Biomass estimations in forests of different disturbance

history in the Atlantic Forest of Rio de Janeiro, Brazil

Andre Lindner

Dietmar Sattler
Received: 21 October 2010 / Accepted: 9 August 2011 / Published online: 18 August 2011
Springer Science+Business Media B.V. 2011
Abstract Tropical forests are large reservoirs of biomass and there is a need for infor-
mation on existing carbon stocks in these ecosystems and especially the effects of logging
on these stocks. Reliable estimates of aboveground biomass stocks within the Atlantic
Forest are rarely available. Past human disturbance is an important factor affecting forest
structure variation and biomass accumulation among tropical forest ecosystems. To sup-
port the efforts of improving the quality of estimations of the current and future biomass
carbon storage capacity of this disturbed forest region we tested a non-experimental small
scale approach to compare the aboveground tree biomass (AGB) of forest sites. Three sites
with known disturbance histories have been investigated: complete cut down, selective
logging and conservation since 70 years. The woody plant community (dbh C 10 cm) was
censused and canopy openness in conjunction with leaf area index has been obtained by
hemispherical photographs at each site. Estimates of aboveground tree biomass have been
carried out using an allometric equation for moist tropical forests already applied for the
study area. Additionally, a FAO standard equation has been employed for crosschecking
our results. We identied signicant differences in recent AGB of the three compared
forest sites. With 313 (48 Mg ha
-1
) the highest AGB-values have been found in the
preserved forest area within a National Park, followed by 297 (83) Mg ha
-1
at the former
clear cut site. Lowest AGB has been calculated for the area with past selective logging: 204
(38) Mg ha
-1
. Values calculated with the FAO standard equation showed the same trend
but at a lower AGB level. Our results based an a small scale approach suggest that biomass
productivity can recover in a forest which was completely cleared 60 years ago to reach
AGB values up to a level that almost represents the situation in a preserved forest.
A. Lindner (&)
Department of Systematic Botany and Functional Biodiversity, Institute for Biology I,
University of Leipzig, Johannisallee 21-23, 04103 Leipzig, Germany
e-mail: alindner@uni-leipzig.de
D. Sattler
Department of Physical Geography and Geo-Ecology, Institute for Geography, University of Leipzig,
Johannisallee 19, 04103 Leipzig, Germany
e-mail: sattler@uni-leipzig.de
1 3
New Forests (2012) 43:287301
DOI 10.1007/s11056-011-9281-9
Selective logging may slow down AGB accumulation and the effect is measurable after
several decades.
Keywords Aboveground live biomass Tropical forests Atlantic Forest Land use
Forest succession
Introduction
Despite the fact that the Atlantic Forest is probably the region with the highest species
diversity and degree of endemism throughout South America (Tabarelli et al. 2005; Silva
and Casteleti 2003), this forest was one of the largest of the subcontinent and covered
about 1.2 million km
2
. Its remaining remnants cover nowadays, depending on the deni-
tion of Atlantic Forest and the spatial methods employed, about 716% of its original
extent (Galindo-Leal and de Gusmao Camara 2003; Tabarelli et al. 2005; SOS Mata
Atlantica 2009; Ribeiro et al. 2009). However, the spatial distribution of these remnants
gives a more meaningful representation of the situation than the total amount of remaining
forest. According to Ribeiro et al. (2009), the remnants of the so called Mata Atlantica
are currently distributed in more than 245,000 forest fragments of which 83.4% are smaller
than 50 ha and only 0.03% are larger than 10,000 ha. In addition to their degree of
fragmentation, these remaining forests are frequently to be found in varying stages of
degeneration caused by anthropogenic disturbance. Forest types with different disturbance
history are very heterogeneous habitats, continuously changing in structure and varying in
species composition (e.g., Williams-Linera et al. 1998; Laurance et al. 2002; Galindo-Leal
and de Gusmao Camara 2003). If these forest remnants cover 716% of the original extent
of the Mata Atlantica biome, do they represent 716% of the original forests biomass?
