Hum Ecol (2011) 39:455–464
DOI 10.1007/s10745-011-9412-0
Population Structure of Brazil Nut (Bertholletia excelsa,
Lecythidaceae) Stands in Two Areas with Different
Occupation Histories in the Brazilian Amazon
Ricardo Scoles & Rogério Gribel
Published online: 8 June 2011
# Springer Science+Business Media, LLC 2011
Abstract Here we hypothesize that the intensity of distur-
bances caused by human activities in Brazil nut stands
(castanhais) is positively related with the regeneration of
Brazil nut trees (Bertholletia excelsa H.B.K., Lecythidaceae)
and consequently with a younger population structure of this
species. In order to test this hypothesis we compared the
population structure of Brazil nut trees in two areas of the
Brazilian Amazon with different histories of land usage by
humans. Archeological and historical data suggest that the
region surrounding the Trombetas River was densely
occupied in pre-Columbian times and experienced depopu-
lation after European contact with Amerindian populations,
especially in the 16th century. The 25 Brazil nut stands
sampled in this region were dominated by old B. excelsa
trees and had scarce recruitment in the understory. These
very mature stands likely owe their origins to the interval
between the depopulation of the indigenous peoples in the
16th–17th centuries and the establishment of quilombos at the
beginning of the 19th century. The second study area was in
R. Scoles
Universidade Federal do Oeste do Pará,
Rua Marechal Rondon s/n. CEP,
68040-070 Santarém, Pará, Brasil
e-mail: ricardscoles@yahoo.es
R. Gribel (*)
Coordenação de Pesquisas em Botânica,
Instituto Nacional de Pesquisas da Amazônia (INPA),
Manaus, AM, Brazil
e-mail: rgribel@jbrj.gov.br
Present Address:
R. Gribel
Diretoria de Pesquisas,
Instituto de Pesquisas do Jardim Botânico do Rio de Janeiro,
Rio de Janeiro, RJ, Brazil
the vicinity of the Madeira River (Capanã Grande Lake),
where the castanhais were more accessible and disturbed. In
this site, a younger population structure and abundant
regeneration of B. excelsa were observed in the 10 sampled
stands. Historical data from this region indicate that
indigenous populations were replaced gradually beginning
in the 18th century, with no evidence of severe depopulation.
We suggest that the different historical and contemporary
land use patterns contributed to the current contrasting
population structures of the castanhais at the two locations.
The data also support the idea that the castanhais, even the
ones considered to be pristine and “native” forests, result
from anthropogenic influences. We found no evidence to
support restrictions on seed harvesting as a means to
improve regeneration rates of Brazil nut stands.
Keywords Amazonia . Anthropogenic forest .
Extractivism . Historical ecology . Regeneration . Brazil nut
Introduction
One of the most important economic non-timber forest
products of the Amazon region is the Brazil nut (Bertholletia
excelsa H.B.K., Lecythidaceae). Stands of Brazil nut trees,
commonly known as castanhais, are aggregations of
hundreds to a few thousand trees in the upland (terra firme)
Amazon forest. The castanhais are long thought to owe their
origin to the pre-Colombian Amerindians (Ducke 1946;
Balée 1989; Mori and Prance 1990). However, there is no
clear historical or ecological evidence of a long-term human
influence on the regeneration or population structure of the
castanhais. The clumped distribution and the abundance of
unusual size classes of B. excelsa have generated discussions
concerning the reasons for these characteristics. For example,
456
it is possible that openings in the forest created by
Amerindian activities permitted the germination of many
seeds that later became a stand (Mori and Prance 1990;
Salomão 1991). On the other hand, agoutis, considered the
sole natural seed dispersers of B. excelsa, may have gathered
and stored seeds in middens (Tuck Haugaasen et al. 2010)
that were later forgotten. The stored seeds may have
germinated, forming stands (Peres and Baider 1997).
The physical proximity of the castanhais to areas with
dark earths and archaeological sites suggests that the
Amerindian peoples were influential in determining their
current pattern of distribution and abundance (Müller et al.
1980; Müller 1981; Posey 1985; Balée 1989; Clement et al.
