BEYOND SPECIES RICHNESS: QUANTIFYING FUNCTIONAL BIODIVERSITY THROUGH MATHEMATICAL ECOLOGY
Abstract
Biodiversity has traditionally been assessed through species richness, yet this approach often fails to capture the functional roles that determine ecosystem processes and resilience. Increasing ecological evidence indicates that ecosystems with similar species counts may differ substantially in functional composition, leading to divergent ecological outcomes. This study aims to develop a mathematical ecology framework that quantifies functional biodiversity by integrating trait-based analysis with nonlinear modeling. The research employs a quantitative design combining secondary ecological datasets, multidimensional trait space construction, and computational modeling to evaluate relationships between functional diversity and ecosystem performance. Results demonstrate that functional richness, evenness, and divergence significantly predict ecosystem productivity and stability, while species richness shows limited explanatory power. Nonlinear analysis reveals threshold effects and complex interactions, indicating that functional trait composition governs ecosystem responses to environmental change. Functional diversity also shapes network structure, enhancing system resilience through redundancy and complementarity among traits. The study concludes that functional biodiversity provides a more comprehensive and predictive measure of ecological complexity than species richness alone. Integration of mathematical ecology with trait-based approaches offers a robust analytical framework for advancing biodiversity research and informing conservation strategies.
Full text article
References
Berman, L. M., Schneider, F. D., Pavlick, R. P., Stenglein, J., Bemowski, R., Dean, M., & Townsend, P. A. (2024). Fractional Richness: An index for camera trap networks. Ecological Indicators, 166, 112266. https://doi.org/https://doi.org/10.1016/j.ecolind.2024.112266
Coro, G., Bove, P., Baneschi, I., Bertini, A., Calvisi, L., & Provenzale, A. (2024). Climate change effects on animal presence in the Massaciuccoli Lake basin. Ecological Informatics, 81, 102644. https://doi.org/https://doi.org/10.1016/j.ecoinf.2024.102644
Deng, X., Carvajal, D. E., Urrutia-Jalabert, R., Machida, W. S., Rosen, A., Zhang-Zheng, H., Galbraith, D., Díaz, S., Malhi, Y., & Aguirre-Gutiérrez, J. (2025). Quantifying the functional composition and potential resilience hotspots across a large latitudinal and environmental gradient in South American forests. International Journal of Applied Earth Observation and Geoinformation, 142, 104704. https://doi.org/https://doi.org/10.1016/j.jag.2025.104704
Dippner, J. W., Carrera, A. F., Kröncke, I., Liskow, I., Loick–Wilde, N., & Voss, M. (2025). Impact of multiple drivers on the trophic position, functional diversity, and ecological memory of benthic macrofauna – analysis of 40 years of data using a complex model hierarchy. Progress in Oceanography, 239, 103561. https://doi.org/https://doi.org/10.1016/j.pocean.2025.103561
Dormann, C. F., & Mello, M. A. R. (2023). Why we need a Canonical Ecology Curriculum. Basic and Applied Ecology, 71, 98–109. https://doi.org/https://doi.org/10.1016/j.baae.2023.05.009
Doxa, A., Adam, C., Nagkoulis, N., Mazaris, A. D., & Katsanevakis, S. (2025). prior3D: An R package for three-dimensional conservation prioritization. Ecological Modelling, 499, 110919. https://doi.org/https://doi.org/10.1016/j.ecolmodel.2024.110919
Eisenhauer, N., Mueller, K., Ebeling, A., Gleixner, G., Huang, Y., Madaj, A.-M., Roscher, C., Weigelt, A., Bahn, M., Bonkowski, M., Brose, U., Cesarz, S., Feilhauer, H., Guimaraes-Steinicke, C., Heintz-Buschart, A., Hines, J., Lange, M., Meyer, S. T., Mohanbabu, N., … Isbell, F. (2024). The multiple-mechanisms hypothesis of biodiversity–stability relationships. Basic and Applied Ecology, 79, 153–166. https://doi.org/https://doi.org/10.1016/j.baae.2024.07.004
