Morphometric diversity of wild poisonous arbila (Phaseolus lunatus) germplasm from Timor and Sumba Islands, Indonesia
Main Article Content
Abstract
Abstract. Simamora AV, Mukkun L, Hahuly MV, Nenotek PS, Mau YS, Kaho NR, Seran YN, Arung ET, Hosang EY. 2026. Morphometric diversity of wild poisonous arbila (Phaseolus lunatus) germplasm from Timor and Sumba Islands, Indonesia. Biodiversitas 27 (5): d270511. https://doi.org/10.13057/biodiv/d270511. Poisonous arbila (Phaseolus lunatus) is an underutilized legume traditionally consumed after detoxification by dryland communities in East Nusa Tenggara, Indonesia. Despite its importance as an emergency food resource, information on its phenotypic variation remains limited. This study aimed to characterize seed morphometric variation in wild and semi-managed arbila germplasm collected from six locations across Timor and Sumba Islands. A total of 108 accessions were evaluated using twelve seed traits, including eight qualitative characters (seed shape, seed coat color and pattern, hilum shape, testa texture, and eye color) and four quantitative characters (seed length, width, thickness, and 100-seed weight). Qualitative traits showed high polymorphism, particularly in seed coat pigmentation and patterning, while quantitative traits exhibited wide ranges (seed length: 10.33-23.70 mm; 100-seed weight: 25.40-56.50 g). Multivariate analyses (principal component analysis and hierarchical clustering) indicated that the first four principal components explained 91.6% of the total variance, with seed size–related traits dominating the primary axis (PC1 = 65.2%). Clustering results identified one major group comprising most accessions and a small number of divergent accessions, consistent with a continuous phenotypic gradient rather than discrete group separation. These results highlight considerable morphometric diversity in arbila and indicate that phenotypic differentiation is primarily structured along this gradient. The findings provide a useful basis for germplasm conservation and highlight the potential of arbila for selection, breeding, and future studies on domestication and genetic diversity.
Article Details
Issue
Section

This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License.
References
Ambika, Aski MS, Gayacharan, Hamwieh A, Talukdar A, Gupta SK, Sharma BB, Joshi R, Upadhyaya HD, Singh K, Kumar R. 2022. Unraveling origin, history, genetics, and strategies for accelerated domestication and diversification of food legumes. Front Genet 13: 932430. https://doi.org/10.3389/fgene.2022.932430.
Antriyandarti E, Barokah U, Rahayu W, Asami A, Laia DH, Sari LD, Pranadita NE, Melati NSK. 2024. Resilience of dryland farm households in the mountains and their adaptability to environmental and social challenges. Environ Chall 17: 101037. https://doi.org/10.1016/j.envc.2024.101037.
Atkinson D. 1994. Temperature and organism size: A biological law for ectotherms?. Adv Ecol Res 25: 1-58. https://doi.org/10.1016/S0065-2504(08)60212-3.
Barros ES, Sarigu M, Lallai A, Nicolau JPB, Benedito CP, Bacchetta G, Torres SB. 2024. Phenotypic identification of landraces of Phaseolus lunatus L. from the Northeastern Region of Brazil using morpho-colorimetric analysis of seeds. Horticulturae 10 (9): 948. https://doi.org/10.3390/horticulturae10090948.
Bohra A, Tiwari A, Kaur P, Ganie SA, Raza A, Roorkiwal M, Mir RR, Fernie AR, Smýkal P, Varshney RK. 2022. The key to the future lies in the past: Insights from grain legume domestication and improvement should inform future breeding strategies. Plant Cell Physiol 63 (11): 1554-1572. https://doi.org/10.1093/pcp/pcac086.
Bria EJ, Suharyanto E, Purnomo. 2019. Variability and intra-specific classification of lima bean (Phaseolus lunatus L.) from Timor Island based on morphological characters. J Trop Biodivers Biotechnol 4 (2): 62-71. https://doi.org/10.22146/jtbb.42547.