2010). The difficulty of naturally dispersing the large,
heavy fruit also favors this hypothesis. If this view is
correct, then Brazil nut stands may be anthropogenic rather
than having natural origins (Posey 1984, 1985, 1986, 1990;
Anderson and Posey 1987; Balée 1989). If humans are
major actors in the geographical distribution and stand
creation of Brazil nuts, the relative roles of agoutis and
humans in the castanhais maintenance and structuring at
local scale remain poorly understood, as is the question of
human intention.
Humans collect and open fruits in the forest and then
transport sacks or baskets of Brazil nut seeds for use and
sale. Seeds occasionally fall during the fruit opening and
transport, affording a dispersal opportunity if the seed
germinates and grows in a favorable location. For example,
forest clearings made by humans may provide the preferred
environment for seed germination, seedling establishment
and sapling development. Unfortunately, records of the
active planting of Brazil nut trees by Amerindians are rare,
possibly because of the long period required for seed
germination (several months).
Here we examine and compare the population structure of
two sets of Brazil nut stands near communities in the Brazilian
Amazon. These two areas are distant from one another and
have different patterns of historical occupation and contem-
poraneous use. We examine whether evidence supports the
hypothesis that the population structure of B. excelsa trees in
the castanhais is due to human activities. We hypothesize that
disturbances caused by continuous human activities in the
forests, such as selective timber extraction, collection of non-
timber products, understory openings, and eventual fires,
increase the regeneration rate of the B. excelsa stands and
shift them toward a younger population structure.
The Species
Bertholletia excelsa is widely distributed in non-flooded
forests (terra firme) in the Amazon and Guiana Shield,
from 14° S to 5° N (Mori and Prance 1990). The species is
found in areas with 1,400–2,800 mm of annual rainfall, 24–
Hum Ecol (2011) 39:455–464
27°C mean annual temperature and an average relative
humidity from 79% to 86% (Diniz and Bastos 1974). The
tree is emergent, reaching up to 50 m in height and a
diameter at breast height (DBH) of >300 cm (Zuidema and
Boot 2002; Salomão 2009). The trunk is straight, with no
branches until near the top. The canopy is formed of well-
spaced branches and may reach up to 50 m in diameter. The
fruit, which contains the Brazil nut, is woody, indehiscent, and
spherical; it weighs 500–2,500 g and has a diameter of >10 cm.
This woody fruit holds 10–25 seeds that are edible after
removal of the woody protective tegument (Mori and Prance
1990). The agouti (Dasyprocta spp.) is thought to be the sole
important disperser of its seeds (Huber 1910; Ortiz 1995,
2002; Peres and Baider 1997; Tuck Haugaasen et al. 2010).
Most reproductive Brazil nut trees are greater than 40 cm
DBH, and the most productive trees are 80–160 cm DBH
(Viana et al. 1998; Zuidema and Boot 2002). Trees may live
for hundreds of years in terra firme forests, and the adults have
high survival rates (Zuidema and Boot 2002). Brazil nut
populations often contain 75–150 trees of >10 cm DBH
(Peres and Baider 1997), with stands of relatively high
densities for humid tropical forests (5–20 trees ha−1) mixed
with areas of low density (0.2 trees ha−1 (Mori and Prance
1990)). Studies of the population structure of B. excelsa have
found few trees >10 cm DBH that are not reproductive
(Salomão 1991; Nepstad et al. 1992; DHV 1993). Popula-
tions are typically dominated by intermediate-sized trees
(Zuidema and Boot 2002; Peres et al. 2003; Salomão 2009).
Bertholletia excelsa is heliophytic (Salomão 1991;
Scoles 2010) and depends on clearings for growth as well
as the survival of seedlings and saplings (Mori and Prance
1990; Viana et al. 1998; Myers et al. 2000; Oliveira 2000).
Plants appear in clearings as if they were pioneer species,
but, unlike pioneers, they remain after the forest has grown
into more advanced successional stages. For this reason, the
species is considered to be a long-lived pioneer tree
(Swaine and Hall 1987; Zuidema 2003). Many experimen-
tal studies have shown that Brazil nut trees grow best in
well-lighted open areas (Fernandes and Alencar 1993;
Yared et al. 1993; Tonini et al. 2008) and that growth is
much slower in forest shade in the understory layer (Kainer et
al. 1998; Myers et al. 2000; Peña-Claros et al. 2002; Scoles
2010).
