Current Forestry Reports (2022) 8:90–110

https://doi.org/10.1007/s40725-022-00158-x

PHYSIOLOGICAL PROCESSES (M MENCUCCINI, SECTION EDITOR)

Advances in Brazil Nut Tree Ecophysiology: Linking Abiotic Factors

to Tree Growth and Fruit Production
Karen Cristina Pires da Costa1 · José Francisco de Carvalho Gonçalves2 · Alexandre Leão Gonçalves2 ·
Adamir da Rocha Nina Junior3 · Roberto Kirmayr Jaquetti2 · Vinícius Fernandes de Souza2 · Josiane
Celerino de Carvalho2 · Andreia Varmes Fernandes2 · Joelma Keith Rodrigues2 · Gleisson de Oliveira Nascimento4 ·
Lúcia Helena de O. Wadt5 · Karen A. Kainer6 · Roberval Monteiro Bezerra de Lima7 · Flávia Camila Schimpl8 ·
Jéssica Pereira de Souza2 · Sabrina Silva de Oliveira2 · Hellen Thaís da Silva Miléo2 · Diego P. Souza2 · Ana Claudia
Lopes da Silva2 · Heloisa Massaco Ito Nascimento2 · Jair Max Furtunato Maia9 · Francisco de Almeida Lobo10 ·
Paulo Mazzafera11 · Marcio Viana Ramos12 · Hector Henrique Ferreira Koolen13 · Ronaldo Ribeiro de Morais7 ·
Karina Martins14 · Niwton Leal Filho15 · Henrique Eduardo Mendonça Nascimento15 · Katharine Duarte Gonçalves2 ·
Yasmin Verçosa Kramer2 · Giordane Augusto Martins15 · Marcelo O. Rodrigues16

Accepted: 17 December 2021 / Published online: 10 February 2022

© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2022

Abstract
Purpose of the Review The Brazil nut tree (Bertholletia excelsa) is a symbolic tree in the Brazilian Amazonian. In a broad
sense, it plays a crucial role in its social, economic, and environmental importance. This species contributes on a large scale
to the equilibrium of the biological processes related to the biogeochemical cycles in the Amazon biome, and its nuts sus-
tain a multi-million-dollar extractive economy, which supports small farmers and traditional populations. Brazil nut is also
becoming one of the most important species in silviculture and is increasingly used in agroforestry systems and the recovery
of degraded areas. In this review, we deepened our understanding of the growth performance of the Brazil nut tree and its
ecophysiological traits, both in native trees and commercial forest plantations. Based on the literature for this species, we
discuss the concepts of plasticity and other functional traits that may help to increase Brazil nut plantation and conservation,
which in turn will increase nut production, forest sustainability, and social welfare.
Recent Findings The Brazil nut tree is a dominant species and is found throughout the Amazon region. Due to its ecophysi-
ological traits, it can be cultivated as a commercial monoculture, in the enrichment of forest plantations, used in the recovery
of degraded areas and the implementation of agroforestry systems. Recent evidence suggests that their dominance of natural
forests and their high functional performance under cultivated conditions may be associated with their physiological plastic-
ity and tolerance to abiotic stresses.
Summary Aspects related to phenotypic variation, genetic diversity, population characteristics, cultivation, and ecophysi-
ological performance of Bertholletia excelsa are revised and linked to growth and nut production. We demonstrate that
Brazil nut exhibits phenotypical plasticity in response to light, water, and nutrient availability. This trait can be explored for
improvements in nut production in native trees and agroforestry plantations. In both cases, the availability of these resources
influences population structure, tree growth, and fruit production. These results reinforce the importance of the use of Brazil
nut tree as an attractive alternative for improving programs that involve the recovery of degraded areas in the continental
Amazon. Lastly, the ecophysiological performance of the Brazil nut tree suggests its resilience to environmental change.

Keywords Forest plantation · Photosynthesis · Plasticity · Resources use efficiency · Tree physiology · Sustainability

This article is part of the Topical Collection on Physiological Introduction

Processes
Historically, the gigantic Brazil nut (Bertholletia excelsa)
* José Francisco de Carvalho Gonçalves
[email protected] tree has contributed to the development of complex socie-
ties in the Amazon region and its domestication dates from
Extended author information available on the last page of the article

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the first millennium of the Common Era [1]. Currently, this this species in natural forests and plantations, directly influ-
species is one of the main non-timber forest products mar- encing the density of individuals, biomass gain, fruit yield,
keted in Brazil. Additionally, it supports the livelihoods of population distribution patterns, and wood production [16,
populations throughout the Amazon basin by producing 20–25]. Water availability, on the other hand, may influence
Brazil nuts, which drives a multi-million-dollar extractive fruit or seed production. Nutrients, especially phosphorus,
economy in Bolivia, Brazil, and Peru [2]. Brazil nut also also seem to be a key determinant of growth and fruit yield
plays a fundamental role in carbon sequestration, ranking in natural forests and plantations [11, 26, 27].
first among economically important Amazonian species In reviewing the relationship between abiotic factors and
while coming in third place among 3458 species evaluated phenotypic characteristics, we posit that the Brazil nut tree
for biomass accumulation [3]. In sum, Brazil nut can be con- has adaptive plasticity to resource availability. This species
sidered a biocultural keystone species [4] due to its critical adjusts its morphophysiological traits (photosynthesis, leaf
multi-faceted role. morphology, resource use efficiency) to favor its survival
This important tree species belongs to the Lecythidaceae and growth under different environmental conditions. In
(Poiteau 1825) family. It was first catalogued in Western addition, the Brazil nut tree also tolerates stresses caused by
science by Alexander von Humboldt and Aimé Bonpland the low availability of water and nutrients in the soil and by
in their natural history expedition to the New World from variations in light availability. These characteristics help us
1799 to 1804. It was subsequently named as Bertholletia understand its vigorous performance and relative dominance
excelsa by Bonpland in 1808 [5]. The genus terminology in the Amazon forest and help explain why this species has
is dedicated to the chemist Claude Louis Berthollet in ref- performed well when planted in pure plantations, agrofor-
erence to his scientific stature, while the epithet refers to estry systems, mixed plantations, enrichment plantings, and
the grandiosity (excelsa = noble) of the adult trees [6]. The plantings to recuperate degraded areas.
social, economic, and ecological importance of Brazil nut Nonetheless, several important gaps in the ecophysiologi-
is due, in part, to its extensive distribution throughout the cal understanding of the Brazil nut tree exist. Because this
Amazon basin and its high rates of survival and growth, species is one of the principal native species targeted for
even in adverse conditions such as sites with acidic soils, restoration in the Amazon, addressing these gaps may guide
low water and nutrient availability, and variations in avail- continued wild tree conservation and production, as well
ability of irradiance [7–11]. In addition, Brazil nut is a large as highlight new silvicultural horizons [28]. Currently, it is
tree with a great capacity to sequester carbon and can play illegal to sell Brazil nut trees because of their traditional and
an important role in mitigating the effects of climate change contemporary non-timber values (i.e., nuts, carbon seques-
on Amazon forests [12, 13]. tration, and storage) (Brazilian Law decree No. 5.975/2006).
This review synthesizes the natural history and the eco- Despite that, some landowners have shown interest in poten-
physiological evidence that may explain the vigorous per- tial future markets for Brazil nut timber, because of its high
formance and relative dominance of Brazil nut in Amazon wood quality.
forests. We first demonstrate that the natural history of this Thus, considering the current and future social, eco-
species indicates phenotypic plasticity, which is defined as nomic, and environmental roles of Brazil nut, we discuss the
an organism’s ability to change, adapt, or respond to stimuli effects of resource availability on the functional and pheno-
or inputs from the environment such as low soil fertility or typic characteristics of this species. Finally, we also review
water, light, and temperature stresses [14]. We then inte- the application and implications of these ecophysiological
grate studies on phenotypic variation, genetic diversity, and characteristics of Brazil nut on its cultivation, closing with
a natural population’s characteristics to infer the influence a brief conclusion of prospects for conservation and culti-
of abiotic factors on phenotypic changes in Brazil nut and vation. The ecophysiological understanding of Brazil nut
the relationships between this ability and its vigorous perfor- can be used to refine and improve management techniques,
mance and relative dominance in Amazon forests. conservation practices, and social sustainability, and to fore-
The variation in the phenotypic characteristics of the cast impacts of climate change in the Brazil nut tree–rich
Brazil nut tree may be attributed partially to human selec- Amazon forests.
tion because humans commonly select phenotypes that have
larger crowns, fruits, and seeds during incipient domesti- The Natural and anthropogenic History of the Brazil
cation phases. Nonetheless, the characteristics of natural Nut Tree Indicates Physiological Plasticity
populations such as seed production, above-ground bio-
mass, regeneration, the density of individuals, and growth in The origin and distribution of Brazil nut throughout the
diameter appear to be directly affected by the availability of Amazon basin still intrigue scientists, and it has been
water, light, and nutrients [15–19]. Light seems to be one of linked with high physiological plasticity. The most wide-
the most critical factors in the physiological performance of spread hypothesis for the location of species origin is based

