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                    UNIVERSIDADE FEDERAL DE ALAGOAS
INSTITUTO DE CIÊNCIAS BIOLÓGICAS E DA SAÚDE
Programa de Pós-Graduação em Diversidade Biológica e Conservação nos
Trópicos

ROBERTA DE ALMEIDA CAETANO

INTERAÇÕES ENTRE O USO DE RECURSOS FLORESTAIS MADEIREIROS E
ALIMENTÍCIOS E SUAS IMPLICAÇÕES PARA A CONSERVAÇÃO BIOCULTURAL

MACEIÓ - ALAGOAS
Março/2024

ROBERTA DE ALMEIDA CAETANO

INTERAÇÕES ENTRE O USO DE RECURSOS FLORESTAIS MADEIREIROS E
ALIMENTÍCIOS E SUAS IMPLICAÇÕES PARA A CONSERVAÇÃO BIOCULTURAL

Tese apresentada ao Programa de PósGraduação
Conservação

em

Diversidade

nos

Trópicos,

Biológica

e

Instituto

de

Ciências Biológicas e da Saúde. Universidade
Federal de Alagoas, como requisito para
obtenção do título de Doutora em CIÊNCIAS
BIOLÓGICAS, na área da Biodiversidade.

Orientadora: Profa. Dra. Patrícia Muniz de
Medeiros
Coorientadores: Prof. Dr. Rafael Ricardo
Vasconcelos da Silva
Profa. Dra. Adriana Rosa Carvalho

MACEIÓ - ALAGOAS
Março/2024

Catalogação na Fonte
Universidade Federal de Alagoas
Biblioteca Central
Divisão de Tratamento Técnico
Bibliotecário: Antonia Izabel da Silva Meyer – CRB-4 - 1558
C128i

Caetano, Roberta de Almeida.
Interações entre o uso de recursos florestais madeireiros e alimentícios e suas
implicações para a conservação biocultural / Roberta de Almeida Caetano. – 2024.
142 f. : il.
Orientadora: Patrícia Muniz de Medeiros.
Co-orientador: Rafael Ricardo Vasconcelos da Silva.
Co-orientadora: Adriana Rosa Carvalho.
Tese (Doutorado em Ciências Biológicas) – Universidade Federal de Alagoas.
Instituto de Ciências Biológicas e da Saúde. Programa de Pós-Graduação em
Diversidade Biológica e Conservação nos Trópicos. Maceió, 2024.
Bibliografia: f.126.
Apêndices: f.127-142.
1. Biodiversidade. 2. Etnobiologia. 3. Conservação biocultural. 4. Produtos da
sociobiodiversidade. 5. Plantas alimentícias. 6. Produtos florestais não madereiros.
I. Título.
CDU: 504

Folha de aprovação
Roberta de Almeida Caetano
Interações entre o uso de recursos florestais madeireiros e alimentícios
e suas implicações para a conservação biocultural
Tese apresentada ao Programa de Pós-Graduação
em Diversidade Biológica e Conservação nos
Trópicos, Instituto de Ciências Biológicas e da
Saúde. Universidade Federal de Alagoas, como
requisito para obtenção do título de Doutor(a) em
CIÊNCIAS
BIOLÓGICAS
na
área
da
Biodiversidade.

Tese aprovada em 27 de março de 2024.

Dr.(a) Patrícia Muniz de Medeiros/UFAL
(orientadora)

Dra. Adriana Rosa Carvalho
(Coorientadora)

Dr. Rafael Ricardo Vasconcelos da Silva
(Coorientador)

Dr. (a) Taline Cristina da Silva

Marcelo Alves Ramos

Dr. (a) Gilberto Costa Justino

Dr. (a) Guilherme Ramos Demétrio Ferreira

MACEIÓ - AL
Março/ 2024

Dr. (a)

DEDICATÓRIA

À minha amada mãe, que com sua ternura, apoiou-me e guiou-me pelos
caminhos que me conduziram até aqui e
À minha querida avó materna (in memoriam).

AGRADECIMENTOS

Chegar ao fim do doutorado me remete ao que posso chamar de prelúdio dessa
jornada, que envolve uma menina cercada por plantas e animais. A menina que
“conversava” com as plantas, que brincava com os soldadinhos e girinos, que adorava
ficar à sombra de árvores como Timbaúba. Curiosa, observava pacientemente o
movimento das borboletas nas flores de hibisco e que, incansavelmente tentava capturálas com as mãos, até conseguir. Aquela menina nem imaginava para onde aquele
contexto a levaria, que iria se encantar ainda mais pela biodiversidade e estudaria para
conhecê-la e conservá-la. Hoje, essa mulher que aqui escreve está contente pela direção
que, deliberadamente, escolheu para a sua vida a anos atrás. Por ter se mantido firme,
mesmo quando alguns não a encorajaram.
E durante essa longa jornada cheia de encantamentos e desafios, algumas
pessoas cruzaram o meu caminho e deixaram suas marcas na forma de apoio intelectual,
emocional, físico e financeiro. A elas, quero expressar a minha gratidão.
A todo corpo docente das Ciências Biológicas e do programa de pós-graduação
em Diversidade Biológica e Conservação nos Trópicos da Universidade Federal de
Alagoas, ao qual tenho imensa gratidão, pois foi nessa instituição que iniciei minha
formação acadêmica (2010 a 2015) e para qual retornei para o doutorado, em 2020.
Agradeço a minha orientadora, Patrícia Medeiros, pelos ensinamentos, conselhos
e incentivos à minha formação acadêmica e pessoal, desde o mestrado. Uma mulher
genial, a quem admiro e respeito por ser empática e pela potência que é como mulher,
cientista, professora e orientadora.
Aos meus coorientadores Rafael Silva e Adriana Carvalho, que com suas skills
deram um plus à minha pesquisa de doutorado ao longo desses anos.
E como uma extensa pesquisa de campo não se faz sem recursos financeiros,
agradeço às duas instituições que apoiaram a minha pesquisa. Ao CNPq pela bolsa de
doutorado a mim concedida e pela taxa de bancada, que foi essencial para aquisição de
alguns materiais para a minha pesquisa. Ao Fundo Brasileiro para a Biodiversidade,

Instituto Humanize e a Eurofins Foundations que, partir do ano de 2022, supriu a maior
parte dos custos de equipamentos, materiais e manutenção de equipe em campo.
A todos os colegas do Laboratório de Ecologia, Conservação e Evolução
Biocultural, pelo suporte durante a minha pesquisa de campo. Uma equipe grande,
animada e eficiente, que tornou essa parte do trabalho mais leve. Agradeço
especialmente a: Jonathan, Fernando, Gabriela, Déborah, Danúbia, Emilly, Daniel, Rita,
Jorge, Jonas, Alisson, Saulo, Álvaro, Alexsander, Élida e seu esposo, Roberto. Também
agradeço a Isabelly, Cícero e João, meus eficientes e dedicados assistentes de campo
durante a etapa das entrevistas.
A Richard Poian, que com muito entusiasmo, participou de várias etapas desse
estudo e até recrutou voluntários para participarem da montagem do herbário de campo.
Gratidão a todos.
Às pessoas da comunidade de Retiro, pelo acolhimento, compartilhamento de
saberes e experiências. De modo especial, agradeço ao Sr. Zé Antônio e Dona das
Dores, que acolheram a mim e a minha equipe como a familiares e nos assistiram durante
todo o desenvolvimento da pesquisa.
À Élida Santos, pela forte parceria em várias atividades do doutorado e pelo apoio
emocional. Dividimos conhecimentos, perrengues e muitos sanduiches em campo.
À Emilly Guedes, pelo suporte nos trabalhos de campo e de escritório, enquanto
bolsista de iniciação científica, e pela gentil companhia e conversas significativas extraacadêmicas.
Às amigas que acompanham e apoiam a minha trajetória acadêmica,
particularmente: Risoneide, Regina, Maíra, Ylana, Jadla, Duyane e Michelle. Obrigada!
Ao querido amigo Bruno Oliveira, por ser um ouvinte ativo dos meus
encantamentos e anseios ao longo do curso e, pelos ótimos conselhos. Agradeço
grandemente.
Agradeço à minha família, pelo apoio e compreensão ao longo desse anos. Por
todo amor emanado da minha mãe para me tranquilizar e animar, principalmente quando,
em meio a crises de ansiedade, eu só conseguia enxergar as “pedras” no meu caminho.
A Deus, pela força e coragem para ousar sonhar e alçar voo nesse vasto mundo
da Ciência.

“Desde que me entendo por gente, andava
mais a minha vó na mata. A gente sempre
colhia cambuí, coco de piaçava, araçá,
jenipapo, caju, ingá e massaranduba. A
mata sempre foi importante pra gente. Aqui
sempre tinha muita água, árvore e muito
passarinho. Mas depois que começaram a
desmatar, começou a sumir as caças, as
frutas... Aí ficou um pouco desvalorizada a
nossa mata”.

AJD, morador da comunidade de Retiro,
Piaçabuçu - Alagoas.

RESUMO
As plantas alimentícias silvestres lenhosas têm alto potencial para a segurança alimentar
e nutricional, mas podem estar sofrendo pressão do uso madeireiro. Por isso, é
importante entendermos se em um contexto de interação de usos, a importância do uso
alimentício pode atuar protegendo as plantas lenhosas de usos destrutivos. Esta tese
está dividida em dois capítulos, os quais convergem sobre a conservação de plantas
lenhosas a partir da interação entre usos distintos, tendo um desses usos um efeito de
proteção. No primeiro capítulo, por meio de uma revisão sistemática, buscamos identificar
espécies prioritárias para a conservação biocultural que, ao mesmo tempo, são
estratégicas para a manutenção da segurança alimentar e nutricional (em termos de
composição de micro e macronutrientes) e potencialmente ameaçadas pelos seus
múltiplos usos madeireiros. Identificamos 42 espécies com aplicação em todas as
categorias de uso madeireiro analisadas neste estudo, sendo consideradas versáteis. A
comparação dos dados etnobiológicos e nutricionais revelou oito espécies versáteis para
as quais estava disponível informação sobre a composição nutricional, entre as quais,
três se destacaram em termos de teor de macronutrientes, nomeadamente Anacardium
occidentale L., Bauhinia cheilantha (Bong.) Steud., e Eugenia pyriformis Cambess.
Encontramos que, muitas espécies versáteis classificadas como ameaçadas ou em
declínio não têm sido foco de estudos nutricionais, o que sinaliza a necessidade de
maiores esforços de investigação nutricional. Também destacamos a necessidade de
investigar se a importância alimentícia exerce algum efeito protetor sobre estas espécies,
reduzindo a pressão do uso madeireiro (hipótese de proteção). No segundo capítulo,
conduzimos um estudo de campo em uma comunidade rural dentro da vegetação de
Restinga, no nordeste do Brasil. Nesse capítulo buscamos preencher uma lacuna na
literatura sobre como a importância doméstica e comercial das plantas alimentícias
lenhosas pode protegê-las da exploração madeireira. Para esse fim, examinamos a
hipótese da proteção a partir de duas perspectivas distintas (proteção generalizada e
proteção direcionada a espécies chave). Nossas descobertas sugerem que não há efeito
protetivo proporcional ao uso alimentício das espécies. No entanto, o uso doméstico de
espécies-chave para alimentação exibiu um forte efeito protetivo. A disponibilidade e a
qualidade percebidas surgiram como preditores importantes para a exploração da
madeira. Assim, discutimos estratégias de conservação biocultural que aumentem a
importância alimentícia das plantas para garantir a sua proteção, juntamente com
medidas para espécies lenhosas não comestíveis sob maior pressão de uso. Por meio
dessa tese, descobrimos que espécies alimentícias com potencial nutricional podem
estar sofrendo pressão de uso devido a seu emprego madeireiro. No entanto, espécies
com alta importância têm o uso madeireiro reduzido, controlando o efeito da qualidade
madeireira e a disponibilidade percebidas. Portanto, é necessária a realização do estudo
ecológico dessas espécies e a adoção de medidas de conservação que incluam a
valorização e o uso sustentável dos recursos vegetais.
Palavras-chave: Produtos da sociobiodiversidade. Usos madeireiros. Interação de usos.
Conservação biocultural. Manejo tradicional. Etnobiologia

Abstract

Wild woody food plants have high potential for food and nutritional security but may be
under pressure from wood use. Therefore, it is important for us to understand whether in
a context of interaction of uses, the importance of food use can act to protect woody plants
from destructive uses. This thesis is divided into two chapters, which converge on the
conservation of woody plants through the interaction between different uses, with one of
these uses having a protective effect. In the first chapter, through a systematic review, we
sought to identify priority species for biocultural conservation that are also strategic for
maintaining food and nutritional security (in terms of micro and macronutrient composition)
and potentially threatened by their multiple wood uses. We identified 42 species with
applications in all categories of wood use analyzed in this study, considered versatile.
Comparison of ethnobiological and nutritional data revealed eight versatile species for
which nutritional composition information was available, among which three stood out in
terms of macronutrient content, namely Anacardium occidentale L., Bauhinia cheilantha
(Bong.) Steud., and Eugenia pyriformis Cambess. We found that many versatile species
classified as threatened or declining have not been the focus of nutritional studies,
signaling the need for greater nutritional research efforts. We also highlight the need to
investigate whether the food importance exerts any protective effect on these species,
reducing the pressure of wood use (protection hypothesis). In the second chapter, we
conducted a field study in a rural community within the Restinga vegetation, in
northeastern Brazil. In this chapter, we sought to fill a gap in the literature on how the
domestic and commercial importance of woody food plants can protect them from wood
exploitation. To this end, we examined the protection hypothesis from two different
perspectives (generalized protection and protection targeted at key species). Our findings
suggest that there is no protective effect proportional to the food use of the species.
However, the domestic use of key food species exhibited a strong protective effect.
Perceived availability and quality emerged as important predictors for wood exploitation.
Thus, we discuss biocultural conservation strategies that increase the food importance of
plants to ensure their protection, along with measures for non-edible woody species under
greater pressure of use. Through this thesis, we discovered that food species with
nutritional potential may be under pressure from use due to their wood employment.
However, species with high importance have reduced wood use, controlling for perceived
wood quality and availability. Therefore, ecological study of these species and adoption
of conservation measures that include the valorization and sustainable use of plant
resources are necessary.

Keyword: Sociobiodiversity products. Wood uses. Use interaction. Biocultural
conservation. Traditional management. Ethnobiology

LISTA DE FIGURAS
Artigo 1

Figure 1. Flowchart with systematic review search and screening steps…………………52
Figure 2. Flowchart with search and screening stages of the systematic review on
nutritional aspects………………………………………………………………………………53
Figure 3. Distribution of studies on food and/or timber plants extracted in this systematic
review. Note that 80 studies distributed throughout Brazil are expressed (two more than
the number mentioned in the text - 78). This occurred because two studies were carried
out in areas of two different states (Pernambuco and Paraiba)…………………………….54
Figure 4. Number of woody food and/or timber species found during the stages of the
ethnobiological and nutritional reviews. Edited in Canva (free version)…………………..66
Figure 5. Commercial fruit pulps of the species (A) S. mombin, (B) A. occidentale and (C)
G. americana. Edited in Canva (free version)………………………………………………..70
Artigo 2
Figure 1. Geographic Location of the Retiro Community in the Municipality of PiaçabuçuAlagoas, Brazil………………………………………………………………………………….85
Figure 2. Stages of ecological and ethnobiological data collection: (1) Participatory
workshop for identifying key species and collection areas; (2) Forest inventory to detect
species co-occurring with key species, accompanied by botanical collection and plant
photography; (3) Checklist-interview utilizing visual stimuli, including photos and
exsiccates (field herbarium). Image edited using Canva (free version)…………………..88
Figure 3. Information collected using a Likert scale on the variables considered in this
study……………………………………………………………………………………………..94
Figure 4. Widespread species model and key species-based model with their variables
and respective measures………………………………………………………………………97
Figure 5. Impact of Predictor Parameters (quality, availability, domestic food use,
commercial food use, domestic use of key species, and commercial use of key species)
on domestic wood utilization of wild edible plants. Left: widespread protection model.
Right: key species-based protection model. The central circles indicate the median
coefficient estimates of the associations, and the horizontal lines delineate the 95%
credibility intervals. The parameter coefficient estimates are plotted along the x-axis, while
the predictor levels are represented on the y-axis. The vertical line intersecting the zero
point on the x-axis (indicating no effect) facilitates comparison of the sizes of positive,
negative, and null effect coefficients. In the parameter level grouping, non-overlapping

horizontal bars denote significant differences. Horizontal bars intersecting the zero line on
the x-axis signify a non-significant effect……………………………………………………..99
Figure 6. Hypothetical example of a trade-off between availability and quality explaining
fuelwood use. In a simplified scenario where these are the only predictors of plant use
within the fuelwood category, the most utilized species would be those exhibiting the
highest trade-offs between availability and quality (represented by the blue dots in the
right and left graphs). When considering the interaction with the food use-category under
the key-species based protection model, the use of wood species with low to intermediate
food importance would be proportional to the trade-off between quality and availability
(graph on the right). However, for species that are considered key food plants (indicated
by the dark green dot), their utilization for fuelwood would be less than what is predicted
by their quality and availability alone………………………………………………………..104

LISTA DE TABELAS
Artigo 1

Table 1. Regions and ecosystems in Brazil where the studies were carried out………….59
Table 2. Woody food plants considered versatile because they are used in the three
categories of wood uses (fuel, construction and technology). Data from the systematic
review of Brazilian ethnobiological studies…………………………………………………..60
Table 3. Nutrients and number of species contain them……………………………………62
Table 4. Macronutrient composition of parts of versatile food woody plants for wood uses.
Data from the systematic review of nutritional studies………………………………………67
Table 5. Composition of mineral content (mg/100g) in parts of versatile food woody plants
for wood uses. Data from the systematic review of nutritional studies…………………….68

Artigo 2
Table 1. Plants that were part of the checklist interview, their citation percentages (general
and by use), and occurrence areas………………………………………………………….115
Table 2. Species recognized for wood purposes and their respective uses in each
category………………………………………………………………………………………..117
Frame 1. Monthly household income and the corresponding percentage of interviewees
who possess it…………………………………………………………………………………..87
Frame 2. Responses from interviewees indicating the protection of key species regarding
wood uses……………………………………………………………………………………..100
Appendix 1. Multicollinearity diagnosis and model average for the widespread protection
model and the key species-based protection model……………………………………….140

SUMÁRIO
1 Apresentação……………………………………………………………………................15
Referências................................................................................................................... ..19
2 Revisão da literatura……………………………………………………………...............22
2.1 Impactos da extração de produtos florestais madeireiros e não
madeireiros…………………………………………………………………….......................20
2.1.1 Uso e conservação de produtos florestais não-madeireiros...................................22
2.2 Evidências do papel de proteção associadas às plantas alimentícias...............27
2.2.1 Proteção
de
plantas
a
partir
do
manejo:
evidências
da
domesticação..................................................................................................................27
2.2.2 Evidências da conservação através do uso de PFNM...........................................29
2.2.3 Evidências da hipótese da proteção.......................................................................32
2.3 Uso e conservação de produtos florestais madeireiros ………………………….33
2.3.1 Critérios de seleção de plantas madeireiras...........................................................36
Referências.....................................................................................................................39
3 Wild food plants with the potential to improve food andnutrition security may be
threatened by timber extraction: a systematic review of the brazilian
context……………………………………………………………………………………….….46
Referências…………………………………………………………………………………......75
4 Conservation of wild food plants from wood exploitation: evidence supporting
the
protection
hypothesis
in
northeastern
Brazil…………………………………………....................................................................80
Referências...................................................................................................................122
5 Conclusões............................................................................................................... 126
6 Apêndices................................................................................................................. 127

15

1 APRESENTAÇÃO

As plantas alimentícias silvestres têm destacada importância ao redor do mundo
do ponto de vista socioeconômico e nutricional (JACOB et al., 2020). Essas plantas
podem desempenhar um papel importante em uma dieta saudável como fonte
alternativa de minerais, vitaminas e antioxidantes, bem como macronutrientes, como
carboidratos, proteínas e lipídios. Por isso, o seu consumo tem sido recomendado
como parte de estratégias globais para combater a desnutrição (HUNTER et al.,
2019), diversificar a dieta humana e os sistemas alimentares como um todo
(BALDERMANN et al., 2016), e gerar renda para pequenos agricultores e extrativistas
(DELANG, 2014). Além disso, a extração de partes comestíveis de plantas é, na
maioria dos casos, menos prejudicial que a extração de madeira, sendo propícia à
aplicação de estratégias de manejo e uso sustentáveis (GAOUE et al., 2016).
No entanto, apesar do grande potencial das plantas alimentícias silvestres para
o manejo sustentável, muitas dessas espécies têm múltiplas aplicações, incluindo a
extração de madeira, que é uma das principais causas do declínio das populações
vegetais (BRUSCHI et al., 2014; ROS-TONEN, 2000; STANLEY et al., 2012). Esta
situação é particularmente preocupante para as plantas alimentícias de alta
versatilidade para os usos madeireiros, pois podem estar enfrentando maior pressão
de uso. E a extinção de espécies representa uma perda irreversível da biodiversidade
e do patrimônio cultural. Para evitar esse problema, é essencial desenvolver
estratégias a partir de uma perspectiva de conservação biocultural, ou seja, buscando
combinar ações de conservação ao conhecimento e uso dos recursos naturais para
reduzir a perda da diversidade biológica e cultural (Gavin et al., 2015).
Nesse sentido, conservar o amplo uso dos recursos alimentícios silvestres pode
ser estratégico porque as plantas silvestres são encontradas em muitas comunidades
locais que ainda enfrentam desafios relacionados à: fome, disponibilidade de
alimentos, pouca diversificação da dieta e a eventos climáticos que impactam a
produção de cultivos (NASCIMENTO et al., 2012, 2011; JACOB et al., 2020).
No Brasil, vários estudos registraram espécies lenhosas com potencial
nutricional (NASCIMENTO et al., 2012, 2011; JACOB et al., 2020; NUNES et al.,
2012). No entanto, a maioria dos estudos não conseguiu analisar até que ponto as
populações de plantas são ameaçadas por outros usos que não o uso alimentício.

