Climatic and elevational drivers of Selaginella species richness in Java, Indonesia
Article Sidebar
Abstract Views: 18
File Views: 9
Main Article Content
Abstract
Abstract. Setyawan AD, Sutarno, Sugiyarto, Sunarto, Sulton MN. 2026. Climatic and elevational drivers of Selaginella species richness in Java, Indonesia. Nusantara Bioscience 18 (1): n180106. https://doi.org/10.13057/nusbiosci/n180106. Understanding environmental factors that shape species richness patterns, yet the determinants of Selaginella diversity in Java, Indonesia, remain poorly understood. Selaginella, an ancient lycophyte genus widely distributed in tropical regions, is highly sensitive to environmental conditions, particularly moisture availability and temperature. This study examined the relationships between Selaginella species richness and elevational and climatic gradients across Java, Indonesia. A total of 1,962 occurrence records representing 21 identified species and one unidentified taxon were compiled from field surveys, herbarium collections, and verified biodiversity databases. Species richness was estimated using a 0.25° × 0.25° grid system and spatially visualized through Inverse Distance Weighting (IDW) interpolation. The influence of elevation, Annual Mean Temperature, Annual Mean Precipitation, Annual Mean Solar Radiation, and Annual Mean Moisture Index on species richness was evaluated using polynomial regression, Pearson correlation analysis, and Principal Component Analysis (PCA). Species richness showed pronounced spatial heterogeneity, with hotspots concentrated in the humid montane regions of West Java. Richness exhibited nonlinear relationships with all environmental variables and peaked at intermediate elevations. Annual Mean Moisture Index (R² = 0.616) and Annual Mean Precipitation (R² = 0.527) were the variables most strongly associated with species richness, whereas Annual Mean Temperature and solar radiation showed negative relationships. Correlation and PCA results further indicated that moisture availability is closely linked to richness patterns, while elevation influences richness indirectly through its effects on local climatic conditions. These findings demonstrate that humid submontane and montane environments are critical for maintaining Selaginella diversity in Java and highlight the importance of conserving climatically suitable habitats under future environmental change.
Article Details
Issue
Section
This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License.
How to Cite
References
Acebey AR, Krömer T, Kessler M. 2017. Species richness and vertical distribution of ferns and lycophytes along an elevational gradient in Los Tuxtlas, Veracruz, Mexico. Flora 235: 83-91. https://doi.org/10.1016/j.flora.2017.08.003. DOI: https://doi.org/10.1016/j.flora.2017.08.003
Alston AHG. 1935. The Selaginellae of the Malay Islands: I. Java and the Lesser Sunda Islands. Bulletin du Jardin Botanique de Buitenzorg, Series III 13: 432-442.
Alston AHG. 1937. The Selaginellae of the Malay Islands II. Sumatra. Bull Jard Bot Buitenzorg III 14: 175-186.
