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Elevation-Related Variation in Vegetation Richness, Composition, and Forage Productivity during the Dry Season in Silvopastoral Systems of Timor Tengah Selatan, Indonesia

Authors

  • Grace Maranatha ORCID Animal Science Study Program, Hasanuddin University, Makassar 85001, Indonesia
  • Budiman Email ORCID Department of Animal Nutrition and Feed Science, Hasanuddin University, Makassar 90245, Indonesia
  • Jasmal Ahmari Syamsu ORCID Department of Animal Nutrition and Feed Science, Hasanuddin University, Makassar 90245, Indonesia

DOI:

https://doi.org/10.30564/re.v8i6.117392
Received: 7 September 2026 | Revised: 30 September 2026 | Accepted: 16 October 2026 | Published Online: 12 November 2026

Abstract

Tropical semi-arid silvopastoral systems undergo seasonal changes in vegetation and forage availability, but their temporal dynamics across elevation classes in Timor, Indonesia, remain insufficiently documented. This study aimed to evaluate species richness, botanical composition, and available forage biomass across two elevation classes during the dry season and to assess monthly variation and elevation × month interactions. An observational 2 × 6 factorial design was applied across four grazing locations in Timor Tengah Selatan, Indonesia, comprising two lowland sites (<500 m above sea level) and two highland sites (>1,000 m above sea level). Monthly observations were conducted from June to November 2025 using temporary 1 × 1 m quadrats. Elevation, month, and their interaction significantly affected species richness, the proportions of Poaceae and Fabaceae, and fresh available forage biomass (p < 0.001). The elevation × month interaction also affected non-Poaceae/non-Fabaceae herbaceous vegetation (p = 0.032) and dry available forage biomass (p = 0.002). Species richness and available forage biomass were generally higher in the highlands, whereas temporal variation was greater in the lowlands. From June to September, fresh available forage biomass decreased by approximately 75% in the lowlands and 37% in the highlands, while dry available forage biomass decreased by 47% and 12%, respectively. Biomass increased again toward November, with different magnitudes between elevation classes. These findings indicate spatial and temporal differences in vegetation and available forage biomass among the four study locations during the dry season, while reflecting associations rather than a universal causal effect of elevation.

Keywords:

Elevation, Forage Productivity, Botanical Composition, Species Richness, Dry Season, Silvopastoral Systems, Timor Tengah Selatan

References

[1] Rusch, G.M., Zapata, P.C., Casanoves, F., et al., 2014. Determinants of grassland primary production in seasonally-dry silvopastoral systems in Central America. Agroforestry Systems. 88, 517–526. DOI: https://doi.org/10.1007/s10457-014-9711-8

[2] Pezzopane, J.R.M., Bosi, C., Nicodemo, M.L.F., et al., 2015. Microclimate and soil moisture in a silvopastoral system in southeastern Brazil. Bragantia. 74(1), 110–119.

[3] Schinato, F., Munka, M.C., Olmos, V.M., et al., 2023. Microclimate, forage production and carbon storage in a eucalypt-based silvopastoral system. Agriculture, Ecosystems & Environment. 344, 108290. DOI: https://doi.org/10.1016/j.agee.2022.108290

[4] Hébert, È.-M., Martin-Guay, M.-O., Maheu, A., et al., 2025. Microclimate, soil moisture and forage yield vary spatially within a temperate tree-based intercropping system: From competition to facilitation. Agricultural Water Management. 319, 109782. DOI: https://doi.org/10.1016/j.agwat.2025.109782

[5] McNichol, B.H., Wang, R., Hefner, A., et al., 2024. Topography-driven microclimate gradients shape forest structure, diversity, and composition in a temperate refugial forest. Plant-Environment Interactions. 5, e10153. DOI: https://doi.org/10.1002/pei3.10153

[6] Moeslund, J.E., Arge, L., Bøcher, P.K., et al., 2013. Topographically controlled soil moisture is the primary driver of local vegetation patterns across a lowland region. Ecosphere. 4(7), 1–26. DOI: https://doi.org/10.1890/ES13-00134.1

[7] Gomes, F.J., Pedreira, B.C., Santos, P.M., et al., 2020. Microclimate effects on canopy characteristics of shaded palisadegrass pastures in a silvopastoral system in the Amazon biome of central Brazil. European Journal of Agronomy. 115, 126029. DOI: https://doi.org/10.1016/j.eja.2020.126029

[8] Grossiord, C., Buckley, T.N., Cernusak, L.A., et al., 2020. Plant responses to rising vapor pressure deficit. New Phytologist. 226(6), 1550–1566. DOI: https://doi.org/10.1111/nph.16485

