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Ecology and management of the Brazil nut tree (Bertholletia excelsa)

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Ecology and management
of the Brazil nut tree
(Bertholletia excelsa)

Pieter A. Zuidema

PROMAB Scientific Series 6

PROMAB (Programa Manejo de Bosques de la Amazonia Boliviana) carries
out research, extension and education activities to support forest users of the
Bolivian Amazon region in achieving sustainable management of forest
resources and biodiversity conservation.

PROMAB, Casilla 107, Riberalta, Beni, Bolivia (e-mail: promab@cotas.net),

or c/o Department of Plant Ecology, Utrecht University, PO Box 80084, 3508
TB Utrecht, the Netherlands (e-mail: promab@bio.uu.nl), Internet:
www.promab.org

P.A. Zuidema
Demography and management of the Brazil nut tree (Ber tholletia excelsa)
ROMAB Scientific Series 6
PROMAB - Riberalta, Bolivia
ISBN 90-393-3390-4

© 2003 PROMAB, Pieter A. Zuidema

All rights reserved. No part of this publication apart from bibliographic data
and brief quotations in critical reviews, may be reproduced, re-recorded or
published in any form including photocopy, microfilm, electronic and
electromagnetic record, without written permission.

This publication was financed by the Ministry of Foreign Affairs of the

Netherlands, projects BO009701 and BO009703, and by Utrecht University,
the Netherlands.

Cover layout: Femke Bulten

Text layout: Marjolein Kortbeek-Smtihuis
Photos: Pieter Zuidema, Arienne Henkemans
Data: Part of the data used in this publication is a product of the
activities of the research and extension staff of PROMAB.
 Preface

This book contains a compilation of information on the ecology and

management of a tropical forest tree of great economic importance: the Brazil
nut tree, Ber tholletia excelsa. The information presented in the book is
obtained from papers published in international scientific journals, “grey”
literature, unpublished reports, and unpublished data. Information on the Brazil
nut tree that was considered to be useful to understand the ecology or assist
the management of the species has been selected. As much as possible, the
information is presented in a quantitative way: a large number of figures and
tables summarize essential research findings. When available, information from
different areas or different sources are presented to allow for comparison.
A book that compiles current knowledge can only be written if sufficient
publications exist on the topic. We therefore thank all those who have
published their research findings on the ecology and management of the Brazil
nut tree. Specific thanks are due to those investigators who have made available
their unpublished research findings, unpublished papers or reports, figures and
plates: Bobbi Angell (taxonomic plate), Claudia Baider, Alan Bojanic,Yaskara
Hayashida Oliver, Karen Kainer, Jhon Leigue Gómez, Scott Mori, Enrique
Ortiz, Marielos Peña-Claros and Lourens Poorter. Without their assistance, the
book not have been filled with a large amount of up-to-date research results.
The information presented in this book is clearly not exhaustive. Certainly
more information than currently available to the author exist on this topic.
Furthermore, there remain some important gaps in our knowledge on the
Brazil nut tree, in spite of the fact that it is a intensively studied species. Some
of the omissions in knowledge are identified and mentioned in the book.
The research findings presented should be interpreted with some care and
extrapolations of certain information is difficult or not possible. Large part of
the information is specific to a certain area, to trees of a certain size, to a
specific study method or experimental treatment, and so on. Therefore, the
information provided in figures, tables and in the text should always be
interpreted in the context of the specific situation.
Lastly, we hope that this book may contribute to an increased
understanding, the sustainable use and proper management of the Brazil nut
tree.

Albert Bokkestijn, General Coordinator PROMAB, Bolivia

René Boot, former General Coordinator PROMAB
Pieter Zuidema, author

February 2003, Riberalta, Bolivia and Utrecht, the Netherlands

Ta b l e o f c o n t e n t s

Preface 3

Table of contents 5

Part A Economic importance of the Brazil nut 8

Chapter 1 Brazil nut exploitation 9

1.1 Past and current exploitation 9
1.2 Socio-economic significance of Brazil nut production 11
Chapter 2 Brazil nut production systems 13
2.1 Phases in the production system 13
2.2 Two important issues for Brazil nut production 15

Part B Ecology of the Brazil nut tree 18

Chapter 3 Taxonomy and geographical distribution 19

3.1 Species description 19
3.2 Geographical distribution 21
Chapter 4 Size structure and spatial distribution of populations 23
4.1 Densities of Brazil nut trees 23
4.2 Spatial distribution 25
4.3 Size distributions 26
Chapter 5 Reproductive ecology 29
5.1 Flowering 29
5.2 Fruiting 31
5.3 Attaining reproductive size 32
5.4 Seed production 33
Chapter 6 Dispersal ecology 41
6.1 Dispersal system 41
6.2 Seed handling by agoutis 43
6.3 Dispersal distances 45
6.4 Germination under natural conditions 47
Chapter 7 Seedling ecology 49
7.1 Density and natural conditions of seedlings 49
7.2 Seedling survival under natural conditions 52
7.3 Seedling growth under natural conditions 53
7.4 Seedling growth in controlled experiments 54
 Chapter 8 Adult ecology 59
8.1 Tree architecture 59
8.2 Tree diameter growth 61
8.3 Tree survival 64
8.4 Tree ages 64
Chapter 9 Population dynamics 67
9.1 Relevance of studying population dynamics 67
9.2 The life cycle 68
9.3 Population matrix models 69
9.4 Methodology of population dynamics study 71
9.5 The population growth rate 72
9.6 Important processes and size classes in the life cycle 73
9.7 The stable population structure 75
Chapter 10 Conclusions:
life history of the Brazil nut tree 79

Part C Management of the Brazil nut tree 82

Chapter 11 Impact of exploitation 83

11.1 Intensity and consequences of Brazil nut extraction 83
11.2 Impact of exploitation on natural regeneration 84
11.3 Impact of exploitation on population dynamics 87
11.4 Impact of exploitation on future resource availability 87
11.5 Impact of exploitation at ecosystem level 88
Chapter 12 Enrichment planting and plantations 91
12.1 Ways to increase Brazil nut production 91
12.2 Germination of Brazil nut seeds under
controlled conditions 92
12.3 Seedling survival and growth in enrichment experiments 93
12.4 Strip planting as an enrichment method 96
12.5 Plantations of Brazil nut trees 99
Chapter 13 Conclusions:
sustainable exploitation and successful enrichment 103

Bibliography On the Brazil nut tree 105

Other literature used in this book 110
Selected Internet sites with relevant information
on the Brazil nut tree 111
PROMAB publications 112
 Part A

Economic importance
of the Brazil nut
 Chapter 1

Brazil nut exploitation

1.1 Past and current exploitation

Brazil nuts have been collected for many ages by Amerindian peoples in
those parts of the Amazon region where Brazil nut trees (Bertholletia excelsa)
occur. Commercial extraction and export outside the region commenced at
the beginning of the 20 th century, after the species was described by the
botanists Humboldt and Bonpland in 1807.
Brazil nuts are commercially collected from lowland rainforests in Brazil,
Bolivia and Peru. Almost all production is harvested from the natural tree
population. Brazil nuts are traded in two ways: as nuts that are still contained
in their shell (in-shell) and as the shelled nuts (kernels). Kernels weigh around
35% of the in-shell nuts. Brazil nuts are traded in five classes based on size and
quality of the nuts.
The export of Brazil nuts has gradually increased since the beginning of
the 20th century, although there are large fluctuations in the production. For
instance, during World War II the production strongly dropped. Fluctuations
in production are due to the price paid for Brazil nuts and to climatic factors
that influence sthe natural fruit production of Brazil nut trees. Since 1970, the
world production averaged 46 tons per year of in-shell nuts (equivalent to 16
tons of shelled nuts), with fluctuations between 28 and 65 tons. Over this
period of time no increasing or decreasing trend in the production is found.
Figure 1.1 shows the development of the total world export of Brazil nuts
since 1970, for the three producing countries. The figure shows that Brazil is
traditionally the largest exporter of Brazil nuts, but that this position is
gradually taken over by Bolivia. Bolivia is currently the largest producer of
shelled nuts (kernels). The lower Brazilian production is probably due to the
comparatively high labour costs and the deforestation in that country.The total
production of Peru has increased over the last 30 years but its share in the
10 • Chapter 1

Brazil nut production

Export [1000 tons]

60 Brazil Bolivia Peru

40

20

0
1970
1972
1974
1976
1978
1980
1982
1984
1986
1988
1990
1992
1994
1996
1998
Year

Figure 1.1
Development of the international Brazil nut trade for the three producing countries. The export values
for the three countries are in thousand tons (1 ton = 1000 kg) of in-shell nuts (i.e. nuts that are not
shelled). Production figures of shelled nuts (kernels) were converted using a conversion factor of 35%
(a shelled nut weighs 35% of an in-shell nut). Information for the year 1992 is missing: the average
of 1991 and 1993 is shown for that year. Source: LaFleur (1992), Man-Producten (1998), Bojanic (2001).

World market is less than 15%. The area of forest rich in Brazil nut trees in
this country is much smaller than in Bolivia and Brazil.
The Brazil nut trade is part of an international market for edible nuts
with an annual value of approximately US$ 2.5 billion. The share of Brazil
nuts in this market is only 1.5%, on average. The world trade in Brazil nuts
amounted to US$ 57 million in 1998.
The international price paid for Brazil nuts fluctuates largely from year
to year. The fluctuation in the price over the last 20 years is shown in Figure
1.2. In general, the international price increased annually by US$ 0.03 per
pound during this period. The price of the Brazil nut is influenced by the
price of other edible nuts (groundnuts, cashew nuts, desiccated coconut,
hazelnuts, almonds, walnuts, macadamia nuts and pistachios).The price of Brazil
nuts strongly depends on that of other nuts, as Brazil nuts are used as a
substitute for other nuts when these become expensive. As a result, price
behaviour is hard to predict.
Half of the World Brazil nut production is imported by European
countries, mainly Great Britain, Germany and the Netherlands; some 40% goes
to the USA and the rest to Australia, South Africa and some Latin American
countries. Brazil nuts are known by different names in different countries:
castanha-do-brasil or castanha-do-pará (Portuguese), castaña, almendra or
nuez del Brasil (Spanish), Paranuss (German), noix du Brésil (French) or
paranoot (Holland). Brazil nuts are sometimes also called Amazonian nuts.
Although the vast majority of Brazil nuts is traded and used as nuts,
different products can be obtained. Brazil nuts can be used to produce oil,
candies, cookies and marzipan. Brazil nut oil is used to make soaps, shampoos
and hair conditioners, the latter being traded internationally. Furthermore,
 Brazil nut exploitation • 11

Brazil nut prices

2

Price FOT [US$/pound]

1,5

0,5

0
1980

1982

1984

1986

1988

1990

1992

1994

1996

1998
Year

Figure 1.2
Development of the international price paid for Brazil nuts. The price is in US dollars paid in the
harbour of Rotterdam. One pound equals 0.454 kg. Source: Man-Producten (1998).

Brazil nuts are used in nut mix, with grains and raisins in muesli (‘Rainforest
crunch’), or surrounded by chocolate. Brazil nuts also have medicinal use: due
to their high content of selenium, they sare used for treating several types of
cancer, among others prostate cancer.

1.2 Socio-economic significance

of Brazil nut production

Brazil nuts are an important source of income in the regions where Brazil
nut trees occur at high densities in the natural forest. For instance, in Northern
Bolivia (department of Pando and part of department of Beni) the regional
economy strongly depends on the collection and processing of Brazil nuts.
This is illustrated in Figure 1.3: Brazil nuts have a large share in the total export
revenues. Since the decrease in rubber trade, this share has strongly increased:
from US$ 3 mln in 1980 to US$ 31 mln in 1998. In Northern Bolivia, a large

Importance of Brazil nut for the Northern Bolivian economy

1980 (US$ 12mln) 1986 (US$ 28 mln) 1998 (US$ 65 mln)
Other Brazil Other Brazil Other Brazil
nut nut nut

Rubber Rubber

Figure 1.3
Share of Brazil nuts in the total export value of the Bolivian Amazon region (Pando and part of Beni).
The total export value is shown in parentheses. “Other products” include timber and palmheart (only
in 1998). Source: Bojanic (2001).
12 • Chapter 1

share of the working population derives part of their income from the
collection, transport or processing of Brazil nuts. Around 10% (5-6 thousand
persons) of the total population of the main town (Riberalta) is employed in
the shelling process in Brazil nut processing plants. The collection of Brazil
nuts in the forest is carried out by an estimated 12-13 thousand persons. Half
of these people are seasonal labourers who move to the forest for 3-4 months
per year, taking along their families.
In short, Brazil nuts provide a substantial input of revenues for both the
regional economy and for a large number of households in Northern Bolivia.
Furthermore, it provides an important source of income to the poor part of
the population.
The great importance of Brazil nuts for the regional economy of
Northern Bolivia is probably rather special. Apart from the region around
Puerto Maldonado, department of Madre de Dios in Peru, there are probably
no other regions where Brazil nuts are such a strong pillar for the regional
economy. In the past, the Brazil nut industry has been important in the
Brazilian regions of Marabá and Tapajos in the state of Pará, the region Abufari
in the state Amazonas, and the state of Acre. But this economic importance
has decreased over the last decades.
 Chapter 2

Brazil nut production systems

2.1 Phases in the production system

Different phases can be distinguished in the production system of Brazil

nuts. Figure 2.1 provides an overview of these phases.
Collection of Brazil nuts is carried out mainly in primary rainforest and
to a smaller extent in secondary forests or degraded areas. Brazil nut collectors
traverse the forest searching for Brazil nut trees. They use a system of narrow
forest trails that connect the different (groups of) Brazil nut trees. This is done
during the wet season, when the fruits of the Brazil nut tree fall from the tree
crowns. Collectors systematically search for fruits in the area below crowns of
Brazil nut trees. As the tough woody fruits remain closed when fallen on the
forest floor, they have to be opened with a machete to remove the seeds (the
actual Brazil nuts). The Brazil nuts are taken out of the fruits and put in large
bags. The bags are carried to a nearby house or settlement, where they may
be stored for some time. Seeds are then further transported to processing plants

In forest In processing plant

• Collection • Drying
- Search and open fruits • Soaking nuts
- Remove seeds (nuts) • Steam boiling
- Carry nuts in bags • Cracking shells
• Transport to Transport • Classifying kernels
gathering points by boat or • Oven-drying kernels
• Transport to settlement truck • Packing
• Storage in storehouses

Trading by brokers;
international transport
Importing countries

Figure 2.1
Phases in the collection, processing and trade of Brazil nuts.
14 • Chapter 2

in towns, sometimes via larger settlements that have access to a river or road.
The forest areas where Brazil nuts are collected may be large concessions
or forest estates owned by companies or individuals (baraccas), small forest
plots owned by individual farmers, communal forest areas, or forest reserves
(extractive reserves). In Bolivia, Brazil nuts are mainly collected in large forest
estates (roughly two third of the production) and in communal forest areas
(one third). In Brazil, a third source is of importance: the so-called “extractive
reserves”, where forest conservation and wise use are combined. In Peru, the
situation is somewhat different: a large area of forest where Brazil nuts are
collected is handed as small concessions (<1000 ha) to individual families.
Forest estates where Brazil nuts are collected are often former rubber
producing areas, as the collection of Brazil nuts has been carried out in
combination with that of rubber (from Hevea brasiliensis) since the early
1930s. As rubber was tapped from April to October, the remaining period of
the year (November to March) could be used for the collection of Brazil nuts
and for the agricultural production.Thus, rubber tapping, Brazil nut collection
and subsistence agriculture were part of an annual ‘agro-extractive cycle’. This
extractive system allowed rubber tappers and their families to stay in the forest
throughout the year. The system began to break down with the decreasing
rubber prices in the late 1980s, causing rubber tappers to move out of the
forest.

The processing of Brazil nuts is mostly carried out in medium-sized to

large towns. Processing contains the following phases (see Figure 2.1). First,
nuts are collected in storehouses where they are air-dried. The actual cracking
of the nuts to remove the woody shell from the kernel (the edible nut itself)
is carried out in two ways: manually and mechanically. The majority of nuts
are cracked manually after the in-shell nuts have been soaked in water, steam-
boiled and immersed in cold water to loosen the shell from the kernel. Then,
the nuts are manually shelled using a small device (a vice) to put pressure on
the shell. This has to be carried out with care in order not to damage the
kernel (damaged kernels have a lower value).
Mechanical cracking is carried out in a few factories only, as it requires
sophisticated machinery. Mechanical cracking involves dry-freezing of the in-
shell nuts, which are then dropped from high altitude (approximately 20 m)
in a large cylinder. When the frozen seeds hit the ground, their shells crack
due to the impact. In Northern Bolivia only two large processing plants are
equipped with the mechanised cracking machines. Initial investments for these
machines are very high, but clearly, labour costs for manual cracking are
reduced to zero.
 Brazil nut production systems • 15

After cracking, the kernels (nuts without shell) are classified on the basis
of their size and degree of damage. Then, nuts are oven-dried and packed for
shipment.
There are large differences in the capacity of processing plants for Brazil
nuts. In Northern Bolivia, factories can be classified in three categories. Large
factories (8 out of 20 factories) process over 2,500 tons of raw nuts per year,
with an average output of 670 tons of kernels (equivalent to 4% of the world
production) and an annual turnover of US$ 2 mln. Medium-sized factories
produce on average 370 tons of kernels, whereas small factories have an output
of 100 tons.

2 . 2 Tw o i m p o r t a n t i s s u e s
for Brazil nut production

In this section, two currently important issues regarding the production

of Brazil nuts are discussed: (1) the problem of aflatoxin in Brazil nuts and (2)
the certification of Brazil nuts. These two issues may greatly influence the
market opportunities for Brazil nuts, and in this way deserve some attention.
1 • The problem of aflatoxins. Aflatoxins are carcinogenic toxins produced
by two species of moulds (fungus) of the genus Aspergillus. Many countries
have set norms for the maximum allowable level of aflatoxins in edible
products, usually between 2 and 50 ppb (parts aflatoxins per billion parts).
Aflatoxins are produced by these moulds on various types of food sources,
among which are Brazil nuts. Brazil nuts are probably infected by these moulds
in the forest, in storehouses, or in the processing factories. In general, only a
small fraction of the nuts contains aflatoxins, but the high aflatoxin
concentration in these seeds may cause the average toxin level in a sample of
nuts to exceed the maximum allowable value.
The possible solution to the aflatoxin problem is to remove the small
amount of infected nuts. As part of the toxins are fluorescent, they may be
identified using ultraviolet light. Nevertheless, there is still the risk that the
sample tested in importing countries by chance contains one infected seed,
thus causing the entire shipment of Brazil nuts to be rejected for import.Thus,
solutions should probably be found in removal of infected nuts in combination
with testing of samples before shipment in the region where Brazil nuts are
processed. Such tests should be carried out by a world-wide acknowledged
laboratory. (Source Williams & Wilson 1999)
2 • Certification of Brazil nuts. The organic nature of collecting Brazil nuts
(gathered in natural forests, no use of fertilisers and pesticides), make Brazil
nut production suitable for certification. Forest products can be certified if
16 • Chapter 2

they are produced in an environmentally sustainable and socially acceptable

way. Forest certification is mostly carried out using the set of Principles and
Criteria for sustainable forest management established by the Forest
Stewardship Council (FSC). Indicators to judge whether Brazil nut production
meets the criteria of sustainable forest management have been developed,
initially for Bolivia, but recently also for Peru and Brazil.The sets of indicators
published by the Bolivian certification organisation show that improvements
in the production process of Brazil nuts are mainly required in the following
fields: land use rights, health conditions and medical care for Brazil nut
collectors, labour contracts for collectors and factory employees, hunting
pressure in Brazil nut forests and working conditions for Brazil nut crackers
in processing factories. (Source: CFV 2001)
 Part B

Ecology of the Brazil nut tree

 Chapter 3

Ta x o n o m y
and geographical distribution

3.1 Species description

The genus of the Brazil nut tree - Bertholletia - belongs to the family of
Lecythidaceae. Species belonging to this family occur in tropical regions
throughout the world. The Brazil nut tree itself, Ber tholletia excelsa, is the

Figure 3.1
Taxonomic plate of the Brazil nut tree showing leaves and inflorescences (A), detail of flowers (cross
section of androecium) with the inward bent hood (B), cross section of ovary (C), ovary and calyx (D),
whole fruit (E), seed in seed coat (F) and seedling with cotyledon (G). Reprinted with permission from
Mori (1992). Illustration by Bobbi Angell.
20 • Chapter 3

Peru



Brazil
Bolivia

Figure 3.2
Distribution of natural populations of the Brazil nut tree (Bertholletia excelsa) in the Neotropics.
Points are locations where taxonomic samples of the species have been collected. Adapted from Kainer
et al. (1999), with permission.

only species in its genus, and was first described taxonomically by the botanists
Humboldt and Bonpland in the begining of the 19th century. Brazil nut trees
are large trees, up to 50 m tall and over 3 m in diameter at breast height (DBH).
Adult trees are usually emergent, i.e. their crowns overtop the forest canopy.
The stem is without buttresses and the bark contains conspicuous longitudinal
crevices. Leaves do not consist of different leaflets and are alternately placed
along branches (i.e. not placed opposite to each others). Flowers are large, c. 3
cm in diameter, and fleshy, and possess an inward bent hood that allows
pollinators to enter the flower (Mori & Prance 1990). Seeds are large, possess
a woody seed coat (also called testa) and are contained in large woody fruits.
The seeds are customary called Brazil nuts. The fruit - called “pod”,
“pyxidium”, or “coco” (Spanish) - has a diameter of >10 cm and possesses a
thick and very tough wall. A plate showing different parts of the Brazil nut
tree is shown in Figure 3.1. After germination, the seed (cotyledon) becomes
part of the stem, a feature that greatly facilitates the identification of seedlings
in the field (see Figure 3.1). Further information on the reproductive ecology
is included in Chapter 5.
 Ta x o n o m y a n d d i s t r i b u t i o n • 21

3.2 Geographical distribution

The Brazil nut tree occurs throughout the Amazon region in South
America. The species can be found in the Guianas, Colombia,Venezuela, Peru,
Bolivia and Brazil (Figure 3.2). However, sufficiently high densities to make
nut collection profitable only occur in parts of Brazil (States of Pará, Amazonas,
Acre and Rondônia), Peru (Department of Madre de Dios) and Bolivia
(Departments of Pando and parts of the Departments of Beni and La Paz).
Brazil nut trees occur in non-flooded (ter ra fir me) forest areas, on
nutrient-poor and well-drained utisol and exisol soils (Peres & Baider 1997
and references therein) and at elevations lower than 800 meters above sea level.
Brazil nut trees are reported to occur in areas with an annual precipitation of
1400-2800 mm, with 2-7 dry months with less than 60 mm per month of
rainfall (Almeida 1963, Diniz & Bastos 1974). Precipitation for the town of
Riberalta, in the Brazil nut region of Northern Bolivia is provided in Figure
3.3.

Precipitation Riberalta - Bolivia

Precipitation [mm month-1]

300

200

100

0
1 2 3 4 5 6 7 8 9 10 11 12
Month

Figure 3.3
Precipitation graph for Riberalta, Beni, Bolivia. Numbers refer to months from January - December.
Data are from AASANA, Riberalta over the period 1948-1998. Vertical lines are standard deviations.
The horizontal line denotes the 60 mm / month value to show that three out of 12 months have a
precipitation lower than 60 mm.
 Chapter 4

Size structure and spatial

distribution of populations

4.1 Densities of Brazil nut trees

The Brazil nut tree occurs at highly varying densities. There are reports
of very high densities (over 26 trees per ha) but also very low figures (much
less than 1 per ha) are mentioned (Mori & Prance 1990, Peres & Baider 1997).
Table 4.1 provides a brief overview of a number of inventories at the scale of
>10 ha. Note that this overview is certainly not comprehensive and should
be considered as a rough reference; not as the basis to extrapolate over larger
areas or in other regions.
As can been seen in the Table, comparison of the densities mentioned in
different sources is difficult as the minimum size is not always the same in the

Table 4.1
Densities of Brazil nut trees in “large” inventories (sample area >10 ha). Note that different minimum
diameter limits have been used. Forest area refers to the total forest area of the type(s) inventoried.
DBH = diameter at breast height, measured at 130 cm height.

