Environmental factors shaping structure and their limited influence on floristic composition in tropical montane forests of southern Ecuador
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Resumen
Tropical montane forests harbor high biodiversity and develop under heterogeneous topographic, climatic, and soil conditions. However, the influence of these environmental factors on their ecological dynamics remains poorly understood. This study was conducted to provide baseline information on the mechanisms underlying biodiversity in this complex mountain landscape. Specifically, we aimed to: (i) determine the floristic composition and forest structure of these ecosystems, and (ii) evaluate the effect of environmental gradients on species floristic composition and structural attributes in southern Ecuador. Across six localities, the tree community was sampled (composition, basal area, density, diameter, and maximum height) and edaphic, climatic, and topographic properties were characterized at the subplot level. Four discrete floristic groups and marked structural differences were identified among the evaluated forests. Environmental factors (mainly elevation and pH) explained a minor fraction of the floristic variation. In contrast, slope, elevation (associated with precipitation, temperature, and evapotranspiration gradients), and soil properties (depth, CEC, texture, and acidity) significantly modulated forest structure. We conclude that environmental factors determine density, biomass, and canopy height in these ecosystems. However, their weak influence on floristic composition suggests that community assembly is primarily mediated by biotic interactions or stochastic processes.
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Esta obra está bajo una licencia internacional Creative Commons Atribución-NoComercial 4.0.
Referencias
Aiba, S. I., Takyu, M., & Kitayama, K. (2005). Dynamics, productivity and species richness of tropical rainforests along elevational and edaphic gradients on Mount Kinabalu, Borneo. Forest Ecology and Management, 220, 279–286. https://doi.org/10.1007/s11284-005-0043-z
Aiba, S. I., & Kitayama, K. (1999). Structure, composition and species diversity in an altitude–substrate matrix of rain forest tree communities on Mount Kinabalu, Borneo. Plant Ecology, 140, 139–157. https://doi.org/10.1023/A:1009710618040
Armenteras, D., Espelta, J. M., Rodríguez, N., & Retana, J. (2017). Deforestation dynamics and drivers in different forest types in Latin America: Three decades of studies (1980–2010). Global Environmental Change, 46, 139–147. https://doi.org/10.1016/j.gloenvcha.2017.09.002
Báez, S., Malizia, A., Carilla, J., Blundo, C., Aguilar, M., Aguirre, N., Farfán-Ríos, W., et al. (2015). Large-scale patterns of turnover and basal area change in Andean forests. PLOS ONE, 10(5), Article e0126594. https://doi.org/10.1371/journal.pone.0126594
Beck, E., & Richter, M. (2008). Ecological aspects of a biodiversity hotspot in the Andes of southern Ecuador. In S. R. Gradstein, J. Homeier, & D. Gansert (Eds.), The tropical mountain forest: Patterns and processes in a biodiversity hotspot (pp. 195–217). Universitätsverlag Göttingen. https://doi.org/10.17875/gup2008-702
Becking, M. (2004). Sistema microregional de conservación Podocarpus: Tejiendo (micro) corredores de conservación hacia la cogestión de una Reserva de Biosfera Cóndor-Podocarpus. Programa Podocarpus. Loja, Ecuador p 35-36.
Brunner, I., & Sperisen, C. (2013). Aluminum exclusion and aluminum tolerance in woody plants. Frontiers in Plant Science, 4, Article 172. https://doi.org/10.3389/fpls.2013.00172
Chase, J. M., & Myers, J. A. (2011). Disentangling the importance of ecological niches from stochastic processes across scales. Philosophical Transactions of the Royal Society B: Biological Sciences, 366(1576), 2351–2363. https://doi.org/10.1098/rstb.2011.0063
Chave, J. (2004). Neutral theory and community ecology. Ecology Letters, 7(3), 241–253. https://doi.org/10.1111/j.1461-0248.2003.00566.x
Condit, R. (1998). Tropical forest census plots: Methods and results from Barro Colorado Island, Panama and a comparison with other plots. Springer-Verlag. https://doi.org/10.1007/978-3-662-03664-8
Cuesta, F., Peralvo, M., & Valarezo, N. (2009). Los bosques montanos de los Andes tropicales: Una evaluación regional de su estado de conservación y de su vulnerabilidad a efectos del cambio climático. Programa Regional ECOBONA–Intercooperation. https://www.fao.org/fileadmin/user_upload/mountain_partnership/docs/los_bosques_montanos_de_los_andes_tropicales.pdf
