{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T05:16:43Z","timestamp":1787894203876,"version":"build-2784847793"},"reference-count":53,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T00:00:00Z","timestamp":1787702400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"The Development and Promotion of Science and Technology Talents Project"},{"DOI":"10.13039\/501100002808","name":"Carlsberg Foundation","doi-asserted-by":"publisher","award":["CF14-0245"],"award-info":[{"award-number":["CF14-0245"]}],"id":[{"id":"10.13039\/501100002808","id-type":"DOI","asserted-by":"publisher"}]},{"award":["CF14-0245"],"award-info":[{"award-number":["CF14-0245"]}],"id":[{"id":"https:\/\/ror.org\/01kpjmx04","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Fabaceae, the third-largest family of angiosperms, occurs in diverse habitats, including limestone areas characterized by karst landscapes, and in Thailand, by frequent human disturbance due to activities such as agriculture and mining. These disturbances affect plant species, which highlights the importance of diversity studies and the use of morphology and anatomy for identification. A study of Fabaceae diversity in a limestone area in Thung Song District, southern Thailand, were conducted along three survey routes. A total of 23 genera and 30 species were encountered, including 2 vulnerable, 1 rare, and 8 introduced species. Morphological, phenological, ecological, and conservation status data were recorded. The leaf epidermis of 23 species was observed under a light microscope. Stomatal size and stomatal index were measured. Phanera tubicalyx, an endemic and vulnerable species restricted to limestone habitats, had multiple papillae on its epidermal cells, which may reduce water loss and enhance photosynthetic efficiency. Both hypostomatic and amphistomatic leaves were observed. Stomatal size ranged from 114.63 \u00b1 17.57 \u03bcm2 to 608.88 \u00b1 16.42 \u03bcm2, while the stomatal index varied from 0.85 \u00b1 0.78 to 23.86 \u00b1 0.16. Peculiar paired crystals and multiple papillae were found in Canavalia cathartica and P. tubicalyx, respectively. These and other morphological and leaf epidermal characteristics were useful for species identification. Human disturbances from agriculture and mining were common in the study area. In conclusion, this study documented 30 Fabaceae species and demonstrated the usefulness of morphological and leaf epidermal characteristics for species identification and conservation.<\/jats:p>","DOI":"10.3390\/plants15172605","type":"journal-article","created":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T15:04:32Z","timestamp":1787756672000},"page":"2605","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Diversity, Leaf Anatomy, and Conservation of Fabaceae in a Thai Limestone Area"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-2986-6440","authenticated-orcid":false,"given":"Wichuda","family":"Manorin","sequence":"first","affiliation":[{"name":"Department of Biology, Section for Ecoinformatics & Biodiversity, Aarhus University, Ny Munkegade 114-116, DK-8000 Aarhus, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1746-1439","authenticated-orcid":false,"given":"Ponprom","family":"Pisuttimarn","sequence":"additional","affiliation":[{"name":"Queen Sirikit Botanic Garden, The Botanical Garden Organization, Chiang Mai 50180, Thailand"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5887-5115","authenticated-orcid":false,"given":"Suparman","family":"Suparman","sequence":"additional","affiliation":[{"name":"Department of Biology Education, Faculty of Teacher Training and Education, Universitas Khairun. Jl. Pertamina Kampus II Unkhair Gambesi, Ternate 97728, North Maluku, Indonesia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Henrik","family":"Balslev","sequence":"additional","affiliation":[{"name":"Department of Biology, Section for Ecoinformatics & Biodiversity, Aarhus University, Ny Munkegade 114-116, DK-8000 Aarhus, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6435-3740","authenticated-orcid":false,"given":"Witsanu","family":"Saisorn","sequence":"additional","affiliation":[{"name":"School of Science, Walailak University, Thasala, Nakhon Si Thammarat 80161, Thailand"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,26]]},"reference":[{"key":"ref_1","unstructured":"LPWG (the Legume Phylogeny Working Group) (2017). A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny. Taxon, 66, 44\u201377."},{"key":"ref_2","unstructured":"Lewis, G., Schrire, B., Mackinder, B., and Lock, M. (2005). Legumes of the World, Royal Botanic Gardens, Kew."},{"key":"ref_3","unstructured":"Sepngang, B.K., Muel, F., Smadja, T., Stauss, W., Stute, I., Simmen, M., and Mergenthaler, M. (2020). Report on Legume Markets in the EU, Deliverable D3.1 of the Eu-Project LegValue, Fachhochschule S\u00fcdwestfalen."