Bhattarai KR, Vetaas OR, Grytnes JA. 2004. Fern species richness along a central Himalayan elevational gradient, Nepal. J Biogeogr 31 (3): 389-400. https://doi.org/10.1046/j.0305-0270.2003.01013.x. DOI: https://doi.org/10.1046/j.0305-0270.2003.01013.x
Bhattarai KR, Vetaas OR. 2003. Variation in plant species richness of different life forms along a subtropical elevation gradient in the Himalayas, east Nepal. Glob Ecol Biogeogr 12 (4): 327-340. https://doi.org/10.1046/j.1466-822X.2003.00044.x. DOI: https://doi.org/10.1046/j.1466-822X.2003.00044.x
Bhattarai P, Vetaas OR, Zhao G. 2025. Do water-energy dynamics drive plant species richness patterns on the high alpine Tibetan plateau? Ecosphere 16 (5): e70285. https://doi.org/10.1002/ecs2.70285. DOI: https://doi.org/10.1002/ecs2.70285
Cavalcante T, Weber MM, Barnett AA. 2022. Combining geospatial abundance and ecological niche models to identify high-priority areas for conservation: The neglected role of broad-scale interspecific competition. Front Ecol Evol 10: 915325. https://doi.org/10.3389/fevo.2022.915325. DOI: https://doi.org/10.3389/fevo.2022.915325
Chen Y, Shan X, Jin X, Yang T, Dai F, Yang D. 2016. A comparative study of spatial interpolation methods for determining fishery resources density in the Yellow Sea. Acta Oceanol Sin 35: 65-72. https://doi.org/10.1007/s13131-016-0966-y. DOI: https://doi.org/10.1007/s13131-016-0966-y
Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré G, Marquéz JRG, Gruber B, Lafourcade B, Leitão PJ, Münkemüller T, McClean C, Osborne PE, Reineking B, Schröder B, Skidmore AK, Zurell D, Lautenbach S. 2013. Collinearity: A review of methods to deal with it and a simulation study evaluating their performance. Ecography 36 (1): 27-46. https://doi.org/10.1111/j.1600-0587.2012.07348.x DOI: https://doi.org/10.1111/j.1600-0587.2012.07348.x
Fick SE, Hijmans RJ. 2017. WorldClim 2: New 1-km spatial resolution climate surfaces for global land areas. Int J Climatol 37 (12): 4302-4315. https://doi.org/10.1002/joc.5086. DOI: https://doi.org/10.1002/joc.5086
Grytnes JA, McCain CM. 2007. Elevational trends in biodiversity. In: Levin SA (eds.). Encyclopedia of Biodiversity. Elsevier, Amsterdam. https://doi.org/10.1002/9780470015902.a0022548. DOI: https://doi.org/10.1016/B978-012226865-6/00503-1
Hammer Ø, Harper DAT, Ryan PD. 2001. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4 (1): 9.
Hawkins BA, Field R, Cornell HV, Currie DJ, Guégan JF, Kaufman DM, Kerr JT, Mittelbach GG, Oberdorff T, O'Brien EM, Porter EE, Turner JRG. 2003. Energy, water, and broad-scale geographic patterns of species richness. Ecology 84 (12): 3105-3117. https://doi.org/10.1890/03-8006. DOI: https://doi.org/10.1890/03-8006
Hernández-Rojas AC, Kluge J, Krömer T, Carvajal-Hernández CI, Lehnert M, Kessler M. 2020. Biogeographical patterns of fern richness and range size along tropical elevational-latitudinal gradients. J Biogeogr 47 (8): 1716-1730. https://doi.org/10.1111/jbi.13841. DOI: https://doi.org/10.1111/jbi.13841
Jolliffe IT, Cadima J. 2016. Principal component analysis: A review and recent developments. Philos Trans R Soc A 374: 20150202. https://doi.org/10.1098/rsta.2015.0202. DOI: https://doi.org/10.1098/rsta.2015.0202
Körner C. 2004. Mountain biodiversity, its causes and function. Ambio 33 (13): 11-17. https://doi.org/10.1007/0044-7447-33.sp13.11. DOI: https://doi.org/10.1007/0044-7447-33.sp13.11
