and INPA/CODAM (Instituto Nacional de with some thermodynamic methods, flow integration within
Pesquisas da Amazônia/Coordenação de Dinâmica Ambiental) trees and scaling up from sample trees to entire forest stands.
for their infra-structure, faculty and staff. We would also like Trees, 18: 529-546.
to thank all the workers at ZF-2 Forest Management Reserve Cermak, J.; Nadezhdina, N. 1998. Sapwood as the scaling parameter:
for their support in fieldwork operations. Many thanks to defining according to xylem water content or radial pattern of
Jorge Freitas, Francisco José Vasconcellos (INPA Xylotheque) sap flow? Annals of Forest Science, 55: 509-521.
and Claudete Catanhede do Nascimento (INPA Laboratory Chavarro-Rincon, D. 2009. Tree transpiration mapping from upscaled
of Wood Artifact Engineering) for giving access to laboratory sap flow in the Botswana Kalahari. Doctoral thesis, University of
equipment and guidance throughout the wood anatomy Twente, Netherlands, 159p.
analyses. Finally, special thanks for the constructive feedback Christman, M.A.; Sperry, J.; Smith, D.D. 2012. Rare pits, large
of anonymous reviewers and the editor for thoroughly revising vessels and extreme vulnerability to cavitation in ring-porous
this manuscript. This present manuscript is based on partial tree species. New Phytologist, 193: 713-720.
data from the first author’s master’s thesis. Clearwater, M.J.; Meinzer, F.C.; Andrade, J.L.; Goldstein, G.;
Holbrook, N.M. 1999. Potential errors in measurement
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This is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use,
distribution, and reproduction in any medium, provided the original work is properly cited.

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SUPPLEMENTARY MATERIAL (only avaliable in the electronic version)

APARECIDO et al. Relevance of wood anatomy and size of Amazonian trees in the determination and allometry of sapwood area

Table S1. Size and characteristics of wood anatomy of all sampled trees. Values are the mean ± standard deviation, when more than one measurement was taken.