},{"key":"ref_4","first-page":"012","article-title":"Soybean (Glycine max L.) isoflavones: Chemical composition and its chemometrics-assisted extraction and authentication","volume":"11","author":"Riswanto","year":"2021","journal-title":"J. Appl. Pharm."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"M\u00e9ndez-L\u00f3pez, L.F., de Le\u00f3n, D.S., Lomel\u00ed, M.L., Gonz\u00e1lez-Mart\u00ednez, B.E., and V\u00e1zquez-Rodr\u00edguez, J.A. (2022). Phytochemicals from Vicia faba Bean as ligands of the aryl hydrocarbon receptor to regulate autoimmune diseases. Front. Nutr., 9.","DOI":"10.3389\/fnut.2022.790440"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Begum, N., Khan, Q.U., Liu, L.G., Li, W., Liu, D., and Haq, I.U. (2023). Nutritional composition, health benefits and bio-active compounds of chickpea (Cicer arietinum L.). Front Nutr., 10.","DOI":"10.3389\/fnut.2023.1218468"},{"key":"ref_7","first-page":"1","article-title":"Leguminosae-Caesalpinioideae","volume":"Volume 4","author":"Smitinand","year":"1984","journal-title":"Flora of Thailand"},{"key":"ref_8","first-page":"131","article-title":"Leguminosae-Mimosoideae","volume":"Volume 4","author":"Smitinand","year":"1985","journal-title":"Flora of Thailand"},{"key":"ref_9","first-page":"221","article-title":"Leguminosae-Papilionoideae","volume":"Volume 4","author":"Chayamarit","year":"2018","journal-title":"Flora of Thailand"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"138","DOI":"10.20531\/tfb.2018.46.2.05","article-title":"The genus Campylotropis (Leguminosae) in Thailand","volume":"46","author":"Satthaphorn","year":"2018","journal-title":"Thai For. Bull. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"23","DOI":"10.58837\/tnh.15.1.103085","article-title":"Taxonomic studies on the genus Phyllodium Desv. (Leguminosae) in Thailand","volume":"15","author":"Saisorn","year":"2015","journal-title":"Trop. Nat. Hist."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"32","DOI":"10.58837\/tnh.18.1.117467","article-title":"The genus Hylodesmum H.Ohashi & R.R.Mill (Leguminosae-Papilionoideae) in Thailand","volume":"18","author":"Saisorn","year":"2018","journal-title":"Trop. Nat. Hist."},{"key":"ref_13","first-page":"788","article-title":"The genus Codariocalyx Hassk. (Leguminosae-Papilionoideae) in Thailand: Taxonomy and anatomy of leaf and stem","volume":"41","author":"Saisorn","year":"2019","journal-title":"Songklanakarin J. Sci. Technol."},{"key":"ref_14","first-page":"72\u221288","article-title":"The genus Ototropis (Leguminosae-Papilionoideae) in Thailand","volume":"20","author":"Saisorn","year":"2020","journal-title":"Trop. Nat. Hist."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"41","DOI":"10.58837\/tnh.21.1.247326","article-title":"Taxonomy of Dendrolobium (Leguminosae) in Thailand","volume":"21","author":"Saisorn","year":"2021","journal-title":"Trop. Nat. Hist."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"231","DOI":"10.11646\/phytotaxa.573.2.4","article-title":"A taxonomic revision of two genera, Pleurolobus and Sohmaea (Leguminosae) in Thailand and Indo-China","volume":"573","author":"Saisorn","year":"2022","journal-title":"Phytotaxa"},{"key":"ref_17","first-page":"39","article-title":"The study of flowering herbs on base area of Khao Wong Phra Chan limestone outcrop, Padang Besar subdistrict, Sadao district, Songkhla province","volume":"9","author":"Satthaphorn","year":"2017","journal-title":"Thai J. Bot."},{"key":"ref_18","first-page":"45","article-title":"Species diversity of subfamily Papilionoideae (Leguminosae) in southern Thailand","volume":"12","author":"Leeratiwong","year":"2020","journal-title":"Thai J. Bot."},{"key":"ref_19","first-page":"1597","article-title":"Review: Biodiversity of forests over limestone in Southeast Asia with emphasis on the Philippines","volume":"21","author":"Tolentino","year":"2020","journal-title":"Biodiversitas"},{"key":"ref_20","unstructured":"Poopath, M., and Tetsana, N. (2018). Limestone Flora in Phu Khieo-Nam Nao, Office of the Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation. [2nd ed.]. (In Thailand)."},{"key":"ref_21","unstructured":"Santisuk, T., Chayamarit, K., Pooma, R., and Suddee, S. (2006). Thailand Red Data: Plants."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"98","DOI":"10.3767\/000651914X684862","article-title":"Derris solorioides (Fabaceae), a new limestone species with true-paniculate inflorescences from North-Central Thailand","volume":"59","author":"Sirichamorn","year":"2014","journal-title":"Blumea"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"62","DOI":"10.20531\/tfb.2018.46.1.09","article-title":"Erythrina calcicola sp. nov. (Fabaceae) from Thailand","volume":"46","author":"Tetsana","year":"2018","journal-title":"Thai For. Bull. Bot."