Articles | Volume 23, issue 14
https://doi.org/10.5194/bg-23-4999-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/bg-23-4999-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Research article
| 
21 Jul 2026
Research article |
| 21 Jul 2026
| 21 Jul 2026
Vegetation patterns and competitive dynamics along elevation gradients: interactions between environmental factors and vegetation in the Central Himalayas
Prashant Paudel
Hawkesbury Institute for the Environment, Western Sydney University, Penrith, NSW, Australia
Stefan Olin
Department of Physical Geography and Ecosystem Science, Lund University, Lund, Sweden
Mark Tjoelker
Hawkesbury Institute for the Environment, Western Sydney University, Penrith, NSW, Australia
Mikael Pontarp
Department of Biology, Lund University, Lund, Sweden
Daniel Metcalfe
Department of Ecology and Environment Science, Umeå University, Umeå, Sweden
Hawkesbury Institute for the Environment, Western Sydney University, Penrith, NSW, Australia
Department of Physical Geography and Ecosystem Science, Lund University, Lund, Sweden
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Cited articles
Acharya, B. K., Chettri, B., and Vijayan, L.: Distribution pattern of trees along an elevation gradient of Eastern Himalaya, India, Acta Oecologica, 37, 329–336, https://doi.org/10.1016/j.actao.2011.03.005, 2011.
Ahmad, M., Luo, Y.-H., Rathee, S., Spicer, R. A., Zhang, J., Wambulwa, M. C., Zhu, G.-F., Cadotte, M. W., Wu, Z.-Y., Khan, S. M., Maity, D., Li, D.-Z., and Liu, J.: Multifaceted plant diversity patterns across the Himalaya: Status and outlook, Plant Divers., 47, 529–543, https://doi.org/10.1016/j.pld.2025.04.003, 2025.
Araghi, A. and Martinez, C. J.: Evaluation of CRU-JRA gridded meteorological dataset for modeling of wheat production systems in Iran, Int. J. Biometeorol., 68, 1201–1211, https://doi.org/10.1007/s00484-024-02659-9, 2024.
Asner, G. P., Anderson, C. B., Martin, R. E., Knapp, D. E., Tupayachi, R., Sinca, F., and Malhi, Y.: Landscape-scale changes in forest structure and functional traits along an Andes-to-Amazon elevation gradient, Biogeosciences, 11, 843–856, https://doi.org/10.5194/bg-11-843-2014, 2014.
Avolio, M. L., Forrestel, E. J., Chang, C. C., La Pierre, K. J., Burghardt, K. T., and Smith, M. D.: Demystifying dominant species, New Phytol., 223, 1106–1126, https://doi.org/10.1111/nph.15789, 2019.
Ayer, S., Yadav, B. K., and Bhatta, K. P.: What drives the regeneration dynamics in Central Himalayan Mountain Forests of Nepal?, Environ. Chall., 21, 101359, https://doi.org/10.1016/j.envc.2025.101359, 2025.
Bertness, M. D. and Callaway, R.: Positive interactions in communities, Trends Ecol. Evol., 9, 191–193, https://doi.org/10.1016/0169-5347(94)90088-4, 1994.
Bi, W. and Zhou, Y.: A global 0.05° dataset for gross primary production of sunlit and shaded vegetation canopies (1992–2020), DRYAD [data set], https://doi.org/10.5061/DRYAD.DFN2Z352K, 2022.
Brewer, J. S.: Disturbance-mediated competition between perennial plants along a resource supply gradient: Disturbances and competition, J. Ecol., 99, 1219–1228, https://doi.org/10.1111/j.1365-2745.2011.01846.x, 2011.
Brooker, R. W. and Kikvidze, Z.: Importance: an overlooked concept in plant interaction research, J. Ecol., 96, 703–708, https://doi.org/10.1111/j.1365-2745.2008.01373.x, 2008.
Bugmann, H., Fischlin, A., and Kienast, F.: Model convergence and state variable update in forest gap models, Ecol. Model., 89, 197–208, https://doi.org/10.1016/0304-3800(95)00135-2, 1996.