Sapwood- Avg vessel

Sapwood Basal Avg vessel Avg vessel
Height Sapwood to-basal frequency
Family/species DBH1 (cm) depth2 area diameter element length
(m) area (cm2) area ratio (vessels/
(cm) (cm²) (μm) (μm)
(%) mm2)
ANNONACEAE
Bocageopsis multiflora (Mart.) R. E. Fries 26.7 26.8 ± 1.1 2.8 ± 0.2 209.9 545.7 38.5 122.0 ± 32.9 6.6 ± 2.4 463.0 ± 107.0
Duguetia flagellaris Huber 13.0 9.4 ± 0.2 4.6 ± 0.5 66.5 66.5 100.0 75.2 ± 24.0 11.8 ± 3.1 448.4 ± 35.1
BIGNONIACEAE
Jacaranda copaia (Aubl.) D. Don 25.1 20.4 ± 0.2 2.5 ± 0.3 137.1 305.4 44.9 151.6 ± 43.6 3.3 ± 1.3 595.9 ± 382.4
BORAGINACEAE
Cordia silvestris Fresen.3 19.8 20.6 ± 4.3 2.1 ± 0.8 124.1 333.3 37.2 105.2 ± 40.1 5.1 ± 3.0 331.6 ± 74.8
BURSERACEAE
Protium heptaphyllum (Aubl.) March. 11.2 8.8 ± 0.8 1.6 ± 0.4 36.8 63.6 57.9 80.4 ± 27.2 22.0 ± 4.1 428.8 ± 109.2
EUPHORBIACEAE
Mabea caudata Pax & K. Hoffm. 16.0 11.5 ± 1.2 1.7 ± 0.4 47.5 89.6 53.0 134.4 ± 28.4 5.6 ± 2.2 701.5 ± 206.3
Mabea caudata Pax & K. Hoffm. 17.2 17.5 ± 0.7 3.4 ± 0.8 152.8 244.4 62.5 126.8 ± 38.2 4.9 ± 1.6 681.2 ± 163.6
Micrandropsis scleroxylon W. Rodr. 28.0 28.1 ± 3.4 3.5 ± 0.7 259.4 586.2 44.2 176.4 ± 51.2 5.1 ± 2.1 575.8 ± 200.5
FABACEAE - CAESALPINOIDEAE
Sclerolobium chrysophyllum Poepp. 25.1 20.9 ± 0.2 2.4 ± 0.4 143.2 350.3 40.9 212.4 ± 29.8 3.1 ± 1.5 482.1 ± 99.6
Tachigali paniculata Aubl. 14.2 8.4 ± 0.5 3.1 ± 0.4 53.9 58.1 92.7 125.6 ± 38.4 7.8 ± 3.4 381.7 ± 97.0
HUMIRIACEAE
Vantanea micrantha Ducke 21.7 20.5 ± 0.5 4.7 ± 2.7 241.9 349.0 69.3 176.4 ± 37.3 3.3 ± 1.2 759.0 ± 143.7
Vantanea micrantha Ducke 23.0 40.6 ± 2.3 1.0 ± 0.3 117.6 1160.5 10.1 136.4 ± 21.0 4.7 ± 1.4 818.0 ± 176.2
LAURACEAE
Ocotea neesiana (Miq.) Kosterm. 20.0 18.8 ± 0.4 1.5 ± 0.1 76.6 253.3 30.3 100.0 ± 30.0 17.6 ± 4.5 578.4 ± 161.3
LECYTHIDACEA
Eschweilera fracta R. Knuth 18.5 21.9 ± 0.9 4.2 ± 0.5 222.2 351.7 63.2 113.2 ± 34.2 6.9 ± 2.4 410.8 ± 119.1
Eschweilera odora (Poepp. ex O. Berg) 22.7 20.7 ± 1.2 2.6 ± 0.4 83.8 334.9 25.0 130.4 ± 42.1 5.9 ± 3.2 470.8 ± 131.5
Eschweilera wachenheimii Sandwith 12.0 10.5 ± 0.7 4.5 ± 0.4 78.4 79.2 99.0 126.0 ± 46.2 8.0 ± 2.9 432.7 ± 142.2
Eschweilera wachenheimii Sandwith 22.5 18.1 ± 0.2 4.2 ± 0.7 186.7 265.9 70.2 112.0 ± 31.4 6.9 ± 2.6 406.0 ± 86.3
Eschweilera wachenheimii Sandwith 19.0 20.8 ± 1.1 1.8 ± 1.8 91.6 259.0 35.4 129.2 ± 48.6 6.4 ± 2.4 443.1 ± 119.6
Eschweilera wachenheimii Sandwith 18.0 16.0 ± 0.0 0.9 ± 0.2 44.6 202.1 22.1 139.2 ± 33.3 5.8 ± 2.2 543.6 ± 105.9
Eschweilera wachenheimii Sandwith 18.0 12.9 ± 0.5 0.8 ± 0.1 31.8 136.0 23.4 103.2 ± 28.1 9.8 ± 4.1 395.2 ± 80.4
Gustavia speciosa (Kunth) DC. 19.0 17.9 ± 0.5 3.0 ± 0.4 133.9 236.7 56.6 66.0 ± 17.3 29.2 ± 4.6 495.2 ± 77.9
MALPIGHIACEAE
Byrsonima crispa A. Juss 16.0 15.9 ± 2.3 2.2 ± 0.6 95.8 208.2 46.0 93.6 ± 19.6 17.7 ± 4.6 857.2 ± 164.9
MORACEAE
Brosimum utile (Kunth) Pittier 27.0 26.7 ± 1.9 2.7 ± 0.8 203.0 544.1 37.3 118.0 ± 28.7 6.5 ± 2.4 452.5 ± 85.8

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Table S1. Continued.

