Abstract
Geometric morphometrics encompasses a set of methods for the acquisition, processing, and quantitative analysis of biological shapes. This study applied this approach to leaf blades of species from Philodendron section Philodendron (Jacq.) Schott, collected in the Amapá State Forest (FLOTA/AP), with the aim of investigating morphological patterns associated with environmental factors. Two species were selected for analysis: Philodendron atabapoense G.S.Bunting and P. ornatum Schott. The results revealed significant variation in leaf morphology, particularly in leaf base and circularity, which were strongly influenced by climatic and altitudinal variables. The findings highlight the value of geometric morphometrics as a complementary tool for species delimitation and for understanding morphological diversity within the genus Philodendron.
References
- Adams, D., Collyer, M., Kaliontzopoulou, A. & Baken, E. (2025) Geomorph: Software for geometric morphometric analyses. R package version 4.0.10. Available from: https://cran.r-project.org/package=geomorph (accessed 16 April 2026)
- Adams, D.C., Rohlf, F.J. & Slice, D.E. (2004) Geometric morphometrics: ten years of progress following the “revolution”. Italian Journal of Zoology 71 (1): 5–16. https://doi.org/10.1080/11250000409356545
- APG IV (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20. https://doi.org/10.1111/boj.12385
- Beck, H.E., Zimmermann, N.E., McVicar, T.R., Vergopolan, N., Berg, A. & Wood, E.F. (2018) Present and future Köppen-Geiger climate classification maps at 1-km resolution. Sci Data 5: 180214. https://doi.org/10.1038/sdata.2018.214
- Baken, E., Collyer, M., Kaliontzopoulou, A. & Adams, D. (2021) geomorph v4.0 and gmShiny: enhanced analytics and a new graphical interface for a comprehensive morphometric experience. Methods in Ecology and Evolution 12: 2355–2363. https://doi.org/10.1111/2041-210X.13723
- Boyce, P.B. & Croat, T.B. (2011 onwards) The Uberlist of Araceae. Totals for published and estimated numbers of species in aroid genera. Available from: http://www.aroid.org/genera/130307uberlist.pdf (accessed 24 March 2022)
- Brasil (1974) Projeto RADAM. Macapá: geologia, geomorfologia, solos, vegetação e uso potencial da terra. Rio de Janeiro, v.1.
- Calazans, L.S.B., Sakuragui, S.M. & Mayo, S.J. (2014) From open areas to forests? The evolutionary history of Philodendron subgenus Meconostigma (Araceae) using morphological data. Flora—Morphology, Distribution, Functional Ecology of Plants 209: 117–121. https://doi.org/10.1016/j.flora.2013.12.004
- Campos, M.A.A. & Uchida, T. (2002) Influência do sombreamento no crescimento de mudas de três espécies amazônicas. Pesquisa Agropecuária Brasileira 37: 281–288. https://doi.org/10.1590/S0100-204X2002000300008
- Castro, A.H.F. & Alvarenga, A.S. (2002) Influência do fotoperíodo no crescimento inicial de plantas de confrei (Symphytum officinale L.). Ciência e Agrotecnologia 26: 77–89.
- Coelho, M.A.N. (2000) Philodendron Schott (Araceae): morfologia e taxonomia das espécies da Reserva Ecológica de Macaé de Cima—Nova Friburgo, Rio de Janeiro, Brasil. Dissertação (Mestrado em Botânica)—Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro. https://doi.org/10.1590/2175-7860200051787902
- Coelho, M.A.N. (2010) A família Araceae na Reserva Natural Vale, Linhares, Espírito Santo, Brasil. Boletim do Museu de Biologia Mello Leitão 28: 41–87.
- Croat, T.B. (1997) A revision of Philodendron subgenus Philodendron (Araceae) for Mexico and Central America. Annals of the Missouri Botanical Garden 84: 311–704. https://doi.org/10.2307/2992022
- Dubreuil, V., Fante, K.P., Planchon, O. & Sant’anna-Neto, J.L. (2018) Os tipos de climas anuais no Brasil : uma aplicação da classificação de Köppen de 1961 a 2015. Journals Open Edition 2018: 37. https://doi.org/10.4000/confins.15738
- Croat, T.B. & Hannon, L.P. (2015) Revision of Chlorspatha (Araceae). Annals of the Missouri Botanical Garden 101: 1–229. https://doi.org/10.3417/2005079
- Croat, T.B. (1999) New Species of Central American Araceae. Novon 9: 491–502. https://doi.org/10.2307/3392149
- Fidalgo, O. & Bononi, V.L. (1984) Techniques for the collection, preservation, and herborization of botanical material. Instituto de Botânica, São Paulo, 62 pp.
- Flora & Funga do Brasil (2025) Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.gov.br/ (accessed 17 April 2025)
- IBGE (2016) Estimativa da População Residente Nos Municípios Brasileiros. Instituto Brasileiro de Geografia e Estatística [S.l.]