[9] López, J., Way, D.A., Sadok, W., 2021. Systemic effects of rising atmospheric vapor pressure deficit on plant physiology and productivity. Global Change Biology. 27(9), 1704–1720. DOI: https://doi.org/10.1111/gcb.15548

[10] Habaora, F., Fuah, A.M., Abdullah, L., et al., 2020. Botanical composition and carrying capacity in various agroecosystems on the Timor Island. Jurnal Ilmu Produksi dan Teknologi Hasil Peternakan. 8(2), 72–79. DOI: https://doi.org/10.29244/jipthp.8.2.72-79

[11] Se’u, V.E., Karti, P.D.M.H., Abdullah, L., 2015. Botanical composition, grass production, and carrying capacity of pasture in Timor Tengah Selatan District. Media Peternakan. 38(3), 176–182. https://doi.org/10.5398/medpet.2015.38.3.176

[12] Jelantik, I.G.N., Malelak, G.E.M., 2023. Study on the impact of season and location on the native pastures’ quality in West Timor, Indonesia. Journal of Agriculture and Ecology Research International. 24(5), 67–72. DOI: https://doi.org/10.9734/jaeri/2023/v24i5543

[13] Maranatha, G., Pandarangga, P., Sobang, Y.U.L., et al., 2025. Forage nutrient fluctuations during the dry season: A case study of tropical grazing land in East Nusa Tenggara, Indonesia. Research in Ecology. 7(4), 71–84. DOI: https://doi.org/10.30564/re.v7i4.9954

[14] Qian, J., Ye, M., Zhang, X., et al., 2024. Characteristics of grassland species diversity and soil physicochemical properties with elevation gradient in Burzin Forest Area. Agriculture. 14(7), 1176. DOI: https://doi.org/10.3390/agriculture14071176

[15] Shaheen, I., Malik, R.A., Sankaran, M., et al., 2025. Differential impacts of grazing on grassland plant diversity, biomass, soil C, and soil N across an elevation gradient. Ecological Applications. 35(3), e70031. DOI: https://doi.org/10.1002/eap.70031

[16] Fenollosa, E., Fernandes, P., Hector, A., et al., 2024. Differential responses of community-level functional traits to mid- and late-season experimental drought in a temperate grassland. Journal of Ecology. 112(10), 2292–2306. DOI: https://doi.org/10.1111/1365-2745.14395

[17] Luo, W., Ma, W., Song, L., et al., 2023. Compensatory dynamics drive grassland recovery from drought. Journal of Ecology. 111(6), 1281–1291. DOI: https://doi.org/10.1111/1365-2745.14096

[18] Chandregowda, M.H., Tjoelker, M.G., Pendall, E., et al., 2022. Root trait shifts towards an avoidance strategy promote productivity and recovery in C3 and C4 pasture grasses under drought. Functional Ecology. 36(7), 1754–1771. DOI: https://doi.org/10.1111/1365-2435.14085

[19] Zhang, M., Delgado-Baquerizo, M., Li, G., et al., 2023. Experimental impacts of grazing on grassland biodiversity and function are explained by aridity. Nature Communications. 14, 5040. DOI: https://doi.org/10.1038/s41467-023-40809-6

[20] Zhang, P., Seabloom, E.W., Foo, J., et al., 2025. Dominant species predict plant richness and biomass in global grasslands. Nature Ecology & Evolution. 9(6), 924–936. DOI: https://doi.org/10.1038/s41559-025-02701-y

[21] Hossain, M.L., Li, J., 2024. Species richness and dominant functional groups enhance aboveground biomass, with no effect on belowground biomass in Qinghai-Tibet Plateau's grasslands. Ecological Informatics. 82, 102688. DOI: https://doi.org/10.1016/j.ecoinf.2024.102688

[22] Hassan, N., Wang, Z., 2024. Paralleled grazing and mowing differentially affected plant community diversity and productivity in a semi-arid grassland. Ecological Processes. 13, 62. DOI: https://doi.org/10.1186/s13717-024-00543-3

[23] Bazzichetto, M., Sperandii, M.G., Penone, C., et al., 2024. Biodiversity promotes resistance but dominant species shape recovery of grasslands under extreme drought. Journal of Ecology. 112(5), 1087–1100. DOI: https://doi.org/10.1111/1365-2745.14288

[24] Liu, L., Zheng, J., Guan, J., et al., 2024. Strong positive direct impact of soil moisture on the growth of Central Asian grasslands. Science of the Total Environment. 954, 176663. DOI: https://doi.org/10.1016/j.scitotenv.2024.176663