Colwell, Robert. K. and Lees, D. C.: The mid-domain effect: geometric constraints on the geography of species richness, Trends Ecol. Evol., 15, 70–76, https://doi.org/10.1016/S0169-5347(99)01767-X, 2000.
Coomes, D. A. and Allen, R. B.: Effects of size, competition and altitude on tree growth, J. Ecol., 95, 1084–1097, https://doi.org/10.1111/j.1365-2745.2007.01280.x, 2007.
De Frenne, P., Graae, B. J., Rodríguez-Sánchez, F., Kolb, A., Chabrerie, O., Decocq, G., De Kort, H., De Schrijver, A., Diekmann, M., Eriksson, O., Gruwez, R., Hermy, M., Lenoir, J., Plue, J., Coomes, D. A., and Verheyen, K.: Latitudinal gradients as natural laboratories to infer species' responses to temperature, J. Ecol., 101, 784–795, https://doi.org/10.1111/1365-2745.12074, 2013.
De Paula, M. D., Forrest, M., Langan, L., Bendix, J., Homeier, J., Velescu, A., Wilcke, W., and Hickler, T.: Nutrient cycling drives plant community trait assembly and ecosystem functioning in a tropical mountain biodiversity hotspot, New Phytol., 232, 551–566, https://doi.org/10.1111/nph.17600, 2021.
DFRS: State of Nepal's Forests, Department of Forest Research and Survey, Kathmandu, ISBN 978-9937-8896-3-6, 2015.
Dhakal, B. P.: Traits and species distribution along the altitudinal gradient of the Himalayas, Nepal, M.Sc. thesis, Wageningen University & Research, Wageningen, the Netherlands, 1–75, https://www.researchgate.net/publication/325780659_Traits_and_species_distribution_along_the_altitudinal_gradient_of_the_Himalayas_Nepal (last access: 10 July 2024), 2018.
Díaz, S., Kattge, J., Cornelissen, J. H. C., Wright, I. J., Lavorel, S., Dray, S., Reu, B., Kleyer, M., Wirth, C., Colin Prentice, I., Garnier, E., Bönisch, G., Westoby, M., Poorter, H., Reich, P. B., Moles, A. T., Dickie, J., Gillison, A. N., Zanne, A. E., Chave, J., Joseph Wright, S., Sheremet'ev, S. N., Jactel, H., Baraloto, C., Cerabolini, B., Pierce, S., Shipley, B., Kirkup, D., Casanoves, F., Joswig, J. S., Günther, A., Falczuk, V., Rüger, N., Mahecha, M. D., and Gorné, L. D.: The global spectrum of plant form and function, Nature, 529, 167–171, https://doi.org/10.1038/nature16489, 2016.
Drollinger, S., Müller, M., Kobl, T., Schwab, N., Böhner, J., Schickhoff, U., and Scholten, T.: Decreasing nutrient concentrations in soils and trees with increasing elevation across a treeline ecotone in Rolwaling Himal, Nepal, J. Mt. Sci., 14, 843–858, https://doi.org/10.1007/s11629-016-4228-4, 2017.
Dyola, N., Sigdel, S. R., Liang, E., Babst, F., Camarero, J. J., Aryal, S., Chettri, N., Gao, S., Lu, X., Sun, J., Wang, T., Zhang, G., Zhu, H., Piao, S., and Peñuelas, J.: Species richness is a strong driver of forest biomass along broad bioclimatic gradients in the Himalayas, Ecosphere, 13, https://doi.org/10.1002/ecs2.4107, 2022.
Earth Resources Observation and Science (EROS) Center: Shuttle Radar Topography Mission (SRTM) 1 Arc-Second Global, USGS [data set], https://doi.org/10.5066/F7PR7TFT, 2000.
Earth Resources Observation and Science (EROS) Center: Global 30 Arc-Second Elevation (GTOPO30), USGS [data set], https://doi.org/10.5066/F7DF6PQS, 2017.