Sapwood- Avg vessel
Sapwood Basal Avg vessel Avg vessel
Height Sapwood to-basal frequency
Family/species DBH1 (cm) depth2 area diameter element length
(m) area (cm2) area ratio (vessels/
(cm) (cm²) (μm) (μm)
(%) mm2)
MYRISTICACEAE
Iryanthera ulei Warb 23.2 19.4 ± 0.5 4.7 ± 1.4 192.7 245.5 78.5 82.0 ± 22.4 10.0 ± 2.6 765.6 ± 142.2
Virola calophylla (Spruce) Warb. 22.5 30.1 ± 1.3 11.0 ± 1.0 648.8 695.6 93.3 87.2 ± 28.5 12.3 ± 4.6 1020.7 ± 198.6
Virola surinamensis (Rol. Ex. Rottb.) Warb 28.5 36.4 ± 1.3 12.9 ± 0.6 950.7 1040.6 91.4 110.4 ± 28.9 6.9 ± 2.6 1176.6 ± 220.8
MYRTACEAE
Caliptanthes spruceana Berg. 21.0 13.9 ± 1.1 2.9 ± 0.4 108.6 171.1 63.5 76.4 ± 19.3 18.5 ± 5.6 479.8 ± 132.8
NYCTAGINACEAE
Neea madeirana Standl. 11.3 7.2 ± 0.3 3.6 ± 0.5 41.6 41.6 100.0 58.8 ± 19.2 12.0 ± 3.8 147.7 ± 37.8
SAPOTACEAE
Chrysophyllum ucuquirana-branca
20.1 18.3 ± 1.1 1.4 ± 0.7 72.2 255.6 28.3 86.0 ± 19.2 15.1 ± 4.3 636.9 ± 173.0
(Aubrev & Pellegr) T. D. Penn.
Pouteria caimito Radlk. 22.6 26.0 ± 1.4 4.0 ± 0.8 262.8 494.0 53.2 79.6 ± 14.9 19.9 ± 4.7 622.8 ± 156.4
URTICACEAE
Cecropia sciadophylla Mart. 15.5 12.2 ± 0.3 3.7 ± 0.8 101.7 122.3 83.2 238.0 ± 44.2 2.5 ± 0.8 441.5 ± 85.8
Cecropia sciadophylla Mart. 17.6 15.1 ± 1.8 6.9 ± 1.4 182.4 184.3 99.0 210.4 ± 37.8 3.2 ± 1.3 500.4 ± 118.1
Cecropia sciadophylla Mart. 16.5 17.3 ± 1.5 7.4 ± 1.4 214.9 217.5 98.8 232.8 ± 53.5 3.1 ± 1.0 448.0 ± 79.6
Pourouma guianensis Aubl. 19.0 27.3 ± 1.1 2.8 ± 0.5 210.8 569.2 37.0 224.0 ± 29.6 2.6 ± 1.2 485.5 ± 111.6
Overall average 19.6 19.2 3.5 171.4 325.3 58.4 127.6 9.1 554.8
Overall standard deviation 4.7 8.4 2.6 177.6 258.5 27.0 49.2 6.5 201.9
Overall minimum 11.2 7.2 0.8 31.8 41.6 10.1 58.8 2.5 147.7
Overall maximum 28.5 40.6 13.0 950.7 1160.5 100.0 238 29.2 1176.6
Diameter-at-breast-height obtained from the measures of the smallest and largest diameter; 2Based on measurement of five radii; 3DBH was obtained through radii
1

measurements of a fragmented stem section.

Table S2. Sapwood data of 34 trees belonging to 26 species sampled in a terra firme forest in the central Brazilian Amazon. Sapwood area depth was estimated from
each of five radii measured (ARn). Sapwood area was estimated considering all five radii estimates (AS). DBH = diameter-at-breast-height (in cm) (average of two
measures); DR = average diameter estimated for each of the five radii (in cm); DR_Avg = average diameter considering all five radii estimates (in cm); DR% = percentage
difference between DBH and each DR; DR_Avg% = total average difference between DBH and each DR. Overall average, standard deviation, minimum and maximum
values of the measurements are also presented at the end of the table. Note: *Diameter was obtained through radii measurements of a fragmented stem section. N/A
represents “non available” data.

Species AR1 AR2 AR3 AR4 AR5 As DBH D1 D2 D3 D4 D5 DR_Avg D1% D2% D3% D4% D5% DR_Avg%

Bocageopsis multiflora 2.5 2.7 3.0 3.0 3.0 2.8 ± 0.2 26.8 ± 1.1 27.4 25.6 26.8 25.0 27.0 26.4 ± 1.0 102 96 100 93 101 99 ± 4

Brosimum utile 3.3 2.2 3.8 2.5 1.9 2.7 ± 0.8 26.7 ± 1.9 29.0 26.0 27.6 21.6 27.4 26.3 ± 2.8 109 98 104 81 103 99 ± 11

Byrsonima crispa 2.3 2.7 2.7 1.6 1.5 2.2 ± 0.6 15.9 ± 2.3 18.4 17.6 15.4 16.0 14.0 16.3 ± 1.8 116 111 97 101 88 102 ± 11