Study Areas
The population structure was studied at two distinct sites: 1)
Trombetas River (hereafter Trombetas), on the northern side
of the Amazon River, near Oriximiná and Obidos in the state
of Pará, Brazil; and 2) Capanã Grande Lake (hereafter Capanã
Grande) on the Madeira River, near Manicoré on the southern
side of the river in the state of Amazonas, Brazil. These two
areas are 800 km from each other (Fig. 1).
Hum Ecol (2011) 39:455–464
Fig. 1 Map showing study areas within the Brazilian Amazon and the location of the sampled Bertholletia excelsa stands
The sampled castanhais in Trombetas are in conservation
areas, the Rio Trombetas Biological Reserve, the Sacará-
Taquera National Forest, the recently created Rio Trombetas
State Reserve, and in Quilombola Territories. The Quilombola
Territories were originally areas to which African slaves fled
from their owners, but they have now become semi-
autonomous communities. The Quilombola areas in this
study are on the Trombetas and Erepecuru Rivers, which
are administered by the Remaining Quilombo Community
Association of Oriximiná (ARQMO). The castanhais in
Capanã Grande all occur within the Lake Capanã Grande
Extractive Reserve, which is a direct-use conservation unit
co-governed by the riparian communities of the lake and the
Chico Mendes Institute for the Conservation of Biodiversity
(ICMBio).
At Capanã Grande, sampling units are located near each
other (1–16 km apart) on the plateaus near the lake and the
communities (100–8,750 m). At Trombetas, the sampled
castanhais extend over the Trombetas River and its western
tributaries’ watersheds. Most sampling units are >10 km
apart from the communities, and more than a third are
difficult to reach due to topography and obstacles, such as
river rapids and ravines. At Trombetas, the greatest distance
between two sampling units is 112 km.
In both areas, the climate is tropical, humid, and has a
short dry season with <100 mm rainfall month−1. Annual
rainfall varies from 2000 to 2500 mm, temperature varies
from 24° to 26°C, and relative humidity is normally >80%
457
(Brasil 1979; SUDAM 1984). In Manicoré, January to
March is usually the rainiest period (monthly rainfall
~300 mm), and only July and August may be considered
dry months (AGRITEMPO 2009). At Trombetas, rainfall is
highest from March–May, and the dry season runs from
August–November (IBAMA 2004). In both regions, the
Brazil nut stands occur in areas with yellow or yellowish red
latosoils which are the predominant soil types associated to
terra firme forest in the lower stretches of the Trombetas and
Madeira rivers (IBGE 2006).
Amerindian occupation and use differ between the two
study areas. The lower Amazon lost most of its native
population during the first two centuries of colonization
(c. 1500–1700). By the early eighteenth century a large
part of the Amerindian population had succumbed to
massacre and epidemics. The remaining Amerindians
fled to less-accessible areas. For example, in Trombetas,
they went upstream above the rapids and waterfalls of
the Trombetas, Cachorro, Mapuera, Acapu and Erepecuru
Rivers. This period was followed by the slow re-
population of the area by neo-indigenous people who
had been uprooted from their original homes and cultures
and were incorporated into Brazilian society by religious
missionaries (Porro 1992). During the 1800s, African
slaves and their descendents fled the towns and farms of
the state of Grão-Pará and established quilombos in the
middle and upper regions of the Trombetas and Erepecuru
Rivers. The inhabitants descended to lower regions when
458
slavery was finally abolished (1886) and commercial trade
in extractive products, such as Brazil nuts and timber
expanded (Acevedo and Castro 1998). Since approximately
1850, the quilombo communities have dominated the lower
and middle regions of the Trombetas and Erepecuru Rivers,
while the indigenous peoples have remained along the upper
regions of these rivers.
The colonial occupation of the Madeira River occurred
later, during the 1800s, with the creation of the states of
Amazonas (1850) and the rubber boom. Previous incur-
sions of colonists were temporary and limited to extractive
commerce and Jesuit missions (Menéndez 1992). In the
1700s, the government, fearing that the Madeira River
would serve as a route for contraband metals from the states
of Minas Gerais and Goiás, prohibited river navigation (in
1733) and spread rumors of dangerous Indians, such as the
Mura, along the river margins (Amoroso 1992). This
strategy deferred more intense contact between the colonists
and the Amerindians. Expansion of extractive commerce
during the latter half of the 1800s marked the beginning of the
colonization of the Madeira River that would give rise to a
society of mostly rural, mixed-race (caboclo) communities,
which endure today.