Falster, D. S., Duursma, R. A., Ishihara, M. I., Barneche, D. R., FitzJohn, R. G., Vårhammar, A., Aiba, M., Ando, M., Anten, N., Aspinwall, M. J., Baltzer, J. L., Baraloto, C., Battaglia, M., Battles, J. J., Bond-Lamberty, B., Van Breugel, M., Camac, J., Claveau, Y., Coll, L., Dannoura, M., Delagrange, S., Domec, J.-C., Fatemi, F., Feng, W., Gargaglione, V., Goto, Y., Hagihara, A., Hall, J. S., Hamilton, S., Harja, D., Hiura, T., Holdaway, R., Hutley, L. S., Ichie, T., Jokela, E. J., Kantola, A., Kelly, J. W. G., Kenzo, T., King, D., Kloeppel, B. D., Kohyama, T., Komiyama, A., Laclau, J.-P., Lusk, C. H., Maguire, D. A., Le Maire, G., Mäkelä, A., Markesteijn, L., Marshall, J., McCulloh, K., Miyata, I., Mokany, K., Mori, S., Myster, R. W., Nagano, M., Naidu, S. L., Nouvellon, Y., O'Grady, A. P., O'Hara, K. L., Ohtsuka, T., Osada, N., Osunkoya, O. O., Peri, P. L., Petritan, A. M., Poorter, L., Portsmuth, A., Potvin, C., Ransijn, J., Reid, D., Ribeiro, S. C., Roberts, S. D., Rodríguez, R., Saldaña-Acosta, A., Santa-Regina, I., Sasa, K., Selaya, N. G., Sillett, S. C., Sterck, F., Takagi, K., Tange, T., Tanouchi, H., Tissue, D., Umehara, T., Utsugi, H., Vadeboncoeur, M. A., Valladares, F., Vanninen, P., Wang, J. R., Wenk, E., Williams, R., De Aquino Ximenes, F., Yamaba, A., Yamada, T., Yamakura, T., Yanai, R. D., and York, R. A.: BAAD: a Biomass And Allometry Database for woody plants: Ecological Archives E096-128, Ecology, 96, 1445–1445, https://doi.org/10.1890/14-1889.1, 2015.
Freschet, G. T., Roumet, C., Comas, L. H., Weemstra, M., Bengough, A. G., Rewald, B., Bardgett, R. D., De Deyn, G. B., Johnson, D., Klimešová, J., Lukac, M., McCormack, M. L., Meier, I. C., Pagès, L., Poorter, H., Prieto, I., Wurzburger, N., Zadworny, M., Bagniewska-Zadworna, A., Blancaflor, E. B., Brunner, I., Gessler, A., Hobbie, S. E., Iversen, C. M., Mommer, L., Picon-Cochard, C., Postma, J. A., Rose, L., Ryser, P., Scherer-Lorenzen, M., Soudzilovskaia, N. A., Sun, T., Valverde-Barrantes, O. J., Weigelt, A., York, L. M., and Stokes, A.: Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs, New Phytol., 232, 1123–1158, https://doi.org/10.1111/nph.17072, 2021.
Gautam, K. H. and Devoe, N. N.: Ecological and anthropogenic niches of sal (Shorea robusta Gaertn. f.) forest and prospects for multiple-product forest management – a review, For. Int. J. For. Res., 79, 81–101, https://doi.org/10.1093/forestry/cpi063, 2006.
Grime, J. P.: Competitive Exclusion in Herbaceous Vegetation, Nature, 242, 344–347, https://doi.org/10.1038/242344a0, 1973.
Guan, Y., Liu, J., Cui, W., Chen, D., Zhang, J., Lu, H., Maeda, E. E., Zeng, Z., and Beck, H. E.: Elevation Regulates the Response of Climate Heterogeneity to Climate Change, Geophys. Res. Lett., 51, e2024GL109483, https://doi.org/10.1029/2024GL109483, 2024.
Hall, A. R., Miller, A. D., Leggett, H. C., Roxburgh, S. H., Buckling, A., and Shea, K.: Diversity–disturbance relationships: frequency and intensity interact, Biol. Lett., 8, 768–771, https://doi.org/10.1098/rsbl.2012.0282, 2012.