Kriticos DJ, Jarošík V, Ota N. 2014. Extending the suite of Bioclim variables: A proposed registry system and case study using principal components analysis. Methods Ecol Evol 5 (9): 956-960. https://doi.org/10.1111/2041-210X.12244. DOI: https://doi.org/10.1111/2041-210X.12244
Kriticos DJ, Webber BL, Leriche A, Ota N, Macadam I, Bathols J, Scott JK. 2012. CliMond: Global high-resolution historical and future scenario climate surfaces for bioclimatic modeling. Methods Ecol Evol 3 (1): 53-64. https://doi.org/10.1111/j.2041-210X.2011.00134.x. DOI: https://doi.org/10.1111/j.2041-210X.2011.00134.x
Krömer T, Kessler M, Kluge J, Acebey A, Gradstein SR. 2013. Effects of altitude and climate in determining elevational plant species richness patterns. Flora 208 (3): 197-210. https://doi.org/10.1016/j.flora.2013.03.003. DOI: https://doi.org/10.1016/j.flora.2013.03.003
Kumar A, Patil M, Kumar P, Singh AN. 2024. Determinants of plant species richness along elevational gradients: Insights with climate, energy and water-energy dynamics. Ecol Process 13: 86. https://doi.org/10.1186/s13717-024-00534-0. DOI: https://doi.org/10.1186/s13717-024-00563-z
Li J, Pandey B, Dakhil MA, Zhang Y, Khan TU, Zhang H, Wang H, Wang Y. 2022. Precipitation and potential evapotranspiration determine the distribution patterns of threatened plant species in Sichuan Province, China. Sci Rep 12: 22418. https://doi.org/10.1038/s41598-022-26171-5. DOI: https://doi.org/10.1038/s41598-022-26171-5
Mauri M, Elli T, Caviglia G, Uboldi G, Azzi M. 2017. RAWGraphs: A visualization platform to create open outputs. Proc 12th Biannual Conference of the Italian SIGCHI Chapter. https://doi.org/10.1145/3125571.3125585. DOI: https://doi.org/10.1145/3125571.3125585
Pouteau R, Meyer JY, Taputuarai R, Stoll B. 2016. Fern species richness and abundance are indicators of climate change on high-elevation islands: Evidence from an elevational gradient on Tahiti (French Polynesia). Clim Change 138: 143-156. https://doi.org/10.1007/s10584-016-1734-x. DOI: https://doi.org/10.1007/s10584-016-1734-x
POWO. 2026. Plants of the World Online. Royal Botanic Gardens, Kew. https://powo.science.kew.org. Accessed 2026.
Rahbek C. 2005. The role of spatial scale and the perception of large-scale species-richness patterns. Ecol Lett 8 (2): 224-239. https://doi.org/10.1111/j.1461-0248.2004.00701.x. DOI: https://doi.org/10.1111/j.1461-0248.2004.00701.x
Rashidi O, Latiff A, Faridah-Hanum I. 2015. Effects of altitude and microclimate on the distribution of ferns in urban and natural environments. J Teknol 77 (30): 125-131. https://doi.org/10.11113/jt.v77.6535. DOI: https://doi.org/10.11113/jt.v77.6876
Setyawan AD, Chikmawati T, Miftahudin, Sutarno, Sugiyarto, Sunarto. 2026a. Revisiting Selaginella diversity in Java (Indonesia) through combined ecological and taxonomic reassessment. Biodiversitas 27 (1): d270150. https://doi.org/10.13057/biodiv/d270150. DOI: https://doi.org/10.13057/biodiv/d270150
Setyawan AD, Chikmawati T, Miftahudin, Sutarno, Sugiyarto, Sunarto. 2026b. Taxonomic reassessment of the Selaginella flora of Java, Indonesia. Biodiversitas 27 (2): d270239. https://doi.org/10.13057/biodiv/d270239. DOI: https://doi.org/10.13057/biodiv/d270239
Setyawan AD, Sugiyarto, Widiastuti A. 2015. Species diversity of Selaginella in the Dieng Plateau, Central Java. Pros Sem Nas Masy Biodiv Indon 1 (5): 980-986. https://doi.org/10.13057/psnmbi/m010504. DOI: https://doi.org/10.13057/psnmbi/m010504
Setyawan AD, Sugiyarto. 2015. Diversity of Selaginella in the Bromo Tengger Semeru National Park, East Java. Pros Sem Nas Masy Biodiv Indon 1 (6): 1312-1317. https://doi.org/10.13057/psnmbi/m010609. DOI: https://doi.org/10.13057/psnmbi/m010609