Caliptanthes spruceana 3.6 3.0 2.7 2.9 2.4 2.9 ± 0.4 13.9 ± 1.1 18.0 16.6 11.4 11.4 16.4 14.8 ± 3.1 129 119 82 82 118 106 ± 22

Cecropia sciadophylla 4.7 4.0 3.1 2.7 3.9 3.7 ± 0.8 12.2 ± 0.3 14.0 14.0 11.6 10.2 12.6 12.5 ± 1.6 115 115 95 84 103 102 ± 13

Cecropia sciadophylla 8.7 7.3 4.7 6.7 7.0 6.9 ± 1.4 15.1 ± 1.8 18.6 15.8 11.8 15.0 15.4 15.3 ± 2.4 123 105 78 99 102 101 ± 16

Cecropia sciadophylla 9.5 7.0 5.9 6.5 8.2 7.4 ± 1.4 17.3 ± 1.5 21.0 16.0 13.2 15.0 18.0 16.6 ± 3.0 122 93 77 87 104 96 ± 17
Chrysophyllum
2.0 1.5 0.6 0.8 2.0 1.4 ± 0.7 18.3 ± 1.1 19.4 19.0 17.2 17.6 17.0 18.0 ± 1.1 106 104 94 96 93 99 ± 6
ucuquirana-branca

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Table S2. Continued.

Species AR1 AR2 AR3 AR4 AR5 As DBH D1 D2 D3 D4 D5 DR_Avg D1% D2% D3% D4% D5% DR_Avg%

Cordia silvestris 3.5 2.0 2.0 1.7 1.5 2.1 ± 0.8 20.6 ± 4.3 26.0 24.0 17.0 16.0 20.0 20.6 ± 4.3 N/A N/A N/A N/A N/A N/A

Duguetia flagellaris 5.1 4.7 4.0 4.2 5.0 4.6 ± 0.5 9.4 ± 0.2 10.2 9.4 8.0 8.4 10.0 9.2 ± 1.0 109 101 86 90 107 98 ± 10

Eschweilera fracta 4.4 4.4 4.3 3.3 4.4 4.2 ± 0.5 21.9 ± 0.9 23.2 22.8 19.8 18.0 22.0 21.2 ± 2.2 106 104 91 82 101 97 ± 10

Eschweilera odora 2.8 2.6 3.1 2.6 2.0 2.6 ± 0.4 20.7 ± 1.2 22.7 22.0 23.8 21.0 22.3 22.3 ± 1.0 110 107 115 107 102 108 ± 5

Eschweilera wachenheimii 4.9 4.8 4.6 4.3 4.0 4.5 ± 0.4 10.5 ± 0.7 10.4 10.6 10.2 9.8 9.2 10.0 ± 1.0 99 101 97 93 88 96 ± 5

Eschweilera wachenheimii 4.4 4.7 4.8 3.7 3.3 4.2 ± 0.7 18.1 ± 0.2 19.4 18.6 17.0 18.6 18.4 18.4 ± 0.9 107 103 94 103 102 102 ± 5

Eschweilera wachenheimii 3.5 2.7 3.1 0.5 0.1 1.8 ± 1.8 20.8 ± 1.1 22.0 18.0 18.8 17.0 16.0 18.4 ± 2.3 106 87 91 82 77 88 ± 11

Eschweilera wachenheimii 0.8 1.3 0.9 1.0 0.7 0.9 ± 0.2 16.0 ± 0.0 18.0 17.6 14.2 14.0 16.4 16.0 ± 1.9 113 110 89 88 103 100 ± 12

Eschweilera wachenheimii 0.8 0.8 0.8 0.8 0.9 0.8 ± 0.1 12.9 ± 0.5 13.6 13.0 13.0 12.6 13.6 13.2 ± 0.4 106 101 101 98 106 102 ± 4

Gustavia speciosa 3.3 2.8 3.4 3.0 2.3 3.0 ± 0.4 17.9 ± 0.5 19.4 19.0 16.8 15.0 16.6 17.4 ± 1.8 109 106 94 84 93 97 ± 10

Iryanthera ulei 7.1 3.9 3.7 5.1 3.9 4.7 ± 1.4 19.4 ± 0.5 25.6 16.8 13.0 19.6 13.4 17.7 ± 5.2 132 87 67 101 69 91 ± 27