Hickler, T., Smith, B., Sykes, M. T., Davis, M. B., Sugita, S., and Walker, K.: Using a Generalized Vegetation Model to Simulate Vegetation Dynamics in Northeastern USA, Ecology, 85, 519–530, https://doi.org/10.1890/02-0344, 2004.
Jackson, J. K..: Manual of Afforestation in Nepal, Second edn., Ministry of Forests and Soil Conservation, Forest Research and Survey Centre, Kathmandu, Nepal, 496 pp., http://frtc.gov.np/downloadfiles/book2-1702615738.pdf (last access: 1 June 2024), 1994.
Jucker, T., Fischer, F. J., Chave, J., et al.: Tallo: A global tree allometry and crown architecture database, Glob. Change Biol., 28, 5254–5268, https://doi.org/10.1111/gcb.16302, 2022.
Khanal, S. and Boer, M. M.: Plot-level estimates of aboveground biomass and soil organic carbon stocks from Nepal's forest inventory, Sci. Data, 10, 406, https://doi.org/10.1038/s41597-023-02314-9, 2023.
Khanal, S., Nolan, R. H., Medlyn, B. E., and Boer, M. M.: Disentangling contributions of allometry, species composition and structure to high aboveground biomass density of high-elevation forests, For. Ecol. Manag., 554, 121679, https://doi.org/10.1016/j.foreco.2023.121679, 2024.
Khanal, S., Nolan, R. H., Medlyn, B. E., and Boer, M. M.: Topoclimatic factors create favourable conditions for carbon-dense forests in the Central Himalayas, Sci. Rep., 15, 35134, https://doi.org/10.1038/s41598-025-19127-y, 2025.
Kumar, P., Kumar, A., Patil, M., Hussain, S., and Singh, A. N.: Factors influencing tree biomass and carbon stock in the Western Himalayas, India, Front. For. Glob. Change, 6, 1328694, https://doi.org/10.3389/ffgc.2023.1328694, 2024.
Lamarque, J.-F., Shindell, D. T., Josse, B., Young, P. J., Cionni, I., Eyring, V., Bergmann, D., Cameron-Smith, P., Collins, W. J., Doherty, R., Dalsoren, S., Faluvegi, G., Folberth, G., Ghan, S. J., Horowitz, L. W., Lee, Y. H., MacKenzie, I. A., Nagashima, T., Naik, V., Plummer, D., Righi, M., Rumbold, S. T., Schulz, M., Skeie, R. B., Stevenson, D. S., Strode, S., Sudo, K., Szopa, S., Voulgarakis, A., and Zeng, G.: The Atmospheric Chemistry and Climate Model Intercomparison Project (ACCMIP): overview and description of models, simulations and climate diagnostics, Geosci. Model Dev., 6, 179–206, https://doi.org/10.5194/gmd-6-179-2013, 2013.
Latombe, G., Burke, A., Vrac, M., Levavasseur, G., Dumas, C., Kageyama, M., and Ramstein, G.: Comparison of spatial downscaling methods of general circulation model results to study climate variability during the Last Glacial Maximum, Geosci. Model Dev., 11, 2563–2579, https://doi.org/10.5194/gmd-11-2563-2018, 2018.
Laurent, L., Mårell, A., Korboulewsky, N., Saïd, S., and Balandier, P.: How does disturbance affect the intensity and importance of plant competition along resource gradients?, For. Ecol. Manag., 391, 239–245, https://doi.org/10.1016/j.foreco.2017.02.003, 2017.
Li, H., Guo, Q., Yang, L., Quan, H., and Wang, S.: Seasonal Eco-Physiology Characteristics of Four Evergreen Rhododendron Species to the Subalpine Habitats, Forests, 13, 653, https://doi.org/10.3390/f13050653, 2022.
Liang, J., Ding, Z., Lie, G., Zhou, Z., Singh, P. B., Zhang, Z., and Hu, H.: Species richness patterns of vascular plants and their drivers along an elevational gradient in the central Himalayas, Glob. Ecol. Conserv., 24, e01279, https://doi.org/10.1016/j.gecco.2020.e01279, 2020.
