Growth and Shading Metrics Reveal Functional Differences Among Similar Inga Species for Forest Restoration
Mayara Monteiro Ferreira, André Cesar Furlaneto Sampaio, Jean Freitag Kramer, Marília Borgo, Nelson Luiz Cosmo, Christopher Thomas Blum, Chaiane Rodrigues Schneider, Alexandre Dal Forno Mastella, Sarah Domingues dos Santos, Alessandro Camargo Angelo
Abstract
Degradation of the Ombrophilous Dense Forest in the Lowlands, driven by cattle ranching and invasive species, limits natural regeneration. Species of the genus Inga are widely used as facilitators in restoration in this ecosystem, but are frequently treated as a uniform group despite their functional differences. We compared the growth and shading capacity of I. edulis, I. marginata, and I. laurina to provide better criteria for species selection in restoration plantings. Using morphometric variables (diameter at breast height, total height) and derived metrics (basal area, volume, and slenderness index), combined with qualitative crown assessments, we found that I. edulis excels in fast growth and high biomass production, I. laurina demonstrates greater structural robustness and denser crowns, and I. marginata shows intermediate performance with adaptability to degraded soils. These interspecific differences highlight the importance of combining species with complementary traits to enhance restoration efficiency in the Atlantic Forest.
Keywords
References
- Alvares CA, Stape JL, Sentelhas PC, Gonçalves JLM, Sparovek G. Köppen’s climate classification map for Brazil. Meteorologische Zeitschrift, 2013; 22(6): 711-728.
- Anten NPR, Schieving F. The role of wood mass density and mechanical constraints in the economy of tree architecture. The American Naturalist, 2010; 175(2): 250-260.
- Atangana A, Chang S, Degrande A, Khasa D. Ecological Interactions and Productivity in Agroforestry Systems. In: Tropical Agroforestry. Springer: Dordrecht; 2014.
- Bila N, Rios RC, Angelo AC, Blum CT, Behling A. Restauração de áreas de Floresta Ombrófila Densa de Terras Baixas degradadas por pastagem de búfalos. Enciclopedia Biosfera 2021; 18(36): 313-327.
- Bishop JP. Tropical forest sheep on legume forage/fuelwood fallows. Agroforestry systems 1983; 1(2): 79-84.
- Bivand R, Altman M, Anselin L, Assunção R, Bera A, Berke O, Blanchet FG, et al. spdep: Spatial Dependence: Weighting Schemes, Statistics. R package version 1.3-10. [cited 2025 Feb 27]. Available from: Available from: https://CRAN.R-project.org/package=spdep
» https://CRAN.R-project.org/package=spdep - Brancalion PHS, Holl KD. Guidance for successful tree planting initiatives. Journal of Applied Ecology 2020; 57(12): 2349-2361.
- Brancalion PHS, Viani RAG, Strassburg BBN, Rodrigues RR. Finding the money for tropical forest restoration. Unasylva 2012; 63(239): 41-50.
- Bruno JF, Stachowicz JJ, Bertness MD. Inclusion of facilitation into ecological theory. Ecology & Evolution 2003; 18(3): 119-125.
- Burkart A. Leguminosas - Mimosoideas Parte I. Flora Ilustrada Catarinense. Trad. R. Klein. Santa Catarina: Itajaí; 1979.
- Burkhart HE, Tomé M. Modeling forest trees and stands. Springer Science & Business Media; 2012.
- Carvalho PER. Espécies arbóreas brasileiras. Brasília: Embrapa Informação Tecnológica; 2006.
- Carvalho PER. Espécies arbóreas brasileiras. Brasília: Embrapa Informação Tecnológica ; 2014.
- Chesson P. Mecanismos de manutenção da diversidade de espécies. Revisão anual de Ecologia e Sistemática 2000; 31(1): 343-366.
- Chave J, Andalo C, Brown S, Cains MA, Chambers JQ, Eamus D, et al. Tree allometry and improved estimation of carbon stocks and balance in tropical forests. Oecologia 2005; 145: 87-99.
- Chave J, Réjou‐Méchain M, Búrquez A, Chidumayo E, Colgan MS, Delitti WBC, et al. Improved allometric models to estimate the aboveground biomass of tropical trees. Global Change Biology: 2014; 20(10): 3177-3190.
- Cheung KC, Marques MCM, Liebsch D. Relação entre a presença de vegetação herbácea e a regeneração natural de espécies lenhosas em pastagens abandonadas na Floresta Ombrófila Densa do Sul do Brasil. Acta Botanica Brasilica: 2009; 23: 1048-1056.