- IEF (2014) Plano de manejo da floresta estadual do Amapá. Instituto Estadual de Florestas do Amapá. Available from: https://ief.portal.ap.gov.br/dados.php?d=739&a=453 (accessed 24 March 2022)
- INPE (2023) Centro de Previsão de Tempo e Estudos Climáticos. Instituto Nacional de Pesquisas Espaciais. Available from: https://previsaonumerica.cptec.inpe.br/#WRF_cpt_07KM/Mais%20recente/empty/0/ (accessed 27 March 2023)
- Jensen, R.J. (2003) The conundrum of morphometrics. Taxon 52: 663–671. https://doi.org/10.2307/3647345
- Jolliffe, I. (2002) Principal component analysis, 2nd edn. Springer, New York.
- Keating, R.C. (2002) Acoraceae and Araceae. In: Gregory, M. & Cutler, D.F. (Eds.) Anatomy of the monocotyledons, Vol. 9. Oxford University Press, Oxford.
- Klich, M.G. (2000) Leaf variations in Elaeagnus angustifolia related to environmental heterogeneity. Environmental and Experimental Botany 44: 171–183. https://doi.org/10.1016/S0098-8472(00)00056-3
- Köppen, W. & Geiger, R. (1928) Klimate der Erde. Verlag Justus Perthes, Gotha, Alemanha (Wall-mapp150cmx200cm.).
- Lima, C.B.S. (2014) Caracterização morfométrica das populações de Melipona subnitida Ducke, 1910 (Apidae, Meliponini) residentes no limite sul da área de sua distribuição natural. Dissertação de Mestrado, Universidade Federal do Recôncavo da Bahia, Cruz das Almas, BA, 50 pp.
- Matsumoto, T.K., Onoue, M., Miyake, T., Ohnishi, K., Takazoe, K., Hirobe, M. & Miyazaki, Y. (2023) Gall midge pollination and ant-mediated fruit dispersal of Pinellia tripartita (Araceae). Plant Ecology 224: 59–72. https://doi.org/10.1007/s11258-022-01278-x
- Mayo, S.J. (1989) Observations of the gynoecial structure in Philodendron (Araceae). Botanical Journal of the Linnean Society 100: 139–172. https://doi.org/10.1111/j.1095-8339.1989.tb01714.x
- Mayo, S.J., Bogner, J. & Boyce, P.C. (1997) The genera of Araceae. 1st ed. Royal Botanical Garden, Kew, London, 370 pp.
- Olsen, A.M. & Haber, A. (2022) StereoMorph: Stereo Camera Calibration and Reconstruction. Version 1.6.5. Available from: https://CRAN.R-project.org/package=StereoMorph (accessed 27 March 2023)
- Olsen, A.M. & Westneat, M.W. (2015) StereoMorph: an R package for the collection of 3D landmarks and curves using a stereo camera set-up. Methods in Ecology and Evolution 6: 351–356. https://doi.org/10.1111/2041-210X.12326
- Pedraza, S. (2024) Leaf traits and leaf‐to‐air temperature differences in tropical plants suggest variability in thermoregulatory capacities across elevations. Biotropica 56: e13332. https://doi.org/10.1111/btp.13332
- Pontes, T.A. (2010) Diversidade de Araceae em fragmentos de Floresta Atlântica de terras baixas ao Norte do Estado de Pernambuco, Brasil. Dissertação (Mestrado)—Universidade Federal de Pernambuco, Recife.
- R Core team. (2022) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.R-project.org/ (accessed 27 March 2023)
- Raven, P.H., Evert, R.F. & Eichhorn, S.E. (1999) Biology of Plants. 6 ed. Worth Publishers, New York.
- Revelle, W. (2022) psych: Procedures for Personality and Psychological Research, Northwestern University, Evanston, Illinois, USA, Version 2.2.9. Available from: https://CRAN.R-project.org/package=psych (accessed 27 March 2023)
- Rodrigues, P., Facundes, A.S. & Pereira, L.A. (2020) Philodendron Schott (Araceae Juss.) no distrito do Carvão, Mazagão, Amapá, Brasil. Boletim do Museu Paraense Emílio Goeldi—Ciências Naturais 15: 421–431. https://doi.org/10.46357/bcnaturais.v15i2.125
- Rohlf, F.J. & Marcus, L.F. (1993) A revolution in morphometrics. Trends in Ecology and Evolution 8: 129–132. https://doi.org/10.1016/0169-5347(93)90024-J
- Rohlf, F.J. & Slice, D. (1990) Extensions of the Procrustes method for the optimal superimposition of landmarks. Systematics Zoology 39: 40–59. https://doi.org/10.2307/2992207
- Rohlf, F.J. (2015) The tps series of software. Hystrix, the Italian Journal of Mammalogy 26: 9–12. https://doi.org/10.4404/hystrix-26.1-11264
- Sakuragui, C.M., Calazans, L.S.B., de Oliveira, L.L., de Morais, É.B., Benko-Iseppon, A.M., Vasconcelos, S., Schrago, C.E.G. & Mayo, S.J. (2018) Recognition of the genus Thaumatophyllum Schott—formerly Philodendron subg. Meconostigma (Araceae)—based on molecular and morphological evidence. PhytoKeys 2: 51–71. https://doi.org/10.3897/phytokeys.98.25044
- Silva Sampaio, M. da, Carvalho Alves, M. de, Carvalho, L.G. de & Sanches, L. (2011) Uso de Sistema de Informação Geográfica para comparar a classificação climática de Koppen-Geiger e de Thornthwaite. XV Simpósio Brasileiro de Sensoriamento Remoto—SBSR. Curitiba-PR: INPE.