Methods
Data were gathered over three years at Trombetas (2007–
2009) and one year at Capanã Grande (2008). A total of 25
(125 ha) transects of 50×1,000 m were established within
stands of Brazil nut trees at the Trombetas River and 10
(49 ha) at Capanã Grande, the latter with one shorter
transect (800 m) due to spatial limitations. In each transect,
10 plots of 10×25 m, randomly placed at 100-m intervals,
were established to measure smaller trees (<10 cm DBH).
Within each plot, we identified all B. excelsa individuals
>10 cm DBH and noted their coordinates within the
transect. We measured the DBH (cm) and the diameter of
the canopy (as twice the length of the longest branch and
the next longest branch approximately perpendicular to the
first (m)), and estimated tree height (m). When the canopy
was irregular, we applied a conversion factor to adjust the
area estimated in quadrants. For example, if the canopy was
irregular because it had a particularly sparse or empty
quadrant, we multiplied by 0.75; if another quadrant was
missing, we multiplied it by 0.50; if most of the canopy was
absent, we multiplied by 0.25.
We also collected data on smaller Brazil nut trees (<10
DBH) in the 10 plots. We measured spatial coordinates, tree
height, the DBH of all saplings >1.3 m in height, and the
diameter at the ground for smaller saplings.
Trees were placed in classes to facilitate analysis.
Classes were young (10–40 cm DBH), young adults (40–
Hum Ecol (2011) 39:455–464
80 cm), productive adults (80–160 cm), mature adults
(160–200 cm), and old adults (>200 cm) (Viana et al. 1998;
Zuidema and Boot 2002). Plants less than 10 cm were
divided into two groups based on the presence of
endosperm. If present, the plant was considered a seedling;
if absent (absorbed), the plant was a sapling (Myers et al.
2000; Zuidema and Boot 2002).
Canopy Openness
The openness of the canopy was measured photographically
(Engelbrecht and Herz 2001). A 19-mm wide-angle lens was
used on a Ricoh GX100 camera. Pictures were taken at
130 cm above the ground using a tripod and with the camera
pointed at the zenith. Pictures were taken early in the
morning or late in the afternoon to avoid direct sunlight. One
photograph was taken from a random location within each of
the 50-m sections of the transects for a total of 20 photos per
transect (Nicotra et al. 1999). Digital images were analyzed
with Miramón 6.0, a program that counts pixels and then
calculates the percentage of white pixels in all pixels in the
photo. The calculated percentage was then considered the
index of canopy openness.
Data Analysis
Experiments in the classical sense are generally impossible to
apply on a large regional scale. The two historical situations
compared here cannot be ‘replicated’ over the Amazon. As a
consequence, the 25 plots in Trombetas and the 10 plots at
Capanã Grande cannot be considered as independent repli-
cations used in classical experimentation; they must instead be
technically classified as ‘spatial pseudo-replications’ (Hurlbert
1984). Because of that, we did not use inferential statistics to
compare the two ‘treatments’ in this unreplicated natural
experiment and our comparisons and conclusions, therefore,
are of an inferential and inductive nature.
Results
Patterns of Population Structure
The large basal area (10.4 and 7.1 m2 ha−1) and the high
proportion of the canopy occupied by Brazil nut trees (29%
and 34%) in Trombetas and Capanã Grande, respectively,
indicate that this species is dominant in both areas, and the
forests can be classified as monodominant or oligarchic (i.e.,
dominated by one or a few species, Peters et al. 1989;
Campbell et al. 2006). However, size distributions of the B.
excelsa trees contrast greatly in the two areas, with
intermediate to large sizes dominating at Trombetas and
smaller sizes (<80 cm DBH) at Capanã Grande (Fig. 2). Tree
Hum Ecol (2011) 39:455–464 459

Fig. 2 Distribution of Brazil nut 1,6

densities in 10 cm size class
intervals at Trombetas
(filled bars) and Capanã Grande
n=609
(white bars)
1,2 n=850

Density (tree ha-1)

0,8

0,4

0,0
10 30 50 70 90 110 130 150 170 190 210 230 250 270 >300

height was on average 3 m greater at Trombetas than in
Capanã Grande. Tree density was lower and the trees were
older at Trombetas, with only 7% of trees too young to
reproduce (Table 1).