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on linguistic and phylogenetic evidence that suggests that the pattern expected by neutral processes [37]. The authors
Brazil nut originated in the central Amazon, with Lecythis argued that factors other than seed dispersal by agoutis may
as ancestor [29, 30]. However, this hypothesis is difficult to have left footprints in the Brazil nut genome. Indeed, it was
prove, considering that the species has very likely occurred confirmed that genetic diversity is more significant in popu-
in the region for at least several hundreds of thousands of lations located near predicted Pleistocene refuges. All this
years [31]. evidence reinforces the need for further studies to disen-
Additionally, theories have been proposed to explain the tangle the relative influences of agouti seed dispersal, local
wide distribution of Brazil nut in the Amazon basin and its adaptation, historic range shifts, and anthropogenic impacts
remarkable abundance in some regions. Adolphe Ducke in in natural populations of Brazil nut throughout the Amazon
1946 was the first to propose that pre-Columbian indigenous [34].
tribes had a significant role in the promotion of clumped
Brazil nut stands. This hypothesis, however, was challenged Phenotypic Variation and Genetic Diversity
by Peres and Baider [32] who suggested that seed preda-
tors, such as short-range agouti (Dasyprocta sp.), are more Bertholletia excelsa is monospecific, but natural populations
effective seed dispersers than humans and thus responsible display phenotypic variation in tree shape and fruit format
for the establishment of new stands. Nonetheless, over the [38]. In 1874, these differences led John Miers to describe
last decade, beginning with Shepard and Ramirez [30] who a species called Bertholletia nobilis. Still, the characteris-
emphasized the role of humans in Brazil nut recruitment, tics of collected specimens were not considered valid for a
several studies using different methodological approaches separate classification in the Lecythidaceae family, and con-
have supported the hypothesis that the humans have been sequently, B. nobilis was later synonymized with B. excelsa
central in species distribution [22, 33–35]. In our view, these [38]. The phenotypic variation is so remarkable in some
hypotheses are not mutually exclusive. Especially in areas of regions that traditional Amazonian communities recognize
anthropogenically disturbed forests such as forest clearings different varieties co-occurring in natural stands [39]. They
for small-scale agricultural plantings, agouti (Dasyprocta identify some morphological differences in wood color and
sp.) is a particularly effective facilitator of Brazil nut seed quality, fruit production, and trunk and crown form. The
placement and burial, and ultimately seedling establishment popular classification varies across Brazilian regions. In
[21]. lower latitudes within Mato Grosso state, Brazil, which is
Range shifts during the Pleistocene may also have the southern limit of the distribution of Brazil nut, the most
strongly influenced the current distribution of Brazil nut well-known types are named cor de rosa (pink), rajada
[31]. Since the last interglacial period, suitable conditions (striped), and mirim (small) [39]. The popular classification
and areas for species establishment emerged in the western is based on wood color, size of fruit, and quantity of seeds
to central Amazon. Following this period, however, when per fruit. When sampled in the same location, the “pink”
cooler and drier weather predominated during the last glacial trees produced heavier fruits and more seeds per fruit than
maximum (about 20,000 years ago), suitable species habitats the other popular types [39]. In distant eastern longitudes,
were reduced considerably, and Brazil nut distribution may in Acre state, the popular classification is based on the color
have been primarily restricted to several potential refuges and quality of the wood (red vs. white) and the size and
across the southern Amazon [31]. During the mid-Holocene, shape of the adult trees [40]. Locals believe the individuals
evidence suggests an expansion of favorable habitats for Bra- that produce reddish wood to be better for construction than
zil nut establishment, which are considered comparable to the white-type wood. These “red” individuals also displayed
current distribution conditions [31]. Today, the species is a larger crown and greater fruit production when compared
widely distributed and grows across various soil types, sea- to the “white” type [40]. Nonetheless, when the consistency
sonality regimes, and topographic conditions [36]. of phenotypical characteristics of trees, fruits, and seeds in
Thus, taking into account the different Brazil nut ori- individuals popularly classified as red or white types and
gin and distribution theories, acclimatization capacity and genotyped adult trees with molecular markers was evalu-
plasticity of this species to various environmental condi- ated, they concluded that the popular classification is not
tions seem important for explaining Brazil nut distribution supported by molecular differences. They also suggested that
throughout the Amazon. Pertinent processes may include the phenotypic differences among the types may represent
agouti seed dispersal, cultivation or favoring by traditional age-related variations [40].
human populations in different regions, and the ability of Phenotypical variations of Brazil nut across Amazonian
the species to survive periods of cooling/warming and dry- regions have been commonly pointed out as a consequence
ing/moistening [30, 31, 34]. One study that characterized of human selection because fruit size, wood quality, and
the genetic structure of natural populations of Brazil nut shape of the crown are considered typical traits selected
across the Brazilian Amazon indicated that it does not follow by humans in incipient domestication of managed species

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[41]. Despite this, the effect of resource availability on the trees, for example, are found throughout a large geographic
phenotypic variation of Brazil nut cannot be ruled out, and range, from 5° N to 14° S latitude, which comprises areas
evidence suggests that, at least in small scales (< 50 m), throughout the Brazilian Amazon and adjacent regions
the availability of resources, especially light and nutrients, of Bolivia, Peru, Colombia, Venezuela, and the Guianas
can directly affect Brazil nut fruit and seed production and (Fig. 1). In general, it is found in terra firme forests, but there
above-ground biomass, as well as the shape and size of the are reports of trees also occurring in floodplains (locally
crown and crop size [20]. known as “várzeas”) [36, 43]. Distribution models estimate
that the species occurs most commonly in areas with eleva-
Brazil Nut Tree Forest Populations tions of 200–400 m [44]. However, studies document that
the species can occur from sea level [31] to up to about
Historical data on the characteristics of natural populations 562 m above sea level [36].
of Brazil nut emanates almost exclusively from the Brazil Brazil nut tree plasticity’s observed extensiveness can at
nut commercial strongholds of Brazil, Peru, and Bolivia least partially explain the wide occurrence of the species in
(Fig. 1), although it occurs naturally throughout the Amazon environments with a wide range of abiotic factors. These
biome. Brazilian data is mostly restricted to Acre, Amapá, include air temperatures that range from approximately
Amazonas, Pará, and Rondônia Brazilian states. 24.3 to 27.2ºC, total annual rainfall variations between 1400
Phenotypic plasticity is a key point in explaining the and 2800 mm, and mean annual relative humidity ranging
dynamics of natural populations [42]. The ability of a plant between 79 and 86% [45]. Edaphic factors, such as particle
to colonize different ecosystems, therefore expanding its size, macroporosity, pH, and nutrient content are factors that
range and adaptability to diverse environments, can be due best predicted spatial variation in Brazil nut occurrence [46].
to greater (or lesser) degrees of plasticity [42]. Brazil nut Additionally, elevation seems to be an important predictor

Fig. 1  Distribution of studies and population densities of Brazil nut in the Amazon region