16

Compreender a sobreposição dos usos das plantas lenhosas é importante porque
estratégias de conservação que abrangem diferentes usos dos recursos vegetais
podem contribuir grandemente para a segurança alimentar e nutricional.
Além disso, é esperado que o uso de plantas alimentícias seja um provável
candidato a conferir um efeito de proteção contra os usos madeireiros. E, dada a
natureza especializada do uso alimentício, sua importância na dieta humana e seu
potencial econômico, é possível que as comunidades locais protejam estas plantas de
danos irreversíveis, como os causados pela maioria dos usos madeireiros, para os
quais estão disponíveis espécies vegetais alternativas devido à sua natureza
generalista. Inclusive, essa hipótese é testada nessa tese e será apresentada mais
adiante.
No contexto de diversas comunidades sul-americanas, os usos especializados
são definidos por uma gama mais restrita de plantas adequadas que atendem a
requisitos específicos, com a premissa de especialização reforçada por observações
de que a disponibilidade de plantas exerce pouca ou nenhuma influência sobre tais
usos (RIBEIRO et al., 2014; SOLDATI et al., 2016). Em contraste, os usos generalistas
acomodam um espectro mais amplo de espécies, sendo as mais utilizadas, muitas
vezes, as mais disponíveis, como acontece com os usos madeireiros (GONÇALVES
et al., 2016).
Embora a qualidade também possa ser um importante preditor da importância
das plantas para usos generalistas (CARDOSO et al. 2015), a natureza generalista do
uso da madeira é apoiada por estudos que investigam a hipótese da aparência
(disponibilidade), que postula uma correlação entre a disponibilidade ambiental e a
utilização das espécies (GONÇALVES et al., 2016; LUCENA et al., 2013; LUCENA et
al., 2007). Portanto, para usos generalistas, há outras opções igualmente vantajosas,
o que permitiria às pessoas pouparem certas espécies para usos especializados,
como o uso alimentício, para a qual menos espécies podem atuar como substitutas.
Diante desse cenário, a hipótese da proteção foi criada para considerar que
plantas de alta importância numa categoria de uso mais especializada (como o uso
medicinal) podem experimentar uma redução na sua utilidade para outras categorias
de natureza mais generalista, como o uso madeireiro (SILVA et al., 2021). Um estudo
que testou essa hipótese demonstrou um efeito pequeno, mas significativo da
importância do uso medicinal sobre o uso madeireiro, indicando evidências favoráveis

17

à hipótese, uma vez que as plantas medicinais de maior importância são utilizadas
com menos frequência para fins madeireiros. Além disso, foi observado que o efeito
protetivo é provavelmente mais forte nas espécies de alta importância medicinal.
Nessa tese, as espécies de alta importância, são denominadas de espécies-chave,
sendo definidas como espécies alimentícias silvestres de alta importância regional
para o consumo e geração de renda.
Investigar como a interação de categorias de uso distintas pode interferir na
conservação das espécies vegetais tem se mostrado uma abordagem promissora
para entender sobre o efeito de proteção em sistemas socioecológicos (SILVA et al.,
2021). Porém, tem sido pouco explorada em estudos dessa natureza, obscurecendo
a nossa compreensão sobre como o valor de uma planta para um determinado
propósito pode protegê-la de outros usos mais prejudiciais.
É essencial entender se existem interações de uso e como elas se comportam
do ponto de vista de conservação, porque, embora o uso madeireiro associado possa
representar uma ameaça maior às espécies vegetais do que somente o uso
alimentício; por outro lado, a planta que tem uso alimentício, pode ser menos usada
para fins madeireiros pelos povos locais do que uma planta de mesma qualidade e
disponibilidade e que não tenha uso alimentício associado.
Com base nessas premissas, esta tese está dividida em dois capítulos, os quais
convergem sobre a conservação de plantas lenhosas a partir da interação entre usos
distintos, tendo um desses usos um efeito de proteção.
O primeiro capítulo é produto de duas revisões sistemáticas que se
complementam: uma de natureza etnobiológica e a outra, de natureza nutricional.
Nesta revisão sistemática, buscamos identificar espécies prioritárias para a
conservação biocultural que, ao mesmo tempo, são estratégicas para a manutenção
da segurança alimentar e nutricional (em termos de composição de micro e
macronutrientes) e potencialmente ameaçadas pelos seus múltiplos usos madeireiros.
De modo geral, nesse capítulo identificamos a partir de estudos etnobiológicos, quais
as espécies lenhosas nativas do Brasil utilizadas para propósitos alimentícios são
também versáteis para os usos madeireiros (nas categorias combustível, construção
e tecnologia) e fornecemos um compilado de informações sobre a composição
química dessas espécies, a partir de estudos nutricionais diversos. Além disso,
discutimos estratégias de conservação biocultural para essas espécies e fazemos

18

recomendações para estudos futuros sobre lacunas identificadas nos estudos
etnobiológicos e nutricionais.
O segundo capítulo é um estudo de caso conduzido em uma comunidade rural
dentro da vegetação de Restinga, no nordeste do Brasil. Nesse capítulo buscamos
preencher uma lacuna na literatura sobre como a importância doméstica e comercial
das plantas alimentícias lenhosas pode protegê-las da exploração madeireira. Para
esse fim, examinamos a hipótese da proteção a partir de duas perspectivas distintas
(proteção generalizada e proteção centrada nas espécies chave). Nosso estudo é o
primeiro a testar a hipótese da proteção considerando o uso alimentício como efeito
protetor para espécies lenhosas e contribui com estratégias de conservação
biocultural que buscam ampliar a importância alimentícia das espécies para garantir a
sua proteção, juntamente com medidas para espécies lenhosas não comestíveis sob
maior pressão de uso. Adicionalmente, apresenta alguns desafios e perspectivas do
teste da hipótese da proteção para estudos futuros.

19

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FARIAS LIMA, J. R.; OLIVEIRA, R. S.; ALVES, C. A. B.; JARDIM, J. G.; LUCENA, R.
F. P. Can ecological apparency explain the use of plant species in the semi-arid
depression of Northeastern Brazil? Acta Botanica Brasilica, v. 28, p. 476–483. DOI:
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MEDEIROS, P. M. Can medicinal use protect plant species from wood uses?
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Sustainable in the Less Developed World? A Systematic Review of the Recent
Economic and Ecological Literature. Ethnobiology and Conservation, n. 1, 2012.

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2 REVISÃO DA LITERATURA

2.1 Impactos da extração de produtos florestais madeireiros e não madeireiros
Os Produtos Florestais Não-Madeireiros (PFNM) ou produtos florestais nãolenhosos, podem ser definidos como produtos vegetais ou animais provenientes de
florestas naturais ou manejadas, com exceção da madeira e suas diferentes
finalidades (SOLDATI, ALBUQUERQUE, 2008). No entanto, existe uma questão
terminológica e conceitual associada aos PFNM que pode causar confusão. Na
literatura inglesa a tradução do termo “timber” pode significar “madeira para
construção” ou “viga” e “madeira bruta”. Outro termo utilizado ao tratar desse tema é
“wood”, que em geral, incorpora a madeira em um sentido mais amplo. Em relação à
questão conceitual relacionada às variações nos termos, é possível encontrar estudos
sobre produtos florestais não-madeireiros que incluem produtos de origem madeireira
que, ora utilizam o termo “non-timber” (DEREBE; ALEMU, 2023; LAARIBYA, 2023),
ora o termo “non-wood” (PURWESTRI et al., 2020; MUSA et al., 2023) e ainda, os que
utilizam

“non-timber”

desconsiderando

usos

madeireiros

(SOLDATI,

ALBUQUERQUE, 2008). É possível que a utilização de termos distintos (“non-timber”
e “non wood”) como sinônimos nesses estudos resulte da confusão conceitual que
esses termos geram. De acordo com o site da Organização das Nações Unidas para
Alimentação e Agricultura (FAO, 2024), o termo “non-wood”, exclui todas as matériasprimas lenhosas. O termo “non-timber”, por sua vez, geralmente inclui lenha e
pequenas madeiras como ferramentas, equipamentos domésticos e esculturas.
Enquanto não há uma definição global dos PFNM, é necessário ter clareza
sobre quais abordagens estão sendo consideradas pelos pesquisadores para não
generalizar equivocadamente resultados favoráveis ou desfavoráveis à conservação
de espécies associadas ao uso desses produtos. No primeiro capítulo dessa tese,
embora tenha sido utilizado o termo “timber”, todas as categorias de usos madeireiros
estão sendo consideradas. Além disso, assim como a definição de Soldati e
Albuquerque (2008), o conceito adotado para produtos florestais não-madeireiros
nesse estudo não inclui os recursos madeireiros.

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2.1.1 Uso e conservação de produtos florestais não-madeireiros
Em se tratando dos aspectos ecológicos associados ao uso de PFNM, este é
considerado como uma forma alternativa de uso da terra, com impactos menores
sobre a cobertura florestal quando comparado à exploração madeireira ou à
agricultura comercial. Isso porque os produtos comercializados podem ser frutas,
fibras vegetais, óleos, resinas, plantas medicinais, sementes, (NAKAZONO;
MAGNUSSUM, 2016) e mel (LOWORE, 2020), entre outros.
Além de serem importantes componentes para a subsistência de famílias,
mitigação ou redução da pobreza e geração de renda a partir da comercialização, eles
fornecem uma rede de segurança em tempos de dificuldades econômicas
(DECARLO; ALI; CERONI, 2020), como a melhoria da segurança alimentar,
principalmente em períodos adversos, além da manutenção das tradições e
conhecimentos culturais (SHACKLETON, 2015). Um reforço à importância econômica
de plantas alimentícias e de subsistência para as populações locais pode ser
encontrado no estudo de Adam, Pretzsch e Pettenella (2013) que foi realizado na
localidade de Rashad, no Sudão, para analisar o papel dos PFNMs em estratégias de
subsistência no desenvolvimento rural. Os resultados revelaram que as funções
desses produtos iam desde ajuda às famílias para garantir o abastecimento de
alimentos, até acumular capital financeiro, de modo que a venda dos frutos de
Adansonia digitata representava uma estratégia de subsistência para a maioria das
famílias de coletores e uma estratégia acumulativa para a maioria das famílias de
comerciantes. Por outro lado, a venda dos frutos de Ziziphus spina-christi e Balanities
aegyptica era uma estratégia de subsistência para a maioria das famílias de coletores
e comerciantes. Essas estratégias tinham relação com o acesso aos mercados e o
valor de mercado de tais produtos.
Os múltiplos benefícios socioeconômicos e socioambientais dos produtos
florestais não-madeireiros, como mencionados acima, fizeram com que estes fossem
visados fortemente a partir dos anos 90 como impulsionadores do manejo sustentável
dos recursos florestais (SHACKLETON; SHACKLETON, 2004; SHANLEY, 2011).
Tendo como principal premissa que, se a floresta apresentasse valor econômico para
as comunidades locais, as pessoas estariam mais propensas a mantê-las (EVANS,
1993).

24

Desde então, vários estudos baseados na premissa de que os produtos
florestais não madeireiros são um caminho sustentável para explorar as florestas, têm
avaliado os impactos ecológicos da extração de diversos produtos florestais nãomadeireiros. Dentre esses estudos, aqueles focados na coleta de frutos com
importância econômica e/ou para a subsistência têm encontrado resultados positivos
(ZUIDEMA; BOOT, 2002; OLIVEIRA; SCARIOT, 2010; BRUSHI et al., 2014;
GIROLDO; SCARIOT, 2015) e negativos (SINHA; BAWA, 2002; AVOCÈVOU-AYISSO
et al., 2009) a respeito da sustentabilidade do uso desse recurso vegetal. Na maioria
dos casos, a coleta de PFNMs não leva à morte dos indivíduos coletados, pois para
várias espécies vegetais, esses produtos são removidos pelas pessoas locais na
própria planta (GAOUE et al., 2016), sem necessidade de cortes ou poda. No entanto,
em situações em que a espécie-alvo é danificada ou morta durante o processo, a
extração parece ser geralmente insustentável (SINHA; BAWA, 2002). Além disso, o
impacto da exploração vai depender da parte da planta que é explorada e do potencial
de regeneração da espécie (CAMPOS et al., 2018). Por exemplo, a extração de frutos
tem sido considerada menos impactante e mais sustentável quando comparada com
a extração de folhas, cascas e raízes (HALL; BAWA, 1993; GAOUE;TICKTIN, 2007;
STANLEY; VOEKS; SHORT, 2012).
Sinha e Bawa (2002) realizaram um estudo nas florestas de Biligri, na Índia,
visando avaliar o impacto das técnicas de coleta dos indígenas Soligas sobre as
espécies frutíferas Phyllathus emblica e Phyllanthus indofischeri e a susceptibilidade
das árvores a infestação por plantas hemiparasitas da família Loranthaceae. Os
autores identificaram que as atuais técnicas de coleta utilizadas pelos Soligas estão
impactando negativamente o recurso que lhes serve de sustento, uma vez que para a
espécie Phyllathus emblica, a poda dos ramos reduziu a produção de frutos no ano
seguinte e, de forma geral, o corte das árvores está comprometendo o uso sustentável
de longo prazo de Phyllanthus spp. No entanto, eles descobriram que a infestação por
plantas hemiparasitárias também tem impacto negativo na produção de frutos das
espécies, sustentando o argumento de que eventos naturais também precisam ser
incluídos nessas avaliações.
Outro aspecto a considerar sobre as práticas de coleta e produção de frutos é
que algumas espécies de frutíferas, mesmo sujeitas a diferentes distúrbios
antropogênicos e extrativismo em escala comercial, conseguem se manter produtivas.

25

Em uma investigação conduzida em uma área de Cerrado lato sensu, Giroldo e Scariot
(2015) realizaram um estudo de estruturação populacional. Eles avaliaram se haveria
alguma evidência de mudança no recrutamento no nível de paisagem em populações
de uma espécie frutífera com importância comercial, Caryocar brasiliense, a qual está
sujeita a diferentes usos e manejos da terra. Giroldo e Scariot (2015) descobriram que
as populações de Caryocar são afetadas pelo uso e manejo da terra, mas a pressão
atual da coleta de frutas nas populações da espécie não comprometia o recrutamento
de mudas na paisagem, concluindo que a coleta de frutos é sustentável. Essa espécie
já foi relatada como tendo coleta máxima sustentável de frutas de cerca de 90%
(OLIVEIRA; SCARIOT, 2010). No entanto, essa informação não pode ser
generalizada, pois estudos têm indicado que a sobrexploração de frutas tem mostrado
impacto na dinâmica de regeneração de árvores frutíferas silvestres (AVOCÈVOUAYISSO et al., 2009; ALMEIDA, 2014). Algumas pesquisas vem apontando que a
insustentabilidade da extração de frutos em algumas comunidades locais está
associada, muitas vezes, ao abandono de estratégias e técnicas de coleta tradicionais
como medida de maximização do retorno econômico (SINHA; BAWA, 2002) e a
superexploração de uma espécie-alvo mediante baixos retornos (CROOK; CLAPP,
1998).
Neste sentido, um estudo realizado por Gaoue et al. (2016) avaliou as taxas
ótimas de coletas de produtos florestais madeireiros e não madeireiros, considerando
dados de pesquisa em ecossistemas tropicais e integrando fatores socioeconômicos
e ecológicos. Os autores classificaram essas coletas, respectivamente, como coleta
letal e coleta não letal. Com base em seus resultados, eles demonstraram que as
estratégias de colheita ideais incluem começar com a coleta não letal de PFNM e adiar
a coleta letal de madeira para começar após alguns anos, embora uma redução
drástica da população seja causada pela coleta letal em todos os cenários de controle
do estudo. Por fim, o estudo indicou que para manter uma população com um declínio
inferior a 10% da sua densidade inicial, as taxas ótimas de coleta letal e não letal, não
devem exceder 40% da densidade populacional total.
No entanto ainda existe um dilema no que diz respeito à sustentabilidade entre
o manejo florestal direcionado de espécies de interesse alimentício e coleta sem
manejo. Por um lado, estudos com PFNM (entre eles as espécies alimentícias) vem
mostrando que a extração sem manejo tem mais chances de trazer consequências

26

negativas para as populações alvo (KUSTERS et al., 2006). Por outro lado, espécies
que experienciaram aumento da demanda nos últimos anos e que tiveram
estabelecidas formas de manejo in situ (ex: plantio em áreas de vegetação nativa),
embora mantenham suas populações, podem trazer impactos para a comunidade
como um todo (FREITAS, 2019; FREITAS et al., 2015; BARROS et al., 2023). Um
exemplo disso é o caso da espécie Euterpe oleraceae Mart. (açaí), cujo aumento do
valor de mercado em escala nacional, impulsionou a intensificação do cultivo da
espécie in situ, o que tem levado à diminuição da diversidade de espécies na floresta
estuarina amazônica (FREITAS, 2019; FREITAS et al., 2015; BARROS et al., 2023).
Como observado, alguns estudos têm sugerido que a coleta de recursos
florestais não madeireiros nem sempre se dá de forma sustentável (BROKAMP et al.,
2014; AVOCÈVOU-AYISSO et al., 2009). Adicionalmente, há o dilema da
sustentabilidade entre a coleta com manejo (KUSTERS et al., 2006) e coleta sem
manejo (FREITAS et al., 2015) atrelada a esses produtos. Por outro lado, a extração
de produtos florestais não madeireiros, especialmente a coleta de frutos silvestres, em
sua maioria, afeta menos a estrutura e a função das florestas do que outros usos
(STANLEY; VOEKS; SHORT, 2012) como os madeireiros.
No tópico seguinte (e seus subtópicos) será abordado um conjunto de
evidências que fornecerá as bases para reforçar a importância do uso alimentício
como fator de proteção. Inicialmente, abordarei sobre evidências históricas de
estratégias de manejo vegetal realizadas por populações locais que promovem a
proteção de certas espécies com características desejáveis. Em seguida, abordarei
sobre a conservação através do uso pautada em produtos florestais não-madeireiros.
Por fim, discorrerei sobre evidências a partir da hipótese da proteção, que leva em
consideração o efeito protetivo para as espécies lenhosas associado a um uso
especializado contra um uso generalista e destrutivo a estas.

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2.2 Evidências do papel de proteção associados às plantas alimentícias

2.2.1 Proteção de plantas a partir do manejo: evidências da domesticação
A literatura vem demonstrando que a relação entre seres humanos e plantas é
contrastante. Alguns exemplos de relação desarmoniosa incluem o abandono de
estratégias e técnicas de coleta tradicionais como medida de maximização do retorno
econômico (SINHA; BAWA, 2002). Adoção dessas novas técnicas podem levar à
coleta destrutiva dos produtos florestais, e à superexploração de certas espécies
vegetais mediante baixos retornos (CROOK; CLAPP, 1998). Por outro lado, um
conjunto de estudos vem demostrando a existência de várias estratégias de manejo
que podem trazer benefícios mútuos para as plantas e pessoas (CASAS et al., 1997;
FONDOUN; TIKI-MANGA, 2000; CASAS et al., 2001; BLANCAS et al., 2010; Blancas
et al., 2013).
A forma de manejo incipiente é um tipo de interação intermediária entre a coleta
e a agricultura, que tem sido documentada por alguns pesquisadores (CASAS et al.,
1997). Essa forma de manejo inclui as práticas realizadas nos ambientes onde as
plantas ocorrem naturalmente, podendo esses ambientes serem as florestas, no caso
de plantas silvestres e ambientes antrópicos quando se trata de plantas espontâneas
(BLANCAS et al., 2010). Esse tipo de manejo é denominado técnicas de manejo in
situ, que podem ser diferenciadas em três processos: tolerância, promoção e proteção
(CASAS et al. 1997). De acordo com Blancas et al. (2010), e, considerando apenas a
ocorrência dessas técnicas em florestas, tem-se que: 1) tolerância é uma técnica por
meio da qual espécies vegetais úteis silvestres ou fenótipos particulares dessas
espécies são deliberadamente deixados sem nenhuma manipulação durante eventos
de desmatamento; 2) a promoção ou aprimoramento ocorre quando atividades são
realizadas com o intuito de aumentar a densidade populacional e a disponibilidade de
espécies vegetais úteis ocorrentes em uma comunidade biótica; e 3) a proteção inclui
ações que favorecem a permanência de certas plantas por meio de cuidados
especiais, que podem envolver a eliminação de competidores e herbívoros, podas,
proteção contra pragas, geadas, radiação solar e outras.
As interações entre humanos e plantas podem ser influenciadas por alguns
fatores. como: a) o papel do recurso vegetal na subsistência humana, seja em termos

28

econômicos ou culturais; b) sua disponibilidade em relação à demanda humana; sua
qualidade; e c) a viabilidade de manejo dos seus propágulos, suas populações ou
comunidades bióticas nas quais esses recursos ocorrem (CASAS et al., 2001). Em
um estudo realizado em quinze aldeias, na zona da floresta úmida do Sul de
Camarões, Fondoun e Tiki-Manga (2000) identificaram as abordagens tradicionais
para a conservação de duas espécies (Garcinia kola e Gnetum africanum) muito
importantes para usos alimentícios, medicinais e econômicos, para geração de renda
de pequenos agricultores. Os resultados mostraram que os agricultores estão
envolvidos na conservação in situ das espécies, usando, dentre outras abordagens, o
desmatamento seletivo durante a preparação da terra para cultivo, a coleta
sustentável de cascas de populações vegetais silvestres mantendo-as de pé e,
evitando o corte de populações das espécies arbóreas durante o desmatamento de
áreas florestais. Outro exemplo das práticas de manejo in situ que beneficiam as
espécies vegetais foi encontrado por Blancas et al. (2010) que, ao revisarem um
conjunto de estudos etnobotânicos conduzidos em treze aldeias do Vale TehuacátanCuicatlán, no México, identificaram as três técnicas de manejo acima descritas para
espécies com usos principalmente alimentício, forrageiro, medicinal e ornamental.
Entre as plantas manejadas com múltiplos usos, o uso alimentício foi o mais frequente
e 60 plantas alimentícias foram registradas entre as espécies com uso exclusivo,
perdendo apenas para as plantas ornamentais.
O tipo e a intensidade e estratégias de manejo de plantas podem variar entre
as espécies conforme as necessidades das pessoas locais. Em uma pesquisa
etnobotânica, Blancas et al. (2013) analisaram a relação entre índices de risco e a
intensidade do manejo de espécies vegetais alimentícias usadas em cinco aldeias de
Coyomeapan, no México. Os resultados indicaram que as pessoas manejam os
recursos vegetais de acordo com o papel que desempenham na subsistência das
famílias, ou a quantidade disponível e a qualidade de seus produtos úteis, sobretudo
o equilíbrio entre a disponibilidade e a demanda de recursos. Os autores encontraram
que, em geral, os menores valores de intensidade de manejo correspondiam a
espécies sujeitas a coleta simples ou tolerância. A maioria delas eram plantas anuais
abundantes, consumidas ocasionalmente por poucas pessoas. Enquanto os maiores
valores de intensidade de manejo foram registrados em espécies com importância

29

econômica, em sua maioria perenes, com variedades reconhecidas, cujo manejo
requer o uso de ferramentas e que são protegidas por regulamentação coletiva.
Embora esses estudos tratem sobre o favorecimento deliberado de plantas
alimentícias de alta importância econômica e cultural nos seus locais de ocorrência a
partir de estratégias de manejo como a tolerância, promoção e proteção, pouco se
sabe sobre como a importância da planta para um fim pode protegê-la de outros usos
mais destrutivos, ou seja, dos efeitos interativos entre as diferentes formas de
aproveitamento das espécies vegetais. Isso limita a nossa compreensão do efeito
protetivo em sistemas socioecológicos e de seu benefício econômico para a
população humana. Essas informações são essenciais para se entender e estabelecer
o arcabouço da conservação biocultural, mostrando os múltiplos benefícios da
manutenção de práticas culturais associadas à biodiversidade.

