Abstract (en):
The distribution of vascular epiphytes on host trees has been linked to deterministic factors associated with host characteristics and microclimatic variation. To evaluate the taxonomic and functional spatial structure of vascular epiphytes in a lower montane cloud forest of the Colombian Andes, epiphytes were sampled following the SVERA protocol, and their position was recorded using Johansson’s zonation scheme. Epiphytes showed higher abundance below the first branching zones (Z1 and Z2), which also exhibited the highest functional richness, encompassing most of the functional space on host trees. In contrast, specialized nutrient uptake strategies and CAM metabolism were more frequent in upper zones, suggesting environmental filtering along the host gradient. These results demonstrate clear taxonomic and functional spatial structuring of vascular epiphyte assemblages in this lower montane cloud forest.
Abstract (es):
La distribución de las epífitas vasculares en los árboles hospederos se ha relacionado con factores determinísticos asociados a las características del hospedero y a la variación microclimática. Para evaluar la estructura espacial taxonómica y funcional de las epífitas vasculares en un bosque nublado montano bajo de los Andes colombianos, las epífitas vasculares se muestrearon siguiendo el protocolo SVERA y su posición se registró usando el esquema de zonación de Johansson. Las epífitas mostraron mayor abundancia por debajo de la primera ramificación (Z1 y Z2), zonas que también presentaron la mayor riqueza funcional, lo que abarcó la mayor parte del espacio funcional disponible en los hospederos. En contraste, las estrategias especializadas de toma de nutrientes y el metabolismo CAM fueron más frecuentes en las zonas superiores, lo que sugiere la presencia de filtros ambientales a lo largo del gradiente del hospedero. Estos resultados demuestran una clara estructuración espacial taxonómica y funcional de los ensamblajes de epífitas vasculares en este bosque nublado montano bajo.
Keywords:
Andean biodiversity, canopy stratification, microclimatic gradients, trait-based ecology, community assembly processes (en)
biodiversidad andina, estratificación del dosel, gradientes microclimáticos, ecología basada en rasgos, procesos de ensamblaje de comunidades (es)
biodiversidad andina, estratificación del dosel, gradientes microclimáticos, ecología basada en rasgos, procesos de ensamblaje de comunidades (es)
References
Acebey, A., & Krömer, T. (2001). Diversidad y distribución vertical de epífitas en los alrededores del campamento río Eslabón y de la laguna. Revista de la Sociedad Boliviana de Botánica, 3(1–2), 104–123.
Bellingham, P. J., & Sparrow, A. D. (2000). Resprouting as a life history strategy in woody plant communities. Oikos, 89(2), 409–416. https://doi.org/10.1034/j.1600-0706.2000.890224.x
Bezzalla, A., Boudjabi, S., & Chenchouni, H. (2018). Seedlings of Argan (Argania spinosa) from different geographical provenances reveal variable morphological growth responses to progressive drought stress under nursery conditions. Agroforestry Systems, 92, 1201–1211. https://doi.org/10.1007/s10457-016-0057-2
Cardelús, C. L., & Chazdon, R. L. (2005). Inner-crown microenvironments of two emergent tree species in a lowland wet forest1. Biotropica, 37(2), 238–244. https://doi.org/10.1111/j.1744-7429.2005.00032.x
da Silva, J. S., Piedade, M. T. F., Klein, V. P., Durgante, F. M., Wittmann, F., & Quaresma, A. C. (2024). Large diameters and tree bark physical attributes drive vascular epiphyte-phorophyte relationships in Amazonian black-water floodplain forest. Plant Ecology, 225(2), 163-173. https://doi.org/10.1007/s11258-023-01387-1
de Cáceres, M. (2023). indicspecies: Relationship between species and groups of sites (Version 1.7.14) [Computer software]. R Foundation for Statistical Computing. https://CRAN.R-project.org/package=indicspecies
de La Rosa‐Manzano, E., Andrade, J. L., Zotz, G., & Reyes‐García, C. (2014). Epiphytic orchids in tropical dry forests of Yucatan, Mexico – Species occurrence, abundance and correlations with host tree characteristics and environmental conditions. Flora, 209(2), 100–109. https://doi.org/10.1016/j.flora.2013.12.002
Francisco, T. M., Couto, D. R., Garbin, M. L., Misaki, F., & Ruiz‐Miranda, C. R. (2021). Role of spatial and environmental factors in structuring vascular epiphyte communities in two neotropical ecosystems. Perspectives in Plant Ecology, Evolution and Systematics, 51, 125621. https://doi.org/10.1016/j.ppees.2021.125621
Garnier, É., Navas, M., & Grigulis, K. (2016). Plant Functional Diversity: Organism traits, community structure, and ecosystem properties. Oxford University Press.
