UNIVERSIDADE ESTADUAL PAULISTA 
“JÚLIO DE MESQUITA FILHO” 

INSTITUTO DE BIOCIÊNCIAS – RIO CLARO unesp 
 

 

 

 

PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS 
(BIOLOGIA VEGETAL) 

 

 

 

 

 

 

 

 

 

INTERAÇÃO LIANA-ÁRVORE EM VEGETAÇÕES COM PADRÕES SAZONAIS 

CONTRASTANTES 

 
 

 

 

 

Betânia da Cunha Vargas 

 
 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Tese apresentada ao Instituto de 
Biociências do Câmpus de Rio 
Claro, Universidade Estadual 
Paulista, como parte dos requisitos 
para obtenção do título de doutor 
em Ciências Biológicas (Biologia 
Vegetal).  



Betânia da Cunha Vargas 

 

 

 

 

 

 

 

 

 

INTERAÇÃO LIANA-ÁRVORE EM VEGETAÇÕES COM PADRÕES 

SAZONAIS CONTRASTANTES 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Rio Claro 

2018 

Tese apresentada ao Instituto de Biociências do 

Câmpus de Rio Claro, Universidade Estadual 

Paulista, como parte dos requisitos para 

obtenção do título de doutor em Ciências 

Biológicas (Biologia Vegetal).  

 

Orientador (a): Profª. Drª. Leonor Patrícia 

Cerdeira Morellato  

 

Co-orientador (a): Drª. Maria Tereza Gromboni-

Guaratini 



V297i
Vargas, Betânia da Cunha
    Interação liana-árvores em vegetações com padrões
sazonais contrastantes / Betânia da Cunha Vargas. -- Rio
Claro, 2018
    133 f. : tabs., mapas

    Tese (doutorado) - Universidade Estadual Paulista
(Unesp), Instituto de Biociências, Rio Claro
    Orientadora: Leonor Patrícia Cerdeira Morellato
    Coorientadora: Maria Tereza Gromboni-Guaratini

    1. Revisão bibliográfica. 2. Biomassa e estoque de
carbono. 3. Atributos funcionais foliares. I. Título.

Sistema de geração automática de fichas catalográficas da Unesp. Biblioteca do
Instituto de Biociências, Rio Claro. Dados fornecidos pelo autor(a).

Essa ficha não pode ser modificada.





 

 

AGRADECIMENTOS 

 

O presente trabalho foi realizado com apoio da Coordenação de Aperfeiçoamento de 

Pessoal de Nível Superior - Brasil (CAPES) - Código de Financiamento 001.  

 

À Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) pelo apoio 

financeiro para o desenvolvimento do projeto (projetos #2013/50155-0 FAPESP-

Microsoft Research, #2010/51307-0 FAPESP-VALE-FAPEMIG e #2009/54208-6 

EMU).  

 

Aos diretores e funcionários do Parque Estadual do Vassununga (Gleba Pé de Gigante), 

Fundação Serra do Japi pelo apoio e suporte para desenvolvimento do trabalho. 

Principalmente ao Sr. Lauro (Fundação Serra do Japi) que nos ajudou com equipamentos 

de campo e uma boa conversa após termos das atividades de coleta.  Incluo também a 

diretora e os funcionários, em especial ao André, do Parque Estadual Porto Ferreira que 

disponibilizou vários dias de seu trabalho para nos ajudar a encontrar uma área adequada 

para desenvolvermos o trabalho. Infelizmente não foi possível, mas o funcionário foi um 

exemplo de dedicação e empolgação com a pesquisa científica.  

 

À Universidade Estadual Paulista Júlio Mesquita Filho (UNESP - Campus Rio Claro, 

SP), à Pró-Reitoria de Pós-graduação da Universidade Estadual e ao Programa de Pós-

graduação em Ciências Biológicas (Biologia Vegetal). Em especial ao coordenador 

Douglas Silva Domingues que sempre esteve disposto a solucionar as eventuais dúvidas 

e problemas que sugiram nos últimos anos. E a ex-coordenadora Alessandra Ike Coan 

que também dispôs muito de seu tempo para que tivesse um bom período enquanto aluna 

deste programa de pós-graduação. Agradeço ainda as funcionárias da seção técnica de 

pós-graduação Ivana Terezinha Brandt e Rosangela pelo bom atendimento prestado 

durante esse período de doutorado. A todos muito obrigado!! 

 

Agradeço à minha orientadora Prof.ª Dr.ª Patrícia Morellato por ter aceitado me orientar 

sem conhecer a mim e ao meu trabalho. Pelas discussões e delimitações de cada parte da 

tese. E por incentivar à tantos alunos a tornarem bons pesquisadores. À Maria Tereza, 

minha co-orientadora, que também aceitou a orientação no escuro, que ajudou em 

exatamente tudo durante a tese e no doutorado. Desde as perguntas pecaminosas que me 



 

 

faziam sair da zona de conforto, as redes de contatos e colaboração que me inseriu me 

unindo a um grupo de pessoas que trabalham e sorriem juntas, por escutar minhas 

lamentações e sempre me dar uma luz de esperança. Com certeza vocês colaboraram 

muito para formação da profissional que sou hoje. Palavras, frases não são suficientes 

para demonstrar a gratidão de trabalhar e conviver com vocês.  

 

Agradeço aos professores e pesquisadores que fizeram parte da banca examinadora como 

membros titulares e suplentes pela valiosa colaboração e sugestões com o trabalho.  

 

Aos professores Gustavo Habermann e Marco Antônio Assis com os quais fiz estágio 

docência e pude aprender muito o conteúdo teórico. Porém, aprendi principalmente o 

modelo de professores, didática e interação com alunos que vocês demonstraram ao longo 

das aulas.  

 

Agradeço também ao Juliano van Mellis que me ajudou muito, muito, muito nas análises 

estatísticas, discussões ao longo do trabalho e colaboração em um dos capítulos. À Mirian 

Cilene Spasini Rinaldi (Instituto de Botânica de São Paulo) que me ensinou, orientou, 

desesperou, ficou até tarde comigo para fazermos as análises laboratoriais da tese. 

Mobilizou seus alunos (e os demais que estavam no laboratório) para me ajudar nas 

incontáveis digestões foliares. À aluna Solange Brandão (Instituto de Botânica de São 

Paulo) que parou seu experimento para me ajudar nas análises de pigmentos. Sol sem sua 

ajuda, paciência e as infindáveis conversas de quando abríamos à Ecologia às 06:30 da 

manhã e ali ficávamos trabalhando, discutindo e rindo muito durante todo o dia. Vocês 

tornaram realidade boa parte desta pesquisa e demonstraram a importância e como são 

saudáveis as parcerias, visando não só minimizar deficiências, mas também somar as 

qualidades de cada um. Serei eternamente grata aos novos amigos que fiz no Instituto de 

Botânica de São Paulo.  

 

Agradeço à Naiara David de Souza (técnica do departamento de Botânica) pela disposição 

a me ajudar durante o estágio docência e todo o período do doutorado. Sem falar nas 

conversas em assuntos em comum que nos levava muitas horas produtivas de 

conhecimento. Ao ex-técnico Rafael Consomagno que esteve em praticamente todos os 

campos comigo, fazendo de tudo o possível e impossível para realizarmos as atividades 

de campo e sempre com a maior boa vontade. Até mesmo para carregar o cilindro de 



 

 

nitrogênio líquido no cerrado e fazer peripécias para coletar algumas lianas. Rafa fez 

muita falta nos campos finais e agora para comemorar os bons frutos.  

 

Serei muito grata ao Renan Borgiani companheiro de trabalho e estudos das lianas. As 

nossas lamentações, conversas, discussões, conquistas, plantas para identificar, planilhas 

para arrumar, pedidos para serem feitos. Ufa foram muitos os tropeços, mas hoje vejo 

como mudamos e melhoramos como estudantes de um grupo de plantas que nos segurava 

em campo até descobrirmos muito de seus mistérios intrigantes. Aos amigos de 

laboratório Amanda, Annia, Bia, Bruna, Bruno, Desirée Diego, Gabi, Gustavo, Irene, 

Léo, Paty, Renan, Rosane, Swanni, Soizig, Valê, Van que tornaram uma família, unida 

que sofria nos campos juntos e sorria de muito mais para não deixa que os momentos 

difíceis superassem a jornada de cada um. Para vocês sempre haverá um grande espaço 

na minha vida.  

 

E o que dizer do quarteto familiar formado por Loris, Paty, Lê e por mim!!! Uma amizade 

que tenho à certeza que foi um reencontro de espíritos amigos. Vocês foram capazes de 

transformar dias, períodos difíceis em um longo aprendizado. Choramos, rimos, 

brincamos, malhamos, discutimos sobre cada coisinha mais mínima que podíamos e 

tenho certeza que uma raiz de cada uma de vocês está presente na minha vida.  

 

Devo um imenso agradecimento à cidade de Rio Claro, os quatro anos que tive aqui me 

fizeram crescer e amadurecer muito. Nossa relação não foi tão boa no começo, mas as 

pessoas que você me apresentou melhorou cada minutinho que estive por aí. Estar em Rio 

Claro não foi apenas uma transformação profissional, mas também um crescimento 

pessoal e espiritual. Muito obrigada Rio Claro.  