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for explaining the occurrence of Brazil nut in some sites, Brazil are associated with protected forest areas (Fig. 1). The
revealing that lowland areas also are suitable for species history of use of the sites and the availability of light have
establishment [36], although this correlation may not be been pointed out as determining factors for the rate of regen-
causal. This species presents better growth performance in eration and seedling densities, and a high mean of regenera-
soils with a clay to heavy clay texture, and its growth is tion and seedlings density has been observed in environ-
impaired in sandy soils [43, 47]. ments where there is greater availability of light [21, 33].
Another indicator of the high level of Brazil nut pheno- Average Brazil nut tree diameters vary from 44.9 cm in
typic plasticity is its long life span that is associated with Alter do Chão to 159 cm in Oriximiná, both in Pará state,
its ability to reach large diameters in natural forests. Large Brazil, where tree populations were among those with the
Brazil nut trees with > 160 cm in diameter at breast height lowest and highest densities of young plants, respectively
(DBH) may be over 300 years old, while those first entering (Table 5). The increasing availability of light favors the Bra-
reproductive sizes (~ DBH > 40 cm) may exceed 100 years zil nut plant’s growth rates [21, 48, 53–55] and can also be
of age [48, 49]. In the eastern Amazon, enormous Brazil associated with the time to reach maturity and the beginning
nut trees (4.46 m DBH) were estimated to be approximately of fruit production. Some reports also have shown that light
800 and 1000 years old [50], and radiocarbon dating sug- does not seem to influence the survival rate of seedlings
gests maximum lifespans can be over 1000 years [51]. The [48], but light does affect population structure. For instance,
main factors responsible for Brazil nut mortality in natural Brazil nut grows best in large forest gaps during the initial
populations appear to be wind, storms, human-ignited fires, developmental stages of its life cycle [38, 53]. The ability of
and liana infestations [49]. Like plants with long life spans, plants to colonize gaps and open areas is an important char-
Brazil nut may have compensatory mechanisms that bestow acteristic of plants that show physiological plasticity [14].
plasticity and maintain their photosynthetic capacity and The lack of standardized inventories also hampers more
vascular complexity, allowing them to grow and survive for precise information on Brazil nut population structure [16],
long periods of time [52]. including the use of sample units of different sizes and
Plastic responses to abiotic factors, such as light, water, shapes, differences in a minimum DBH limits, and criteria
and nutrients, can be observed in several population char- for plot locations [16]. Purposefully or unwittingly, inven-
acteristics. For example, Brazil nut trees often are found in tories may include areas that were modified by humans, and
groves of 5–100 individuals known as castanhais that are characteristics such as crown shape and height may result
separated from other groves by gaps of about 1 km in which from anthropogenic activity over time [33], reflecting the
the species is entirely absent [38]. However, a random distri- way the forest was occupied and/or exploited by humans.
bution pattern has also been reported [16, 18, 48]. In Acre, Sufficient and timely rainfall also increases growth
individual trees were separated by distances ranging from 1 rates in relation to diameter and has a strong relationship
to 233 m in locations considered to be rich in Brazil nut [16]. with fruit production, but to date, there is no evidence that
Groves versus more scattered distribution patterns may be drought causes changes in survival rates of Brazil nut tree
partially due to differences in forest types, and light seems populations [49, 56]. Phenotypic plasticity is beneficial in
to exercise an important effect [16]. Perhaps, open forest periods of water stress and favors the emergence and growth
provides more consistent favorable regeneration conditions, of plants even in places where water availability is limited.
such as high light intensity, which results in a more scat- Nutrient availability is also related to water, and Brazil nut
tered spatial distribution pattern and more constant seedling can grow in both low and high fertility soils. However, the
establishment over time [16]. Evidence suggests that a grove increased availability of phosphorus seems to favor species
distribution pattern may be influenced by human dispersal, growth and photosynthetic rates [11].
while the more scattered pattern may result from greater Tree diameter growth and canopy shape (related to can-
agouti-mediated seed dispersal [34]. opy size) are positively related to Brazil nut fruit yield [16].
Additionally, light availability has been associated with The variation in annual fruit yield of individuals is very
several other population characteristics of Brazil nut, such as large and can range from 9 to 218.5 fruits per tree (Table 6).
seedling density, diameter growth, population structure, and On average, the annual nut production is 15 kg t­ ree−1 with a
fruit production [18, 21, 33, 48, 53–55]. In natural popula- variation from 3.8 to 39.4 kg ­tree−1 (Table 6), which trans-
tions, for example, Brazil nut seedling densities vary from lates to 30 kg ­ha−1 with variation from 8 to 116 kg ­ha−1.
extremes of 0.005 trees h­ a−1 in Oriximiná to 25 trees h­ a−1 Several factors can be considered alone and in association,
in the Caxiunã Reserve, both recorded in the state of Pará such as (1) genetic background, (2) climate, (3) soil fertility,
(Fig. 1). In Brazil, the largest populations are concentrated in (4) biological competition, (5) tree size and architecture, (6)
the states of Acre, Amazonas, Amapá, Pará, and Roraima. In canopy position, (7) and level of liana infestation (Table 1).
Bolivia, recorded densities averaged about 1.7 trees h­ a−1 in Lianas have a negative effect on nut production based on
Beni to 3.3 trees h­ a−1 in Pando. Nowadays, high densities in studies located in the Chico Mendes Reserve in Acre state,

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Table 1  Effects of availability Availability of resource

of resources on the population
characteristics of Brazil nut Population Soil attributes
traits
Rain
Light Elevation Clay Silt Porosity Water CEC P Zn Cu
fall
Occurrence - - - - -
Survival - - - - - - - - -
Aggregate
- - - - - - - - - -
spatial
distribution
Population - - - - - - - - - - -
density
Regeneration - - - - - - - - - -
Diameter - - - - - - - -
growth
Life span - - - - - - - - - -
Age of - - - - - - - - - -
maturity
Fruit or seed - - - -
production
= indicates positive effects; = indicates a predominantly positive trend; = indicates a
predominantly negative trend; = indicates no effects.

Brazil [17, 20, 26]. The elimination of lianas increased nut including monoculture or pure plantations, agroforestry sys-
production significantly (3 ×). This increase was attributed to tems, mixed plantations, and enrichment plantings (Fig. 2).
reduced competition for light and below-ground resources, Brazil nut plantations have been established throughout and
such as water and nutrients [17]. Cation exchange capacity outside the species’ natural range, with the highest concen-
(CEC) and foliar phosphorus have been positively associated tration of Brazil nut plantations occurring in the Brazilian
with fruit production [26]. On the other hand, in lowland Legal Amazon (Fig. 2).
areas, where individuals can spend months in flooded con- The wide distribution of Brazil nut plantations throughout
ditions, fruit production seems lower [43, 56]. Other soil very different climatic conditions beyond Amazon biome
attributes, especially the availability of phosphorus ­(PO43−), borders reinforces inferences about phenotypic plasticity. In
exchangeable aluminum (Al), pH, and potassium (K), seem Lavras, Minas Gerais state, where Brazil nut tree species is
to influence fruit yield [26, 56, 57], but nutritional research also planted (Fig. 2), mean air temperatures range from 18
has been limited. to 20 °C, and total annual rainfall ranges between 1300 and
While there are still gaps regarding which abiotic factors 1700 mm [63]. These variations in temperature and rainfall
determine growth and fruit production of Brazil nut trees are very different from the conditions of the natural species
in natural forests, recurring results converge on several abi- occurrence whereby the mean air temperature ranges from
otic factors that seem to influence Brazil nut populations 24.3 to 27.2 °C and total annual rainfall ranges between 1400
(Table 1). Cumulative research to date indicates that irradi- and 2800 mm [45]. Brazil nut plants exhibit robust physi-
ance in natural populations seems to be the factor that most ological processes (for example photosynthesis) capable to
influences Brazil nut tree regeneration, diameter growth, handle functional adjustments in order to ensure higher sur-
landscape distribution patterns, and time to reach repro- vival rates and growth in different environments [64].
ductive maturity, as supported by several authors [21, 38, Researchers and farmers considered several economic
58–61]. and technical aspects before Brazil nut plantations became
widespread in the Amazon. Observed low fruit production
Cultivating the Brazil Nut Tree in previously established plantations coupled with pest and
disease risks associated with pure plantations were the main
Brazil nut cultivation is not a recent activity, and there are concerns. “Growing Bertholletia excelsa in plantations has
records of plantings by humans dating from the first millen- not been very successful, probably due to the lack of effi-
nium of the Common Era [1]. Contemporary planting has cient pollinators and the risk of inbreeding” [38, 60] and
been attributed to Japanese immigrant settlers in the Brazil- “Bertholletia excelsa monoculture plantations are an option
ian cities of Tomé-Açu, Pará state, and Parintins, Amazo- for already capitalized investments, but in the future, they
nas state, beginning in the 1930s [62]. The Brazil nut tree may be limited by the attack of pests and disease.” [65] Lit-
is currently being cultivated in different planting systems, tle was known about B. excelsa pollinators and fears that