Region Forest Sample Min Nr of Mean

area area DBH trees density
[ha] [ha] [cm] [ha-1 ]

1 Beni, Bolivia - El Tigre reserve 834 834 50 900 1.1

2 Beni, Bolivia - El Tigre reserve 12 12 10 20 1.7
3 Pará, Brazil - Madeira/Tapajos region 3,700,000 252 25 97 0.38
4 Pará, Brazil - Tapajos/Xingu region 2,500,000 415 25 198 0.48
5 Pará, Brazil - Planalto de Curuá-Una region 60,000 309 45 316 1.0
6 Pará, Brazil - Pinkaití (transects ) c. 950 < 22 10 1.3
7 Pará, Brazil - Pinkaití (in Brazil nut stand ) 28.5 28.5 10 137 4.8
8 Acre, Brazil - Xapurí 51 51 40 98 1.9

Sources. 1: PROMAB (1999: unpublished report, census on Brazil nut trees); 2: Zuidema & Boot (2002);
3-5: FAO inventory cited in Salomao (1991); 6-7: Peres & Baider (1997); 8: Viana et al. (1998).
24 • Chapter 4

studies. In certain cases, even no reference is made to the minimum size at all,
making the comparison of values difficult.
Another cause of differences in the densities in Table 4.1 is that the
selection of study plots has presumably not been carried in the same way in
all studies. In the case of study 2 and 7, the plots have been deliberately placed
within stands of Brazil nut trees. Studies 3-5 are standard forest inventories
for which a large number of 1-ha plots have been established according to a
certain sampling design. Studies 1 and 6 are landscape-wide inventories, that
attempt to provide an estimate for an area of a number of square kilometres.
Lastly, the mean values that are presented in Table 4.1 are based on counts
of trees in a certain number of plots. Such counts typically have a highly
skewed distribution, i.e. with frequent occurrence of low values and infrequent
occurrence of high values. Figure 4.1 gives an example of such a distribution
for the Brazil nut tree. The data do not follow a normal distribution which
can be conveniently described using the average value, but a so-called Poisson
distribution. The most convenient descriptor of the abundance of trees for
such types of datasets is not the mean value, but probably the category that
has the highest abundance, or the median value of the distribution. But the
best way to describe such data is by making a graph of the distribution of the
tree counts as in Figure 4.1. This provides the most reliable information on
densities.
Tree counts - Bolivia Tree counts - Brazil
184 plots Mode Median 51 plots
Mode
Proportion of plots

0,2 Median 0,2 Mean

0,1 0,1

0 0
0 1 2 3 4 5 6 7 8 9 10 11 0 1 2 3 4 5 6 7 8 9 10 11 12 13
Trees per plot Trees per plot

Figure 4.1
Typical distribution of Brazil nut tree counts in plots in Bolivia and Brazil. Shown is the proportion of
study plots in which a certain number of trees is found. It becomes clear that different descriptors of
such datasets provide very different values: the mode (the most frequent category) equals 1 (left) or
4, the median (the category which encompasses the midpoint of all data) equals 3 (left) and 5 (right)
and the ordinary mean 3.2 (left) and 5.5 (right). One should therefore be careful with the use of the
mean values in this type of tree count data. Data are counts of Brazil nut trees >60 cm DBH from 184
plots (0.5 ha) in Northern Bolivia (DHV 1993; left graph) and counts of trees > 1 m height in 51 ha in
Acre, Brazil (Viana et al. 1998; right graph).
 Population distributions • 25

4.2 Spatial distribution

An important part of the variation in densities of Brazil nut trees that is

reported (see Table 4.1) may be explained by the clumped occurrence of the
species. Spatial clumping of Brazil nut trees seems to occur in large part of
the distribution area of the species (Peru: Sánchez 1973, Pará, Brazil: Peres &
Baider 1997). At the landscape-level, Brazil nut trees seem to occur in stands
with high densities interspersed with areas with much lower densities of the
species (Peres & Baider 1997). These stands are referred to as “groves”,
“manchales” (Spanish), “castañales” (Sp). One illustration of the landscape level
distribution of Brazil nut trees is provided by Peres & Baider (1997). In a large
inventory in Pará, Brazil, they found all Brazil nut trees along only 13 of the
total of 56 transect segments which they established. The remaining segments
did not contain any Brazil nut tree.
Brazil nut stands are reported to contain around 50-100 individuals (Mori
& Prance 1990), or 5-100 individuals (Sánchez 1973). However, actual counts
of the number of individuals in stands are scarce. Two stands of over 25 ha in
Pará, Brazil, were intensively inventoried and contained 75 and 149 trees of
>10 cm DBH (Peres & Baider 1997). Within the stands, the position of Brazil
nut trees is not clumped: in a stand of almost 30 ha in Pará, Brazil, the spatial
distribution of trees was random (Peres & Baider 1997), i.e. neither clumped
nor overdispersed.
Within stands, Brazil nut trees are often dominant features in the forest
due to their large adult stature (emergent and large maximum DBH) and their
relatively high density. They often account for a large proportion of the total
basal area in a forest. In a number of 1-ha plots in Acre, Brazil, Brazil nut trees
often account for 10 to even over 25% of the total basal area of trees > 10 cm
DBH (Salomao 1991). In 4 ha of permanent sample plots in Northern Bolivia
(Reserve El Tigre, Beni), Brazil nut trees accounted for 11% of the total basal
area of trees >10 cm DBH (PROMAB, unpublished data).
The finding that Brazil nut trees occur in stands has led to hypotheses on
the origin of these high-density areas. It has been argued that the stands are
the result of Amerindian planting activities (Mori & Prance 1990 and
references therein), i.e. that the current spatial distribution of stands reflects
that of ancient indigenous “plantations”. There are two arguments against this
hypothesis. First, there are large areas which have been inhabited by
Amerindians but where Brazil nut trees do not occur (Mori & Prance 1990).
Furthermore, a large share of tropical tree species have a clumped spatial
distribution at a certain scale and to a certain degree. In a tropical forest in
Panama, clumped spatial distributions were found for 44 out of 66 tree species
(Hubbell 1979).
26 • Chapter 4

An alternative explanation of the clumped distribution uses the system

of seed dispersal of the Brazil nut tree. This explanation is more likely and is
further described in Section 6.3.

4.3 Size distributions

Size distributions (also called: population structures) for the Brazil nut
tree have been established for a number of areas. These structures provide
information on the abundance of Brazil nut trees of different sizes. They may
be an important tool to obtain estimates of the seed production in a certain
area. Using these structures together with knowledge on the seed production
per size category, an estimate of the total seed production can be made. Below,
some population structures that include seedlings and saplings are shown,
followed by a larger number of structures that focus on adult and pre-adult
trees (>30 cm DBH).
Two population structures from the Bolivian Amazon and one from Acre
in Brazil are presented in Figure 4.2. The Brazilian population structure starts
at 1 m height, which makes the comparison somewhat difficult. In all three
structures of Figure 4.2 it is clear that the number of seedlings or saplings is
much higher than that of reproductive and juvenile trees. For the two Bolivian
populations, the number of seedlings (<70 cm height) per reproductive tree
amounted to 8 in the Pando site and 40 in the Beni site. The population
structures have the shape of a reverse J, i.e. a continuously declining abundance
with increasing size, but with somewhat higher densities again for medium-
sized adult trees (see Figure 4.3 and below).
Unfortunately, little information on seedling abundance of the Brazil nut
tree has been published. This may be due to the fact that Brazil nut seedlings
occur at relatively low densities in primary forests. Even within stands of Brazil
nut trees the abundance of plants <1.5 m height is only 25-50 per hectare (see
Figure 4.2).
More information is available on the abundance of larger individuals. In
Figure 4.3, population structures for trees >30 cm DBH are presented from
Brazil and Bolivia. The general pattern that can be observed in these graphs is
that the density of trees of 30-70 cm DBH and >150 cm DBH is typically
low, and that the density is higher at intermediate sizes (100-150 cm DBH).
This pattern is very clear from the large scale Bolivian inventories (middle
graphs), and for the Pando and Acre populations. One of the Pará sites (the
left upper graph) gives a less clear pattern, although the lower abundance of
pre-adult individuals 50-70 cm DBH is also clear in this population structure.
 Population distributions • 27

Pando - Bolivia Beni - Bolivia

20 40
6 ha 12 ha
Tree abundance [ha-1] n=179 n=152
15 30

10 20

5 10

0 0
1 3 5 7 9 11 13 15 1 3 5 7 9 11 13 15
Size category Size category

Xapuri - Brazil
3
51 ha
Tree abundance [ha-1]

n=271
2

0
1 3 5 7 9 11 13 15 17 19
Size category

Figure 4.2
Population structures of Brazil nut trees in Bolivia and Brazil. Black bars refer to size categories with
seedlings, saplings and pole-sized trees (< 10 cm); the arrows point to the first category starting with
60 cm DBH. Data are from Zuidema & Boot (2002) for left and middle graph, and from Viana et al.
(1998) for right graph.

Instead of the peak in the other structures, the Pará population has a longer
tail to the right in its size distribution. Indeed, more very large trees are found
there: around 3% of all measured trees had a DBH >220 cm.
Several interpretations have been given to the occurrence of the “hump”
in the population structure of the Brazil nut tree. It has been argued that the
accumulation of individuals of certain sizes reflects the Amerindian cultivation
of the species (Clay 1997), i.e. the trees in this clump have approximately the
same age as they were planted in the same period of time. Alternatively, this
typical distribution has been attributed to the occurrence of large-scale
disturbances that are necessary for regeneration of the species (Mori & Prance
1990, Peres & Baider 1997). The background of this hypothesis is similar to
the previous: the trees in the clump have approximately the same age, which
is the time since a certain large-scale disturbance. An alternative explanation
of the low and high densities on the basis of information on DBH growth is
provided in Section 9.7.
28 • Chapter 4

Pará - Brazil Pando - Bolivia

15% 15%
Percentage of all trees

28.5 ha
n =179 n =117

10% 10%

5% 5%

0% 0%
Northern Amazon - Bolivia Beni - Boilivia
10% 15%
158 ha
Percentage of all trees

834 ha
n = 1060 n = 900

10%
5%
5%

0% 0%
Pará - Brasil Acre - Brasil
0,60 0,20
9 ha 51 ha
Tree abundance [ha-1]

n = 30
n = 11
0,15
0,40

0,10
0,20
0,05

0,00 0,00
100
120
140
160
180
200
220
40
60
80

100
120
140
160
180
200
220
40
60
80

DBH [cm] DBH [cm]

Figure 4.3
Population structures of adult and pre-adult Brazil nut trees in Bolivia and Brazil. DBH categories are
10 cm wide, i.e. the category denoted by 40 includes all individuals of 30-40 cm DBH. The four upper
graphs show the relative abundance of different-sized trees, the lower two show absolute abundance.
The dots in the middle right graph denote missing data. Sources: Upper left graph: Peres & Baider
(1997) (grove I only); upper right: Zuidema & Boot (2002); middle left: DHV (1993) (based on
elaboration of database owned by ZONISIG - La Paz); middle right: PROMAB (1999); lower left:
Salomao (1991); and lower right: Viana et al. (1998).
 Chapter 5

Reproductive ecology

5.1 Flowering

Flowers of the Brazil nut tree appear at the beginning of the rainy season.
Figure 5.1 shows the distribution of dropped flowers in a Northern Bolivian
forest. The peak of flowering at this site is December and January, whereas this
seems to be earlier (November- December) in Peru (Ortiz 2002).Variation in
flowering period is also reported in Brazil, where the western (Acre)
populations flower before the eastern (Pará) populations (Moritz 1984),
probably related to the distribution of rain over the year.
Flower production in a natural stand of Brazil nut trees has been studied
in detail in a Bolivian primary forest population (Leigue Gómez & Boot in
pressa).The number of flowers per tree was found to be approximately 100,000
(range: 28,000-161,000) for the complete flowering period, based on counts
for six reproductive trees (DBH 72-134 cm).

Flowering and fruiting - Bolivia

80 Flowers
Fruits
60
%

40

20

0
Nov Dec Jan Feb
Month

Figure 5.1
Periods of flowering and fruiting of the Brazil nut tree. Shown is the percentage of the total
production of flowers and fruits that is dropped in each of the months. Data are from 40 reproductive
trees an undisturbed primary forest site in Beni, Bolivia, and were collected in 1994-95. Source: Leigue
Gómez & Boot (in press a).
30 • Chapter 5

Table 5.1
Comparison of the number of seeds per fruit for natural populations of the Brazil nut tree in different
regions. SD = standard deviation.
Sources. 1&3: Zuidema & Boot (2002); 2. Leigue Gómez & Boot (in press a); 4: Peres & Baider (1997);
5: Viana et al. (1998); 6: Sánchez (1973). Sample sizes. 1&3: 150 fruits; 2&5: unknown; 4: 361 fruits;
10 fruits.

Region Average SD Minimum Maximum

1 Beni, Bolivia 18.1 3.5 8 26

2 Beni, Bolivia 17.4 1.9 11 24
3 Pando, Bolivia 19.4 2.9 9 26
4 Pará, Brazil 17.1 7 28
5 Acre, Brazil 18.0 2.6 13 25
6 Madre de Dios, Peru 16.8 2.5 12 21

Flowers of the Brazil nut tree are open only for a few hours: they open
before sunrise and fall early in the afternoon (Moritz 1984, Mori & Prance
1990, Ortiz in press). Flowers are reported to be pollinated by large-bodied
bees, especially Euglossinae bees. Only large insects are able to pollinate the
flowers of the Brazil nut tree, as they are able to lift the “hood” of the flower
to reach the pollen and nectar inside (see Figure 5.1b; Nelson et al. 1985).
Basic information on pollinators of Brazil nut trees in natural forest is still
scarce, in spite of its importance for the establishment of plantations and
enrichment planting. Studies in Brazil have focussed on pollination of new
varieties of the Brazil nut tree in experimental plantations (Moritz 1984).
The relatively low seed production of isolated trees and plantations of the
Brazil nut tree is thought to be related to the limited visitation by certain
pollinators. As these pollinators seem to restricted to (primary) forest
conditions, pollination may be a limiting factor in the production of seeds.
This is especially important as Brazil nut trees rely on cross-fertilisation for
the development of seeds. However, some inbreeding may occur (O’Malley et
al. 1988).
The proportion of flowers that finally yield a fruit is very low: only 0.28%
of the total number of flowers will develop into a fruit (based on flower and
fruit counts of six reproductive trees in a Bolivian primary forest, Leigue
Gómez & Boot in press a). Large part of this loss is probably due to lack of
pollination, although fruit abortion may also be important: aborted fruits are
found regularly on the forest floor during the ripening period of the fruits.
 Reproductive ecology • 31

Table 5.2
Comparison of seed fresh weight and seed size for natural populations of the Brazil nut tree in
different regions. Av = average, SD = standard deviation.

Region Weight: in-shell Weight: shelled Length Width

[g] [g] [cm] [cm]
Av SD Av SD Av SD Av SD

1 Beni, Bolivia 7.3 0.9 3.5 0.63

2 Acre, Brazil 9.8 2.0 4.2 0.5 2.0 0.2
3 Acre, Brazil 4.2 1.8
4 Pará, Brazil 5.8 0.9 4.8 0.6 2.1 0.3
5 Madre de Dios, Peru 9.0 1.3

Sources. 1: Leigue Gómez & Boot (in press a); 2: Kainer et al. (1999a); 3: Viana et al. (1998); 4: Peres
& Baider (1997); 5: Sánchez (1973). Sample sizes. 1&3: unknown; 2: 600 seeds; 4: 50-100 seeds; 5: 168
seeds.

5.2 Fruiting

Brazil nut trees produce fruits each year, although some individual trees
may not produce in a given year. The development of seeds takes very long,
in many cases more than 12 months (Moritz 1984,Ortiz 2002). This implies
that flowers and fruits are often present at the same time. The distribution of
fruit fall in a Bolivian site (Figure 5.1) shows a main peak in fruiting in
November, but this peak often occurs later, in December - February (Mori &
Prance 1990, Zuidema & Boot 2002, Ortiz 2002).
The number of seeds per fruit varies roughly between 10 and 25.Values
of different primary forest populations are included in Table 5.1. For trees in
a Brazilian experimental plantation, the number of seeds per fruit are similar
to those in the Table: average number varies from 17.5 to 18.8 for four different
varieties (Moritz 1984).
As for seed number, the size and weight of seeds are also rather variable.
For instance, Brazil nut trees from the region of Abufari Amazonas are known
to produce much larger fruits and seeds than elsewhere. Differences in seed
size and form are also present at smaller spatial scales (Kainer et al 1999a).
Fresh weight and dimensions of seeds from different areas are summarised in
Table 5.2. The data from Acre (2) include information from 10 different trees
which were located at some distance of each others. Seed sizes were strongly
different among these trees: seed weight ranged from 7.0-12.3 g (means of 60
seeds each; Kainer et al 1999a).
32 • Chapter 5

Reproduction vs. DBH - Data Reproduction vs. DBH - Model

yes 100%
90%

Reproductive trees
Reproductive?

50%
50%

10%
no 0%
0 50 100 150 200 0 20 40 60 80 100
DBH [cm] DBH [cm]
Figure 5.2
Relation between DBH (diameter at breast height) and reproductivity for the Brazil nut tree. The left
graph shows the original data of 364 trees in two populations in Beni and Pando departments, Bolivia.
The right graph shows a logistic regression model that describes the size-dependent pattern in
reproduction: the drawn line indicates the average percentage of reproductive trees for any value of
DBH. This model had a very good fit and explained c. 93% of the variation in the data. The stippled
lines indicate the tree size at which 10% (at DBH=25 cm) and 90% (at DBH=60) of the trees is
reproductive; the broken line indicates the tree size at which 50% of the trees is reproductive (at
DBH=43 cm). Source: Zuidema & Boot (2002).

5.3 Attaining reproductive size

To obtain estimates on the (potential) productivity of Brazil nut trees, it

is important to know the minimum reproductive size. Information on the
relation between tree size and reproduction has been collected in two primary
forest sites in Bolivia. Figure 5.2 shows the relation between tree diameter
(DBH) and reproductive status of Brazil nut trees.There is a strong correlation
of DBH and reproductive status, as can be seen by the clear distinction between
non-reproductive and small trees and reproductive and large trees. As much as
98% of the individuals of >40 cm DBH are reproductive, whereas only 3.5%
of the trees of 1-40 cm DBH is reproductive. The typical relation of DBH and
reproductive status is shown in the right graph of Figure 5.2, and was not
different for the two Bolivian forest sites. According to this model, over 90%
of the Brazil nut trees of 60 cm DBH are reproductive. Brazil nut trees of this
size are roughly 140 yr old (see Section 8.5 on Tree ages).
Studies in Northern Bolivia also revealed that a certain proportion of
trees that are in the reproductive size categories do not bear any fruits in a
given year. A large-scale census in over 800 ha of mainly primary forests in
Beni showed that 16% of a total of 900 trees (ranging in diameter from 50-
230 cm) was not reproductive (PROMAB 1999). In another study, it was found
that c. 8% of 275 trees did not produce any fruit during one or two out of
three years (Zuidema & Boot 2002).
Reproduction in the Brazil nut tree is also strongly related to the
availability of light. Trees with crowns that overtop the canopy of a forest have
 Reproductive ecology • 33

Reproduction vs . light
100%

Reproductive trees
50%

0%
Below In canopy Emergent
canopy
Crown position

Figure 5.3
Relation between crown illumination and reproductive status for the Brazil nut tree. The position of
the tree crown related to the forest canopy is crucial for the reproductive status of the tree. The data
are from 364 trees in two populations in Beni and Pando departments, Bolivia. Source: Zuidema &
Boot (2002).

a very high probability of being reproductive. Figure 5.3 shows that 97% of
those trees are reproductive, whereas this percentage is much lower for trees
of which the crown is in the canopy. Brazil nut trees never seem to produce
fruits when their crown is below the existing forest canopy.
In plantations or disturbed areas with much lower (forest) canopy,
reproductive size may be attained at much smaller tree size than in primary
forest. The smallest reproductive tree in the primary forest studies in Bolivia
measured 33.3 cm DBH. In disturbed areas along roads and in an experimental
garden in Northern Bolivia, Brazil nut trees of much smaller diameter (20-25
cm DBH) have been observed with fruits. First reproduction of grafted Brazil
nut trees in experimental gardens in Brazil have been reported to first produce
fruits 6 years after grafting, certainly at a diameter much lower than 20 cm
(Müller 1981).

5.4 Seed production

An adequate estimate of the (potential) productivity of Brazil nut trees

in a certain forest area requires information on the nut production per tree.
Several studies have shown that fruit production of Brazil nut trees is highly
variable among individual trees. The fruit production of a tree may range from
a few fruits to 1032 (Zuidema & Boot 2002), 1126 (Leigue Gómez & Boot
in press a), or even over 2000 (Ortiz 2002). Figure 5.4 shows the variation in
fruit production among trees for two Bolivian sites. It becomes clear that most
trees only produce a small amount of fruits: the largest share of adult trees
produce less than 100 fruits. Nevertheless, a substantial part (around 25%) of
the adult trees produce 100-200 fruits. Less than 10% of all reproductive Brazil
34 • Chapter 5

Variation in fruit production - Beni Variation in fruit production - Pando

60% 1996
1994
1997
Percentage of trees

1995
1996 1998
40%
1997
1998
20%

0%
0
1-100

>900
101-200
201-300
301-400
401-500
501-600
601-700
701-800
801-900

1-100

>900
101-200

201-300

301-400

401-500

501-600

601-700

701-800

801-900
Fruit production Fruit production

Figure 5.4
Variation in fruit production among adult trees in two forest sites in the Bolivian Amazon. Shown is
the percentage of trees that produces a certain number of fruits. Data of different years are presented
with different bar types. The data are from 40 trees in Beni, Bolivia, and 48-58 trees in Pando, Bolivia.
The Pando data are only those fruits that have been opened by Brazil nut collectors, and thus slightly
underestimate the total fruit production. Sources: Leigue Gómez & Boot (in press a) for Beni and
Zuidema & Boot (2002) for Pando.

nut trees produce over 500 fruits. The high maximum fruit production figures
that are often mentioned, should thus be regarded as highly exceptional.
In Table 5.3 the average production of Brazil nuts in three Bolivian and
Brazilian forest sites is shown. The average number of fruits is around 140-
180, but the median number (a more suitable parameter when part of the
values are very high) is substantially lower: 100-140. For seed weight figures
there are no median values available, but the median would probably be around
17-18 kg per tree. That fruit production is highly variable among adult trees
can be seen by the very high standard deviation values of fruit production and
seed weight.
Considering the large variation in production of Brazil nuts, an important
question is raised: Which factors determine the number of fruits produced by

Table 5.3
Fruit production of three natural populations of the Brazil nut tree in different regions. The values
for the two Bolivian sites are averages over five and three years for the Beni and Pando site,
respectively. Av = average, SD = standard deviation; Min = Minimum; Max = Maximum.