 

Agradeço a minha família Natália (mãe), Almir (pai), Lívio, (irmão), Káritas (irmã), Fran 

e Pedro (sobrinhos), Mariano (cunhado), Celina (avó), Nilton (tio) e Leide (tia). Essas 

pessoas depositaram em mim uma confiança que nem eu tinha, sorriam e vibravam com 

cada vitória, choravam e tinham esperança em cada tropeço. Fizeram o doutorado junto 

comigo, cada um do seu jeito, a quilômetros ou milhas de distância. Mas, nossa conexão 

é tão forte que eu tive sempre vocês onde quer que eu estivesse. Amo muito, muito, muito 

vocês e agradecer é muito pouco de tudo que fizeram por mim!! 

 



 

 

Resumo 
Lianas são plantas que alocam a biomassa primordialmente para a produção de folhas em 

detrimento da estrutura de suporte com alta densidade de madeira. Como consequência disso, 

lianas precisam de outros componentes florestais, geralmente árvores, para alcançar e permanecer 

no dossel. Uma vez estabelecida essa interação caracteriza-se como antagônica e duradoura, 

sendo que as lianas são os competidores mais eficientes acima e abaixo do dossel.  Isso ocorre, 

pois, as características foliares e radiculares garantem às lianas rápida absorção e transporte de 

água e nutrientes, principalmente em períodos de escassez hídrica. Este trabalho teve como 

objetivo avaliar a interação entre árvores e lianas ao longo de comunidades florestais neotropicais 

e foi dividido em três capítulos. No capítulo um, fizemos uma revisão sistemática sobre os estudos 

com lianas na região Neotropical (26ºN-26ºS). Para realizar esta revisão, pesquisamos em bancos 

de dados especializados, artigos que continham os termos liana*, “climb* plant*”, vine*, trepad*, 

sem limitação de país, e incluímos somente artigos científicos. Os artigos foram classificados em 

categorias pré-estabelecidas. Esta revisão encontrou 425 artigos com lianas para a região 

Neotropical, sendo a maior quantidade destes focados na interação entre árvores e lianas (255) e 

os demais exclusivos sobre lianas (170). A maioria dos artigos derivou de pesquisas no Brasil e 

Panamá, em florestas úmidas e floresta sazonal, e os temas de fitossociologia e ecofisiologia 

prevaleceram nos trabalhos. No capítulo dois, realizamos um levantamento fitossociológico em 

duas áreas de cerrado e uma floresta semidecidual no estado de São Paulo e incluímos áreas 

sazonais e não-sazonais disponibilizadas no banco de dados de Alwyn H. Gentry. Constatamos 

que o modelo proposto de biomassa e acúmulo de carbono foi bem representado e  que os fatores 

climáticos não exerceram efeito direto sobre a biomassa de lianas e nem no estoque de carbono. 

A biomassa de lianas tem efeito direto e significativo somente com a biomassa arbórea e 

abundância de lianas, e biomassa arbórea é o único fator com efeito direto e significativo para o 

estoque de carbono. Por fim, no terceiro capítulo avaliamos as variações sazonais das 

propriedades foliares e espectrais de árvores e lianas e a possibilidade de distinguir árvores de 

lianas no cerrado. Nossos resultados mostraram que por meio do espectro de reflectância não foi 

possível distinguir árvores e lianas. Enquanto que, a área foliar específica e a concentração 

nutricional nas folhas são fortes atributos para separar árvores e lianas. Portanto, consideramos a 

importância de incluir estudos em áreas sazonalmente extremas, pois nestes ambientes, assim 

como em cerrado, lianas podem ter padrões e respostas distintos do que previsto para florestas 

úmidas. Além disso, destacarmos a relevância do componente arbóreo em detrimento dos 

fatores climáticos nas florestas neotropicais. 

 

Palavra-chave: revisão, interação antagonista, lianas, estoque de carbono, características 

funcionais, Neotropical. 



 

 

Abstract 

Lianas are plants that invest biomass primarily to leaf production in detriment of woody 

components. The consequence of this is that lianas need other forest components, usually 

trees, to reach and remain in the canopy. Once established, this interaction is characterized 

as antagonistic and lasting, as lianas are more efficient in below and aboveground. This 

is due to the foliar and root characteristics of lianas that are known for fast absorption and 

transportation of water and nutrients, especially in drought period. We carried out an 

ecological research, that is divided in 3 chapters, on the antagonistic interaction between 

trees and lianas along neotropical forests. In chapter I, we performed a review on liana 

studies in the Neotropical region (26ºN-26ºS. To do this review we searched, in 

specialized databases, for articles that used the terms liana *, "climb * plant*", vine *, 

trepad *, without country limitation, but restricting ourselves only to scientific articles. In 

this process, we found 425 articles regarding lianas on the Neotropical region. Most of 

them were focused on the interaction between trees and lianas (255), although some 

talked exclusively about lianas (170). The majority of the articles came from studies in 

Brazil and Panama, in rainforests and seasonal forest, with dominance of phytosociology 

and ecophysiology themes. In Chapter 2, we performed a phytosociological survey in two 

cerrado and a semidecidual forest in São Paulo, where we followed a standard 

methodology suggested by Alwyn H. Gentry. We have also included seasonal and no 

seasonal areas available in this author's database. We found that the model well 

represented and the climatic factors had no direct effect on lianas biomass or carbon stock. 

Liana biomass has a direct and significant effect only with tree biomass and abundance 

of lianas. While tree biomass is the only factor with a direct and significant effect on the 

forest carbon stocks. In Chapter 3, we evaluated the seasonal variations of foliar and 

spectral properties of trees and lianas, and identified if it is possible to distinguish these 

groups in cerrado. As we concluded, the reflectance spectrum was not a good parameter 

to distinguish the lianas and trees. However, the specific leaf area and nutritional 

concentration in the leaves are strong attributes to separate trees and lianas. Therefore, 

we consider that studies with lianas need to be conducted in areas with extreme 

seasonality of precipitation. Because in these environments, like in cerrado, lianas may 

have differeny patterns from those predicted for rainforests. In addition, we highlight the 

relevance of the arboreal component to the detriment of the climatic factors in the 

Neotropical forests. 



 

 

Key-words: review, antagonistic interaction, lianas, carbon stocks, functional traits, 

Neotropical.  

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 



 

 

SUMÁRIO 

 

Introdução Geral......................................................................................................10-13 

Referências............................................................................................................14-18 

Capítulo I: Lianas research in the Neotropics: state of the art and future 

perspectives...............................................................................................................19-59 

Abstract..................................................................................................................21-22 

Introduction...........................................................................................................23-25 

Criteria for the inclusion of studies, vegetation classification and research 

topics......................................................................................................................25-26 

Review on liana research in the Neotropical region...............................................26-30 

Liana research over time and topics addressed among the research on 

lianas.................................................................................................................26-29 

Liana research in the Neotropics:  geographical distribution and vegetation 

types..................................................................................................................29-30 

Lianas: its antagonistic relation to trees and importance to biodiversity...............30-32 

Global change and the future of lianas...................................................................32-34 

Control, management and conservation of lianas..................................................34-37 

Future perspectives and conclusions......................................................................37-38 

References.............................................................................................................38-53 

Tables and figures..................................................................................................54-59 

 

Capítulo 2: Biomassa de lianas e estoque de carbono em florestas 

neotropicais...............................................................................................................60-94 

Resumo.......................................................................................................................62 

Introdução..............................................................................................................62-65 

Material e Métodos................................................................................................65-69 

Amostragem florística e estrutural.........................................................................65 

Áreas de estudo e banco de dados Alwyn H. Gentry.........................................65-66 

Biomassa e estoque de carbono.........................................................................66-67 

Variáveis ambientais..............................................................................................67 

Justificativa e hipóteses do modelo hipotético...................................................67-68 

Análises dos dados..................................................................................................69 

Resultados..............................................................................................................69-70 



 

 

Discussão...............................................................................................................70-72 

Considerações finais....................................................................................................73 

Referências............................................................................................................74-84 

Taelas e figuras......................................................................................................85-89 

Material suplementar.............................................................................................90-94 

 

Capítulo 3: Características foliares entre lianas e árvores em cerrado sensu stricto 

(Itirapina, SP).........................................................................................................95-127 

Resumo.......................................................................................................................97 

Introdução............................................................................................................98-100 

Material e Métodos....................................................................................................101 

Área de estudos: Itirapina -  cerrado sensu stricto.................................................101 

Constituição dos grupos estudados e amostragem foliar................................101-102 

Espectro de reflectância foliar e análise espectral..........................................102-103 

Conteúdo de pigmentos: Clorofila (a, b, total) e Carotenóides..............................103 

Conteúdo nutricional (fósforo e nitrogênio), área foliar específicas, massa foliar por 

unidade de área..............................................................................................103-104 

Análises dos dados................................................................................................104 

Resultados..........................................................................................................104-106 

Discussão...........................................................................................................106-109 

Considerações finais..........................................................................................109-110 

Referências........................................................................................................111-120 

Tabelas e Figuras...............................................................................................121-128 

 

Considerações finais.............................................................................................129-132 

Referências........................................................................................................130-132 

 

 

 

 

 

 

 

 

 

 

 



9 

 

Introdução Geral  

 

Lianas (trepadeiras lenhosas) são plantas heliófilas, ou seja, demandantes de luz 

para o desenvolvimento e o seu estabelecimento (Stevens 1987; Gerwing et al. 2006) e 

que investem em crescimento rápido em altura em prejuízo de sua sustentação mecânica 

(Castellanos et al. 1992; van der Heijden et al. 2013). Este grupo de plantas é 

caracterizado como parasita estrutural por utilizar o suporte físico de outras plantas para 

ascenderem ao dossel da floresta, mantendo suas raízes no solo ao longo de sua vida 

(Schnitzer & Bongers 2002; 2011). Nesse sentido, as lianas usam outros componentes 

florestais, principalmente árvores, como estrutura de suporte para maximizar a obtenção 

de luz (Hergaty 1991; Laurance et al. 2014).  