[25] Zhang, T., Liu, Y., Zimini, Y., et al., 2024. Effects of soil moisture and atmospheric vapor pressure deficit on the temporal variability of productivity in Eurasian grasslands. Remote Sensing. 16(13), 2368. DOI: https://doi.org/10.3390/rs16132368

[26] Novick, K.A., Ficklin, D.L., Grossiord, C., et al., 2024. The impacts of rising vapour pressure deficit in natural and managed ecosystems. Plant, Cell & Environment. 47(9), 3561–3589. DOI: https://doi.org/10.1111/pce.14846

[27] John, A., Pradhan, K., Case, M.J., et al., 2024. Forest canopy cover affects microclimate buffering during an extreme heat event. Environmental Research Communications. 6, 091015. DOI: https://doi.org/10.1088/2515-7620/ad7705

[28] Demir, G., Michalzik, B., Filipzik, J., et al., 2022. Spatial variation of grassland canopy affects soil wetting patterns and preferential flow. Hydrological Processes. 36(12), e14760. DOI: https://doi.org/10.1002/hyp.14760

[29] Greiser, C., Hederová, L., Vico, G., et al., 2024. Higher soil moisture increases microclimate temperature buffering in temperate broadleaf forests. Agricultural and Forest Meteorology. 345, 109828. DOI: https://doi.org/10.1016/j.agrformet.2023.109828

[30] Ye, J., Ji, Y., Wang, J., et al., 2024. Climate factors dominate the elevational variation in grassland plant resource utilization strategies. Frontiers in Plant Science. 15, 1430027. DOI: https://doi.org/10.3389/fpls.2024.1430027

[31] Su, J., Zhang, Y., Xu, F., 2023. Divergent responses of grassland productivity and plant diversity to intra-annual precipitation variability across climate regions: A global synthesis. Journal of Ecology. 111(9), 1921–1934. DOI: https://doi.org/10.1111/1365-2745.14149

[32] Zhang, Y., Zhou, T., Liu, X., et al., 2023. Crucial roles of the optimal time-scale of water condition on grassland biomass estimation on Qinghai–Tibet Plateau. Science of the Total Environment. 905, 167210. DOI: https://doi.org/10.1016/j.scitotenv.2023.167210

[33] Lin, X., Zhao, H., Zhang, S., et al., 2024. Global response of different types of grasslands to precipitation and grazing, especially belowground biomass. Agriculture, Ecosystems & Environment. 363, 108852. DOI: https://doi.org/10.1016/j.agee.2023.108852

[34] Shi, L., Zhou, Y., He, W., et al., 2024. Mapping critical soil moisture thresholds of water stress for global grasslands. Journal of Hydrology. 644, 132090. DOI: https://doi.org/10.1016/j.jhydrol.2024.132090

[35] Cao, F., Li, W., Jiang, Y., et al., 2024. Effects of grazing on grassland biomass and biodiversity: A global synthesis. Field Crops Research. 306, 109204. DOI: https://doi.org/10.1016/j.fcr.2023.109204

[36] Duan, D., Tian, Z., Wu, N., et al., 2023. Drought neutralizes positive effects of long-term grazing on grassland productivity through altering plant–soil interactions. Functional Ecology. 37(7), 1827–1840. DOI: https://doi.org/10.1111/1365-2435.14346

[37] Zhu, X., An, Y., Qin, Y., et al., 2024. How do short-term and long-term factors impact the aboveground biomass of grassland in Northern China? Carbon Research. 3, 54. DOI: https://doi.org/10.1007/s44246-024-00134-z

[38] Su, J., Bai, Y., Xu, F., 2022. Multiple global changes drive grassland productivity and stability: A meta-analysis. Journal of Ecology. 110(12), 2850–2869. DOI: https://doi.org/10.1111/1365-2745.13983

[39] Jackson, J., Middleton, S.L., Lawson, C.S., et al., 2024. Experimental drought reduces the productivity and stability of a calcareous grassland. Journal of Ecology. 112(4), 917–931. DOI: https://doi.org/10.1111/1365-2745.14282

[40] Zhu, J., Zhang, Y., Wu, J., et al., 2024. Herbivore exclusion stabilizes alpine grassland biomass production across spatial scales. Global Change Biology. 30(1), e17155. DOI: https://doi.org/10.1111/gcb.17155

[41] Cordeiro, A.D.A.C., Klanderud, K., Villa, P.M., et al., 2023. Patterns of species richness and beta diversity of vascular plants along elevation gradient in Brazilian páramo. Journal of Mountain Science. 20, 1911–1920. DOI: https://doi.org/10.1007/s11629-022-7480-9

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2026 Vol.8 No.6

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Copyright © 2026 Grace Maranatha, Budiman, Jasmal Ahmari Syamsu

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