At Trombetas, most trees were in the 80–160 cm DBH
range, whereas at Capanã Grande, most trees were <100 cm
DBH (76%), and approximately 26% were not reproductive
(DBH 10–40 cm). The distribution of tree sizes at Capanã
Grande was asymmetric, favoring small sizes (Fig. 2). At
Capanã Grande, the most productive trees (80–160 cm DBH)
were less well represented in the total population than at
Trombetas, even though their absolute density was greater
(4.5 versus 3.7 trees ha−1). The density of trees <130 cm
DBH was higher at Capanã Grande, and the density of
trees >130 cm DBH was higher at Trombetas (Fig. 2).
Capanã Grande thus exhibited a younger population
structure, with few trees >160 cm DBH (4%). At
Trombetas, 27% of trees were >160 cm DBH, and 10% of
trees were old adults (DBH>200 cm, Fig. 3). Old adult
trees were almost absent at Capanã Grande (1%). At
Trombetas, three trees were >300 cm DBH, whereas the
largest tree at Capanã Grande had a DBH of 267 cm.
Recruitment and regeneration were much greater at Capanã
Grande than at Trombetas (Table 1). Recruitment was
estimated by seedling density and regeneration by seedling,
sapling and juveniles densities. Recruitment is thus a subset

Table 1 Comparative analysis

of population structure (means ± Parameter Trombetas River Capanã Grande
standard deviation, if%, maxi-
mum and minimum) of B. excelsa Tree Density DBH>10 (tree ha−1) 6.8±4.1 12.5±7.8
in two regions of Brazilian Tree Density DBH>40 cm (tree ha−1) 6.3±3.9 9.2±4.7
Amazon: Trombetas River Tree Density DBH 10–40 cm (tree ha−1) 0.5±0.8 3.2±3.4
Region (Oriximiná, Obidos, PA)
and Capanã Grande Lake Seedlings ha−1 4.8±8.7 24.8±19.9
(Manicoré, AM) Saplings ha−1 1.0±3.1 4.4±5.3
% juveniles (DBH 10–40/DBH>10) 7 (0.0, 33) 18 (0, 41)
Seedlings adult−1 1.0±2.2 2.9±1.9
Saplings adult−1 0.1±0.4 0,5±0.6
Juveniles adult−1 0.1±0.1 0.3±0.2
Tree DBH (cm) 128.5±55.5 73.1±44.2
Trunk height (m) 22.5±4.9 19.2±4.0
Basal area (m2 ha−1) 10.4±5.6 7.1±2.5
Crown area (m2 ha−1) 2.865±1.250 3.385±1.351
Occupation canopy forest (%) 28.7 (11.8, 51.3) 33.8 (7.2, 51.0)
Mean Canopy Opening (%) 6.5 (49.5, 1.9) 6.6 (2.3, 30.1)
Proportion of canopy gap forest (>20% canopy opening) 0.01 0.02
460
Fig. 3 Ancient Brazil nut tree at Trombetas river region with >2 m DBH
of and an indicator of potential regeneration. The seedling
density at Trombetas (mean 4.6 seedlings ha−1) was five
times lower than at Capanã Grande (24.4 ha−1). Similarly, the
density of saplings (1.0 and 4.4 saplings ha−1 at Trombetas
and Capanã Grande respectively) and juvenile trees (0.5 and
3.2 juveniles ha−1 respectively) were also much lower at
Trombetas.
Both regions have very shaded understories with only 6–
7% light entrance. At Capanã Grande, however, there were
more small clearings (2% of total ground area with>20%
light) than in Trombetas (1%) (Table 1). Canopy openings
with 10–20% light entrance were twice as common at
Capanã Grande, and thus the probability of light reaching
the forest floor was higher in Capanã Grande.
Discussion
In the Amazon basin, the density of Brazil nut trees varies
from 1 to 23 trees ha−1. The density of B. excelsa at Capanã
Grande (12.5 trees.ha−1) is thus among the highest recorded
in studies for this species in the Amazon, with the
exception of stands in Cajari, Amapá state, Brazil (Table 2).