Lianas próximas filogeneticamente compartilham mecanismos de escalada 

semelhantes e colonizam árvores com características funcionais equivalentes (Zulqarnain 

et al. 2016). Contudo, o estabelecimento desse contato é marcado pela competição entre 

árvores e lianas, em todos os estágios de vida (Vleut & Pérez-Salicrup 2005; Toledo-

Aceves 2015), por água, luz e nutrientes, sendo as lianas mais eficientes que as árvores 

(Schnitzer & Bongers 2002; Schnitzer & Bongers 2011; Tobin et al. 2012; Rios et al. 

2014). Consequentemente, lianas podem reduzir as taxas de recrutamento, crescimento e 

produtividade das árvores (Schnitzer & Carson 2010; Álvarez-Casino et al. 2015). Em 

contrapartida, árvores podem apresentar atributos restritivos à colonização por lianas, por 

exemplo, casca lisa, crescimento rápido em altura, caule flexível, diâmetro menor, 

ramificações altas no tronco, folhas grandes e copa menos iluminadas (Putz 1984; Nabe-

Nielsen 2001; Pérez-Salicrup & Meijere 2005; Reddy & Parthasarathy 2006; van der 

Heijden et al. 2008; Schnitzer & Carson 2010; Sfair et al. 2013). Sendo a combinação de 

uma ou mais estratégias a forma eficiente para limitar a ocupação dessas plantas (Sfair et 

al. 2016). Assim, têm-se que lianas podem alterar a dinâmica e a diversidade da 

comunidade arbórea, atuando então como uma força seletiva (Pérez-Salicrup et al. 2001; 

Phillips et al. 2005). 

Lianas em conjunto com as demais trepadeiras herbáceas, são um grupo 

polifilético de plantas, uma vez que, durante o processo evolutivo das angiospermas, esta 

forma de vida surgiu várias vezes em eventos independentes (Burnham 2009). Um estudo 

realizado com 442 famílias de angiospermas, mostrou que 159 (36 % do total), possuem 

ao menos uma espécie de plantas escaladoras em sua composição (Gianoli 2015). O 

hábito trepador é considerado a inovação evolutiva para as angiospermas, uma vez que 



10 

 

táxons compostos por estas plantas apresentam maior riqueza de espécies do que seus 

grupos irmãos onde estas estão ausentes (Gianoli 2004; Gianoli et al. 2016).  

Lianas apresentam uma diversidade de atributos ecológicos e morfológicos, tais 

como, modo de dispersão de semente e forma de ascender à copa. Tais plantas, de maneira 

geral, desenvolvem-se no interior de florestas em estádios finais de sucessão (Gentry 

1991), são predominantemente anemocóricas (Morellato & Leitão-Filho 1998; Vargas et 

al. 2018) e têm pico de floração nas transições das estações seca e chuvosa e frutificação 

no final da estação seca (Croat 1975; Morellato & Leitão-Filho 1996). 

Lianas têm uma distribuição ampla em diversos ecossistemas, mas é na região 

tropical, principalmente em formações florestais de baixa altitude, que essas plantas 

atingem maior riqueza, diversidade e abundância (Emmos & Gentry 1983; Gentry 1991; 

Schnitzer & Bongers 2002). Em florestas tropicais 50 % das árvores carregam lianas em 

sua estrutura (Acevedo-Rodrigues 2005), o que caracteriza as lianas como responsáveis 

por cerca de 2-15 % da biomassa foliar, 5 % da biomassa lenhosa, 24-30 % serapilheira 

(Hergaty 1991; Gerwing & Farias 2000; Hora et al. 2008) e 15-45 % da diversidade 

florística (Pérez-Salicrup et al. 2001; Schnitzer & Bongers, 2002; Gallagher 2015).  

Lianas produzem mais folhas com menor massa foliar em relação as árvores, e 

elevada concentração de nitrogênio foliar, o que lhes assegura maior eficiência 

fotossintética (Kazda & Salzer 2000; Kazda et al. 2009), e por essa razão formam uma 

extensa camada de folhas sobre a copa das árvores (Hergaty & Caballé 1991; Avalos et 

al. 2007). Em Barro Colorado, Avalos et al. (1999) compararam as propriedades foliares 

de árvores e lianas, estas diferenciaram apenas quanto à transmitância, que foi 

significativamente menor nas folhas de lianas. Tal abordagem foi ampliada por Castro-

Esau et al. (2004) e Sánchez-Azofeifa et al. (2009) que buscaram distinguir os padrões 

ecofisiológicos destes grupos em florestas tropicais com graus de sazonalidade 

contrastantes. Os resultados obtidos pelos trabalhos acima citados se opuseram ao 

encontrado por Avalos et al. (1999) pois revelaram que as propriedades ópticas entre estes 

grupos diferem somente quando são oriundas de ambientes florestais com um período 

seco marcante. Além disso, suas características foliares marcantes de elevada 

transmitância e reflectância e baixa absorbância reduzem a perda de calor, a troca gasosa 

e o estresse hídrico favorecendo as taxas fotossintéticas (Sánchez-Azofeifa et al. 2009).  

Dentro deste contexto, torna-se relevante avaliar as características e as respostas foliares 

de árvores e lianas em ambientes submetidos a diferentes graus de sazonalidade (Anser 

& Martin 2012).  



11 

 

O interesse por lianas nos estudos ecológicos é crescente (Schnitzer et al. 2015), 

tendo em vista que existem evidências que estas estão aumentando em densidade, 

abundância e biomassa nas florestas tropicais (Laurance et al. 2014; Schnitzer 2015) e 

ocasionando alterações na estrutura florestal (Schnitzer & Bongers 2011; Tymen et al. 

2016). A intensidade das atividades antrópicas provocam a formação de clareiras no 

interior da floresta e o de bordas associadas a fragmentação, ambientes que favorecem a 

germinação, ocupação e super-dominância de lianas (Balch et al. 2011; Schnitzer et al. 

2014). Estas plantas mecanicamente dependentes, em uma floresta úmida na Guiana 

Francesa, aumentaram em biomassa 60 % mais rápido que as árvores, e reduziram em 4.6 

% a abundância da comunidade arbórea, enquanto incrementaram 1.8 % em sua 

abundância (Chave et al. 2008). Em Barro Colorado (Panamá), em um período de dez 

anos a proporção de copas colonizadas por lianas aumentaram de 32 % para 47 %, no 

qual aquelas com domínio severo por lianas atingiram 75 % do número de árvores 

(Ingwell et al. 2010). É documentado também em Barro Colorado, uma maior densidade 

de lianas (75-140 %), principalmente daquelas com diâmetro maior que 5 cm (Schnitzer 

et al. 2012).  

As alterações do clima são um dos fatores relevantes que ocasionam as alterações 

na dinâmica em florestas tropicais (Phillips & Lewis 2013). O rápido aquecimento global 

desde meados do século 20 alterou o padrão de circulação de ar atmosférico, 

estabelecendo áreas com secas mais intensas (Dai 2011; Trenberth et al. 2013). Os 

modelos climáticos prevêem secas severas em diversas localidades depois da metade 

deste século, tendo por consequência a diminuição da precipitação e/ou aumento da 

demanda evaporativa (Dai 2013). O período de menor disponibilidade de água é oportuno 

para o desenvolvimento das lianas (DeWalt et al. 2010; Cai et al. 2009). Isto porque, 

lianas, supostamente, possuem sistema radicular que tem ampla capacidade de 

forrageamento, tendo acesso a água e nutrientes que geralmente estão escassos durante 

os períodos mais secos (Schnitzer 2005) e um sistema vascular que garante um transporte 

rápido e eficiente de água (Andrade et al. 2005; Rosell & Olson 2014). Com isso, durante 

o período de escassez de água, quando os demais componentes florestais estão em 

dormência de suas funcionalidades, lianas são capazes de crescer 7 vezes mais que seus 

competidores (Schnitzer et al. 2005). As alterações climáticas previstas, então 

intensificam o aumento da abundância, densidade e biomassa de lianas em florestas 

tropicais secas (Gentry 1991; DeWalt et al. 2010; Yorke et al. 2013). Contudo, além das 

lianas contribuírem pouco em biomassa lenhosa (Gentry 1991; van der Heijden et al. 