Setyawan AD, Supriatna J, Darnaedi D, Rokhmatuloh, Sutarno, Sugiyarto. 2016. Diversity of Selaginella across altitudinal gradient of the tropical region. Biodiversitas 17 (1): 384-400. https://doi.org/10.13057/biodiv/d170152. DOI: https://doi.org/10.13057/biodiv/d170152
Setyawan AD, Supriatna J, Nisyawati, Nursamsi I, Sutarno, Sugiyarto, Sunarto, Pradan P, Budiharta S, Pitoyo A, Suhardono S, Setyono P, Indrawan M. 2021. Projecting the expansion range of Selaginella zollingeriana in the Indonesian archipelago under future climate conditions. Biodiversitas 22 (4): 2088-2103. https://doi.org/10.13057/biodiv/d220458. DOI: https://doi.org/10.13057/biodiv/d220458
Setyawan AD, Supriatna J, Nisyawati, Nursamsi I, Sutarno, Sugiyarto, Sunarto, Pradhan P, Budiharta S, Pitoyo A, Suhardono S, Setyono P, Indrawan M. 2020b. Anticipated climate changes reveal shifting in habitat suitability of high-altitude Selaginellas in Java, Indonesia. Biodiversitas 21 (11): 5482-5497. https://doi.org/10.13057/biodiv/d211157. DOI: https://doi.org/10.13057/biodiv/d211157
Setyawan AD, Supriatna J, Nisyawati, Nursamsi I, Sutarno, Sugiyarto, Sunarto, Pradhan P, Budiharta S, Pitoyo A, Suhardono S, Setyono P, Indrawan M. 2020a. Predicting potential impacts of climate change on the geographical distribution of mountainous Selaginellas in Java, Indonesia. Biodiversitas 21 (10): 4866-4877. https://doi.org/10.13057/biodiv/d211053. DOI: https://doi.org/10.13057/biodiv/d211053
Setyawan AD, Supriatna J, Nisyawati, Sutarno, Sugiyarto, Nursamsi I. 2018. Predicting impacts of future climate change on the distribution of the widespread selaginellas (Selaginella ciliaris and S. plana) in Southeast Asia. Biodiversitas 19 (5): 1960-1977. https://doi.org/10.13057/biodiv/d190548. DOI: https://doi.org/10.13057/biodiv/d190549
Setyawan AD, Sutarno, Sugiyarto. 2013. Species diversity of Selaginella in Mount Lawu, Java, Indonesia. Biodiversitas 14 (1): 1-9. https://doi.org/10.13057/biodiv/d140101. DOI: https://doi.org/10.13057/biodiv/d140101
Setyawan AD. 2011. Recent status of Selaginella (Selaginellaceae) research in Nusantara. Biodiversitas 12 (2): 112-124. https://doi.org/10.13057/biodiv/d120209. DOI: https://doi.org/10.13057/biodiv/d120209
Umair M, Hu X, Cheng Q, Ali S, Ni J. 2023. Distribution patterns of fern species richness along elevations on the Tibetan Plateau in China: Regional differences and effects of climate change variables. Front Plant Sci 14: 1178603. https://doi.org/10.3389/fpls.2023.1178603. DOI: https://doi.org/10.3389/fpls.2023.1178603
Weststrand S, Korall P. 2016. A subgeneric classification of Selaginella (Selaginellaceae). Am J Bot 103 (12): 2160-2169. https://doi.org/10.3732/ajb.1600288 DOI: https://doi.org/10.3732/ajb.1600288
Wong KM. 1982. Critical observations on Peninsular Malaysian Selaginella. Gard Bull Singap 35: 107-135.
Wong KM. 2010. Selaginellaceae. In: Kiew R, Chung RCK, Saw LG, Soepadmo E (eds.). Flora of Peninsular Malaysia Series I: Ferns and Lycophytes. Vol. 1. Forest Research Institute Malaysia, Kepong.
Zhang XC, Nooteboom HP, Kato M. 2013b. Selaginellaceae. In: Wu ZY, Raven PH, Hong DY (eds). Flora of China. Vol. 2-3. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.
Zuur AF, Ieno EN, Elphick CS. 2010. A protocol for data exploration to avoid common statistical problems. Methods Ecol Evol 1 (1): 3-14. https://doi.org/10.1111/j.2041-210X.2009.00001.x. DOI: https://doi.org/10.1111/j.2041-210X.2009.00001.x