Fan, X., Zang, Z., Tang, J., Zhao, L., Xu, W., & Ouyang, Z. (2025). Ecological integrity assessment system for Wuyishan national park. Ecological Indicators, 178, 113910. https://doi.org/https://doi.org/10.1016/j.ecolind.2025.113910
Feng, K., Wang, Q., Tao, K., Zhang, Y., Wang, Y., Yuan, J., Li, Z., Hugueny, B., & Er?s, T. (2025). Ecological threshold responses of fish communities to lake eutrophication. Journal of Environmental Management, 384, 125599. https://doi.org/https://doi.org/10.1016/j.jenvman.2025.125599
Ferré, C., Mascetti, G., Gentili, R., Musetti, M., Agaba, S., & Comolli, R. (2025). The role of pedodiversity in shaping biodiversity: a quantitative analysis of an alpine pasture. CATENA, 261, 109511. https://doi.org/https://doi.org/10.1016/j.catena.2025.109511
Fu, L., Zhan, C., Wang, H., Song, S., Xu, P., Zhang, L., Zhao, J., & Gao, D. (2024). Functional traits mediate the elevational patterns of functional diversity and community structure of mosses in a tropical mountain area. Global Ecology and Conservation, 55, e03259. https://doi.org/https://doi.org/10.1016/j.gecco.2024.e03259
Gholizadeh, H., Rakotoarivony, M. N. A., Hassani, K., Johnson, K. G., Hamilton, R. G., Fuhlendorf, S. D., Schneider, F. D., & Bachelot, B. (2024). Advancing our understanding of plant diversity-biological invasion relationships using imaging spectroscopy. Remote Sensing of Environment, 304, 114028. https://doi.org/https://doi.org/10.1016/j.rse.2024.114028
Ghosh, M., Mondal, S., Fajriyah, R., Mondal, K. C., & Roy, A. (2025). Association of IUCN-threatened Indian mangroves: A novel data-driven rule filtering approach for restoration strategy. Ecological Informatics, 89, 103164. https://doi.org/https://doi.org/10.1016/j.ecoinf.2025.103164
Grajales Noreña, S., Bernal, G., Cardona, O. D., Rincón, D. F., & Carreño, M. L. (2024). Holistic evaluation of climate risk to prioritise adaptation measures for ecosystems. International Journal of Disaster Risk Reduction, 109, 104593. https://doi.org/https://doi.org/10.1016/j.ijdrr.2024.104593
Huang, X., Li, S., Li, C., Lai, J., Ge, S., Gerong, Q., Tang, R., Zhang, R., & Su, J. (2025). Multiple dimensions of biodiversity jointly drive community completeness in Baima Snow Mountain typical forest. Ecological Indicators, 179, 114197. https://doi.org/https://doi.org/10.1016/j.ecolind.2025.114197
Inostroza, P. A., Jessen, G. L., Li, F., Zhang, X., Brack, W., & Backhaus, T. (2025). Multi-compartment impact of micropollutants and particularly antibiotics on bacterial communities using environmental DNA at river basin-level. Environmental Pollution, 366, 125487. https://doi.org/https://doi.org/10.1016/j.envpol.2024.125487
Lan, J., Lei, X., He, X., Gao, W., & Guo, H. (2023). Multiple mechanisms drive biodiversity-ecosystem service multifunctionality but the dominant one depends on the level of multifunctionality for natural forests in northeast China. Forest Ecology and Management, 542, 121101. https://doi.org/https://doi.org/10.1016/j.foreco.2023.121101
Lei, Z., Tingting, Z., Ziyi, Z., Beibei, L., Wuhui, L., & Guobao, C. (2025). Biodiversity and spatial heterogeneity of fish communities in response to geo-environmental disturbances. Estuarine, Coastal and Shelf Science, 320, 109322. https://doi.org/https://doi.org/10.1016/j.ecss.2025.109322
Liu, X., Frey, J., Munteanu, C., Still, N., & Koch, B. (2023). Mapping tree species diversity in temperate montane forests using Sentinel-1 and Sentinel-2 imagery and topography data. Remote Sensing of Environment, 292, 113576. https://doi.org/https://doi.org/10.1016/j.rse.2023.113576
Liu, Y., Wang, J., Dai, W., Wang, Z., & Wu, Z. (2024). Mapping terrestrial mammals in China based on species richness of functional groups. Ecological Indicators, 166, 112401. https://doi.org/https://doi.org/10.1016/j.ecolind.2024.112401
Lueders, M. B., McManamay, R. A., Bouille, D. Le, Garrett, K., Roush, L., Powers, S., Young, D., Bean, P., Mayes, K., Smith, R., & Trungale, J. F. (2025). BBEST practices: Intentional efforts to identify elusive flow-ecology thresholds. Ecological Indicators, 172, 113311. https://doi.org/https://doi.org/10.1016/j.ecolind.2025.113311