2.2.2 Conservação através do uso de PFNM
A promoção dos produtos florestais não-madeireiros tem sido proposta como
uma maneira sustentável de explorar as florestas tropicais, intensificando-se a partir
dos anos 90 (BELCHER; RUIZ-PÉREZ, 2004). Isso se deve, sobretudo, à possibilidade
de coleta de partes da planta, que seria menos destrutiva do que a exploração
madeireira e a importância desses recursos para muitas comunidades rurais ao redor
do mundo. Além disso, os PFNM têm sido associados ao alívio da pobreza (ZENTENO
et al., 2013).
Com a promoção e utilização dos PFNM, muitos pesquisadores incorporaram
uma abordagem chamada “conservação através do uso” aos seus estudos. Essa
abordagem

foi

posteriormente

denominada

hipótese

da

conservação

pela

comercialização (EVANS, 1993) e postula que as florestas podem ser conservadas se
as comunidades locais obtiverem retorno econômico direto da coleta desses produtos
(EVANS, 1993).
No entanto, não é qualquer PFNM que pode alcançar retornos econômicos
suficientes e de forma segura do ponto de vista sustentável. O consumo de produtos
florestais não-madeireiros ganha visibilidade econômica quando seu valor de mercado
é superior ao valor para uso doméstico, possibilitando a geração de renda (VENTER;
WITKOWSKI, 2013). Vários fatores podem influenciar o sucesso desta atividade

30

econômica como sistemas de coleta, taxa de regeneração de espécies, disponibilidade,
procura, diversificação de produtos, mercado, cadeia de valor, entre outros
(SHACKLETON et al., 2024). No contexto de recursos alimentícios como PFNM, para
que a comercialização seja disseminada em uma base sustentável, o local deve ter:
pessoas que precisem gerar renda, mercados disponíveis e espaço para melhorar a
base da dieta. Além disso, as atividades comerciais precisam ser realizadas em
pequena escala, serem específicas para o local e considerarem os lucros somente para
os produtores (EVANS, 1993).
Exemplos de casos exitosos considerando essa abordagem foram encontrados
em vários estudos (PETERS; GENTRY; MENDELSOHN, 1989a; PETERS; GENTRY;
MENDELSOHN, 1998b; WADT et al., 2008; CHAGAS et al., 2021). Por exemplo,
Zuidema e Boot (2002), realizaram o estudo da demografia de Bertholletia excelsa
(castanha-do-pará) em duas florestas primárias do norte da Bolívia, das quais a
castanha do Pará é extraída há décadas. Os autores concluíram que devido ao
contínuo rejuvenescimento da população, o seu tamanho estável, a elevada idade na
maturidade e o longo período reprodutivo, a extração da castanha-do-pará poderia
durar por décadas. Mais tarde, Wadt et al. (2008) analisaram as populações exploradas
de castanha-do-pará (Bertholletia excelsa) em áreas protegidas no oeste da Amazônia,
a partir de um estudo comparativo da estrutura da espécie em três locais com contextos
ecológicos e socioeconômicos distintos. Os autores encontraram que a estrutura
populacional em todos os três locais foi representada por uma distribuição de classe de
tamanho J reversa, tendo maior representação em classes de tamanho menores e
representação proporcional ligeiramente decrescente com o aumento do tamanho, uma
estrutura que sugere populações demográficas saudáveis. O estudo concluiu que a
regeneração é suficiente para a persistência a médio prazo das populações de plantas
nesses locais.
Outro exemplo da conservação através do uso é destacado por Chagas et al.
(2021). O estudo foi realizado em áreas da Mata Atlântica brasileira e avaliou a estrutura
e a dinâmica populacional de Euterpe edulis Mart. (palmeira juçara), em três tipos de
florestas: florestas secundárias protegidas, florestas secundárias manejadas e
agroflorestas. Também foram realizadas avaliações financeiras e ecológicas da
produção da polpa da fruta e palmito para comparar os resultados da colheita de frutas
nessas áreas florestais. Os resultados revelaram que árvores frutíferas maduras eram

31

mais comuns em florestas secundárias e agroflorestas do que em áreas protegidas,
apesar da falta de plantas de tamanho intermediário. As projeções demográficas com
dados de campo, indicaram que a produção manejada de frutas em florestas
secundárias e agroflorestas tem resultados de longo prazo superiores na dinâmica
populacional de E. edulis, lucratividade e disponibilidade de frutos para wildlife, em
comparação com a maioria dos cenários de coleta de palmito. As conclusões do estudo
indicam que agroflorestas e florestas secundárias são eficazes na conservação e
restauração das populações de E. edulis, enquanto geram lucro sustentável para os
agricultores. A coleta de frutos é destacada como uma estratégia que maximiza tanto a
conservação quanto os benefícios financeiros, demonstrando um equilíbrio entre
objetivos de conservação e econômicos.
Com o passar do tempo, no entanto, a partir de revisões da literatura, com base
nos estudos de caso realizados, algumas análises e críticas surgiram, contestando a
viabilidade das atividades de coleta dos PFNM (CROOK; CLAPP, 1998; BELCHER;
RUIZ-PÉREZ, 2004; NEWTON, 2008). Esse contexto crítico inclui: desconsiderar que
os lucros a partir da venda desses produtos muitas vezes não são diretamente
destinados aos extrativistas, mas divididos com intermediários; desconsiderar perdas
após a colheita e o impacto nos preços dos produtos diante da intensificação nas
atividades extrativistas; desconsiderar restrições relacionadas à posse da terra que
pode limitar o acesso ao recurso, a escala comercial desses produtos, entre outras
(CROOK; CLAPP, 1998).
A partir disso, um conjunto de soluções e desafios foi posto em perspectiva
para obtenção de sucesso na promoção e venda dos PFNM pelos povos locais. Esse
conjunto inclui: conhecimento técnico sobre conservação, processamento e
armazenamento de produtos sazonais (ex. frutos), educação financeira, estratégias de
marketing, certificação dos produtos, coleta sustentável, utilização de parte dos
retornos para ações práticas de conservação destinadas a combater outras ameaças
às espécies, etc. (EVANS, 1993; NEWTON, 2008). Isto resultaria em maiores retornos
financeiros, os quais seriam incentivos à conservação das áreas florestais ou espécies
vegetais.
Entretanto, é preciso atentar sobre qual conceito de PFNM é assumido pelo
pesquisador, porque há muitos estudos que consideram, além de frutos, alimentos
silvestres como carne de caça e de outros animais e até a extração de madeira de alto

32

valor como parte desses produtos, muitas vezes incluindo mais de um tipo desses
PFNM. Por exemplo, na revisão de Newton (2008) são exemplificadas dez espécies
arbóreas de interesse para a conservação em vários estudos, nos quais a exploração
madeireira é mais citada do que a obtenção de produtos resina, nozes, folhas e flores
cuja extração é menos destrutiva. Em nove desses estudos a coleta de madeira foi
considerada insustentável ou pouco sustentável, enquanto o uso de nozes, folhas e
flores foi considerado sustentável.
Diante das críticas associadas à abordagem da conservação através do uso de
PFNM, faz-se necessária a realização de estudos que usem ferramentas sistemáticas
para colocar em prova a ideia de conservação pelo uso. Uma das principais hipóteses
a serem testadas, nesse sentido, é a hipótese da proteção. O próximo subtópico,
fornece argumentos sobre como a interação de usos especializados e generalistas
pode ser importante para estudos voltados à conservação de espécies lenhosas.
2.2.3 Evidências da hipótese da proteção

Quando se compara usos alimentícios a usos madeireiros quanto ao número
de espécies disponíveis para uso e o grau de dano em termos do impacto da coleta
das partes usadas, tem-se que o uso alimentício é considerado pela literatura como
especializado e com menores impactos nas populações vegetais. Já os usos
madeireiros costumam ser considerados generalistas e causadores de maiores
impactos à estrutura vegetal (MEDEIROS et al., 2011), devido a coletas geralmente
destrutivas. Os usos especializados são definidos por uma quantidade restrita de plantas
adequadas que atendem a requisitos específicos. A ideia de especialização é reforçada,

pelo fato de a disponibilidade das plantas possuir influência menor ou nula sobre esse
uso (RIBEIRO et al., 2014; SOLDATI et al., 2016).
Uma evidência empírica favorável ao efeito de proteção de usos especializados
sobre usos generalistas foi registrada por Silva et al. (2021), que testou a hipótese da
proteção para a qual a importância de usos medicinais (usos especializados) tinha um
efeito de proteção sobre usos madeireiros (usos generalistas). O estudo demonstrou
um efeito pequeno, porém significativo da importância do uso medicinal sobre o uso
madeireiro (lenha) para espécies da Caatinga.

33

Assim, de um lado, a alta importância das plantas alimentícias para as
populações locais (consumo direto e geração de renda) pode impulsionar as pessoas
a protegerem e a manterem essas espécies que tem uma forte especialização. Do
outro, usos madeireiros podem causar danos estruturais às populações de plantas
lenhosas que também fornecem recursos alimentícios de elevada importância.
Entretanto, como o uso madeireiro é generalista, há outras opções de espécies
igualmente vantajosas que podem ser usadas pelas pessoas, permitindo poupar
certas espécies vegetais para utilização exclusivamente alimentícia. Tendo em vista
isso, as plantas alimentícias com importância doméstica e/ou comercial se constituem
como um modelo viável para testar o efeito de proteção desse uso sobre usos
madeireiros.
Para melhor compreender as diferenças dos usos alimentícios e madeireiros,
discutirei no tópico a seguir sobre os usos e impactos ecológicos nos recursos
florestais madeireiros.

2.3 Uso e conservação de produtos florestais madeireiros

A madeira pode ser usada para diversos fins e os principais usos madeireiros
estão distribuídos principalmente nas categorias combustível, construção e tecnologia
(RAMOS; MEDEIROS; ALBUQUERQUE, 2010). Na categoria combustível está a
madeira que é destinada à geração de energia para o cozimento de alimentos,
aquecimento da água ou de ambientes (RAMOS; MEDEIROS; ALBUQUERQUE,
2010). Nesse caso, a madeira é utilizada na forma de lenha ou carvão. Os elementos
de madeira que são parte de uma estrutura para delimitação territorial, moradia ou
abrigo de animais e armazenamento de objetos fazem parte da categoria construção.
Nessa categoria, a madeira é utilizada para a construção de cercas, mourões, linhas
de casa, caibros, ripas, portas, janelas, entre outros (RAMOS; MEDEIROS;
ALBUQUERQUE, 2010). A categoria tecnologia, por sua vez, comporta os elementos
que sofrem manipulação, contudo, não são destinados para delimitar espaços. Entram
nessa categoria, os cabos de ferramentas, bancos, mesas, cadeiras, canoas, remos
etc. (RAMOS; MEDEIROS; ALBUQUERQUE, 2010).
Esses recursos florestais madeireiros representam grande importância para as
populações locais, com destaque para o uso combustível, principalmente em países

34

subdesenvolvidos. Por exemplo, no nordeste da Índia, Saha e Sundiyal (2012)
registraram que a dependência de recursos florestais foi de 100% para a lenha e para
materiais de construção de casas. Na aldeia de Santiago Quiotec de Oxaca, no
México, 464 toneladas de lenha são extraídas anualmente para cocção (PÉREZNEGRÓN; CASAS, 2007). A lenha representa a principal fonte de energia doméstica
para a população de Muda, em Moçambique (BRUSCHI et al., 2014). Além disso, a
biomassa de madeira extraída nessa região para a produção de carvão vegetal é de
cerca de 26.000 m³ por ano, sendo um dos principais responsáveis pelos índices de
desmatamento da região. No Brasil, a lenha ainda tem utilização significativa pelas
pessoas com maior vulnerabilidade social (HORA et al., 2021; GONÇALVES et al.,
2021; BRITO, 1997). Particularmente no semiárido, as estratégias de sobrevivência
das populações rurais dependem de espécies lenhosas que fornecem material para
construção e usos tecnológicos (ALBUQUERQUE; ANDRADE, 2002).
Embora o uso da madeira seja de grande importância para as populações
locais, sobretudo para as que se encontram em países subdesenvolvidos, ela é o
produto cuja extração é mais danosa às populações vegetais. Avaliando os padrões
de coleta e uso de madeira entre as diferentes categorias, Medeiros et al. (2011)
observaram que esses padrões diferiam substancialmente, de modo que a categoria
combustível abrangia grandes quantidades de madeira, apresentava curto tempo de
reposição e a coleta era baseada em galhos, troncos e em indivíduos mortos e vivos.
As categorias construção e tecnologia, no entanto, apresentaram tempos de
reposições longos e a coleta era baseada em troncos de indivíduos vivos, com um
maior volume de madeira utilizada para construção do que para tecnologia.
A maioria dos estudos sobre produtos florestais madeireiros tem considerado
ora a avaliação conjunta de várias categorias de uso e seus impactos na conservação
dos recursos vegetais, ora especificamente o uso combustível devido ao fato de ser o
uso madeireiro mais difundido em diferentes comunidades. No estudo etnobotânico
com amostragem da vegetação de Miombo (floresta tropical), realizado por Bruschi et
al. (2014) em quatro comunidades de Muda-Serração, em Moçambique, as plantas
lenhosas mais citadas, depois das plantas alimentícias, foram as plantas artesanais
(38 espécies) e domésticas (37 espécies), que foram relatadas como tendo coletas
destrutivas, como cortes de galhos de árvores ou caules principais de madeira. As
plantas classificadas como artesanais são utilizadas, em sua maioria, como postes

35

para construção, objetos domésticos e carpintaria. Dentre os usos domésticos, está o
uso de lenha (10 espécies) e carvão (11 espécies). Os autores indicaram que os
principais riscos para a conservação dos recursos vegetais estão concentrados nos
usos para fins madeireiros, sobretudo de lenha, carvão e carpintaria, que apresentam
um forte impacto tanto pela sobre-exploração de espécies quanto por terem coleta
destrutiva. Estudos sobre o uso e preferência de lenha em comunidades rurais,
revelaram que as pessoas selecionam e utilizam as espécies pela qualidade
madeireira percebida, ao analisarem as propriedades físicas da madeira das plantas
mencionadas como preferidas pelas pessoas locais (RAMOS et al., 2008; CARDOSO
et al., 2015). Quando essa preferência associada ao uso efetivo se concentra em um
pequeno número de espécies pode comprometer a sustentabilidade do uso dessas
espécies a longo prazo.
Esses usos também têm sido tratados na literatura como causadores de
distúrbios antropogênicos crônicos, referindo-se a distúrbios relacionados aos usos
de recursos florestais em proporções pequenas, mas contínuos que vão se
acumulando a longo prazo, ao ponto de seus impactos ecológicos se tornarem visíveis
(RIBEIRO et al., 2019; GONÇALVES, MEDEIROS, ALBUQUERQUE, 2021). Um
exemplo disso é o estudo de Gonçalves, Medeiros e Albuquerque (2021) realizado no
Parque Nacional do Catimbau, em Pernambuco. Os autores descreveram os padrões
de coleta de lenha e construção de cercas e avaliaram os efeitos da coleta de madeira
para esses usos na estrutura da comunidade arbórea. Eles encontraram que as
pessoas tinham um comportamento generalista na coleta de lenha, mas um
comportamento mais especializado na coleta de madeira para construção de cercas.
Embora, de modo geral, a coleta de madeira doméstica tenha demostrado pouco
impacto na estrutura da comunidade, os autores alertam que a coleta de madeira para
construção de cerca pode afetar a estrutura populacional das espécies mais usadas
no futuro, devido ao caráter especializado da sua coleta.
Muitas vezes as plantas lenhosas podem ter múltiplos usos e isso pode
influenciar na avaliação do status de conservação de tais espécies. Por exemplo,
Oliveira et al. (2007) sugerem que os índices de prioridade de conservação podem ser
fortemente influenciados pelo fato de que usos medicinais estão associados com usos
concorrentes de madeira. É possível imaginar que o mesmo aconteça para plantas
lenhosas que também sejam úteis para a finalidade alimentícia, uma vez que os usos

36

madeireiros por serem mais danosos, podem estar reduzindo o uso alimentício, como
sugerido por Brushi et al. (2014). Contudo, os usos madeireiros têm sido classificados
na literatura como sendo generalistas. Para usos generalistas há um repertório maior
de espécies úteis e, geralmente, uma relação forte com a disponibilidade. Desse
modo, as espécies mais usadas são justamente as mais disponíveis, como ocorre
com a maioria dos usos madeireiros (MEDEIROS et al., 2011; GONÇALVES;
ALBUQUERQUE; MEDEIROS, 2016). Essa característica possibilita outras opções
igualmente

vantajosas

de

espécies

lenhosas

para

propósitos

madeireiros

relacionados às necessidades das pessoas locais, o que permitiria que certas
espécies fossem poupadas para usos nos quais dificilmente há equivalentes para
substituição (SILVA et al, 2021), como o uso alimentício.
As evidências empíricas da maioria dos estudos com produtos florestais
madeireiros e não madeireiros explanadas acima, indicam a comum característica da
dependência das populações locais sobre esses recursos vegetais e a notável
distinção em termos da sustentabilidade do uso desses recursos pelas populações
locais. No entanto, ainda são necessários estudos nos remanescentes da Mata
Atlântica que possam contribuir para a ampliação das informações sobre
sustentabilidade dos usos madeireiros e usos alimentícios. Além disso, em um
contexto em que a grande parte das plantas conhecidas e usadas apresentam múltiplo
usos, destaca-se a importância de investigações com vistas à conservação que
avaliem o impacto de um uso sobre outro. Ou seja, estudos que considerem a
interação entre usos de plantas lenhosas, os quais podem indicar, além da possível
influência negativa de certos usos sobre outros, o potencial efeito de proteção que
essa interação pode promover.
No próximo tópico serão explanados alguns fatores que influenciam na seleção
de espécies usadas para fins madeireiros.

2.3.1 Critérios de seleção de plantas madeireiras

Um conjunto de estudos etnobiológicos realizados em distintos contextos
socioambientais têm mostrado que a seleção de plantas madeireiras pelas
populações humanas pode ser influenciada por fatores socioeconômicos (RAMOS et
al., 2008a; ARRUDA et al., 2019; CRUZ et al., 2020), físicos e ecológicos (RAMOS et

37

al., 2008a,b; CARDOSO et al., 2015; SILVA et al., 2017; HORA et al., 2021; CRUZ et
al., 2020). Dentre esses fatores, a qualidade e a disponibilidade têm se destacado em
termos de maior poder preditivo entre os estudos.
A qualidade vem sendo estudada em termos de suas propriedades físicas
(RAMOS et al., 2008; CARDOSO et al., 2015) e da percepção local (HORA et al.,
2020; SILVA et a., 2017). A disponibilidade, por sua vez, vem sendo estudada a partir
da percepção local e/ou sendo aferida por meio de ferramentas ecológicas (HORA et
al., 2020; GALEANO, 2000; ALBUQUERQUE; LUCENA, 2009). Muitos estudos
avaliaram a influência da disponibilidade ambiental (aferida por parâmetros
fitossociológicos) a partir da hipótese da aparência e, ora foi encontrada relação entre
a disponibilidade ambiental e o uso de plantas madeireiras (GALEANO, 2000;
GONÇALVES et al., 2016), ora não (ALBUQUERQUE; LUCENA, 2005). E ambos os
fatores são avaliados ou isoladamente, ou em conjunto, sendo a categoria de uso
combustível a mais presente nesses estudos.
As propriedades físicas da madeira são indicadores da sua qualidade.
Componentes físicos da madeira como densidade, teor de água, teor de umidade, teor
de cinzas e valor calorífico são aferidos como determinantes da combustibilidade de
uma planta. Geralmente, a combustibilidade é estimada por meio do Índice de valor
Combustível (IVI). Alguns trabalhos sobre uso e preferência de lenha que utilizaram
esse índice como indicador de qualidade da madeira, identificaram que as espécies
com maiores valores de IVI são as preferidas pelas pessoas locais, sugerindo que as
pessoas selecionam e utilizam as espécies pela sua qualidade (RAMOS et al., 2008;
CARDOSO et al., 2015). Silva et al. (2017) examinaram a influência da qualidade na
seleção de recursos madeireiros. O estudo foi realizado em uma comunidade
quilombola no município de União dos Palmares, em Alagoas. O estudo testou a
influência da qualidade percebida sobre o uso de lenha em ambiente de Mata Atlântica
com escassez florestal. Os autores encontraram que a qualidade percebida exerce
influência sobre a frequência de uso, mas não sobre o número de usuários de plantas
para lenha. Eles inferiram que o contexto de escassez florestal interfere mais
fortemente no número de pessoas que utilizam recursos de alta qualidade do que na
frequência de uso desses recursos por pessoas que podem acessá-los. Assim,
evidenciou-se a necessidade de que as estratégias de conservação levem em conta
o acesso diferencial a recursos de alta qualidade em contextos de escassez.

38

Uma pesquisa realizada por Hora et al. (2021) em uma comunidade rural do
município de Altinho, em Pernambuco, avaliou a influência de fatores ecológicos
(percepção da dificuldade de aquisição e disponibilidade ambiental) e de fatores
biológicos (dentre eles, a qualidade percebida geral) no uso de plantas para fins
combustíveis. O resultado revelou influência da disponibilidade ambiental, qualidade
percebida (geral) e durabilidade da madeira no uso da lenha. Os fatores que exibiram
influência no uso de carvão foram a qualidade percebida (geral) e a durabilidade. O
estudo mostra a importância de avaliar diferentes fatores de forma combinada para
obter uma resposta mais completa sobre o comportamento de coleta de plantas para
fins combustíveis e, consequentemente direcionar estratégias de conservação mais
realistas.
Embora o efeito das variáveis disponibilidade e qualidade sobre o uso de plantas
para fins madeireiros esteja sendo amplamente estudado em diferentes contextos
socioecológicos, a literatura científica carece de estudos que considerem interações
entre categorias de uso ao avaliar os critérios de seleção dos recursos vegetais. Isso
é importante para entender até que ponto múltiplos fatores podem atuar como
impulsionadores ou não da seleção de plantas lenhosas usadas para fins alimentícios
e madeireiros. Além disso, esses estudos também poderão informar quando a
importância de uma planta para determinado fim supera o seu uso potencial para outro
fim.
Diante de todas as evidências e argumentos apresentados nesta revisão, neste
estudo busquei investigar as interações entre os usos alimentícios e madeireiros e
suas implicações para a conservação biocultural. Esse estudo é o primeiro framework
a testar a hipótese da proteção para plantas alimentícias silvestres e a utilizar
ferramentas sistemáticas para isso.

39

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46

CAPÍTULO 1

3. WILD FOOD PLANTS WITH THE POTENTIAL TO IMPROVE FOOD AND
EXTRACTION: A SYSTEMATIC REVIEW OF THE BRAZILIAN CONTEXT
Article published in Ethnobiology and Conservation

Roberta de Almeida Caetano1*, Élida Monique da Costa Santos1, Richard Zago Poian2, Adriana
Rosa Carvalho3, Rafael Ricardo Vasconcelos da Silva1, Patrícia Muniz de Medeiros1
1

Ecology, Conservation and Biocultural Evolution Laboratory (LECEB), Postgraduate Program

in Biological Diversity and Conservation in the Tropics (DIBICT-ICBS), Campus of
Engineering and Agricultural Sciences (CECA), Federal University of Alagoas, Maceió, AL,
Brazil
2

Ecology, Conservation and Biocultural Evolution Laboratory (LECEB), Campus of

Engineering and Agricultural Sciences (CECA), Federal University of Alagoas, Maceió, AL,
Brazil
3

Department of Ecology, Federal University of Rio Grande do Norte, s/n, Lagoa Nova, PO box

1524, Natal, RN, 59098-970, Brazil
* Correspondence: robertacaetano1991@gmail.com
BR-104, Km 85, s/n, Rio Largo – AL, CEP 57100-000, Brazil
Significance Statement
The interaction of uses is an important tool to capture potentially threatened woody plants.
However, it has been little explored in studies on biodiversity conservation. Our systematic
review provides a compilation of ethnobiological and nutritional information on versatile food
woody plants for the main timber uses, uses considered to be the most destructive to the
structure of plant populations. We indicate food species native to Brazil that are strategic for
food and nutritional security, due to their high nutritional potential and, at the same time,
strategic for conservation, due to the probable risk of loss of their natural populations by
logging. We discuss possible biocultural conservation strategies for these species. We also

47

make recommendations for future studies on identified gaps in both ethnobiological and
nutritional studies.

Abstract
Wild food plants can contribute to improving the food and nutrition security of local populations
by promoting diet diversification and increasing the intake of micro- and macronutrients.
However, many of these plants are also used as timber. Wild food species need to be identified
and their food–wood use interactions need to be well understood for the development of
conservation strategies, as species with the potential to improve food and nutrition security may
be threatened by destructive extraction. This systematic review recorded and compiled
nutritional information on woody plant species native to Brazil that are used by local
populations for food and timber purposes, seeking to identify which species have a high overlap
between food and timber uses as well as a high nutritional potential. A total of 635 woody
species with timber and/or food uses were identified. Of this total, at least 42 species find
application in all timber use categories analyzed in this study, being considered versatile.
Comparison of ethnobiological and nutritional data revealed nine versatile species for which
nutritional composition information was available, among which three stood out in terms of
macronutrient contents, namely Anacardium occidentale L., Bauhinia cheilantha (Bong.)
Steud., and Eugenia pyriformis Cambess. Many versatile species classified as threatened or in
decline have not been the focus of nutritional studies, which signals the need for greater
nutritional research efforts. It is also necessary to investigate whether food importance exerts
any protective effect on these species, reducing timber use pressure (protection hypothesis).

Keywords: Biocultural conservation, Ethnobotany, Food use, Nutritional composition, Timber
use, Use interaction.
1. Introduction
In various parts of the world, studies have underscored the importance of wild food
plants for the food and nutrition security of local populations, particularly during periods of
food shortage (do Nascimento et al., 2012, 2011; Medeiros Jacob et al., 2020; Medeiros et al.,
2021; Shackleton et al., 2015; Shackleton and Shackleton, 2004). Wild plants can play an
important role in a healthy diet as alternative sources of minerals, vitamins, and antioxidants
(Bacchetta et al., 2016; Jacob et al., 2022; Rico et al., 2016), as well as macronutrients such as
carbohydrates, proteins, and lipids (Medeiros Jacob et al., 2020). Consumption of wild food
plants has been recommended as part of global strategies to manage malnutrition (Hunter et al.,
2019) diversify the human diet (Baldermann et al., 2016), improve food systems, and generate
income for small-scale farmers and extractivists (Delang, 2014).