12 

 

2013) e acarretarem o aumento da mortalidade arbórea (Laurance et al. 2001; van der 

Heijden et al. 2008), estas também diminuem o estoque de carbono florestal (Durán & 

Gianoli 2013; Schnitzer et al. 2014). Esta redução ocorre porque os recursos deslocados 

para alta produção de folhas são rapidamente dispensados com alta produtividade de 

serapilheira, portanto, o tempo de residência do carbono em florestas dominadas por 

lianas é baixo (Tymen et al. 2016).  

Tendo em vista a importância das lianas no cenário atual, esta tese teve como 

objetivo realizar uma pesquisa ecológica sobre a interação antagônica entre árvores e 

lianas ao longo de comunidades florestais neotropicais. Para isso, associamos base de 

dados climáticos, estruturais, biomassa, acúmulo de carbono e características foliares para 

compreender esta interação entre árvores e lianas. 

No capítulo I, intitulado “Lianas research in the Neotropics: state of the art 

and future perspectives”, fizemos uma revisão sistemática sobre o estado da arte dos 

estudos de liana na região Neotropical de 1950 a 2016.  Os trabalhos foram classificados 

por ano, país, tipo de vegetação e categoria de estudos. Nosso objetivo foi ampliar o 

interesse dos pesquisadores pelas lianas, e discutir o futuro das lianas em termos de 

controle, conservação e legislação para possibilitar a regulação e o manejo de destas 

espécies nativas que pode se tornar super-dominantes e dificultar a regeneração florestal.  

No capítulo II, intitulado “Biomassa de lianas e estoque de carbono em florestas 

neotropicais”, estabelecemos transectos em três unidades de conservação (dois cerrado 

sensu stricto e uma floresta estacional semidecidual) seguindo a metodologia proposta 

por Gentry (1982). Complementamos nossa amostragem com o banco de dados desse 

autor e calculamos parâmetros fitossociológicos, estruturais, biomassa e acúmulo de 

carbono. Com isso, determinamos os fatores climáticos e estruturais que estão 

relacionados direta e indiretamente com a biomassa de lianas e acúmulo de carbono 

florestal.  

No capítulo III, intitulado “Características foliares entre lianas e árvores em 

cerrado sensu stricto (Itirapina, SP)”, analisamos as características ecofisiológicas 

foliares entre lianas e árvores durante as estações seca e chuvosa de um fragmento de 

cerrado. Buscando distinguir estes grupos estruturais por meio dos parâmetros 

ecofisiológicos.  

 

 

 

 



13 

 

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17 

 

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African Journal of Botany 103: 108-125. 

 

 



 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Capítulo 1 

 

LIANAS RESEARCH IN THE NEOTROPICS: STATE OF THE ART AND 

FUTURE PERSPECTIVES 

 

 

 

 

 

 

 

 

 

 



19 

 

Lianas research in the Neotropics: state of the art and future perspectives 

 

Betânia Cunha Vargas1,2, Maria Tereza Gromboni-Guarantini3, Leonor Patricia Cerdeira 

Morellato1 

 

1. UNESP - Universidade Estadual Paulista, Instituto de Biociências, 

Departamento de Botânica, Laboratório de Fenologia, Rio Claro, São Paulo, Brasil. 

2. UNESP - Universidade Estadual Paulista, Instituto de Biociências, Programa de 

Pós-graduação em Biologia Vegetal, Rio Claro, São Paulo, Brasil. 

3. Instituto de Botânica, Núcleo de Pesquisa em Ecologia, São Paulo, São Paulo, 

Brasil.  

 

 

 

 



20 

 

Abstract 

          Lianas are key components of species composition, structure and dynamics of tropical 

'forests. Current global warming scenario, however, are favoring increases in the 

abundance and density of lianas in tropical forests, affecting tree growth and fertility, and 

in the number of tree injuries, which increases tree mortality over time. Here we present 

a review of studies on Neotropical lianas aiming to (i) establish the current state of 

ecological research, identifying knowledge gaps and determining new directions and 

perspectives for future studies; (ii) offer baseline knowledge to support the control, 

management and conservation of lianas and (iii) boost the interest of researchers towards 

lianas and their intricate relation with trees. We systematically reviewed the literature on 

lianas (woody climbers) since 1950 through the databases Web of Science, Google 

Scholar and Scielo using the terms liana*, “climb* plant*”, vine*, trepad*. We organized 

the literature by country, vegetation type, topic addressed and whether the study focused 

exclusively on lianas or lianas and trees.  We surveyed 425 studies in the Neotropical 

region: 255 focused on the interaction between lianas and trees and 170 exclusively on 

lianas. Studies bloomed in the 90’s, but mostly after 2000, and were carried out mainly 

in rainforests (194 – 42%) and seasonal forests (120- 26%), the majority in the Brazilian 

Amazon (162- 38%) and BCI-Panamá (74- 17%), respectively. Our review demonstrates 

the importance of lianas in seasonal forests and the lack of studies in seasonally dry 

tropical forests, woody savannas and especially extremely dry vegetations as the 

Caatinga. The main topics addressed were: phytosociology (104- 19% studies), 

ecophysiology (59- 11%), biomass (48- 9%), control and management (46- 9%), floristics 

(41- 8%), functional traits (42- 7%), phenology (39- 7%) and plant-animal interactions 

(33- 6%), and more recently global change (23- 4%), phylogeny (15- 3%), and recent 

technologies (14- 2%). Regardless of their remarkable importance in the Neotropics and 

worldwide, and their contribution for diversity, biomass and carbon flux, lianas are rarely 

used in global vegetation models and have been overlooked in restoration and 

management studies. We must consider the relevance of lianas in maintaining diversity 

and microclimate, and as resources for native animals, such as pollinators, herbivores and 

seed dispersers, as well as for traditional human communities. Research on ecophysiology 

and functional spectral traits, and the control, restoration and management of lianas are 

among the key areas of study in the Anthropocene.  

 



21 

 

Keywords: biomass; liana; climate change; ecological restoration; management; 

tropical forest 

  



22 

 

1. Introduction 

  

Climbing plants have been of interest to naturalists since the 17th century. Plumier 

(1693), Molh (1827), Palm (1827), Darwin (1865), Beal (1870), and Schenk (1892) were 

the first to describe habit-related features of climbing plants, such as using other plants as 

support and the variety of morphological mechanisms used to reach tree crowns and thus 

occupy forest canopies. Climbing plants grow fast in height but invest little in support  

relying on other plants to reach the forest canopy (Gentry 19991). However, climbers 

remain rooted to the soil during their entire life cycle (Putz and Chai, 1987), which has 

earned them the designation of structural parasites (Steven 1987; Phillips et al., 2002, 

Tobin et al., 2012). Climbing plants are classified into woody climbers (lianas), which 

show secondary growth, and herbaceous climbers (Gentry, 1991, Gerwing et al., 2006). 

According to Hergaty (1991), climbing mechanisms comprise: use of tendrils, including 

stem and leaf tendrils, twining using stems, branches or petioles and petiolules, scandent, 

which can be assisted by hooks and thorns, and the use of adventitious roots. These 

mechanisms are related to characteristics of the supporting plant, such as stem diameter, 

bark thickness and roughness, and height (Malizia and Grau, 2006, Campanello et al., 

2007b). 

Climbing plants are common across many climate and plant communities, but 

reach the highest abundance, diversity and species richness in tropical forests (Emmons 

and  Gentry, 1983, Pérez-Salicrup et al., 2001b). However, in the tropics, the contribution 

of lianas (woody climbers) to both community structure and composition is greater in 

relation to herbaceous climbers or vines (Putz, 1984b, Phillips et al., 2005, Durigon et al., 

2014).  The importance of lianas (woody climbers) to the structure and dynamics of 

tropical environments is noteworthy in the amount of food resources they offer to animals, 

including fleshy leaves, flowers and fruits, mostly during periods when tree resources are 

scarce (Morellato and Leitão-Filho, 1996, Chaves et al., 2015). Lianas (woody climbers) 

also play a role facilitating animal displacement (Nabe-Nielsen, 2001) across forest 

canopy and maintaining the microclimate of seasonal forests during the dry season, period 

in which many trees are leafless (Reston and Nepstad, 2001, Campanello et al., 2007a). 