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Fig. 2  Distribution of studies and plantation systems of Brazil nut

concentrated plantings might provide favorable conditions predominantly reproduces by cross-pollination and viable
for pest and/or disease development were widespread, as seeds are mostly produced after crossing among unrelated
had been observed with the rubber tree and the pathogen trees [73–75]. A study conducted in the Peruvian Amazon
Microcyclus ulei [66]. demonstrated that fruit yield was negatively affected by con-
Brazil nut trees often are widely recommended for com- specific proximity, probably because of biparental inbreed-
mercial plantations beyond the geographic boundaries of ing and resource competition [76].
the Amazon region. So far, pests and/or diseases affect- Low germination rates (of about 25%) and delayed seed-
ing growth or fruit production by this species have been ling emergence (from 12 to 15 months) also discouraged
described [67, 68]. Only one disease (brown spot) caused Brazil nut tree plantations in the past [77]. However, removal
by the fungi Colletotrichum sp. has been detected, infecting of the external integument (testa) from the seeds and treat-
leaves, but with apparently no damage to growth or fruit ment with fungicides before sowing have proven to raise
production [67, 69]. Species with high levels of genetic plas- germination rates to over 90% [77].
ticity are more tolerant to pests and are also able to recover The Brazil nut tree has several silvicultural characteristics
more quickly after disturbances caused by diseases and pests that make it one of the most frequently recommended native
[70, 71]. forest species for restoration of degraded areas, agroforestry
One study investigated several aspects of Brazil nut tree systems, forest enrichment, and monoculture plantations for
pollination ecology in plantations [72], but the relationship the production of fruits and wood [61, 78–80]. Diameter
between pollination success and fruit yield is still unclear. growth in plantations varies from 9.6 mm ­year−1 in mixed
Evidence from mating system studies indicates that lower plantations used to restore degraded areas to 31.6 mm ­year−1
fruit yield in plantations may be due to genetic incompat- in agroforestry systems, and plant survival is over 80%
ibility among trees and inbreeding. The Brazil nut tree (Table 7) [61, 79–81]. In these planted environments, little

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is known about fruit production, but light availability has species has a high capacity both to support and to recover
been reported to have a strong positive influence on growth from disturbances caused by changes in the availability of
rates (diameter and height) [64]. light, water, and nutrients that may occur due to silvicultural
Seedling establishment in enrichment planting (increas- interventions such as thinning [11].
ing the number of a specific tree in disturbed or undisturbed Among the native species, Brazil nut is one of the most
areas) has also been examined and Brazil nut trees support promising for reforestation and recovery of degraded
the variation in light availability that occurs according to the areas from different activities such as mining and graz-
different sizes of clearings, but better results were seen in ing of animals, which are among the main types of deg-
large clearings, where the availability of light is more signifi- radation in the Amazonian forests [9, 11, 27, 61, 62].
cant. In Bolivia, in an enrichment planting, after 4 years, the The recommendation to use Brazil nut for the recovery
survival rate of seedlings varied between 59 and 94% [59]. of degraded areas is mainly due to its capacity to tol-
The results obtained in Trombetas experimental plantations, erate stress caused by the extreme conditions imposed
Pará state, Brazil, followed the pattern observed in trees of by these adverse environments, such as high irradiance
enrichment experiments, where the degree of light exposure and low availability of water and nutrients. In addition to
was the primary determinant for the best performance of tolerating the stresses caused by the harsh conditions of
the plant, increasing the growth with the size of the canopy the degraded sites, the species also possesses efficiency
opening [82]. in the use of the most limiting resources such as water
In agroforestry systems, it has been observed that Brazil and nutrients. In the case of mined areas, the species’
nut trees invested a substantial part of their biomass and capacity to tolerate high levels of heavy metals in soils
nutrients on branches and crowns, it shows a tendency of demonstrates the high potential of the species for phy-
Brazil nut trees to develop spreading crowns with very toremediation [80].
large and heavy branches when grown under agroforestry The fruit production of Brazil nut trees growing in
conditions [9]. In addition, the availability of Ca and Mg forests normally begins at ages over 70 years [49, 83]. In
was associated with higher biomass production in Bra- plantations, the production generally starts between 12 and
zil nut trees in agroforestry systems. The changes in the 15 years [68], but there are reports of Brazil nut trees that
biomass allocation pattern in plants are an essential strat- fruit at 10 years [38], 8 years [68], 6 years [62], and even at
egy of acclimatization and plasticity that gives plants the 5 years [84]. In plantations, annual fruit production by tree
ability to increase efficiency in capturing or using more varies from 1 to 132 fruits ­tree−1 [81, 85], and annual seed
limited resources for their survival and growth. production varies from 7.5 to 24.0 kg t­ ree−1 [68, 81, 86]. The
Additionally, differences between the growth rates of the evaluation of plantations and fruit production in Rondônia
Brazil nut tree in agroforestry systems were mainly attrib- showed a variation from 73 to 1,792 fruits h­ a−1 over 14 years
uted to the availability of nutrients in the soil and also the of monitoring [86]. Production did not show a clear relation-
historical use of the areas, since the growth of the species ship with the ages of the trees during this period and did not
can be favored primarily by the levels of phosphorus and exhibit differences between the production of trees in mixed
organic matter in the soil [68]. Similar results were observed plantations and pure plantations [86].
in fertilized agroforestry systems, in which trees responded In general, ecophysiology studies of Brazil nut trees have
to increased levels of nitrogen, phosphorus, and lime, with been carried out under different forest site conditions and
significantly improved foliar nutrition and growth. Plastic- these edaphoclimatic variations make it difficult to restore
ity is partially responsible for determining the responses of the linking abiotic factors with tree growth and fruit produc-
plant performance to soil fertilization. The rationale is that tion in natural conditions. For Brazil nut plantations, we
functional traits reflect the plants’ capacities for resource found recurrent information in published studies that showed
capture and adaptations to environmental changes. us that light availability is the most investigated resource
The effects of silvicultural interventions, such as fertiliza- regarding the survival and growth of Brazil nut plantations.
tion and thinning, on the growth rates of the Brazil nut were Light availability favors plant growth in enrichment planta-
also tested in pure plantations formed by the species [11]. As tions, agroforestry systems, and monocultures, especially in
a monoculture, the species has shown high growth rates and, diameter, but it does not affect the survival of the seedlings
in general, in the Amazon region these plantations have been in the initial phase of the plantation (Table 2). In addition,
established without the adoption of silvicultural interven- edaphic factors, mainly the availability of Ca and Mg, were
tions such as the application of fertilizer or correction of soil highlighted for biomass production in Brazil nut trees in
pH and the control of ants [11]. Even in these conditions, agroforestry systems.
the species has a high rate of survival and growth when
compared to other native and even exotic species planted
under the same site conditions. In these planting systems, the

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Table 2  Effects of the

availability of resources on
Availability of resources
Plantation’s traits
plantation traits of Brazil nut Light Water CEC pH Ca Mg P
Survival - - - - -
Diameter growth - - - - -
Height growth - - - - -
Biomass - - - - -
Wood production - - - - -
Fruits production - - - - - -
= indicates positive effects; = indicates a predominantly positive trend; = indicates no effects.