Gotelli, N. J., & McCabe, D. J. (2002). Species co‐occurrence: a meta‐analysis of JM Diamond's assembly rules model. Ecology, 83(8), 2091–2096. https://doi.org/10.1890/0012-9658(2002)083[2091:SCOAMA]2.0.CO;2
Guzmán‐Jacob, V., Guerrero‐Ramírez, N. R., Craven, D., Paterno, G. B., Taylor, A., Krömer, T., Wanek, W., Zotz, G., & Kreft, H. (2022). Broad and small‐scale environmental gradients drive variation in chemical, but not morphological, leaf traits of vascular epiphytes. Functional Ecology, 36(8), 1858–1872. https://doi.org/10.1111/1365-2435.14084
Hietz, P., & Hietz‐Seifert, U. (1995). Structure and ecology of epiphyte communities of a cloud forest in central Veracruz, Mexico. Journal of Vegetation Science, 6(5), 719–728. https://doi.org/10.2307/3236443
Hietz, P., & Briones, O. (1998). Correlation between water relations and within-canopy distribution of epiphytic ferns in a Mexican cloud forest. Oecologia, 114(3), 305–316. https://doi.org/10.1007/s004420050452
Janzen, T., Zotz, G., & Etienne, R. S. (2020). Community structure of vascular epiphytes: a neutral perspective. Oikos, 129(6), 853–867. https://doi.org/10.1111/oik.06537
Johansson, D. R. (1974). Ecology of vascular epiphytes in West African rain forest [Doctoral dissertation, Uppsala University].
Kembel, S. W., Cowan, P. D., Helmus, M. R., Cornwell, W. K., Morlon, H., Ackerly, D. D., Blomberg, S. P., & Webb, C. O. (2010). Picante: R tools for integrating phylogenies and ecology. Bioinformatics, 26(11), 1463–1464. https://doi.org/10.1093/bioinformatics/btq166
Klimešová, J., Ottaviani, G., Charles‐Dominique, T., Campetella, G., Canullo, R., Chelli, S., Janovský, Z., Lubbe, F. C., Martínková, J., & Herben, T. (2021). Incorporating clonality into the plant ecology research agenda. Trends in Plant Science, 26(12), 1236–1247. https://doi.org/10.1016/j.tplants.2021.07.019
Krömer, T., Kessler, M., & Gradstein, S. R. (2007). Vertical stratification of vascular epiphytes in submontane and montane forest of the Bolivian Andes: the importance of the understory. Plant Ecology, 189(2), 261–278. https://doi.org/10.1007/s11258-006-9182-8
Laliberté, E., Legendre, P., & Shipley, B. (2024). FD: Measuring functional diversity from multiple traits, and other tools for functional ecology (Version 1.0-12.1) [Computer software]. R Foundation for Statistical Computing. https://CRAN.R-project.org/package=FD
Liu, F., Liu, J., & Dong, M. (2016). Ecological consequences of clonal integration in plants. Frontiers in Plant Science, 7, 770. https://doi.org/10.3389/fpls.2016.00770
Marí, M. L. G., De Toledo, J. J., Nascimento, H. E. M., & Zartman, C. E. (2016). Regional and fine scale variation of holoepiphyte community structure in central Amazonian white-sand forests. Biotropica, 48(1), 70–80. https://doi.org/10.1111/btp.12300
Mason, N. W. H., Mouillot, D., Lee, W. G., & Wilson, J. B. (2005). Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos, 111(1), 112–118. https://doi.org/10.1111/j.0030-1299.2005.13886.x