48

The importance of wild plants extends well beyond socioeconomic and nutritional
factors (Medeiros Jacob et al., 2020). From a conservation perspective, studies have argued that
people who rely on utilitarian and/or economic returns from forests are less likely to carry out
activities that generate changes in land use in forest areas. This concept became known as the
"conservation by commercialization hypothesis" (Evans, 1993; Lowore, 2020). It is also
noteworthy that the extraction of edible parts of plants is, in most cases, less harmful than timber
extraction, being conducive to the application of sustainable management and use strategies.
However, despite the great potential of wild food plants for sustainable management, many of
these species have multiple applications, including timber extraction, which is a major cause of
decline among plant populations (Bruschi et al., 2014; Ros-Tonen, 2000; Stanley et al., 2012).
Timber uses include fuel, construction, and technological applications (Ramos et al. 2010).
Several plants found in forests and other natural ecosystems can be used both as food
and timber. Such species need to be identified and their food–wood use interactions understood
to guide the development of conservation actions. Populations of species with high potential to
improve food and nutrition security may be threatened by more destructive uses than food
extraction. This situation is particularly worrisome for woody species that combine multiple
timber uses (versatile species), as they may be facing greater use pressure. In Brazil, several
studies described woody species with nutritional potential (do Nascimento et al., 2012, 2011;
Medeiros Jacob et al., 2020; Nunes et al., 2012). However, most studies have failed to analyze
the intersection between nutritional potential and the extent to which plant populations are
threatened by uses other than human consumption. Understanding the overlap of uses of woody
plants is important because conservation strategies encompassing different uses of plant
resources can greatly contribute to food and nutrition security.
Species extinction represents an irreversible loss of biodiversity and cultural heritage.
To avoid this problem, it is essential to develop strategies from a biocultural conservation
perspective, that is, seeking to combine environmental conservation actions with knowledge on
the use of natural resources to reduce the loss of biological and cultural diversity (Gavin et al.,
2015). Preserving the widespread use of wild food resources might be strategic because wild
plants are found in many local communities that still face challenges related to hunger, food
availability, diet diversification, and climatic events impacting crop production (do Nascimento
et al., 2012, 2011; Medeiros Jacob et al., 2020).
On the one hand, timber extraction tends to be harmful to plant populations, which
explains its use as a threat indicator. On the other hand, it is important to emphasize that (i)

49

timber extraction can be carried out sustainably, as has been proposed in different
socioecological contexts (Bahru et al., 2021; Cavalcanti et al., 2015; Lucena et al., 2007; Tabuti
et al., 2011), and (ii), in some cases, food extraction can be more damaging to plant communities
than timber extraction, depending on forest management intensity. A prominent example is açaí
(Euterpe oleracea Mart.), whose management has led to the simplification of estuarine
communities in the Amazon Forest (Freitas et al., 2021). In this study, we approach the topic
with awareness of possible contrasting effects. Nevertheless, it is understood that woody
species with multiple timber uses may be exposed to greater use pressure and, consequently,
higher management intensity.
In this systematic review, we aimed to identify priority species for biocultural
conservation that are, at the same time, strategic for the maintenance of food and nutrition
security (in terms of micro- and macronutrient composition) and potentially threatened by
multiple timber uses. For this, we identified woody species native to Brazil that are used by
local populations for food and timber purposes and compiled nutritional information available
in the scientific literature to answer the following questions: (i) Which woody food species have
high versatility as timber? and (ii) Which food and timber species have high potential to
contribute to food and nutrition security?
This study presents the results of two systematic reviews. The first, of an ethnobiological
nature, summarizes information on wild plants with overlapping applications as food and
timber, and the second, of a nutritional nature, compiles information on the chemical
composition of these species.

2. Methods
2.1. Ethnobiological systematic review
This systematic review was conducted based on the PRISMA guidelines (see Additional
File 1). Figure 1 shows a flowchart of the research steps.

2.1.2. Eligibility criteria

Studies were selected according to the following eligibility criteria: (i) articles of an
ethnobotanical nature, (ii) original studies, and (iii) studies assessing food and/or timber plants
native to Brazil. Duplicates, articles focused on herbaceous plants only, studies not indexed in
major databases, and review studies were excluded. Priority was given to studies with complete

50

floras. Studies assessing plants from only one botanical family or only a few species (<5) were
excluded.
2.1.3. Information sources
Searches were carried out between January and February 2022 in three databases: Web
of Science, Scopus, and SciELO. The first two databases were chosen because they contain the
largest number of articles published in international journals and achieved excellent
performance in systematic reviews (Bramer et al., 2017). SciELO was included to reach a
greater number of studies published in Brazilian journals. Additional articles were identified by
screening the reference list of articles identified from database searches.

2.1.4. Search strategy

Database searches were performed on two occasions, hereafter referred to as B1 and B2.
An additional search was performed via other sources (B3). After the initial search (B1), a
second search (B2) was performed to expand the retrieval of articles not identified through the
initial keywords. The reference list of all articles selected in B1 was screened for potentially
relevant titles, and these newly identified articles were examined. We recorded and identified
the most frequent keywords used in these articles, including keywords that had not been used
in B1, in order to conduct a new cycle of searches (B2) in the three databases. A third search
(B3) was performed through other sources, in which articles retrieved in B2 were consulted to
identify new keywords and then submitted to the selection processes described in the next
section.
The same search terms were used for the three databases, with the inclusion of
Portuguese keywords for searches in the SciELO database. Search efforts were directed to
article titles, abstracts, and keywords by using the different fields available in each database, as
follows: topic (Web of Science); title, abstract, and keywords (Scopus); and all indexes
(SciELO). The search strategies used in each database are available in Additional File 2.

2.1.5. Study selection

Search records were saved in RIS format and imported into the Mendeley reference
manager, which automatically identifies and deletes duplicates. After that, references were
exported to an Excel spreadsheet. In Excel, we were able to identify and manually delete some

51

duplicates that had not been identified by Mendeley's automatic check, possibly because of
errors in the references or titles written in different languages.

52

Figure 1. Flowchart with systematic review search and screening steps

53

Figure 2. Flowchart with search and screening stages of the systematic review on nutritional aspects

54

Figure 3. Distribution of studies on food and/or timber plants extracted in this systematic
review. Note that 80 studies distributed throughout Brazil are expressed (two more than the
number mentioned in the text - 78). This occurred because two studies were carried out in areas
of two different states (Pernambuco and Paraíba). Elaborated by Klebson da Silva.

The first author (RAC) selected articles individually according to previously mentioned
eligibility criteria. First, titles and abstracts were screened, and those that did not meet the
eligibility criteria or had already been retrieved in previous searches (B1 or B2) were excluded.
In case of uncertainty regarding eligibility, another author was consulted (PMM). In the next
step, potentially eligible texts were read in full, and once again analyzed according to eligibility
criteria, that is, whether they presented a list of plants with indications of timber and/or food

55

uses, analyzed more than 5 species, and did not have authors in common analyzing the same
dataset. For B3 articles, as an additional quality filter before these procedures, we checked if
the journals were indexed in official databases (SciELO or Scopus).

2.1.6. Data extraction

Data were extracted from selected articles to an Excel spreadsheet. The first author
(RAC) was responsible for this procedure. The following information was collected: (a) article
data (authors, year of publication, and journal), (b) plant species (without authority), (c) timber
and/or food uses, and (d) use categories (food, fuel, construction, and technology).
Study quality was assessed based on the identification of botanical materials, including
only articles that reported having deposited a voucher specimen in herbaria and/or indicated the
voucher number in tables. This procedure was undertaken because poor identification or
absence of records in herbaria could lead to the inclusion of information mistakenly linked to
certain species (Medeiros et al., 2014). Although other ethnobiological studies used
interviewees' samples as a criterion for risk of bias (De Medeiros et al., 2013), we chose to not
apply this method, given that, for our approach, the fact that a study did not use a representative
sample does not make it unfeasible to compile information on useful plants. Our approach,
therefore, integrated different findings and was not aimed at comparing different studies.

2.1.7. Synthesis of results

Only angiosperms classified as food and/or timber were included in the species survey.
Scientific nomenclature, habits, origin, botanical families, and identifiers were obtained using
the flora package of R software, which is based on information from the Flora and Funga do
Brasil website (Jardim Botânico do Rio de Janeiro, 2022). All taxa were updated to currently
accepted nomenclature at the species level. Taxa identified only at the genus or family level
were excluded, and taxa with subspecies or variety information are presented only at the species
level.
For cases in which the flora package returned no result for the species, we manually
consulted Flora and Funga do Brasil and World Flora Online (WFO, 2022). This review
included only plants classified as native and woody (i.e., plants classified as "shrub" and/or
"arboreal" in the "life form" field). Thus, our research does not cover the entire universe of wild

56

food plants, being limited to native woody plants, which are the species of interest for
conservation strategies at the food–wood interface.
Timber uses were classified according to a previous study (Ramos et al. 2010). When
studies categorized or specified the uses of timber species, but such a classification merged fuel,
construction, or technological applications under a different denomination (e.g., handcraft), we
reclassified the use category in the spreadsheet under a new column labeled "Updated
category." Species grouped together in the manufacture/handicraft categories that did not
contain these specifications were excluded. This procedure did not lead to the exclusion of
entire studies.
2.1.8. Species of high importance for biocultural conservation
Food species included in the four timber use categories (food, technology, construction,
and fuel), referred to herein as versatile species, were considered of high importance from a
biocultural conservation perspective.
2.2. Nutritional systematic review
The nutritional systematic review also followed PRISMA guidelines (see Additional
File 3). It included only species considered versatile in the ethnobiological systematic review.
A flowchart illustrating the steps in the nutritional systematic review is presented in Figure 2.
2.2.1. Eligibility criteria

Original articles focused on human food plants and assessing the nutritional composition
of the selected plant species were screened by reading the title and abstract.

2.2.2. Information sources
A review of the scientific literature was conducted in the same three databases used in
the previous review (Web of Science, Scopus, and SciELO) in addition to a specific database
for nutritional composition information (Brazilian Biodiversity Information System, SiBBr)
(SiBBr, 2022).
2.2.3. Search strategy

57

Search strings were constructed by combining the currently accepted scientific name of
each species (without the authority) + nutritional. For species whose scientific names were
recently modified or whose alternative nomenclatures, despite not being currently accepted,
were or still are widely used in studies, alternative terms were included in the search (see
Additional File 4).
The word "nutricional" was used in the SciELO database to search for articles written in
Portuguese. In the SiBBr database, we used only the scientific name of species. All database
searches were carried out between August and September 2022.
All procedures performed in Mendeley and Excel for the ethnobiological systematic review
were also used in the nutritional systematic review. To identify additional studies from other
sources, we screened the reference list of review articles directly related to the nutritional
composition of the species of interest. We checked, moreover, the reference list of articles
retrieved from the four databases and screened the keywords of these studies to enrich our
search strategy. However, the most frequent keywords were very similar to those already in
use, precluding the need for new searches.
2.2.4. Study selection
Duplicates, articles not indexed in official databases, and review papers were excluded.
Articles analyzing mixtures of ingredients, enriched products, or quality parameters during food
storage and processing were also excluded.
2.2.5. Data extraction

The following information was extracted from selected studies: (a) article data (authors,
year of publication, and journal), (b) species names (without the authority), (c) part of the plant
analyzed, (d) type of preparation, (e) ecosystem, (f) place of collection, (g) macronutrient
composition (proteins, carbohydrates, and lipids), and (h) micronutrient composition (minerals
and vitamins). At first, we chose to include the following micronutrients: calcium (Ca),
potassium (K), phosphorus (P), magnesium (Mg), sodium (Na), iron (Fe), zinc (Zn), copper
(Cu), manganese (Mn), molybdenum (Mo), chromium (Cr), selenium (Se), sulfur (S), cobalt
(Co), and boron (B). However, the nutritional tables presented in the results include only the
following data: available macronutrients (proteins, carbohydrates, and lipids), total energy

58

value, and mineral contents reported in at least two articles. Thus, the tables include information
on the macrominerals Ca, K, P, Mg, and Na and the microminerals Fe, Zn, Cu, and Mn.
Macrominerals are defined as minerals for which the recommended daily intake is greater than
100 mg. For microminerals, the recommended daily intake is less than 100 mg (Almeida et al.,
2009). Information on carbohydrates was extracted from selected articles by searching for the
term "carbohydrate," including different denominations, such as "total carbohydrates,"
"available carbohydrates," and "calculated carbohydrates."
Given the diversity of information found in some articles, we adopted criteria for the
recording of nutritional information. For example, a study on uvaia (Eugenia pyriformis
Cambess.) analyzed several accessions of the species. We opted to include data on the most
common accession, as it is the most widely known. Another situation occurred when extracting
data from a study on juá (Ziziphus joazeiro Mart.), which analyzed specimens collected in
different regions of the country. In this case, we recorded information on the sample that had
the highest values in all nutrient categories.
For the generation of nutritional tables, when more than one study analyzed the same
plant species, food part, and type of preparation, we selected only one study for data extraction,
whereas when studies analyzed the same species but different food parts or types of
preparations, we included data from all studies. When there was an overlap between nutritional
information, whether of macro- or micronutrients, plant species, or parts, we prioritized studies
analyzing raw materials collected in Brazil. In case of information overlap between Brazilian
studies, we selected the most recent publication.
The first author (RAC) participated in all stages of the nutritional systematic review,
together with two collaborators (RZP) and (AJRCS), under the supervision of PMM.
3. Results

3.1. Ethnobiological systematic review: General aspects
A total of 145 full papers were assessed for eligibility. Of this total, 78 were included in
the ethnobiological systematic review (see Figure 1) after the exclusion of (i) publications
analyzing less than five species, (ii) articles without indication of timber/food uses, (iii) articles
that did not mention the inclusion of voucher specimens in herbaria, and (iv) studies that used
the same dataset and had at least one author in common.

59

Most studies included in the review are of a descriptive nature and can be classified into
three groups: studies on food plants (n =18), studies on timber plants (n = 15), and studies on
plants with multiple uses (n = 45). The selected studies were conducted between 1992 and 2021
in all regions of the country, especially in the Northeast and Southeast. A map of Brazil showing
the occurrence frequency of studies on plant groups included in this review is presented in
Figure 3.
Most studies are concentrated in the Caatinga (n = 27) and Atlantic Forest (n = 27)
biomes. The ecosystems with the lowest number of studies were Cerrado (4) and Pantanal (2).
Additional information is presented in Table 1.
Table 1. Regions and ecosystems in Brazil where the studies were carried out

a

Brazilian regions

Number of studies

Northeast

40

Southeast

22

Midwest

6

North

5

South

4

South/Southeast

1

Ecosystems

Number of studies

Caatinga

27

Atlantic forest

27

Cerrado

4

Pantanal

2

Amazon

5

Atlantic Forest and Cerrado

2

Pantanal and Cerrado

1

Variousa

3

Missing information

7

When there are more than two informed ecosystems

3.1.2. Versatile woody food species used as timber

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A total of 635 native woody angiosperms were recorded, of which 167 are used
exclusively for food, 328 exclusively as timber, and 140 for both purposes. Of the woody food
species used in all timber applications (fuel, construction, and technology), 42 are native to
Brazil. However, given that many studies did not indicate the specific type of timber
application, it is possible that the number of versatile species is much higher.
We observed that half of the versatile species occur in the Caatinga, supported by the
fact that most studies specifying the type of timber use were carried out in this biome. The most
represented botanical families in number of species were Anacardiaceae (n = 7), Fabaceae (n =
6), and Myrtaceae (n = 4). Other six families were represented by two species each, namely
Lauraceae, Euphorbiaceae, Capparaceae, Celastraceae, Burseraceae, and Bignoniaceae. The
major genera, represented by two species each, were Spondias, Handroanthus, Monteverdia,
Ocotea, and Eugenia (Table 2).

Table 2. Woody food plants considered versatile because they are used in the three categories
of wood uses (fuel, construction and technology). Data from the systematic review of Brazilian
ethnobiological studies
Family
Scientific name
Anacardiaceae

Anacardium occidentale L.
Astronium urundeuva (M.Allemão) Engl.
Schinus terebinthifolia Raddi
Spondias mombin L.
Spondias tuberosa Arruda
Tapirira guianensis Aubl.
Thyrsodium spruceanum Benth.

Araliaceae

Didymopanax morototoni (Aubl.) Decne. & Planch.

Bignoniaceae

Handroanthus impetiginosus (Mart. ex DC.) Mattos
Handroanthus serratifolius (Vahl) S.Grose

Burseraceae

Commiphora leptophloeos (Mart.) J.B.Gillett
Protium heptaphyllum (Aubl.) Marchand

Cactaceae

Pilosocereus pachycladus F.Ritter

Capparaceae

Cynophalla flexuosa (L.) J.Presl
Neocalyptrocalyx longifolium (Mart.) Cornejo & Iltis

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Celastraceae

Monteverdia obtusifolia (Mart.) Biral
Monteverdia rigida (Mart.) Biral

Combretaceae

Combretum leprosum Mart.

Euphorbiaceae

Croton heliotropiifolius Kunth
Manihot dichotoma Ule

Fabaceae

Amburana cearensis (Allemão) A.C.Sm.
Anadenanthera colubrina (Vell.) Brenan
Bauhinia cheilantha (Bong.) Steud.
Copaifera langsdorffii Desf.
Inga thibaudiana DC.
Libidibia ferrea (Mart. ex Tul.) L.P.Queiroz

Lauraceae

Ocotea glomerata (Nees) Mez
Ocotea odorifera (Vell.) Rohwer

Lecythidaceae

Eschweilera ovata (Cambess.) Mart. ex Miers

Malpighiaceae

Byrsonima sericea DC.

Moraceae

Brosimum guianense (Aubl.) Huber

Myrtaceae

Eugenia pyriformis Cambess.
Eugenia uniflora L.
Myrcia splendens (Sw.) DC.
Psidium guineense Sw.

Rhamnaceae

Ziziphus joazeiro (Mart.)

Rubiaceae

Genipa americana L.

Sapindaceae

Talisia esculenta (Cambess.) Radlk.

Sapotaceae

Sideroxylon obtusifolium (Roem. & Schult.) T.D.Penn.

Simaroubaceae

Simarouba amara Aubl.

Urticaceae

Cecropia pachystachya Trécul

Ximeniaceae

Ximenia americana L.

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The plant parts most frequently mentioned were fruits (n = 26), flowers (n = 4), and
seeds (n = 4). Leaf, root, and pseudofruit/floral peduncle were cited twice each, and exudate
and resin were cited once. The parts of interest of six species were not specified in any study.

3.2.1. Nutritional systematic review: General aspects

After the first stage of screening, 52 articles were retained and read in full (see Figure
2). Of these, 36 articles were excluded for the following reasons: (i) raw material of commercial
origin (in these studies, pulps/fruits were obtained commercially and information on place of
origin or possible mixtures with other materials or parts, such as nectar, peel + pulp, or
mesocarp + exocarp, pericarp, was not provided), (ii) absence of nutritional information (e.g.,
studies on bioactive compounds), (iii) plant parts with no ethnobotanical uses identified in our
previous review, and (iv) plant parts and ingredients mixed and/or analyzed in combination
(e.g., pulp and peel, sweetened nectar). Therefore, 16 articles were included for compilation of
nutritional data. These studies were published between 1986 and 2020.
Of the 16 studies included in this review, only 4 concerned plant material collected in
countries other than Brazil (Nigeria and India). The plant species with the most nutritional
studies were Anacardium occidentale L. (n = 5), Spondias mombin L. (n = 4), and Pilosocereus
pachycladus F.Ritter (n = 2). Five species were addressed in a single study, namely Genipa
americana L., Bauhinia cheilantha (Bong.) Steud., E. pyriformis Cambess., Manihot dichotoma
Ule, and Z. joazeiro (Mart.).
After application of eligibility criteria, 11 articles were selected to generate the
nutritional tables, 10 of which concerned material collected in Brazil. Four of these studies
provided information on both macro- and micronutrients. The nutrients analyzed in nutritional
tables and the number of species investigated are described in Table 3.

Table 3. Nutrients and number of species contain them
Nutrient type

Number of plant species

Carbohydrates

6

Lipids

7

Proteins

8

63

Calcium

4

Iron

4

Zinc

2

Potassium

3

Phosphorus

4

Sodium

2

Copper

2

Magnesium

3

Manganese

2

The nutritional tables (Tables 4 and 5) contain information on eight plant species. Two
studies analyzed different plant parts of A. occidentale (nut and pseudofruit) and P. pachycladus
(cladode and fruit). The main plant tissue analyzed was the fruit (n = 5). Other parts included
cladode, root, seed, nut, and fruit/pseudofruit (floral peduncle of cashew).
3.2.2. Macro- and micronutrient contents in versatile wild food plants used as timber
3.2.3. A. occidentale

The nut and fruit/pseudofruit of the species were analyzed, with more than one type of
preparation reported. The analyzed studies provided information on the macronutrient
composition of the plant. The highest macronutrient content was found in cashew nut. The
major macronutrients were lipids in both roasted (47.79 g) and raw (47.4 g) cashew nuts. These
values are similar to the lipid content of peanut (Arachis hypogaea L.), as reported by (Ayoola,
P. B, Adeyeye, 2010). The lipid content of roasted cashew nut is higher than that of roasted
peanut (40.60 g).
A. occidentale nut also had the highest micromineral contents, including K (roasted,
556.16 mg; raw, 540 mg), P (roasted, 1101.04 mg; raw, 470 mg), and Mg (roasted, 277.09 mg;
raw, 240 mg). Regarding microminerals, the species is rich in Zn (roasted, 4.98 mg; raw, 5.0
mg). The micronutrient contents of cashew nut are higher than those of peanut (Ayoola, P. B,
Adeyeye, 2010).
3.2.4. B. cheilantha

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The macronutrient content of different preparations of B. cheilantha seed flour was
assessed. Protein was the major macronutrient (soaked flour, 36.0 g; raw seed flour, 35.9 g;
heated seed flour, 31.5 g). The protein content of the seed flour is similar to that of peeled
soybean (Glycine max (L.) Merr.) seed (37.8 g) (TBCA, 2022). In our review of the literature,
no data were found on the micronutrient content of B. cheilantha.
3.2.5. E. pyriformis

The fruit pulp of the species is rich in carbohydrates (53.651 g), having a higher content
than the fruit pulp of Mangifera indica L. (16.0 g), according to the Brazilian Food Composition
Table (TBCA, 2022). The carbohydrate content of E. pyriformis fruit pulp is also higher than
that of a well-known banana variety (Musa acuminata Colla × Musa balbisiana Colla), which,
raw, contains 32.1 g of carbohydrates (TBCA, 2022).
E. pyriformis had the highest Ca content (341.33 mg) among all evaluated species. This
micronutrient content is higher than that of raw orange (Citrus sinensis (L.) Osbenk., 34.6 mg)
(TACO, 2011). E. pyriformis ranked second in P (134.00 mg) and Mg (41.00 mg) contents and
third in K content (134.00 mg). Compared with C. sinensis, the micronutrient contents of E.
pyriformis are high, except that of K, which is higher in orange (170 mg) (TACO, 2011).
The major microminerals in E. pyriformis are Fe (5.37 mg), Cu (0.58 mg), and Mg
(3.05 mg). The Fe content of the species is higher than that of conventional fruits, such as
plantain, orange, strawberry (Fragaria × ananassa), and Hass avocado (Persea americana L.)
(Motalab et al., 2022; Rozan et al., 2021; TACO, 2011). Furthermore, the Mn content of E.
pyriformis is higher than that of avocado pulp (0.30 mg) (Motalab et al., 2022).

3.2.6. M. dichotoma

M. dichotoma root flour has high carbohydrate content (24.2 g), although lower than
that of the more common species of the same genus Manihot esculenta Crantz (87.9 g) (TACO,
2011). However, it is similar to raw yam (Colocasia esculenta L.) in terms of carbohydrate
content (23.2 g) (TACO, 2011).
Although it is widely used as both fodder and human food, especially in periods of
scarcity, M. esculenta is reported to have toxic properties (do Nascimento et al., 2012; Nunes
et al., 2018). Adequate preparation is necessary before consumption to avoid poisoning, which

65

has been reported by local peoples over the years. In our review, no data on micronutrients were
found for this species.

3.2.7. S. mombin

The fruit does not have an expressive macronutrient content; nevertheless, it has gained
relevance for its micronutrient content. The major macromineral was Na (5.551 mg), with the
highest value among the analyzed species. S. mombin ranked second in K content (288.276 mg)
and third in P (32.849 mg) and Mg (15.095 mg) contents. Compared with avocado, a reference
fruit in terms of these macronutrients (K, 514.6 mg; P, 60.5 mg; Mg, 27.7 mg), S. mombin fruit
has low contents (Rozan et al., 2021). However, these values are higher than those of raw C.
sinensis fruit (TACO, 2011).
S. mombin fruit had the second-highest Cu (0.118 mg) and Mn (0.025 mg) contents,
behind only E. pyriformis. Its Cu content is higher than that of C. sinensis (0.04 mg) (TACO,
2011).