Brazil nut Tree Functional Traits and Ecophysiology common photosynthetic behavior for the species. The dark
(Rd) respiration rates found for the Brazil nut vary from
The description of leaf phenological stages is essential 0.12 to 2.9 µmol ­m−2 ­s−1, and the range of values for sto-
for understanding the functional traits and ecophysiology. matal conductance (gs) and transpiration (E) is from 0.02
These measurements are generally carried out at the leaf to 0.59 mol ­m−2 ­s−1 and from 0.57 to 6.5 mmol ­m−2 ­s−1,
level and according to strict data collection protocols. Most respectively [8, 10, 27, 88]. In general, the lowest Pn, Rd,
commonly, in the field, it is possible to identify at least gs, and E values have been observed in seedlings subjected
four leaf phenological stages in the Brazil nut tree: recently to severe water deficit [10]. In contrast, the highest values
released leaves, new leaves, mature leaves, and old leaves. have been observed in plants exposed to direct sunlight and
The youngest leaves always occur at the base of the branches fertilized [8, 10, 27, 64, 88].
and will differentiate into older leaves over time. A typical photosynthesis curve can indicate a light com-
Newly released and new leaves are tender and can be pensation point ranging from 2.11 to 71 µmol ­m−2 ­s−1, satu-
larger than mature and old leaves, but this depends on the ration point ranging from 190 to 1.032 µmol ­m−2 ­s−1, and
plant’s growing environment. The main difference between quantum yield ranges between 0.026 and 0.08 μmol μmol−1
the newly released leaves and the new leaves is their color, as [8, 27, 64, 87, 88]. An unusual quantum yield value
recently released leaves have a light brown color. The differ- (0.12 mmol ­mmol−1) was also already found for Brazil nut
ence between new and mature leaves is mainly in the texture seedlings in controlled conditions [64]. These values can
and color of the leaves. The mature leaves are less tender vary according to the light environment, water availability,
than the new ones and have a more intense and brighter soil fertility, and leaf age. There is not a large variation in
green color. Regarding the difference between mature leaves the photosystem II photochemical efficiency (­ FV/FM), which
and old leaves, the main difference is the leathery appear- ranges from 0.58 to 0.85, with lower values being found in
ance and the more opaque green color in the older leaves. plants under full sunlight and in nutrient-impoverished soils
The mature leaves show the leaf area (LA) ranges from [7, 8, 10, 84, 85].
74.8 to approximately 400 c­ m2 according to their age and Concentrations of macro- and micronutrients in leaves
the environmental conditions, with higher values occurring of Brazil nut trees at different ages, growing in different
in shaded plants [8, 27, 64, 87, 88]. The specific leaf area conditions across the Amazon region show little varia-
(SLA) ranges from 91.3 to approximately 200 ­cm2 ­g−1, with tion (Table 3), with higher values being observed in young
higher values found in shaded plants and with lower values plants in the greenhouse and the lowest in plantations in
observed in plants under full sunlight [8, 27, 64, 87, 88]. degraded areas. In general, the order of nutrient concentra-
In general, changes in leaf area occur due to changes in the tion in the leaves of the Brazil nut tree is N (18 g ­kg−1) > Ca
light environment and differences in soil fertility. On the (7 g ­kg−1) > K (5 g ­kg−1) > Mg (2 g ­kg−1) > P (1 g ­kg−1) > Mn
other hand, changes in specific leaf area values have been (102 mg ­kg−1) > Fe (68 mg ­kg−1) > Zn (28 g ­kg−1) (Table 3).
observed relating to changes in the light environment and The highest concentrations of macro- and micronutrients
water availability at the site and less from changes in soil are observed in the leaves + fine branches, and the crowns
fertility [8, 27, 64, 87, 88]. represented about 57% of the total nutrient stock in Brazil
In the literature, it is possible to find photosynthe- nut trees [89].
sis values (Pn) for Brazil nut trees that vary from 0.5 to Brazil nut is considered a tolerant species to the low soil
15 µmol ­m−2 ­s−1 [8, 10, 27, 88]. However, in general, Pn nutrient availability, but fertilization favors the ecophysi-
values between 7 and 10 µmol ­m−2 ­s−1 represent the most ological performance and growth of Brazil nut [27]. Higher

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Table 3  Concentrations of macro- and micronutrients in the leaves of Brazil nut at different ages and under different growth conditions in the
Amazon region
Growth condition Amazon region N P K Ca Mg Fe Zn Mn
g ­kg−1 mg ­kg−1
Pure plantation, 8 ­years1 Itacoatiara, AM, Brazil 17.5 0.7 6.2 4.3 2.4 57.5 25.7 94.5
Vegetation house, 9 ­months2 Itacoatiara, AM, Brazil 20.2 1.1 5.5 4.2 1.5 39.3 36.7 52.9
Clonal (606) ­plantation3 Itacoatiara, AM, Brazil 19.2 1.0 5.7 5.6 2.1 64.2 29.0 107.4
Clonal (609) ­plantation3 Itacoatiara, AM, Brazil 19.7 0.9 4.4 8.4 2.2 67.4 29.2 134.5
Clonal (ARU) ­plantation3 Itacoatiara, AM, Brazil 18.8 1.0 4.7 6.1 2.3 66.2 30.1 99.9
Clonal (Manuel Pedro) ­plantation3 Itacoatiara, AM, Brazil 19.0 1.0 4.8 6.3 2.6 70.3 28.5 124.1
Clonal (Santa Fé) ­plantation3 Itacoatiara, AM, Brazil 19.3 1.0 4.4 7.8 2.6 68.7 30.2 149.2
RAD plantations, 1 ­year4 Manaus, AM, Brazil 10.7 1.2 3.1 11.1 2.4 119.6 19.6 32.6
Mixed plantations, 10 ­years5 Manaus, AM, Brazil 18.5 0.7 4.1 8.8 2.7 62.3 28.2 123.2
Agroforestry, 7 ­years6 Manaus, AM, Brazil 19.4 1.1 7.0 9.5 2.7 - - -
Pure plantation, 17 ­years7 Itacoatiara, AM, Brazil 17.2 0.9 3.6 5.5 2.5 97.1 20.5 95.2
Pure plantation, 12 ­years8 Itacoatiara, AM, Brazil 17.1 0.7 4.6 5.6 3.7 36.6 18.9 62.4
Pure plantation, 29 ­years8 Claudia, MT, Brazil 19.3 1.1 8.4 5.2 2.7 46.4 18.7 32.4
[1]=Costa et al. 2015; [2]=Maia et al. 2015; [3] Ferreira et al. 2015; [4]=Ferreira et al. 2013; [5]=Morais et al. 2007; [6]=Schroth et al.
2015; [7]=Lopes 2018, unpublished data; [8]=Castro 2017, unpublished data

photosynthesis, growth rates, and biomass accumulation irradiance availability. The species is tolerant to the low
in seedlings of Brazil nut have been related to the positive water availability and is highly efficient in the use of nutri-
effects of fertilization on stomatal conductance, respira- ents, especially phosphorus (Table 4). These responses
tion and the efficiency of uptake, and use of other primary were associated with the plasticity of functional traits that
resources, such as water and light [27, 64]. the species develops, favoring the capture and use of these
The functional traits of mature leaves of Brazil nut trees resources [10, 11, 27, 64, 87, 88].
growing under controlled and field conditions, thus sub- The specific leaf area and the nutrient and water use
jected to variations in the availability of light, water, and efficiency are improved in the Brazil nut trees at high light
nutrients, indicate that plants benefit from an increased availability, allowing it to keep the stomata open and thus

Table 4  Physiological responses of Brazil nut to the variation in the availability of light, water, and nutrients
Physiological traits Light1 Water2 Nutrients3
Shade leaves Sun leaves Dry season Rainy season Without fertili- Fertilized
zation

Pn (µmol ­m−2 ­s−1) 10.34 12.61 11.21 11.74 8.23 11.60

gs (mol ­m−2 ­s−1) 0.24 0.30 0.28 0.24 0.26 0.37
E (mmol ­m−2 ­s−1) 4.14 4.77 4.62 4.29 3.57 4.71
Rd (µmol ­m−2 ­s−1) 1.03 1.72 1.47 1.28 0.82 0.95
SLA ­(cm−2 ­g−1) 115 95 115 95 142 150
Chl a/Chl b 2.92 3.05 2.88 3.08 3.46 3.29
Chltotal/Car 2.83 2.65 2.72 2.77 3.07 2.50
FV/FM 0.82 0.78 0.81 0.80 0.74 0.76
NUE (mmol mol-1 ­s−1) 0.56 0.70 0.61 0.66 0.66 0.62
PUE (mmol mol-1 ­s−1) 11.02 14.45 10.83 14.64 9.27 10.30
WUE (µmol ­CO2 ­mmol−1 ­H2O) 2.63 2.69 2.53 2.79 2.28 2.50

Obs.: The values are means calculated from literature data. Pn photosynthesis, gs stomatal conductance, E transpiration, Rd respiration, SLA
specific leaf area; ­Chltotal chlorophyll total, Car carotenoids, FV/FM photochemical efficiency, NUE nitrogen use efficiency, PUE phosphorus use
efficiency, WUE water use efficiency. Authors: 1 = [83, 84]; 2 = [7, 8, 27, 75, 83]; 3 = [85, unpublished data].