Luitel, D. R., Jha, P. K., Siwakoti, M., Shrestha, M. L., and Munniappan, R.: Climatic Trends in Different Bioclimatic Zones in the, Climate, 8, 1–18, https://doi.org/10.3390/cli8110136, 2020.
Maharjan, S. K., Sterck, F. J., Dhakal, B. P., Makri, M., and Poorter, L.: Functional traits shape tree species distribution in the Himalayas, J. Ecol., 109, 3818–3834, https://doi.org/10.1111/1365-2745.13759, 2021.
Maletha, A., Maikhuri, R. K., Bargali, S. S., Sharma, A., Negi, V. S., and Rawat, L. S.: Vegetation dynamics and soil nutrient availability in a temperate forest along altitudinal gradient of Nanda Devi Biosphere Reserve, Western Himalaya, India, PLoS ONE, 17, 1–22, https://doi.org/10.1371/journal.pone.0275051, 2022.
Måren, I. E., Karki, S., Prajapati, C., Yadav, R. K., and Shrestha, B. B.: Facing north or south: Does slope aspect impact forest stand characteristics and soil properties in a semiarid trans-Himalayan valley?, J. Arid Environ., 121, 112–123, https://doi.org/10.1016/j.jaridenv.2015.06.004, 2015.
MoFSC: Conservation Landscapes of Nepal, Ministry of Forests and Soil Conservation, Kathmandu, Nepal, https://lib.icimod.org/records/2h0bn-8rw98 (last access: 14 June 2024), 2016.
Monteux, S., Blume-Werry, G., Gavazov, K., Kirchhoff, L., Krab, E. J., Lett, S., Pedersen, E. P., and Väisänen, M.: Controlling biases in targeted plant removal experiments, New Phytol., 242, 1835–1845, https://doi.org/10.1111/nph.19386, 2024.
Muñoz Mazón, M., Klanderud, K., Finegan, B., Veintimilla, D., Bermeo, D., Murrieta, E., Delgado, D., and Sheil, D.: How forest structure varies with elevation in old growth and secondary forest in Costa Rica, For. Ecol. Manag., 469, 118191, https://doi.org/10.1016/j.foreco.2020.118191, 2020.
Murtagh, F. and Legendre, P.: Ward's Hierarchical Agglomerative Clustering Method: Which Algorithms Implement Ward's Criterion?, J. Classif., 31, 274–295, https://doi.org/10.1007/s00357-014-9161-z, 2014.
Myneni, R., Knyazikhin, Y., and Park, T.: MYD15A2H MODIS/Aqua Leaf Area Index/FPAR 8-Day L4 Global 500m SIN Grid V006, NASA Land Processes Distributed Active Archive Center [data set], https://doi.org/10.5067/MODIS/MYD15A2H.006, 2015.
Naud, L., Måsviken, J., Freire, S., Angerbjörn, A., Dalén, L., and Dalerum, F.: Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra, Ecol. Evol., 9, 4783–4795, https://doi.org/10.1002/ece3.5081, 2019.
Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P., and Kassem, K. R.: Terrestrial Ecoregions of the World: A New Map of Life on Earth, BioScience, 51, 933, https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2, 2001.
Paudel, P., Olin, S., and Smith, B.: LPJ-GUESS Code for Himalaya (Elevation gradient), Zenodo [code and data set], https://doi.org/10.5281/zenodo.17214801, 2025a.
Paudel, P., Olin, S., and Smith, B.: Forcing data and model output for Himalayas Elevation Gradient, Zenodo [data set], https://doi.org/10.5281/zenodo.17214851, 2025b.
Paudel, P., Olin, S., Tjoelker, M., Pontarp, M., Metcalfe, D., and Smith, B.: Impacts of Tree Allometry on Structure, Composition, Functioning and Competitive Interactions in Savanna Ecosystems on the Northern Australian Tropical Transect, J. Biogeogr., 53, e70131, https://doi.org/10.1111/jbi.70131, 2026.