Probably not, because there are indications that the biomass of a large number of small
forest fragments left over after habitat fragmentation is much less then in a continuous
forest of the same overall size (Groeneveld et al. 2009).
Compared to other neotropical forests (e.g., Amzonian Rain Forests), reliable estimates
of aboveground biomass stocks within the Atlantic Forest are rarely available. The few
existing studies estimating aboveground biomass in this region have been carried out
mainly in relatively undisturbed and protected Atlantic Forest of different elevations (e.g.,
Rolim et al. 2005; Alves et al. 2010). Two studies known to the authors have been carried
out in secondary forests of the Mata Atlantica region using direct measurements by har-
vesting and weighting dry biomass (Burger and Delitti 2008; Tiepolo et al. 2002). Nev-
ertheless, particularly the effects of different disturbance histories on forest structure and
biomass stocks seem to be essential in the Amazon (Nogueira et al. 2008; Kellner and
Asner 2009), but are poorly understood in fragments of the Atlantic Forest (Kindel et al.
1999).
There is an alarming loss of biomass due to logging activities (Houghton 2005) and
nowadays about 60% of all tropical forests are secondary or degraded (ITTO 2002). Hence,
it becomes increasingly important to evaluate biomass stocks of recovering disturbed forest
(Frolking et al. 2009; Saldarriaga and Uhl 1991; Uhl and Buschbacher 1985). In our study
region, many of the forest remnants are in fact secondary re-growth of formerly cut down
or strongly disturbed Atlantic Forest (Pedreira et al. 2009).
With this study we address the hypothesis that even small scale, non-experimental
approaches using existing information on past disturbance can be used to reveal differences
in aboveground tree biomass (AGB) accumulation. Using the existing heterogeneous
288 New Forests (2012) 43:287301
1 3
situation we compared the AGB of forest sites with three distinct disturbance histories:
complete cut down, selective logging and conservation since 70 years. In addition to
historic information we used structural parameters such as stem density per hectare, tree
height classes, leaf area index (LAI) and canopy openness as possible proxies for the
disturbance history. Because of the fact that large tree individuals are known to provide a
disproportionally high contribution regarding aboveground biomass in tropical forests
(Vieira et al. 2004; Midgley and Niklas 2004; Chambers et al. 2001), we analyzed forest
composition with a special regard on these structural elements. As indirect AGB estima-
tions are carried out using allometric models, the results depend essentially from the
calculation method applied. To verify the AGB values calculated for the three different
forest sites with an allometric model based on dbh only, we performed an additional
standard FAO calculation including average wood density.
Study area
The study was conducted in two areas within the Atlantic Forest (Mata Atlantica) of the
state of Rio de Janeiro, Brazil (Fig. 1). The rst research area is the 5,500 ha NGO reserve
Reserva Ecologica de GuapiacuREGUA (22 25
0
53 S, 42 45
0
20 W), located in the
municipality of Cachoeiras de Macacu about 100 km northeast of Rio de Janeiro. Mean
annual temperature is about 23C with a mean annual rainfall of about 2,560 mm (Kurtz
and de Araujo 2000). The recently enlarged national park Parque Nacional da Serra dos
O

rgaos (PARNASO), founded in 1939 and now covering an area of 20,024 ha within the
municipalities of Petropolis, Guapimirim, Mage and Teresopolis, was our second research
area (2227
0
24
00
S, 4259
0
48
00
W). Annual rainfall is 2,821 mm in combination with high
relative humidity and a mean annual temperature of 17.8C (Rizzini 1954; Guimaraes and
Arle 1984). The regional climate is characterized by a hot and rainy season from October
to March and a cooler and drier season from April to September (Kurtz and de Araujo
Fig. 1 Study area in south-eastern Brazil in the state of Rio de Janeiro. REGUANGO reserve Reserva
Ecologica de Guapiacu, located in the municipality of Cachoeiras de Macacu. PARNASOnational park
Parque Nacional da Serra dos O

rgaos, research site located in the municipality of Teresopolis

New Forests (2012) 43:287301 289
1 3
2000). The natural vegetation can be classied as dense ombrophilous forest (Veloso et al.