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increasing photosynthesis rates [64, 88]. Additionally, due Apart from the high protein content (15.7%), Brazil nuts
to high light intensity, the species invests in photoprotective may be considered oilseeds (70.7%) and also an important
strategies, with the possibility of increasing the concentra- source of P, Mg, and Zn [90]. Storage proteins derived from
tion of carotenoids in the leaves [64, 87]. On the other hand, albumin isoforms [91] and selenoproteins [92] are important
under shady conditions, a larger leaf area and an increase components, in addition to fatty acids [93] and their respec-
of chlorophylls are observed, allowing an improvement of tive triacylglycerides and phospholipids [94]. Inorganic
light capture [64, 87, 88]. It has also been observed that phosphate and organic phosphate derivatives (phosphatidic
shaded plants increase their biomass allocation in leaves and acid and phosphatidylinositol lipids) accumulate in the Bra-
branches as a strategy for capturing more light, while plants zil nuts [94].
in full sun invest a greater amount of biomass in the roots, Besides being crucial for the biosynthesis of ATP, phos-
which can favor the capture of water and nutrients [64, 88]. phate is also important in the biosynthesis of amino acids,
Another interesting aspect of the Brazil nut tree is its abil- terpenes, phytosteroids and tocopherols (mainly as organic
ity to recover from stress caused by sudden changes in the diphosphates), and simple phenolic compounds, flavonoids,
availability of light, which is represented by the recovery of and tannins [93, 95]. The abundant availability of such
the photosystem II photochemical efficiency (­ FV/FM) value metabolites qualifies Brazil nut as a promising resource of
at levels close to 0.8, such as that which can occur naturally biologically-active substances, with industrial applications
in forests with the opening of clearings or in plantations after in the cosmetic, food, and biotechnological fields.
thinning [64]. Some research has attributed this response to
the efficient capacity of this species to release excess energy
in the form of heat and maintain electron transport rates in Conclusion and Future Prospects
the transport chain [11]. The maintenance of electron trans-
port rates, even under stress conditions due to high irradi- The importance of the Brazil nut tree in the social, eco-
ance, seems to be related to the species’ ability to develop nomic, and environmental framework of the Amazon can be
photoprotective strategies such as increasing iron and carot- attributed, in part, to its plasticity and other biological abili-
enoid concentrations in its leaves [11]. ties discussed in this review. How to deal more efficiently
The Brazil nut tree is tolerant to water deficit [10]. The with different abiotic factors and plant stresses or cope with
water potential values observed for Brazil nut trees range their consequences may be the significant differential of
from − 0,19 to − 4.7 MPa [10, 79]. Brazil nut trees under this species, explaining the Brazil nut tree’s physiological
water deficiency increase the concentration of osmoregula- performance.
tory solutes, such as potassium and proline in the leaves, Thus, following an ecophysiological approach, we aimed
allowing stomata opening [10]. The Brazil nut tree also to enhance our understanding of the effects of resource avail-
alters the allocation of carbon with the increase in root ability on the Brazil nut tree’s growth performance to high-
growth, thus favoring the capture of water by the roots [10]. light the plasticity and other eco-functional characteristics
Although the species can also occur in flooded areas, there is that are potentially useful for improving the production of
still no information in the literature regarding the effects of the species. As a result, we confirm that B. excelsa shows
flooding on the physiological characteristics of the species. phenotypic plasticity in response to light, water, and nutrient
availability. In both natural forests and plantations, the avail-
Seeds Chemical Components with Economic ability of these resources influences population structure,
Relevance and Bioperspectives tree growth, and fruit production.
Light represents a major factor in determining the struc-
The climatic conditions of the Amazon Rainforest and the ture of populations and growth in both natural forests and
different extreme conditions of cultivation lead to the acti- plantations. The Brazil nut tree shows tolerance to stresses
vation of the functional plasticity of Brazil nut, in particu- caused by the reduction of soil water and nutrient availability
lar, a robust photosynthetic apparatus that is a result of its and variations in irradiance. This supports using this species
complex and specific genome. This qualifies it as a large as an alternative in programs to recover degraded areas in
“factory” for the production and storage of important chemi- the continental Amazon. We also found robust evidence that
cal components, some of them “compartmentalized” in the increases in irradiance may lead to increased fruit produc-
Brazil nuts (fruits), which are the main commodity. Among tion; however, fruit production is also affected by competi-
chemical components in the Brazil nuts are the primary tion with lianas and edaphic factors, especially phosphorus
metabolites (proteins, lipids, and carbohydrates), second- availability and soil cationic exchange capacity.
ary metabolites (terpenes, phytosteroids, flavonoids, tannins, Although this review has compiled several reports on
and tocopherols) and other nutrients such as selenium and the physiology of the Brazil nut tree, such as photosynthe-
inorganic phosphate (Fig. 3). sis, biomass gain, vegetative propagation, susceptibility

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Fig. 3  Illustrative scheme of primary and secondary metabolic pathways applied to Brazil nut seeds

to climate change, plantation management and modeling, inferences about how to improve the silvicultural interven-
genetic enhancement, ecology, fruit production, Pi accu- tions. Additionally, research well designed and executed may
mulation in the nuts, and implications on seed metabolic help to potentiate synergies for sustainable practices, mini-
pathway, we believe that we are still far from understanding mizing the impacts of climate change on both native trees
the physiological behavior of this species under the different and commercial forest plantations.
system cultivation possibilities. Further studies are neces-
sary to assist management practices, which will undoubt-
edly improve Brazil nut production and allow making robust

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Appendix

Table 5

Table 5  Population traits of Brazil nut in natural forests in different Amazon regions
Amazon region Density Juveniles DBH References
(Tree ­ha−1) (%) (cm)

Pinkaiti Reserve, Pará, Brazil 3.30 - - Baider 2000

O Deserto, lower Xingú River, Pará, Brazil 1.00 - - Campbell et al. 1986
Tapajós Reserve, Pará, Brazil 0.67 - - Carvalho 1981
Chico Mendes Reserve—Fallow, Acre, Brazil 12.7 - - Cotta et al. 2008
Chico Mendes Reserve—Forest, Acre, Brazil 5.30 - - Cotta et al. 2008
Cajari, Amapá Reserve—Forest, Brazil 7.00 21.0 112 Guedes et al. 2014
Cajari, Amapá Reserve—Capoeira, Brazil 11.00 63.0 57 Guedes et al. 2014
Tambopata Reserve, Peru 1.30 - - Gentry 1988
Alto Cajari—Forest, Amapá, Brazil 6.70 - - Neves et al. 2015
Alto Cajaria—Capoeira, Amapá, Brazil 11.00 - - Neves et al. 2015
Filipinas, Acre, Brazil 1.80 28.0 74.6 Neves et al. 2016
Cachoeria, Acre, Brazil 2.70 17.0 88.4 Neves et al. 2016
Água Branca, Amapá, Brazil 6.80 22.0 92.9 Neves et al. 2016
Sororoca, Amapá, Brazil 11.20 10.0 108.6 Neves et al. 2016
Tahuamanu Reserve, Madre de Dios, Peru 0.86 - - Nunes et al. 2012
Tambopata Reserve, Madre de Dios, Peru 0.40 - - Nunes et al. 2012
Indigenous land Kayapó, Pará, Brazil 1.30 - - Peres and Baider 1997
Pinkaiti Kayapó land, Pará, Brazil 3.30 43.3 72.6 Peres et al. 2003
Kranure Kayapó land, Pará, Brazil 3.40 52.5 65.7 Peres et al. 2003
Saracá-Taqüera land, Pará, Brazil 1.50 1.6 134.8 Peres et al. 2003
Marabá, Pará, Brazil 4.30 33.3 119.7 Peres et al. 2003
Tapajós Reserve, Pará, Brazil 0.70 35.7 73.8 Peres et al. 2003
Alto Cajarí Reserve, Amapá, Brazil 12.00 0.7 156.4 Peres et al. 2003
Iratapuru Reserve, Amapá, Brazil 9.40 0.9 154.1 Peres et al. 2003
Aventura, Lago Uauaçú, Amazonas, Brazil 6.80 24.6 102.3 Peres et al. 2003
Ussicanta, Lago Uauaçú, Amazonas, Brazil 8.50 3.8 133.5 Peres et al. 2003
Lago Cipotuba, Rio Aripuanã, Amazonas, Brazil 1.80 22.4 116.6 Peres et al. 2003
Amanã Reserve, Amazonas, Brazil 1.40 5.3 123.5 Peres et al. 2003
Rio Cristalino, Mato Grosso, Brazil 4.90 40.7 90.7 Peres et al. 2003
Cláudia, Mato Grosso, Brazil 3.60 31.2 71.0 Peres et al. 2003
Nova Esperança, Acre, Brazil 3.10 47.2 73.9 Peres et al. 2003
Colocação Tucumã, Acre, Brazil 1.40 12.2 108 Peres et al. 2003
Colocação Rio de Janeiro, Acre, Brazil 1.40 32.4 89.8 Peres et al. 2003
Encontro, Acre, Brazil 1.40 25.1 91.4 Peres et al. 2003
Oculto, Madre de Dios, Peru 0.70 10.9 109.9 Peres et al. 2003
Limón, Madre de Dios, Peru 0.10 10.6 108.2 Peres et al. 2003
El Tigre, Beni, Bolivia 1.70 25.0 102.1 Peres et al. 2003
El Sena, Pando, Bolivia 3.30 21.7 111.3 Peres et al. 2003
Alter do Chão, Pará, Brazil 23.00 75.6 44.9 Peres et al. 2003
Rio Ouro Preto Reserve, Rondônia, Brazil 2.00 4.5 127.7 Peres et al. 2003
Kikretum Kayapó land 1.7 - - Ribeiro et al. 2014
A’Ukre Kayapó land 3.5 - - Ribeiro et al. 2014
Moikarakô Kayapó land 2.4 - - Ribeiro et al. 2014
Brazil nuts conssesions, Madre de Dios, Peru 0.62 - 126.3 Rockwell et al. 2015