Region Fruit number per tree Seed weight per tree [kg]
Av SD Min Max Median Av SD Min Max

1 Beni, Bolivia 184 122 0 1126 141 23.3 21.2 0.0 90.7
2 Pando, Bolivia 139 104 0 1032 103
3 Acre, Brazil 24.0 27.5 1.5 105.0
4 Madre de Dios, Peru 100 20

Sources. 1: Leigue Gómez & Boot (in press a); 2: Zuidema & Boot (2002); 3: Viana et al. (1998); 4:
Ortiz (2002). Sample sizes. 1: 40 trees; 2: 53 trees; 3: 144 trees; 4: estimation.
 Reproductive ecology • 35

a Brazil nut tree? Several factors have been put forward (by Viana et al. 1998,
Leigue Gómez & Boot in press a,b): (1) tree size, (2) crown position of the
tree, (3) liana infestation, (4) inherent temporal variation, (5) climatic factors,
(6) genetic factors, (7) soil conditions, (8) interactions with pollinators, and
(9) interactions with fruit predators (when the fruit is still in the tree). Below
the importance of several of these factors is discussed.
1 • Tree size. In two Bolivian sites, fruit production has been related to the
diameter of adult trees and to their crown area (i.e. the area of the crown when
projected on the forest floor). Fruit counts can be obtained by: (a) counting
the number of opened fruits shortly after the end of the harvesting period;
(b) systematically searching for fruits before Brazil nut collectors enter the
forest; or (c) by counting fruits with binoculars in the crown of the tree during
periods that the tree is changing leaves. The latter method can be during
periods that Brazil nut trees shed large part of their leaves during the dry
season.
The relation of DBH and fruit production is shown in Figure 5.5. It
becomes clear that there is a lot of scatter around the regression line that shows
the average relation between DBH and fruit production. Only a small
proportion of the total variation in fruit production is explained by the DBH
of the tree: for the Beni site this is 25%, for the Pando site only 9%.The higher
percentage of explained variance for the Beni site is probably due to the fact
that 5 years of fruit production data have been averaged, whereas in Pando

Reproduction and DBH - Beni Reproduction and DBH - Pando

800 500
Fruit production per tree

400
600
300
400
200
200
100

0 0
40 80 120 160 40 80 120 160 200

DBH [cm] DBH [cm]

Figure 5.5
Relation between fruit production and diameter at breast height (DBH) for Brazil nut trees in two
Bolivian forest sites. Shown is the fruit production for adult trees of different sizes. Fruit production
was averaged over 5 years for Beni and 3 years for Pando. The straight line is the linear regression
of fruit production against DBH. The curved stippled line is the results of linear regression on
logarithmically transformed data which adjust for the skewed distribution of fruit production. The
data are from 40 trees in Beni, Bolivia, and 52 trees in Pando, Bolivia. The Pando data are only those
fruits that have been opened by Brazil nut collectors, and thus slightly underestimate the total fruit
production. Sources: Leigue Gómez & Boot (in press a) for Beni and Zuidema & Boot (2002) for Pando.
36 • Chapter 5

Reproduction and crown area

800

Fruit production per tree

600

400

200

0
0 200 400 600 800

Crown area [m2]

Figure 5.6
Relation between fruit production and crown area for 40 Brazil nut trees in a Bolivian forest site.
Shown is the fruit production for adult trees with different crown areas. Fruit production was
averaged over 5 years. Crown areas where determined as vertical projection of the crown on the
forest floor. The straight line is the linear regression of fruit production against crown area. Source:
Leigue Gómez & Boot (in press a).

this was only 3 years. For fruit counts of one year only, it is often difficult to
find a relation with DBH.
Based on the straight lines in Figure 5.5, one may expect the average fruit
production to increase by 8-25 fruits when DBH increases with 10 cm (8 fruits
for the Pando data, and 25 fruits for the Beni data). Using the curved regression
lines, these figures are somewhat different. For the Beni graph, the increase is
4 fruits per 10 cm DBH increase for small trees, but much larger for large trees
(> 120 cm DBH) which may gain over 40 fruits by having a 10 cm bigger
diameter. For the Pando data, the increase is lower: large trees gain only 10
fruits when they grow 10 cm in DBH.
The relation between fruit production and crown area has been
investigated at one Bolivian site. In Figure 5.6, this relation is shown for the
same 40 trees as in the left graph of Figure 5.5. Also here, the straight line
shows the average relation between fruit production and crown area. Almost
50% of the variation in fruit production can be explained by the value of
crown area. When comparing Figures 5.5 and 5.6, it becomes clear that the
crown area is a much better predictor of fruit production than DBH.
Unfortunately, determining crown area is much more labour intensive than
measuring DBH. Information on crown area of Brazil nut trees is therefore
very scarce.
2 • Crown position.The position of the crown in relation to the forest canopy
may also co-determine the production of fruits of the Brazil nut tree. However,
the influence of this parameter may be small as, most reproductive Brazil nut
trees are emergent, i.e. their crowns are positioned above the existing forest
canopy. In a study in Northern Bolivia, the influence of crown position on
 Reproductive ecology • 37

Lianas and fruit production

200

More than
expected
100
Difference in fruit
production 0
-100
Less than
expected
-200
-300
-400
0 300 600 900 1200
2
Liana basal area [cm ]

Figure 5.7
The impact of liana infestation on the fruit production of the Brazil nut tree. Shown is the difference
between the expected number of fruits that a tree of a given DBH produces and its actual production.
A positive value indicates that actual fruit production is higher than expected on the basis of the
relation in the left graph of Figure 3.5; a negative value indicates less fruits than expected. The line
shows that with more liana infestation, fruit production becomes smaller than expected. This indicates
a negative effect of lianas on fruit production. Regression: Difference = -75 × Ln(liana basal area) +
374 (R 2 = 0.21). Source: Leigue Gómez & Boot (in press b), based on a study on 18 Brazil nut trees
with liana infestation.

fruit production was investigated for 100 trees (Zuidema & Boot 2002). First,
fruit production was related to DBH, then to crown position. DBH had an
effect on fruit production, but no additional effect of crown position was
found.
3 • Liana infestation. Lianas may influence the fruit production in Brazil nut
trees in case they overtop a significant part of the tree crown. In that way, they
reduce the amount of light intercepted by leaves in the tree crown and reduce
the amount of carbohydrates produced by photosynthesis. As a consequence,
the production of fruits may also decrease.
In a population of 40 adult Brazil nut trees in Bolivia, 18 were infested
by lianas, with a varying number of size of the lianas (Leigue Gómez & Boot
in press b). An inventory in the same area, but at a larger scale revealed that
45% of 900 Brazil nut trees of >50 cm DBH were infested by lianas
(PROMAB 1999). The basal area of the lianas in the 18 Brazil nut trees of the
first study (both those that climbed on the Brazil nut tree itself and those that
climbed on neighbouring trees but which overtopped the crown of the Brazil
nut tree) ranged from 60 to 1200 cm2, with an average of 350 cm2. The latter
figure is equivalent to a the basal area of a tree of 21 cm DBH, which is quite
substantial. Detailed observations on the fruit production of these 18 infested
trees over a period of 5 years, showed a negative relation between fruit
production and the basal area of lianas in Brazil nut trees (Figure 5.7). This
was determined by calculating the difference between the expected fruit
production of a given tree on the basis of its DBH (see Figure 5.5) and the
38 • Chapter 5

Variation in fruit production

1500

Fruit production per tree

1200

900

600

300

0
1994 1995 1996 1997 1998 1999 2000
Year

Figure 5.8
Variation in fruit production among years for individual Brazil nut trees in a natural population in
Northern Bolivia. Symbols that are connected are fruit production figure from the same tree in
different years. “Year” refers to the year in which fruiting started, e.g. 1997 refers to the production
in the fruiting period late 1997 - early 1998. Fruit production was measured by counting Brazil nut
fruits on the forest floor for 7 years and for 40 trees. Source: Leigue Gómez & Boot (in press a).

actual observed fruit production of that tree. It then appeared that the observed
fruit production was lower for trees with a high degree of liana infestation.
4 • Inherent temporal variation. There are numerous observations on the
variation in fruit production of the Brazil nut tree over time. Brazil nut
collectors frequently mention the fact that trees that produce abundantly
during one year, hardly produce any fruits during the following year. Also there
is evidence for this variation from detailed fruit counts (Leigue Gómez & Boot
in press a, Zuidema & Boot 2002, Ortiz 2002). An extreme example of the
high temporal variability in fruit production is the a tree of 165 cm DBH that
produced 31 fruits in 1997 followed by 1003 fruits in the next year (Zuidema
& Boot 2002). Such differences among years are certainly not exceptional.
In a 7-yr study on the fruit production in a Bolivian population of the
species, large year-to-year variation in the number of fruits produced was
found. Figure 5.8 shows the variation in fruit production of 40 Brazil nut trees.
It becomes clear that a year with a high production is often preceded and
followed by years with substantially lower production.
The question is: what causes the large temporal variation in fruit
production in the Brazil nut tree? Considering the large variation in fruit
production in time the question arises as to what are the causes of this
variation. First of all, strong temporal variation in fruit production occurs very
frequently in tropical tree species (Janzen 1978). In certain tree species,
temporal variation is extreme in the sense that during most years there is no
reproduction at all, and at certain years, mast fruiting occurs. Also, many
tropical tree species have a regular cycle of 2 or 3 years in which fruiting
occurs once. And even in the large group of tree species that produce fruits
each year, high levels of year-to-year variation in fruit production are found.
 Reproductive ecology • 39

Average fruit production of trees

500

Fruit production
400
300
200
100
0
Fruit production of population
12000
10000
Fruit production

8000
6000
4000
2000
0
1994 1995 1996 1997 1998 1999 2000
Year

Figure 5.9
Variation in fruit production among years for a population of 40 Brazil nut trees in Northern Bolivia.
The upper graph show the median production (the mid-value of the dataset, or 50% percentile), the
lower point of the vertical line is the 25% percentile and the upper point of the vertical line is the
75% percentile. The horizontal line is the average production over 7 years. The lower graph shows
the total production over 7 years. The horizontal line in this graph also denotes the average
production. Source: Leigue Gómez & Boot (in press a).

The temporal variation in fruiting in the Brazil nut tree is therefore not
exceptional when compared to other tropical tree species.
Two types of explanations may be given for the temporal variation in
fruiting in the Brazil nut tree. The first is a matter of space limitation on
branches of the tree. As fruit development in this species often takes over a
year, flowers for the next year’s fruits are present when the ripening fruits of
this year are still on the tree. As flowers and fruits are present on the final part
of small branches, they may have to “compete” for space. A high density of
fruits on the branches in a high-production year would then leave less space
for flowers in the next year. However, this reasoning is very speculative, as no
information on detailed observations of flowering and fruiting on branches of
natural Brazil nut trees are available.
The second explanation of the fruiting variability in time has an
evolutionary background. It has been suggested that producing large amounts
of seeds or fruits with a lower frequency than once per year, may be more
successful for the regeneration of the species.The reason for this is that animals
that feed on the seeds (in this way killing them) consume a smaller proportion
of the total seed production when the moment of fruiting is unpredictable
and when very large amounts of fruits are produced in a short period of time.
40 • Chapter 5

The strong variation in fruit production in the Brazil nut tree may thus be an
adaptation to support the regeneration of the species.
At the population level, the variation in fruit production in time is much
less than that at the individual level. The Beni study showed that the total
production of 40 Brazil nut trees over 7 years ranged from 4792-10749 fruits,
with a coefficient of variation of 22. Figure 5.9 shows the total production
over the five observation years, as well as the median production in the
population. Thus, large individual variation over years and among individuals
tends to average out over a population of trees. The finding that the degree of
variation in fruiting at the population-level is less than that among individual
trees is important: estimates on fruit production over larger areas should
therefore be based on fruit counts of a substantial number of trees, not just a
few.
5 • Climatic factors. As stated above, the reasons behind the large year-to-
year variation in fruit production of individual Brazil nut trees remain unclear.
Nevertheless, in spite of the large and unpredictable variation in fruiting of
individual trees, it seems that at the level of the population, climatic factors
may play a role in determining total fruit production. Both Brazil nut collectors
and the officials of the processing plants speak about “good” and “bad”
production years. Substantial variation in fruit production at the population
level also has become clear from the 7-yr fruit production study on 40 trees
in the Bolivian Amazon (Leigue Gómez & Boot in press a). The population-
level fruit production in Figure 5.9 shows a significant drop in production
during 1998. This year coincided with an El Niño year with a greatly reduced
precipitation.Thus, climatic conditions are likely to influence fruit production.
In order to better understand annual fluctuations in fruit production of
individual trees and populations of Brazil nut trees, it is necessary that more
reproduction studies are carried out, preferably over long periods of time.
Little information is available on the other factors that may influence the
fruit number of Brazil nut trees. Genetic factors (6) will certainly play a role,
but it is unclear how large this is. Similarly, the influence of soil conditions
(7) on fruit production has not been quantified, although it may have an effect
on fruiting. Interactions with pollinators (8) certainly influence fruiting of the
Brazil nut tree. An indication for this is that isolated trees in agricultural fields
are often found to produce less fruits than equally large trees in the forest.
Nevertheless, other factors may also explain this difference (e.g. damage by
burning of agricultural fields; wind damage of branches).
Fruit predators (9) are able to damage the developing fruits in the tree.
In the Peruvian Amazon, it was found that close to 5% of the total fruit crop
was damaged by birds, macaws (Ortiz 2002). Macaws were observed to visit
up to 75% of the trees in one Brazil nut tree population.
 Chapter 6

Dispersal ecology

6.1 Dispersal system

The Brazil nut tree has a complex dispersal ecology that probably largely
depends on the scatterhoarding activity of agoutis. A schematic chart of the
different phases in the dispersal system of the Brazil nut tree is provided in
Figure 6.1.
The first phase of the dispersal system consists of the primary dispersal:
fruits drop to the forest floor by gravity. As the fruits have an extremely hard
and thick woody wall, they are not damaged by the impact of the fall and
remain closed. Most fallen fruits can be found at short distances from the trunk
of the adult tree, almost always below the crown of the trees. The left graph
in Figure 6.2 shows the distances of closed fruits to adult trees from a
population of the Brazil nut tree in Northern Bolivia.

†
Predation
by animals

Primary Secondary "Tertiary"

Seeds Seeds in dispersal Seeds dispersal Seeds
dispersal
in fruit, fruit, on burried in burried in
in tree by gravity forest floor by agoutis caches by agoutis caches

Germination
Germination in soil
in fruit

Seedlings

Figure 6.1
Schematic overview of dispersal system of the Brazil nut tree. Circles are stages within the process,
arrows are the processes. Further explanation in the text.
42 • Chapter 6

Fruits of the Brazil nut tree do not possess an opening that is sufficiently
large to let the large seeds escape the fruit. Seeds can thus only be released by
active opening of the fruit. This is mainly done by the agoutis (Dasyprocta
spp.). Agoutis are large cavylike rodents that feed on fruits and nuts. They are
reported to open over 80% of all Brazil nut fruits that are opened by animals
(Ortiz 2002). Two other rodents, the paca (Agouti paca) and a squirrel may
also open the fruits. In addition, monkeys are occasionally observed to open
Brazil nut fruits when these are still in the tree.
Agoutis gnaw open the thick and tough woody pericarp of the fruit.
Agoutis sometimes move the fruit before opening it. This behaviour explains
the larger distances from opened Brazil nut fruits to the nearest adult tree in
the right graph of Figure 6.2. It can be seen in this graph that almost 50% of
the fruits opened by agoutis is found beyond the crown area of Brazil nut
trees, that is: at distances over 20 m from the tree trunk. For comparison, only
2% of the fruits not opened by agoutis were found at this distance (see left
graph in Figure 6.2). There are examples of fruits being carried for over 300m
by agoutis (Ortiz 1995).
Agoutis eat part of the seeds after opening the fruit. This kills the seeds,
thus making them unavailable for the rejuvenation of the tree. Those seeds
that are not eaten by agoutis, are carried away from the location where the
fruit is opened and buried at a depth of a few centimetres. This is the second
phase in the dispersal system of the Brazil nut tree: the secondary dispersal
(see Figure 6.1). The process of moving and burying seeds is called
scatterhoarding and is also commonly practised by other rodents.
Scatterhoarded seeds are stored to serve as food sources in periods of food
scarcity, e.g. during the dry season when little fruits are available. Usually, each
seed is scatterhoarded individually at one spot, at a short distance from the
location where the fruit was opened.
The seeds buried by agoutis may be recovered by these or other animals
to serve as food. In some cases, these seeds may be reburied at another location.
This would then be the next phase in the dispersal system: the “tertiary”
dispersal. It is not clear how frequently this reburying occurs in reality.
Alternatively, the stored seeds may be predated upon by other species, especially
rodents. This leads to the death of the seed, which then cannot contribute
anymore to the regeneration of the tree.
The stored seeds that are not found and consumed within a certain period
of time may germinate and become seedlings. Presumably, the vast majority
of seedlings stem from seeds that were buried by agoutis. However, on rare
occasions seeds may also be found germinating within an intact pod on the
forest floor (Peres & Baider 1997, P.A. Zuidema, personal obser vation). Also
clumps of seedlings may be found surrounded by the remainders of a pod (P.A.
 Dispersal ecology • 43

Closed fruits Opened fruits

60% 60%
Percentage of all fruits

40% 40%

20% 20%

0% 0%
5

5
15

25

35

45

55

65

15

25

35

45

55

65
Distance to nearest adult tree [m] Distance to nearest adult tree [m]

Figure 6.2
Distance at which fruits of the Brazil nut tree are found relative to the position of adult trees in a
study site in Northern Bolivia. The distance values are the upper limits of distance categories, e.g. 15
includes all distances of 11-15 m. The left graph shows the distances for 263 unopened fruits, the
right graph that of 51 fruits opened by agoutis (with tooth marks of agoutis). Source: Zuidema &
Boot (2002).

Zuidema, personal observation). These seedlings probably stem from seeds that
have germinated after the woody wall of the fruit has completely disintegrated.
From the above description of the dispersal system of the Brazil nut tree
it becomes clear that seed germination under natural conditions depends
heavily on the presence and the activity of agoutis. These animals are crucial
for the natural regeneration of the Brazil nut tree.

6.2 Seed handling by agoutis

An important question concerning the dispersal system of the Brazil nut

is: what is the proportion of seeds that is available for germination? In other
words: which part of the total amount of seeds that is available to agoutis, may
finally become a seedling (see Figure 6.1). This question is important in order
to understand the natural regeneration process of the Brazil nut tree, but even
more, it is important to foresee the consequences of seed collection on the
regeneration of the tree. Although several studies have dealt with the dispersal
system of the Brazil nut tree, there is still not sufficient information to answer
this question.
There are two important methodological constraints that hamper dispersal
studies. First, the fate of seeds needs to be monitored in order to know whether
they are eaten, buried or left without handling. Although marking is possible
with vinyl tread and flags, it is an extremely laborious task. Furthermore, it
should be followed by intensive searching for the removed seeds in a large
area, which is very time-consuming as well.
44 • Chapter 6

Fruit opening - Beni Fruit opening - Pando

100%
Percentage of all
80%

60%
fruits

40%

20%

0%
0 1 2 0 1 2
Time since fruit fall [yr] Time since fruit fall [yr]

Figure 6.3
Percentages of Brazil nut fruits opened by agoutis in relation to the time since fruit fall. Shown are
the results for two primary forest sites in Northern Bolivia. The proportion of opened fruits are
significantly different among years. Source: Zuidema & Boot (2002).

Second, in order to mark the seeds, they have to be removed from the
fruit: marking seeds that are still in the fruit is simply impossible. As a
consequence, in seed dispersal experiments, seeds have been put barely on the
forest floor, without the fruit. This makes the seed handling much easier, as
agoutis do not have to open the fruit before obtaining the seeds. In this way,
the rate at which seeds are eaten, buried and removed is probably higher than
under natural conditions. An important and unanswered question is whether
this method changes seed handling when compared to the natural situation.
In spite of the methodological constraints on dispersal studies, the main
results of these studies are briefly stated below. First, in all studies it was found
that groups of Brazil nut seeds placed on the forest floor are rapidly taken
away (either consumed or dispersed) once they are encountered by agoutis or
other animals. In a Brazilian study, over 85% of the seeds were either eaten or
dispersed in 14 days after the start of the experiment (Peres & Baider 1997).
In a Bolivian experiment using higher density of seeds, 60-80% of seeds was
removed after three weeks (Bouwman & van Dijk 1999). Similarly high rates
of removal were found in secondary forests of different ages in Bolivia, where
80% of the seeds were removed after two weeks (Peña-Claros & de Boo 2002).
In contrast to rapid removal of seeds in seed dispersal experiment, are
observations on fruit handling under natural conditions in two Bolivian forest
sites. Figure 6.3 shows the proportion of Brazil nut fruits that is opened by
agoutis, in relation to the time since fruit fall. It appears that a large part of
the fruits is not opened during the first year after fruit fall. The reason for the
late opening of Brazil nut fruits may be that in older fruits the woody wall of
the fruit has softened somewhat, thus making fruit opening easier and more
efficient for agoutis. However, this idea is still hypothetical, and should be
further studied.
 Dispersal ecology • 45

Seed handling - Brazil Seed handling - Bolivia

Left Eaten Left Eaten

Not
buried
Not
buried

Buried Buried

Figure 6.4
Seed handling of Brazil nuts by agoutis in Bolivia and Brazil. Shown are the proportions of seeds that
are eaten, dispersed (and either buried or not buried) or left by agoutis. The data are derived from
seed dispersal experiments in which (groups of) marked Brazil nut seeds were put around Brazil nut
trees). Only those marked seeds that were found back are included. Number of seeds: 424 for the
Brazil study and 59 for the Bolivia study. Sources: Peres & Baider 1997 for left graph, and Bouwman
& van Dijk (1999) for right graph.

Second, the part of Brazil nuts that is consumed and the part that is
scatterhoarded is variable. The results of two dispersal studies in Brazil and
Bolivia are shown in Figure 6.4. In spite of the variation, it becomes clear that
a substantial part of the total number of seeds is scatterhoarded, and may thus
finally become available for germination. In both studies this is around 50%
of the seeds. Thus, in general less Brazil nuts are eaten than scatterhoarded. It
should be kept in mind, though, that conclusions from these results should be
drawn with care, as a substantial part of the marked seeds in all experiments
was not recovered after they were removed (Brazilian study: 40%; Bolivian
study: 57%).
In the Bolivian study, Brazil nuts were offered at two densities (only the
results of the low-density treatment are shown in Figure 6.4), with a tenfold
difference in the number of seeds per unit area. It was found that for each seed
that is consumed, more seeds were scatterhoarded in the high-density treatment
compared to the low-density treatment. This suggests that there is some kind
of satiation of the agoutis: when the density of seeds is very high, agoutis tend
to scatterhoard a larger part of all seeds, as they are probably satiated after
eating a small fraction of the larger number of seeds available.

6.3 Dispersal distances

In order to understand the clumped spatial distribution of the Brazil nut

tree (see Section 6.2), it is important to have information on the distances at
which seeds of the species are dispersed by agoutis. Such distances have been
measured in two studies in natural stands of the Brazil nut tree. In these
46 • Chapter 6

Percentage of all seeds Seed dispersal - Bolivia Seed dispersal - Bolivia

60% high low
density density

40%

20%

0%
1
2
3
4
5
6
7
8
>8

>8
Dispersal distance [m] Dispersal distance [m]

Seed dispersal - Brasil

Percentage of all seeds

60%

40%

20%

0%
0

10

15

20

25

30

Dispersal distance [m]

Figure 6.5
Distance at which seeds of the Brazil nut tree are dispersed by agoutis. These graphs show the results
of three seed dispersal studies in natural forest. Dispersal distance is the distance at which marked
seeds were found back, measured from the location where seeds were put at the start of the
experiment. The distance values are the upper limits of distance categories, e.g. 15 includes all
distances of 11-15 m. The upper left graph is based on the results of 217 marked seeds which were
placed at a low density (45 seeds around each of 10 Brazil nut trees) in a forest where no Brazil nut
collection takes place. The upper right and lower left graphs are based on 283 and 323 marked seeds,
placed at two densities (340 and 3400 seeds around each of 17 Brazil nut trees) in a forest where
Brazil nuts are collected. Due to the differences in study design, only the general patterns can be
compared. Sources: Peres & Baider 1997 for upper left graph; Bouwman & van Dijk (1999) for upper
left and lower right graph.

experiments, a large number of seeds has been put on the forest floor close to
adult Brazil nut trees. The seeds were marked with a vinyl thread with a flag
at the end of it.
The dispersal distances observed in three experiments in Bolivia and
Brazil are shown in Figure 6.5. From the graphs it becomes clear that the vast
majority of seeds is dispersed at very short distances. However, a small
proportion of seeds does reach locations far away from the parent tree. The
maximum dispersal distance found for the three studies shown in Figure 6.5
was 25 m for the Brazilian study and 20.2 m for the Bolivian studies. Intensive
searches in larger areas than those searched in these studies would most
probably reveal dispersal over longer distances. Also, the reburying of
scatterhoarded seeds at other locations may result in further dispersal of seed
from parent trees. Dispersal distances up to 50 m have also been mentioned.
 Dispersal ecology • 47

The dispersal distances found for the Brazil nut tree may help to explain
the clumped spatial distribution of the species at the level of the landscape
(see Section 4.2). It has been suggested that the short dispersal distances of
seeds lead to a clumped distribution of trees (Peres & Baider 1997). The very
few seeds that reach sites at long distances from a Brazil nut tree would then
be the possible sources for the development of new stands of Brazil nut tree.
Long-distance dispersal may be due to reburying of scatterhoarded seeds, or
by other animals. Birds, and especially macaws, are thought to play a role in
this long-distance dispersal: they are reported to open Brazil nut fruits and
may carry the seeds over large distances. Although this explanation remains to
be proven, it is consistent with the clumped distribution of adult trees, the
random distribution of trees within Brazil nut tree stands and the generally
short dispersal distances. It is certainly a good alternative to the hypothesis
that the clumped distribution of Brazil nut trees can be attributed largely to
the activity of indigenous peoples in the Amazon region. Nevertheless, the
relatively high abundance of small Brazil nut trees in disturbed areas (secondary
forests) suggests that human agricultural activity may influence the abundance
and distribution of the species to some extent, but probably this influence is
at a relatively small spatial scale.