Culmsee, H., & Leuschner, C. (2013). Consistent patterns of elevational change in tree taxonomic and phylogenetic diversity across Malesian mountain forests. Journal of Biogeography, 40(10), 1997–2010. https://doi.org/10.1111/jbi.12138
Culmsee, H., Pitopang, R., Mangopo, H., & Sabir, S. (2011). Tree diversity and phytogeographical patterns of tropical high mountain rain forests in Central Sulawesi, Indonesia. Biodiversity and Conservation, 20, 1103–1123. https://doi.org/10.1007/s10531-011-0005-2
Duivenvoorden, J. F. (1995). Tree species composition and rain forest–environment relationships in the middle Caquetá area, Colombia, NW Amazonia. Vegetation, 120(2), 91–113. https://doi.org/10.1007/BF00034341
Dufrêne, M., & Legendre, P. (1997). Species assemblages and indicator species: The need for a flexible asymmetrical approach. Ecological Monographs, 67(3), 345–366. https://doi.org/10.1890/0012-9615(1997)067[0345:SAAISN]2.0.CO;2
Fine, P. V. A., Daly, D. C., Villa Muñoz, G., Mesones, I., & Cameron, K. M. (2005). The contribution of edaphic heterogeneity to the evolution and diversity of Burseraceae trees in the western Amazon. Evolution, 59(7), 1464–1478. https://doi.org/10.1111/j.0014-3820.2005.tb01796.x
Food and Agriculture Organization of the United Nations, & United Nations Environment Programme. (2020). The state of the world’s forests 2020: Forests, biodiversity and people. Food and Agriculture Organization of the United Nations. https://doi.org/10.4060/ca8642en
Gentry, A. H. (1988). Changes in plant community diversity and floristic composition on environmental and geographic gradients. Annals of the Missouri Botanical Garden, 75(1), 1–34. https://doi.org/10.2307/2399464
Graefe, S., Hertel, D., & Leuschner, C. (2008). Fine root characteristics of an Ecuadorian tropical montane forest along an altitudinal gradient. European Journal of Forest Research, 127(2), 97–107. https://doi.org/10.1007/s10342-007-0186-1
Grubb, P. J., & Whitmore, T. C. (1966). A comparison of montane and lowland rain forest in Ecuador: II. The climate and its effects on the distribution and physiognomy of the forests. The Journal of Ecology, 54(2), 303–333. https://doi.org/10.2307/2258396
Hamilton, L. S. (1995). Mountain forest conservation and research: A synopsis. Mountain Research and Development, 15(3), 259–266. https://doi.org/10.2307/3673899
Homeier, J., Breckle, S. W., Günter, S., Rollenbeck, R. T., & Leuschner, C. (2010). Tree diversity, forest structure and productivity along altitudinal and topographical gradients in a species-rich Ecuadorian montane rain forest. Biotropica, 42(2), 140–148. https://doi.org/10.1111/j.1744-7429.2009.00562.x
Jørgensen, P. M., Ulloa Ulloa, C., León, B., León-Yánez, S., Beck, S. G., Nee, M., Zarucchi, J. L., Celis, M., Bernal, R., & Gradstein, S. R. (2011). Regional patterns of vascular plant diversity and endemism. In S. K. Herzog, R. Martínez, P. M. Jørgensen, & H. Tiessen (Eds.), Climate change and biodiversity in the tropical Andes (pp. 192–203). Inter-American Institute for Global Change Research (IAI) & Scientific Committee on Problems of the Environment (SCOPE). https://doi.org/10.13140/2.1.3718.4969
Kappelle, M., & Brown, A. D. (Eds.). (2001). Bosques nublados del neotrópico. Instituto Nacional de Biodiversidad (INBio).
Keddy, P. A. (1992). Assembly and response rules: Two goals for predictive community ecology. Journal of Vegetation Science, 3(2), 157–164. https://doi.org/10.2307/3235676
Kochian, L. V., Piñeros, M. A., & Hoekenga, O. A. (2005). The physiology, genetics and molecular biology of plant aluminum resistance and toxicity. Plant and Soil, 274(1-2), 175–195. https://doi.org/10.1007/s11104-004-1158-7
Körner, C. (2007). The use of altitude in ecological research. Trends in Ecology & Evolution, 22(11), 569–574. https://doi.org/10.1016/j.tree.2007.09.006
Kraft, N. J. B., Adler, P. B., Godoy, O., James, E. C., Fuller, S., & Levine, J. M. (2015). Community assembly, coexistence and the environmental filtering metaphor. Functional Ecology, 29(5), 592–599. https://doi.org/10.1111/1365-2435.12345
Ließ, M., Glaser, B., & Huwe, B. (2011). Functional soil-landscape modelling to estimate slope stability in a steep Andean mountain forest region. Geomorphology, 132(3–4), 287–299. https://doi.org/10.1016/j.geomorph.2011.05.015