1991) and is typical for the lower and medium elevations 501,500 m asl. of the coastal
mountain range of the Rio de Janeiro state (Morellato and Haddad 2000; Oliveira-Filho
and Fontes 2000). Within the Mata Atlantica Biome, the dense ombrophilous forest hosts
more than half (60%) of the plant species richness and the vast majority (80%) of endemic
plants (Stehmann et al. 2009). Characteristic canopy tree species of the lower elevations of
this forest (up to 500 m asl.) are Vochysia tucanorum (Vochysiaceae), Miconia thaezans,
Tibouchina granulosa (Melastomataceae), Xylopia brasiliensis (Annonaceae) and Pla-
thymenia foliosa (Fabaceae), while montane elevations (up to 1,500 m asl.) are charac-
terized by Vochysia laurifolia (Vochysiaceae), Talauma ovata (Magnoliaceae), Cariniana
estrellensis (Lecythidaceae), Cedrela angustifolia (Meliaceae) and Clethra brasiliensis
(Clethraceae). Myrtaceae, Rubiaceae and Melastomataceae are the most abundant and
typical families both in the sub-canopy and the understory layer (Rambaldi et al. 2003). In
the study area there are several stages of succession of forest and non-forest vegetation
representing degradation after disturbance. The very early stages of forest succession
following disturbance can be dominated by only a few woody species such as Miconia
albicans (Melastomataceae), Attalea humilis (Arecaceae) and Gochnatia polymorpha
(Asteraceae) as reported by Lima et al. (2006). The comparison of sites within these two
areas may be disadvantageous because they are representing an elevation difference of
around 900 m. We had to accept this because of the fact that the occurrence of mature
forest with low or no anthropogenic inuence is mainly restricted to conservation units in
higher elevations of the Atlantic Forest of this region.
Methods
Plot design, census and forest structure
At the NGO reserve area our research plots were established in two different forest sites: in
continuous forest where selective logging took place until the late 1970s at 500 m asl. and
in forests re-growing after clear-cut about 60 years ago at 235 m asl. (Locke, N. pers.
comm.). A 100 9 100 m (1 ha) research plot with a 10 9 10 m grid of marked points was
set up at each study site. Five 20 9 20 m subplots were randomly chosen within those
plots with the constraint that subplots do not overlap. Data from the preserved forest areas
in the national park was acquired in ve 20 9 20 m plots distributed at elevations between
1,165 and 1,300 m asl. In each 20 9 20 m plot at every investigation site the woody plant
community was censused according to Condit (1998). All woody plants with a diameter at
breast height (dbh) C 10 cm were tagged (including multiple stemmed individuals
exceeding 10 cm in sum). Diameter in breast height was measured by using a calliper or a
measuring tape when the diameter exceeded 50 cm. The stem-height of each individual
was estimated using four height classes:
1. \5 m
2. 510 m
3. 1020 m
4. [20 m.
Canopy openness and LAI were measured with hemispherical photographs, taken in
series at the end of the dry and wet season respectively (2004/05 PARNASO, 2007/08
REGUA) to capture seasonal effects and inter-annual variability. This variability is a
290 New Forests (2012) 43:287301
1 3
typical pattern for forests of the region and needs to be taken into account as it develops
when forests are disturbed and/or close to edges (Sattler et al. 2007). However, the vari-
ability does not seriously affect biomass calculations due to the comparably low contri-
bution (\2%) of seasonally varying leaf biomass to total AGB (Chave et al. 2008).
For hemispherical photography we used a Nikon
TM
Coolpix 4500 digital camera with a
Nikon
TM
FC-E8 sheye converter mounted on a leveled tripod at 1.30 m above ground.