However, lianas’ most outstanding feature is related to its increasing abundance in areas 

of early forest succession, forest gaps, edges and fragments (Phillips et al., 2002, 

Schnitzer et al., 2014, Barry et al., 2015). The unprecedented state of fragmentation of 

natural environments has created micro-environmental conditions with increased light 



23 

 

availability, which is suitable for the establishment and rapid development of lianas 

(Schnitzer and Carson, 2010, Yorke et al., 2013). As a consequence, liana abundance in 

disturbed and fragmented areas in the tropics has increased (Schnitzler et al., 2012, 

Magnano et al., 2016), causing negative impacts on the community. In such areas lianas 

may be considered as a potentially harmful super-dominant species, which requires 

management actions (Pivello et al. 2018) 

The current global warming scenario, with increased atmospheric CO2 and 

temperatures, and concurrent increases in the duration of the dry season in many regions, 

has favored the performance of lianas in tropical forests (Wrigth et al., 2004, Schnitzer, 

2015). Lianas have been increasing in abundance, basal area and/or biomass in several 

tropical forest sites (Schnitzer et al., 2012, Philips and Lewis, 2014) and have already 

been considered as an indicator of global change effects in such ecosystems (Phillips et 

al., 2002, Pan et al,. 2011). As a result, trees sustaining lianas are affected, for example, 

by stems break and injury, favoring the attack of herbivores and pathogens (Putz, 1991, 

van der Heijden and Phillips, 2009). Those damages may lead to tree mortality, biomass 

reduction and, consequently, a decline in CO2 storage (Schnitzer et al., 2014, van der 

Heijden et al., 2015). The dominance of lianas in the canopy of tropical forests can also 

affect light availability in the understory (Avalos et al., 1999, 2007), seedling 

establishment, development and recruitment (Campanello et al., 2007b, Schnitzer and 

Carson et al., 2010, DeWalt et al., 2015), sap flow velocity (Tobin et al., 2012), canopy 

height (Pérez-Salicrup, 2001b, Tymen et al. 2016), and overall tree fecundity (Kainer et 

al., 2006, Grogan and  Landis, 2009).  

Nevertheless, the effects of lianas in tropical tree communities are not uniform 

(Schnitzer et al., 2016). Liana abundance, density, biomass, and basal area are usually 

positively related to dry season duration and severity and negatively related to mean 

annual precipitation (Gentry, 1991, Schnitzer, 2005, DeWalt et al., 2010). However, 

Laurance et al. (2014) detected an increase in liana density in the Amazonian rainforest 

that was not related to precipitation variation but rather to increasing tree turnover, 

mortality and recruitment. Conversely, Marvin et al. (2015) did not find significant 

differences between trees and lianas in relation to CO2 increase and fertilization, both in 

the dry and rainy seasons. Therefore, including lianas in vegetation models and improving 

the quality of data regarding this plant group is crucial to better understand tropical forest 

dynamics and forecast the possible changes of ecosystems worldwide (Durán et al., 2015, 

Schnitzer et al., 2016). 



24 

 

Here we present a review of studies on the biology of lianas in the Neotropical 

region aiming to (i) establish the current state of ecological research regarding lianas, 

identify the knowledge gaps and determine new directions and perspectives for future 

studies, (ii) offer baseline knowledge to support the control, management and 

conservation of lianas and (iii) boost the interest of researchers towards lianas and their 

intricate relation with trees. We systematically reviewed the literature on lianas (woody 

climbers) regarding studies conducted in the Neotropics (26º N – 26º S) since 1950. We 

organized the literature by country, vegetation type, topic addressed by the study and 

whether it focused exclusively on lianas or on the relationship between lianas and trees. 

We drew some considerations about lianas, their importance and relevance to vegetation 

worldwide, and the key areas of study in the Anthropocene.  

 

2. Criteria for the inclusion of studies, vegetation classification and research 

topics 

 

In this systematic review, we searched through the databases of Web of Science 

using the terms liana*, “climb* plant*”, vine*, trepad* with no limitation by country or 

year. We also searched the online data banks of Google Scholar and Scielo using the same 

terms. From all studies surveyed, we selected only those conducted in the Neotropical 

region (26ºN - 26ºS) between 1950 and 2018 (august). Additionally, we excluded all 

studies that addressed agricultural and cultivated species.  

Each paper surveyed was classified by country, vegetation type, topic and whether 

it focused exclusively on lianas or on the relationship between trees and lianas. To assign 

a vegetation category to each of our study site, we adopted the nine vegetation types used 

by Mendoza et al. (2017), based on the ecoregions of Olson et al. (2001), as follows: 

rainforests, seasonal forests, tropical dry forests, cerrado woodlands, desert and xeric 

shrublands, open grassy savannas, temperate evergreen forests, montane formations and 

seasonally flooded forests. We added the ecotone type when the study site was located in 

transition zones (between ecoregions). We classified the sites into the “wide distribution” 

category when a study used databases involving several countries in the Neotropics, and 

into the “no vegetation” category when the vegetation type of the study was not disclosed 

(Table 1). If a study included more than one country or vegetation formation we 

considered each site as an independent dataset (e.g. Mendoza et al., 2017), even when the 

vegetation types showed the same geographical coordinates. 



25 

 

We established 15 topics that could have been addressed in a given study: 

taxonomy, phylogeny, floristics, ontogeny and development, anatomy, phytosociology, 

biomass, control and management, functional traits (habit, seed dispersal, climbing mode, 

morphology of supporting trees), ecophysiology (structure, chemistry and physiology of 

leaf and canopy, and spectral reflectance), phenology, ethnobotany, plant-animal 

interaction, new technologies (satellite data), and global change. Each study could be 

classified in one or more topics. 

 

3. Review on liana research in the Neotropical region 

3.1. Liana research over time and topics addressed among the research on lianas 

 

We found 1060 articles using the search criterion described above. We identified 

425 studies that were carried out in the Neotropical region, of which most (255- 60%) 

focused on the interaction between lianas and trees and 170 (40%) focused exclusively 

on lianas (Table S.1, supplementary-digital), and are here described over time (Fig. 1). 

The main topics addressed among these studies surveyed were: phytosociology (104-19% 

studies), ecophysiology (59-11%), biomass (48-9%), control and management (46-9%), 

and functional traits (42-8%) (Table 1). Such descriptive studies are followed by 

ecological research on floristic (41-8%), phenology (39-7%), and plant-animal 

interactions (33-6%) (Table 1). 

Up to the early 90s, research on lianas showed only a slight increase over time, 

however, after 2000 the number of studies addressing lianas increased exponentially (Fig. 

1). Scholander et al. (1957) carried out the first study with lianas in the Neotropics, which 

focused on the physiology of Tetracera sp. (Dilleniaceae) in Barro Colorado Island. 

However, it was only in the 1970s that new studies on the ecology of lianas in the 

Neotropics were published:  Janzen (1971) addressed strategies to avoid seed predation 

of Dioclea sp. and the classic study by Gentry (1974) on the phenology of Central 

American Bignoniaceae. Up to the early 1990s, lianas were nearly neglected and 

understudied by the scientific community, except by the pioneer work of Gentry (1985, 

1987, 1991, 1992) and the studies by Putz (1980), Putz (1984b), Putz and Chai (1987) 

and Boom and  Mori (1982), describing the mechanisms in which trees avoid lianas. In 

fact, the 1990s mark the flourishing of papers regarding lianas, which began with the 

seminal book by Putz and Mooney (1991) “The biology of vines”. A very rich and 

diversified literature was published during the 90s, including papers that addressed 



26 

 

ethnobotany and the importance of lianas for traditional communities (Arenas and  

Gilbert, 1987, Gentry, 1992, Phillips and  Gentry, 1993, Paz y Niño et al. 1995), and 

papers describing the contrasting and complementary phenology patterns between lianas 

and trees (Morellato and Leitão-Filho, 1996) and their role as food resource to the fauna 

(Chiarello, 1998, Galleti et al., 1994, Wallace et al. 1998).  

During the 2000s, the number of studies increased by more than three folds, 

establishing lianas as a key field of study in the Neotropics (Fig. 1). The exponential 

growth in scientific papers with lianas observed in our survey is similar to that shown by 

Schnitzer et al. (2015). The number of publications involving liana ecology increased 

more than publications involving the ecology of other groups (Schnitzer et al., 2015, 

Mendoza et al., 2017). Among the topics (Table 1), research focusing on the interaction 

between trees and lianas followed the same proportion of topics as for the total summary. 

While we found that studies focusing exclusively on lianas are rather basic research and 

combine most of the research on phytosociology and taxonomy (29-14%, each), anatomy 

(21-10%), floristic (19- 9%), ecophysiology (17- 8%), functional traits, phylogeny and 

phenology (12-4%, each), and biomass and plant-animal interaction (11- 5% each). 

The phytosociology studies, mostly describing the importance of lianas as a 

structural component of tropical communities (Ibarra-Marínquez and Martínez-Ramos, 

2002, Phillips et al., 2002, Burnham, 2004, Gerwing, 2004; Franci et al., 2016). The 

studies have focused too mainly on liana dynamics in gaps and forest areas of different 

successional stages and have demonstrated how well these plants grow in such 

environments (Gerwing and Uhl, 2002: Madeira et al., 2009; Yorke et al., 2013, Barry et 

al. 2015). It was during 2000s that Schnitzer (2005) discussed the importance of the dry 

season for the development of lianas, and Schnitzer et al. (2006) proposed an allometric 

equation to calculate liana biomass after demonstrating how different types of diameter 

measurements affect liana biomass, basal area and abundance estimates. Subsequently, 

the first studies proposing general guidelines and protocols to standardize the methods 

employed with lianas (Gerwing et al., 2006, Schnitzer et al., 2008).  