Nieder, J., Engwald, S., Klawun, M., & Barthlott, W. (2000). Spatial distribution of vascular epiphytes (including Hemiepiphytes) in a Lowland Amazonian Rain Forest (Surumoni Crane Plot) of Southern Venezuela. Biotropica, 32(3), 385–396. https://doi.org/10.1111/j.1744-7429.2000.tb00485.x
Oksanen, J., Simpson, G. L., Blanchet, F. G., Kindt, R., Legendre, P., Minchin, P. R., O'Hara, R. B., Solymos, P., Stevens, M. H. H., Szoecs, E., Wagner, H., Barbour, M., Bedward, M., Bolker, B., Borcard, D., Carvalho, G., Chirico, M., de Cáceres, M., Durand, S., ... Weedon, J. (2022). vegan: Community Ecology Package (Version 2.6-4) [Computer software]. Comprehensive R Archive Network (CRAN). https://CRAN.R-project.org/package=vegan
Pillar, V. D., Blanco, C. C., Müller, S. C., Sosinski, E. E., Joner, F., & Duarte, L. D. (2013). Functional redundancy and stability in plant communities. Journal of Vegetation Science, 24(5), 963–974.
Pérez-Harguindeguy, N., Díaz, S., Garnier, E., Lavorel, S., Poorter, H., Jaureguiberry, P., Bret-Harte, M. S., Cornwell, W. K., Craine, J. M., Gurvich, D. E., Urcelay, C., Veneklaas, E. J., Reich, P. B., Poorter, L., Wright, I. J., Ray, P., Enrico, L., Pausas, J. G., de Vos, A. C., ... Cornelissen, J. H. C. (2016). Corrigendum to: New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany, 64(8), 715. https://doi.org/10.1071/bt12225_co
Petter, G., Wagner, K., Wanek, W., Delgado, E. J. S., Zotz, G., Cabral, J. S., & Kreft, H. (2016). Functional leaf traits of vascular epiphytes: vertical trends within the forest, intra- and interspecific trait variability, and taxonomic signals. Functional Ecology, 30(2), 188–198. https://doi.org/10.1111/1365-2435.12490
Pla, L., Casanoves, F., & Di Rienzo, J. (2012). Functional diversity indices. In L. Pla, F. Casanoves, & J. Di Rienzo (Eds.), Quantifying functional biodiversity (pp. 27–51).
Quaresma, A. C., & Jardim, M. A. G. (2014). Floristic composition and spatial distribution of vascular epiphytes in the restingas of Maracanã, Brazil. Acta Botanica Brasilica, 28(1), 68–75. https://doi.org/10.1590/s0102-33062014000100007
Richards, J. H., & Damschen, E. I. (2022). Leaf economics in a three‐dimensional environment: Testing leaf trait responses in vascular epiphytes to land use, climate and tree zone. Functional Ecology, 36(3), 727–738. https://doi.org/10.1111/1365-2435.13978
Sanford, W. W. (1968). Distribution of epiphytic orchids in Semi-Deciduous tropical forest in southern Nigeria. Journal of Ecology, 56(3), 697. https://doi.org/10.2307/2258101
Stuefer, J. F. (1998). Two types of division of labour in clonal plants: benefits, costs and constraints. Perspectives in Plant Ecology, Evolution and Systematics, 1(1), 47–60. https://doi.org/10.1078/1433-8319-00051
Tewari, L. M., Tewari, G., Nailwal, T., & Pangtey, Y. P. S. (2009). Bark factors affecting the distribution of epiphytic ferns communities. Nat Sci, 7, 76–81.