Mateus-Aguilar, B., Díaz-Salazar, A. F., Andrade-Rivas, F., Batista, N. M., Cárdenas-Navarrete, A., Arenas, A. D., Hernandez, K. V., Herrera-R, G. A., Langhans, K. E., Levey, D. R., Neill, A., Nguyen, O., Ocampo-Peñuela, N., Suárez-Castro, A. F., Zapata, F., & Echeverri, A. (2025). Assessing Biocultural Diversity Across Scales Using Ecological Indicators. Ecological Indicators, 176, 113616. https://doi.org/https://doi.org/10.1016/j.ecolind.2025.113616
Morelli, F. (2025). CAI ? Community Aggregated Index: a tool to rank and compare species assemblages combining diversity indices. Journal for Nature Conservation, 88, 127017. https://doi.org/https://doi.org/10.1016/j.jnc.2025.127017
Mou, A. T., Uddin, M. T., & Rahman, M. H. (2023). Empirical assessment of species vulnerability for biodiversity conservation: A case study on Chalan beel of Bangladesh. Heliyon, 9(4), e15251. https://doi.org/https://doi.org/10.1016/j.heliyon.2023.e15251
Nie, H., Zhao, Y., Zhu, J., Ning, A., & Zheng, W. (2024). Ecological Security Pattern Construction in Typical Oasis Area Based on Ant Colony Optimization: A Case Study in Yili River Valley, China. Ecological Indicators, 169, 112770. https://doi.org/https://doi.org/10.1016/j.ecolind.2024.112770
Nothaaß, D., & Huth, A. (2025). Community recomposition caused by species extinction in the colonization-competition trade-off model for vegetation. Ecological Modelling, 499, 110906. https://doi.org/https://doi.org/10.1016/j.ecolmodel.2024.110906
Ohlmann, M., Matias, C., Poggiato, G., Dray, S., Thuiller, W., & Miele, V. (2023). Quantifying the overall effect of biotic interactions on species distributions along environmental gradients. Ecological Modelling, 483, 110424. https://doi.org/https://doi.org/10.1016/j.ecolmodel.2023.110424
Perdomo-González, A., Pérez-Reverón, R., Goberna, M., López, H., Arribas, P., Reyes-Betancort, J. A., Andújar, C., & Díaz-Peña, F. J. (2025). The hidden shift: The role of exotic plantations in modulating soil arthropod communities in an arid island. European Journal of Soil Biology, 125, 103724. https://doi.org/https://doi.org/10.1016/j.ejsobi.2025.103724
Pommerening, A., & Särkkä, A. (2025). Towards a generalisation of the spatial species mingling diversity index. Ecological Indicators, 181, 114429. https://doi.org/https://doi.org/10.1016/j.ecolind.2025.114429
Pompeo, P. N., Oliveira Filho, L. C. I., Alexandre, D., Lovatel, A. C., Martins da Silva, P., Sousa, J. P., Klauberg-Filho, O., & Baretta, D. (2023). How does the subtropical landscape configuration influence the ecomorphological traits and community composition of ground-dwelling beetles in southern Brazil? Applied Soil Ecology, 189, 104949. https://doi.org/https://doi.org/10.1016/j.apsoil.2023.104949
Potapov, A., Lindo, Z., Buchkowski, R., & Geisen, S. (2023). Multiple dimensions of soil food-web research: History and prospects. European Journal of Soil Biology, 117, 103494. https://doi.org/https://doi.org/10.1016/j.ejsobi.2023.103494
Qin, T., Valente, L., & Etienne, R. S. (2025). Impact of evolutionary relatedness on species diversification and tree shape. Journal of Theoretical Biology, 598, 111992. https://doi.org/https://doi.org/10.1016/j.jtbi.2024.111992
Rawat, Y. S., & Tekleyohannes, A. T. (2025). Ecological assessment and sustainable utilization of plant resources in the periphery of George, Garden route area, South Africa. Journal of Environmental Management, 380, 124828. https://doi.org/https://doi.org/10.1016/j.jenvman.2025.124828
Rudaya, N., Kuzmina, O., Frolova, L., Nazarova, L., Nigmatullin, N., Syrykh, L., Vnukovskaya, Y., Cao, X., Zhilich, S., Novikov, V., Karachurina, S., & Darin, A. (2025). Holocene environments and the forest dynamics enigma in the arid Altai highlands. Quaternary Science Reviews, 367, 109548. https://doi.org/https://doi.org/10.1016/j.quascirev.2025.109548