Photographs were only taken under overcast conditions to avoid overexposure which could
subsequently lead to inaccurate LAI and canopy openness calculations. In the NGO reserve
area photos were taken at every grid-point within the 1 ha plots (100 photos per plot per
series), whereas at the preserved forest sites 16 photos were taken per series in each of the
20 9 20 m plots.
Data analysis
We used an allometric equation (1) given in Chave et al. (2001) for moist tropical forests
and already applied for the Atlantic forest (Rolim et al. 2005), where AGB for each tree
was calculated as: (1) ln(AGB) = -2.19 ? 2.54 (ln dbh), valid for a dbh range from 10 to
150 cm. As palm trees are especially abundant in the National Park plots, AGB of palms
was calculated separately using an allometric model suggested by Hughes et al. (1999) and
modied by Vieira et al. (2008): (2) exp((5.7236 ? 0.9285 (ln dbh
2
))*1.05001)/10
3
.
Following Alamgir and Al-Amin (2008) models using basal area (dbh respectively) alone
were found to be the best predictor of biomass stock in trees because of high coefcient of
determination. Aboveground tree biomass is measured in kg and dbh in cm. As the
equation by Chave et al. (2001) is based on only one independent variable (dbh) we also
applied the standard equation by the Food and Agriculture Organisation of the United
Nations (3) as described in Garzuglia and Saket (2003), which additionally includes wood
density and a biomass expansion factor. This methodological comparison was employed to
reconrm our AGB calculation with a different and commonly used calculation approach.
Aboveground biomass for each tree was then calculated as:
AGB BV BEF; with BV VOB
10
WDand BEF Expf3:213 0:506 lnBVg
1
where BV = dry woody biomass of inventoried volume, VOB
10
= stem volume of living
trees dbh C 10 cm, WD = wood density and BEF = biomass expansion factor.
The total basal area (calculation based on individual dbh) per 20 9 20 m plot was
scaled up to get the basal area per hectare per forest site. The stem volume (VOB
10
) was
calculated using ve-level stem height classes. Since the specic wood density data of
many of the censused tree species could not be acquired or showed only small variation
(Chave et al. 2009), we used a constant WD = 0.603 g cm
3
as suggested by Vieira et al.
(2008) for the Atlantic forest of South East Brazil. A One-Way-ANOVA with Student
NewmanKeuls multiple comparison procedure was used to analyze differences in AGB
between the differently disturbed sites on the one hand, and differences between the results
of both AGB estimation methods for each site on the other.
The WinScanopy 2005ab software (Regent Instruments Inc. 2005) was used to analyze
the hemispherical photographs and to derive data about canopy openness (CO) and leaf
area index (LAI). Canopy openness is dened as the proportion of open sky area in a 180
hemisphere monitored from a centre point. Pixels of digital images are to be classied as
either canopy or sky based on the grayscale threshold value. Leaf are index is the
New Forests (2012) 43:287301 291
1 3
total one-sided area of leaf tissue per ground unit. The LI-COR LAI2000 modied algo-
rithm (assuming random leaf distribution, transmittance is equivalent to gap fraction,
modelled by the Poisson model) was used to calculate LAI values (Welles and Norman
1991).
A One-Way ANOVA on ranks was used to check for signicant differences in CO and
LAI between the different forest types. Due to the different sample size of hemispherical
photographs a Dunns all multiple comparison procedure was performed to differentiate
between the groups. Additionally, we compared the seasonal difference at each site in
those canopy structure parameters with a repeated-measure ANOVA on ranks. Statistical
analyses were performed using the SigmaStat software package (version 3.0.1, SPSS Inc.,
2003).
Remarks on methodology
As there are many possible environmental an biological factors inuencing the develop-
ment of a forests AGB, the realistic prediction of the AGB development of different forest
sites over time is highly uncertain (Asner et al. 2009). A way out of this unsatisfactory
situation would be an experimental plot design which is very complicated due to the
required time scale and the inevitable destruction of forest. Nonetheless, our small scale
non-experimental approach provides valuable results on a eld of research in which data is
still scarce. Among others (McWilliam et al. 1993; DeWalt and Chave 2004; Vilela et al.