3.2.8. G. americana

Genipap pulp ranked third in the macromineral Ca (45.82 mg) and micromineral Fe
(0.80 mg). Genipap has a higher Fe content than strawberry (Fragaria × ananassa), reported
as 0.41 mg by (Motalab et al., 2022), and a higher Ca content than Hass avocado pulp (P.
americana), reported as 13.4 mg (Rozan et al., 2021).
The macro- and micronutrient contents are listed in Tables 4 and 5.

3.3. Woody species with the potential to promote food and nutrition security
Of the species for which macro- and micronutrient data were available, six stood out for
their nutritional value, namely A. occidentale (roasted and raw cashew nuts), E. pyriformis (fruit
pulp), S. mombin (fruit pulp), M. dichotoma (root flour), G. Americana (fruit pulp), and B.
cheilantha (seed flour). Although all eight versatile species have some nutritional value and are
versatile in terms of timber applications, demonstrating their importance for further
conservation and nutritional studies, we consider these six species as priorities for future

66

studies. A summary of the results on woody food species used as timber identified in this
systematic review is presented in Figure 4.

Figure 4. Number of woody food and/or timber species found during the stages of the
ethnobiological and nutritional reviews. Edited in Canva (free version).

67

Table 4. Macronutrient composition of parts of versatile food woody plants for wood uses. Data from the systematic review of nutritional studies
Species name
Analyzed
Preparation
Protein
Carbohydrates
Lipids
Sources
part
Pseudofruit
(peduncle)
Anacardium occidentale L.

Cashew nut
Cashew nut

Bauhinia cheilantha (Bong.) Steud.

Seed

type
Pulp

1.130*

-

0.666*

Singh et al. (2019)

22.67

19.86

47.79

Oliveira Sousa et al. (2011)

Fresh (Raw)

20.2

20.9

47.40

Rico et al. (2015)

Raw flour

35.9

6.2

8.7

Soaked flour

36.0*

7.0*

8.6*

Heated flour

31.5

12.8

8.6

Ground
(roasted)

Teixeira et al. (2013)

Eugenia pyriformis Cambess.

Fruit

Pulp

2.617

53.651

0.924

Silva et al. (2019)

Genipa americana L.

Fruit

Pulp

0.68*

-

0.35*

Figueiredo et al. (1986)

Manihot dichotoma Ule

Root

Flour

0.1*

24.2*

0.4*

Nascimento et al. (2012)

Cladode

Crushed

0.25

4.75

0.53

Nascimento et al. (2011)

Fruit

Pulp

1.15*

-

-

Souza et al. (2015)

Spondias mombin L.

Fruit

Pulp

1.06

13.9

0.62

Tiburski et al. (2011)

Ziziphus joazeiro (Mart.)

Fruit

Pulp

1.68

-

0.17

Oliveira et al. (2020)

Pilosocereus pachycladus F.Ritter

*Nutritional values in which the unit of macronutrient composition is percentage.

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Table 5. Composition of mineral content (mg/100g) in parts of versatile food woody plants for wood uses. Data from the systematic review of nutritional studies
Species name

Analyzed

Preparation

part

type
Ground

Anacardium

Cashew

occidentale L.

nut

Eugenia pyriformis

Fruit

(roasted)

Ca

Fe

Zn

K

P

Mg

Cu

Mn

Na

Oliveira
64.05*

3.89*

4.98*

556.16*

1101.04*

277.09*

3.08*

Sousa et al.
(2011)

Fresh (Raw)

28.0

5.1

5.0

540.0

470.0

240.0

-

-

Pulp

341.33*

5.37*

1.03*

134.00*

134.00*

41.00*

0.58*

3.05*

Cambess.

-

Rico et al.
(2015)
Silva et al.

-

(2019)

Genipa americana L.

Fruit

Pulp

45.82*

0.80*

-

-

33.50*

-

-

-

-

Spondias mombin L.

Fruit

Pulp

11.038*

0.327*

-

288.276*

32.849*

15.095*

0.118*

0.025*

5.551*

*Medium values.

Sources

Figueiredo
et al. (1986)
Tiburski et
al. (2011)

69

4. Discussion
4.1. Versatile woody food species used for fuel, construction, and technological applications
In this systematic review, we identified a significant number of native species that may
be suffering from intensive use (n = 42), because they are traditionally used as food and also
for timber purposes (fuel, construction, and technology) (see Table 2). These numbers might be
underestimated, given that many studies did not detail the categories of timber use, hampering
analysis of versatility. Studies on the harvesting patterns of timber resources have shown that
there are differences in replacement time, parts of plants used, harvest volume, and the state in
which the plant material is preferentially harvested (alive or dead) according to timber use (De
Medeiros et al., 2012). However, it is not yet possible to identify which usage is more harmful
to plant populations. Various studies indicated the potential effect of chronic anthropogenic
disturbances on gradual extinction of local species and alteration of vegetation structure
(Ribeiro et al., 2019, 2015). Therefore, the greater the timber versatility of a given species, the
greater the chances that it is somehow affected by unsustainable management strategies.
Of the identified versatile species, at least five are characterized as preferential fuel
wood in the Brazilian semiarid because of their physical properties, namely Anadenanthera
colubrina, B. cheilantha, Astronium urundeuva, A. occidentale, Z. joazeiro, M. dichotoma, and
Eugenia sp. (Ramos et al., 2008). It is possible that these species are suffering high use pressure
from local populations, given their good timber qualities.
A total of 31 of the 42 identified species are classified as least concern in the Red List
of the International Union for Conservation of Nature (IUCN, 2022). Moreover, eight are not
on this list, and three are considered important from a conservation point of view. Amburana
cearensis and Handroanthus serratifolius are classified as endangered, with a declining
population trend. Handroanthus impetiginosus is classified as near threatened, with a declining
population. Other species, despite being classified as least concern, have records of decreasing
population trends in some areas. This is the case of A. colubrina, Protium heptaphyllum, and P.
pachycladus. A. urundeuva is overexploited as timber and is classified as data deficient. This
category indicates that more information is needed for possible reclassification of the species
and that potential threats are not excluded.
Although there was no information on the nutritional characteristics of these species,
except for P. pachycladus, we believe that they must be included in conservation strategies. It
is also necessary to quickly identify whether any of these species have strategic importance for

70

food security, as this factor would be an additional argument for the creation of public
conservation policies.
4.2. Strategic species for food and nutrition security and biocultural conservation
Of the eight species with available nutritional data, six stood out in terms of
macronutrient (proteins, lipids, and carbohydrates) and/or mineral contents: A. occidentale, E.
pyriformis, S. mombin, M. dichotoma, G. americana, and B. cheilantha. Some of these are wellknown for their food applications, such as A. occidentale, S. mombin, and G. americana, whose
pulps are sold in various regions of Brazil. Figure 5 shows photographs of fruit pulps marketed
by a company in Alagoas State. In addition to occurring naturally in Brazilian ecosystems, these
three species are also domestically grown and cultivated on a small/medium scale (Araújo et
al., 2010; Mattietto and Matta, 2011; Rocha et al., 2015), contributing to the reduction of
conservation pressures.

Figure 5. Commercial fruit pulps of the species (A) S. mombin, (B) A. occidentale and (C) G.
americana. Edited in Canva (free version)

Species with high nutritional potential but little used as food (E. pyriformis, M.
dichotoma, and B. cheilantha) deserve special attention in conservation strategies, because they
are obtained almost entirely by extraction practices; there are few reports of their cultivation in
agroforestry systems (Florentino et al., 2007; Freitas et al. 2016). For species with low food
popularity, biocultural conservation strategies could include dissemination of their food
potential, which could contribute to income generation and reduction of timber extraction. In
fact, a study conducted in the Brazilian semiarid demonstrated that species with high medicinal
potential were less used for timber purposes than would be expected considering their
availability and wood quality (Silva et al., 2021). It is necessary, however, to test the hypothesis
of protective effects from food use in other socioenvironmental contexts.

71

Additional strategies that involve the entire plant community and not only species of
food interest are required, given that, if one species is protected from extraction, other timber
species may be targeted compensatorily, intensifying anthropogenic pressures on the latter.
Because low income in the countryside in certain regions of Brazil has been one of the greatest
intensifiers of logging for domestic purposes (De Medeiros et al., 2012; Specht et al., 2015),
strategies that generate income from the marketing of wild food plants could have a secondary
effect on timber use.
Sustainable extraction of species with nutritional importance can be achieved by their
inclusion in agroforestry systems, enhancing the supply of these products, increasing the
chances of successful food–timber management, and linking agriculture and extractivism. In
some regions of the country, however, small farmers are reluctant to make the transition from
conventional cropping to agroforestry, especially because of uncertainties regarding the success
of such systems, a possible decrease in the yield of the main crop, lack of successful models,
and limited knowledge on the subject (Sagastuy and Krause, 2019). Some of the difficulties
encountered by agroforestry farmers include the marketing of agroforestry products and
absence of public policies (Shennan-Farpón et al., 2022). Thus, a public approach toward
agroforestry production is essential to increase the cases of success and the number of small
producers and extractivists involved in the practice.
The biocultural conservation strategies discussed herein should be aimed not only at the
few versatile species with available nutritional studies but at all species with overlapping timber
and food potentials. Four of the five versatile food species with timber applications that did not
have nutritional information in the literature are classified as least concern in the IUCN Red
List (IUCN, 2022), namely Byrsonima sericea, Combretum leprosum, Monteverdia rigida, and
Ocotea glomerata. Although these species are currently classified in this category, our first
systematic review identified that these species are used as food and timber. In other words,
these species may be suffering from anthropogenic pressure with regard to timber extraction in
local and regional contexts, which could compromise their populations over time, hampering
their use in food systems.
Even for species with available data, it is necessary to carry out local diagnostics to
identify the real use pressure on these resources, the feasibility of incorporating them into
regional agroforestry systems, and the potential for expanding associated demand and
production chains. Studies focused on consumer behavior can be strategic for identifying
potential demands (Barbosa et al., 2021).

72

4.3. Recommendations for future ethnobiological studies

It is important to fill the knowledge gap on the sustainability of timber extraction of
species at the food–wood interface. In addition to biological conservation, the cultural
importance of these species must be investigated and preserved, given that these aspects are
inseparable (Gavin et al., 2015).
Many ethnobiological studies on food plants do not identify the edible parts that are
consumed or used in traditional preparations. This makes it difficult to carry out nutritional
analyses focused on specific parts of the plant that are appreciated or of commercial value to
local communities. This lack of information is also observed in timber research, as various
studies do not indicate the type of application of timber species. Considering the heterogeneity
of use dynamics among different timber categories (Walters, 2005) and the need to understand
the versatility of timber species, we recommend that further studies provide more details on the
timber uses of target species.

4.4. Study limitations

Given that this is a systematic review, it is important to highlight that species not
contemplated here may also be versatile and, therefore, could be included in the group of
keystone species. The key species identified might be biased by the research effort, which was
greater in certain ecosystems of the country, such as the Caatinga. This is due to the fact that
there is a higher proportion of ethnobiological studies in the Caatinga biome and that these
studies provided more information on timber uses. It is possible that there are many other
priority species for conservation in other ecosystems but that have not been as widely studied
as Caatinga species. The current study can be seen as a preliminary effort, which will need to
be augmented with new species through further investigations.

5. Conclusion
We identified a representative number of native woody plants that have overlapping
uses as food and timber. Such results underscore the need to assess sustainability and propose
conservation strategies for these species to ensure the continued existence of potential resources
for food and nutrition security. On the basis of ethnobiological and nutritional data available in

73

the literature, as well as ecological profiles, we recommend that E. pyriformis and B. cheilantha
be the target of ecological studies and popularization strategies because they are versatile in
terms of timber uses and have high nutritional relevance.
Acknowledgements
We thank the collaborators André José Rubião Cavalcante da Silva (AJRCS) and Emilly
Luize Guedes da Silva (ELGS) for their participation in the data collection stages of this
systematic review.
Funding
This work was funded by the Brazilian Fund for Biodiversity – FUNBIO, by
HUMANIZE and Eurofins Foundation (FUNBIO - Conserving the Future Scholarships,
granted to RAC, nº 025/2022), National Council for Scientific and Technological Development
(CNPq) (Scholarship doctorate to RAC, n° 141873/2020-5).
Conflicts of interest
The authors have no relevant financial or non-financial interests to disclose and have no
conflicts of interest to declare that are relevant to the content of this article.
Ethical approval and consent to participate
Not applicable for this research.
Consensus for publication
Not applicable for this research.
Other information
We declare that this review has not been registered.
Supplementary information
Additional File 1: PRISMA Protocol Checklist – Ethnobiology review
Additional File 2: Search strategies – Ethnobiology review
Additional File 3: PRISMA Protocol Checklist – Nutritional review

74

Additional File 4: Search strategies – Nutritional review
Availability of data and materials
Datasets that support the conclusions of this article are included in the article (and its
appendices). Other data referring to the list of articles of systematic reviews can be made
available on request.
Author statement
RAC – Conceptualization; Investigation; Methodology, Data curation, Writing original draft. EMCS – Organization and creation of figures; Writing - revision and editing.
RRVS e ARC – Supervision; Writing - revision and editing. PMM – Conceptualization;
Methodology; Writing - revision and editing. RZP – Methodology; Data curation, Writing revision and editing.

75

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CAPÍTULO 2

4. CONSERVATION OF WILD FOOD PLANTS FROM WOOD EXPLOITATION:
EVIDENCE SUPPORTING THE PROTECTION HYPOTHESIS IN
NORTHEASTERN BRAZIL

Manuscript submitted to Journal of Ethnobiology and Ethnomedicine

Roberta de Almeida Caetano¹,3*, Emilly Luize Guedes da Silva3, Luis Fernando Colin
Nolasco²,3, Rafael Ricardo Vasconcelos da Silva¹,3, Adriana Rosa Carvalho4, Patrícia Muniz de
Medeiros¹,3*

¹Postgraduate Program in Biological Diversity and Tropical Conservation (DIBICT-ICBS),
Federal University of Alagoas, Avenida Lourival Melo Mota, S/N, Tabuleiro do Martins,
Maceió – AL, 57072-900, Brazil
²Postgraduate Program in Ethnobiology and Nature Conservation (PPGEtno), Federal Rural
University of Pernambuco, Rua Dom Manoel de Medeiros, S/N – Dois Irmãos, Recife - PE,
52171-900, Brazil
3

Laboratory of Biocultural Ecology, Conservation and Evolution (LECEB), Campus of
Engineering and Agricultural Sciences (CECA), BR-104, Km 85, S/N, Rio Largo – AL, 57100000, Brazil
4

Federal University of Rio Grande do Norte, Department of Ecology, University Campus s/n,
Lagoa Nova, Natal - RN, 59098-970, Brazil

*Correspondence. Email: Roberta de Almeida Caetano (robertacaetano1991@gmail.com).
Address: Avenida Lourival Melo Mota, S/N, Tabuleiro do Martins, Maceió – AL, 57072-900,
Maceió - AL, Brazil.
Patrícia Muniz de Medeiros (patricia.medeiros@ceca.ufal.br). Address: Avenida Lourival
Melo Mota, S/N, Tabuleiro do Martins, Maceió – AL, 57072-900, Maceió - AL, Brazil

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Abstract
Background: The interplay between different uses of woody plants remains underexplored,
obscuring our understanding of how a plant's value for one purpose might shield it from other,
more harmful uses. This study examines the protection hypothesis by determining if food uses
can protect woody plants from wood exploitation. We approached the hypothesis from two
distinct perspectives: 1) the protective effect is proportional to the intensity of a species' use for
food purposes, and 2) the protective effect only targets key species for food purposes.
Methods: The research was conducted in a rural community within Restinga vegetation in
Northeast Brazil. During a participatory workshop, we pinpointed three food species vital for
both consumption and income (key species), along with their natural occurrence and collection
areas. A floristic survey in three distinct areas identified additional species coexisting with the
key species. Using a field herbarium and species photographs as visual stimuli, participants
assessed the species for wood quality, perceived availability, and usage. We employed
Cumulative Link Mixed Models (CLMMs) to evaluate the hypotheses “the food uses
(specialized) protect plants from wood exploitation (generalist)” from two different
perspectives (generalized protection and protection targeted at key species).
Results: Findings suggest there is no proportional protective effect from food uses across
species. However, domestic food use of key species exhibited a marked protective effect.
Perceived availability and utility emerged as notable predictors for wood exploitation.
Conclusion: We advocate for biocultural conservation strategies that enhance the food value
of plants for their safeguarding, coupled with measures for non-edible woody species under
higher use-pressure.
Keywords: Conservation through use. Ethnobiology. Traditional management. Wild Edible
plants. Wood uses.

Background
A growing body of research points to the potential effects of chronic anthropogenic
disturbances leading to the gradual extinction of local species and alterations in vegetation
structure(1,2). Among these disturbances, the impact of forest product utilization has been
highlighted, demonstrating that while wood use is crucial for local communities, especially in
developing countries, it often results in more pronounced impacts on plant populations ((3,4).
While wood exploitation exerts considerable pressure on plant resources, the
sustainability of non-timber forest product (NTFP) collection is also not always assured (5).
Nevertheless, the extraction of NTFPs, particularly the harvesting of wild fruits, generally has

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a lesser impact on forest structure and ecosystem functions than other uses (6). Moreover, the
consumption of NTFPs fulfills multiple roles, frequently underpinning rural livelihoods and
local economies, aiding food security, fostering trade, and preserving cultural traditions and
knowledge (7).
Hence, some researchers argue that discouraging the commercialization of wild food
plants may adversely affect the income of local populations, potentially leading to greater
reliance on other forest resources with more harmful consequences than food collection itself
(8). Conversely, the commercial value of NTFPs, coupled with the opportunity for income
generation, may incentivize conservation efforts among local communities for the forests that
supply these resources (9).
Since the 1990s, investigations into the sustainability of food plant use have sought to
determine the impact on species populations without conclusively addressing whether such use
confers protective benefits. In contrast, research on plant domestication supports the notion that
significant food value may lead to conservation practices, such as tolerance, protection, and
promotion (10). Plants with desirable traits may be maintained during deforestation or other
disturbances, promoted through distribution and dispersal, and specifically safeguarded against
competitors and herbivory (10).
However, the extent to which a plant’s significance for one use can shield it from more
destructive applications, namely the interaction effects among different utilization types,
remains underexamined. This gap hinders our comprehension of protective dynamics in socioecological systems and their economic benefits for humans. Such insights are vital for shaping
biocultural conservation frameworks that recognize the multifaceted advantages of maintaining
cultural practices intertwined with biodiversity.
It is conceivable that certain NTFP uses, including for food, may exert a protective effect
against more damaging activities such as wood exploitation. Although overharvesting of fruits

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has been shown to affect the regeneration of wild fruit trees adversely (11,12), food use is
typically seen as specialized, with minimal impact on plant populations, whereas wood
exploitation is often deemed generalist, posing broader threats (3).
The classification of plant uses as specialized or generalist may vary depending on the
social-ecological context. In the context of several South American communities, specialized
uses are defined by a narrower range of suitable plants meeting specific requirements, with the
specialization premise reinforced by observations that plant availability exerts little to no
influence on such uses (13,14). In contrast, generalist uses accommodate a broader spectrum of
species, with the most utilized often being the most accessible, as with many wood exploitation
practices (15).
Although quality may also be an important predictor of plant importance for generalist
uses (16), the generalist nature of wood use is supported by studies investigating the apparency
(availability) hypothesis, which posits a correlation between environmental availability and
species utilization (15,17,18). Therefore, for generalist applications, alternatives may spare
certain species for specialized uses, such as food, where fewer species can act as substitutes.
The protection hypothesis, initially proposed by Silva et al. ((19)), analyzed Caatinga
woody plants utilized domestically for medicinal and fuel purposes to assess whether medicinal
significance (specialized use) impacts wood exploitation (generalist use). Their findings
revealed a modest yet significant medicinal use effect on wood exploitation, providing
supportive evidence for the hypothesis as plants of greater medicinal value saw less wood
utilization. Moreover, Silva et al. (19) suggested that the protective effect could be more
pronounced in species with high medicinal importance.
The use of plants for food is also considered a likely candidate for conferring protective
effects against wood exploitation. Wild food plants are often crucial for providing essential
nutrients or for supplementing diets, playing a vital role in ensuring food security and offering

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economic benefits through the trade of these resources. Given food use’s specialized nature,
dietary importance, and economic potential, there is a presumption that communities may prefer
to preserve these plants from irreversible harm, such as wood exploitation. For the latter,
alternative species are available due to the generalist nature of wood use.
In this context, we investigate the protection hypothesis from two distinct perspectives.
We hypothesize that food uses (specialized) protect plants from wood exploitation (generalist).
We examined: 1) whether the protective effect is proportional to the intensity of a species' use
for food purposes, and 2) if a protective effect only targets key species for food purposes. Here,
'key species' denotes wild food plants of high regional importance for consumption and income.
This study is the inaugural inquiry into the protection hypothesis concerning the
protective effect stemming from food use. Moreover, unlike Silva et al. (19), our study
incorporates the commercial relevance of woody plants, providing income for the local
population. Methodologically, we refine hypothesis testing by employing the checklistinterview technique (20) to boost respondent recall, ensuring all associated uses (food and
wood) are considered.

Materials and methods
Study area and socioeconomic characterization

The research was carried out in a rural community within the coastal Restinga vegetation
of Piaçabuçu, situated on the southern coast of Alagoas state. Piaçabuçu spans an area of
243.686 km², housing a population of 15,908 individuals (21). It features a tropical 'As' climate
in the Köppen and Geiger classification, with an average annual temperature of 25.3ºC and an
annual rainfall average of 1283 mm (22). Notably, the municipality is designated with two

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Environmental Protection Areas (APAs): the federally instituted APA Piaçabuçu, established
in 1983, and the state-sanctioned APA Marituba do Peixe, created in 1988, both of which are
classified as sustainable use Conservation Units.
The APA Marituba do Peixe spans 18,556 hectares and extends over portions of the
Alagoan municipalities of Piaçabuçu (45%), Feliz Deserto (43%), and Penedo (6%) (23). This
area boasts diverse vegetation, including native Restinga, Várzea, and other forest formations
(23). Within the Indirect Influence Area of APA Marituba do Peixe lies the village of Retiro
(depicted in Figure 1), which was the focal point for the ethnobiological segment of this study.

Fig. 1 Geographic Location of the Retiro Community in the Municipality of PiaçabuçuAlagoas, Brazil.

The Retiro community is structured with a residents' association and a family farmers'
association. It is equipped with a primary healthcare unit and a municipal elementary school.

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The predominant faith among residents is Christianity, represented by two Catholic and two
evangelical churches. Currently, the community comprises approximately 288 families, a
decrease of 81 families since before the pandemic, as reported by Gomes et al.(24). This
discrepancy may be partly due to some families not being documented, a requirement for health
unit registration.
Retiro was selected for this study due to the local reliance on plant resources for both
food and wood. The community's economy is significantly driven by the extraction and
commercialization of wild food plant fruits (24), along with shrimp and fish (25). Wood
resource extraction for personal use and commerce, particularly firewood, charcoal, and
materials for fencing, is also prevalent. These resources are marketed through open markets or
direct orders in Piaçabuçu and Penedo, whereas wood products are solely distributed by order.
Firewood is the primary cooking fuel in the community, though some households use
both cooking gas and firewood. Meals are typically prepared on traditional clay or makeshift
brick stoves. Firewood also serves in roasting shrimp and baking cakes from rice straw, a
common bait for shrimp in local fishing gear known as cóvu.
Architecturally, many taipa houses (rammed earth) are present within the community,
often serving as dwellings for individuals from other regions staying temporarily in the area.
The primary livelihoods include gathering, particularly collecting edible fruits, as well
as retirement, fishing, and agriculture, with some engaging in multiple occupations. A variety
of other professions are represented to a lesser extent. The demographic profile of interviewees
spans ages 18 to 82. Most interviewees are literate (76.65%) are literate, of whom 73.91% have
completed or partially completed basic education, and 1.74% have higher education
qualifications. The number of people occupying the residences ranges from one to seven
residents. However, the majority of houses are occupied by: two or three residents (29.57%),

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followed by one or four resident(s) (15.65%). Household incomes show substantial variation,
as show the frame 1.
Frame 1 Monthly household income and the corresponding percentage of interviewees who
possess it.
Monthly household income
% of household
Under one minimum wage
28.70
One minimum wage
14.78
One and a half to two minimum wages
41.74
Up to three minimum wages
13.04
More five minimum wages
1.74

Ethical and legal aspects of the research

This research project received approval from the Research Ethics Committee by Federal
University of Alagoas (UFAL), Nº 1998673, securing authorization for studies involving
human participants as per the stipulations of National Health Council Resolution 466/2012.
Additionally, scientific activities involving the collection and transport of botanical specimens
within the Marituba do Peixe Environmental Protection Area were duly registered with Chico
Mendes Institute for Biodiversity Conservation/Biodiversity Authorization and Information
System (ICMBio/SISBIO), Nº 87112-1.
To ensure ethical compliance, all community members aged 18 and older who
comprehended the objectives of the research and consented to participate were asked to provide
a signature or thumbprint on the Informed Consent Form (ICF), as well as on the image use
authorization form.
Data Collection
Data collection was carried out in three distinct phases: a participatory workshop, a
forest inventory, and checklist-interviews, all of which are depicted in Figure 2.