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Table 5  (continued)

Amazon region Density Juveniles DBH References

(Tree ­ha−1) (%) (cm)
Carajás, Pará, Brazil 1.30 - 134.8 Salomão 1991
Marabá, Pará, Brazil 4.20 - 131.4 Salomão 1991
Platô Almeida, Oriximiná, Pará, Brazil, 1.49 1.2 - Salomão 2009
Platô Aviso, Oriximiná Pará, Brazil 0.005 71.4 - Salomão 2009
Platô Bacaba, Oriximiná Pará, Brazil 0.3 - - Salomão 2009
Platô Bela Cruz, Oriximiná Pará, Brazil 0.023 - - Salomão 2009
Trombetas River, Pará, Brazil 6.80 7.0 128.5 Scoles and Gribel 2011
Capanã Grande, Amazonas, Brazil 12.50 18.0 73.1 Scoles and Gribel 2011
River Trombetas Region, Pará, Brazil 6.80 - 128.4 Scoles and Gribel 2012
Oriximiná, Pará, Brazil 2.00 4.6 159 Scoles et al. 2016
Caxiuanã Reserve, Pará, Brazil 25.00 54.5 64.9 Sousa et al. 2014
North Bolivia 1.70 - - Stoian 2004
Caracaraí-Itã, Roraima, Brazil 13.50 - 82.7 Tonini et al. 2014
Caracaraí-Cujubim, Roraima, Brazil 6.50 - 118.8 Tonini et al. 2014
São João da Baliza, Roraima, Brazil 3.70 - 65.9 Tonini et al. 2014
São João Baliza, Roraima, Brazil 3.70 35.3 65.9 Tonini et al. 2008
Caracaraí, Roraima, Brazil 12.90 26.7 74.6 Tonini et al. 2008
Caracaraí-Itã, Roraima, Brazil 13.00 - 81.8 Tonini and Baldoni 2019
Caracaraí-Cujubim, Roraima, Brazil 6.00 - 112.6 Tonini and Baldoni 2019
Itaúba, Mato Grosso, Brazil 15.00 - 59.9 Tonini and Baldoni 2019
Chico Mendes Reserve, Acre, Brazil 1.35 25.5 86.1 Wadt et al. 2005
Cachoeira, Acre, Brazil 2.50 - 93 Wadt et al. 2008
Pindamonhangaba, Acre, Brazil 2.20 - 71.6 Wadt et al. 2008
Filipinas, Acre, Brazil 1.50 - 70.9 Wadt et al. 2008
El Tigre Reserve, Beni, Bolivia 3.00 - 107.5 Zuidema and Boot 2002
El Sena Reserve, Pando, Bolivia 2.40 - 126.9 Zuidema and Boot 2002
*DBH Diameter at breast height measured at 1.3 m above ground level.

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Table 6  Fruit and seed production of Brazil nut trees in natural forests
Local DBH Fruit production Seed production References
−1 −1 −1
(Fruit ­tree ) (kg ­ha ) (kg ­tree )

Pinkaiti Reserve, Pará, Brazil - 184.3 - - Baider 2000

RESEX Cajari, Amapá, Brazil – Capoeira 57 9 - - Guedes et al. 2014
RESEX Cajari, Amapá, Brazil – Forest 112 22 - - Guedes et al. 2014
Chico Mendes Reserve, Acre, Brazil - 65.5 - 9.7 Kainer et al. 2006
Chico Mendes Reserve, Acre, Brazil - 72.0 - 10.07 Kainer et al. 2006
Chico Mendes Reserve, Acre, Brazil - 66.2 12.5 9.3 Kainer et al. 2007
Alto Cajari—Forest, Amapá, Brazil - 165.0 - - Neves et al. 2015
Alto Cajaria—Capoeira, Amapá, Brazil - 62.0 - - Neves et al. 2015
Tahuamanu Reserve, Madre de Dios, Peru - - 15.0 17.14 Nunes et al. 2012
Tambopata Reserve, Madre de Dios, Peru - - 14.3 35.7 Nunes et al. 2012
Kayapó land, Pará, Brazil - 207.0 - Ribeiro et al. 2014
Madre de Dios, Peru 125 218.5 - 39.4 Rockwell et al. 2015
Plato Almeida, Pará, Brazil 122 29 - - Salomão et al. 2006
São João Baliza – Roraima 65.9 24.8 32.7 4.3 Tonini et al. 2008
Caracaraí – Roraima 74.6 18.6 8.48 3.8 Tonini et al. 2008
Caracaraí-Itã, Roraima, Brazil 82.7 - 34.8 - Tonini et al. 2014
Caracaraí-Cujubim, Roraima, Brazil 118.8 - 116.18 - Tonini et al. 2014
São João da Baliza, Roraima, Brazil 65.9 - 16 - Tonini et al. 2014
RESEX Chico Mendes, Acre, Brazil 86.1 - - 10.28 Wadt et al. 2005
Pindamonhangaba, Acre, Brazil - 86.5 - - Wadt et al. 2008
Filipinas, Acre, Brazil - 79.6 - - Wadt et al. 2008
Beni, Bolivia 184 - - - Zuidema 2003
Pando, Bolivia 139 - - - Zuidema 2003

*DBH Diameter at breast height measured at 1.3 m above ground level.

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Table 7  Brazil nut under different plantation systems within (Northern Brazil and Peru) and outside the Amazon region (Southeastern Brazil)
Region Plantation Age Spacing Height DBH MAI Survival References
system
DBH Height
(years) (m) (m) (cm) (cm ­ano ) (m ­ano−1) (%)
−1

Lavras, MG, Brazil Agroforestry 16 3×3 12 17.4 83.5 Caetano 2012

Manaus, AM, Brazil Agroforestry 12 12 × 12 20.9 38 3.16 1.74 78 Costa et al. 2009
Manaus, AM, Brazil RAD mono- 8 2.5 × 1.5 11.47 9.57 1.20 1.43 - Costa et al. 2015
culture
West Amazon, AC, Brazil Enrichment 5 5×5 2.85 1.51 0.53 10 d’Oliveira 2000
Manaus, AM, Brazil Monoculture 10 3×3 Fernandes Alencar 1993
Nova Califórnia, RO, Brazil Agroforestry 16.5 10 × 4 19.9 36.1 1.39 1.5 69.4 Condé et al. 2013
Cantá, RR, Brazil Mixed 10 - 14 26 - - 98.6 Ferreira and Tonini 2009
Madre de Dios, Peru Silvopastures 3 10 × 10 4.1 4.8 98.1 Frank and Cruz 1995
Madre de Dios, Peru Silvopastures 2 15 × 15 2.1 2 97.5 Frank and Cruz 1996
Porto Velho, RO, Brazil Agroforestry 25 - 20.6 41 - - - Locatelli et al. 2013
Nova Mamoré, RO, Brazil Monoculture 35 - 23.9 44 - - - Locatelli et al. 2013
Machadinho do Oeste, RO, Brazil Monoculture 28 12 × 12 28.3 60.09 86 Locatelli et al. 2015
Machadinho do Oeste, RO, Brazil Mixed 28 12 × 12 28.6 59.75 80 Locatelli et al. 2015
El Tigre, Bolivia—Close canopy Enrichment ~3 10 × 10 0.68 86.5 Peña-Claros et al. 2002
El Tigre, Bolivia—Open canopy Enrichment ~3 10 × 10 3.86 98.4 Peña-Claros et al. 2002
Oriximiná, PA, Brazil RAD 19 - 14.7 19.4 1.02 0.77 Salomão et al. 2006
Manaus, AM, Brazil Agroforestry ~ 10 × 10 5.4 8.4 Schroth et al. 1999
Manaus, AM, Brazil Agroforestry 7 10.5 × 10.5 20.6 3.0 97 Schroth et al. 2015
Manacapuru, AM, Brazil Agroforestry 10 - - - 3.1 1.6 - Soares et al. 2004
Manacapuru, AM, Brazil Enrichment 10 - - - 1.8 1.3 - Soares et al. 2005
Saracá-Taquera, PA, Brazil RAD mixed 30 10 × 50 15.3 19 0.96 0.78 Tonini et al. 2008
Machadinho do Oeste, RO, Brazil Mixed 10 12 × 12 12.95 21 3.2 2.23 89.63 Vieira et al. 1998
Machadinho do Oeste, RO, Brazil Monoculture 10 12 × 12 12.25 22 3.1 2.13 95.38 Vieira et al. 1999
Belterra, PA, Brazil Monoculture 6.5 3×3 7.5 12 1.8 1.2 66.7 Yared et al. 1988
Manaus, AM, Brazil Monoculture 40 10 × 10 23.9 69 - - - Yared et al. 1993
Porto Velho, RO, Brazil Monoculture 30 - 22 40 - - - Yared et al. 1993
Macapá, AP, Brazil Monoculture 30 10 × 10 20.4 45 - - - Yared et al. 1993
Tomé-Açu, PA, Brazil Monoculture 49 20 × 20 20.6 80 - - - Yared et al. 1993
Cláudia, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Santa Carmem, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
São José do Rio Claro, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Rosário Oeste, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Sinop, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Terra Nova do Norte, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Nova Bandeirantes, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Alta Floresta, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Juína, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
Paranaíta, MT, Brazil Monoculture - - - - - - - Baldoni et al. 2019
*DBH Diameter at breast height measured at 1.30 m above ground level.