Peng, S., Terrer, C., Smith, B., Ciais, P., Han, Q., Nan, J., Fisher, J. B., Chen, L., Deng, L., and Yu, K.: Carbon restoration potential on global land under water resource constraints, Nat. Water, 2, 1071–1081, https://doi.org/10.1038/s44221-024-00323-5, 2024.
Pierce, S., Brusa, G., Vagge, I., and Cerabolini, B. E. L.: Allocating CSR plant functional types: the use of leaf economics and size traits to classify woody and herbaceous vascular plants, Funct. Ecol., 27, 1002–1010, https://doi.org/10.1111/1365-2435.12095, 2013.
Pokhrel, S. and Sherpa, C.: Analyzing the relationship, distribution of tree species diversity, and above-ground biomass on the Chitwan-Annapurna landscape in Nepal, Int. J. For. Res., 2020, https://doi.org/10.1155/2020/2789753, 2020.
Poudel, A. S., Shrestha, B. B., Joshi, M. D., Muniappan, R., Adiga, A., Venkatramanan, S., and Jha, P. K.: Predicting the Current and Future Distribution of the Invasive Weed Ageratina adenophora in the Chitwan-Annapurna Landscape, Nepal, Mt. Res. Dev., 40, R61–R71, https://doi.org/10.1659/MRD-JOURNAL-D-19-00069.1, 2020.
Pugh, T. A. M., Arneth, A., Kautz, M., Poulter, B., and Smith, B.: Important role of forest disturbances in the global biomass turnover and carbon sinks, Nat. Geosci., 12, 730–735, https://doi.org/10.1038/s41561-019-0427-2, 2019.
Rabin, S. S., Melton, J. R., Lasslop, G., Bachelet, D., Forrest, M., Hantson, S., Kaplan, J. O., Li, F., Mangeon, S., Ward, D. S., Yue, C., Arora, V. K., Hickler, T., Kloster, S., Knorr, W., Nieradzik, L., Spessa, A., Folberth, G. A., Sheehan, T., Voulgarakis, A., Kelley, D. I., Prentice, I. C., Sitch, S., Harrison, S., and Arneth, A.: The Fire Modeling Intercomparison Project (FireMIP), phase 1: experimental and analytical protocols with detailed model descriptions, Geosci. Model Dev., 10, 1175–1197, https://doi.org/10.5194/gmd-10-1175-2017, 2017.
Satyakam, Zinta, G., Singh, R. K., and Kumar, R.: Cold adaptation strategies in plants – An emerging role of epigenetics and antifreeze proteins to engineer cold resilient plants, Front. Genet., 13, 909007, https://doi.org/10.3389/fgene.2022.909007, 2022.
Scherstjanoi, M., Kaplan, J. O., and Lischke, H.: Application of a computationally efficient method to approximate gap model results with a probabilistic approach, Geosci. Model Dev., 7, 1543–1571, https://doi.org/10.5194/gmd-7-1543-2014, 2014.
Shah, S., Shrestha, K. K., and Scheidegger, C.: Variation in Plant Functional Traits along Altitudinal Gradient and Land Use Types in Sagarmatha National Park and Buffer Zone, Nepal, Am. J. Plant Sci., 10, 595–614, https://doi.org/10.4236/ajps.2019.104043, 2019.
Shrestha, K. B., Hofgaard, A., and Vandvik, V.: Recent treeline dynamics are similar between dry and mesic areas of Nepal, central Himalaya, J. Plant Ecol., 8, 347–358, https://doi.org/10.1093/jpe/rtu035, 2015.
Sigdel, S. R., Liang, E., Wang, Y., Dawadi, B., and Camarero, J. J.: Tree-to-tree interactions slow down Himalayan treeline shifts as inferred from tree spatial patterns, J. Biogeogr., 47, 1816–1826, https://doi.org/10.1111/jbi.13840, 2020.
Sigdel, S. R., Liang, E., Rokaya, M. B., Rai, S., Dyola, N., Sun, J., Zhang, L., Zhu, H., Chettri, N., Chaudhary, R. P., Camarero, J. J., and Peñuelas, J.: Functional traits of a plant species fingerprint ecosystem productivity along broad elevational gradients in the Himalayas, Funct. Ecol., 383–394, https://doi.org/10.1111/1365-2435.14226, 2022.