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Fig. 2 Stages of ecological and ethnobiological data collection: (1) Participatory workshop for
identifying key species and collection areas; (2) Forest inventory to detect species co-occurring
with key species, accompanied by botanical collection and plant photography; (3) Checklistinterview utilizing visual stimuli, including photos and exsiccates (field herbarium). Image
edited using Canva (free version).

1st Data Collection Stage: Participatory Workshops

Participatory workshops with the residents of the Retiro community aimed to identify
significant wild food plant species for consumption and commercial use, as well as their
harvesting locations. These workshops were facilitated by local leaders and a researcher from

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the Laboratory of Biocultural Ecology, Conservation, and Evolution (LECEB), who had
previously interviewed community members.
In the inaugural workshop, residents listed wild food plants harvested for sale or
personal use. They documented the common names on cardboard, selecting eleven for further
discussion. Participants then evaluated these plants for their economic significance within the
community, ultimately ranking the top five in importance.
Additionally, the workshop served to note wood resources tied to food plants and their
utilization for consumption and commerce within the community.
Thirteen women and three men, ranging in age from 31 to 82, contributed to this first
workshop. While all were identified as gatherers, some also engaged in agriculture and fishing.
A follow-up workshop sought to enrich this data with contributions from another set of
gatherers (n=17), including eight newcomers. This session, comprising thirteen women and
four men from the same age bracket, validated the initial findings regarding species and
harvesting sites.
Participants utilized a detailed satellite image from Google Earth to denote areas
frequented for food and wood collection. An overlay of transparent acetate allowed them to
make corrections directly on the map.
Subsequent to pinpointing these areas, we selected those most frequented for the
harvesting of both food plants and wood, prioritizing locations where the ranked key species
were prevalent. Among the listed species - Myrciaria floribunda H. West ex Willd.) O. Berg,
Genipa americana L., Psidium guineense Sw., Spondias mombin L., and Tamarindus indica L.
- only the first three were deemed key for this study due to their abundance in native and less
disturbed habitats.

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Three sites were thus chosen for the forest inventory: two with a natural predominance
of key species and one characterized by a more generalized distribution of various plant species,
including those bearing edible fruits.
In addition to these participatory workshops, our study relied on complementary
previous information reinforcing the high economic and consumption importance of these three
plant species in the Retiro community. This information was obtained from: 1) direct
observations of residents' routines and 2) a previous doctoral research study (24).
The research design we adopted gave precedence to examining species that occurred
alongside the key species; consequently, not all food and wood plants were included in our
scope. Notably, Schinus terebinthifolia Raddi, while not a key species within Retiro, is a
significant commercial species in the community and the most important commercial plant in
neighboring areas, such as the Fazenda Paraíso settlement (24).

2nd Data Collection Stage: Forest Inventory and Field Herbarium

The research included a forest inventory as part of a larger investigation of our research
group into plant resource utilization within the region. For the purposes of this study, the forest
inventory was only used to identify species that co-occur with the key species. The sites selected
for the inventory were privately owned yet accessible to local gatherers. Two of these sites fell
within the Marituba do Peixe APA boundaries in Piaçabuçu, while the third was in the
municipality of Penedo, outside the APA but still proximal to the community.
We established five permanent plots, each measuring 50 x 20 meters, and further divided
these into 50 smaller subplots of 10 x 10 meters situated within the primary native vegetation
gathering sites designated during the workshops. This amounted to 0.5 hectares per area, with
a total of 1.5 hectares surveyed across all areas.

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During the inventory, we collected at least three reproductive samples of each plant
species within the plots for identification and to assemble a field herbarium for use in
subsequent interviews. Certain species, commonly referred to as Ingá and Pau d'arco, lacked
fertile material at the time of collection, leading us to categorize them as ethnospecies for the
purposes of this study. Consequently, in our identification records, we referred to these simply
as Ingá and Pau d'arco, acknowledging that these common names might represent multiple
botanical species. Furthermore, the ethnospecies Cambuí, although biologically uniform –
belonging to the species Myrciaria floribunda (H. West ex Willd.) O. Berg – was recognized
by some residents as having different ethnovarieties – a distinction not universally
acknowledged. In our analysis, we accounted for each mention of Cambuí by participants, even
though the general data summary did not differentiate between ethnovarieties. For instance, if
Interviewee A identified two types of Cambuí (Yellow and Red) and Interviewee B referred to
one (a general Cambuí), we recorded two entries for A and one for B in our database.
For the field herbarium, we mounted exsiccates from species with more than 15
individuals in the surveyed areas onto duplex paper of dimensions 42 x 29.7 cm and stored them
in folders of matching size. The herbarium included 24 species in total and 2 taxa that were
treated as ethnospecies.
Photography of each species was conducted in situ, capturing images that emphasized
the plants' distinguishing features: overall appearance, flowers and/or fruits, branches, and
stems. These photographs were compiled into folders on a tablet, which was employed to
display the images during interviews. Both the exsiccates and the photo folders were
numerically coded to correspond with the identifiers on the interview forms, ensuring that
interviewees were unaware of the plant names and assisting the interviewer.

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The botanical collection phase commenced in November 2021 and concluded in April
2023, an extended period due to intermittent interruptions from COVID-19 peaks and flooding
that hindered fieldwork.
A local guide with extensive knowledge of the vegetation provided assistance for all
fieldwork involving local vegetation access. We adhered to standard botanical collection
protocols, and the exsiccate samples were deposited at the IPA Herbarium (Instituto
Agronômico de Pernambuco) - Dárdano de Andrade Lima.

3rd Data Collection Stage: Checklist Interview

Before commencing the interviews (third stage), we mapped all Retiro households in
May 2023. This mapping was imperative for sample size calculation due to the absence of a
census record; the health unit's data was limited to registered families. We determined that
household heads (one per household) aged 18 or older present during our visit would be
interviewed. Considering that some individuals reside in the community only for short periods,
we established an inclusion criterion that only families living in the area for more than one year
would be eligible for the study.
We ascertained the number of residences, including both occupied and vacant, to be
361, initially yielding a sample size of 187 residences based on a 95% confidence level and a
5% margin of error. Subsequently, we conducted a simple random selection.
As every house in the community was recorded, including unoccupied ones, some
selected residences were vacant. Additionally, given the research's focus on potentially harmful
wood resource use within the APA, some families were reluctant to participate. Therefore, from
the 187 chosen residences, we could only conduct interviews in 81 interviews of them. To

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overcome refusals, flood-affected houses, and temporary residents, additional draws were
made.
After all draws, we excluded unoccupied houses (n=82), residences on flood-impacted
streets (n=12), households temporary inhabitants (n=18) and refusals or unavailability (n=74)
from the sample. After three unsuccessful attempts to locate a household head, we inferred their
non-participation.
A notable number of individuals opted out of the study, a figure aligned with
expectations for wood use research in protected areas, mirroring findings from Medeiros et al.
(3). The considerable number of unoccupied houses in the community can be primarily
attributed to their use as summer residences by individuals from nearby municipalities, taking
advantage of the community's closeness to the beach. Additionally, a number of these houses
are situated in areas susceptible to flooding during the rainy season, which also contributes to
their vacancy.
The final sample consisted of 115 individuals—81 women and 34 men. Interviews were
conducted from May to July 2023. During interviews, we applied the checklist-interview
technique (20) to ensure uniform visual stimuli across all informants, enhancing recall of all
plant-associated uses.
Interviewees were shown photos of each species and queried on whether they
recognized the species. Affirmative responses led to further questions on the plant's name, its
uses (food and wood), whether the interviewee actually used the species, parts utilized,
commercial harvesting, and collection and sale sites. For recognized plants, a Likert scale rated:
perceived availability (only for those interviewees that often frequent vegetation areas), wood
quality by use category (fuel, construction, technology), domestic use for wood and food, and
commercial use. The ratings and responses are presented in Figure 3.

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Fig. 3 Information collected using a Likert scale on the variables considered in this study.

During interviews, we outlined the three primary categories of wood use considered in
our study: fuelwood, construction, and technology. This classification facilitated the synthesis
of scoring for perceived wood quality, allowing individuals to assign ratings by category rather
than for each specific use. If a participant identified a plant as useful for wood but did not
personally use it, we probed for the reasons behind this choice. We also asked if there were any
of the mentioned plants that, despite being good for wood uses, the interviewee did not collect.
These questions were posed to obtain qualitative responses about the protection of food plants
from wood uses. Additionally, we gathered socio-economic data from all informants through
structured interviews, including gender, age, occupation, income, place of origin, education,
and length of residence.

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Only for the ethnospecies Ingá and Pau d'arco, instead of showing the photos and
exsiccate, we asked directly if the person knew them for food or wood uses. In case of a positive
answer, we asked the same cycle of questions conducted for the other species. This was done
because we did not obtain sufficient fertile material for the taxonomic identification of all
species of Ingá and Pau d'arco during the various collection events.

Data Analysis

For statistical analyses, we removed from the database any instances where species were
identified for food purposes but not for wood purposes, as the focus of the research was on
criteria for selecting wood plants. Consequently, non-wood plants were disregarded. Similarly,
we excluded data from individuals who did not frequent forest environments to ensure that our
information on species availability came from realistic assessments.
Our response variable, domestic wood use, was ordinal, as depicted in Figure 4.
Therefore, we utilized Cumulative Link Mixed Models (CLMMs), incorporating the
interviewee as a random effect to account for the non-independence of information from the
same individual. The CLMMs were executed using the clmm function from the R package
ordinal.
To evaluate the stability of our models and check for multicollinearity, we used the
omcdiag function from the mctest package in R. We determined an absence of multicollinearity
if none or at most one of the six indicators were positive. To circumvent multicollinearity, we
constructed two models. The first model, termed the widespread protection model, assigned
domestic and commercial food use values on a 5-point Likert scale based on reported usage
intensity. For the key species-based protection model, food use was a binary variable: it took

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the value of 1 if the mention included the use of a key species, and 0 if the mention involved a
key species only known but not used, or non-key species, regardless of usage.
Model selection was based on the most parsimonious option, as indicated by the lowest
Akaike Information Criterion corrected for small sample sizes (AICc). We interpreted a ΔAICc
(difference from the lowest AICc) of less than 2 as substantial support for the model's inclusion
among the best set of models, following Burnham and Anderson (26). Following model
selection, we computed a model average, which considered the average beta of all variables
within the parsimonious models. Since the variables were standardized via z-standardization,
we compared the relative effect sizes of all variables.
The variable 'commercial wood use' was not included in the models due to its limited
mentions (n=5) within the community and only six citations of species that are commercially
traded for wood, exclusive of domestic use.
In addition to the explanatory variables related to food use, both models incorporated
control variables for availability and quality, as previously identified in the literature as
predictors of wood use (16–18). Our quality indicator was the maximum perceived quality. It
was determined by the highest Likert scale quality rating given by an interviewee for a species
across the three categories of wood use. For example, if, for a given species, values of 3, 4, and
5 were assigned by an interviewee to the categories of construction, technology, and fuelwood,
respectively, the maximum perceived quality would be recorded as 5.

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Fig. 4 Widespread species model and key species-based model with their variables and
respective measures.

Results
Wood and food uses: general aspects

All plants were recognized to varying extents by the interviewees. The most recognized
species/ethnospecies were: Genipa americana L. (Jenipapo), Inga spp. (Ingá), Myrciaria
floribunda (H. West ex Willd.) O. Berg (Cambuí), Manilkara salzmannii (A.DC.) H.J.Lam
(Massaranduba), Psidium guineense Sw. (Araçá), Mouriri sp. (Cruirí), and Bignoniaceae spp.
(Pau d’arco), with recognition rates of 56.5% or higher during interviews. The first five species
achieved high recognition levels, exceeding 80%. Notably, G. americana, M. floribunda, and
P. guineeense were identified as key species during the workshops. A comprehensive list of all

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species included in the checklist, along with their recognition and citation frequencies, is
presented in Table 1.
Over half of the species on the checklist (58.33%) were recognized for both food and
wood uses. Within the three categories of wood use addressed in this study, fuelwood (37.89%)
and construction (36.93%) had the highest citation percentages. Technology accounted for only
25.18% of wood citations. Within the fuelwood category, firewood led with the highest
percentage (62.82%) of citations relative to the total uses in the category, followed by charcoal
(37.18%). The construction category comprised 25 wood uses, with over half (50.55%) the
citations pertaining to fences, and the remainder divided among uses such as line (11.98%) and
rafter (10.74%). The technology category included 67 wood uses, featuring lower usage
percentages compared to the other categories. Uses such as hoe handle (11.92%) and hoe shaft
(10.30%) were the only applications exceeding 10% of citations in relation to the total uses
within this category. All wood uses attributed to the species are detailed in Table 2.

Please, put the table 1 and 2 here.

Widespread protection model
In the widespread protection model, domestic and commercial food use did not
significantly influence domestic wood use when controlling for availability and quality
variables (Figure 4). Quality and availability were significant predictors of domestic wood use
in the model. This suggests that, within the local context, there is a tendency to use woody
plants for wood purposes based on their higher quality and greater availability.
Key species-based protection model

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We observed a pronounced protective effect on key species, where the domestic use
variable was more influential than both perceived availability and wood quality (see Figure 5).
However, the variable indicating commercial use did not significantly affect the use of wood
for domestic purposes.
Within the model focusing on key species, both availability and wood quality
(considered as control variables) had a significant impact on wood use. Consequently, our
findings imply the existence of a threshold level of importance for the protective effect of food
use on wood uses. This indicates that only those plants with substantial domestic food
importance are shielded from being utilized for wood by the local population. The complete
statistical results are available in the Additional file 1.

Fig. 5 Impact of Predictor Parameters (quality, availability, domestic food use, commercial
food use, domestic use of key species, and commercial use of key species) on domestic wood
utilization of wild edible plants. Left: widespread protection model. Right: key species-based
protection model. The central circles indicate the median coefficient estimates of the
associations, and the horizontal lines delineate the 95% credibility intervals. The parameter
coefficient estimates are plotted along the x-axis, while the predictor levels are represented on
the y-axis. The vertical line intersecting the zero point on the x-axis (indicating no effect)
facilitates comparison of the sizes of positive, negative, and null effect coefficients. In the
parameter level grouping, non-overlapping horizontal bars denote significant differences.
Horizontal bars intersecting the zero line on the x-axis signify a non-significant effect.

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Evidence of protection based on qualitative data
When inquiring whether individuals refrained from harvesting any of the recognized
plants for wood purposes, despite acknowledging their suitability for such use, we gathered
responses that support a tendency to protect certain species with dual edible and wood
functions. The key species identified during the participatory workshop as significant to the
local community, and which garnered substantial recognition in the checklist, were notably
prominent in this context.
Out of the 60 respondents to this question, 37 reported no restraint in harvesting plants
suitable for wood purposes. Among the 23 participants that chose not to collect certain plants,
12 indicated not collecting species had both edible and wood uses.
Of all mentions of plants with both edible and wood applications, seven pertained to key
species (as shown in Frame 2). The primary rationale for sparing these species from wood
harvesting, or only using their dry branches, is their provision of edible fruits valued within the
community. This rationale is illustrated by the testimonies concerning P. guineense, G.
americana, and M. floribunda.
Frame 2 Responses from interviewees indicating the protection of key species regarding wood
uses.
Key species
Psidium guineense Sw.

Genipa americana L.
Myrciaria floribunda H. West ex
Wild.

Explanations
"Because if you take the wood, it will dry, and the plant
will stop bearing fruits”.
"They are good (as wood), but they are also fruits."
"Because the fruit is good and sought after by the people,
if it's green, I don't take it, only if it's dead and dry."
"I don't take the female one because it bears fruit."
"Because I really like the fruit, and I find it very beautiful."
"Because it's a nostalgic, good fruit."
"I avoid taking them because they are fruits. I only collect
the dry branches."

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Manilkara salzmannii though having a limited role in commerce, garners mixed views
on its suitability for consumption within the community. Nonetheless, six interviewees
mentioned the species, with two specifically expressing their intent to conserve it from being
used for wood purposes: 1) "I don't take it, thinking about the fruits and the future. I don't like
to take it (wood) while it's still green, I only pick up the dry branches that have fallen on the
ground." 2) "Because it's a plant that bears fruit, and it doesn't sprout again if you cut it."
Despite M. salzmannii not being designated as a key species during the participatory
workshop, it nonetheless received noteworthy acknowledgment in the checklist-interview. This
suggests that M. salzmannii may possess a certain degree of importance for food-related uses
within the community.

Discussion
In our widespread protection model, neither commercial nor domestic food use
significantly explains domestic wood use. By contrast, in the key species-based protection
model, domestic use emerges as the primary explanatory variable. In both models, perceived
availability and quality significantly explain wood use, with quality being more important than
availability.
Consistent with our hypothesis, we identify a protective effect of food use on wood use.
This effect is not directly proportionate to the food use of the species but is confined to plants
with considerable domestic food importance. Research conducted in the Brazilian Caatinga
region, which initially tested the protection hypothesis using medicinal (specialized) and wood
(generalist) use, suggested this possibility (19). Although they observed a modest yet significant
linear trend supporting the hypothesis, the authors graphically demonstrated that the protective
effect intensified specifically among highly valued medicinal plants. This study furnishes
statistical substantiation for what was previously inferred graphically.

102

Given that the protective effect is selective for key species, it indicates that merely
having intermediate or low food importance is insufficient for wild food plants to evade wood
use. Protection is afforded only to those species recognized as highly important. Indeed, key
species not only receive high acknowledgment in the checklist (>80%) but are also extensively
consumed and increasingly traded within the community, in forms such as fresh fruit, juice
pulp, and in the manufacture of alcoholic beverages, ice pops, among other products. Literature
highlights that elevating the value of non-timber forest products for local populations acts as an
incentive for forest species conservation (9,27).
Our findings suggest that protection is predominantly correlated with domestic
consumption. The domestic use of non-timber forest products can be a way for poorer local
populations to save money (28), as is the case with edible fruits that can replace commercially
purchased foods. Although wild food plants serve only as supplementary food resources within
the community—with staple crops like rice and beans constituting the primary plant food
intake—the cultural significance of key wild food plants likely motivates the observed
protection behaviors.
The absence of protective behaviors reported in interviews could stem from various
factors. Not all protective actions are necessarily conscious. Additionally, individuals may
inadvertently omit mention of such behaviors in response to indirect inquiries like those posed
in our study. Furthermore, protection may not be universally practiced within the community,
and while the pressure to use wood from wild food plants may not be entirely eliminated, it
could be reduced by fewer community members intensively exploiting key food plants for wood
purposes.
Wood quality and species availability are significant determinants of wood use. It
appears that, aside from key food species—whose utilization for wood is limited due to their
value as food—other species are more likely to be used for wood purposes when they offer

103

better trade-offs between availability and quality. Most studies that investigate the drivers of
wood use tend to analyze quality or availability indicators separately, rather than in
combination. These studies have found that either quality or availability can influence wood
use (16–18).
Studies that consider multiple predictors of wood use have yielded divergent results.
While availability seems to be a consistent predictor across different contexts, quality may or
may not be a determinant of wood use (29,30).
In various social-ecological contexts, research has indicated that trade-offs between
multiple variables act as drivers of plant resource use (29,31). However, these trade-offs are
often considered within a single use-category (e.g., the trade-off between quality and
availability to explain fuelwood use). Therefore, the evidence of protection underscores the
necessity of considering interactions between use-categories when evaluating criteria for plant
resource selection (Figure 6).

104

Fig. 6 Hypothetical example of a trade-off between availability and quality explaining fuelwood use. In a simplified scenario where these are the
only predictors of plant use within the fuelwood category, the most utilized species would be those exhibiting the highest trade-offs between
availability and quality (represented by the blue dots in the right and left graphs). When considering the interaction with the food use-category
under the key-species based protection model, the use of wood species with low to intermediate food importance would be proportional to the
trade-off between quality and availability (graph on the right). However, for species that are considered key food plants (indicated by the dark
green dot), their utilization for fuelwood would be less than what is predicted by their quality and availability alone.

105

1

Recommendations for conservation strategies for plant species

2

The practical implication of a protective effect that acts solely on species of high food

3

importance is that species recognized as having intermediate or lower importance remain

4

unprotected, as do wood species without any associated food use. Moreover, if only a few

5

species are highly valued for food, they might experience intense pressure from their use as

6

food or be protected at the expense of other species. Therefore, we recommend that

7

conservation strategies be mindful of these dynamics.

8

For species with intermediate or lower food importance, popularization strategies could

9

prove beneficial to enhance their perceived value. Programs aimed at popularizing such species

10

are crucial, as they may significantly contribute to food and nutritional security, while their use

11

as food might concurrently protect them from being exploited for wood. These programs should

12

establish incentives that encourage community members to use these resources sustainably.

13

However, the effectiveness of this approach should be continuously monitored, as if cultural

14

significance is the primary factor driving protective behaviors, integrating other wild food

15

plants into the set of key species may prove challenging.

16

Although the commercial importance of key species did not lead to protection in this

17

study, the inclusion of certain species in local markets could also positively influence domestic

18

use. Thus, popularization strategies could extend beyond local communities, emphasizing the

19

importance of these plants for diet diversification and their potential nutritional value to

20

generate demand for products sourced from local communities. One method to achieve this is

21

through marketing campaigns that raise awareness about the significance of these plants in local

22

markets and across social and conventional media platforms (32).

106

23

However, it is crucial to approach the popularization of highly important food species

24

with caution to prevent the oversimplification of the plant community, as observed with açaí

25

(Euterpe oleracea Mart.), where management practices have simplified estuarine communities

26

in the Amazon Rainforest (33).

27

For wood species that lack an associated food use, conservation strategies must be

28

implemented to mitigate the pressure on their exploitation. Considering that the primary wood

29

uses in the community are for fuel (firewood) and construction (fencing), conservation efforts

30

should be tailored to these applications. Firewood is the most commonly cited use in the Retiro

31

community, and due to its characteristics regarding short replenishment time and large volume

32

of wood used, it poses a significant threat to species conservation, depending on the collection

33

method (green or dry). For people with greater social vulnerability, firewood is an important

34

resource for cooking. To address this, we recommend the use of efficient wood stoves. These

35

stoves, through their structural configuration, reduce cooking time and, consequently, the daily

36

volume of wood used and deforestation compared to traditional stoves (34).

37

Although there is controversy in the literature regarding the long-term economic costs

38

and benefits of improved stove use in developing countries (35) and their efficiency(36), several

39

studies have shown significant reductions in firewood use with the adoption of this technology

40

(34,37,38). For instance, a study based on an improved stove intervention in the Chalaco

41

District, Northern Andes of Peru, recorded a 46% reduction in firewood consumption

42

(approximately 650 kg of firewood per household throughout the rainy season) among

43

households that properly used improved stoves during winter (38). Similarly, Bensch and Peters

44

(37), who evaluated the impact of these stoves in rural Senegal through a randomized clinical

45

trial, found a total 31% reduction in firewood consumption over one week. Additionally, the

107

46

use of efficient stoves can contribute to a higher quality of life for users by reducing smoke

47

from wood combustion, which can cause respiratory diseases (39). However, for successful

48

implementation of efficient stoves, besides local community interest, factors influencing long-

49

term adoption, such as maintenance costs, need to be considered.

50

An alternative to replacing firewood use is increased investment in public policies that

51

ensure access to Liquefied Petroleum Gas (LPG). While families receiving the gas voucher

52

through the federal government program (Bolsa Família) still use a mix of LPG and firewood

53

in the community, education, health, and human well-being initiatives, combined with these

54

public policies, may have a better response in the community during the transition from

55

firewood to LPG use. This is especially important considering that the use of firewood, for the

56

most part, spans generations. The same applies to the transition from traditional or makeshift

57

brick stoves to efficient stoves.