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106 Current Forestry Reports (2022) 8:90–110

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of Higher Education Personnel (CAPES – Brazil- Finance Code 001), 2013. https://​doi.​org/​10.​1111/​geb.​12092.
and the National Council for Scientific and Technological Development 13. Lutz JA, Furniss TJ, Johnson DJ, Davies SJ, Allen D, Alonso
(CNPq, Brazil) for their financial support for the research. Many thanks A, et al. Global importance of large-diameter trees. Global Ecol
to Empresa Agropecuária Aruanã S.A. for its support and cooperation. Biogeogr. 2018. https://​doi.​org/​10.​1111/​geb.​12747.
J.F.C Gonçalves, F.A. Lobo, M. V. Ramos, Hector H. F. Koolen, and P. 14. Gratani L. Plant phenotypic plasticity in response to environmen-
Mazzafera thank (CNPq-Brazil) for PQ research fellowships. tal factors. Adv Bot. 2014. https://​doi.​org/​10.​1155/​2014/​208747.
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Authors and Affiliations

Karen Cristina Pires da Costa1 · José Francisco de Carvalho Gonçalves2 · Alexandre Leão Gonçalves2 ·
Adamir da Rocha Nina Junior3 · Roberto Kirmayr Jaquetti2 · Vinícius Fernandes de Souza2 · Josiane
Celerino de Carvalho2 · Andreia Varmes Fernandes2 · Joelma Keith Rodrigues2 · Gleisson de Oliveira Nascimento4 ·
Lúcia Helena de O. Wadt5 · Karen A. Kainer6 · Roberval Monteiro Bezerra de Lima7 · Flávia Camila Schimpl8 ·
Jéssica Pereira de Souza2 · Sabrina Silva de Oliveira2 · Hellen Thaís da Silva Miléo2 · Diego P. Souza2 · Ana Claudia
Lopes da Silva2 · Heloisa Massaco Ito Nascimento2 · Jair Max Furtunato Maia9 · Francisco de Almeida Lobo10 ·
Paulo Mazzafera11 · Marcio Viana Ramos12 · Hector Henrique Ferreira Koolen13 · Ronaldo Ribeiro de Morais7 ·
Karina Martins14 · Niwton Leal Filho15 · Henrique Eduardo Mendonça Nascimento15 · Katharine Duarte Gonçalves2 ·
Yasmin Verçosa Kramer2 · Giordane Augusto Martins15 · Marcelo O. Rodrigues16

Karen Cristina Pires da Costa Diego P. Souza

[email protected] [email protected]
Alexandre Leão Gonçalves Ana Claudia Lopes da Silva
[email protected] [email protected]
Adamir da Rocha Nina Junior Heloisa Massaco Ito Nascimento
[email protected] [email protected]
Roberto Kirmayr Jaquetti Jair Max Furtunato Maia
[email protected] [email protected]
Vinícius Fernandes de Souza Francisco de Almeida Lobo
[email protected] [email protected]
Josiane Celerino de Carvalho Paulo Mazzafera
[email protected] [email protected]
Andreia Varmes Fernandes Marcio Viana Ramos
[email protected] [email protected]
Joelma Keith Rodrigues Hector Henrique Ferreira Koolen
[email protected] [email protected]
Gleisson de Oliveira Nascimento Ronaldo Ribeiro de Morais
[email protected] [email protected]
Lúcia Helena de O. Wadt Karina Martins
[email protected] [email protected]
Karen A. Kainer Niwton Leal Filho
[email protected] [email protected]
Roberval Monteiro Bezerra de Lima Henrique Eduardo Mendonça Nascimento
[email protected] [email protected]
Flávia Camila Schimpl Katharine Duarte Gonçalves
[email protected] [email protected]
Jéssica Pereira de Souza Yasmin Verçosa Kramer
[email protected] [email protected]
Sabrina Silva de Oliveira Giordane Augusto Martins
[email protected] [email protected]
Hellen Thaís da Silva Miléo Marcelo O. Rodrigues
[email protected] [email protected]

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110 Current Forestry Reports (2022) 8:90–110

1 9
Faculty of Agricultural Sciences, Institute of Studies University of State of Amazonas (UEA), Av. Djalma Batista,
in Agrarian and Regional Development – IEDAR, Federal Manaus, AM 247069.050‑010, Brazil
University of South and Southeast of Pará (UNIFESSPA), 10
Faculty of Agronomy and Zootechny, Federal University
Folha 31, Quadra 07, Nova Marabá, Marabá, PA 68507‑590,
of Mato Grosso, Mato Grosso (UFMT), Cuiabá,
Brazil
MT 78060‑900, Brazil
2
Laboratory of Plant Physiology and Biochemistry, National 11
Department of Plant Biology, Institute of Biology,
Institute for Amazonian Research (MCTI-INPA), Manaus,
University of Campinas, Campinas, Brazil and Department
Amazonas, Brazil
of Crop Science, Luiz de Queiroz College of Agriculture –
3
Federal Institute of Education, Science and Technology ESALQ/University of São Paulo, Piracicaba, Brazil
of Amazonas (IFAM) – Campus Humaitá, BR230 Highway, 12
Federal University of Ceara, Ceará, Brasil (UFC), Ceará,
km 07, Humaitá, AM 69.800‑000, Brazil
Brazil
4
Multidisciplinary Center, Federal University of Acre 13
Metabolomics and Mass Spectrometry Research
(UFAC), Cruzeiro do Sul, São Paulo, AC 69980‑000, Brazil
Group, Amazonas State University (UEA), Manaus,
5
Centro de Pesquisa Agroflorestal de Rondônia, Amazonas 690065‑130, Brazil
Brazilian Agricultural Research Corporation (Embrapa), 14
Centro de Ciências Humanas E Biológicas, Departamento de
BR 364, km 5,5, Caixa Postal 127, Porto Velho,
Biologia, Universidade Federal de São Carlos (UFSCar), SP
Rondônia CEP 76815‑800, Brazil
264, km 110, Itinga, Sorocaba, SP 18052‑780, Brazil
6
School of Forest, Fisheries, and Geomatics Sciences, 15
Campus III – V8, Coordenação de Biodiversidade,
and Center for Latin American Studies, University of Florida,
Instituto Nacional de Pesquisas da Amazônia,
P.O. Box 110410, Gainesville, FL 32611‑0410, USA
Av. André Araújo, 2.936, Petrópolis, Brasil, Manaus,
7
Embrapa Western Amazon, Research and Development, Rod AM Cx. Postal 2223 – CEP 69080‑971, Brazil
AM 010 km 29, Manaus, Amazonas CEP 69010‑970, Brazil 16
LIMA‑Laboratório de Inorgânica E Materiais, Universidade
8
Federal Institute of Education, Science and Technology de Brasília – UNB, Campus Universitário Darcy Ribeiro,
of Amazonas (IFAM) – Campus - Presidente Figueiredo, P. O. Box 4478, Brasília, Distrito Federal 70904‑970, Brazil
Amazonas, Brazil

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