6.4 Germination under natural conditions

Not much is known about the germination of Brazil nut seeds under
natural conditions. Germination has been reported to occur after a period of
12-18 months (Müller 1981), based on one study in Brazil. However, more
extensive germination experiments under natural conditions are needed to
obtain additional estimates of the germination period and success, and on the
biophysical conditions that influence germination.
Although the vast majority of Brazil nut seeds probably germinate after
being scatterhoarded by agoutis, a small fraction of seedlings found in natural
populations appear to have germinated in the fruit itself. This may occur in
two ways. First, seeds may germinate after the woody fruit wall has completely
rotten (Sanchez 1973, Peres & Baider 1997, Zuidema & Boot 2002). The
complete rot of the fruit takes around three years. Secondly, seeds may
germinate when still in the intact pod, and seedlings may grow through the
small hole in the fruit wall. A very early experiment reported in 1901 (Watson
1901) already showed that Brazil nuts are able to germinate within the fruit.
In a demographic study in Bolivia, the number of new seedlings was
determined per fruit opened by agoutis (Zuidema & Boot 2002). In two
primary forest sites where Brazil nuts are collected, 0.8-3 new seedlings were
48 • Chapter 6

found per fruit. Taking the average number of 18 seeds per fruit, this implies
that one of each of 6-23 seeds in fruits opened by agoutis will germinate and
produce a seedling. More estimates of this ratio in populations of the Brazil
nut tree are needed to better understand the natural germination and to better
quantify the impact of exploitation on the regeneration.
 Chapter 7

Seedling ecology

7.1 Density and natural conditions of seedlings

The information on densities and conditions of naturally occurring

seedlings of the Brazil nut tree is scarce. Studies in Acre, Brazil and in Northern
Bolivia provide information on the density of different-sized seedlings. Figure
7.1 shows the abundance of seedlings of the Brazil nut tree in two Bolivian
forests.The most abundant size category is that containing seedling of less than
35 cm height, which occur at a density of 20-40 seedlings per ha. Summing
the data of all seedlings smaller than 140 cm height, yields a density of 25-50
seedlings per ha. Another study in one of these sites revealed a similar density
of 53 seedlings per ha for plants <130 cm height (Myers et al. 2000). A study
in Acre, Brazil that included large seedlings (or saplings) of over 1 m in height
reported a seedling abundance of 2.5 individuals per ha (Viana et al. 1998).
This abundance is similar to that of the largest category in Figure 7.1. Clearly,
the number of seedlings found in one hectare strongly depends on the
abundance of adult trees in that area.
One of the reasons that little information is available on the abundance
of seedlings, is the time-consuming activity of searching large areas in order

Seedling abundance - Bolivia Figure 7.1

Abundance of seedlings of the Brazil nut tree in
40 Beni, Bolivia
Seedling abundance [ha-1]

two primary forest sites in Northern Bolivian

Pando, Bolivia
where Brazil nuts are collected. Open bars refer
30
to densities in Beni, black bars to those in
20 Pando. These data area based on searches in 4.5
(Beni) and 1.4 ha (Pando). A total of 97 (Beni)
10 and 56 (Pando) seedlings were found in these
areas. Source: Zuidema & Boot (2002).
0
10-35 35-70 70-140 140-230
Seedling height [cm]
50 • Chapter 7

Figure 7.2
Position of new seedlings Distribution of seedlings of the Brazil nut tree
30%
Percentage of seedlings

around parent trees in a forest in Northern

Bolivia. Shown is the percentage of seedlings
20% that occurs at a certain distance from the closest
adult tree. Distance values are the upper
boundaries of 5-m distance categories, e.g. the
10%
value 20 includes all distances of 15-20 m. This
graph is based on information of the spatial
0% distribution of 75 seedlings of less than 1 yr age
5

15

25

35

45

55

65
in relation to the position of 22 adult trees.
Distance to nearest adult tree [m] Source: Zuidema & Boot (2002).

to obtain reliable estimates of the number of seedlings per hectare. In the two
Bolivian studies in Figure 7.1, almost 6 ha have been systematically searched
for seedlings. These searches were carried out in subplots (of 20 by 20 m, or
25 by 25 m). In a third of these subplots, not any seedling was found. In the
other Bolivian study, 0.3 ha has been searched. In the Brazilian study, 51 ha
were searched for large seedlings of >1 m height. In conclusion, large sample
areas are needed to obtain good estimates of densities.
Recently germinated seedlings occur at some distance from the closest
adult trees. In Figure 7.2, the distances of new seedlings to adult Brazil nut
trees is shown based on information about the position of 75 young (<1 yr)
seedlings in relation to 22 adult trees in a site in Northern Bolivia. The spatial
distribution of these seedlings appeared to be random in relation to the adult
trees. This means that they are not clumped around the adult trees.
Small seedlings of the Brazil nut tree mainly occur at dark locations in
the understorey of the forest. Figure 7.3 shows the percentage of seedlings that
does not receive direct light due to small holes in the canopy of the forest.
They mainly receive indirect light. This figure also shows that the proportion

Figure 7.3
Light conditions of seedlings
60% Light conditions of seedlings of the Brazil nut
Percentage of seedlings

tree in two primary forest sites in Northern

No direct light
Substantial direct light Bolivia. The percentage of seedlings of different
40%
heights that do not receive direct light is shown
(black bars), as well as the percentage receiving
20% substantial direct light. Light conditions were
assessed using a modification of the Dawkins
crown position index (Dakwins & Field 1978,
0%
Zuidema & Boot 2002), which can be easily
10-35 35-70 70-140 140-230
determined by the observer in the field,
Seedling height [cm]
without the use of light measuring instruments.
The information in the graph is based on light
determination of 242 seedlings in Beni and
Pando, Bolivia. Source: Zuidema & Boot (2002).
 Seedling ecology • 51

Saplings and gaps - Data Saplings and light - Model

yes 100%
90%
Sapling present?

Sapling present?
50%
50%

10%
no 0%
0 10 20 30 40 0 10 20 30 40
Light level [GSF] Light level [GSF]

Figure 7.4
Occurrence of large seedlings (saplings) of the Brazil nut tree in relation to light availability in small
to large canopy gaps. The left graph shows the occurrence in relation to the light level in the middle
of canopy gaps. GSF is a parameter that indicates the amount of direct and indirect light that a plant
receives at a certain location. The right graph shows the model that relates the presence of saplings
to the GSF value. It can be seen that an increase in light results in an increase in the probability to
find a Brazil nut tree sapling. The statistical model in the right graph (logistic regression) explains
39% of the variation in occurrence of saplings. Data on saplings were collected in 50 gaps of different
sizes. Source: Myers et al. (2000), re-analysed in PROMAB (in press).

of seedlings that receive substantial amounts of direct light (through small holes
in the canopy) gradually increases with seedling size.
Large seedlings of the Brazil nut tree appear to occur more in gaps than
in the understorey of the forest. An inventory of gaps in a Bolivian forest
revealed that the probability to encounter a large seedling (>130 cm height)
in a gap increases with the light availability in that gap. Figure 7.4 shows this
relationship for 50 gaps, of which 15 contained a large seedling of the Brazil
nut tree.
Small seedlings (<130 cm height) of the Brazil nut tree may benefit from
higher light availability in gaps. On average these seedlings are almost 70%
taller when in a gap, compared to an understorey location. Heights of small
and large seedlings in gaps and forest understorey are shown in Figure 7.5.

Figure 7.5
Seedling height and gaps
400 Height of seedlings of the Brazil nut tree in
Seedling height [cm]

Understorey relation to their position in the forest. Small

Gap
300 seedlings are <130 cm in height, large seedlings
> 130 cm height. No large seedlings were found
200
in the understorey of the forest. On average
small seedlings were larger in gaps than in
100
understorey conditions. Data on seedlings was
0 collected in 50 gaps of different sizes and 20
Small seedling Large seedling understorey locations. Source: Myers et al.

Seedling category (2000).

52 • Chapter 7

Figure 7.6
Seedling survival
100% Survival of seedlings of the Brazil nut tree in
Survival during one year

two primary forest sites in Northern Bolivia

75% (black bars: Pando; white bars: Beni). The
information in the graph is based on
50%
measurements on 163 (Beni) and 79 (Pando)
seedlings during one year. Source: Zuidema &
25%
Boot (2002).
0%
10-35 35-70 70-140 140-230
Seedling height [cm]

7.2 Seedling survival under natural conditions

The information on seedling dynamics in natural populations of the Brazil

nut tree is scarce. To date, the results of only one study have been published.
This study was carried out in two primary forest sites in Northern Bolivia.
By marking and annually remeasuring the size of seedlings, information on
the growth and survival of seedlings was obtained and could be related to the
initial size of the seedlings and to their light conditions.
The survival probability for seedlings of the Brazil nut tree is relatively
high compared to that in other tropical trees. Figure 7.6 shows the percentage
of seedlings that survive, for different-sized seedlings and two forest areas. Even
for the smallest seedlings, at least 50% survive per year. From the graphs in
Figure 7.6 it becomes clear that the survival probability steeply increases with
the size of the seedlings. The survival chance for seedlings of 35-70 cm height
is at least one third higher than that for smaller seedlings (10-35 cm height).
Seedlings of over 70 cm height have an annual survival probability of almost
100%. A substantial part of the mortality of small seedlings is probably caused
by agoutis or other rodents that eat the cotyledon (seed reserve) of seedlings
of the Brazil nut tree. As the cotyledon is a part of the stem (see Figure 3.1),

Figure 7.7
Seedling survival and drought
100% Survival of seedlings (<230 cm height) of the
Survival during one year

Normal year
Brazil nut tree during a normal and a dry year.
Dry year
The difference in survival between the dry and
normal year is statistically significant only for
50%
the Beni site; in the Pando site, survival is the
same in the two years.. Data are from two
primary forest sites in Northern Bolivia. The
0% information in the graph is based on
Beni Pando measurements on 355 (Beni) and 202 (Pando)
Forest site seedlings during two years. Source: Zuidema &
Boot (2002).
 Seedling ecology • 53

this kills the seedling. High seedling mortality due to cotyledon predation has
also been reported from Peruvian (Ortiz 2002) and Brazilian studies (Baider
2002).
In the same Bolivian study, a lower seedling survival was found during an
exceptionally dry year (concurring with an “El Niño” episode). Figure 7.7
compares the survival of seedlings during this dry year with that of a normal
year. In one of the study sites a statistically significant difference in survival
was found. At this site mortality was 15% higher during the dry year.

7.3 Seedling growth under natural conditions

Height growth of Brazil nut seedlings in natural populations generally

increases with seedling size. Figure 7.8 shows the relation between growth and
initial height of seedlings in two natural populations in Northern Bolivia.
When seedlings become 1 m taller, their height growth rate increases by
approximately 6 cm yr -1. In this study, no difference between forest sites was
found in seedling growth.
Seedling growth was highly variable, as can be seem by the large standard
deviations for height growth in Figure 7.8. Part of this variation was due to
the large difference in light availability experienced by the seedlings. Seedlings
that received some direct light attained a substantially higher growth rate than
seedlings that received only indirect light.
Seedling growth was found to be lower during an exceptionally dry El
Niño year in two natural populations in Northern Bolivia. Rainfall during

Seedling growth - Beni Seedling growth - Pando

60 60
Height growth [cm yr-1]

40 40

20 20

0 0

-20 -20
0 50 100 150 200 0 50 100 150 200
Seedling height [cm] Seedling height [cm]

Figure 7.8
Height growth of seedlings of the Brazil nut tree in relation to height in two natural populations in
Northern Bolivia. The open dots denote the average height growth for seedlings in a size category.
Size categories are 10-35, 35-70, 70-140 and > 140 cm height. Vertical lines denote the standard
deviation of the growth rate. The closed dots represent the maximum growth rates observed for a
size category. The stippled lines are linear regressions indicating the trend of increasing growth with
increasing seedling size. Source: Zuidema & Boot (2002).
54 • Chapter 7

Figure 7.9
Seedling growth and drought
Height growth of seedlings of the Brazil nut
Normal year
Height growth [cm yr-1]

10 tree in a normal and a dry year. Data are from

Dry year
a study in natural populations in two forest
sites in Northern Bolivia, during a normal and a
5 dry year. Seedling growth was significantly
lower during the dry year. Data on seedlings
was collected in plots of 6 and 12 ha in
0 exploited forests; data of 40-100 seedlings per
Beni Pando site were used for the analysis. Vertical bars
Forest site (error bars) denote the standard deviation.
Source: Zuidema & Boot (2002).

the Niño year 1998 was considerably lower than normal. In Figure 7.9 height
growth of seedlings is compared between years: the difference in growth rate
was 4.0 cm yr -1 for Beni and 2.5 cm yr -1 for Pando. Dry years such as 1998
occur at an approximate frequency of once per eight years in Northern Bolivia.

7.4 Seedling growth in controlled experiments

A relatively large amount of information exists on the growth of seedlings

of the Brazil nut tree under experimental conditions. Seedling experiments
have been carried out in shade houses and at different conditions in the field.
The results of experimental seedling studies that are carried out in the field
as trials of enrichment planting activities are discussed in more detail in Part

Seedling growth vs. light Seedling growth vs. light

60 60
Height growth [cm yr-1]

40 40

20 20

0 0
0 25 50 75 100 0 25 50 75 100
Light availability [%] Light availability [%]

Figure 7.10
Height growth of seedlings of the Brazil nut tree in relation to light availability. Left graph: data
from a shade house experiment in which seedlings were put at 3, 6, 12, 25, 50 and 100% of full
daylight, during a period of 172 days. Right graph: data from a shade house experiment (using large
leaves to produce shading) with light availability of ca. 1, 10 and 100% of full daylight, during a
period of 98 days. The initial biomass of seedlings was similar: 45 cm for the left graph and 47 for
the right graph. The symbols denote the average height growth per light treatment; the stippled
lines show the general trend. Sources: Poorter (1999) (left graph) and Zuidema et al. (1999) (right
graph).
 Seedling ecology • 55

C, Chapter 12. In this section the results of experiments under controlled

conditions are discussed.
In seedling experiments, growth of Brazil nut seedlings has been related
to three parameters: light availability, water availability and initial seedling size.
Increasing light availability generally does not lead to higher height
growth. In one of the experiments included in Figure 7.10 (left graph), it is
shown that height growth tends to decline with increasing light availability.
In the other experiment (right graph in Figure 7.10), both very low and very
high light levels result in low height growth. This may be due to the very low
light availability in the treatment with 1% of full daylight. At high light levels
Brazil nut seedlings tend to produce many branches, thus investing more
biomass in horizontal spread than in vertical growth.
As opposed to height growth, growth in biomass generally increases with
higher light availability. In both experiments that are shown in Figure 7.11,
an increase in light availability always leads to an increase in biomass growth.
However, this increase in growth rate is much higher for low light levels than
for high light levels. For instance, increasing the light level from 3 to 6% (the
left-most symbols in the left graph) has a much larger impact on growth rate
than an increase from 25 to 50% light availability.
Combining the results of Figures 7.10 and 7.11 it may be concluded that
the most suitable light conditions for fast growth of Brazil nut seedlings is
probably at intermediate light levels. At a light availability of 10-25% both
height growth and biomass growth are relatively high. Both types of growth
are important when seedlings are used for enrichment planting or plantations.

Seedling growth vs. light Seedling growth vs. light

80 80
Biomass growth [g yr-1]

60 60

40 40

20 20

0 0
0 25 50 75 100 0 25 50 75 100
Light availability [%] Light availability [%]

Figure 7.11
Biomass growth of seedlings of the Brazil nut tree in relation to light availability. Left graph: data
from a shade house experiment in which seedlings were put at 3, 6, 12, 25, 50 and 100% of full
daylight, during a period of 172 days. Right graph: data from a shade house experiment (using large
leaves to produce shading) with light availability of ca. 1, 10 and 100% of full daylight, during a
period of 98 days. The initial size of seedlings was similar: 13 g for the left graph and 12 for the right
graph. The symbols denote the average biomass growth per light treatment; the stippled lines show
the general trend. Sources: Poorter (1999) (left graph) and Zuidema et al. (1999) (right graph).
56 • Chapter 7

Figure 7.12
Seedling growth and water
60 Height growth of seedlings of the Brazil nut
3% Light
Height growth [cm yr-1]

tree in relation to light and water availability.

50 25% Light
Data are from an experiment with three
40
watering treatments (low, medium and high),
30 and two light regimes (3 and 25% of full
20 daylight). Watering treatments differed in the
frequency of watering: low = once per 15 days;
10
medium = once per 6 days; high = once per 3
0
days. Watering was done by giving 0.5 l of
Low Medium High
water to each plant; this is equivalent to 28 mm
Water availability
of rain. A total of 36 plants were followed for
110 days. An analysis of variance of these data
showed that only light had a significant effect
on growth; there was no influence of water
availability on seedling growth. Source:
Hayashida-Oliver et al. (2001).

Good height development is important as this allows seedlings to compete

with neighbouring plants and reach higher layers in the forest with higher
levels of light. Good biomass growth is important as this implies good
development of branches and leaves.
A second factor that influences seedling growth is the availability of water.
As relatively dry seasons occur throughout the distribution area of the Brazil
nut tree, it is important that seedlings of this species tolerate periods of
(moderate) drought. In the previous section it was shown that an exceptionally
dry year caused seedling growth and survival to be reduced (Figures 7.7 and
7.9). The effect of water availability was tested in a controlled experiment in
shade houses in Bolivia. Figure 7.12 shows the results of this experiment.
Contrary to the field measurements (Section 7.2) no effect was found of water
availability on height growth of Brazil nut seedlings. This can probably be
explained by the fact that the lowest water availability that was simulated in
this experiment (watering every 15 days) was not sufficiently low to reduce
seedling growth. In fact, rainless periods of this length occur on average twice
per year in Northern Bolivia (Hayashida-Oliver et al., 2001). Nevertheless,
from the experiment it becomes clear that growth of Brazil nut seedlings is
not affected by short to moderately long periods without rain.
The third factor that influences seedling growth is the initial seedling size.
An experiment to test this relation was carried out under controlled conditions
in Bolivia. Brazil nut seedlings that were used for the experiment differed in
age (and thus in size): 2 months (14 cm height), 5-6 months (47 cm height)
and 24 months (80 cm height). The results of this experiment are presented
in Figure 7.13. Height growth was highest for intermediate-sized seedlings,
and lower for small and large seedlings (left graph of Figure 7.13). Biomass
 Seedling ecology • 57

Seedling growth vs. size Seedling growth vs. size

60
30

Biomass growth [g yr-1]

Height growth [cm yr-1]

40 20

20 10

0 0
0 20 40 60 80 0 10 20 30
Initial height [cm] Initial biomass [g]

Figure 7.13
Growth of seedlings of the Brazil nut tree in relation to initial size. Left graph: height growth of
different sized seedlings; right graph: biomass growth of different-sized seedlings Data are from a
shade house experiment (using large leaves to produce shading) with light availability of ca. 10% of
full daylight, during a period of 98 days. The symbols denote the average biomass growth per light
treatment; the stippled lines show the general trend. Source: Zuidema et al. (1999).

growth generally increased with seedling size, although it was slightly lower
for large plants (right graph of Figure 7.13). In practice this implies that
seedlings of around 40-60 cm height are probably more suitable for the
establishment of plantations or for enrichment planting than smaller and larger
seedlings. Apart from attaining higher growth rates than smaller or larger
seedlings, these seedlings also suffer less from cotyledon predation when
transplanted to natural situations. Cotyledon predation is an important cause
of seedling mortality, especially for small seedlings.
 Chapter 8

Adult ecology

8 . 1 Tr e e a r c h i t e c t u r e

Brazil nut trees can attain very large sizes. They can reach heights of up
to 50 m, and a diameter at breast height (DBH) of over 3 meters. Adult trees
(of >60 cm DBH) have a straight unbranched stem of 23 m height; at that
height 2-4 main branches are formed that are often strongly curved. Full crown
development usually takes place when the tree crown has reached the emergent
layer in the forest (i.e. crown is positioned above the canopy).
Initial growth of Brazil nut trees seems to be directed more towards height
growth than diameter growth. This can be concluded from the relation
between diameter and height, shown in Figure 8.1. Up to 20-25 cm DBH,
the increment in height per cm increment in DBH is large (steep slope of the
curve), whereas trees of over 40-50 cm do not grow much anymore in height.
For example, trees between 2 and 20 cm DBH have an increment of 1 m in
height per cm increment in DBH. For trees between 20 and 40 cm DBH, the
height increment is 0.5 m per cm DBH increment. For larger trees, this figure
is even smaller.This growth pattern is probably an advantage to reach the forest
canopy as fast as possible.

Height and DBH Figure 8.1

Relation between diameter at breast height
40
(DBH) and tree height of Brazil nut trees in two
Total height [m]

natural populations in Northern Bolivia. The

30
stippled line is a logarithmic regression that
20 explains 94% of the variation in tree height on
the basis of the value of DBH. The graph is
10 based on measurements of 201 trees in two
natural populations (in Beni and Pando, Bolivia).
0
Source: Zuidema & Boot (2002).
0 50 100 150 200
DBH [cm]
60 • Chapter 8

DBH and crown area Figure 8.2

800 Relation between diameter at breast height

(DBH) and crown area of Brazil nut trees in a
Crown area [m2]

600 natural population in Northern Bolivia. The

stippled line is an exponential regression that
400 explains 52% of the variation in crown area on
the basis of the DBH value. The graph is based
200 on measurements of 40 trees in a natural
population in Beni, Bolivia. Source: Leigue
0
Gómez & Boot (in press a).
0 50 100 150

DBH [cm]

The late development of the crown in Brazil nut trees can also be seen
in Figure 8.2. In this Figure, the relation between crown area and DBH is
shown for 40 trees in Beni, Bolivia. Crown area increases rapidly with DBH
in trees with a DBH of over 100 cm DBH. Trees of this size are all emergent
(see below): this implies that there is no space limitation to the development
of a large crown.This graph also shows that crown growth continues for larger
trees (trees >150 cm DBH).
The canopy position of Brazil nut trees in natural forests is closely related
to the DBH of the tree. Figures 8.3 shows this relation for a natural population
in Northern Bolivia. It can be seen that trees that reach the canopy have a
DBH of between 20 and 30 cm. The following transition from canopy to
emergent, takes place for trees between 40-60 cm DBH. These relations are
important as it has been shown previously (Figure 5.2) that the reproductive
status is strongly correlated with the canopy position of trees: the vast majority
of reproductive trees is emergent.
Similar results were obtained for a large population in Acre, Brazil (Figure
8.4). Canopy openness of trees in different size categories strongly increased
with DBH. The proportion of trees smaller than 40 cm DBH which receive
full light (canopy openness of 100%) is very low (<5%), whereas this is over
Figure 8.3
Trees and light - Bolivia
Relation between diameter at breast height
5
Crown illumination score

Emergent (DBH) and crown illumination of Brazil nut trees

4 Canopy in two natural populations in Northern Bolivia.
Crown illumination values are Dawkins scores:
3 1=no direct light; 2=some lateral direct light;
3=substantial lateral direct light and some
2 Sub-canopy
overcast light (sub-canopy); 4=full overcast
light (in canopy); 5=full overcast and lateral
1
crown illumination (emergent). Graph is based
0 50 100 150 200 250 300
on measurements of 364 trees in two natural
DBH [cm]
populations (in Beni and Pando, Bolivia).
Source: Zuidema & Boot (2002).
 Adult ecology • 61

Trees and light - Brazil Figure 8.4

1,00
Canopy Relation between diameter at breast height
openness
Proportion trees

0,75 100% (DBH) and canopy openness for Brazil nut

75% trees in Acre, Brazil. The canopy openness
0,50 50% was estimated for each tree in a natural
25% population in 51 ha of undisturbed forest.
0,25
0% The total number of trees included in the
0,00 study is 299. Source: Viana et al. (1999).
40 80 120 160 >160
DBH category [cm]

75% for trees of more than 40 cm DBH. For larger trees, the increment in the
proportion of trees that receive full light is much smaller: almost all are
emergent.