- Cook-Patton SC, Leavitt SM, Gibbs D, Harris NL, Lister K, Anderson-Teixeira KJ, Brancalion PHS, et al. Mapping carbon accumulation potential from global natural forest regrowth. Nature 2020; 585: 545-550.
- De Lima MS, Araujo MM, Aimi SC, de Oliveira VV, Berghetti ÁL, Nascimento NF, Tarouco CP. Uso de atributos fisiológicos para selecionar espécies florestais nativas para restauração florestal no bioma Mata Atlântica Sul, Brasil. Ecologia e Manejo Florestal: 2021; 501: 119-659.
- De Lima RAF, Souza VC, de Siqueira MF, Steege H. Defining endemism levels for biodiversity conservation: tree species in the Atlantic Forest hotspot. Biological Conservation 2020; 252:108825.
- De Lima RB, Rutishauser E, Da Silva JAA, Guedes MC, Herault B, De Oliveira CP, et al. Accurate Estimation of Commercial Volume in Tropical Forests, Forest Science 2021; 67(1):14-21.
- Drobyshev I, Linderson H, Sonesson K. Relationship between crown condition and tree diameter growth in Southern Swedish oaks. Environ Monit Assess: 2007; 128: 61-73.
- Empresa Brasileira de Pesquisa Agropecuária - Embrapa Solos. Sistema Brasileiro de Classificação de Solos. 5. ed. Brasília, DF: Embrapa; 2018 [cited 2022 jun. 14]. Available from: Available from: https://www.embrapa.br/solos/sibcs
» https://www.embrapa.br/solos/sibcs - Elias TS. Mimosoideae. In: Polhil RM, Raven PH, editor. Advances in Legume Systematics. Kew: Inglaterra. Royal Botanic Gardens: 1981.
- Engel VL, Gandara FB. Restauração ecológica de ecossistemas naturais. Botucatu: Editora Fepaf; 2003.
- Fernandes JM, Almeida AAD, Cruz KR, Lopes CRAS. Taxonomia de Inga obtusata (Leguminosae): uma espécie quase desconhecida nos estados de Mato Grosso e Rondônia, Brasil. Scientific Electronic Archives 2022; 15(2): 35-40. https://doi.org/10.36560/15220221504
» https://doi.org/10.36560/15220221504 - Ferretti AR, De Britez RM. Restauração ecológica, sequestro de carbono e conservação da biodiversidade: a experiência da Sociedade de Pesquisa em Vida Selvagem e Educação Ambiental (SPVS) na Mata Atlântica do Sul do Brasil. Journal for Nature Conservation 2006: 14(3-4), 249-259.
- Fonseca Lima PA, Gatto A, De Albuquerque LA, Vitoria Malaquias J, De Gois Aquino F. Crescimento de mudas de espécies nativas na restauração ecológica de matas ripárias. Neotropical Biology & Conservation 2016: 11(2), 72-79.
- Fundação SOS Mata Atlântica, Instituto Nacional de Pesquisa Espaciais INPE. Atlas da evolução dos remanescentes florestais da Mata Atlântica e ecossistemas associados no período 2022-2023.[cited 2023] Available from: Available from: https://www.sosma.org.br/sobre/relatorios-e-balancos
» https://www.sosma.org.br/sobre/relatorios-e-balancos - Fundação SOS Mata Atlântica; Instituto Nacional de Pesquisas Espaciais INPE. Atlas da evolução dos remanescentes florestais da Mata Atlântica e ecossistemas associados no período 2019-2020. [cited 2010]. Available from: Available from: http://mapas.sosma.org.br/site_media/download/atlas-relatorio2008-2010parcial.pdf
» http://mapas.sosma.org.br/site_media/download/atlas-relatorio2008-2010parcial.pdf - Garcia FCP. Relações sistemáticas e fitogeografia do gênero Inga Miller (Leguminosae, Mimosoideae, Ingeae) nas florestas da costa sul e sudeste do Brasil. Tese de Doutorado, São Paulo: Universidade Estadual Paulista; 1998.