2000) our approach conrms the feasibility of small scale studies to assess questions of
forest structure and biomass in tropical forests.
Using the standard equations by the FAO (Garzuglia and Saket, 2003) for reconrming
our AGB calculation with a different approach we applied a constant wood density of
0.603 g cm
3
. Needless to say that wood gravity differs between species and therefore
between sites with a different species composition, but our approach has been validated by
Wiemann and Williamson (2002), Chave et al. (2006) and Vieira et al. (2008).
Results
Biomass and stand structure
Stem density and tree height was signicantly higher in the preserved forest plots, whereas
both parameters did not differ between the sites with different disturbance history in the
NGO reserve area. We noted a non-signicant trend of higher basal area of the forest stand
in the preserved forest plots.
The composition of the forest regarding dhb-size classes of trees revealed a different
pattern (Fig. 2), especially in the fraction up to a dbh of 30 cm. There were signicantly
less small trees (dbh 1020 cm) in the area which was completely cleared 60 years ago,
whereas trees with a dbh of 2030 cm were signicantly less represented in the preserved
forest plots. We found no differences in the number of trees per plot with a dbh C 30 cm
between investigation areas. When looking at the distribution of the total basal area within
different dbh-classes (Fig. 3) the same pattern was identied. In general big trees
(dbh C 30 cm) do contribute more to total basal area in the preserved forest plots, even
though no signicances were found due to a high variability (Lindner 2010). Nevertheless
we identied a distinct coherence between the basal area of the largest tree and the total
basal area per plot (Fig. 4).
292 New Forests (2012) 43:287301
1 3
Biomass calculations using two different approaches resulted in a comparable trend
pattern. We identied the highest AGB-values in the preserved forest area within the
National Park (313 48 Mg ha
-1
). Intermediate results for AGB were found at the for-
mer complete cut down site (297 83 Mg ha
-1
) and lowest AGB was calculated for the
area with past selective logging (204 38 Mg ha
-1
). The results indicate that AGB
values for the complete cut down site in the NGO reserve area and those in the preserved
forest area are not signicantly different (q = 0.72, P = 0.62). In contrast, the low AGB
Fig. 2 Diameter distribution of all individuals dbh C 10 cm sampled in ve plots per forest type (complete
cut down: n = 109, selective logging: n = 133, conservation area: n = 207)
Fig. 3 Diameter distribution of total basal area (ba) of all individuals C 10 cm dbh sampled in ve plots of
20 9 20 m per forest type (complete cut down: ba = 6.10 m
2
, selective logging: ba = 4.72 m
2
,
conservation area: ba = 7.60 m
2
)
New Forests (2012) 43:287301 293
1 3
value for the forest which was selectively logged is signicantly different to the other sites
(clear cut q = 3.42, P = 0.03; preserved forest q = 4.14, P = 0.03). The crosscheck
performed by applying the FAO calculation method revealed lower AGB for both the
selectively logged and the complete cut down site but higher AGB in the preserved forest
(Table 1). As the FAO calculation does not treat the palm trees separately it leads to an
overestimation of AGB in the preserved forest due to high palm abundance at these sites.
Canopy structure and dynamics
We detected several differences in canopy structure parameters depending on location and
climate seasonality (Table 1). The lowest LAI of all sites was measured in the former clear
cut plots, whereas a higher LAI was recorded in plots in the selectively logged area and the
preserved forest plots of the national park. This interval becomes even more obvious and
signicant during the dry season. Seasonal changes in LAI were signicant only for plots
in the preserved forest and the selectively logged area. Despite higher LAI measures we
found higher canopy openness in the preserved forest plots too. Canopy openness was
subject to strong seasonal uctuations at both sites in the NGO reserve area REGUA,
which is in contrast to the preserved forest plots where no signicant seasonal changes in
this parameter were found.