Chacón-Sánchez MI, Martínez-Castillo J. 2017. Testing domestication scenarios of lima bean (Phaseolus lunatus L.) in Mesoamerica: Insights from genome-wide genetic markers. Front Plant Sci 8: 1551. https://doi.org/10.3389/fpls.2017.01551.
Chen X, Zhou G, Pang J, Srinives P. 2021. Editorial: Domestication of agronomic traits in legume crops. Front Genet 12: 707600. https://doi.org/10.3389/fgene.2021.707600.
Dave K, Kumar A, Dave N, Jain M, Dhanda PS, Yadav A, Kaushik P. 2024. Climate change impacts on legume physiology and ecosystem dynamics: A multifaceted perspective. Sustainability 16 (14): 6026. https://doi.org/10.3390/su16146026.
Fagbédji RF, Djedatin LG, Nanoukon C, Kinhoegbe G, Havivi A, Yédomonhan H, Agbangla C. 2023. Insight into the genetic diversity of cultivated lima bean (Phaseolus lunatus L.) in Benin. Am J Mol Biol 13 (1): 32-45. https://doi.org/10.4236/ajmb.2023.131003.
Hammer Ø, Harper DAT, Ryan PD. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontol Electron 4 (1): 1-9. https://palaeo-electronica.org.
Heredia-Pech M, Chávez-Pesqueira M, Ortiz-García MM, Andueza-Noh RH, Chacón-Sánchez MI, Martínez-Castillo J. 2022. Consequences of introgression and gene flow on the genetic structure and diversity of lima bean (Phaseolus lunatus L.) in its Mesoamerican diversity area. PeerJ 10: e13690. https://doi.org/10.7717/peerj.13690.
International Board for Plant Genetic Resources (IBPGR). 1982. Lima Bean Descriptors. IBPGR Secretariat, Rome.
Islam NS, Dhaubhadel S. 2023. Proanthocyanidin biosynthesis and postharvest seed coat darkening in pinto bean. Phytochem Rev 24 (4): 2445-2461. https://doi.org/10.1007/s11101-023-09895-8.
Jaiswal SK, Dakora FD. 2024. Seed-coat pigmentation plays a crucial role in partner selection and N2 fixation in legume-root-microbe associations in African soils. Plants 13 (11): 1464. https://doi.org/10.3390/plants13111464.
Ku YS, Contador CA, Ng MS, Yu J, Chung G, Lam HM. 2020. The effects of domestication on secondary metabolite composition in legumes. Front Genet 11: 581357. https://doi.org/10.3389/fgene.2020.581357.
Mau YS, Oematan SS, Arsa IGBA, Ndiwa ASS, Bere AC. 2025. Genetic diversity of local cowpea (Vigna unguiculata) germplasms of East Nusa Tenggara Province, Indonesia, revealed by seed characteristics. Biodiversitas 26 (3): 1279-1288. https://doi.org/10.13057/biodiv/d260327.
Mousseau TA. 1997. Ectotherms follow the converse of Bergmann’s rule. Evolution 51 (2): 630-632. https://doi.org/10.2307/2411138.
Naisali H, Witoyo JE, Utoro PAR. 2023. Local legumes from the dry land of East Nusa Tenggara: Diversity, nutritional composition, and their use in society: A literature study. Jurnal Ilmiah Teknologi dan Industri Pangan UNISRI 8 (2): 155-166. https://doi.org/10.33061/jitipari.v8i2.9054. [Indonesian]
Paipilla G, Escobar MC, Ovalle MF, Vanegas BS, García T, Chacón-Sánchez MI. 2025. Genome-wide discovery of short sequence repeat markers useful for genetic diversity studies in lima bean (Phaseolus lunatus L.). Sci Rep 16 (1): 1419. https://doi.org/10.1038/s41598-025-31509-w.
Penha JS, Oliveira GCX, Lopes ACA, Gomes RLF, Costa MF, Pinheiro JB, Zucchi MI, Assunção-Filho JR, Silvestre EA, Dequigiovanni G, Martínez-Castillo J. 2025. Genetic diversity and population structure of lima bean (Phaseolus lunatus L.) based on cpSSR markers. Braz J Biol 85: e298660. https://doi.org/10.1590/1519-6984.98660.