In other studies of castanhais, size class distributions are
usually more similar to that found at Trombetas, with
frequencies being greatest in intermediate classes (Zuidema
Table 2 Comparison of Brazil
nut tree density (>10 cm DBH) Location
in different regions of the
Amazon basin, by increasing Forest Reserve El Tigre, Beni, Bolívia
order of density Indigenous Area Pinkaiti, Pará State, Brazil
Saracá-Taquera Forest, Pará State, Brazil
River Trombetas Region, Pará State, Brazil
Cajari, Amapá State, Brazil
Lake Capanã Grande, Amazonas State, Brazil
Hum Ecol (2011) 39:455–464
2003). This trend was found also in Bolivia (DHV 1993;
Zuidema and Boot 2002), in the Brazilian state of Pará
(Salomão 1991, 2009) and in most of the studies examined
in the meta-analysis carried out by Peres et al. (2003).
The castanhais at Capanã Grande are in other ways
strikingly different, with a distribution strongly skewed to
the left, favoring trees with DBH<80 cm (Fig. 2). While
this pattern is atypical, it is not unknown in the Amazon. At
Nova Esperança, Xapuri (in the state of Acre), 58% of
Brazil nut trees have <80 cm DBH. At the Chico Mendes
Extractive Reserve in Acre, ~60% of trees have <100 cm
DBH (Wadt et al. 2005). Interestingly, on the Madeira
River, those castanhais that are farthest from communities
tend to have size distributions similar to that of the stands at
Trombetas (Scoles 2010).
At Capanã Grande, trees tended to be smaller (mean
DBH=73 cm) than at Trombetas (129 cm), which indicates
a younger population at the former location. Populations of
B. excelsa in the Amazon, like that of Trombetas, usually
have an average DBH >100 cm (Peres et al. 2003).
Conversely, stands closer to Amerindian or Extractive
Reserves or that are known to be persistently used by local
communities tend to have an average DBH similar to that
of Capanã Grande. For example, mean DBH at the Pinkaiti
Indigenous Area in Pará was 73±4 cm; at Nova Esperança,
Acre, it was 74±4 cm, and at the Chico Mendes Extractive
Reserve, Acre, it was measured 86±5 cm (Peres et al. 2003;
Wadt et al. 2005).
Seedling density at Trombetas (mean, 4.6 seedlings ha−1)
is the lowest published to date (Baider 2000; Zuidema and
Boot 2002; Zuidema 2003; Wadt et al. 2008). The ratio of
seedlings to adults is also the lowest reported to date in the
literature. In contrast, seedling density at Capanã Grande
(24.4 ha−1) is similar to densities reported elsewhere
[29.8 ha−1, Baider (2000) in Para; 30–52 ha−1, Zuidema
(2003) in Bolivia].
Thus, the castanhais at Capanã Grande seem to have a
greater regenerative capacity than those at Trombetas
because of the much higher density of juveniles, saplings,
and seedlings. Castanhais in Capanã Grande are intermedi-
ate in terms of regeneration (18% young), which is within
the range of 1–52% found in other areas (Peres et al. 2003).
Yet, that percentage increases inversely with distance from
Area (ha) Density Source
12 1.7 Zuidema and Boot (2002)
28.5 4.8 Peres and Baider (1997)
203.7 5.6 Salomão (2009)
125 6.8 This study
22.6 12.2 Baider (2000)
49 12.5 This study
Hum Ecol (2011) 39:455–464
communities; closer castanhais have 21–40% young plants
(in comparison to the usual 18%) and are similar to studied
areas that have the greatest proportion of non-reproductive
trees. This regeneration trend found in the Capanã Grande
area is clearly contrary to the conclusions of Peres et al.
(2003), which postulated that more persistently harvested
castanhais exhibited poor regeneration and that these
intensely exploited populations may eventually succumb
to recruitment bottleneck.
The entirety of the population structure parameters found
here shows that Brazil nut stands are younger and have
greater regeneration potential at Capanã Grande. In Trombetas,
the canopy was more closed and higher, and the stand
contained less-dense, older trees. What are the most likely
explanations for differences in the population structure
between the two areas? Although climate, soils and other
abiotic factors may have some unmeasured importance, we
believe that two anthropic factors seem most likely to explain
these different patterns. The first relates to the current trends in
occupancy and use of the castanhais, and the second relates to
their historical occupation.