The studies with ecophysiology was also relevant, with many studies focusing on 

how lianas are affected by light and water availability (Sanches and Válio, 2002, Reston 

and Nepstad, 2004). For instance, Molina-Freaner et al. (2004) observed a decrease in 

leaf parameters (total and specific leaf area) with a decrease in precipitation and an 

increase in water stress and described how such factors could limit the local distribution 

of lianas. Ecophysiological studies provided evidence, in terms of leaf-level traits (leaf 



27 

 

area index, nutrient content and leaf dry mass) of the competitive advantages lianas have 

over trees (Selaya and Anten, 2008, Deurwaerder et al. 2018). Through these leaf 

distinctions, remote sensing studies start to map and monitor forest areas dominated by 

lianas, producing large-scale biomass estimates and revealing changes on forest structure 

across landscapes and biomes (Kalácska et al., 2007, Foster et al., 2008, Ledo et al., 2016, 

Marvin et al., 2016, Tymen et al., 2016). News technologies with terrestrial laser scanner 

and imagens hiperespectrais, detected structural changes related to lianas dominance 

(Sánchez-Azofeifa et al. 2017; Moorthy et al. 2018). Such new sets of data allow the 

identification of areas that need liana management and control, potentially enhancing the 

local experiences (e.g. César et al., 2016). 

Research has shown that lianas have become increasingly important in forest 

ecosystems (Schnitzer et al., 2012, Laurance et al., 2014b), occupying forest canopy 

(Ingwell et al., 2010, van der Heijden et al., 2010, Campanello et al., 2012) and producing 

structural and economic damage to trees (Nascimento et al., 2013, Staudhammer et al., 

2013). Likewise, studies began to evaluate the effect of experimentally removing lianas 

as a mean to manage forest remnants (Schnitzer et al., 2010, César et al., 2016), evaluate 

phenological parameters and reproductive advantages, such as production of fruits and 

seeds for species of economic potential (Kainer et al., 2006, Fonseca et al., 2009, Grogan 

and Landis, 2009), the fast regeneration rates of lianas, even after removal (Campanello 

et al., 2012), and how the increase of liana abundance (Nascimento et al., 2013, Laurance 

et al., 2014b) and biomass (Laurance et al., 2001, Tobin et al., 2012, van der Heijden et 

al., 2015) affect community structure throughout time.Questions concerning how climate 

change will affect forests and fragments and liana dominance also began to be raised by 

researchers (Durán et al., 2015, van der Heijden et al., 2015, Collins et al., 2016). 

Likewise, quantify the overall negative effects of lianas on forest structure, tree biomass 

and carbon storage in forested ecosystems (Durán et al., 2015, Zelarayán et al., 2015). 

All taxonomic studies focused exclusively on lianas, and include both reviews 

(Nee 2007, Pool, 2009, Trethowan et al., 2015) and new species descriptions (Camargo 

and Tozzi, 2014,  Aguirre-Morales et al., 2016). Most of the taxonomic studies (14) were 

on a single family, the Bignoniaceae (Hauk, 1997, Pool 2007, Udulutsch et al., 2009, 

Zuntini and Lohmann, 2014, Fonseca et al., 2016), stressing the relevance of this group 

of lianas (Gentry, 1991, Santos et al., 2009). Bignoniaceae is among the richest families 

in lianas in Neotropical (Gentry 1995). In addition, the largest tribe in this family, 

Bignonieae, is predominate by lianas (Lohmann 2006). Worth mentioning the studies that 



28 

 

described lianas floral pattern (Alcantara et al. 2013) and the association between 

molecular analysis and palynology to identify liana species (Milward-Azevedo et al. 

2014). Anatomical studies were practically exclusive with lianas and have focused on 

secondary growth and cambial activity, mainly of species from the most representative 

families, such as Sapindaceae and Bignoniaceae (Tamaio et al., 2011, Angyalossy et al., 

2012, Pace et al., 2015), including dendroecology and dendrochronology of lianas, and 

the importance of rainfall on liana growth rates (Lima et al., 2010, Brandes et al., 2011). 

Floristc also was important to lianas, those combining floristics with functional traits 

(habit, dispersal mode and climbing mechanisms) were frequent and have resulted in 

important identification keys (Araújo and Alves, 2010, Villagra and Roumaniuc Neto, 

2011).  

 

3.2. Liana research in the Neotropics:  geographical distribution and vegetation 

types  

  

Our review showed that research on lianas has been conducted in 20 countries, 

mainly in Central America (particularly Panama, Costa Rica), South America (especially 

in Brazil and other Amazon countries such as Bolivia, Ecuador, Peru, and the Guianas) 

and Mexico (North America, Fig. 2;3). Most studies were carried out in Brazil (162- 

38%), Panama (74- 17%), and Mexico (45- 10%), or were placed into the wide 

distribution category (35- 8%, Fig. 2). The studies in Brazil are mainly in the Amazon 

forest and the Atlantic forest from southeastern Brazil (Fig. 3). The Panamanian studies 

were predominantly conducted at Barro Colorado Island. In contrast with liana research 

in Brazil, the Panamanian studies have long-term data (frequency, density, dynamics and 

biomass) from permanent plots (Wright et al., 2004, Barry et al., 2015, Ledo et al., 2016). 

In the wide distribution category, most studies focused exclusively on lianas (Fig. 2), 

including phylogenetic and taxonomic reviews, some addressing exclusively the 

Bignoniaceae family (Hauk, 1997, Pool, 2009, Alcantara et al., 2013).  

Our survey detected a dominance of studies conducted in rainforests (194- 42%) 

and seasonal forests (120- 26%), both included into the moist broadleaf forest biome 

according to Olson et al. (2001, Fig. 3, Table 2) and occupying the largest area within the 

Neotropical region. Although the rainforest is the vegetation type where most tropical 

research, including those on lianas, have been conducted up to today, we observed a 



29 

 

significant amount of studies conducted in vegetations types that undergo some degree of 

climate seasonality with a marked dry season: seasonal forests (120), tropical dry forests 

(30- 6%) and cerrado woodlands (11- 2%) (Table 1). Both seasonality and a lower total 

annual precipitation have been pointed out as key climatic factors favoring high liana 

abundance and basal area in tropical forests (Gentry, 1991, Schnitzer, 2005). However, 

severe dry seasons, as occurs in the Brazilian semiarid (Caatinga dry forest and xeric 

shrublands), restricts liana distribution and abundance. Other climatic factors limiting 

liana abundance are altitude and minimum temperatures, as they limit water conduction 

in the xylem, eventually leading to embolism, and thus restrict the occurrence of woody 

lianas in high elevations (Alves et al., 2011). Therefore, although lianas have wide and 

long vessel elements that ensure efficient water transport, these characteristics also make 

lianas vulnerable to cold temperatures and water stress (Hu et al., 2010).  

 

4.  Lianas: its antagonistic relation to trees and importance to biodiversity 

 

Lianas face the trade-off between investing little in secondary growth, lacking 

thereby self-support, and investing in rapid growth in height, to reach the forest overstory 

where light availability is abundant (Castellanos et al., 1992, van der Heijden et al., 2013). 

Given this trade-off and the fact that these plants need abundant sunlight, lianas need the 

structural support of trees and can be defined as structural parasites (Ingwell et al., 2010, 

Tang et al., 2012, Stewart and Schnitzer, 2017). This dependency on trees may occur as 

seedlings or in later stages of their life cycles (Vleut and Pérez-Salicrup, 2005, Avalos 

and Mulkey, 2014). Lianas reach the canopy efficiently mostly through specialized 

organs, such as tendrils and adventitious roots (Hergaty, 1991, Putz and Holbrook, 1991). 

However, the effect of lianas on tree communities is not uniform given that trees show 

diverse ecological and morphological strategies, such as rapid growth in height, height, 

bark diameter, thickness and roughness, stem flexibility, to avoid or reduce liana 

colonization (Putz, 1980, van der Heijden et al., 2010, Garbin et al., 2014), thus 

restraining the competitive interaction between lianas and trees (Sfair et al., 2016). 

Alternatively, lianas may neutralize such mechanisms by using other climbers as support 

(facilitated colonization) or by using neighboring plants to reach the canopy (Putz, 1984b, 

van der Heijden et al., 2008, Yorke et al., 2013). 

Once the interaction liana-tree is established, competition for resources (water, 

light and nutrients) begins, where lianas are usually more efficient (Schnitzer and 



30 

 

Bongers, 2002, Schnitzer and Bongers, 2011, Tobin et al., 2012), especially in seasonally 

dry forests. Schnitzer (2005) suggests that lianas have a root and vascular system that is 

highly efficient in water transport and nutrient uptake, even in periods of water shortage, 

and Rosell and Olson (2014) observed that lianas have a higher density of vessels than 

self-supporting plants. Therefore, during periods when many plant growth forms exhibit 

some degree of dormancy, ceasing or reducing their metabolic activity, lianas continue 

photosynthetically active, assimilating carbon, and producing and maintaining their 

leaves and reproductive structures. Thus, lianas can be aggressive competitors that forage 

efficiently for resources, expanding and dominating the forest canopy during drier periods 

(Schnitze et al., 2005). 

Nonetheless, lianas lose their competitive advantage in forests where the dry 

season is too severe (Carvalho et al., 2016), given their roots become more superficial 

with limiting water availability (Andrade et al., 2005, Johnson et al., 2013; De 

Deurwaerder et al. 2018). Additionally, lianas do not tolerate water stress and the high 

vapor pressure deficit (Schnitzer et al., 2005), shedding its leaves at the beginning of the 

dry season to avoid water stress and damages to the xylem (Carvalho et al., 2016). 