8 . 2 Tr e e d i a m e t e r g r o w t h

Diameter growth (DBH) of Brazil nut trees is generally high compared

to other tropical tree species. DBH growth was studied in two natural
population in Bolivia over a period of 2-3 year. Maximum diameter growth

DBH growth - Beni, Bolivia

2,0
DBH growth [cm yr-1]

Mean growth
1,5 Maximum growth

1,0

0,5

0,0

DBH growth - Pando, Bolivia

DBH growth [cm yr-1]

2,5
2,0
1,5
1,0
0,5
0,0
20-30

30-40

40-50

50-60

60-80

80-100

>160
100-120

120-140

140-160
1-4

4-9

9-20

DBH Category
. [cm]

Figure 8.5
Growth of diameter at breast height (DBH) for Brazil nut trees belonging to different DBH categories
in two natural populations in northern Bolivia. Bars represent the average growth rate; triangles
denote the maximum observed growth rate for a category. Data are averages over 2-3 yr, for 167
trees in Beni and 130 trees in Pando. It should be noted that DBH growth can be highly variable in
certain categories. Source: Zuidema & Boot (2002).
62 • Chapter 8

DBH growth model - Beni, Bolivia

DBH growth [cm yr-1]

1,0

0,5

0,0
0 25 50 75 100 125 150 175 200

DBH growth model - Pando, Bolivia

1,5
DBH growth [cm yr-1]

1,0

0,5

0,0
0 25 50 75 100 125 150 175 200
DBH [cm]

Figure 8.6
Relation between growth in DBH (diameter at breast height) and initial DBH for Brazil nut trees in
two natural populations in northern Bolivia. The curves represent the outcome of a regression model
using the Hossfeld IV growth equation. This equation is frequently used in forestry studies to relate
DBH or height growth to the size or age of trees. More information on this equation can be found in
a publication by Zeide (1993). The shown curves explain 23% of the variation in DBH growth in the
Beni site; and 41% of the variation in the Pando site. The data used for the model are growth
measurements over 2-3 yr, for 167 trees in Beni and 130 trees in Pando. Source: Zuidema & Boot (2002).

rates were often over 1.5 cm yr-1; the absolute maximum rate was 2.6 cm yr-1
for a tree of 43 cm DBH. Figure 8.5 shows the diameter growth rate of
different size categories in the two studied populations. The general pattern
that is observed in these graphs is that DBH growth is low - generally less
than 0.5 cm yr -1 - for small and large trees, and much higher for trees of
intermediate size. Also, it is shown that the maximum growth rate in a category
is often two times that of average growth rate.This implies that Brazil nut trees
have high potential growth rate (see also Section 12.5 in Part C on tree growth
in plantations).
The pattern of lower DBH growth for small and large trees, and high
growth at intermediate tree sizes can be described using growth equations
used in forestry research. A number of such equations have been developed to
describe patterns of tree growth (in diameter or height) in relation to tree size
or tree age. One such equation - the Hossfeld IV equation - was used to relate
DBH growth to initial DBH for the data shown in Figure 8.5. The resulting
curves are presented in Figure 8.6. The typically higher growth rates in
intermediate-sized trees are found especially in the dataset from Pando (lower
 Adult ecology • 63

Figure 8.7
DBH growth and light
1,5 The influence of crown position on DBH growth
Crown in or above canopy
DBH growth [cm yr-1]

of Brazil nut trees. The two lines denote the

Crown below canopy
average growth of trees of a certain DBH which
1,0
have their crown either below the forest canopy
or in/above the canopy. The growth rate of
0,5 trees that are in or above the canopy is on
average 0.36 cm yr-1 higher than that of trees
below the canopy. Note that only trees <40 cm
0,0
DBH are included. Data are from a natural
0 10 20 30 40
population of 130 Brazil nut trees in Beni,
Initial DBH [cm]
Bolivia. Source: Zuidema & Boot (2002).

graph in Figure 8.6.). This peak is less clear for the growth data from Beni.
The reason for this difference -which was found to be statistically significant
- is not known. Differences in soil conditions or other a-biotic factors may
play a role. Forest structure and composition of the two sites are comparable.
Apart from size (DBH) also light plays a role in determining diameter
growth. In the mentioned Bolivian study on tree growth, crown position of
Brazil nut trees was found to influence DBH growth for trees with equal DBH.
Figure 8.7 shows the influence of both DBH and crown position on DBH
growth. On average, DBH growth increases by 0.4 cm yr -1 when the crown
of a tree is located in or above the canopy, as compared to when the crown is
located below the canopy.
Compared to other tree species, the Brazil nut tree is a fast growing
species. Figure 8.8 shows the median DBH growth for a set of 32 tree species
in Beni, Bolivia, and compares this with the growth rate of the Brazil nut tree.
Whereas most tree species in this area grow at rates between 0.04 and 0.2 cm
in DBH per year, the median diameter growth rate of Brazil nut trees amounts
Figure 8.8
Growth compared to other trees
Median DBH growth [cm yr-1]

0,5 Median diameter growth rate of the Brazil nut

tree (black triangle) in Bolivia compared to that
0,4
of 35 Bolivian tree species. The horizontal line
0,3 Brazil nut tree in the box indicates the median value of the
growth rates of the 35 Bolivian tree species; the
0,2 upper and lower boundary of the box are the
75% and 25% percentiles (indicating that
0,1
respectively 75% and 25% of the growth rates
0 is below that value); and the ends of vertical
Beni, Bolivia lines indicate the 90 and 10% percentiles. Open
circles are the extreme values. There is one
species with a median DBH growth rate of 9.5, which falls outside the borders of the graph. The
values for the Brazil nut tree were obtained from 3-yr measurements in a natural population in Beni,
Bolivia; for the other tree species rates were obtained from 4-yr measurements in a permanent sample
plot at 1-km distance of the Brazil nut tree population. Source: Zuidema & Boot (2002) (for Brazil nut
trees); Poorter et al. (2001) (other species).
64 • Chapter 8

to 0.34 cm per year. Also maximum DBH growth of the Brazil nut tree (over
2.5 cm per year) are high in comparison with those of other tropical tree
species. The majority of tropical tree species do not reach a maximum DBH
growth rate of 2.0 cm per year.

8 . 3 Tr e e s u r v i v a l

Very little is known about mortality rates of Brazil nut trees. A

demographic study at two sites in Bolivia showed that mortality in Brazil nut
trees is very low: only 4 out of 374 trees of >1 cm DBH died in two years
(Source: Zuidema & Boot 2002). This implies an annual mortality rate of 0.5
%. Thus, 99.5% of all trees of this size survive per year. As these figures are
only based on a few observed deaths, it is good to know how variable this
figure of 99.5% is. It was calculated that there is a 95% chance that survival
probability lies in the range of 98.7-99.9% per year. Presumably, the survival
rate of small trees (less than approximately 10 cm DBH) and large trees (over
approximately 100 cm DBH) is lower. The former are more susceptible to die
when they are hit by a falling tree or branch. The latter may gradually senesce.
In fact, in the same study it was found that almost 7% of the trees larger than
80 cm DBH had lost one or more of their main branches. As mature Brazil
nut trees typically have 2-4 main branches, loosing one implies the loss of a
large part of the total leaf area of the tree.
In Peru, the main cause of mortality of Brazil nut trees in natural
populations is reported to be strong winds after heavy rains and lighting, both
causing trees to be uprooted or snapped off. Ortiz (in press) describes a massive
tree fall of 400 Brazil nut trees in several hundreds hectares after hurricane-
like winds following several days of rain in 1991.

8 . 4 Tr e e a g e s

Age estimates of Brazil nut trees have been determined in two studies,
using two different techniques. In the first study, an estimate of tree age was
obtained using a carbon isotope ( 14C) that is frequently used to date timber.
This was done for one Brazil nut tree of 225 cm DBH, being the largest tree
from a natural population of 45 trees in 9 ha, Marabá, Brazil. The age of this
tree was found to be 440 year with a standard deviation (measure of
uncertainty) of 60 years. This implies that the average DBH increment over
the complete life of this tree has been 0.6 cm yr-1.
 Adult ecology • 65

Tree age - to reach DBH category

400
Estimated age [yr]
300

200

100

Tree age - average in DBH category

Estimated age [yr]

400
300
200
100
0
20-30
30-40
40-50
50-60
60-80
80-100

>160
100-120
120-140
140-160
1-4
4-9
9-20

DBH Category [cm]

Figure 8.9
Age estimates of Brazil nut trees in different DBH categories. Age estimates shown in bars are based
on demographic models (matrix population models). The estimates shown as filled circles are based
14
on one tree that has been dated using C isotopes (see text for further explanation). The estimates
in bars are from a demographic study in a natural population of the Brazil nut tree in Pando, Bolivia.
Age estimates of a second research site in Beni (Bolivia) were very similar. Further explanation of
these models is provided in Chapter 7. The error bars (vertical lines) denote the standard deviation
(a measure of variability) of the estimates. Source: Zuidema & Boot (2002) (bars) and Camargo et al.
(1994) (dots).

In the other study, age estimates have been obtained using a certain type
of models that describe population dynamics (matrix population models).
These models are frequently applied to analyse the demography of tree species.
Using an extension of these models, age estimates can be obtained for different
size categories. The results of these analyses are presented in Figure 8.9. Both
the age at reaching a certain DBH category and the average age in a certain
category are shown. It can be seen that the age at reaching the first category
with a high proportion of reproductive trees (60-80 cm DBH) is around 125
year, with a standard deviation of 50 year. The average age in the last size
category (>160 cm DBH) amounts to over 350 year (standard deviation of
100 year).
Figure 8.9 also shows that the results of the two ageing studies are
comparable. When applying the 0.6 cm annual DBH increment that was
predicted from the first age estimate to the size categories used in the second
model, very similar age estimates are obtained. Nevertheless, it would be very
useful to obtain more 14C age estimates, of a large set of trees, in order to be
able to better verify the age estimates produced by demographic models.
 Chapter 9

Population dynamics

9.1 Relevance of studying population dynamics

Studies on population dynamics analyse the changes in the size of

populations of a certain species in time. Such studies can determine the future
prospects for populations of exploited species. Despite the economic
importance of the Brazil nut tree, there is only limited information on its
population dynamics.To date, the only published study on population dynamics
of the Brazil nut tree is based on a study in two natural populations in the
Bolivian Amazon region, where Brazil nuts are collected (Zuidema & Boot
2002).
Why are studies on population dynamics needed for exploited trees such
as the Brazil nut tree? The answer is that such studies are an important tool
to determine the impact of exploitation on the maintenance of populations
of a certain tree species, and the future availability of the resource that is
obtained from that tree species. Thus, these studies can be used to determine
the effect of harvesting nuts, leaves, bark or fruits, on the continued existence
of the species from which they are harvested, and on the availability of these
products in the future.
How do studies on population dynamics reach that goal? First, they help
to find out what the most important processes are in the life cycle of a tree.
This is done by analysing the natural population dynamics of the species.
Secondly, they evaluate how the extraction of certain plant parts (nuts, leaves,
etc.) influences certain processes in the life cycle and how this affects the size
of the entire population in time. For example, the collection of Brazil nuts has
an influence on the availability of seeds for germination, since a large
proportion of seeds is taken away. If in a population dynamics study, it is found
that the availability of seeds is very important for the maintenance of the
68 • Chapter 9

population of Brazil nut trees, a large impact of exploitation may be expected

on the whole population.
In this Chapter the results of a population dynamics study on the Brazil
nut tree are presented. This chapter focuses on the analysis of the population
dynamics, without looking at the impact of exploitation. In Chapter 11 of
Part C of this book, the impact of nut collection on population dynamics is
discussed, based on the same study. The following three Sections deal with the
methodology of population dynamics studies (the concepts, the model and the
methods of the field study), and the last three Sections treat the results of the
population dynamics study on the Brazil nut tree.

9.2 The life cycle

The concept of the life cycle is important for studies on population

dynamics.The life cycle shows how an individual tree moves through all phases
in the course of its life: from seed to seedling, from seedling to juvenile tree,
and eventually to a reproductive tree. An example life cycle for an imaginary
tree species is shown in Figure 9.1. The arrows in this diagram denote the
processes in the life cycle: growth, survival and reproduction. The circles are
the different classes in which individuals can be grouped, in this case seeds,
seedlings, juvenile trees and two categories with adult trees. These classes can
be based on size, or on age. For tree species, usually size is used to classify
individuals. In this case, different parameters can be used to distinguish
categories, e.g. diameter, height, but also number of leaves. The number of
classes in the life cycle can be varied. In the example life cycle a small number
of classes is chosen for the sake of clarity.
The essence of studies on population dynamics is to quantify the various
processes in the life cycle. When all processes are quantified, the influence of
growth, survival and reproduction of all individuals together can be used to

Life cycle of a tree

Seed production

Sd Sdl Juv Ad1 Ad2

Dispersal & Growth & Growth & Growth &

germination survival survival survival

Figure 9.1
Schematic representation of the life cycle of a tree species. Circles denote the different classes in the
population. Arrows are the processes in the life cycle. Sd = Seeds, Sdl = Seedling; Juv = Juvenile tree;
Ad1 = Small adult tree; Ad2 = Large adult tree.
 Population dynamics • 69

determine the development of the size of the population in time. Different

types of models may be used to calculate the consequences of growth, survival
and reproduction of individuals for the entire population. One such type of
model - a population matrix model - has been used for the study on the Brazil
nut tree.

9.3 Population matrix models

Population matrix models are frequently used for the analysis of

population dynamics of plant species, including exploited plant species. They
use the information on survival, growth and reproduction per class in the life
cycle to calculate the development of population size and population structure
in time.
The basic idea of population matrix models is that the processes in the
life cycle are considered as transitions among categories. An example: to study
the survival, growth and reproduction of a population of trees, this population
is measured two times, with one year between the measurements. During this
year, individuals may die, produce seeds, and/or grow to the next size class.
For instance, an individual that is classified as a seedling may grow to the next
category (Juveniles) during this year. By studying and measuring all individuals
in the population repeatedly in time, information is obtained on all transitions
among categories. These transitions are used to obtain the average probability
for each size category. Such transition probabilities can be ordered in a table,
which is called a transition matr ix. This matrix is the basis of the population
matrix model.
Figure 9.2 shows how different transitions in the life cycle of a tree can
be ordered in a transition matrix. Three groups of transitions can be
distinguished: growth (G) means that an individual survives and grows to the
next category; remaining (R) implies that an individual survives, but stays in
the same category; seed production (S) is the number of seeds that is produced
by an individual of a certain size. Individuals that die are not specifically
included in the life cycle graph or in the transition matrix. But implicitly,
mortality is taken into account: the sum of growth (G) and remaining (R)
transitions equals the survival probability in a category.
The transition matrix summarises all transitions among the categories.
The columns in the transition matrix represent the situation (i.e. the class) at
the start of the year, and the rows represent the situation after one year. Each
type of transition (G, R and S) has a certain position within the matrix. All R
(remaining in category) transitions are on the diagonal of the matrix, as the
situation at the start and after one year is the same. For instance, the probability
70 • Chapter 9

Life cycle and matrix model

G rowing to next category
R emaining in category
S eed production

Sd Sdl Juv Ad1 Ad2

In class at start
In class after one year

Sd Sdl Juv Ad1 Ad2

Sd R 0 0 S S
Sdl G R 0 0 0
Juv 0 G R 0 0
Ad1 0 0 G R 0
Ad2 0 0 0 G R

Figure 9.2
An example of the codification of life cycle processes in a transition matrix. Circles in the life cycle
denote the different classes in the population. Sd = Seeds, Sdl = Seedling; Juv = Juvenile tree; Ad1 =
Small adult tree; Ad2 = Large adult tree. Arrows are the transitions among life cycle classes. R = the
probability than an individual survives and remains in the same category during a certain period of
time (usually one year); G = the probability that an individual survives and moves to the next category
during one year; S = the seed production of an adult tree during one year. In the transition matrix
(the table), columns represent the class at the start, and rows denote the situation after a certain
period of time (usually one year). In this way, all processes in the life cycle can be included in the
transition matrix, which then contains a summary of the dynamics of the population.

that a seedling will remain a seedling is put in the second column and second
row in the transition matrix. All growth (G) transitions are in the below the
diagonal as this implies the transition from one class to the next class. For
instance, the probability that a juvenile grows into a an adult-1 tree is noted
in the third column and the fourth row. All seed production (S) transitions are
on the first row, and only for those columns that represent adult classes:
individuals in an adult class at the start, will have produced a certain number
of seeds after one year. Thus, the number of seeds produced by an adult-1
individual is put in the fourth column and the first row.
When filled with information on transitions among categories, the
transition matrix contains important information on the dynamics of the study
population. Now, matrix algebra can be used to simulate the development of
the population in time. In essence, this is done by multiplying the transition
matrix with the population structure (in the form of a vector) at the start of
the study. For these calculations, the population structure is presented in the
form of a vector (i.e. one column of information), which contains the number
or abundance of individuals in each size category. By multiplying the transition
 Population dynamics • 71

matrix with the population structure, another population structure is obtained:

the population structure after one year.This process can be continued for many
years to simulate the population development over longer time spans. In this
way, a population growth rate can be calculated. Also, the development of the
population structure in time can also be determined.

9.4 Methodology of population dynamics study

To analyse the population dynamics of a tree species and to develop a

matrix population model, a field study has to be carried out in which
measurements on the same trees are repeated in time. This Section gives an
example of the methods used to study two populations of the Brazil nut tree
in the Bolivian Amazon.
In two sites, at a distance of c. 140 km, the demography of the Brazil nut
tree was studied in permanent plots in primary forest where Brazil nuts have
been collected over several decades. Permanent plots of 6 and 12 hectares were
established. For the two sites, a total of 600 individuals of the Brazil nut tree
were identified, measured and re-measured over a period of 2-3 yr. Individuals
of all sizes were included: from recently germinated seedlings to large emergent
adult trees. To increase the number of trees included in the study, trees outside
the permanent sample plot were also measured. As adult Brazil nut trees occur
at densities of around 2-3 tree per ha, an area of 6 or 12 ha is not sufficiently
large to obtain a reasonable number of individuals. Thus, trees were also
searched for in an area of several square kilometres surrounding the plots.
Within the plots, sub-plots were established where seedlings were marked and
measured.
Figure 9.3 shows the layout of plots and sub-plots in one of the two
populations. Six subplots per ha were selected at random. In these subplots,
seedlings and pole-sized trees (<5 cm DBH, diameter at breast height) were
searched. In total, the area searched for seedlings and pole-sized trees was 4.5
ha in the 12-ha plot and 1.4 ha in the 6-ha plot. The Figure also shows the
location of adult and juvenile Brazil nut trees in the study plots.
During a period of 2-3 year the following measurements were carried
out. Of seedlings and saplings (up to 1 cm DBH), total plant height was
measured each year to calculate growth rate. Of all individuals larger than 1
cm DBH, the diameter at breast height was measured. The location at the tree
where the diameter was measured was painted to assure that all following
measurement would be taken at exactly the same position. Also, it was noted
whether trees were reproductive.
72 • Chapter 9

Study plots for demography study - Beni, Bolivia

100 m

9 10 11 12
300 m

5 6 7 8

1 2 3 4

100 m
400 m

Subplot of 25 x 25 m in which seedlings of the Brazil nut tree have been searched

Figure 9.3
Map of the study plots in Beni, Bolivia, used for a population dynamics study on the Brazil nut tree.
The total area of the study plots is 12 ha. Numbers refer to hectares. The stippled cells indicate
subplots of 25 ×25 m where seedlings were studied. Dots represent reproductive Brazil nut trees (>60
cm DBH). Triangles are trees of 1-60 cm DBH. Source: Zuidema & Boot (2002).

Fruit production was determined for part of the study trees (40 trees per
study site). All fruits opened by Brazil nut collectors were counted for these
trees. The amount of fruits that was not opened by the collectors was
determined by searching fruits in the seedling subplots.

9.5 The population growth rate

The population growth rate of the two study populations of Brazil nut
trees indicated that the population size is probably stable, i.e. no large increases
or decreases in population size were found. In numbers, the two studied
populations were found to slightly grow: 0.7 to 1.7% per year. These were the
figures for a year with normal precipitation. However, during an exceptionally
dry year, the population was found to decline in one of the study sites (by
0.5% per year, Beni) and grow less in the other site (by 0.4% instead of 0.7%
per year in Pando). When sequences of dry and normal years are simulated in
the population models, fluctuations in population size are found. Figure 9.4
 Population dynamics • 73

Figure 9.4
Population growth - Bolivia
1500 Simulations of the development in time of a
Simulated population size

Normal Brazil nut tree population. Three scenarios were

Alternating
Dry simulated: only years with normal precipitation
(‘Normal’), only years with low precipitation
1000 (‘Dry’) and sequences of normal and dry year
(‘Alternating’), during which dry years occur at
random in time at a frequency of one dry year
every eight years. These simulations are based
500 on a population matrix model for a natural
0 20 40 60 80 100 population of the Brazil nut tree in Pando,
Simulation time [yr] Bolivia. Source: Zuidema & Boot (2002).

shows an example of these fluctuations for one of the study populations: when
dry years occur the population growth rate is reduced. Over long periods
during which dry years occur at a frequency of one out of eight years, the
population is growing nor declining in size.
The finding that the size of populations where Brazil nuts are collected
does not notably changes in time, is an indication that the exploitation of this
species can be sustained over longer time periods. The sustainability of Brazil
nut extraction is discussed in more detail in Chapter 11 in Part C.

9.6 Important processes and size classes

in the life cycle

Not all processes in the life cycle of the Brazil nut tree are equally
important for the maintenance and growth of the population. Certain processes
in the life cycle may be crucial for population growth, whereas others are of
minor importance. Similarly, different stages of the life cycle may have a
different impact for population growth. Using matrix population models (see
Section 9.3), the importance of the different processes and different stages in
the life cycle can be analysed. This analysis is called sensitivity analysis: it
considers how sensitive the population growth is rate for changes in certain
processes in the life cycle.
The results of the sensitivity analysis for the Brazil nut tree population
models show that especially the survival of juvenile and adult individuals of
the Brazil nut tree is important for the maintenance of the population. Figures
9.5 and 9.6 provide the results of the model analysis. In Figure 9.5 it is shown
that the juvenile and small adult categories are of special importance for the
maintenance and growth of the population. Changes in the growth and survival
of individuals in these categories have a strong impact on the population
growth. For instance, a 10% change in the survival of adult trees leads to a
change in population growth rate of 1.1% per year: if adult survival is decreased
74 • Chapter 9

Importance for population growth

15
Measure of importance Seedling Juvenile Adult

10

0
35-70

20-30
30-40
40-50
50-60
60-80
<35

70-140

80-100

>160
140-230

100-120
120-140
140-160
1-4
4-9
9-20

Height [cm] DBH [cm]

Figure 9.5
The importance of different classes in the life cycle of the Brazil nut tree for the maintenance and
growth of the population. The first four classes are based on height; the remaining classes on diameter
at breast height (DBH). These results are based on a matrix population model which was established
for a natural population of the Brazil nut tree in Beni, Bolivia. The values are the output of a specific
type of sensitivity analysis: ‘elasticity analysis’. Source: Zuidema & Boot (2002).

by 10%, the population growth rate will decrease by 1.1%. Seedlings and large
adult trees are of less importance. A change in survival of small seedlings by
10%, causes the population growth rate to decrease by only 0.11%.
Survival appears to be the most critical process in the life cycle of the
Brazil nut tree. The survival of juvenile and adult trees have by far the highest
‘importance values’ for the population growth rate. Figure 9.6 shows that they
together make up 90% of all summed ‘importance values’. The growth of
individuals is much less important, and reproduction is the least important
process.