MacArthur, R. H. (1965). Patterns of species diversity. Biological Reviews, 40(4), 510–533. https://doi.org/10.1111/j.1469-185X.1965.tb00815.x
Merckx, V. S. F. T., Hendriks, K. P., Beentjes, K. K., Mennes, C. B., Becking, L. E., Peijnenburg, K. T. C. A., Buang, M. M., et al. (2015). Evolution of endemism on a young tropical mountain. Nature, 524(7565), 347–350. https://doi.org/10.1038/nature14949
Myers, N., Mittermeier, R. A., Mittermeier, C. G., da Fonseca, G. A. B., & Kent, J. (2000). Biodiversity hotspots for conservation priorities. Nature, 403(6772), 853–858. https://doi.org/10.1038/35002501
Ohsawa, M., Nainggolan, P. H. J., Tanaka, N., & Anwar, C. (1985). Altitudinal zonation of forest vegetation on Mount Kerinci, Sumatra: With comparisons to zonation in the temperate region of East Asia. Journal of Tropical Ecology, 1(3), 193–216. https://doi.org/10.1017/S0266467400000359
Oksanen, J., Blanchet, F. G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P. R., O’Hara, R. B., Simpson, G. L., Solymos, P., Stevens, M. H. H., Szoecs, E., Wagner, H., Barbour, M., Bedward, M., Bolker, B., Borcard, D., Carvalho, G., Chirico, M., ... Weedon, J. (2019). vegan: Community Ecology Package (Version 2.5-6) [R package]. https://CRAN.R-project.org/package=vegan
Pitman, N. C. A., Terborgh, J., Silman, M. R., Núñez V., P., Neill, D. A., Cerón, C. E., Palacios, W. A., & Aulestia, M. (2001). Dominance and distribution of tree species in upper Amazonian terra firme forests. Ecology, 82(8), 2101–2117. https://doi.org/10.1890/0012-9658(2001)082[2101:DADOTS]2.0.CO;2
Quinn, G. P., & Keough, M. J. (2002). Experimental design and data analysis for biologists. Cambridge University Press. https://archive.org/details/experimentaldesi0000quin
Quesada, C. A., Phillips, O. L., Schwarz, M., Czimczik, C. I., Baker, T. R., Patiño, S., Fyllas, N. M., Hodnett, M. G., Herrera, R., Almeida, S., & Dávila, E. A. (2012). Basin-wide variations in Amazon forest structure and function are mediated by both soils and climate. Biogeosciences, 9(6), 2203–2246. https://doi.org/10.5194/bg-9-2203-2012
Richter, M. (2008). Tropical mountain forests: Distribution and general features. In S. R. Gradstein, J. Homeier, & D. Gansert (Eds.), The tropical mountain forest: Patterns and processes in a biodiversity hotspot (pp. 7–24). Universitätsverlag Göttingen.
Rosindell, J., Hubbell, S. P., & Etienne, R. S. (2011). The unified neutral theory of biodiversity and biogeography at 10 years. Trends in Ecology & Evolution, 26(7), 340–348. https://doi.org/10.1016/j.tree.2011.03.024
Sale, P. F. (1977). Maintenance of high diversity in coral reef fish communities. The American Naturalist, 111(978), 337–359. https://doi.org/10.1086/283164
Salinas, N., Cosio, E. G., Silman, M., Meir, P., Nottingham, A. T., Roman-Cuesta, R. M., & Malhi, Y. (2021). Editorial: Tropical montane forests in a changing environment. Frontiers in Plant Science, 12, 712748. https://doi.org/10.3389/fpls.2021.712748
Schawe, M., Gerold, G., Bach, K., & Gradstein, S. R. (2010). Hydrometeorological patterns in relation to montane forest types along an elevational gradient in the Yungas of Bolivia. In S. R. Gradstein, J. Homeier, & D. Gansert (Eds.), The tropical mountain forest: Patterns and processes in a biodiversity hotspot (pp. 199–207). Universitätsverlag Göttingen.
Tanner, E. V. J., Vitousek, P. M., & Cuevas, E. (1998). Experimental investigation of nutrient limitation of forest growth on wet tropical mountains. Ecology, 79(1), 10–22. https://doi.org/10.1890/0012-9658(1998)079[0010:EIONLO]2.0.CO;2
The Angiosperm Phylogeny Group. (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), 1–20. https://doi.org/10.1111/boj.12385
Tuomisto, H., Ruokolainen, K., & Yli-Halla, M. 2003. Dispersal, environment, and floristic variation of western Amazonian forests. Science, 299(5604), 241–244. https://doi.org/10.1126/science.1078466
Walsh, C., & Mac Nally, R. (2013). hier.part: Hierarchical partitioning (Version 1.0-4) [R package]. R Foundation for Statistical Computing. https://CRAN.R-project.org/package=hier.part
Whittaker, R. H. (1960). Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs, 30(3), 279–338. https://doi.org/10.2307/1943563
Yaguana Puglla, C. A. (2020). Florestas tropicais pluviais montanas do sul do Equador: Fatores condicionantes da estrutura, composição e dinâmica [Doctoral dissertation, Universidade Estadual Paulista]. Repositório Institucional UNESP. http://hdl.handle.net/11449/202357
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