Moreover, climbing species from Vitaceae and Cucurbitaceae family (important in arid 

and semi-arid ecosystems) use the Crassulacean acid metabolism (CAM) pathway when 

submitted to water stress in order to improve water use efficiency (Rundel and Franklin, 

1991). 

Competition for resources between trees and lianas is highest at canopy levels. 

Once lianas reach tree crowns, they form a layer of leaves, intercepting light before liana-

supporting trees (Avalos et al., 1999, 2007, Castro-Esau et al., 2004). Thus, canopy 

spectral signatures may be altered in areas dominated by lianas (Castro-Esau et al. 2004, 

Sanchéz-Azofeifa and Castro-Esau, 2006). This is particularly noticeable in seasonally 

dry forests as liana leaves, in contrast to tree leaves, are narrower, containing more 

aerenchym, water, nitrogen and phosphorous, and low concentrations of photosynthetic 

pigments (Sanchéz-Azofeifa and Castro-Esau, 2006, Sanchéz-Azofeifa et al., 2009a). 

These leaf traits, coupled with its higher specific leaf area and water use efficiency, give 

lianas a higher capacity to allocate resources and promote higher photosynthetic rates in 

relation to trees. On the other hand, trees show resource conservation strategies in order 

to avoid water stress and survive periods of leaf shedding and reduced growth (Pooter 

and De Jong’s, 1999, Cai et al., 2009, Sanchéz-Azofeifa et al., 2009a, Ball et al., 2015). 

Moreover, although liana leaves are less tolerant to the dry season than tree leaves, lianas 



31 

 

maintain cell turgidity through osmotic control even in dry conditions, enabling plant 

growth (Márechaux et al., 2017). 

Lianas are important floristic, functional and structural components of tropical 

forests (Gentry, 1991, Morellato and Leitão-Filho, 1996). Lianas contribute greatly to 

carbon and nitrogen dynamics in these ecosystems as the litter layer contains around 24-

30% of liana leaves and, in a smaller proportion, flowers, fruits, seeds and stems (Hergaty, 

1991, Hora, 2008). Liana phenology is complementary to tree phenology, becoming an 

alternative food source during periods of resource scarcity for animals (Morellato and 

Leitão-Filho, 1996, Dunn et al., 2012). For instance, lianas may account for nearly 30% 

of a monkey’s diet (Galetti and Pedroni, 1994, Souza-Alves et al., 2011) and be the main 

food source for bees (Odegaard et al., 2000). 

 Regarding the contribution to plant species diversity, lianas appeared multiple 

times throughout the evolution of angiosperms and are present in nearly 159 botanical 

families (Gentry, 1991).  The life form is considered a key innovation given that clades 

containing lianas are more diverse in relation to those where lianas are absent (Gianoli, 

2004, Gianoli et al., 2016). Thus, lianas represent 18-25% of the taxonomic diversity in 

tropical forests (Gentry, 1991, Schnitzer and Bongers, 2002, Schnitzer et al., 2012) and 

may represent up to 44% of the diversity in the Amazonian forests (Pérez-Salicrup et al., 

2001b). 

 

5.  Global change and the future of lianas 

 

The climate change forecast predicts an average global surface temperature 

increase between 1 to 4 oC by 2100 (IPCC, 2013). In tropical forests, such temperature 

raise may result in longer and more intense dry seasons (Joetzjer et al., 2013) and irregular 

or extreme rainfall regimes (Cusack et al., 2016). The dry season has been pointed out as 

a key factor driving liana abundance, basal area, density and biomass (Gentry, 1991, 

Schnitzer, 2005, DeWalt et al., 2010, Sfair et al., 2016). Therefore, climate change may 

favor lianas in detriment of trees, especially in seasonally dry forests (Yorke et al., 2013, 

Schnitzer et al., 2014, Durán et al., 2015, Maréchaux et al., 2017). Collins et al. (2016) 

compared roots of lianas and trees and concluded that the association with mycorrhiza is 

less frequent in lianas than in trees. The authors state that the increasing temperatures due 

to climate change may accelerate nutrient mineralization in the soil, thus the advantages 



32 

 

bestowed to trees from the association with mycorrhiza (nutrient uptake, capability to 

avoid water loss and pathogens) would become insignificant, benefiting lianas.  

Aside from changes in temperature, climate change research has demonstrated the 

increase of atmospheric CO2 (Cramer et al., 2001, Eissenhauer et al., 2011). Eamus (1991) 

observed that an increase in CO2, coupled with elevated temperatures, reduce stomatal 

conductance and plants can obtain more carbon with less water loss. Additionally, 

elevated CO2 leads to increments in photosynthetic rates resulting in plant growth 

(Eamus, 1991) and, therefore, promotes a positive response from both trees and lianas 

(Phillips and Gentry, 1994). However, resource acquisition in lianas is extremely efficient 

given their leaf characteristics (high nitrogen and phosphorous concentrations, low 

carbon concentration, low tissue density) and root systems (deep roots and efficient water 

and nutrient transport) (Sanchéz-Azofeifa and Castro-Esau, 2006, Sanchéz-Azofeifa et 

al., 2009a, Collins et al., 2016). Thus, climate change favors higher photosynthetic rates 

in lianas in relation to trees (Granados and Körner, 2002, Wrigth et al., 2004, Phillips et 

al., 2004), which leads to higher growth, fecundity and recruitment rates in lianas (Phillips 

et al., 2002, Hättenschwiller and Köner, 2003, Mohan et al., 2006, Yorke et al., 2013) and 

increases their biomass contribution in forests (Granados and Köner, 2002, Phillips et al., 

2002, van der Heijden et al., 2015). 

Liana leaf and root systems alter water and nutrient availability in the soil, increase 

litter production and accelerate decomposition, thus altering biogeochemical cycles 

(Sayer et al., 2011, Collins et al., 2016). Lianas also promote structural changes in forest 

communities reducing forest carbon stocks by 35 to 50% (Durán and Gianoli, 2013, van 

der Heijden et al., 2015) as lianas increase tree mortality (Ingwell et al., 2010) and reduce 

tree fecundity (Kainer et al., 2006, Staudhammer et al., 2013) and biomass (van der 

Heijden et al., 2009, Tobin et al., 2012, Phillips and Lewis, 2014, van der Heijden et al., 

2015). Additionally, climate change increases treefall gaps, which also favors lianas 

(Clark et al., 2010, Schnitzer et al., 2014). 

In contrast, Schnitzer (2015) states that the direct effect of elevated CO2 in the 

antagonistic relationship between trees and lianas is debatable due to the lack of 

experimental studies that evaluate leaf traits associated with size, age and species identity 

of both trees and lianas. Moreover, Marvin et al. (2015) simulated the effect of CO2 

concentrations in seedlings of trees and lianas in both rainy and dry conditions and did 

not observe significant physiological differences favoring lianas in relation to trees. They 

conclude that raising CO2 is not the main mechanism related to the increase of lianas in 



33 

 

tropical forests. The elevation of CO2 coupled with the severity of the dry season 

(Schnitzer, 2005, Cai et al., 2009, Tobin et al., 2012), the increase in temperature (Anser 

and Martin, 2012) and in nutrient availability in the soil (Schnitzer et al., 2011) are likely 

driving the dominance of lianas in tropical forests.  

 

6.  Control, management and conservation of lianas 

 

We have highlighted the effect of climate change as a factor favoring lianas in 

tropical forests. However, human-related activities such as agriculture, anthropogenic 

fires and fragmentation may also be associated to the super-dominance lianas (Balch et 

al., 2011, Schnitzer, 2015). Between 1980-2010 Latin American forest had deforestation 

grater than global average (Armenteras et al. 2017). In Brazil, between 1970 and 2015, 

lost 768, 938 Km² of forest area (Burton 2017). This has led to disturbed environments, 

such as edges and gaps, where light availability and temperatures are higher, favoring the 

recruitment and establishment of species typical of early successional stages, including 

lianas (Schnitzer and Carson, 2001, Schnitzer and Carson, 2010). Such conditions favor 

liana abundance, density and distribution (Malizia et al., 2010, Schnitzer et al., 2014). 

Clonal reproduction (Schnitzer et al., 2012, Yorke et al., 2013) also allows lianas to 

rapidly and efficiently colonize gaps and disturbed areas (Dalling et al., 2012, Ledo and 

Schnitzer, 2014). Lianas associated to forest canopies can regenerate quickly after 

treefalls, rooting and developing new stems from ramets (Putz, 1984a, Schnitzer and 

Bongers, 2011). Therefore, increasing disturbances caused by changes in rainfall, 

atmospheric CO2 concentration or in the duration and severity of the dry season can 

stimulate lianas to colonize such altered environments through clonal reproduction, which 

may explain the increased liana abundance and biomass in tropical forests (Schnitzer and 

Bongers, 2011, Yorke et al., 2013). 

  Fires are frequently used to clear forest areas for agriculture, reducing biomass 

and preventing recruitment of new trees (Balck et al., 2011, Devisscher et al. 2016). 