58

To reduce the use of species employed in the construction of dead fences - where trunks

59

and branches of woody plants are cut green for use - we recommend a gradual replacement with

60

species used as living fences, which are kept alive. This strategy has been indicated as effective

61

as it represents a gene bank of native species and contributes to the maintenance of these species

62

(40). "Rompe gibão" (Phyllanthus sp.) and "Cruirí" (Mouriri sp.) were mentioned by some

63

interviewees as species used for living fences, and "Peroba" (Tabebuia elliptica (DC.)

64

Sandwith) was mentioned as having the ability for its stake to remain green in a humid

65

environment. They are considered hard and resistant woods ("fixe") by the interviewees who

66

recognized them on the checklist. These species could potentially be used for this purpose, but

67

they need to be evaluated in terms of their characteristics and ecological status.

108

68

Finally, although our results admit that there is a protective effect on species with high

69

food importance (key species) regarding wood uses, it is necessary to investigate the ecological

70

status of these species to assess whether harvesting is being done sustainably and if

71

overexploitation of these species is not occurring, as has been identified in other studies with

72

non-timber forest products (5,11,12).

73
74

Recommendations for future ethnobiological studies

75

Some challenges for testing the protection hypothesis in future studies include:

76

Studies should account for the interactions not only between two use-categories but also

77

among all use-categories associated with the plant species. For instance, a plant might be

78

protected from wood uses not solely due to its food or medicinal value, but because it serves

79

multiple purposes. Thus, protection may only become apparent when evaluating the full

80

spectrum of plant use dynamics.

81

Gender and age variables ought to be incorporated into the tests of the protection

82

hypothesis, given that individuals of different ages or genders may protect plants for varied

83

purposes.

84

Studies could delve into the affective aspects of protection, as these may inspire

85

individuals to spare certain species from wood uses due to resources that evoke positive

86

affective memories. For example, a fruit that was greatly cherished during one's childhood or

87

that constituted the main sustenance for a person's family might be protected. While affective

88

reasons are personal, common patterns may surface, especially among individuals with similar

89

cultural or community backgrounds who may share collective memories.

109

90

It's necessary to further investigate the influence of social organization on the protective

91

behaviour of local peoples towards wild food species. For instance, in contexts where there are

92

associations of fruit gatherers or cooperatives, protective behaviour may increase compared to

93

rural communities where social organization is poorly established or absent. Alternatively,

94

protective behaviour may be directed on an individual basis.

95

Research designs should enhance the methodological approach concerning qualitative

96

evidence for protection. The questioning should be crafted to elicit precise responses without

97

leading the participant, yet still addressing the core issue effectively. Our study utilized indirect

98

questions that may not have fully captured our main objective. We propose that future research

99

adopting discourse analysis techniques (underpinned by multiple theoretical frameworks)

100

would yield valuable insights.

101
102

Limitations of this study

103

For two groups of plants treated in this study as ethnospecies (Pau d'arco and Ingá), we

104

were unable to elucidate their taxonomies despite our efforts. Our results suggest that these

105

ethnospecies are not under the protective effect of food use, and the lack of botanical

106

identification complicates the targeting of conservation strategies, especially for future studies

107

in this region. Although we don't know the quantity and specific species, we suspect they are at

108

risk of threat due to logging, especially for Pau d'arco (Bignoniaceae spp.). At least two species

109

of Pau d'arco are listed in the International Union for Conservation of Nature Red List with

110

concerning ecological statuses: Handroanthus impetiginosus (Mart. ex DC.) Mattos (Pau

111

d'arco rosa) listed as near-threatened and Handroanthus serratifolius (Vahl) S. Grose (Pau

112

d'arco amarelo), listed as endangered (41). Both were assessed for the list in 2020. As

110

113

respondents mentioned three types of Pau d'arco (roxo, amarelo, and branco), it's possible that

114

species of this genus are included. Through botanical identification, we identified that a plant

115

known in the community as Peroba is the species Tabebuia elliptica (DC.) Sandwith (White

116

Pau d'arco), specified on the Red List with a status of least concern. This makes this area an

117

interesting occurrence for this plant group. In light of this, we acknowledge this limitation in

118

our study and invite other researchers specializing in these plant groups to direct research efforts

119

in this region and clarify the taxonomy of these species.

120

Our data on the perceived quality of wood were collected from a single Likert-scale

121

value considering all wood uses of the plant reported by the interviewee for each wood use

122

category, instead of considering the quality for each reported wood use in each category. This

123

optimized data collection. However, the heterogeneous nature of categories such as technology,

124

where the wood quality of the plant can vary significantly among uses (e.g., tools, furniture,

125

boat), can be challenging for the interviewee to assign a single rating considering various

126

distinct uses. This may have biased our results with very generic perceptions of species quality.

127

Given that wood use is diverse, future studies could consider a more meticulous design, such

128

as focusing on the most relevant uses within each category in the local community and assessing

129

their perceived quality independently.

130

The inclusion of species for the composition of the checklist interview was based on

131

their availability in areas of co-occurrence with key species. Although greater availability of

132

species is a potential indicator of higher use, it is not universal. There may be species in the

133

sampled vegetation areas that are less available precisely because they are under greater use

134

pressure or due to other environmental or intrinsic factors not considered in this study.

135

Therefore, it is essential to also consider ecological approaches in research to have an overall

111

136

assessment of the impact of such uses on the plant community structure, even if the focus of

137

the research is on the most important plant species.

138
139

Conclusion

140

We have identified a protective effect of domestic food use against wood exploitation,

141

specifically directed towards key species. Consequently, we encourage future studies to test the

142

protection hypothesis within various socio-environmental contexts and we suggest considering

143

two distinct perspectives: generalized protection and protection targeted at key species.

144

In light of our findings, we advise that species demonstrating an overlap between food

145

and wood uses, yet possessing intermediate or lower food importance as detailed in Table 1,

146

should be prioritized in popularization strategies to raise their significance. Moreover, species

147

solely used for timber, which do not benefit from food-related protection, also require attention

148

through biocultural conservation strategies. Given that the protective effect is limited to a select

149

number of plant species, these species warrant further ecological investigation to determine

150

their conservation status within their natural habitats, to identify whether they face increased

151

pressure from their use as food, and to ascertain if their prominence is leading to a reduction in

152

plant diversity.

153

Abbreviations

154
CLMMs
NTFPs
APAs
UFAL
SISBIO

Cumulative Link Mixed Models
Non-Timber Forest Products
Environmental Protection Areas
Universidade Federal de Alagoas (Federal University of
Alagoas)
Sistema de Autorização e Informação em Biodiversidade
(Biodiversity Authorization and Information System)

112

ICMBio
ICF
LECEB
AICc
LPG

155

Instituto Chico Mendes de Conservação da Biodiversidade
(Chico Mendes Institute for Biodiversity Conservation)
Informed Consent Form
Laboratory of Biocultural Ecology, Conservation, and Evolution
Akaike Information Criterion
Liquefied Petroleum Gas

156

Supplementary Information

157

Additional file 1 - Multicollinearity diagnosis and model average for the widespread protection

158

model and the key species-based protection model.

159
160

Declarations

161

Ethics approval and consent to participate:

162

This study was approved by the Ethics Committee in research by Federal University of

163

Alagoas (UFAL), Nº 1998673. Furthermore, all participants previously signed the Informed

164

Consent Form (ICF).

165

Consent for publication

166

Not applicable

167
168
169

Data Availability
Data is provided within the manuscript or supplementary information files.

170
171

Competing interests

113

172

I declare that the authors have no competing interests as defined by BMC, or other

173

interests that might be perceived to influence the results and/or discussion reported in this

174

paper.

175
176

Funding

177

This study was funded by the Brazilian Biodiversity Fund - FUNBIO, the HUMANIZE

178

and Eurofins Foundations (FUNBIO - Fellowships Conserving the Future, granted to RAC, Nº

179

025/2022), the National Council for Scientific and Technological Development (CNPq)

180

(Doctoral fellowship for RAC, Nº 141873/2020-5 and productivity grant to PMM, Nº

181

304866/2020-2) and the Research Support Foundation of the State of Alagoas (FAPEAL)

182

(Granted to PMM, Nº APQ2022021000027).

183
184

Authors' contributions

185

RAC - Conceptualization; Investigation; Methodology; Writing - original draft and

186

final. ELGS - Writing - revision and editing. LFCN - Writing - revision and editing; Data

187

compilation; RRVS and ARC - Supervision; Writing - revision and editing. PMM -

188

Conceptualization; Methodology; Writing – final version and editing. All authors read and

189

approved the final version.

190
191

Acknowledgments

192

We extend our gratitude to the residents of the Retiro community for their support in

193

this research, particularly to Mr. José Antônio and Mrs. Maria das Dores, who warmly

194

welcomed us and facilitated our requests within the community. Our appreciation also goes to

114

195

Dr. Marcos Sobral, a Myrtaceae expert, who graciously identified the species of this family in

196

our study. Likewise, we express our thanks to the researcher from the Agronomic Institute of

197

Pernambuco, Maria Olivia de Oliveira Cano, for her receptiveness and efficiency in assisting

198

us during all visits throughout the COVID-19 pandemic.

115

Table 1 Plants that were part of the checklist interview, their citation percentages (general and by use), and occurrence areas.
Nº of
%
% Food % Wood
Popular name
Family
Scientific name
Areas1
Voucher
citations general
use
use
Jenipapo
Rubiaceae
Genipa americana L.
C
94826
114
99.1
57.4
99.1
Ingá*
Leg. Mim.
Inga spp.
A, B and C
112
97.4
67.8
97.4
Cambuí
Myrtaceae
Myrciaria floribunda (H. West ex Wild.) O. Berg
A and B
94056
103
89.6
41.7
89.6
Massaranduba
Sapotaceae
Manilkara salzmannii (A.DC.) H.J.Lam
A and B
93971
96
83.5
77.4
82.6
Araçá
Myrtaceae
Psidium guineense Sw.
C
94807
93
80.9
30.4
80.9
Cruiri
Melastomataceae Mouriri sp.
C
94822
75
65.2
51.3
64.3
Pau d'arco*
Bignoniaceae
Bignoniaceae spp.
A and B
66
57.4
57.4
0
Carrapatinho
Rutaceae
Esenbeckia grandiflora Mart.
A and B
93988
27
23.5
23.5
0
Banana de papagaio Clusiaceae
Kielmeyera rugosa Choisy
A and B
93982
26
22.6
22.6
0
Murici comum
Malpighiaceae
Byrsonima sericea DC.
A, B and C 94066
22
19.1
18.3
13.9
Murta roxa
Myrtaceae
Neomitranthes obtusa Sobral & Zambom
A and B
94767
18
15.7
15.7
10.4
Peroba
Bignoniaceae
Tabebuia elliptica (DC.) Sandwith
A and B
94765
18
15.7
15.7
0
Orelha d'onça
Polygonaceae
Coccoloba laevis Casar.
A
94000
18
15.7
15.7
7.8
Louro
Lauraceae
Ocotea notata (Nees & Mart.) Mez
A and B
94064
16
13.9
13.9
0.9
Sicupira
Leg. Caes.
Diptychandra aurantiaca Tul.
A
93999
14
12.2
12.2
0
Espinho branco
Rubiaceae
Machaonia acuminata Bonpl.
C
94813
12
10.4
10.4
0
Meiru
Annonaceae
Xylopia laevigata (Mart.) R.E.Fr.
A and B
94067
9
7.8
7.8
0
Murici de vaqueiro Malpighiaceae
Byrsonima bahiana W.R.Anderson
A
93989
9
7.8
6.1
7.8
Pirunga
Myrtaceae
Myrcia arenaria L.L.Santos et al.
A and B
93995
9
7.8
6.1
7.0
Murta branca
Myrtaceae
Eugenia punicifolia (Kunth) DC.
A and B
94068
8
7.0
7.0
4.3
Açoita égua
Myrtaceae
Myrcia loranthifolia (DC.) G.Burton & E.Lucas
A, B and C 93994
7
6.1
6.1
3.5
Camarão
Flacourtiaceae
Casearia Sylvestris Sw.
A and C
94821
7
6.1
6.1
0

116

Piranha
Nyctaginaceae
Guapira opposita (Vell.) Reitz
A and B
93993
Rompe gibão
Euphorbiaceae
Phyllanthus sp.
C
Sete casco
Euphorbiaceae
Pera glabrata (Schott) Baill.
A and B
93976
Murta amarela
Myrtaceae
Neomitranthes sp.
A, B and C 93991
*Ethnospecies
1
Occurrence areas of plant species - local denominations: (A) - Carrasco, (B) - Zé Marinho, (C) - Lalo (Patos).

5
4
3
1

4.3
3.5
2.6
0.9

0.9
0
0
0

4.3
3.5
2.6
0.9

117

Table 2 Species recognized for wood purposes and their respective uses in each category
Family
Cientific Name Popular name
Fuel
Construction
Technology
Annonaceae
Rafter, Fence, Window, Line, Handle for hoe, Handle for broom, Fishing hook,
Xylopia laevigata (Mart.) R.E.Fr. (Meiru) Charcoal, Firewood
Door
Handle for cóvu, Handle for covú
Bignoniaceae
Washstand alpendre, Horse stall, Scaffolding, Cabinet, Bench, Boat, Handle for
Rafter,
Fence,
House hoe, Handle for sickle, Handle for axe, Handle for
construction, Post, Line, Roof hammer, Handle for drag net, Handle for brush
Bignoniaceae spp. (Pau d'arco)
Charcoal, Firewood timber, Mourão, Door, Portal, cutter, Handle for broom, Chair, Canoe, Shelf,
Slat, Gate board, Corral board, Fish string, Wardrobe, Table, Oar, Boat rhombus,
Fill board (rammed earth house), Boat plank, Canoe board, Cart board, Stool,
Stick for rammed earth house
Handle for cóvu
Cabinet, Boat, Handle for digging stick, Handle
Brabo, Rafter, Fence, Post, Line, for hoe, Handle for small hoe, Handle for sickle,
Tabebuia elliptica (DC.) Sandwith (Peroba) Charcoal, Firewood Mourão, Door, Portal, Slat, Fill Handle for axe, Handle for hammer, Handle for
board (rammed earth house)
drag net, Handle for broom, Chair, Coffin, Bed,
Wooden spoon, Wood for sofa, Pestle hand, Table,
Furniture, Oar, Handle for cóvu, Handle for covú
Clusiaceae
Kielmeyera rugosa Choisy (Banana de
Bench, Handle for broom, Cork for fishing net,
Charcoal, Firewood Fence, Post, Line
papagaio)
Oar
Euphorbiaceae
Pera glabrata (Schott) Baill. (Sete casco)

Charcoal, Firewood Rafter, Fence, Post,

118

Phyllanthus sp. (Rompe gibão)

Charcoal, Firewood Fence

Flacourtiaceae
Casearia Sylvestris Sw. (Camarão)

Charcoal, Firewood Rafter, Fence

Bench, Handle for hoe, Handle for small hoe,
Chair, Coffee table, Handle for cóvu

Lauraceae
Ocotea notata (Nees & Mart.) Mez (Louro)

Boat, Handle for hoe, Handle for sickle, Handle for
Barrote, Rafter, Fence, Post, Line,
axe, Handle for broom, Chair, Canoe, Table, Oar,
Charcoal, Firewood Door, Portal, Slat, Fill board
Boat plank, Canoe board, Table board, Handle for
(rammed earth house)
cóvu

Leg. Caes.
Diptychandra aurantiaca Tul. (Sicupira)

Rafter, Fence, Corral, Post, Line, Handle for hoe, Handle for small hoe, Handle for
Charcoal, Firewood Mourão, Fill board (rammed earth sickle, Handle for axe, Chair, Boat cave, Ibone,
house)
Table, Handle for cóvu

Leg. Mim.
Inga spp. (Ingá)
Malpighiaceae
Byrsonima bahiana W.R.Anderson (Murici
de vaqueiro)
Byrsonima sericea DC. (Murici comum)
Melastomataceae

Horse stall, Rafter, Fence, House Handle for hoe, Handle for tool, Handle for sickle,
Charcoal, Firewood construction, Post, Line, Mourão, Handle for broom, Handle for cóvu, Handle for
Fill board (rammed earth house) covú
Fence, Stick for rammed earth
house
Door jamb, Brabo, Rafter, Fence, Handle for hoe, Handle for axe, Oar, Boat
Charcoal, Firewood
Window, Line, Door
rhombus, Handle for cóvu, Handle for covú
Charcoal, Firewood

119

Mouriri sp. (Cruiri)
Myrtaceae
Eugenia punicifolia (Kunth) DC. (Murta
branca)
Myrcia arenaria L.L.Santos et al. (Pirunga)

Fence, Pigsty, Corral, Post, Scaffolding, Bench, Handle for hoe, Handle for
Mourão, Fill board (rammed earth sickle, Handle for hammer, Handle for drag net,
Charcoal, Firewood
house), Stick for rammed earth Canoe cave, Pitchfork, Pestle, Oar, Boat rhombus,
house
Handle for cóvu, Handle for covú
Charcoal, Firewood Rafter, Fence

Handle for hoe, Handle for sickle, Handle for
broom, Handle for cóvu

Charcoal, Firewood Fence

Handle for cóvu, Handle for covú

Myrciaria floribunda (H. West ex Wild.) O.
Charcoal, Firewood Fence, Line, Mourão
Berg (Cambuí)
Myrcia loranthifolia (DC.) G.Burton &
E.Lucas (Açoita égua)

Charcoal, Firewood Rafter, Woodshed, Fence

Neomitranthes obtusa Sobral & Zambom
(Murta roxa)

Charcoal, Firewood

Rafter, Fence,
Mourão, Portal

Charcoal

Fence

Neomitranthes sp. (Murta amarela)
Psidium guineense Sw. (Araçá)

Post,

Handle for hoe, Handle for sickle, Handle for
shuttlecock, Handle for broom, Fishing hook,
Handle for cóvu, Handle for covú, Yard broom
Handle for hoe, Handle for broom, Handle for
cóvu
Bench, Handle for digging stick, Handle for hoe,
Handle for small hoe, Handle for sickle, Handle
Line,
for axe, Handle for brush cutter, Handle for broom,
Chair, Coffee table, Handle for cóvu, Handle for
covú

Charcoal, Firewood Fence, Corral

Nyctaginaceae
Guapira opposita (Vell.) Reitz (Piranha)
Polygonaceae

Charcoal, Firewood

Rafter, Fence, Line, Slat, Fill
board (rammed earth house)

Scaffolding, Handle for shuttlecock, Spinning top,
Handle for cóvu, Handle for covú

120

Coccoloba laevis Casar. (Orelha d'onça)

Charcoal, Firewood Rafter, Fence

Skewer for roasting corn

Rubiaceae

Genipa americana L. (Jenipapo)

Machaonia acuminata Bonpl. (Espinho
branco)
Rutaceae

Zabumba arch, Cabinet, Bench, Boat, Boat tiller,
Handle for digging stick, Handle for hoe, Handle
for strainer, Handle for sickle, Handle for axe,
Alpendre, Washstand alpendre,
Handle for hammer, Handle for drag net, Chair,
Rafter, Fence, Corral, Line,
Charcoal, Firewood
Bed, Canoe, Wooden spoon, Pestleant prop, Shed,
Mourão, Door, Slat, Door board,
Cupboard, Mast, Table, Furniture, Chair leg,
Stick for rammed earth house
Camiseiro leg, Pestle, Oar, Boat rhombus, Boat
plank, Canoe board, Meat board, Board for rice
husking machine, Stool board, Stool
Charcoal, Firewood Fence

Barrier for caiçara, Fishing hook, Handle for cóvu

Needle for cast net, Handle for hoe, Handle for
Fence, Slat, Fill board (rammed
sickle, Handle for drag net, Handle for broom,
Esenbeckia grandiflora Mart. (Carrapatinho) Charcoal, Firewood earth house), Stick for rammed
Skewer for barbecue, Handle for cóvu, Handle for
earth house
covú
Sapotaceae

121

Manilkara salzmannii (A.DC.) H.J.Lam
(Massaranduba)

199

Scaffolding, Cabinet, Bench, Boat, Handle for
Barrote, Door jamb, Rafter,
digging stick, Cleaver, Handle for hoe, Handle for
Fence,
Pigsty,
House
small hoe, Handle for strainer, Handle for
construction,
Corral,
Post,
machete, Handle for sickle, Handle for axe,
Window, Line, Mourão, Door,
Charcoal, Firewood
Handle for hammer, Handle for broom, Chair,
Slat, Corral board, Fill board
Bed, Chest of drawers, Canoe, Boat cave, Shelf,
(rammed earth house), Stable
Table, Furniture, Oar, Boat rhombus, Boat plank,
board, Door board, Stick for
Canoe board, Furniture board, Stool, Handle for
rammed earth house
cóvu, Handle for covú

122

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for the selection of wild food plants for consumption and sale in Alagoas, Brazil.
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25.

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migratórias na Área de Proteção Ambiental de Piaçabuçu, Alagoas, Brasil. Rev Bras Zool.
2006;23(3):865–9.

26.

Burnham KP, Anderson DR. Multimodel inference: Understanding AIC and BIC in model
selection. Sociol Methods Res. 2004;33(2):261–304.

27.

Evans, M. I. (1993). “Conservation by commercialization” in Tropical forests, People and
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M. Hladik, A. Hladik, O. F. Linares, H. Pagezy, A. Semple, and M. Hadley ( 815–822.
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fuelwood purposes: A case study in the Brazilian semiarid region. J Arid Environ [Internet].
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5481885627b6ab166cd

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Marquez-Reynoso MI, Ramírez-Marcial N, Cortina-Villar S, Ochoa-Gaona S. Purpose,
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Intensification of açaí palm management largely impoverishes tree assemblages in the
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35.

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36.

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39.

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40.

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agricultural landscapes and their conservation role in an area of caatinga (dryland vegetation)
in northeast Brazil. Environ Dev Sustain. 2009;11(5):1005–29.

41.

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126

5. Conclusões
O primeiro capítulo desta tese mostrou que uma gama de espécies vegetais
que tem potencial alimentício, muitas vezes inexplorado, pode estar ameaçada pelos
seus usos madeireiros, correndo risco de terem sua disponibilidade amplamente
reduzida antes mesmo de ter o seu potencial explorado. No entanto, o segundo
capítulo mostrou que espécies de alta importância alimentícia (espécies-chave) são
poupadas dos usos madeireiros pelos povos locais. Desse modo, a valorização do
potencial alimentício pode contribuir para a conservação de espécies vegetais, aliada
a estratégias de manejo mais holísticas, que protejam as espécies sem potencial
alimentício e evitem a simplificação de comunidades vegetais a partir de um manejo
para fins alimentícios.
Por isso, as espécies com sobreposição entre os usos alimentícios e
madeireiros são prioritárias à conservação a fim de garantir a existência contínua de
recursos potenciais para a segurança alimentar e nutricional, especialmente aquelas
versáteis para os usos madeireiros e com alto potencial nutricional. E, considerando
que o efeito protetivo do uso alimentício sobre os usos madeireiros está direcionado
para as espécies-chave, as espécies com baixa ou média importância alimentícia
devem ser priorizadas em estratégias de popularização para elevar sua importância.
No entanto, é crucial realizar mais pesquisas ecológicas para entender o estado de
conservação das espécies-chave em seus habitats naturais. Isso inclui investigar se
essas espécies estão sob maior pressão devido ao uso como alimento e se seu
manejo está contribuindo para a redução da diversidade vegetal. Além disso, espécies
lenhosas não comestíveis, que não se beneficiam da proteção associada ao uso
alimentício, também precisam de atenção por meio de estratégias de conservação
biocultural.

127
6. Apêndices
Artigo 1: Supplementary material

Add File 1. PRISMA Protocol Checklist – Ethnobiology review
Section and Topic

Item
#

Checklist item

Location where
item is reported

TITLE
Title

1

Identify the report as a systematic review.

1

ABSTRACT
Abstract

2

See the PRISMA 2020 for Abstracts checklist.

2

INTRODUCTION
Rationale

3

Describe the rationale for the review in the context of existing knowledge.

2-4

Objectives
METHODS
Eligibility criteria

4

Provide an explicit statement of the objective(s) or question(s) the review addresses.

5

5

Specify the inclusion and exclusion criteria for the review and how studies were grouped for the syntheses.

5

Information sources

6

Specify all databases, registers, websites, organisations, reference lists and other sources searched or consulted to
identify studies. Specify the date when each source was last searched or consulted.