- Garcia FCP, Fernandes JM. Inga in: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro; 2015 [cited 2025 Jul 22]. Available from: Available from: http://floradobrasil2015.jbrj.gov.br/FB23000
» http://floradobrasil2015.jbrj.gov.br/FB23000 - Garcia FCP, Bonadeu F. Inga in Flora do Brasil. Jardim Botânico do Rio de Janeiro; 2020 [cited 2025 Jul 22]. Available from: Available from: https://floradobrasil2020.jbrj.gov.br/FB23000
» https://floradobrasil2020.jbrj.gov.br/FB23000 - Garnier E, Navas M-L, Grigulis K. Plant functional diversity: organism traits, community structure, and ecosystem properties. Oxford: Oxford University Press; 2016.
- Hardy NG, Kuebbing SE, Duguid MC, Ashton MS, Sheban KC, Inman SE, et al. Non‐native invasive plants in tropical dry forests: a global review of presence, impacts, and management. Restoration Ecology 2025; 33(1):14288.
- McMahon T. Size and shape in biology. Science 1973; 179(4079): 1201-1204.
- Moorcroft PR, Hurtt GC, Pacala SW. A method for scaling vegetation dynamics: the ecosystem demography model. Ecological Monographs 2001: 71(4), 557-586.
- Instituto Brasileiro de Geografia e Estatística (IBGE). Brasil em Números. 2rd ed. Rio de Janeiro: Rio de Janeiro; 2012.
- Kageyama P, Gandara FB. Recuperação de áreas ciliares. In: Rodrigues RR, Leitão Filho HF, editores. Matas ciliares: Conservação e recuperação. 2.ed. São Paulo: Universidade de São Paulo, FAPESP; 2004.
- King DA, Davies SJ, Tan S, Noor NSM. The role of wood density and stem support costs in the growth and mortality of tropical trees. Journal of Ecology 2006; 94(3): 670-680.
- Koptur S. Floral and extrafloral nectars of Costa Rica Inga trees: A comparison of their constituents and composition. Biotropica 1994; 26(3): 276-284.
- Lamb D, Erskine PD, Parrotta JA. Restoration of degraded tropical forest landscapes. Science 2005; 310(5754): 1628-1632.
- Larjavaara M. A mechanistic model for the probability of tree and branch failure under wind loading [thesis]. Helsinki: University of Helsinki; 2008.
- Laughlin DC. Applying trait-based models to achieve functional targets for theory-driven ecological restoration. Ecology Letters 2014; 17(7): 771-784.
- Lim TK. Inga edulis. In Edible medicinal and non-medicinal plants. Volume 2, fruits. Dordrecht: Springer Netherlands; 2012.
- Lorenzi H. Árvores brasileiras: manual de identificação e cultivo de plantas arbóreas nativas do Brasil. 2 nd ed. Nova Odessa: Plantarum; 1998.
- Melo FPL, Pinto SRR, Brancalion PHS, Castro PS, Rodrigues RR, Aronson J, Tabarelli M. Priority setting for scaling-up tropical forest restoration projects: early lessons from the Atlantic Forest Restoration Pact. Environmental Science & Policy 2013; 33: 395-404.
- Niklas KJ, Spatz HC. Worldwide correlations of mechanical properties and green wood density. American Journal of Botany 2010; 97(10), 1587-1594.
- Ogle DH, Doll JC, Wheeler AP, Dinno A. FSA: Simple Fisheries Stock Assessment methods. R package version 0.9.6. [cited 2025 Feb 27]. Available from: Available from: https://CRAN.R-project.org/package=FSA
» https://CRAN.R-project.org/package=FSA - Ostertag R, Warman L, Cordero RA, Sachs JP, Barber AL, Litton CM, Cabral JS, Scowcroft PG. Using plant functional traits to restore Hawaiian rainforest. Journal of Applied Ecology 2015; 52(4): 805-809.
- Paciencia MLB. Diversidade de Pteridófitas em gradientes de altitude na Mata Atlântica do Estado do Paraná [thesis]. São Paulo: Universidade de São Paulo; 2008.
- Pélico Netto S, Brena DA. Inventário Florestal. Curitiba: Edição autores; 1997.
- Piratelli AJ. Comportamento alimentar de Beija-flores em flores de Inga spp. (Leguminosae, Mimosoideae) e Jacaratia spinosa (Caricaceae) em um fragmento florestal do sudeste brasileiro. IPEF 1993; 46: 47-51.
- Poorter L, Wright SJ, Paz H, Ackerly DD, Condit R, Ibarra-Manríquez G. et al. Are functional traits good predictors of demographic rates? Evidence from five neotropical forests. Ecology 2008; 89(7): 1908-1920.