At Trombetas, the castanhais are less often visited by
local people. This pattern is certainly due to the greater
distance of the stands from the communities, and thus the
trees are only visited during harvesting (≤4 month yr−1). In
addition, at Trombetas, many of the castanhais are near
rapids that must be crossed by canoe or in steep and
difficult terrain. Thus, gathering and transporting the
harvest is much more difficult. Additionally, staging sites
for transport are often very far from where seeds are
harvested. As a result, the product must be carried over
relatively great distances. Finally, legal regulations restrict
access to harvesting periods in 60% of the area within the
Biological Reserve of Rio Trombetas. In the ten castanhais
within the domain of the quilombos territories, people tend
not to visit the stands outside of the harvest season due to
limited accessibility.
The situation is very different at Capanã Grande, where
castanhais are near the communities. Comparatively, these
stands are much more accessible, and collection and
transport of Brazil nuts is easier. Several stands are near
small rivers that can be used during high water. Aside from
Brazil nut collection, other extractive activities occur
when Brazil nuts are not being harvested, including rubber
tapping, fruit collection, collection of fiber from vines and
tree barks, and timber exploitation. Also, many stands are
in forest fragments that are separated by land used for
agriculture. This fragmentation increases the availability
of light in the understory and creates edge effects and
other perturbations. Despite the similarity of average light
levels in the two forests, the distribution of light is very
different because more openings are found at Capanã
Grande.
461
In other studies in the southwestern Amazon, the size
distribution (i.e., a large proportion of young trees, average
70–80 cm DBH) is similar to that at Capanã Grande but has
a lower total tree density (Viana et al. 1998; Wadt et al.
2005). In other stands, seedlings are always present despite
the low density of reproductive trees (Viana et al. 1998;
Zuidema and Boot 2002; Wadt et al. 2005). The presence of
younger plants of B. excelsa is usually attributed to open
forests with plenty of light, as compared to closed, denser
forests where light may be limiting (IBGE 2004; Wadt et al.
2005). Hence, this mechanism may explain recruitment in
our study area at Capanã Grande, where human activities
and settlements have opened the forest, in comparison with
Trombetas.
The history of occupation is also very different in the two
study areas and may explain the contrasting population
structures of the B. excelsa stands. The Trombetas-
Erepecuru watershed is a region particularly rich in
archaeological sites (Magalhaes 2007), black earths (Kern
and Kämpf 1989), pottery (Hilbert 1955), and rock paintings
(Pereira 2003; Pereira et al. 2009). These archaeological
findings suggest that this region probably supported dense
Amerindian populations and, consequently, experienced
extensive forest disturbance during pre-Columbian times. It
is quite likely that the Brazil nut, which grows well in
disturbed areas, was favored by the slash-and-burn agriculture
of the pre-Colombian societies (Posey 1984, 1986, 1990;
Peters 2000; Roosevelt 2000; Campbell et al. 2006).
It is notable that a tree-ring study of 22 Brazil nut trees
(87–145 cm DBH) from the Trombetas region measured
ages from 228 to 502 years (Schöngart et al. unpublished
data). These data suggest that the biggest trees found in the
Trombetas area, those with DBH >300 cm, may reach ages
of 600 years or more. Interestingly, the estimated age of the
majority of the Brazil nut trees in this study was between
300 and 450 years, which probably coincides with a period
of depopulation of the Amerindian people following
European contact, especially along the lower and middle
stretches of the main rivers (Porro 1992). The current
population structure of B. excelsa likely owes its origins to
the interval between land abandonment due to Amerindian
depopulation (sixteenth-seventeenth centuries) and the
establishment of the quilombos (nineteenth century). Thus,
it appears that the abandonment of the Trombetas area
resulted in the development of the current B. excelsa
population structure, with a high proportion of old trees
and scarce regeneration.