Jacaranda copaia 3.0 2.5 2.6 2.5 2.1 2.5 ± 0.3 20.4 ± 0.2 20.8 20.6 20.6 18.4 18.2 19.7 ± 1.3 102 101 101 90 89 97 ± 7

Mabea caudata 2.2 1.6 1.9 1.2 1.5 1.7 ± 0.4 11.5 ± 1.2 12.0 10.0 10.4 11.0 10.0 10.7 ± 0.8 108 90 93 99 90 96 ± 8

Mabea caudata 4.0 2.3 2.8 4.0 4.0 3.4 ± 0.8 17.5 ± 0.7 23.0 17.0 13.6 15.6 19.0 17.6 ± 3.6 131 97 78 89 109 101 ± 20

Micrandropsis scleroxylon 4.5 3.8 3.0 3.0 3.0 3.5 ± 0.7 28.1 ± 3.4 31.0 24.0 27.0 26.6 28.0 27.3 ± 2.5 110 85 96 95 100 97 ± 9

Neea madeirana 4.2 4.1 3.4 3.3 3.2 3.6 ± 0.5 7.2 ± 0.3 8.4 8.2 6.8 6.6 6.4 7.3 ± 0.9 117 114 94 92 89 101 ± 13

Ocotea neesiana 1.6 1.5 1.4 1.4 1.5 1.5 ± 0.1 18.8 ± 0.4 19.0 19.0 16.8 17.0 18.0 18.0 ± 1.1 101 101 90 91 96 96 ± 5

Pourouma guianensis 2.5 2.8 2.1 3.5 3.0 2.8 ± 0.5 27.3 ± 1.1 30.4 25.0 25.8 25.4 28.0 26.9 ± 2.3 112 92 95 93 103 99 ± 8

Pouteria caimito 4.9 2.9 3.6 4.3 4.1 4.0 ± 0.7 26.0 ± 1.4 32.0 27.0 24.6 18.4 23.4 25.1 ± 5.0 123 104 95 71 90 96 ± 19

Protium heptaphyllum 2.0 1.9 1.4 1.7 0.9 1.6 ± 0.4 8.8 ± 0.8 10.6 9.6 7.0 8.2 9.6 9.0 ± 1.4 121 110 80 94 110 103 ± 16
Sclerolobium
2.4 3.0 2.6 2.4 1.8 2.4 ± 0.4 20.9 ± 0.2 25.4 21.0 18.0 19.2 22.0 21.1 ± 2.9 122 101 86 92 106 101 ± 14
chrysophyllum
Tachigali paniculata 3.8 3.2 2.8 3.0 2.9 3.1 ± 0.4 8.4 ± 0.5 9.6 8.8 8.6 8.6 7.4 8.6 ± 0.8 115 105 103 103 89 103 ± 9

Vantanea micrantha 1.0 1.0 0.5 1.3 1.2 1.0 ± 0.3 40.6 ± 2.3 49.0 40.2 36.4 31.6 35.0 38.4 ± 6.7 121 99 90 78 86 95 ± 16

Vantanea micrantha 9.0 3.2 2.1 3.7 5.5 4.7 ± 2.7 20.5 ± 0.5 29.0 19.0 15.0 20.4 22.0 21.1 ± 5.1 142 93 73 100 108 103 ± 25

Virola calophylla 12.5 9.7 10.8 10.8 11.3 11.0 ± 1.0 30.1 ± 1.3 33.0 30.0 28.0 27.2 30.6 29.8 ± 2.3 110 100 93 90 102 99 ± 8

Virola surinamensis 13.0 12.1 13.0 13.7 13.2 12.9 ± 0.6 36.4 ± 1.3 38.0 35.0 36.0 36.4 36.0 36.3 ± 1.1 104 96 99 100 99 100 ± 3

Average 4.0 3.3 3.1 3.1 3.2 3.5 ± 0.7 19.2 ± 0.9 21.5 18.7 16.9 16.7 18.2 19.0 ± 2.2 114 100 91 92 97 99 ± 9

Standard Deviation 2.6 1.9 1.9 2.1 2.5 2.6 8.4 8.5 6.8 7.0 6.0 6.9 7.4 10 9 10 8 10 4±1

Minimum 0.8 0.8 0.5 0.5 0.1 0.8 7.2 8.4 8.2 6.8 6.6 6.4 7.3 99 79 67 71 69 88 ± 13