According to Nelson (1994), fire plays an important role in the dominance of lianas in the 

Brazilian Amazon forest. Pinard (1999) and Gerwing (2002) agree and have observed an 

increase in liana recruitment after fires, which limits the regeneration of the tree 

component.   

Lianas can be a hazard for both logging and silviculture practices as they can 

damage barks and stems of species of economic interest (Putz, 1991, Kainer et al., 2007, 



34 

 

Grogan and Landis, 2009). Thus, removing lianas is an alternative to reduce their 

competitive effects, benefitting the tree community. Although lianas super-dominance 

impacts the community structure and diversity, no legislation regulates the management 

of native super-dominant species (Pivello et al. 2018).  Reid et al. (2015) observed that 

cutting lianas can alter soil humidity according to soil depth: in shallow soils, liana 

removal increases soil humidity by 6.5 % during drier periods, while in deeper soils, liana 

removal can increase soil humidity up to 25 %. While monitoring the response of adult 

individuals of Senna multijuga after the removal of lianas, Pérez-Salicrup and Barker 

(2000) observed that lianas limit tree growth by reducing the amount of water available. 

Removing lianas from trees can increase tree sap velocity by 60% (Álvarez-Casino et al., 

2015) and tree fruiting probability by up to 25% (Fonseca et al., 2009). For instance, 

Bertholletia excelsa trees not associated to lianas produce more fruits and heavier nuts 

than liana-infested trees (Kainer et al., 2007). Likewise, biomass, basal area, canopy 

openings and seedling growth show a significant increase when lianas are removed 

(Venturoli et al., 2015, César et al., 2016). 

The detrimental effects of lianas on trees is widely recognized, thus managing and 

monitoring liana growth is needed. New technologies, such as satellite derived 

information and remote sensing monitoring, have been used to detect areas where lianas 

prevail (Li et al, 2011, Marvin et al., 2016). Lianas can be detected by an increase in the 

normalized difference vegetation index (NDVI) and in the photosynthetic vegetation 

index (PV) in areas where timber harvesting occurs as they are marked by a great number 

of gaps (Broadment et al., 2006).  

The greater spectral chlorophyll indices from liana leaves in relation to tree leaves 

is another parameter that makes distinguishing them possible (Castro-Esau et al. 2004, 

Sanchéz-Azofeifa and Castro-Esau, 2006, Sanchéz-Azofeifa et al., 2009b). The LIDAR 

dataset (a cloud of laser echoes originating from ground and vegetation), the high-

resolution color-infrared videography (high-resolution viodeography) and the 

hyperspectral imagery from EO-1 Hyperion (sensor hyperspectral aboard the EO-1 

satellite) have been employed to detect and monitor lianas (Foster et al., 2008, Tymen et 

al., 2016). Areas dominated by lianas appear in hyperspectral imagery as closed areas 

with a higher brightness and greenness index in comparison to canopies lacking lianas 

(Kalácska et al., 2007, Foster et al., 2008, Tymen et al., 2016). This occurs because lianas 

modify canopy architecture as they form a continuous and extensive layer of leaves on 

top of trees, thus reducing shaded areas (Avalos et al., 1999, 2007, Kalácska et al., 2007).  



35 

 

Alternatively, tree leaves usually do not form a horizontal angle and their orientation 

shifts throughout the day (Gamon et al., 2005).  

On the other hand, the ecological and ecophysiological characteristics that allows 

lianas to dominate tropical forests are the same traits that make lianas suitable for forest 

restoration. In a recent review, Campbell et al. (2015) show that the fast growth and high 

leaf productivity of lianas may reduce the establishment of shade intolerant plants, 

minimize the time for canopy closure, and contribute to the formation of wildlife 

corridors, facilitating animal displacement and reducing the financial cost of ecological 

restorations (Pérez-Salicrup et al., 2001a). Lianas may also efficiently close fragment 

edges that are being restored, thus avoiding damages caused by winds, reducing the 

proliferation of weeds and the incidence of fires (Santiago et al. 2010, Balch et al., 2011). 

Liana leaves have high nutrient concentrations and are major contributors to litter 

production, their leaves decompose quickly and return a great amount of nutrients to the 

soil, which helps maintain soil microfauna and reduces the risk of erosion (Chalker-Scott, 

2007, Tang et al., 2012). 

Lianas are also an important non-timber forest product (NTFP), being a significant 

economic source for traditional communities (Bezecry, 2005, Guadagnin and Gravato, 

2013). For instance, stems of lianas may be used to produce handicraft pieces (Gentry 

1992, Tamaio 2011) and stems and fruits of Odontocarya asarifolia (Menispermaceae) 

are widely consumed in the diet of indigenous families of the Chaco region in South 

America (Arenas and Gilbert 1987). Gentry (1992) highlights the importance of species 

of Bignoniaceae for human populations in South and Central America, where lianas have 

been used as ornamental (Podranea ricasoliana, Pyrostegia venusta), as spices (species 

of Mansoa- scent resembles garlic and Tynanthus sp.- sweet scent that resembles clove), 

to manufacture sieves and filters (species of Fridericia sp.), in construction, as dyes for 

handicraft pieces and body paintings (Fridericia chica, Cybistax antisyphilitica) and 

during religious rituals (Bignonia aequinoctialis, Cuspidaria inaequalis). Gentry (1992) 

also points out that around 27 genera of lianas have some medicinal property, such as 

clove, garlic and almond scented species that are used as insect repellents and in the 

treatment of conjunctivitis, hepatitis, tuberculosis, rheumatism and fever (stems of 

Stizophyllum sp.); additionally, some species may be used as hallucinogens (Bignonia 

noctura, Bignonia hyacinthina) or as tools for fishing (Adenocalymma allamandiflorum) 

and hunting crabs (Dolichandra quadrivalvis). Tynanthus fasciculatus (Bignoniaceae) is 

a source of drinking water and may be used as an antihelminthic and as ornamental (Lopes 



36 

 

et al. 2008). Davilla nitida (Dilleniaceae) is used to manufacture fishing lines and as a 

source of water to treat eye and stomach infections, while Dioclea sp. (Fabaceae) is 

antiophidic but is also applied on bug bites (DeWalt et al., 1999).  

 Therefore, aside from being important for forest composition and diversity, lianas 

are also extremely relevant for traditional communities, where they are used in a variety 

of daily activities. Additionally, the sale of manufactured goods made with liana stems 

generates an income of around USD 1926/year for these communities (Guadagnin and 

Gravato, 2013). 

 

7.  Future perspectives and conclusions 

 

This review shows that liana neotropical studies are concentrated in moist forests, 

particularly in Brazil (across Amazon and Atlantic forests) and Panama (Barro Colorado 

Island). Tropical dry forests are still understudied although they show favorable climatic 

conditions for the establishment and development of lianas. The best studied tropical dry 

forests are in Mexico, however, of the 45 studies surveyed in Mexico, only twelve were 

in tropical dry forests. Studies in environments as the Brazilian dry caatinga and the 

seasonally flooded forests are still scarce (Table 1). Cavalho et al. (2016) points out that 

understanding how lianas establish and develop in extreme environments (under severe 

dry seasons) may uncover different responses from what is known from moist 

environments. We also suggest diversifying the research, targeting species from other 

liana-rich families, such as Apocynaceae (Sales et al., 2006), Cucurbitaceae (Nee, 2007, 

Gomes-Costa et al., 2015), Passifloraceae (Cervi and Linsingen, 2008) and Fabaceae 

(Carmargo and Tozzi, 2014). 

Lianas are increasing in density, abundance and basal area with consequences to 

carbon stocks, tree survival, recruitment and growth, and detrimental effects on the 

productivity of species of economic interest. Remote sensing techniques have improved 

the identification of areas that urgently need liana control and management (Marvin et 

al., 2016). Even though lianas affect forest carbon stocks, they are rarely included in 

tropical forest studies that address that issue (Durán and Gianoli, 2013, Durán et al., 

2015). Therefore, it is extremely important to consider lianas in global vegetation models 

to ensure accurate predictions of future changes on carbon stocks and carbon cycles 

(Schnitzer 2016). 



37 

 

Management and restoration actions targeting only tree communities are 

inadequate given lianas’ importance for tropical forests structure and function, increasing 

diversity, maintaining the microclimate and providing resources for animals during 

periods of scarcity. Lianas are also important for traditional communities, being used with 

multiple purposes: nutritional, cultural, structural and pharmaceutical, generating income 

when handicraft pieces using liana stems are commercialized. Therefore, promoting 

adequate conservation and management actions ensures forest maintenance, but also 

respects traditional uses as many populations still use the forest as their main resource 

source. 

 

Acknowledgements 

 

            Our research was supported by FAPESP, the São Paulo Research Foundation (grants 

#2013/50155-0 FAPESP-Microsoft Research, #2010/51307-0 FAPESP-VALE-

FAPEMIG and grant #2009/54208-6 EMU). BCV received a doctoral fellowship and 

additional financial support from CAPES. LPCM receives a Research Productivity 

Fellowship from CNPq, the National Council for Scientific and Technological 

Development. We have also benefited from funds from CAPES-PROAP (Coordination 

for the Improvement of Higher Education Personnel).  We are also grateful to Talita Zupo 

for English review.  

 

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