Vegetation composition at REGUA NGO reserve area
Although no standard oristic survey was conducted, we recognized different character-
istics in terms of tree species composition in the investigated sites. Especially two species
showed particularly noticeable patterns of dominant occurrence. The Jussara-palm tree
Euterpe edulis (Arecaceae) represented 38% of all individuals and 10.5% of total basal
area in the forest plots of the national park and was the dominating species with a
dbh B 20 cm (mean dbh 11.6 1.1 cm). This palm tree is endemic to the Atlantic Forest,
where it provides an economically important resource for palm heart harvesting (Galetti
and Fernandez 1998). Harvesting the apical meristemic cone (palm heart) means killing the
Fig. 4 The relationship between basal area of the largest tree and total basal area in the complete data set
294 New Forests (2012) 43:287301
1 3
individual. Thus, it is no surprise that E. edulis was completely absent in the disturbed
NGO reserve plots. The other conspicuous tree species was Piptadenia gonoacantha
(Mimosaceae), dominating the tree community with a dbh C 20 cm in the plots in NGO
reserve area which were completely cleared 60 years ago (mean dbh 43.4 11.6 cm).
This pioneer tree species is typical for early and mid successional stages of the Atlantic
Forest (Borem and Oliveira-Filho 2002; Thier and Wesenberg 2009) and represented
18.5% of all individuals and 44% of total basal area in the NGO plot with past clear-cut
and could only be recorded here. Surprisingly, no individuals of P. gonoacantha with a
dbh B 20 cm or, in other words, no actual regeneration of this species were found.
Discussion
Aboveground tree biomass, forest structure and disturbance history
Our results in biomass calculations based on a small scale approach are well in range of
published data for other moist neotropical forests (Chave et al. 2001; DeWalt and Chave
2004; Urquiza-Haas et al. 2007; Groeneveld et al. 2009). Compared to AGB estimates for
Table 1 Stand structure, aboveground tree biomass (AGB) and canopy structure parameters of all study
sites
REGUA PARNASO P
Complete cutdown Selective logging Preserved forest
Stand structure
a
AGB (Mg ha
-1
)
b
297 (83) a 204 (38) b 313 (48) a 0.031
AGB (Mg ha
-1
)
c
207 (40) a 155 (18) b 336 (27) c \0.001
P 0.06 0.031 0.42
Stem density per ha 545 (33) a 665 (128) a 1,010 (219) b \0.001
Basal area (m
2
ha
-1
) 30.5 (6.9) 23.6 (2.5) 38.0 (13.1) 0.07
Tree hight (classes) 2.89 (0.15) a 2.81 (0.14) a 3.00 (0.12) b \0.001
Canopy structure
LAI
Wet season 3.49 (0.34) 3.66 (0.58) 3.64 (0.47) 0.078
Dry season 3.55 (0.29) a 3.93 (0.54) b 4.08 (0.70) b \0.001
P 0.11 \0.001 \0.001
CO (%)
Wet season 6.45 (0.58) a 6.36 (0.64) a 6.91 (1.06) b 0.012
Dry season 5.82 (0.39) a 5.24 (0.53) b 6.57 (1.00) c \0.001
P \0.001 \0.001 0.12
Letters (a, b, c) indicate the horizontal grouping within the table resulting of a One-Way ANOVA with
StudentNewmanKeuls multiple comparison procedure for the AGB estimates and a One-Way ANOVA on
ranks with Dunns all multiple comparison procedure for the canopy structure parameters
Tree height classes: (1) \5 m; (2) 510 m; (3) 1020 m; (4) [20 m
a
Interpolated from ve 400 m
2
plots per forest type
b
Calculated using allometric equation (Chave et al. 2001)
c
Calculated using FAO-standards (Garzuglia and Saket 2003)
New Forests (2012) 43:287301 295
1 3
other disturbed and undisturbed forest sites in Brazils Atlantic Forest, our results coincide
as well with data for the respective forest condition, even though data published for mature
and/or undisturbed forests are from lower elevations (Table 2). The FAO equation
crosscheck of our AGB calculation revealed once more the inuence of the methodological
approach in estimating AGB of a given forest site (e.g., Brown et al. 1989 and Chave et al.