- Schott, H.W. (1829a) Für Liebhaber der Botanick. Wiener Zeitschrift für Kunst, Literatur, Theater und Mode 100: 828. Available from: https://anno.onb.ac.at/cgi-content/anno?aid=wzz&datum=18290820&seite=8&zoom=33 (accessed 16 April 2026)
- Schott, H.W. (1829b) Für Liebhaber der Botanick. Wiener Zeitschrift für Kunst, Literatur, Theater und Mode 94: 779–780. Available from: https://anno.onb.ac.at/cgi-content/anno?aid=wzz&datum=18290806&seite=7&zoom=33 (accessed 16 April 2026)
- Schott, H.W. (1856) Synopsis Aroidearum Complectens Enumerationem Systematicam Generum et Specierum Huju Ordinis. Typis Congregationis Mechitharisticae, Vienna, 140 pp.
- Scoles, R., Gribel, R. & Klein, G.N. (2011) Growth and survival of Brazil nuts (Bertholletia excelsa Bonpl.) in different environmental conditions in region River Trombetas, Oriximiná, Pará, Brazil. Boletim do Museu Paraense Emílio Goeldi—Ciências Naturais 6: 273–293. https://doi.org/10.46357/bcnaturais.v6i3.610
- Silva, M.F.S., Andrade, I.M. & Mayo, S.J. (2012) Geometric morphometrics of leaf blade shape in Montrichardia linifera (Araceae) populations from the Rio Parnaíba Delta, northeast Brazil. Botany Journal of the Linnean Society 170: 554–572. https://doi.org/10.1111/j.1095-8339.2012.01309.x
- Slice, D., Bookstein, F., Marcus, L. & Rohlf, F.J. (2003) A Glossary for Geometric morphometrics. Available from: http://life.bio.sunysb.edu/morph/glossary/gloss1.html (accessed 27 March 2023)
- Slice, D. (2005) Modern Morphometrics in Physical Anthropology. Springer, New York. https://doi.org/10.1007/0-387-27614-9
- Spani, F., Locato, V. & Gara, L. (2025) Unveiling Nature’s Architecture: Geometric Morphometrics as an Analytical Tool in Plant Biology. Plants 14: 808. https://doi.org/10.3390/plants14050808
- SUDAM (1984) Climatological Atlas of the Brazilian Amazon. Projeto de Hidrologia e Climatologia da Amazônia, Belém-PA, 1, 125 pp.
- Thiers, B.M. (2010 onwards) Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 7 April 2023)
- Viscosi, V. & Cardini, A. (2011) Leaf morphology, taxonomy and geometric morphometrics: a simplified protocol for beginners. PLoS ONE 6: e25630. https://doi.org/10.1371/journal.pone.0025630
- Viscosi, V., Fortini, P., Slice, D.E., Loy, A. & Blasi, C. (2012) Geometric morphometric analyses of leaf variation in four oak species of the subgenus Quercus (Fagaceae). Plant Biosystems 146: 611–621. https://doi.org/10.1080/11263504.2011.650729
- Liu, Y., Li, Y., Song, J., Zhang, R., Yan, Y., Wang, Y. & Du, F.K. (2018) Análises morfométricas geométricas do formato das folhas em duas espécies simpáticas Carvalhos chineses: Quercus dentata Thunberg e Quercus aliena Blume (Fagaceae). Anais de Ciência Florestal 75: 90. https://doi.org/10.1007/s13595-018-0770-2
- Watanabe, A. (2018) How many landmarks are enough to characterize shape and size variation? PLoS ONE 13: e0198341. https://doi.org/10.1371/journal.pone.0198341
- Wei, T. & Simko, V. (2024) corrplot: Visualization of a Correlation Matrix. R package version 0.95. Available from: https://github.com/taiyun/corrplot (accessed 27 March 2023)
- Wheeler, B. & Torchiano, M. (2016) lmPerm: Permutation Tests for Linear Models. R package version 2.1.0. Available from: https://CRAN.R-project.org/package=lmPerm (accessed 27 March 2023)
- Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. Available from: https://ggplot2.tidyverse.org (accessed 27 March 2023)