Maximum 12.5 9.7 10.8 10.8 11.3 13 40.6 49.0 40.2 36.4 31.6 35.0 38.4 142 119 115 107 118 108 ± 13

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Table S3. Descriptors of the physical characteristics observed on each sampled tree log cross-section surface (n = 34). For heartwood visibility: V = completely visible,
P = partially visible.
Heartwood Sapwood reaches
Species Vessel type Exudate Tylose Hollow stem
visibility heartwood
Bocageopsis multiflora (Mart.) R. E. Fries Diffuse V Sap - - X
Brosimum utile (Kunth) Pittier Diffuse V Resin - - -
Byrsonima crispa A. Juss Diffuse P - - X -
Caliptanthes spruceana Berg. Diffuse - - X - -
Cecropia sciadophylla Mart. Diffuse - - - - X
Cecropia sciadophylla Mart. Diffuse - - - X X
Cecropia sciadophylla Mart. Diffuse - - - X X
Chrysophyllum ucuquirana-branca (Aubrev & Pellegr) T. D. Penn. Diffuse V Latex - - -
Cordia silvestris Fresen. Diffuse P - X - -
Duguetia flagellaris Huber Diffuse V - - - X
Eschweilera fracta R. Knuth Diffuse V - X X1 -
Eschweilera odora (Poepp. ex O. Berg) Diffuse V - X X -
Eschweilera wachenheimii Sandwith Diffuse V - X X X
Eschweilera wachenheimii Sandwith Diffuse V Resin X X X
Eschweilera wachenheimii Sandwith Diffuse V - X X -
Eschweilera wachenheimii Sandwith Diffuse V - - X -
Eschweilera wachenheimii Sandwith Diffuse V - X - -
Gustavia speciosa (Kunth) DC. Diffuse P - - - -
Iryanthera ulei Warb Diffuse - Resin X - X
Jacaranda copaia (Aubl.) D. Don Diffuse V - X - X
Mabea caudata Pax & K. Hoffm. Diffuse - - X - -
Mabea caudata Pax & K. Hoffm. Diffuse V - X - -
Micrandropsis scleroxylon W. Rodr. Diffuse V Latex X - -
Neea madeirana Standl. Diffuse - - - - -
Ocotea neesiana (Miq.) Kosterm. Diffuse V - - X X
Pourouma guianensis Aubl. Diffuse P Latex - - X
Pouteria caimito Radlk. Diffuse - - X - -
Protium heptaphyllum (Aubl.) March. Diffuse P - - - X
Sclerolobium chrysophyllum Poepp. Diffuse V Latex - - X
Tachigali paniculata Aubl. Diffuse - - - X X
Vantanea micrantha Ducke Diffuse V - X X X
Vantanea micrantha Ducke Diffuse P - X - -
Virola calophylla (Spruce) Warb. Diffuse - - - - X
Total trait occurrences (n) 34 22 9 17 13 18
% occurrences 100 65 26 50 38 53
Diseased.
1

Table S4. Descriptive statistical parameters of exponential power model fits using sapwood area and tree size [diameter-at-breast-height (DBH), total height (Ht)],
volume (estimated using DBH and Ht), and anatomical variables (vessel diameter, element length and frequency). Significance is set at 5%. * P<0.05; ** P<0.01; ***
P<0.001; ns = non-significant. a and b = model coefficients (Y=aXb); R2 = coefficient of determination; r = Pearson correlation coefficient; † = fitted with an exponential
decay model (Y=a∙e(-bx)). Note: only the best allometric model fits are shown.
Variables a b R² r F
DBH 0.823 1.781 0.46*** 0.68 27.32
Total height 0.013 3.110 0.33*** 0.58 16.08
Volume (DBH, Ht) 376.95 0.817 0.48*** 0.70 29.96
DBC 0.188 2.307 0.71*** 0.84 39.52
Vessel diameter 11.848 0.499 0.001ns 0.03 0.02
Vessel element length 0.000007 2.628 0.65*** 0.80 58.63
Vessel frequency† 197.96 0.016 0.01ns -0.10 0.32

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Figure S1. Microscopic wood proprieties classified according to their vessel dimensions as proposed by Copant (1974) and the percentage of trees included in each
category. A. Vessel-lumen diameter (µm); B. vessel frequency (vessels per mm2); C. vessel element length (µm); D. vessel groupings (solitary or multiples). This figure
is in color in the electronic version.

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