- Possette RFDS, Rodrigues WA. O gênero Inga Mill. (Leguminosae-Mimosoideae) no estado do Paraná, Brasil. Acta Botanica Brasilica 2010; 24: 354-368.
- R Core Team. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna; 2023 [cited 2025 Jul 22]. Available from: Available from: http://www.R-project.org/
» http://www.R-project.org/ - Ragusa-Netto J, Fecchio A. Plant food resources and the diet of a parrot community in a gallery forest of the Southern Pantanal (Brazil). Brazilian Journal of Biology 2006; 66(4): 1021-1032.
- Rezende CL, Scarano FR, Assad ED, Joly CA, Metzger JP, Strassburg BBN et al. From hotspot to hopespot: An opportunity for the Brazilian Atlantic Forest. Perspectives in Ecology and Conservation 2018; 16(4): 208-214.
- Rodrigues RR, Lima RAF, Gandolfi S, Nave AG. On the restoration of high diversity forests: 30 years of experience in the Brazilian Atlantic Forest. Biological Conservation : 2009; 142(6): 1242-1251.
- Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hirota MM. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 2009; 142(6):1141-1153.
- Rodrigues RR, Brancalion PHS, Isernhagen I. Pacto pela Restauração da Mata Atlântica: referencial dos conceitos e ações de restauração florestal. São Paulo LERF/ESALQ : Instituto BioAtlântica; 2009.
- Rodrigues RR, Gandolfi S. Conceitos, tendências e ações para recuperação de florestas ciliares. In: Rodrigues RR, Leitão-Filho HF, editor. Matas ciliares: conservação e recuperação. São Paulo: EDUSP; 2004.
- Santos BA, Peres CA, Oliveira MA, Grillo A, Alves-Costa CP, Tabarelli M. Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil. Biological Conservation 2008; 141(1): 249-260.
- Scarano FR. Structure, function and floristic relationships of plant communities in stressful habitats marginal to the Brazilian Atlantic rainforest. Annals of Botany 2002; 90(4): 517-524.
- Schaffer LH, Mattar EA, Nakajima NY, Silva SA, Borges RA, Borges AVP et al. Comportamento de seis espécies arbóreas na recuperação de áreas degradadas por pastagens em relevo de planície no litoral do Paraná, Brasil. Pesquisa Florestal Brasileira 2020; 40: 2-9.
- Schneider CR, Mastella ADF, Mattar EA, Angelo AC, Behling A, Rios RC, et al. Different Species Proportions Influence Silvicultural Heterogeneity of Trees in a Restoration of a Ombrophilous Dense Forest in Lowlands. Forests 2024; 15(3): 447.
- SPVS - Sociedade de Pesquisa em Vida Selvagem e Educação Ambiental (1rd ed). Reservas Naturais da SPVS - 20 anos de história. Curitiba: InVerso; 2020.
- Tabarelli M, Aguiar AV, Ribeiro MC, Metzger JP, Peres CA. Prospects for biodiversity conservation in the Atlantic Forest: lessons from aging human-modified landscapes. Biological Conservation 2010; 143(10): 2328-2340.
- Tilman D, Knops J, Wedin D, Reich P, Ritchie M, Siemann E. The influence of functional diversity and composition on ecosystem processes. Science 1997; 277(5330): 1300-1302.
- Vasconcelos GL. A Tribo Ingeae Benth. (Mimosoideae, Leguminosae) no Estado da Paraíba - Brasil [Dissertation]. Minas Gerais: Programa de Pós-Graduação em Botânica-UFV; 2014.
- Vieira M, Da Mata Mendonça YC. Desmatamento da Mata Atlântica paranaense: análise espacial para o período 2014 e 2019. Revista Catarinense de Economia 2021; 5(1): 46-57.
- Wickham H. Data analysis. In: ggplot2: elegant graphics for data analysis. Springer International 2016;
- Wickham H, François R, Henry L, Müller K, Vaughan D. dplyr: A Grammar of Data Manipulation. R package version 1.1.1. [cited 2025 Jul 22]. Available from: Available from: https://CRAN.R-project.org/package=dplyr
» https://CRAN.R-project.org/package=dplyr - Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, et al. The worldwide leaf economics spectrum. Nature 2004; 428(6985): 821827.
- Zuur AF, Ieno EN, Elphick CS. A protocol for data exploration to avoid common statistical problems. Methods in Ecology and Evolution 2010; 1:3-14.
Submitted date:
08/04/2025
Accepted date:
06/10/2026