2008) as the values obtained with this calculation (Table 1) are up to 30% lower than the
ones calculated with the allometric equation of Chave et al. (2001). Nevertheless, the
relative trend of AGB within the differently disturbed forest sites remains the same. The
selectively logged forest contains the lowest AGB within the investigated areas in our
study and this result is independent from the applied biomass calculation method. Here the
selective removal of trees with a diameter appropriate for timber production indicates an
underrepresentation of trees with dbh C 50 cm (Fig. 3), although this demographic dif-
ference was not signicant it is still reected in the outcome of the calculated AGB.
Signicantly more AGB was estimated for the forest regrowth after complete clearing
60 years ago, which is mainly due to the contribution of a single mid-successional species:
P. gonoacantha. Within 60 years after complete clear cut the dominance of this species
within a re-growing forest led to a higher AGB than in a selectively logged forest. Overall
tree diversity is of course much lower at the clear-cut site. Furthermore, the fate of this
forest and further succession is rather unpredictable, especially after the disappearance of
P. gonoacantha which is likely to happen because neither seedlings nor young individuals
of this species were found within the plots.
Big trees (dbh C 30 cm) contribute more into total basal area and hence in calculated
AGB, especially for the protected forest area. For the National Park Parque Nacional da
Serra dos O

rgaos a contribution of 78% to total AGB is reported for such big trees
(Lindner 2010), which conrms their importance in terms of carbon storage capacity.
We also recognized a gap in dbh size class distribution between the different areas:
there were obviously less individuals with a dbh 2030 cm in the national park area. For
Table 2 Atlantic forest AGB estimates from literature in comparison to data of this study
Reference Forest type Forest
condition
Elevation
a.s.l. (m)
DBH
range
(cm)
AGB
estimation
AGB
(Mg ha
-1
)
Alves
et al.
(2010)
Lowland to lower
montane Atlantic
rain forest
Late
secondary/
mature
1001,000 5156 Allometric
model
243.7
(33.2)
Cunha
et al.
(2009)
Atlantic moist forest Secondary
(40 years)
600900 1070 Allometric
model
166.8
Rolim
et al.
(2005)
Lowland, semi-
deciduous Atlantic
moist forest
Relatively
undisturbed
2865 C 10 Allometric
model
334.5
(11.3)
Burger
(2005)
Atlantic moist forest Secondary
(30 years)
570 1.647.8 Destructive
sampling
245
This
study
Montane, dense
ombrophilous forest
Late
secondary/
mature
1,1651,300 1070 Allometric
model
313.5
(48.2)
This
study
Submontane, dense
ombrophilous forest
Secondary 235500 1070 Allometric
model
250.4
(77.9)*
* Mean value and STD from merged data of both selective logging and complete cutdown sites
296 New Forests (2012) 43:287301
1 3
our study, this difference might be due to an earlier successional stage indicating that this
size class provides a major contribution to total biomass within areas exposed to distur-
bance. The higher stem density in the protected forest plot may be due to the higher
elevation (Moser et al. 2007). No signicant difference in stem density was found in the
disturbed NGO reserve plots, independent from disturbance history. This conrms the
results by Vilela et al. (2000) which investigated the effect of selective logging in a
seasonally dry Brazilian Atlantic Forest. The most trees with a dbh B 20 cm have been
recorded within the plots of the protected forest area and were mainly represented by E.
edulis. The frequent occurrence of the Jussara-palm tree made the most noticeable dif-
ference when compared the NGO plots, where it was completely missing. In general high
density stands seem to be important in terms of carbon sequestration (Redondo-Brenes
2007).