Figure 9.6
The importance of different processes in the life

Importance for population growth cycle of the Brazil nut tree for the maintenance
and growth of the population. Stippled slices in
Growth Seed
production the pie refer to survival, the white slice refers
Seedling
survival to the summed importance of seedling, juvenile
and adult growth, and the black slice to the
importance of seed production. The proportion
of the total importance values that is accounted
Juvenile for by seed production equals 0.5%. These
survival
Adult results are based on a matrix population model
survival which was established for a natural population
of the Brazil nut in Beni, Bolivia. The values are
the output of a specific type of sensitivity
analysis: ‘elasticity analysis’. Source: Zuidema &
Boot (2002).
 Population dynamics • 75

The reason that survival of juveniles is crucial for the growth of the
population is probably as follows. It takes a very long time before a Brazil nut
tree produces fruits. As only the production of new individuals through seeds
can assure the growth and continued existence of the population, it is very
important that trees reach the reproductive stage.To do so, they need to survive
for at least 110 yr (the time needed for a pole-sized tree of 1 cm diameter to
reach a diameter of 60 cm, see Figure 8.9). Surviving this period of time is
therefore of crucial importance. Small changes in survival probability of
juvenile trees will have a great impact on the number of juveniles that will
finally reach the adult stage. When juvenile survival is decreased from 0.99 to
0.98, this implies that out of 100 trees of 1 cm diameter, only 13 trees will
survive for 110 years instead of 37.
The finding that seed production has a very small influence on population
growth is an indication that the collection of seed (Brazil nuts) from the
population probably has a small impact on the future size of the population
and the future production of Brazil nuts. This is further discussed in Section
11.4 in Part C.

9.7 The stable population structure

Apart from the population growth rate and the sensitivity analysis,
population matrix models also provide information on the population structure
after many simulation years, the stable population structure. This stable
population structure contains the relative abundance of individuals in different
size categories. It can be compared to the population structure that has been
obtained in the field study. If the two structures are similar, this is an indication
that the model is realistic. Thus, the stable population structure can be
considered as a tool to validate the model.
Figure 9.7. shows the stable population structure for the two study
populations of the Brazil nut tree for which population matrix models were
constructed.The stable population structure (dots) is compared to the observed
population structure (bars). Although there are some differences between the
two structures, in general the pattern is similar. This implies that the model
represents the real population dynamics well.
In one of the study sites (Pando), the population structure that is obtained
from the matrix model shows a slight increase in the abundance of reproductive
trees (> 60 cm DBH). That is, the abundance values of simulated population
structure (dots in Figure 9.7) are somewhat higher for trees of 60-120 cm
DBH, compared to smaller and larger trees. This slight “hump” in the
76 • Chapter 9

Population structures: observed and simulated

10
Beni

Abundance [ha-1]
Observed population structure
Simulated population structure
5

10
Pando
Abundance [ha-1]

0
35-70

20-30
30-40
40-50
50-60
60-80
70-140

80-100

>160
140-230

100-120
120-140
140-160
1-4
4-9
9-20

Height DBH [cm]

Figure 9.7
A comparison of the population structure of the Brazil nut tree as observed in two natural
populations, and that obtained from simulations using a model to analyse population. The observed
and simulated population structures are rather similar, which indicates that the population dynamics
are well described in the matrix population model. These results are based on a matrix population
model which was established for a natural population of the Brazil nut in Beni, Bolivia. Source:
Zuidema & Boot (2002).

population structure has been frequently found for the Brazil nut tree. Figure
4.3 shows examples of this typical population structure from Bolivia and Brazil
(the general pattern is discussed in Section 4.2). All population structures
except one from Pará, Brazil (the left upper graph in Figure 4.3) show such a
pattern.
In Section 4.3 two explanations for this “hump” in the population
structure were mentioned: (1) that Brazil nut trees have been planted by
Amerindians in certain period of time, and the trees in the hump are of the
same age; and (2) that the regeneration Brazil nut trees requires large-scale
disturbances which only occur sporadically and thus lead to cohorts of trees
of the same age and size. The simulated population structure resulting from
the matrix model, provides a third and more likely explanation of the typical
“hump”. That is, the hump may be the result of the inherent characteristics
of the Brazil nut tree, as the tree typically grows rapidly in certain size
categories, thus causing the abundance to be low, and it grows slowly in other
categories, thus causing a higher abundance.
Figure 9.8 compares the growth in diameter (DBH) of different-sized
trees with the abundance of trees that is simulated using a population matrix
model. It is clear that the low abundance of trees of 40-80 cm DBH coincides
 Population dynamics • 77

DBH growth and relative abundance

1,5 0,4
DBH growth [cm yr-1]

Abundance [ha-1]
1,0

0,2

0,5

DBH growth
Abundance
0,0 0,0
20 60 100 140 180
DBH [cm]

Figure 9.8
A comparison of the size-dependent pattern in diameter (DBH) growth of the Brazil nut tree and the
relative abundance of individuals in different size categories. The comparison shows that the peak in
DBH growth coincides with low values for tree abundance. Thus, trees tend to rapidly pass through
these stages, and are therefore present at low densities. The size-dependent DBH growth curve is
the same as in Figure 6.6 (lower graph - Pando). The abundance values are from the stable population
structure that is obtained using matrix population models. This model was established for a natural
population of the Brazil nut tree in Pando, Bolivia. Source: Zuidema & Boot (2002).

with high DBH growth rates. Furthermore, the reduction in DBH growth in
trees of over 50 cm DBH, causes a gradual increase in the abundance of trees.
This abundance again drops for trees larger than 120 cm DBH, probably as a
result of an increased mortality risk. This comparison thus shows that the
species-specific pattern of diameter growth may explain the typical population
structure of the Brazil nut tree with low abundance of juvenile trees and
relatively high abundance of adults. In mathematical terms, it can also be shown
that a large decrease in growth at increasing tree size, leads to a simulated
population structure that contains a “hump” of the size classes with slow-
growing trees.
Lastly, the typical population structure of the Brazil nut tree is observed
for many more tropical tree species. The size-dependent pattern in DBH
growth can also be found for many tropical tree species.
 Chapter 10

Conclusions: life history

of the Brazil nut tree

The previous chapters have dealt with the ecology of the Brazil nut tree.
A summary of current knowledge was provided, covering taxonomy,
distribution, reproduction, dispersal, and the ecology of seedlings, adults as
well as the entire population. On the basis of this knowledge base, some
conclusions may be drawn with respect to the life history of the Brazil nut
tree.That is, we can answer questions such as: what type of species is the Brazil
nut tree? Is it a long- or short-lived tree, a slow- or fast-growing tree, a pioneer
or climax species? Answers to these questions are important to determine the
impact of Brazil nut collection on the continued existence of the species and
the future availability of nuts. Furthermore, it is important to evaluate the
potential of enrichment planting of the Brazil nut tree in order to increase
the production capacity. The topics of sustainability of Brazil nut harvest and
forest enrichment with Brazil nut trees will be dealt with in Part C of this
book.
The Brazil nut tree reaches large sizes: trees frequently are over 40 m tall,
and have a diameter at breast height (DBH) of over 2 m (which means a
circumference of over 6.3 m). As a result, Brazil nut trees are often dominant
in the forest structure. The density of Brazil nut trees is highly variable; they
tend to occur in clusters.
The Brazil nut tree is a relatively fast growing species. Seedlings may grow
rapidly in height per year, when experiencing rather high light conditions.
Average height growth of seedlings may be as high as 0.5 m per year, when
water availability is sufficient (see Figure 7.9). In natural populations the
maximum height growth may also attain 0.5 m per year (see Figure 7.7).
Seedlings strongly respond to increased light availability: height growth in the
field increases when seedlings receive more indirect light. Large seedlings of
the Brazil nut tree seem to need high light conditions for continued growth:
80 • Chapter 10

they are (almost) absent in the forest understorey and occur more frequently
in large than in small canopy gaps (see Figure 7.4 and 7.5).
Diameter growth of Brazil nut trees is also high in comparison with other
tropical tree species. The median diameter (DBH) growth rate of Brazil nut
trees in a natural population in Beni, Bolivia amounted to 3.4 mm per year,
whereas most tree species in this area grow at rates between 0.4 and 2.0 mm
per year. Maximum growth rates are also exceptionally high compared to other
tree species.
Brazil nut trees may reach high ages. Age estimates show that it takes over
100 year to reach reproductive size, and that large adult trees may be as old as
350 years. A large (220 cm DBH) Brazil nut tree was estimated to be 440 years,
using 14C dating techniques.
Thus, the Brazil nut tree appears to be a long-lived tree species, that may
attain very high growth rates and requires high light conditions for growth
into the canopy. The Brazil nut tree has been classified as a gap-dependent or
light-demanding species. In the classification of tropical tree species it may
therefore be placed in the intermediate category between pioneer (or: light
demanding) species and climax (or: non-pioneer, shade-tolerant) species.
Pioneer trees are short-lived and typically dominate during the first phase of
succession in canopy gaps or in secondary forests. Their seedlings cannot
survive under low-light conditions. Climax trees enter at later stages in the
succession and are generally long-lived. Their seeds may germinate in the
understorey and seedlings may survive for some years under low light
conditions. An intermediate group is that of the long-lived pioneers (or: late
secondary species). They may germinate and survive in the forest understorey,
but can also be very successful in early stages of forest succession.They strongly
respond to increased light conditions by rapid growth, and require high light
conditions to reach adult size. Thus, they have characteristics of both pioneer
and climax species. On the basis of the available ecological information, the
Brazil nut tree can be classified as a long-lived pioneer species.
 Part C

Management
of the Brazil nut tree
 Chapter 11

Impact of exploitation

11.1 Intensity and consequences

of Brazil nut extraction

In areas where Brazil nuts are collected, often a large part of the seeds are
taken away. In Peru and Brazil over 60% of the total seed production is taken
away by Brazil nut collectors (Ortiz 2002). In two exploited populations in
Bolivia, this was found to be much higher: in two study areas (6 and 12 ha in
size) and three years, an average of 93% of the total seed production was
removed by Brazil nut collectors. Estimates for the entire Brazil nut region of
Bolivia suggest that around 50% of the total seed production is collected. The
large differences between these estimates are most probably caused by
differences in scale: in small populations close to settlements (as in the two
Bolivian study sites) the collection is likely to be more intensive as compared
to a much larger area which encompasses considerable tracks of remote and
inaccessible forest where no collection takes place. Nevertheless, it becomes
clear that in exploited populations of the Brazil nut tree, a substantial to large
part of the seeds are removed.
In order to determine the possible impacts of this intensive seed removal
for populations of the Brazil nut tree, it is necessary to define where in the
life cycle this collection has an effect. Figure 11.1 shows a simplified life cycle
of the Brazil nut tree and indicates which processes in the life cycle are
changed as a result of the seed harvest. In this life cycle, the process of seed
production is changed: a large part of these are collected and thus not available
for dispersal and germination.The number of seeds in the seed class is therefore
reduced.
Knowing where in the life cycle Brazil nut collection has an effect, four
questions can be asked. (1) What is the influence of Brazil nut harvest on the
regeneration of the population, i.e. on the appearance of new seedlings in the
84 • Chapter 11

Influence of Brazil nut collection in the life cycle

Harvested seeds
Seed production

Sd Sdl Juv
Juv Ad1
Ad1 Ad2
Dispersal & Growth & Growth & Growth &
germination survival survival survival

Figure 11.1
The influence of Brazil nut collection in the life cycle of the Brazil nut tree. This figure is similar to
Figure 9.1, but showing the processes in the life cycle (the stippled lines: seed production) that are
changed due to Brazil nut collection. Often a large part of the total seed production is harvested in
exploited populations of the Brazil nut tree. Circles denote the different classes in the population.
Arrows are the processes in the life cycle. Sd = Seeds, Sdl = Seedling; Juv = Juvenile tree; Ad1 = Small
adult tree; Ad2 = Large adult tree.

population? (2) What is the influence of Brazil nut harvest on the population
dynamics of the Brazil nut tree? (3) What is the impact of Brazil nut harvest
on the future availability of seeds (Brazil nuts)? And, (4) what is the impact of
Brazil nut harvest at the level of the ecosystem.These questions will be tackled
in the following four Sections.

11.2 Impact of exploitation

on natural regeneration

The collection of Brazil nuts may have consequences for the regeneration
(i.e. the appearance of new individuals) of the tree in three ways. (1) The
amount of seeds available for germination is strongly reduced, causing less new
seedlings to appear. (2) The seed dispersing activity of the dispersing animals
(agoutis) may change due to the reduced availability of seeds. (3) The density
of seed dispersing animals may decrease as a result of hunting by the Brazil
nut collectors. And, (4) there may be a positive influence of exploitation when
Brazil nut collectors unintentionally disperse seeds when opening the fruits
or transporting the seeds in bags. Below, the importance of each of these
changes for tree regeneration is discussed. Figure 11.2 shows the possible effects
of each of these factors for natural regeneration of the population.
1 • In general, it is logical that the number of germinated seeds will decrease
when the availability of seeds is lower. But whether a 50% reduction in the
number of seeds available results in a 50% reduction of new seedlings in the
population is not clear. This depends on the influence of seeds availability on
the predation and dispersal of seeds. It may also depend on density-dependent
mortality of seedlings: seedling mortality may be higher when many seedlings
 Impact of exploitation • 85

Figure 11.2
Effect of Brazil nut harvest on regeneration
The possible effects of Brazil nut harvesting on
Possible effect on regeneration
natural regeneration of the Brazil nut tree. The
Negative Positive
potential influence of four factors is shown,
(1) Reduction of indicating whether a positive or negative effect
available seeds
for natural regeneration is to be expected.
(2) Change in seed
handling by animals
(3) Hunting of ? ?
seed dispersers
(4) 'Seed dispersal'
by harvesters

of the same species occur within a small area. If, for instance, the predation
(i.e. the consumption by animals) of seeds is higher if seeds occur at a high
density, a lower density of seeds may actually result in a higher proportion of
germinated seeds (see below). Thus, although in general a lower seed
availability can be assumed to result in a reduced number of new seedlings, it
is not clear how strong this reduction will be.
2 • The activity of seed dispersing animals (agoutis) was found to be
dependent on the availability of seeds. In Bolivia, a field experiment was
carried out which Brazil nuts (i.e. the seeds that had been removed from the
fruits) were put around Brazil nut trees at two densities: 340 and 3400 seeds
per tree in groups of 17 seeds (the average number of seeds per fruit). The
density of seeds in the high-density treatment reflects that for an average-
producing tree of which no seeds are harvested. The seed density in the low-
density treatment is 10% of that in the high-density treatment, which resembles
the density in an area where 90% of the Brazil nuts is collected. The number
of seeds eaten and the number of dispersed seeds was determined in both
treatments. It was found that for each seed that is consumed, more seeds were
scatterhoarded in the high density treatment compared to the low density
treatment. This suggests that there is some kind of satiation of the agoutis:
when the density of seeds is very high, agoutis tend to scatterhoard a larger
part of all seeds. They are probably satiated after eating a small fraction of all
seeds available. Thus, in harvested populations of the Brazil nut tree there may
be relatively less seed dispersal and more seed consumption (predation) than
in undisturbed populations. This finding has important consequences for seed
dispersal in areas where Brazil nuts are collected. As Brazil nut collection
greatly reduces the availability of seeds (or in fact fruits), this result suggests
that of the fraction of seeds left by Brazil nut collectors even a smaller
proportion is scatterhoarded and more seeds are consumed. Thus, a negative
impact of this changed seed handling behaviour may be expected.
3 • Hunting of agoutis may result in a reduction of the number of dispersed
Brazil nut seeds. Agoutis are commonly hunted for meat in areas where Brazil
86 • Chapter 11

nuts are collected. In Bolivia, it has been calculated that the agouti that is
dispersing Brazil nut seeds (Dasyprocta var iegata) is among the ten most
hunted mammals by Brazil nut collectors. An estimated 0.8 individuals is killed
by Brazil nut collectors per km 2 per harvest season (Santivañez unpublished
data). Nevertheless, information from Peru suggests that densities of agoutis
in exploited Brazil nut forests remain high in spite of hunting, probably as
large predators of the agoutis disappear from the area due hunting or the
presence of humans (Ortiz 2002). Thus, it is not clear whether densities of
agoutis generally decrease or increase in exploited Brazil nut populations.
In addition to the uncertainty about agouti densities, the effect of a
changes in agouti abundance on the natural regeneration of the Brazil nut tree
is also unclear. If the density of agoutis is greatly reduced a much smaller
amount of Brazil nut fruits will be opened. As the probability that seedlings
emerge from closed fruits is very low, many seeds will finally rot in the fruits
without germinating. On the other hand, the remaining agoutis may have a
larger number of Brazil nut fruits at their disposal, which again may increase
the proportion of seeds they disperse in relation to the seeds they consume.
In case the density of agoutis is increased, their competition for food sources
may be higher. This may again lead to more consumption and less dispersal of
Brazil nuts.
4 • Brazil nut collectors may leave seeds of the Brazil nut tree when opening
the fruits and emptying these in large bags. Also, seeds may drop from their
bags while transporting the seeds in the forest. Although there is no field-
evidence of this ‘seed dispersing activity’ by Brazil nut collectors, it clearly has
a positive effect on the natural tree regeneration.

Which of the above factors is most important, is not clear. It will depend
on the intensity of the harvest, the harvesting and transport technique, the
hunting pressure and the natural density of seed-dispersing animals. In certain
cases the net effect of Brazil nut harvest on the natural tree regeneration may
be negative, and in other cases it may be positive.
Information on the natural regeneration in an exploited population of
the Brazil nut tree is available for two study sites in Bolivia. In these
populations, where approximately 90% of the seeds are removed, a rather large
number of new seedlings were found. In a study site in Beni, 75 new seedlings
(less than 1 year old) were found in an area of 1.5 ha: in a study site in Pando,
51 new seedlings were found in 1.4 ha. When expressed per adult tree in the
study site, 20 and 5 new seedlings appeared per adult tree (per year), in the
two sites respectively. Thus, it appears that in spite of the large reduction in
the amount of available seeds, there is a natural regeneration of the Brazil nut
in these populations. Another indication for this is that no size categories were
 Impact of exploitation • 87

completely missing in these populations, i.e. seedlings and saplings of different

sizes are found in exploited populations.

11.3 Impact of exploitation

on population dynamics

The impact of Brazil nut collection on the population dynamics of the

Brazil nut tree can be assessed using models for population dynamics. Such
models were constructed for two exploited populations of the Brazil nut tree
in Bolivia. The results of these models were shown in Chapter 9 of Part B.
Three results from the population models indicate that the influence of
Brazil nut harvesting on the population dynamics is small. Firstly, it was shown
that both populations where 90% of all seeds from the Brazil nut tree are
harvested, have a stable population size. This implies that, in spite of the
intensive harvest, no reduction in the size of the populations is expected over
a long period of time.
Secondly, the rate of population growth hardly changes when the amount
of seed production is reduced in the model. Thus, the process in the life cycle
that is most affected by Brazil nut harvest, seed production, has little influence
on the population growth rate. Also large changes in seed production have a
very small impact on the population growth rate.
Thirdly, the population model predicts that the future size distribution of
exploited populations will be similar to the present distribution. Thus, seed
removal has little impact on the relative number of individuals over different
size categories.
In short, population models for two exploited populations in Bolivia
indicate that the impact of intensive harvest of Brazil nuts has little impact on
the future development of the population. Clearly, the above results are based
on studies in only two forest areas in Northern Bolivia, and may not be
representative for exploited populations of the Brazil nut tree elsewhere.
Nevertheless, the results from the two studies are rather similar, in spite of the
140 km distance between the sites.

11.4 Impact of exploitation

on future resource availability

The question of the impact of Brazil nut harvest on the future availability
of Brazil nuts is partly answered in the previous Section. It was stated there
that exploited populations do not decrease in size and that distribution of
88 • Chapter 11

individuals over size categories does not change. This implies that the future
abundance of reproductive Brazil nut trees remains more or less the same, and
that the production of Brazil nuts does not change.
Two other study results suggest that the availability of Brazil nuts will be
similar in the future. Firstly, it takes a long time before Brazil nut trees become
reproductive: trees that start to produce fruits are on average 125 year (see
Figure 8.9).This implies that, if the collection of Brazil nuts reduces the natural
regeneration of the population, it would take a long time until this would
result in a lower production of Brazil nuts. For instance, if the natural
regeneration of Brazil nut trees has been reduced since 25 years, this would
mean that a reduction in the abundance of adult trees is expected after 100
yr from now. Thus, if there is any impact of current harvest on the future
availability of Brazil nuts, it will become apparent only after a long period,
most probably only after more than 100 year.
Secondly, it has been found that recruitment does take place in exploited
populations (see Section 11.2): there is input of new seedlings and individuals
of all size categories can be found in exploited populations. Furthermore,
population dynamics models for do not predict shifts in the size distribution
of the population. Thus, it may be expected that current extraction practices
do not lead to future changes in size distribution.
In short, the results of this study suggest that high levels of Brazil nut
extraction may be sustained at least for over 50 years without reducing the
production potential and that there are good prospects for continued
regeneration of exploited populations, which would safeguard productivity for
much longer periods.

11.5 Impact of exploitation at ecosystem level

The collection of Brazil nuts may have consequences for the ecosystem
in two ways. Firstly, a large amount of a very nutritious food source is taken
out of the forest. This may have consequences for those animals that feed on
Brazil nuts. These are mostly those rodents (agoutis) that can open the tough
woody fruit of the Brazil nut tree. It is unclear whether a reduced availability
of Brazil nuts will have consequences for the survival of agoutis. Fruits of the
Brazil nut tree fall during the wet season when the availability of fruits and
seeds is high. But agoutis may store Brazil nuts for consumption during periods
of lower food availability in the dry season. In this way, Brazil nuts may be a
crucial food source when food is scarce. However, how important Brazil nut
storage is for agoutis, remains unclear.
 Impact of exploitation • 89

Table 11.1
Hunting in a forest in Northern Bolivia where Brazil nuts are collected. Shown is the number of
individuals hunted per km 2 of hunting area, during the period of Brazil nut collection (usually 90
days). This value is based on two years of detailed observations in a forest estate in Pando, Bolivia.
The black spider monkey is included in CITES Appendix II for threatened species.

Common name Scientific name Extraction rate

[ind km-2 year-1 ]

Paca Agouti paca 2.15

South American coati Nasua nasua 1.46
White-lipped peccary Tayassu pecari 1.24
Brown capuchin monkey Cebus apella 1.21
Red brocket deer Mazama americana 0.98
Brown agouti Dasyprocta variegata 0.82
Black spider monkey Ateles paniscus 0.82

Source: Santivañez unpublished data.