Plestenjak E, Neji M, Sinkovič L, Meglič V, Pipan B. 2025. Genomic insights into genetic diversity and seed coat color change in common bean composite populations. Front Plant Sci 15: 1523745. https://doi.org/10.3389/fpls.2024.1523745.
Purwanti E, Fauzi A. 2019. The morphological characteristics of Phaseolus lunatus L. in different areas of East Java, Indonesia. IOP Conf Ser Earth Environ Sci 276: 012017. https://doi.org/10.1088/1755-1315/276/1/012017.
Pusat Perlindungan Varietas Tanaman dan Perizinan Pertanian (PPVTPP). 2021. Panduan Penggunaan Aplikasi Pendaftaran Varietas Lokal dan Hasil Pemuliaan. PPVTPP, Indonesian Ministry of Agriculture, Jakarta. [Indonesian]
Puspita D, Palimbong S, Toy B, Notosoedarmo S. 2017. Identifikasi legum lokal di Pulau Timor yang berpotensi untuk pengembangan inovasi pangan lokal. Prosiding Seminar Nasional Pengembangan Sumber Daya Perdesaan dan Kearifan Lokal Berkelanjutan VII 324-335. Purwokerto, 17-18 November 2017. [Indonesian]
Quilichini TD, Gao P, Yu B, Bing D, Datla R, Fobert P, Xiang D. 2022. The seed coat's impact on crop performance in pea (Pisum sativum L.). Plants 11 (15): 2056. https://doi.org/10.3390/plants11152056.
Riptanti EW, Masyhuri M, Irham I, Suryantini A. 2019. The ability of dryland farmer households in achieving food security in the food-insecure areas of East Nusa Tenggara, Indonesia. AIMS Agric Food 5 (1): 30-45. https://doi.org/10.3934/agrfood.2020.1.30.
Sandrine MAD, Kevin KK, Clémence KL. 2020. Preliminary diversity assessment of lima beans (Phaseolus lunatus) cultivated in Côte d’Ivoire. Am J Plant Sci 11: 2059-2065. https://doi.org/10.4236/ajps.2020.1112145.
Sita K, Sehgal A, HanumanthaRao B, Nair RM, Vara Prasad PV, Kumar S, Gaur PM, Farooq M, Siddique KHM, Varshney RK, Nayyar H. 2017. Food legumes and rising temperatures: Effects, adaptive functional mechanisms specific to reproductive growth stage, and strategies to improve heat tolerance. Front Plant Sci 8: 1658. https://doi.org/10.3389/fpls.2017.01658.
Smýkal P, von Wettberg EJB, McPhee K. 2020. Legume genetics and biology: From Mendel’s pea to legume genomics. Intl J Mol Sci 21 (9): 3336. https://doi.org/10.3390/ijms21093336.
Souza JDS, Rodrigues LN, Ferreira LT, Unêda-Trevisoli SH, Nascimento NFFD. 2023. Future-ready crops, genetic variability in lima bean seeds. Acta Sci Agron 46 (1): e66767. https://doi.org/10.4025/actasciagron.v46i1.66767.
Tudor EP, Lewandrowski W, Krauss S, Veneklaas EJ. 2024. Local adaptation to climate inferred from intraspecific variation in plant functional traits along a latitudinal gradient. Conserv Physiol 12 (1): coae018. https://doi.org/10.1093/conphys/coae018.
Wang Z, Zhao W, Huang Y, Zhao P, Yang K, Wan P, Chu L. 2023. Progress in adzuki bean seed coat color studies. Plants 12 (18): 3242. https://doi.org/10.3390/plants12183242.
Zhang H, Mascher M, Abbo S, Jayakodi M. 2022. Advancing grain legumes domestication and evolution studies with genomics. Plant Cell Physiol 63 (11): 1540-1553. https://doi.org/10.1093/pcp/pcac062.
Zhang JY. 2024. Legume evolution: From wild ancestors to modern crops. Legume Genom Genet 15 (3): 93-104. https://doi.org/10.5376/lgg.2024.15.0011.