6-7

Search strategy

7

Present the full search strategies for all databases, registers and websites, including any filters and limits used.

7, 8 and
additional file 2

Selection process

8

Specify the methods used to decide whether a study met the inclusion criteria of the review, including how many
reviewers screened each record and each report retrieved, whether they worked independently, and if applicable,
details of automation tools used in the process.

8

Data collection process

9

Specify the methods used to collect data from reports, including how many reviewers collected data from each
report, whether they worked independently, any processes for obtaining or confirming data from study
investigators, and if applicable, details of automation tools used in the process.

8-9

128

Section and Topic
Data items

Item
#
10a

Checklist item
List and define all outcomes for which data were sought. Specify whether all results that were compatible with
each outcome domain in each study were sought (e.g. for all measures, time points, analyses), and if not, the
methods used to decide which results to collect.

Location where
item is reported
9-10

10b

List and define all other variables for which data were sought (e.g. participant and intervention characteristics,
funding sources). Describe any assumptions made about any missing or unclear information.

8-9

Study risk of bias
assessment

11

Specify the methods used to assess risk of bias in the included studies, including details of the tool(s) used, how
many reviewers assessed each study and whether they worked independently, and if applicable, details of
automation tools used in the process.

9

Effect measures

12

Specify for each outcome the effect measure(s) (e.g. risk ratio, mean difference) used in the synthesis or
presentation of results.

NA

Synthesis methods

13a

Describe the processes used to decide which studies were eligible for each synthesis (e.g. tabulating the study
intervention characteristics and comparing against the planned groups for each synthesis (item #5)).

8

13b

Describe any methods required to prepare the data for presentation or synthesis, such as handling of missing
summary statistics, or data conversions.

8-9

13c

Describe any methods used to tabulate or visually display results of individual studies and syntheses.

9-10

13d

Describe any methods used to synthesize results and provide a rationale for the choice(s). If meta-analysis was
performed, describe the model(s), method(s) to identify the presence and extent of statistical heterogeneity, and
software package(s) used.

9-10

13e

Describe any methods used to explore possible causes of heterogeneity among study results (e.g. subgroup
analysis, meta-regression).

NA

13f

Describe any sensitivity analyses conducted to assess robustness of the synthesized results.

NA

Reporting bias
assessment

14

Describe any methods used to assess risk of bias due to missing results in a synthesis (arising from reporting
biases).

NA

Certainty assessment
RESULTS
Study selection

15

Describe any methods used to assess certainty (or confidence) in the body of evidence for an outcome.

NA

16a

Describe the results of the search and selection process, from the number of records identified in the search to the
number of studies included in the review, ideally using a flow diagram.

14 and Fig.1

16b

Cite studies that might appear to meet the inclusion criteria, but which were excluded, and explain why they were
excluded.

14

129

Study characteristics

Item
#
17

Cite each included study and present its characteristics.

Location where
item is reported
14-16

Risk of bias in studies

18

Present assessments of risk of bias for each included study.

NA

Results of individual
studies

19

For all outcomes, present, for each study: (a) summary statistics for each group (where appropriate) and (b) an
effect estimate and its precision (e.g. confidence/credible interval), ideally using structured tables or plots.

NA

Results of syntheses

20a

For each synthesis, briefly summarise the characteristics and risk of bias among contributing studies.

16-18

20b

Present results of all statistical syntheses conducted. If meta-analysis was done, present for each the summary
estimate and its precision (e.g. confidence/credible interval) and measures of statistical heterogeneity. If
comparing groups, describe the direction of the effect.

NA

20c

Present results of all investigations of possible causes of heterogeneity among study results.

NA

20d

Present results of all sensitivity analyses conducted to assess the robustness of the synthesized results.

NA

Reporting biases

21

Present assessments of risk of bias due to missing results (arising from reporting biases) for each synthesis
assessed.

NA

Certainty of evidence

22

Present assessments of certainty (or confidence) in the body of evidence for each outcome assessed.

NA

DISCUSSION
Discussion

23a

Provide a general interpretation of the results in the context of other evidence.

26-30

23b

Discuss any limitations of the evidence included in the review.

26 and 30

23c

Discuss any limitations of the review processes used.

30

23d

Discuss implications of the results for practice, policy, and future research.

30

Provide registration information for the review, including register name and registration number, or state that the
review was not registered.

NA

24b

Indicate where the review protocol can be accessed, or state that a protocol was not prepared.

NA

24c

Describe and explain any amendments to information provided at registration or in the protocol.

NA

Support

25

Describe sources of financial or non-financial support for the review, and the role of the funders or sponsors in the
review.

31

Competing interests

26

Declare any competing interests of review authors.

32

Section and Topic

OTHER INFORMATION
Registration and
24a
protocol

Checklist item

130

Section and Topic
Availability of data,
code and other materials

Item
#
27

Checklist item
Report which of the following are publicly available and where they can be found: template data collection forms;
data extracted from included studies; data used for all analyses; analytic code; any other materials used in the
review.

Location where
item is reported
32

From: Page MJ, McKenzie JE, Bossuyt PM, Boutron I, Hoffmann TC, Mulrow CD, et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ
2021;372:n71. doi: 10.1136/bmj.n71
7. For more information, visit: http://www.prisma-statement.org/

131

Add File 2. Search strategies – Ethnobiology review
Search 1 (B1)
Key words - SCOPUS, WEB OF SCIENCE AND
SCIELO

Search 2 (B2)
Key word in Portuguese - SCIELO

("plantas comestíveis" OR "plantas alimentícias" OR
("edible plants" OR "food plants" OR "wild food plants"
"plantas alimentícias silvestres" OR "plantas alimentares"
OR "non-timber forest products" OR "food uses") AND
OR "produtos florestais não madeireiros" OR "usos
ethnob* AND Brazil
alimentícios") AND etnob*

Key words - SCOPUS, WEB
OF SCIENCE AND SCIELO
("local knowledge" OR
"traditional knowledge")
AND ethnob* AND Brazil

("wood plants" OR "wood uses" OR "wood products"
("plantas madeireiras" OR "usos madeireiros" OR
OR "wood resources" OR "wood forest products" OR
"produtos madeireiros" OR "recursos madeireiros" OR
"timber forest products" OR "timber resources") AND
"produtos florestais madeireiros") AND etnob*
ethnob* AND Brazil
"useful plants" AND ethnob* AND Brazil

“plantas úteis” AND etnob*

("woody plants" OR "woody species" OR "woody forest ("plantas lenhosas" OR "espécies lenhosas" OR "recursos
resources") AND ethnob* AND Brazil
florestais lenhosos") AND etnob*
"multipurpose plants" AND ethnob* AND Brazil

("plantas multiuso" OR "plantas de múltiplos usos") AND
etnob*

"wild food plants" AND ethnob* AND Brazil

"plantas alimentícias silvestres" AND etnob*

"food plants" AND ethnob* AND Brazil

"plantas alimentícias" AND etnob*

Key words in Portuguese SCIELO
("conhecimento local" OR
"conhecimento tradicional")
AND etnob*

132

"edible plants" AND ethnob* AND Brazil

"plantas alimentares" AND etnob*

"non-timber forest products" AND ethnob* AND Brazil "plantas comestíveis" AND etnob*
"food uses" AND ethnob* AND Brazil

"produtos florestais não madeireiros" AND etnob*

("wood plants" OR "wood species") AND ethnob* ("plantas madeireiras" OR "espécies madeireiras") AND
AND Brazil
etnob*
"wood uses" AND ethnob* AND Brazil

"usos madeireiros" AND etnob*

"wood products" AND ethnob* AND Brazil

"recursos madeireiros" AND etnob*

("wood resources" OR "timber resources") AND "produtos madeireiros" AND etnob*
ethnob* AND Brazil
"wood forest products" AND ethnob* AND Brazil

"produtos florestais madeireiros" AND etnob*

"timber forest products" AND ethnob* AND Brazil lenha AND etnob*
NOT "non-timber"
("fuelwood species" OR "fuelwood plants" OR "uso combustível" AND etnob*
"firewood use" OR firewood OR fuelwood) AND
ethnob* AND Brazil
("fuel use" OR "woodfuel") AND ethnob* AND Brazil ("carvão vegetal" OR carvão) AND etnob*
(charcoal OR coal) AND ethnob* AND Brazil

"categoria tecnologia" AND etnob*

"construction category" AND ethnob* AND Brazil

"categoria construção" AND etnob*

133

"fuelwood category" AND ethnob* AND Brazil

"categoria combustível" AND etnob*

"technology category" AND ethnob* AND Brazil

cercas AND etnob*

fence AND ethnob* AND Brazil

134
Add File 3. PRISMA Protocol Checklist – Nutritional review
Section and
Topic

Ite
m#

Checklist item

Location
where
item
is reported

TITLE
Title
ABSTRACT

1

Identify the report as a systematic review.

1

Abstract
INTRODUCTION

2

See the PRISMA 2020 for Abstracts checklist.

2

Rationale

3

Describe the rationale for the review in the context of existing knowledge.

2-4

Objectives
METHODS

4

Provide an explicit statement of the objective(s) or question(s) the review addresses.

5

Eligibility criteria

5

Specify the inclusion and exclusion criteria for the review and how studies were grouped for the syntheses.

10

Information
sources

6

Specify all databases, registers, websites, organisations, reference lists and other sources searched or consulted to identify studies. Specify the
date when each source was last searched or consulted.

12

Search strategy

7

Present the full search strategies for all databases, registers and websites, including any filters and limits used.

Fig 2 and
page 12

Selection process

8

Specify the methods used to decide whether a study met the inclusion criteria of the review, including how many reviewers screened each
record and each report retrieved, whether they worked independently, and if applicable, details of automation tools used in the process.

12-13

Data collection
process

9

Specify the methods used to collect data from reports, including how many reviewers collected data from each report, whether they worked
independently, any processes for obtaining or confirming data from study investigators, and if applicable, details of automation tools used in the
process.

13-14

10a

List and define all outcomes for which data were sought. Specify whether all results that were compatible with each outcome domain in each study 18-19
were sought (e.g. for all measures, time points, analyses), and if not, the methods used to decide which results to collect.

10b

List and define all other variables for which data were sought (e.g. participant and intervention characteristics, funding sources). Describe any
assumptions made about any missing or unclear information.

18-19

Study risk of bias
assessment

11

Specify the methods used to assess risk of bias in the included studies, including details of the tool(s) used, how many reviewers assessed each
study and whether they worked independently, and if applicable, details of automation tools used in the process.

NA

Effect measures

12

Specify for each outcome the effect measure(s) (e.g. risk ratio, mean difference) used in the synthesis or presentation of results.

NA

Data items

135
13a

Describe the processes used to decide which studies were eligible for each synthesis (e.g. tabulating the study intervention characteristics and
comparing against the planned groups for each synthesis (item #5)).

18-19

13b

Describe any methods required to prepare the data for presentation or synthesis, such as handling of missing summary statistics, or data
conversions.

12

13c

Describe any methods used to tabulate or visually display results of individual studies and syntheses.

12

13d

Describe any methods used to synthesize results and provide a rationale for the choice(s). If meta-analysis was performed, describe the
model(s), method(s) to identify the presence and extent of statistical heterogeneity, and software package(s) used.

NA

13e
13f

Describe any methods used to explore possible causes of heterogeneity among study results (e.g. subgroup analysis, meta-regression).
Describe any sensitivity analyses conducted to assess robustness of the synthesized results.

NA
NA

Reporting bias
assessment

14

Describe any methods used to assess risk of bias due to missing results in a synthesis (arising from reporting biases).

NA

Certainty
assessment
RESULTS

15

Describe any methods used to assess certainty (or confidence) in the body of evidence for an outcome.

NA

16a

Describe the results of the search and selection process, from the number of records identified in the search to the number of studies included in
the review, ideally using a flow diagram.

16b

Cite studies that might appear to meet the inclusion criteria, but which were excluded, and explain why they were excluded.

18
and
Fig.3
18

Study
characteristics

17

Cite each included study and present its characteristics.

18-19

Risk of bias in
studies

18

Present assessments of risk of bias for each included study.

NA

Results of
individual studies

19

For all outcomes, present, for each study: (a) summary statistics for each group (where appropriate) and (b) an effect estimate and its precision
(e.g. confidence/credible interval), ideally using structured tables or plots.

NA

Results of
syntheses

20a

For each synthesis, briefly summarise the characteristics and risk of bias among contributing studies.

20-25

20b

Present results of all statistical syntheses conducted. If meta-analysis was done, present for each the summary estimate and its precision (e.g.
confidence/credible interval) and measures of statistical heterogeneity. If comparing groups, describe the direction of the effect.

NA

20c

Present results of all investigations of possible causes of heterogeneity among study results.

NA

20d

Present results of all sensitivity analyses conducted to assess the robustness of the synthesized results.

NA

21

Present assessments of risk of bias due to missing results (arising from reporting biases) for each synthesis assessed.

NA

Synthesis
methods

Study selection

Reporting biases

136
Certainty of
evidence
DISCUSSION

Discussion

22

23a
23b
23c

Present assessments of certainty (or confidence) in the body of evidence for each outcome assessed.

NA

Provide a general interpretation of the results in the context of other evidence.
Discuss any limitations of the evidence included in the review.
Discuss any limitations of the review processes used.

26-30
26 and 30
30
30

23d
Discuss implications of the results for practice, policy, and future research.
OTHER INFORMATION
Registration and
protocol

24a

Provide registration information for the review, including register name and registration number, or state that the review was not registered.

NA

24b

Indicate where the review protocol can be accessed, or state that a protocol was not prepared.

NA

Support

24c
25

Describe and explain any amendments to information provided at registration or in the protocol.
Describe sources of financial or non-financial support for the review, and the role of the funders or sponsors in the review.

NA
31

Competing
interests

26

Declare any competing interests of review authors.

32

Availability of
data, code and
other materials

27

Report which of the following are publicly available and where they can be found: template data collection forms; data extracted from included
studies; data used for all analyses; analytic code; any other materials used in the review.

32

From: Page MJ, McKenzie JE, Bossuyt PM, Boutron I, Hoffmann TC, Mulrow CD, et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ
2021;372:n71. doi: 10.1136/bmj.n71

For more information, visit: http://www.prisma-statement.org/

137
Add File 4. Search strategies – Nutritional review
Scopus, Web of science and Scielo
(English)
"Amburana cearensis" AND nutritional

Scielo (Portuguese)

SiBBr

"Amburana cearensis" AND nutricional Amburana cearensis
"Anacardium occidentale" AND
"Anacardium occidentale" AND nutritional
Anacardium occidentale
nutricional
"Anadenanthera colubrina" AND
"Anadenanthera colubrina" AND nutritional
Anadenanthera colubrina
nutricional
("Astronium urundeuva" OR
"Myracrodruon urundeuva") AND
nutritional
"Bauhinia cheilantha" AND nutritional
"Brosimum guianense" AND nutritional
"Byrsonima sericea" AND nutritional
"Cecropia pachystachya AND nutritional
"Combretum leprosum" AND nutritional
"Commiphora leptophloeos" AND
nutritional
"Copaifera langsdorffii" AND nutritional
("Croton heliotropiifolius" OR "Croton
rhamnifolius") AND nutritional
("Cynophalla flexuosa" OR "Capparis
flexuosa") AND nutritional
("Didymopanax morototoni" OR
"Schefflera morototoni") AND nutritional

("Astronium urundeuva" OR
"Myracrodruon urundeuva") AND
Astronium urundeuva
nutricional
"Bauhinia cheilantha" AND nutricional Bauhinia cheilantha
"Brosimum guianense" AND nutricional Brosimum guianense
"Byrsonima sericea" AND nutricional
Byrsonima sericea
"Cecropia pachystachya AND
Cecropia pachystachya
nutricional
"Combretum leprosum" AND
Combretum leprosum
nutricional
"Commiphora leptophloeos" AND
Commiphora leptophloeos
nutricional
"Copaifera langsdorffii" AND
Copaifera langsdorffii
nutricional
("Croton heliotropiifolius" OR "Croton
Croton heliotropiifolius
rhamnifolius") AND nutricional
("Cynophalla flexuosa" OR "Capparis
Cynophalla flexuosa
flexuosa") AND nutricional
("Didymopanax morototoni" OR
"Schefflera morototoni") AND
Didymopanax morototoni
nutricional

Myracrodruon
urundeuva
Croton rhamnifolius
Capparis flexuosa
Schefflera morototoni

138
("Eschweilera ovata" OR "Eschweilera
luschnathii") AND nutritional
("Eugenia pyriformis" OR "Eugenia
uvalha") AND nutritional
"Eugenia uniflora" AND nutritional
"Genipa americana" AND nutritional

("Eschweilera ovata" OR "Eschweilera
luschnathii") AND nutricional
("Eugenia pyriformis" OR "Eugenia
uvalha") AND nutricional
"Eugenia uniflora" AND nutricional
"Genipa americana" AND nutricional
("Handroanthus impetiginosus" OR
("Handroanthus impetiginosus" OR
"Tabebuia impetiginosa") AND
"Tabebuia impetiginosa") AND nutritional
nutricional
("Handroanthus serratifolius" OR
("Handroanthus serratifolius" OR "Tabebuia
"Tabebuia serratifolia") AND
serratifolia") AND nutritional
nutricional
"Inga thibaudiana" AND nutritional
"Inga thibaudiana" AND nutricional
"Libidibia ferrea" AND nutritional
"Libidibia ferrea" AND nutricional
"Manihot dichotoma" AND nutritional
"Manihot dichotoma" AND nutricional
("Monteverdia obtusifolia" OR
("Monteverdia obtusifolia" OR "Maytenus
"Maytenus obtusifolia") AND
obtusifolia") AND nutritional
nutricional
("Monteverdia rigida" OR "Maytenus
("Monteverdia rigida" OR "Maytenus
rigida") AND nutritional
rigida") AND nutricional
"Myrcia splendens" AND nutritional
"Myrcia splendens" AND nutricional
("Neocalyptrocalyx longifolium" OR
("Neocalyptrocalyx longifolium" OR
"Capparis jacobinae") AND nutritional
"Capparis jacobinae") AND nutricional
"Ocotea glomerata" AND nutritional
"Ocotea glomerata" AND nutricional
"Ocotea odorifera" AND nutritional
"Ocotea odorifera" AND nutricional
"Pilosocereus pachycladus" AND
"Pilosocereus pachycladus" AND
nutritional
nutricional
"Protium heptaphyllum" AND
"Protium heptaphyllum" AND nutritional
nutricional
"Psidium guineense" AND nutritional
"Psidium guineense" AND nutricional

Eschweilera ovata

Eschweilera luschnathii

Eugenia pyriformis

Eugenia uvalha

Eugenia uniflora
Genipa americana

-

Handroanthus
impetiginosus

Tabebuia impetiginosa

Handroanthus serratifolius

Tabebuia serratifolia

Inga thibaudiana
Libidibia ferrea
Manihot dichotoma
Monteverdia obtusifolia

Maytenus obtusifolia

Monteverdia rigida

Maytenus rigida

Myrcia splendens
Neocalyptrocalyx
longifolium
Ocotea glomerata
Ocotea odorifera

-

Pilosocereus pachycladus
Protium heptaphyllum
Psidium guineense

Capparis jacobinae
-

139
("Sarcomphalus joazeiro" OR "Ziziphus
joazeiro") AND nutritional

("Sarcomphalus joazeiro" OR "Ziziphus
joazeiro") AND nutricional
"Schinus terebinthifolia" AND
"Schinus terebinthifolia" AND nutritional
nutricional
"Sideroxylon obtusifolium" AND
"Sideroxylon obtusifolium" AND
nutritional
nutricional
"Simarouba amara" AND nutritional
"Simarouba amara" AND nutricional
"Spondias mombin" AND nutritional
"Spondias mombin" AND nutricional
"Spondias tuberosa" AND nutritional
"Spondias tuberosa" AND nutricional
"Talisia esculenta" AND nutritional
"Talisia esculenta" AND nutricional
"Tapirira guianensis" AND nutritional
"Tapirira guianensis" AND nutricional
"Thyrsodium spruceanum" AND
"Thyrsodium spruceanum" AND nutritional
nutricional
"Ximenia americana" AND nutritional
"Ximenia americana" AND nutricional

Sarcomphalus joazeiro
Schinus terebinthifolia
Sideroxylon obtusifolium
Simarouba amara
Spondias mombin
Spondias tuberosa
Talisia esculenta
Tapirira guianensis
Thyrsodium spruceanum
Ximenia americana

Ziziphus joazeiro
-

140

Artigo 2: Supplementary material

Appendix 1. Multicollinearity diagnosis and model average for the widespread protection model and the
key species-based protection model
Overall Multicollinearity Diagnostics for the widespread protection model
MC Results detection
Determinant |X'X|:
0.8793
0
Farrar Chi-Square:
82.5421
1
Red Indicator
0.1465
0
Sum of Lambda Inverse:
4.2647
0
Theil's Method
0.0200
0
Condition Number
13.5726
0
Overall Multicollinearity Diagnostics for the key species-based protection model
MC Results detection
0.8407
0
Determinant |X'X|:
Farrar Chi-Square:
111.3357
1
Red Indicator
0.1645
0
Sum of Lambda Inverse:
4.3748
0
Theil's Method
0.0694
0
Condition Number
11.5739
0
1 -->
Collinearity is detected by the test
0 -->
Collinearity is not detected by the test
Model average for the widespread protection model
Component models
Df
LogLik
AiCc
Delta
Weight
1234
9-684.03
1389.34
0.00
0.33
123
8-685.25
1386.72
0.38
0.27
12
7-686.31
1386.79
0.45
0.26
124
8-685.89
1388.00
1.66
0.14
Term codes
1 – Availability
2 - Maximum_quality
3 - Commercial_food_use
4 - Domestic_food_use
Model-averaged coefficients:
(Full average)
Estimate
Variables
Error
Z
P
Std.
<2e1|2
4.29845
0.61263
7.016
16***
<2e2|3
5.48143
0.63207
8.672
16***

141

3|4

6.45156

0.65265

9.885

4|5

6.69560

0.65840

10.169

0.32415
0.07260
0.62826
0.11316
-0.12221
0.13921
0.05383
0.08789
(Conditional average)

4.465
5.552
0.878
0.648

1|2

4.29845

0.61263

7.016

2|3

5.48143

0.63207

8.672

3|4

6.45156

0.65265

9.885

4|5

6.69560

0.65840

10.169

Availability
Maximum_quality

0.32415
0.62826

Availability
Maximum_quality
Commercial_food_use
Domestic_food_use

0.07260
4.465
0.11316
5.552
Commercial_food_use
-0.20465
1.640
0.20465
Domestic_food_use
0.11476
0.08789
1.306
Model average for the key species-based protection model
Component models
Df
LogLik
AiCc
Delta
234
8-678.03
1372.29
0.00
1234
9-677.25
1372.78
0.49
Term codes
1Commercial_key_species_use
2 - Availability
3 - Maximum_quality
4Domestic_key_species_use
Model-averaged coefficients:
(Full average)
Estimate
Variables
Error
Z
Std.
1|2

4.29086

0.60813

7.056

2|3

5.51657

0.62948

8.764

3|4

6.50263

0.65073

9.993

4|5

6.74713

0.65643

10.279

Availability

0.35829

0.07267

4.930

<2e16***
<2e16***
8e-06***
3e-08***
0.380
0.517
<2e16***
<2e16***
<2e16***
<2e16***
8e-06***
3e-08***
0.101
0.192

Weight
0.56
0.44

P
< 2e-16
***
< 2e-16
***
< 2e-16
***
< 2e-16
***
8e-07
***

142
Maximum_quality

0.62038

0.11237

5.521

1.03517

0.30352

3.410

-0.26085
0.43358
(Conditional average)

0.602

1|2

4.29086

0.60813

7.056

2|3

5.51657

0.62948

8.764

3|4

6.50263

0.65073

9.993

4|5

6.74713

0.65643

10.279

Availability

0.35829

0.07267

4.930

0.62038

0.11237

5.521

-1.03517

0.30352

3.410

0.47973

1.239

Domestic_key_species_use
Commercial_key_species_use

Maximum_quality
Domestic_key_species_use

Commercial_key_species_use
-0.59433
Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1

< 2e-16
***
0.000648
***
0.547435
< 2e-16
***
< 2e-16
***
< 2e-16
***
< 2e-16
***
8.2e-07
***
3.0e-08
***
0.000648
***
0.215388