Sitch, S., Smith, B., Prentice, I. C., Arneth, A., Bondeau, A., Cramer, W., Kaplan, J. O., Levis, S., Lucht, W., Sykes, M. T., Thonicke, K., and Venevsky, S.: Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model, Glob. Change Biol., 9, 161–185, https://doi.org/10.1046/j.1365-2486.2003.00569.x, 2003.
Smith, B. and Wilson, J. B.: A Consumer's Guide to Evenness Indices, Oikos, 76, 70, https://doi.org/10.2307/3545749, 1996.
Smith, B., Prentice, I. C., and Sykes, M. T.: Representation of vegetation dynamics in the modelling of terrestrial ecosystems: Comparing two contrasting approaches within European climate space, Glob. Ecol. Biogeogr., 10, 621–637, https://doi.org/10.1046/j.1466-822X.2001.t01-1-00256.x, 2001.
Smith, B., Wårlind, D., Arneth, A., Hickler, T., Leadley, P., Siltberg, J., and Zaehle, S.: Implications of incorporating N cycling and N limitations on primary production in an individual-based dynamic vegetation model, Biogeosciences, 11, 2027–2054, https://doi.org/10.5194/bg-11-2027-2014, 2014.
Thakur, D. and Chawla, A.: Functional diversity along elevational gradients in the high altitude vegetation of the western Himalaya, Biodivers. Conserv., 28, 1977–1996, https://doi.org/10.1007/s10531-019-01728-5, 2019.
Thakur, G., Kumar, P., Bhardwaj, D. R., Prakash, P., and Poonam: Dynamics of aboveground vegetation biomass and carbon stocks along the altitudinal gradients and overstorey composition types in the temperate Himalayan region, Trees For. People, 16, 100553, https://doi.org/10.1016/j.tfp.2024.100553, 2024.
Thakur, R. B. and Phulara, N. K. (Eds.): A compendium of tree species of Nepal, 2nd edn., D. K. Agencies (P) Ltd., Kathmandu, Nepal, 375 pp., ISBN 9789937275583, 2014.
Thorne, J. H., Choe, H., Dorji, L., Yangden, K., Wangdi, D., Phuntsho, Y., and Beardsley, K.: Species richness and turnover patterns for tropical and temperate plants on the elevation gradient of the eastern Himalayan Mountains, Front. Ecol. Evol., 10, 1–15, https://doi.org/10.3389/fevo.2022.942759, 2022.
Tito, R., Vasconcelos, H. L., and Feeley, K. J.: Mountain Ecosystems as Natural Laboratories for Climate Change Experiments, Front. For. Glob. Change, 3, 38, https://doi.org/10.3389/ffgc.2020.00038, 2020.
Tsunoda, Y., Tsuda, T., and Ohno, Y.: Influence of stand structure on tree height: A comparative study across 46 species, For. Ecol. Manag., 578, 122474, https://doi.org/10.1016/j.foreco.2024.122474, 2025.
Whittaker, R. H.: Dominance and Diversity in Land Plant Communities: Numerical relations of species express the importance of competition in community function and evolution, Science, 147, 250–260, https://doi.org/10.1126/science.147.3655.250, 1965.
Zhu, L., Zhang, Y., Ye, H., Li, Y., Hu, W., Du, J., and Zhao, P.: Variations in leaf and stem traits across two elevations in subtropical forests, Funct. Plant Biol., 49, 319–332, https://doi.org/10.1071/FP21220, 2022.
Short summary
We used computer simulations to study how vegetation changes from low to high elevations in the Himalayas. We found that competition between plants is more important in warm lowlands, while cold conditions limit plant growth at high elevations. These differences influence plant diversity, vegetation structure, and the amount of plant growth across mountain landscapes, showing how climate and plant strategies together shape ecosystems.
We used computer simulations to study how vegetation changes from low to high elevations in the...
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