In contrast, Capanã Grande likely owes the formation of its
castanhais to the growth in human presence that intensified in
the late nineteenth century, following colonization of the
Madeira River and the beginning of the rubber boom. This
timing may explain the domination of trees <80 cm DBH
(which are approximately 150 years old (Zuidema and Boot
462
2002)) in this population. The early riverine communities
were probably established around existing stands of rubber
(Hevea brasiliensis) and Brazil nut trees, which would have
then favored the rejuvenation of the castanhais as previously
described.
Therefore, we hypothesize that when native Amazonian
populations were decimated post-conquest in the Trombetas,
forest disturbance was reduced, allowing for the shift in
demography toward over-mature stands. Those human
populations in Trombetas never fully recovered; rather,
they were replaced by low-density black and caboclo
populations, each with different subsistence strategies. In
the Madeira, the decimation occurred later, and the
Amerindian replacement by caboclos was more gradual and
complete. The Madeira region probably recovered pre-
conquest population densities, whereas the Trombetas most
likely did not. The contemporaneous stands in Capanã Grande
probably have a younger population profile, not only because
the castanhais are expanding but also because tree death is
higher due to the more intensive use of fire and other
disturbances in the landscape.
Forests once considered primary and pristine, such as the
‘native’ castanhais in Trombetas, are now being recognized
as regenerated forests that have recovered after the
abandonment of lands that were previously used for
agriculture. In tropical America, Africa and Asia, areas that
were once considered virgin or pristine have been discovered
to have been occupied and significantly perturbed by earlier
human settlements that, once abandoned, allowed forests to
regrow (Willis et al. 2004). In tropical Latin America, many
monodominant forests are the consequence of management
and manipulation of the original forests by pre-Colombian
and subsequent societies (Anderson 1990; Peters 2000;
Campbell et al. 2006). Today, it is thought that the
occurrence of many high-density stands of fruit trees,
monospecific stands of palms, and other anthropogenic
forests are evidence that at least 11% of the Brazilian
Amazon was anthropogenically modified (Balée 1989).
Here, we suggest that the differences in the population
structures of the Brazil nut stands in Trombetas and
Capanã Grande may be a consequence of pre- and post-
colonial human occupation and manipulation of the
forest. Confirmation of this hypothesis will require
archaeological studies which would improve understand-
ing of the impact of Amerindians on pre-colonial tropical
forests. The Brazil nut stands at Capanã Grande exhibit a
younger structure likely due to more recent human
perturbations. In contrast, at the Trombetas River, trees
are older, and the current recruitment is much lower. The
lower recruitment is in part due to the greater density of
big trees (and the consequent low-light conditions in the
understory for the past several centuries) and in part due
to few recent perturbations.
Hum Ecol (2011) 39:455–464
Regeneration of the heliophytic, long-lived Brazil nut is
therefore likely to be most influenced by ecological factors
that allow light to reach the understory, such as low-
intensity forest disturbances mediated by humans. In
addition, the ages of the stands and population structure
indicate a history of human influence, both old (pre-
colonization) and recent (post-colonization). Hence, we
suggest that these stands are anthropogenic forests that are
distinctly influenced by humans. Finally, we found no
scientific evidence that there should be restrictions on seed
harvesting as a means to improve regeneration rates of the
Brazil nut stands.
Acknowledgments We thank the Protected Areas Program of the
Chico Mendes Institute for Biodiversity Conservation (ICMBio), the
National Council for Scientific and Technological Development
(CNPq), the Scholarship Program of the International Education
Institution of Brazil (IIEB), the Brazil Nut Germplasm Bank project of
the National Institute for Amazonian Research (INPA), the Mineração
Rio do Norte Company (MRN), and the Chico Mendes Biodiversity
Institute (ICMBio) for their logistical and financial support. We
sincerely thank the team of ICMBio at Porto Trombetas and Manicoré
for their collaboration and logistical support during field work,
especially Gilmar N. Klein, Altemar Lopes Silva, Vivian Mara Uhlig
and Valmir Raimundo Lopes da Silva. Thanks to Paulo M. Alencastro
and to Carlos Palacio for the map, Charles Clement and André
Junqueira for the literature on historical ecology and helpful
discussions, David Bertran for suggestions for the analyses, Jochen
Schöngart for the data on the tree-ring studies, and James J. Roper and
Maristerra R. Lemes for constructive comments on the manuscript.
Our heartfelt thanks go to all the families of the communities that were
involved in this project. Research Grant to RG was provided for
CNPq.
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