A second and probably more important ecological effect of Brazil nut

exploitation are the hunting practices associated with the periods of nut
collection. In the Brazil nut region in Bolivia and Peru large monkeys are rare
or have disappeared, as a result of hunting. A detailed study on hunting
practices during two years has been carried out in a Brazil nut collection area
in Northern Bolivia (Santivañez unpublished data). Hunting in the Bolivian
study area is done to obtain extra protein sources, but is clearly not sufficient
to cover the daily required amount of proteins. It was calculated that Brazil
nut collectors consume only 11.5 grams of bush meat per day during the
harvest period. The most hunted mammals are mentioned in Table 11.1.
The hunting pressure during periods of Brazil nut collection was found
to be higher than that during logging operations. Hunting of the species in
Table 11.1 (as well as some other less hunted species) is most probably not
sustainable, indicating that the populations of these species are declining and
the species will become absent in the area. It has been noted that the forest
around old centros - the isolated houses in the forest where Brazil nut
collectors live during the harvest period - contains much less mammals
compared to areas around new centros.
Many of the hunted mammals and bird species are important consumers
and dispersers of seeds and in this sense of great importance for the
regeneration of many tree species. A severe reduction in the abundance of
these animal species certainly has an impact on the dispersal of sees and the
regeneration of trees in the forest. However, it is completely uncertain how
and to what extent this may finally influence the structure, composition and
functioning of the entire forest ecosystem. Research results from Peru indicate
90 • Chapter 11

that the level of biodiversity in forests exploited for Brazil nuts (where no
logging takes place) is similar to that of undisturbed forests. Diversity of
mammals and trees were not influences by Brazil nut collection (Ortiz, 2002).
 Chapter 12

Enrichment planting
and plantations

12.1 Ways to increase Brazil nut production

In view of the considerable income generated by the collection of Brazil

nuts, there is interest in techniques to increase the production potential of
Brazil nuts. An increase in production can be achieved in two ways: (1) by
increasing the nut production of existing Brazil nut trees, or (2) by increasing
the number or density of Brazil nut trees.
1 • An increase in production of existing trees can be achieved by eliminating
certain factors that reduce reproduction. An important measure is the removal
of lianas from Brazil nut trees. In Section 5.4 it was shown that lianas reduce
the production of Brazil nuts in natural populations. When Brazil nut trees are
infested by large amounts of lianas, their seed production is considerably
reduced. The problem of liana infestation can be eliminated relatively easily
by cutting the stems of those lianas climbing into Brazil nut trees. This is an
activity that does not require much time and can probably be combined with
the collection of Brazil nuts. Large-scale trials with liana cutting have been
carried out Beni, Bolivia.
2 • The density and number of Brazil nut trees can be increased by
enrichment planting, i.e. planting seedlings of the species. Several enrichment
planting trials have been carried out in primary and secondary forests and in
agricultural and pasture areas. Results of these trials are presented in Sections
12.3 and 12.4. Another way to increase the number of Brazil nut tree is the
establishment of plantations. Some results of Brazil nut tree plantations are
provided in Section 12.5. As enrichment and plantation initiatives require the
availability of large numbers of seedlings, the following section provides some
information on controlled germination of Brazil nuts.
92 • Chapter 12

12.2 Germination of Brazil nut seeds

under controlled conditions

Brazil nut tree has been frequently used for planting in agricultural fields,
enrichment planting in secondary forests and to establish plantations. Although
this has predominantly been carried out in experimental trials and at a
relatively small scale, planting programs at a larger scale are now underway.
For all these activities seedlings of the Brazil nut are needed, which in most
cases will have to be obtained from germinated seeds under controlled
conditions. In this Section some methods for controlled germination of Brazil
nut seeds are presented and compared. Table 12.1 provides an overview of
germination success of Brazil nuts using different seed treatments.

The results of two of the studies mentioned in Table 12.1 (those by Kainer
et al. 1990 a&b) reveal that seeds from different parent trees differ largely in
the percentage germination and time to germination. For instance, after 8
months the germination success (% of germinated seeds) varied between 30%
and 70%, depending on the parent tree. These differences were not related to
the differences in size or weight of the seeds. Apparently, other factors play an
important role in determining germination time and success.
Moist storage of seeds has a strongly positive influence on the germination
time and success of Brazil nut seeds. In a germination experiment, part of the
Brazil nut seeds collected in natural populations were sown shortly after
collection and another part was stored for 5.5 months under moist conditions.
As becomes clear from the data in Table 12.1, germination of stored seeds

Table 12.1
Germination of seeds of the Brazil nut tree under different seed treatments. The time to the
germination of the first seed (‘Start’), the percentage of germinated seeds and the period after which
this is achieved are shown. The most important difference in seed treatment are whether the seed
coat is removed, whether fungicide is applied, and whether seeds are stored.

Treatment of seeds before germination Start Germinated after

[days] [%] [months]

1 “Natural conditions” (leave seed coat, no treatments) 30 12

2 Scarify seed coat at appendices, leave seed coat 41 18
3 Soak seeds 24 h, remove seed coat, fungicide 30 78 5
4 Soak seeds 24 h, remove seed coat, no fungicide,
in sterilised coarse sand 60 55 9
5 Most storage of seeds for 5.5 months, soak seeds 24 h,
remove seed coat, no fungicide, in sterilised coarse sand 14 75 3

Sources. 1: Sanchez (1973); 2: Müller (1981); 3: Müller (1982); 4: Kainer et al. (1999a); 5: Kainer et al.
(1999b).
 Enrichment planting and plantations • 93

started much earlier than that of non-stored seeds, and a larger part of the
stored seeds germinated in a much shorter period. Therefore, it seems that a
certain period of time under moist conditions is necessary for rapid and
successful germination. For efficient and continued production of Brazil nut
seedlings, it is therefore recommended to store sufficient quantities of Brazil
nuts under moist conditions. For the experiment mentioned in Table 12.1,
seeds were stored in a circular cement tank, on top of a layer of 25 cm washed
fine sand. The seeds were covered with a second 25-cm layer of fine sand. The
tank was equipped with a drainage pipe at the bottom to discharge excess
water. The tank was watered until water dripped out from the drainage pipe.
Watering was regularly in order to maintain the tank contents humid (in the
dry season watering was done once per week).

12.3 Seedling survival and growth

in enrichment experiments

Enrichment planting of the Brazil nut tree has been carried out in
agricultural fields, pasture areas, and in primary and secondary forests. In
secondary forests, Brazil nut seedlings have been planted in strips where the
original vegetation was cleared. The results of that study are presented in the
next Section. Information on enrichment planting in primary forest,
agricultural fields and pastures is provided in this Section. The results of three
enrichment studies are presented.
Brazil nut seedlings have been planted in gaps in primary forest in two
experiments carried out in Acre, Brazil. In one of these experiments (Study
1), small seedlings (7 cm height) were planted in small gaps; the other
experiment (Study 2) was carried out with larger seedlings (20 cm height)
and in large logging gaps or skidding trails.The survival and growth of planted
seedlings in the two experiments is compared in Figure 12.1. There is a large
difference in height growth between the experiments: smaller seedlings planted
in small gaps (Study 1) had a much lower growth rate than the larger seedlings
in larger gaps (Study 2). This can be explained by the differences in seedling
size and light availability between the experiments. Larger seedlings may attain
larger height growth rates, as they have a larger leaf area to capture light. And,
higher light availability may result in more photosynthesis and thus in higher
growth rates. As a result, the large seedlings in large gaps were able to attain a
height of almost 3 metres within 5 years.
Another difference in the results in Figure 12.1 is the lower survival rate
in Study 2. The high mortality in the first year of Study 2 was mostly due to
rodents that kill the seedlings to eat the nutritious cotyledon (seed reserve).
94 • Chapter 12

Seedling survival in gaps Seedling growth in gaps

100% 3
Study 1
Surviving plants [%]

Study 2

Plant height [m]

2
50%
1
Study 1
Study 2
0% 0
0 1 2 3 4 5 0 1 2 3 4 5
Time after planting [yr] Time after planting [yr]

Figure 12.1
Survival and growth of Brazil nut seedlings planted in forest gaps and skidding roads in primary
forests in Acre, Brazil. Survival is the percentage surviving seedlings at a given time after planting.
Study 1 was carried out using 125 seedlings in 5 forest sites over a period of 2 years. Small seedlings
were used (7 cm height at the start of the experiment) and seedlings were planted in small gaps (15-
50m 2 ). Study 2 was carried out with an initial number of 162 larger seedlings (20 cm height), which
were planted in large logging gaps (100-800 m 2 ) and on skidding roads. Sources: Kainer et al. (1999)
(Study 1) and d’Oliveira (2000) (Study 2).

At the site of Study 1, the abundance of rodents was probably much lower due
to intensive hunting. From both studies it becomes clear that less than a third
of the planted seedlings survives up to two years. This high mortality rate is
important to keep in mind when planning enrichment activities in gaps.
Enrichment planting of Brazil nut seedlings in agricultural fields has been
carried out in Acre, Brazil (Study 1 as above) and in Beni, Bolivia (Study 3).
The results of these studies are shown in Figure 12.2. Again, planted seedlings
in Study 1 were much shorter (7 cm) than those in the other study (Study 3;
23 cm). The initial size difference probably explains the difference in growth
rate and mortality between the studies. Small seedlings generally experience
higher mortality. In fact, mortality may be much higher than found in Study
1, in case the abundance of rodents is high in the area: in a trial experiment
with small seedlings planted in secondary forests in Bolivia, the vast majority
of seedlings died after several months due to the consumption of the cotyledon
by rodents.Very small seedlings such as used in Study 1 have intrinsically lower
growth rates than somewhat larger seedlings.
Brazil nut seedlings planted in pasture areas survived well but grew poorly.
Figure 12.3 shows the results of one study in Acre, Brazil, where small seedlings
were planted with fencing in pastures with grazing animals and without
fencing in ungrazed areas. When compared to forest gaps and agricultural
fields, height growth was low, but survival was high.
Above, the different enrichment sites for the Brazil nut tree have been
compared in biological sense. Apart from this, it is also important to consider
the social suitability of the different enrichment sites. In the above mentioned
Study 1, the suitability of three planting sites (forest gaps, pasture areas and
 Enrichment planting and plantations • 95

Seedling survival in fields Seedling growth in fields

100%
Surviving plants [%] 4

Plant height [m]

3

50% 2

1
Study 1 Study 1
Study 3 Study 3
0% 0
0 1 2 3 4 5 0 1 2 3 4 5
Time after planting [yr] Time after planting [yr]

Figure 12.2
Survival and growth of Brazil nut seedlings planted in agricultural fields in Acre, Brazil (Study 1) and
Beni, Bolivia (Study 3). Survival is the percentage surviving seedlings at a given time after planting.
Study 1 was carried out using 125 small seedlings (7 cm height) in 5 agricultural fields over a period
of 2 years. Study 3 was carried out with an initial number of 32 larger seedlings (25 cm height), which
were planted in 4 agricultural fields. Sources: Kainer et al. (1999) (Study 1) and Peña-Claros et al.
(2002) (Study 3).

agricultural fields) was determined by including the opinions of local people

that assisted in the experiments. Forest gaps appeared to be suitable sites from
a social point of view, as the required input of labour is low. From a forest-
biological viewpoint, these sites were the least suitable as seedling growth was
low due to limited availability of light.The second enrichment site, agricultural
fields, was found to be the most suitable environment from both a social and
biological point of view: growth and survival were high in these areas, and
enrichment activities (planting, weeding) can be combined with the activities
required for the cultivation of crops. The third enrichment site, pasture areas
was least suitable from a social point of view: planted seedlings should be
protected from animals by fences, fences should be checked frequently and
weeding should be carried out frequently. Nevertheless, the high survival rate

Seedling survival in pasture Seedling growth in pasture

100% 0,4
Surviving plants [%]

Plant height [m]

50% 0,2

Study 1 Study 1
0% 0
0 1 2 0 1 2
Time after planting [yr] Time after planting [yr]

Figure 12.3
Survival and growth of Brazil nut seedlings planted in pasture areas in Acre, Brazil. Survival is the
percentage surviving seedlings at a given time after planting. The study (Study 1) was carried out
using 125 small seedlings (7 cm height) in 5 pasture areas over a period of 2 years. Source: Kainer et
al. (1999).
96 • Chapter 12

and good growth of planted seedlings in pasture areas suggest that this site is
suitable for enrichment in the biological sense.
In conclusion, when combining the social and biological suitability of the
three enrichment sites, agricultural fields seem to be the best option for
successful enrichment of the Brazil nut tree. Another good option for
enrichment planting which was not included in the above studies, is strip
planting in secondary forests: this method is presented in the next Section.
Some important practical recommendations can be derived from the
results of the three presented enrichment studies. Firstly, it is recommended
to plant seedlings of at least 20-25 cm height, as these seedlings are less likely
to be killed by rodents that consume the cotyledon (the seed reserve in the
stem). The cotyledon in seedlings of that size is protected by the woody stem.
Secondly, it is recommended to clear the vegetation around planted
seedlings.This was done in the three discussed experiments: in some cases only
once after planting, and in other cases at regular intervals. The removal of the
existing vegetation, decreases the competition of planted seedlings with
neighbouring plants, and thus increases their growth rate and their survival
probability.

12.4 Strip planting as an enrichment method

A method for enrichment planting that is frequently used in forestry

practices is strip or line planting. This implies that seedlings of the desired
species are planted in strips in which the existing vegetation has been removed
to increase the light availability and reduce competition for light, water and
nutrients. As the canopy in secondary forests is lower than that in primary
forests, relatively high light levels may be achieved by cutting rather narrow
strips. Furthermore, secondary forests are suitable for this type of enrichment
planting as the removal of the vegetation is less time consuming than in
primary forests, where large trees have to be logged to create strips. In this
Section the results of an strip cutting experiment with seedlings of the Brazil
nut tree are discussed.
Figure 12.4 provides a schematic representation of a cleared strip with a
Brazil nut seedling. This Figure also shows different measures that can be used
to determine the angle of canopy openness of the planted seedlings. This
technique was developed to assess the light level of planted seedlings in strips
without the use of sophisticated equipment (hemispherical lenses, light
sensors). The angle of canopy openness is calculated using the height of the
tallest tree in the vegetation bordering the strip, the strip width and the height
of the planted seedling. The angle is calculated as β = 180 - (α1+α2), where α1
 Enrichment planting and plantations • 97

Figure 12.4
Strip planting in secondary forests
vegetation Schematic representation of enrichment
planting of the Brazil nut tree in strips in
vegetation secondary forest. Seedlings of the Brazil nut
tree are planted in strips (lines) of a certain
width, in which the existing vegetation has
β been cleared. The arrows indicate different
α1 α2 height measures (x = seedling height; y = height
of tallest tree in vegetation) and angles ( α =
y1
Brazil nut x y2 angle from seedling to highest point in the
seedling adjacent vegetation; β = angle of open canopy
above seedling). Adapted from: Peña-Claros et

line al. (2002).

width

and α 2 are calculated as the cotangent of the height difference between the
Ber tholletia plant (x) and the tallest tree next to it (y1 and y2, respectively)
divided by half of the line width.
The calculated angle of canopy openness is closely related to the canopy
openness when this is measured using hemispherical photographs. Figure 12.5
shows the relation between these two variables. Roughly, the canopy openness
expressed in percentage opening increases with 1% when the calculated angle
of openness increases by 5 degrees. The fact that there is a good relation
between these variables implies that using simple measurements (plant height
and strip width), a reasonable estimate of the canopy openness can be obtained.
The strip planting experiment was carried out using strips of different
width: 2, 4 and 6 m wide. For comparison seedlings were also planted in the
forest understorey. The vegetation around the planted seedlings was removed

Canopy openness and angle Figure 12.5

Relation between the measured canopy
Measured canopy openness [%]

50 openness (along y-axis) and the angle of canopy

openness ( β , along x-axis) for seedlings of the
Brazil nut tree planted in strips in secondary
forests. The canopy openness percentages are

25 determined using hemispherical photographs;

the angle values are calculated using the
equation mentioned in the text and different
height measurements indicated in Figure 2.4.
The correlation between the two measures for
0
light availability is good: 45% of the variation
0 50 100 150 200
in the measured canopy openness is explained
Calculated openness angle [β]
by the calculated angles. The regression
equation is: y = 0.20x +12, indicating that an
increase of 5 degrees in the angle of canopy
openness results in an increase of 1% in
measured canopy openness. Source: Peña-Claros
et al. (2002).
98 • Chapter 12

Canopy openness in strips Figure 12.6

Canopy openness in strips in secondary forests
Canopy openness [%]

40
in Bolivia, where seedlings of the Brazil nut tree
30 have been planted. US = understorey treatment,
i.e. no vegetation was cleared. Canopy
20 openness was determined using hemispherical

10 photographs at 1 m height shortly after the

experiment started. Bars denote the average
0 value, vertical lines are the standard deviation.
US 2 4 6 Canopy openness is statistically different in all
Strip width [m] treatments except for the 2 and 4 m wide strips,
Source: Peña-Claros et al. (2002).

regularly during the period of the experiment.The planted Brazil nut seedlings
had an average height of 25 cm.
The initial light availability differed among treatments, as is shown in
Figure 12.6. Canopy openness varied from 15% in the understorey treatment
to almost 40% in the 6-m wide strips. This was the situation at the start of the
experiment, just after the strips had been cut. As the vegetation next to the
strips gradually grew, the light availability decreased in time. In the strips of 2
and 4 meters the decrease in light availability for the planted seedlings was
45-50%; in the 6-m treatment light level decreased by 30%.
Survival of the planted Brazil nut seedlings was high: 87-98% of the
seedlings survived each year. Seedling survival was lower for the understorey
treatment (87%) than in the strip treatments (96-98%), as is shown in Figure
12.7 (left graph). Large differences among the treatments were found for height
growth: seedlings in the 6-m wide strips grew almost 6 times faster than in
the forest understorey (83 compared to 15 cm per year). The right graph in
Figure 12.7 shows the height growth rates during two periods: the first and
the fourth year after the seedlings were planted. The graph shows that height
growth of seedlings in the 2- and 4-m wide strips is much lower in the fourth
than in the first year. Seedling growth in the 6-m wide strips, on the other
hand, remains the same.This reason for this is that narrow strips gradually close
because of lateral in-growth of the vegetation that borders the strips.
The results of this experiment show that the Brazil nut tree is a suitable
species for enrichment planting in secondary forests. Planted seedlings (of 25
cm height) have a high survival probability and attain high growth rates. This
study also shows that in wide strips a high light availability is maintained and
consequently high growth and survival rates are achieved. It is therefore
recommended to establish wide strips for the enrichment planting of the Brazil
nut tree. Strips that result in a canopy openness of 25-40% (or an angle of
canopy openness of 60-90 degrees) are most suitable. Wider strips are costly
without resulting in much higher growth rates, and more narrow strips require
intensive maintenance to keep the strips open.
 Enrichment planting and plantations • 99

Survival in strips Growth in strips

Height growth [cm yr-1]

100
125 First year
Survival [% yr-1]

75 Fourth year
100
50 75
50
25
25
0 0
US 2 4 6 US 2 4 6
Strip width [m] Strip width [m]
Figure 12.7
Survival and growth of planted seedlings of the Brazil nut tree in strips in secondary forests in Bolivia.
US = understorey treatment, i.e. no vegetation was cleared. Survival rates are calculated over 4 years
of measurements, growth is the annual growth during the first and the fourth year after the seedlings
had been planted. 90-100 seedlings were included in the study. Source: Peña-Claros et al. (2002).

12.5 Plantations of Brazil nut trees

Several plantations of Brazil nut trees have been established (most in

Brazil) and a substantial amount of research has been carried out to facilitate
the establishment of plantations. Brazil nut plantations may contain high
densities of trees: up to 100 trees per ha. Brazil nut trees in plantations are
reported to produce fruits within 10 yr, and when grafted even within 4 yr.
Normal production can be expected after approximately 25 years for normal
trees and approximately 11 years for grafted trees (Mori & Prance 1990). Brazil
nut seedlings should be planted at 10-20 m distance from each other, and may
be combined with other crops. They should preferably be located close to
native forests, in order to ensure pollination to take place.
In spite of the research focus on plantations, the amount of large-scale
commercial plantations is low and the vast majority (over 99%) of the world
Brazil nut production is collected from natural populations. Most Brazil nut
plantations are small and have been established for experimental purpose in
stead of commercial nut production. Nevertheless, some large plantations exist:
Fazenda Aruanã in Amazonas (Brazil) is probably the largest with 318,660
planted Brazil nut trees on 3,341 hectares in 1990 (Mori 1992). Brazil nut
trees in this plantation are grafted using a high yielding variety. In 1999 the
oldest trees were producing around 100 fruits per tree, thus yielding an average
of 2500 fruits or 500 kg in-shell nuts per hectare. Establishment of large
commercial Brazil nut plantations, such as the one in Amazonas, is difficult
due to the high initial costs, the long time to nut production and the low
return in general.
The results of two plantation experiments that lasted for 10 years in
Amazonas and Rondônia, Brazil, are presented below. The two experiments
100 • Chapter 12

DBH in plantation - Amazonas DBH in plantation - Rondonia

Average Monoculture
30 30
Maximum Mixed cropping
DBH [cm]

20 20

10 10

0 0
0 5 10 0 5 10
Time since planting [yr] Time since planting [yr]

Figure 12.8
Diameter growth of Brazil nut trees in a plantation in two experimental plantations in Brazil. The
left graph shows data from a plantation in Amazonas where trees were planted at high density (3 m
distance). Open circles in the left graph represent the average size of the plants, vertical lines the
standard deviation (measure of variation in size), and the black dots are the maximum DBH values
reached. The right graph shows data from a plantation in Rondônia where trees were planted in
monoculture (circles) and mixed culture with crops and small trees (triangles), both with at 12 m
distance between Brazil nut tree. Sources: Fernandes & Alencar (1993) (left graph) and Vieira et al.
(1998) (right graph).

differ in the space between the planted trees (at 3×3 m intervals in Amazonas
and 12 ×12 m in Rondônia) and in the type of cultivation (a monoculture of
Brazil nut trees in Amazonas and both mono- and mixed cultures in
Rondônia). Mixed cropping was done in combination with banana plants and
small Theobroma trees that yield fruits (called cupuazu).
Diameter growth of Brazil nut trees in the two experimental plantations
is shown in Figure 12.8. From the Amazonas graph it becomes clear that the
rate of DBH growth gradually decreases in time. This is due to the narrow
spacing of Brazil nut trees: when trees are planted at a distance of 3 m they
strongly compete for space as they grow. Some trees may outcompete others
and grow larger. This is also shown by the high maximum diameter after 10
years, which is 50% larger than average. In the Rondônia plantation diameter
growth rates are much higher: trees reach a diameter of over 20 cm in 10 years.
Furthermore, there is no decrease in diameter growth over time. The two
planting systems (monoculture or mixed with other crops and small trees) yield
trees of very similar size. Apparently, the presence of other crops is not limiting
the growth of Brazil nut trees.
Height growth of planted Brazil nut trees was also higher in the spaced
Rondônia plantation than in the dense Amazonas plantation, as can be seen
in Figure 12.9. Height development in agricultural fields in Bolivia was very
similar to that in the Rondônia plantation. Planting other crops and trees in
combination with Brazil nut trees did not influence the height growth of Brazil
nut trees, as can be concluded from the Rondônia experiment.
A comparison of rates of annual diameter growth is shown in Figure
12.10. This reveals that annual increments may be very high: over 4-5 cm per
 Enrichment planting and plantations • 101

Plantation height growth - AM Plantation height growth - RO

30 30
Plantation: average Monoculture
Plantation: maximum Mixed cropping
Enrichment
Height [m]

20 20

10 10

0 0
0 5 10 0 5 10
Time since planting [yr] Time since planting [yr]

Figure 12.9
Height growth of Brazil nut trees in a plantation in two experimental plantations in Brazil. The left
graph shows data from a plantation in Amazonas where trees were planted at high density (3 m
distance). Open circles in the left graph represent the average size of the plants, vertical lines the
standard deviation (measure of variation in size), and the black dots are the maximum DBH values
reached. Tree heights are compared with those found in enrichment planting of Brazil nut trees in
agricultural fields in Bolivia (triangles, see Section 12.3). The right graph shows data from a plantation
in Rondônia where trees were planted in monoculture (circles) and mixed culture with crops and small
trees (triangles), both with at 12 m distance between Brazil nut tree. Sources: Fernandes & Alencar
(1993) (plantation, left graph), Peña-Claros et al. (2002) (enrichment, left graph) and Vieira et al.
(1998) (right graph).

year in the case of the Rondônia plantation. The comparison also shows that
there exists a large variation in DBH growth among the different experimental
plantations. The Rondônia data seem to show that larger tree have a lower
DBH growth rate.
From the above results it becomes clear that spacing of trees in plantations
is important for good growth of Brazil nut trees: spacing of at least 10 m
between trees is often recommended, and used in large-scale plantations. It
also is shown that intercropping with other species does not affect the growth
of Brazil nut trees. The advantage of mixed cropping is that other products
can be obtained from the plantation after shorter periods of time, before the
Brazil nut trees start to produce.