A factor indirectly inuencing forest biomass calculations is the abundance of lianas.
They play a key role for forest dynamics and processes. Lianas can reduce the host tree
growth rate and suppress gap phase regeneration (Schnitzer and Bongers 2002). Moreover it
is known that lianas increase the mortality of host trees (Ingwell et al. 2010). Although
lianas represent a small fraction of AGB that has not been taken into account in our study,
their negative effects on tree growth can contribute to the low stand biomass in the selective
logging plot where lianas tend to be more abundant than in the other plots (Faske 2009).
General implications
As incipiently mentioned there is a general uncertainty about biomass calculation and
therefore for carbon stock estimations in tropical forests (Houghton et al. 2001; Fearnside
et al. 2009). Nevertheless, information and data availability differs regionally with some
regions (e.g., Amazonian Forests) being investigated more intensely. Especially in the
Atlantic Forest studies are scarce. Our study provides data that help to ll the data-gap for
the Atlantic Forest and contributes knowledge for a largely under-sampled area, especially
with regard to forests representing different disturbance histories.
While the estimation of biomass directly from remote sensing data still has been proved
illusive as a tool for large-scale estimates (Fearnside et al. 2009), reference points on the
ground are essential to improve the results of meta-studies (Saatchi et al. 2007) and to
supply data for ecological models like FORMIND (Kohler and Huth 1998; Groeneveld
et al. 2009). Most of the remaining area of the Atlantic Forest is highly fragmented
(Ribeiro et al. 2009) and the inuence of fragmentation and its effect on biomass condi-
tions is an important aspect to study (Saunders et al. 1991; Groeneveld et al. 2009).
However, disturbance history of forest patches and continuous forest is often not known
and generalization of ecosystem responses may lead to indistinct outcomes for upcoming
investigations. Midgley and Niklas (2004) drew attention on the fact that different dis-
turbance regimes determine the max size that trees can achieve and therefore strongly
inuence the total AGB of a forest. Therefore, small scale structural approaches as tested in
our study are able to form an initiating framework of more detailed results and help to
improve estimates on biomass and carbon storage.
Conclusions
Our small scale, non-experimental approach using existing information on past disturbance
and actual forest stand characteristics was applicable to reveal signicant differences in
New Forests (2012) 43:287301 297
1 3
AGB formation within the studied forest sites. Furthermore, the results of this study lead to
the conclusion that biomass productivity can recover in a forest which was completely
cleared 60 years ago, even if the details of past and further successional development
remain rather unexplained. On the other hand the outcome of our study shows that
gentle inuences like selective logging can have, even if stopped several decades ago. In
other studies selective logging is described to have little effect on forest structure and
dynamics (Deccker and de Graaf 2003), but our results suggest otherwise. These ndings
can contribute to the discussion on the value of different forest management strategies
within the context of the reductions in emissions from deforestation in developing coun-
tries (REDD) debate (Gibbs et al. 2007; Shevliakova et al. 2009; Reyer et al. 2009).
Acknowledgments We are grateful to Jens Wesenberg for kindly providing the oristic and forest
structure data for the national park area. We are thankful to Christian Wirth and anonymous reviewers for
valuable comments on earlier versions of the manuscript. Furthermore we would like to thank the national
park Serra dos Orgaos, the Brazilian Institute of Environment and Renewable Natural Resources
(IBAMA) and the National Counsel of Technological and Scientic Development (CNPq) for issuing
research permission. We are grateful to the Reserva Ecologica de Guapiacu (REGUA) and the whole
staff, especially to Mr. Nicholas Locke for logistic support and the permission to work on the property. For
funding we are thankful to the German Federal Ministry of Education and Research (BMBF). This study was
conducted within the framework of the Brazilian-German joint research project Climate change, landscape
dynamics, land use and natural resources in the Atlantic Forest of Rio de Janeiro (FKZ 01LB0801B).
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