Sedmák, R., Scheer, ?., Kúdela, P., Vencúrik, J., Modranský, J., Daniš, D., & Fabrika, M. (2025). Multicriteria optimization of ecosystem services as a base for participative forest management promoting recreation near tourist centers and cities. Trees, Forests and People, 21, 100905. https://doi.org/https://doi.org/10.1016/j.tfp.2025.100905
Selvan, S. U., Windorfer, L., Saroglou, S. T., Calbi, M., Culshaw, V., Roccotiello, E., Grobman, J. Y., & Barath, S. (2025). Multispecies design decision-making: A hybrid computational model for ecological building envelope optimization and evaluation. Journal of Building Engineering, 108, 112565. https://doi.org/https://doi.org/10.1016/j.jobe.2025.112565
Shen, X., Rezaei, T., Kachenchart, B., Tanhan, P., & Chaiyarat, R. (2024). Optimal region connection: Establishing effective ecological corridors for biodiversity conservation in Yunnan Province, China. Ecological Indicators, 169, 112918. https://doi.org/https://doi.org/10.1016/j.ecolind.2024.112918
Spencer, K. L., Deere, N. J., Aini, M., Avriandy, R., Campbell-Smith, G., Cheyne, S. M., Gaveau, D. L. A., Humle, T., Hutabarat, J., Loken, B., Macdonald, D. W., Marshall, A. J., Morgans, C., Rayadin, Y., Sanchez, K. L., Spehar, S., Suanto, Sugardjito, J., Wittmer, H. U., … Struebig, M. J. (2023). Implications of large-scale infrastructure development for biodiversity in Indonesian Borneo. Science of The Total Environment, 866, 161075. https://doi.org/https://doi.org/10.1016/j.scitotenv.2022.161075
Steadman, H. (2025). Resigned reductionism: Reconceptualising digital imaginaries of automated natural capital. Geoforum, 162, 104292. https://doi.org/https://doi.org/10.1016/j.geoforum.2025.104292
Vu Ho, K., Kröel-Dulay, G., Tölgyesi, C., Bátori, Z., Tanács, E., Kertész, M., Török, P., & Erd?s, L. (2023). Non-native tree plantations are weak substitutes for near-natural forests regarding plant diversity and ecological value. Forest Ecology and Management, 531, 120789. https://doi.org/https://doi.org/10.1016/j.foreco.2023.120789
Wan, D., Yang, J., Liu, X., Liu, Q., Chen, B., Chen, Y., Zhao, D., & Liu, J. (2025). Identifying freshwater wetland suitable habitat through a hydrological-biological connectivity framework: A case study of Naoli River wetlands, China. Ecological Indicators, 170, 112993. https://doi.org/https://doi.org/10.1016/j.ecolind.2024.112993
Wiebe, R. A., Wilcove, D. S., & Tarnita, C. E. (2025). Trajectories of biodiversity loss and extinction from trade globalization. Current Biology, 35(17), 4096-4105.e4. https://doi.org/https://doi.org/10.1016/j.cub.2025.07.030
Yang, X., Lei, S., Xu, J., Zhao, Y., Tian, Y., & Guo, Y. (2025). Mapping alpha diversity of plant species using scale effects of remote sensing. Ecological Informatics, 86, 102993. https://doi.org/https://doi.org/10.1016/j.ecoinf.2025.102993
Zhang, F., Xie, C., Liu, H., Zhou, T., Shan, B., & Chen, J. (2024). Ecological conservation redline delineation in flood detention areas: A case study in Anxiang County, Yangtze River Basin, China. Journal of Cleaner Production, 462, 142662. https://doi.org/https://doi.org/10.1016/j.jclepro.2024.142662
Zhang, L., Zhao, L., Lan, S., Chen, L., & Han, G. (2025). Genotypic richness of Phragmites australis negatively impacts ecosystem multifunctionality in the coastal wetland of Yellow River Delta, China. Global Ecology and Conservation, 60, e03609. https://doi.org/https://doi.org/10.1016/j.gecco.2025.e03609
Zhang, Y., Zhu, X., & Guo, Z. (2025). How do urban green space designs shape avian communities: Optimizing distribution, size and shape. Journal of Environmental Management, 389, 126131. https://doi.org/https://doi.org/10.1016/j.jenvman.2025.126131
Zhou, T., Liang, G., Reich, P. B., Delgado-Baquerizo, M., Wang, C., & Zhou, Z. (2024). Promoting effect of plant diversity on soil microbial functionality is amplified over time. One Earth, 7(12), 2139–2148. https://doi.org/https://doi.org/10.1016/j.oneear.2024.10.004
Authors
Copyright (c) 2026 Yang Xiang, Kaito Tanaka, Lena Hoffmann
This work is licensed under a Creative Commons Attribution-ShareAlike 4.0 International License.