Figure 12.10
DBH growth in plantations
Comparison of diameter (DBH) growth rates of
Rondonia
DBH growth [cm yr-1]

6 Beni Brazil nut trees in three plantations. Data are

Amazon from plantations in Amazonas, Brazil, where
4 trees were at 3 m distance (circles); Rondônia ,
Brazil, where trees were planted at 12 m
2 distance (triangles) and Beni, Bolivia, where
trees were planted at intermediate but varying
0
distance (square, with vertical lines being the
0 5 10 15 20 25 30
standard deviation). Sources: Fernandes &
DBH [cm]
Alencar (1993) (Amazonas), Vieira et al. (1998)
(Rondônia) and Zuidema, unpublished data
(Beni).
 Chapter 13

Conclusions:
sustainable exploitation
and successful enrichment

In the previous two chapters, two topics related to the management of

the Brazil nut tree have been covered: (1) the sustainability of Brazil nut
harvest, and (2) the increase of Brazil nut production.
1 • In Chapter 11 the question was asked whether harvesting Brazil nuts can
be considered to be sustainable. That is, to what extent the collection of Brazil
nuts has a negative impact on the regeneration of the Brazil nut tree, on its
population dynamics and on the forest ecosystem. It was shown that little
impact of nut collection is expected for the natural regeneration of the tree
and the population dynamics. In exploited populations, new seedlings occur,
and population models suggest that the size of exploited populations remains
stable and that there will be a continuous recruitment in the population. At
the ecosystem level, an important indirect effect of Brazil nut harvest is the
hunting pressure on mammals (mostly rodents, monkeys and ungulates). As a
result of hunting, certain mammals are strongly reduced or absent in forests
where Brazil nuts are collected.
Furthermore, the question is raised whether Brazil nut collection can be
sustained in the future. This depends on the impact of current harvesting on
the future availability of nuts. Population models demonstrated that the size
of harvested populations will remain rather constant in time, and that the
proportion of reproductive trees will not change. This implies that no change
in the production of nuts is expected. Moreover, in case nut collection would
lead to reduced regeneration, the production of nuts would only decrease after
more than 50 years, as it takes a long time before Brazil nut trees become
reproductive. Thus, the continued production of Brazil nuts in exploited
populations is safeguarded for a long period.
As a consequence of the above, there are very good prospects for
certification of Brazil nuts as a environmentally friendly product that can is
harvested in a sustainable way. As mentioned in Section 2.2, organisations in
104 • Chapter 13

Bolivia, Brazil and Peru are preparing norms for certification of Brazil nuts.
(CFV 2001)
2 • In Chapter 12 different ways to increase the production of Brazil nuts
were evaluated. It was shown that some increase in productivity can be
obtained by removing lianas from reproductive Brazil nut trees. Substantial
increases in Brazil nut production can be obtained by planting Brazil nut trees
at small scales in existing vegetations (enrichment planting) 0r at a larger scale
in plantations. Enrichment planting was shown to be most successful in
agricultural fields and in strips in secondary forests. Growth and survival of
planted Brazil nut seedlings was high in these areas, and these types of
vegetations are considered to be suitable by farmers. Both types of vegetation
are usually located close to farmer’s houses or settlements. This is convenient
for planting and maintenance of planted seedlings, and for harvesting Brazil
nuts once the trees start to produce fruits. Strips in secondary forests in which
the existing vegetation is removed should not be too narrow as they may close
due to lateral ingrowth of the vegetation. Strips that have a canopy openness
of 25-40% (or an angle of canopy openness of 60-90 degrees) are expected to
be most effective.
Large-scale plantations may yield slarge amount of Brazil nuts per hectare.
However, high initial costs and long time to return make the establishment of
such large commercial plantations rather difficult. Alternatively, small-scale
plantations (actually similar to enrichment planting in agricultural fields) may
be established close to farmer’s houses. Mixed cropping with small fruit trees
is a good option for small-scale plantations.
In conclusion, Brazil nuts may be harvested without putting at risk the
continued existence of the species and the future availability of the product.
Planting of Brazil nut trees in agricultural fields, secondary forests or
plantations has proven to be a successful way to increase Brazil nut production.
 Bibliography
The bibliography is divided in three parts: (1) an extensive list of published
papers and unpublished reports on the ecology and management of the Brazil
nut tree; (2) other references used in this book which are not related to the
Brazil nut tree; and (3) some addresses of internet sites with information on
the Brazil nut tree or organisations where studies on the Brazil nut tree have
been carried out. For those publications on the Brazil nut tree that are available
on the Internet, the Internet addresses are provided in parentheses at the end
of the reference.
Publications marked with * can be obtained without costs by sending a
request to promab@bio.uu.nl. Some of these publications are available at
www.promab.org.

On the Brazil nut tree

Almeida, C.P. 1963. Castanha do Pará: Sua exportacao e importancia na economia Amazonica.
Estudios Brasileiros 19:1-86.
Assies, W. 1997a. Going nuts for the rainforest. Non-timber forest products, forest conservation
and sustainability in Amazonia. Thela, Amsterdam. 96 pp.*
Assies, W. 1997b. The extraction of Non-timber Forest Products as a conservation strategy in
Amazonia: European Review of Latin American Studies 62: 33-54.*
Assies, W. 1998. Muchas cascaras y pocas nueces: extraccion y desarrollo sustantable en la
Amazonia. Debate Agrario 26: 41-57.*
Baider, C. 2001 . Demografia e Ecologia de Dispersão de Frutos de Bertholletia excelsa
Humb. & Bonpl. (Lecythidaceae) em Castanhais Silvestres da Amazônia Oriental. PhD
Thesis University of Sao Paolo, Brazil.
Beekma, J., A. Zonta, and B. Keijzer. 1996. Base ambiental para el desarollo del departamento
de Pando y la Provincia de Vaca Diez. SNV Bolivia Cuadernos de trabajo 3, La Paz,
Bolivia.
Bojanic, A. 2001. Balance is beautiful: Assessing sustainable development in the rainforest of the
Bolivian Amazon. PhD thesis Utrecht University, Utrecht, the Netherlands.*
Boom, B.M. 1987. Ethnobotany of the Chacobo Indians, Beni, Bolivia. Advances in Economic
Botany 4:1-68.
Boot, R.G.A., and R.E. Gullison. 1995. Approaches to developing sustainable extraction
systems for tropical forest products. Ecological Applications 5:896-903.*
Bouwman, M., and S. van Dijk. 1999. Removal and fate of Brazil nut seeds (Bertholletia
excelsa) Exposed in two contrasting seed densities in a Bolivian moist forest. MSc Thesis,
Utrecht University, Utrecht. 12 pp.
106 • Bibliography

Broekhoven, G. 1996. Non-timber forest products. Ecological abd economic aspects of

exploitation in Colombia, Ecuador and Bolivia. IUCN, Gland and Cambridge. 125 pp.*
Buckley, D.P., D.M. O’Malley,V. Apsit, G.T. Prance, and K.S. Bawa. 1988. Genetics of Brazil nut
(Bertholletia excelsa Humb. & Bonpl.: Lecythidaceae). 1. Genetic variation in natural
populations. Theoretical and Applied Genetics 76:923-928.
Camargo, P.B.D., R.P. Salomao, S. Trumbore, and L.A. Martinelli. 1994. How old are large
Brazil-nut trees (Bertholletia excelsa) in the Amazon? Scientia Agricola 51:389-391.
CFV. 2001. Estandares para la certificacion forestal de castaña (Bertholletia excelsa). Quinto
borrador. CFV, Santa Cruz, Bolivia.
(bolfor.chemonics.net/bolfor/cfv/castaña.htm#Contenido)
Clay, J.W. 1995. An overview of harvesting, forest processing and transport of non-wood forest
products. FAO Report of the international expert consultation on non-wood forest
products. Non-wood forest products 3. FAO, Rome.
(www.fao.org/DOCREP/V7540e/V7540e00.htm).
Clay, J.W. 1997. Brazil nuts. The use of a keystone species for conservation and development.
Pages 246-282 in C.H. Freese, editor. Harvesting wild species - Implications for
Biodiversity and Conservation. John Hopkins University Press, Baltimore.
DHV. 1993. Estudios agro-ecologicos, forestales y socio-economicos en la region de la castaña
de la Amazonia Boliviana. Forest Resource Inventory. Banco Mundial / Gobierno de
Holanda, Amersfoort.
Dijkman, W., Stoian, D., Henkemans, A.B., Assies, W. and Boot, R. 1998. Temporal and spatial
dynamics in the extraction of non-timber forest products in the Northern Bolivian
Amazon. In: Tropenbos (ed.) Seminar proceedings ‘Research in tropical rain forests: its
challenges for the future’, 25-26 November 1997. The Tropenbos Foundation,
Wageningen, the Netherlands, pp. 149-158.*
Diniz, T.D.A.S., and T.X. Bastos. 1974. Contribucao ao conhecimento do clima tipico da
castanha do Brasil. Boletin Tecnico IPEAN 64:59-71.
Dominguez, J. 1994. Precios y costos en la produccion de castaña (Bertholletia excelsa, Hump
& Bonpl.) en Peru y Bolivia: su importancia para la conservacion del bosque tropical.
Revista Forestal del Peru 21:23-38.
Fernandez, N.P., and J.C. Alencar. 1993. Desenvolvimento de arvores nativas em ensaios de
especies 4. Castanha-do-Brasil (Bertholletia excelsa) H.B.K.), dez anos apos o plantio.
Acta Amazonica 23:191-198.
Fifer, J.V. 1970. The empire builders: a history of the Bolivian rubber boom and the rise of the
house of Suarez. Journal of Latin American Studies 2:147-173.
Goubitz, S. and T. Nijkamp. 1996. Reproductive traits of the Brazil nut tree: Bertholletia
excelsa.. A study contributing to a sustainable forest exploitation system. MSc thesis
Utrecht University, Utrecht. 29 pp.
Hayashida-Oliver,Y., R.G.A. Boot, and L. Poorter. 2001. Influencia de la disponibilidad de
agua y luz en el crecimiento y la morfologia de plantines de Swietenia macrophylla,
Cedrela odorata y Bertholletia excelsa. Ecología en Bolivia 35: 51-60.*
 Bibliography • 107

Homma, A.K.O. 1992. The dynamics of extraction in Amazonia: a historical perspective.

Advances in Economic Botany 9:23-32.
Homma, A.K.O. 1996. Modernisation, and technological dualism in the extractive economy in
Amazonia. Pages 59-81 in M. Ruiz-Peres, and J.E.M. Arnold, editors. Current issues in
non-timber forest products research. CIFOR, Bogor.
Kainer, K.A., and M.L. Duryea. 1992. Tapping women’s knowledge: plant resource use in
extractive reserves, Acre, Brazil. Economic Botany 46:408-425.
Kainer, K.A., M. Matos Malavasi, M.L. Duryea, and A. Rodrigues da Silva. 1999a. Brazil nut
(Bertholletia excelsa) seed characteristics, preimbition and germination. Seed Science and
Technology 27:731-745.
Kainer, K.A., M.L. Duryea, M. Matos Malavasi, A. Rodrigues da Silva, and J. Harrison. 1999b.
Moist storage of Brazil nut seeds for improved germination and nursery management.
Forest Ecology and Management 116:207-217. (abstract available at
www.elsevier.com/locate/foreco)
Kainer, K.A., M.L. Duryea, N. Costa de Macedo, and K. Williams. 1998. Brazil nut seedling
establishment and autoecology in extractive reserves of Acre, Brazil. Ecological
Applications 8:397-410.
Kitamura, P.C., and C.H. Muller. 1984. Castanhais nativos de Maraba-PA: Fatores de
depredacao e bases para a sua preservacao. EMRRAPA/CPATU Documentos 30:1-32.
LaFleur, J.R. 1992. Marketing of Brazil nuts. Food and Agricultural Organisation of the United
NationsAO, Rome.
Leigue Gómez, J.W., and R.G.A. Boot. in press a. Dinámica reproductiva de árboles de castaña
(Bertholletia excelsa) en un bosque alto tropical de tierra firme, de la Provincia Vaca
Diez del departamento del Beni. *
Leigue Gómez, J.W., and R.G.A. Boot. in press b. Competición de árboles contiguos y lianas
en la producción de frutos de castaña (Bertholletia excelsa) en un bosque del Norte de
Bolivia. *
Man-Producten. 1998. Edible nut market report. No 145. March 1998. Man-Producten BV,
Rotterdam.
Mori, S.A. 1992. The Brazil nut industry - past, present and future. Pages 241-251 in M.
Plotkin, and L. Farmocare, editors. Sustainable harvest and marketing of rain forest
products. Island Press, Washington. (www.nybg.org/bsci/braznut)
Mori, S.A., and G.T. Prance. 1987. Species diversity, phenology, plant-animal interactions, and
their correlation with climate, as illustrated by the Brazil nut family (Lecythidaceae).
Pages 69-89 in R.E. Pickinson, editors. The geophysiology of Amazonia.Mori, S.A., and
G.T. Prance. 1990. Flora Neotropica. Lecythidaceae - Part II. New York Botanical
Garden, New York.
Mori, S.A., and G.T. Prance. 1990. Taxonomy, ecology, and economic botany of the Brazil nut
(Bertholletia excelsa Humb.& Bonpl.: Lecythidaceae). Advances in Economic Botany
8:130-150.
108 • Bibliography

Moritz, A. 1984. Estudios biologicaos da castanha-do-Brasil (Bertholletia excelsa H.B.K.).

EMRRAPA/CPATU Documentos 29:1-82.
Mostacedo, B., and T.S. Fredericksen. 1999. Regeneration status of important tropical forest
tree species in Bolivia: assessment and recommendations. Forest Ecology and
Management 124:263-273.
Muller, C.H. 1981. Castanha-do-Brasil: estudios agronomicos. EMRRAPA/CPATU
Documentos 1:1-25.
Muller, C.H. 1982. Quebra de dormencia da semente e enxertia em castanha-do-Brasil.
EMRRAPA/CPATU Documentos 16:40.
Muller, C.H., I.A. Rodrigues, A.A. Muller, and N.R.M. Muller. 1980. Castanha do Brazil:
resultados de pesquisas. EMRRAPA/CPATU Miscelanea 2:1-25.
Myers, G.P., A.C. Newton, and O. Melgarejo. 2000. The influence of canopy gap size on
natural regeneration of Brazil nut (Bertholletia excelsa) in Bolivia. Forest Ecology and
Management 127:119-128. (abstract available at www.elsevier.com/locate/foreco)*
Nelson, B.W., M.L. Absy, E.M. Barbosa, and G.T. Prance. 1985. Observation on flower visitors
to Bertholletia excelsa H.B.K and Couratari tenuicarpa A.C. Sm. (Lecythidaceae). Acta
Amazonica 15:225-234.
Nepstad, D.C., I.F. Brown, L. Luz, A. Alechandre, and V.Viana. 1992. Biotic impoverishment of
Amazonian forests by rubber tappers, loggers and cattle ranchers. Advances in Economic
Botany 9:1-14.
Oliveira, M.V.N. 2000. Artificial regeneration in gaps and skidding trails after mechanised forest
exploitation in Acre, Brazil. Forest Ecology and Management 127:67-76. (abstract
available at www.elsevier.com/locate/foreco)
O’Malley, D.M., D.P. Buckley, G.T. Prance, and K.S. Bawa. 1988. genetics of Brazil nut
(Bertholletia excelsa Humb.& Bonpl.: Lecythidaceae). Theoretical and Applied Genetics
76:929-932.
Ortiz, E. 1995. Survival in a nutshell. Americas 47:7-12.
Ortiz, E., 2002. Brazil nut (Bertholletia excelsa). In: A. Guillen, S.A. Laird, P. Shanley and A.R
Pierce (eds.) Tapping the green market: certification and management of non-timber
forest products. Earthscan.
Peña-Claros, M. 2001. Secondary forest succession: processes affecting the regeneration of
Bolivian tree species. PhD thesis Utrecht University, Utrecht, the Netherlands. 170 pp.*
Peña-Claros, M., and H. de Boo. 2002. Seed removal of nine tree species in tropical rainforests
of the Bolivian Amazon: differences among areas differing in successional stage. Journal
of Tropical Ecology 18: 261-274.
Peña-Claros, M., R.G.A. Boot, and J. Dorado-Lora. 2002. Enrichment planting of Bertholletia
excelsa in secondary forest in the Bolivian Amazon: effect of cutting line width on
survival, growth and crown traits. Forest Ecology and Management 161: 159-168.
(abstract available at www.elsevier.com/locate/foreco)*
 Bibliography • 109

Peres, C.A., and C. Baider. 1997. Seed dispersal, spatial distribution and population structure of
Brazilnut trees (Bertholletia excelsa) in southeastern Amazonia. Journal of Tropical
Ecology 13:595-616.
Peres, C.A., L.C. Schiesari, and C.L. Dias-Leme. 1997.Vertebrate predation of Brazil-nuts
(Bertholletia excelsa, Lecythidaceae), an agouti-dispersed Amazonian seed crop: a test of
the escape hypothesis. Journal of Tropical Ecology 13:69-79.
PESACRE. 1996. Cultuvando a castanheira. No rocado, no campo e na mata. PESACRE,
Grupo de Pesquisa e Extensao em Sistemas Agroflorestais do Acre, Rio Branco, Brasil.
Pires, J.M. 1984. The Amazonian forest. Pages 581-602 in H. Sioli, editors. Limnology and
landscape ecology of a mighty tropical river and its basin. Junk, Dordrecht, the
Netherlands.
Poorter, L. 1998. Seedling growth of Bolivian rain forest tree species in relation to light and
water availability. Ph.D. Dissertation, Utrecht University.195 pp.*
Poorter, L. 1999. Growth responses of 15 rain-forest tree species to a light gradient: the relative
importance of morphological and physiological traits. Functional Ecology 13:396-410.*
PROMAB. 1999. Censo forestal para castaña (Bertholletia excelsa) en la reserva ecologico El
Tigre, Provincia Vaca Diez. PROMAB Informe Tecnico 3-99, PROMAB, Riberalta,
Bolivia.
Richards, M. 1993. The potential of non-timber forest products in sustainable natural forest
management in Amazonia. Commonwealth Forestry Review 72:21-27.
Salomao, R.P. 1991. Estructura e densidade de Bertholletia excelsa H. & B. (“Castanheira”) nas
regioes de Carajas, e Maraba, Estado do Para. Boletem do museu Paraense Emilio Goeldi
serie botanica 7:47-68.
Sanchez, J.S. 1973. Explotacion y comercializacion de la castaña en Madre de Dios. Informe
Nro 30, Ministerio de Agricultura, Peru, Lima, Peru.
Sokolov, R. 1985. Tough Nuts. Natural History 3:78-80.
Stoian, D. 2000. Shifts in forest product extraction: the post-rubber era in the Bolivian
Amazon. International Tree Crops Journal 10: 277-297.*
Stoian, D. and Henkemans, A.B. 2000. Between extractivism and peasant agriculture:
differentiation of rural settlements in the Bolivian Amazon. International Tree Crops
Journal 10: 299-319.*
Tabarelli, M., and W. Mantovani. 1996. Remocao de sementes de Bertholletia excelsa
(Lecythidaceae) por animais em uma floresta de terra firme na Amazonia Central, Brasil.
Revista Brasilea de Biologia 56:755-760.
Terborgh, J., E. Losos, M.P. Riley, and M. Bolaños Riley. 1993. Predation by vertebrates and
invertebrates on the seeds of five canopy species of an Amazonian forest.Vegetatio
107/108:375-386.
110 • Bibliography

Van Rijsoort, J., S. Ugueto, and P. Zuidema. 1993. The Brazil nut tree (Bertholletia excelsa):
population structures in tropical rain forest and growth response of seedlings to different
light intensities. MSc Thesis, Utrecht University, Utrecht.Viana,V.M., R.A. Mello, L.M.
Moraes, and N.T. Mendes. 1998. Ecologia e manejo de populacoes de castanha-do-Pará
em reservas extrativistas Xapuri, Estado do Acre. Pages 277-292 in C. Gascon, and P.
Mountinho, editors. Floresta Amazonica: dinamica, regeneracao e manejo. INPA, Manaus.
Vieira, A.H., M. Locatelli, and V.F. de Souze. 1998. Crescimento de castanha-do-brasil em dois
sistema de cultivo. EMBRAPA Boletim de Pesquisa 22, EMBRAPA Rondônia.
Watson, W. 1901. Germination of seeds of Bertholletia excelsa. Annals of Botany 15:99-102.
Williams, J., and D. Wilson. 1999. Informe sobre el problema de aflatoxinas de la castaña
(Bertholletia excelsa) en Bolivia. BOLFOR Documento Tecnico 71/99, Santa Cruz,
Bolivia. bolfor.chemonics.net/DOCUMENT/dt71.pdf)
Zuidema, P.A. 2000. Demography and exploitation of tree species in Northern Bolivia. Phd
thesis Utrecht University, Utrecht. (www.promab.org)*
Zuidema, P.A., W. Dijkman, and J. van Rijsoort. 1999. Crecimiento de plantines de Bertholletia
excelsa H.B.K. en funcion de su tamaño y la disponibilidad de luz. Ecologia en Bolivia
33:23-35. (abstract available at go.to/revistaecologia.bo)*
Zuidema, P.A. and R.G.A. Boot. 2002. Demography of the Brazil nut tree (Bertholletia
excelsa) in the Bolivian Amazon: impact of seed extraction on recruitment and
population dynamics. Journal of Tropical Ecology 18:1-31.*

Other literature used in this book

Hubbell, S.P. 1979. Tree dispersion, abundance, and diversity in a tropical dry forest. Science
203: 1299-1309.
Janzen, D.H. 1978. Seeding patterns of tropical trees. Pp. 83-128 in Tomlinson, P.B. &
Zimmermann, M.H. (eds). Tropical trees as living systems. Cambridge Univ. Press,
Cambridge.
Peters, C.M. 1996. The ecology and management of non-timber forest products. World Bank
Technical Paper 22, World Bank, Washington. 168 pp.
Poorter, L., R.G.A. Boot,Y. Hayashida-Oliver, J. Leigue-Gómez, M. Peña-Claros and P.A.
Zuidema. 2001. Dinamica de especies arboreas en un bosque humedo tropical en el norte
de la Amazonia Boliviana. In: Regeneracion y silvicultura de bosques tropicales en
Bolivia. B. Mostacedo, T.S. Fredericksen (eds). BOLFOR, Bolivia. pp 53-76.*
Zeide, B. 1993. Analysis of growth equations. Forest Science 39: 594-616.
 Bibliography • 111

Selected Internet sites with information on

the Brazil nut tree

EMBRAPA/CPATU - The Eastern Amazonian Research Centre (CPATU) of the Brazilian

Governmental Agricultural Research Organisation (EMBRAPA)
www.cpatu.embrapa.br
EMBRAPA/CPAFRO - The Rondônia Research Centre (CPAFRO) of the Brazilian
Governmental Agricultural Research Organisation (EMBRAPA)
www.cpafro.embrapa.br
Brazil nut home page - Includes list of literature on the Brazil nut tree and information on a
Brazil nut project in Peru.
www.bertholletia.org
PROMAB - Programa Manejo de la Amazonia Boliviana - Sustainable forest management
programme in Northern Bolivia.
www.promab.org
Bolivian Council for Voluntary Forest Certification / Consejo Boliviano para la certificación
Forestal Voluntaria (CFV) - Bolivian forest certification organisation.
www.angelfire.com/pq/cfv
 112 • PROMAB Publications

PROMAB publications

PROMAB Scientific Series

1. Poorter, L., 1998. Seedling growth of Bolivian rain Forest tree species in relation to ligth
and water availability. PhD thesis, Utrecht University. 195 pp.
2. Zuidema, P. A., 2000. Demography of exploited tree species in the Bolivian Amazon. PhD
thesis, Utrecht University. 240 pp.
3. Peña-Claros, M., 2001. Secondary forest succession - Processes affecting the regeneration of
Bolivian tree species. PhD thesis, Utrecht University. 170 pp.
4. Bojanic Helbingen, A., 2001. Balance is beautiful: Assessing sustainable development in the
rain forest of the Bolivian Amazon. PhD thesis, Utrecht University. 256 pp.
5. Henkemans A. B., 2002. Tranquilidad and hardship in the forest: Livelihoods and percep-
tions of Camba forest dwelers in the northern Bolivian Amazon. PhD thesis, Utrecht
University. 285 pp.

P R O M A B Te c h n i c a l S e r i e s

1. Peña, M., P. Zuidema, A. Zonta, A. Goncalvez, G. Chapi, 2001. Producción de palmito: limit-
caciones del manejo sostenible de poblaciones naturales de Asai (Euterpe precatoria) y el
potential del cultivo del tembe (Bactris gasipaes) como fuente alternativa.
PROMAB/IPHAE, Bolivia. 47 pp.
2. Poorter, L. R. Boot,Y. Hayashida, J. Leigue, M. Peña, 2001. Estructura y dinámica de un
bosque húmedo tropical en el norte de la Amazonía Boliviana. PROMAB, Bolivia. 54 pp.

These publications can be obtained free of charge from www.promab.org or by sending a

request to promab@bio.uu.nl. Recent information on PROMAB publications can be found at
www.promab.org.

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