Van Leerdam, A., Zagt, R., & Veneklaas, E. J. (1990). The distribution of epiphyte growth-forms in the canopy of a Colombian cloud-forest. Vegetatio, 87(1), 59–71. https://doi.org/10.1007/bf00045656
Vilela, A. E., Agüero, P. R., Ravetta, D. A., & González-Paleo, L. (2012). Long-term plasticity in growth, storage and defense allocation produces drought-tolerant juvenile shrubs of Prosopis alpataco RA Philippi (Fabaceae). Flora-Morphology, Distribution, Functional Ecology of Plants, 207(6), 436–441. https://doi.org/10.1016/j.flora.2012.02.006
Vinod, N., Slot, M., McGregor, I. R., Ordway, E. M., Smith, M. N., Taylor, T. C., Sack, L., Buckley, T. N., & Anderson-Teixeira, K. J. (2023). Thermal sensitivity across forest vertical profiles: Patterns, mechanisms, and ecological implications. New Phytologist, 237(1), 22–47. https://doi.org/10.1111/nph.18539
Vittoz, P., & Engler, R. (2007). Seed dispersal distances: a typology based on dispersal modes and plant traits. Botanica Helvetica, 117, 109–124. https://doi.org/10.1007/s00035-007-0797-8
Wagner, K., Mendieta-Leiva, G., & Zotz, G. (2015). Host specificity in vascular epiphytes: A review of methodology, empirical evidence and potential mechanisms. AoB Plants, 7, plu092. https://doi.org/10.1093/aobpla/plu092
Wang, X., Long, W., Schamp, B. S., Yang, X., Kang, Y., Xie, Z., & Xiong, M. (2016). Vascular Epiphyte Diversity Differs with Host Crown Zone and Diameter, but Not Orientation in a Tropical Cloud Forest. PLOS ONE, 11(7), e0158548. https://doi.org/10.1371/journal.pone.0158548
Werner, F. (2011). Reduced growth and survival of vascular epiphytes on isolated remnant trees in a recent tropical montane forest clear-cut. Basic and Applied Ecology, 12(2), 172–181. https://doi.org/10.1016/j.baae.2010.11.002
Wolf, J. H. D., Gradstein, S. R., & Nadkarni, N. M. (2009). A protocol for sampling vascular epiphyte richness and abundance. Journal of Tropical Ecology, 25(2), 107–121. https://doi.org/10.1017/s0266467408005786
Woods, C. L., Cardelús, C. L., & DeWalt, S. J. (2015). Microhabitat associations of vascular epiphytes in a wet tropical forest canopy. Journal of Ecology, 103(2), 421–430. https://doi.org/10.1111/1365-2745.12357
Zotz, G., & Büche, M. (2000). The epiphytic filmy ferns of a tropical lowland forest-species occurrence and habitat preferences. Ecotropica, 6, 203-206
Zotz, G., & Vollrath, B. (2003). The epiphyte vegetation of the palm Socratea exorrhiza - correlations with tree size, tree age and bryophyte cover. Journal of Tropical Ecology, 19(1), 81–90. https://doi.org/10.1017/s0266467403003092
Zotz, G. (2007). Johansson revisited: the spatial structure of epiphyte assemblages. Journal of Vegetation Science, 18(1), 123–130. https://doi.org/10.1111/j.1654-1103.2007.tb02522.x
Zotz, G., & Schultz, S. (2007). The vascular epiphytes of a lowland forest in Panama - species composition and spatial structure. Plant Ecology, 195(1), 131–141. https://doi.org/10.1007/s11258-007-9310-0
Zotz, G. (2013). ‘Hemiepiphyte’: a confusing term and its history. Annals of Botany, 111(6), 1015–1020. https://doi.org/10.1093/aob/mct085
Zotz, G. (2016). Plants on Plants - The Biology of Vascular Epiphytes. Springer.
How to Cite
Arias Martínez, L. L., González Echeverry, J. C., & Zuluaga Tróchez, A. (2026). Functional and taxonomic spatial structure of vascular epiphytes in a neotropical montane cloud forest. Biota Colombiana, 27, e1306. https://doi.org/10.21068/2539200X.1306
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