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"4","DOI":"10.20531\/tfb.2018.46.1.02","article-title":"Sophora huamotensis, a new species of Sophora (Fabaceae-Papilionoideae-Sophoreae) from Thailand","volume":"41","author":"Mattapha","year":"2018","journal-title":"Thai For. Bull. Bot."},{"key":"ref_25","first-page":"1","article-title":"Environmental impacts of limestone-shell rock mining and processing: Assessment and monitoring approaches","volume":"58","author":"Bukayev","year":"2025","journal-title":"Acta Innov."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1111\/sjtg.12203","article-title":"Impacts of artisanal gold mining systems on soil and woody vegetation in the semi-arid environment of northern Ethiopia","volume":"38","author":"Meaza","year":"2017","journal-title":"Singap. J. Trop. Geogr."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1186\/s40068-020-00171-8","article-title":"Effects of stone mining on woody plant species diversity and selected soil properties in northern Ethiopia","volume":"9","author":"Belay","year":"2020","journal-title":"Environ. Syst. Res."},{"key":"ref_28","unstructured":"Chamchumroon, V., Suphuntee, N., Tetsana, N., Poopath, M., and Tanikkool, S. (2017). Threatened Plants in Thailand, Department of National Parks, Wildlife and Plant Conservation. [1st ed.]."},{"key":"ref_29","unstructured":"Wichiennopparat, W., Visaratana, T., Kuldelok, C., Wannapinit, M., and Luangviriyasaeng, V. (2008, January 17\u201320). Effects of Soil Improvement on Growth of Acacia Plantation in Nakhon Ratchasima, Thailand. Proceedings of the FORTROP II: Tropical Forestry Change in a Changing World, Commercial Plantation Forestry, Kasetsart University, Bangkok, Thailand."},{"key":"ref_30","unstructured":"Santisuk, T. (2012). Forest of Thailand, Office of the Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation. [3rd ed.]. (In Thailand)."},{"key":"ref_31","first-page":"113","article-title":"Species diversity of family Leguminosae in Ngao Waterfall National Park, Ranong province","volume":"12","author":"Pinjom","year":"2020","journal-title":"Thai J. Bot."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1139\/cjb-2022-0060","article-title":"Micromorphology of leaflets of Senna series Bacillares (Leguminosae) and its taxonomic significance","volume":"100","author":"Souto","year":"2022","journal-title":"Botany"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"43","DOI":"10.3329\/bjpt.v21i1.19264","article-title":"Stomatal and trichome diversity in Senna Mill. from Bangladesh","volume":"21","author":"Begum","year":"2014","journal-title":"Bangladesh J. Plant Taxon."},{"key":"ref_34","first-page":"1","article-title":"Leaf epidermal anatomy of some Crotalaria L. (Fabaceae) species in Thailand","volume":"24","author":"Ninkaew","year":"2024","journal-title":"Trop. J. Hist."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"51","DOI":"10.3897\/phytokeys.201.85154","article-title":"Micromorphological leaf epidermal traits as potential taxonomic markers for infrageneric classification of Oxytropis (Fabaceae)","volume":"201","author":"Zhao","year":"2022","journal-title":"PhytoKeys"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"237","DOI":"10.11646\/phytotaxa.606.4.1","article-title":"Taxonomic implications of leaf epidermal characteristics in subfamily Caesalpinioideae (Leguminosae)","volume":"606","author":"Pandey","year":"2023","journal-title":"Phytotaxa"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2688","DOI":"10.1111\/nph.71399","article-title":"A drought stress-induced MYB transcription factor regulates pavement cell shape in leaves of European aspen (Populus tremula)","volume":"251","author":"Liu","year":"2026","journal-title":"New Phytol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1016\/j.bjp.2018.04.012","article-title":"Leaf morphoanatomy of \u201cmoror\u00f3\u201d (Bauhinia and Schnella, Fabaceae)","volume":"28","author":"Pereira","year":"2018","journal-title":"Rev. Bra. Farmacogn."},{"key":"ref_39","first-page":"3","article-title":"Taxonomic value of the leaf micro-morphology and quantitative phytochemistry of Clitoria ternatea and Centrosema molle (Papilionoideae, Fabaceae)","volume":"20","author":"Chukwuma","year":"2014","journal-title":"Phytol. Balc."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1086\/337019","article-title":"Leaflet anatomy of Phaseoleae (Leguminosae: Papilionoideae) and its relation to taxonomy","volume":"139","author":"Lackey","year":"1987","journal-title":"Bot. Gaz."},{"key":"ref_41","first-page":"1757","article-title":"Microscopic characteristics and leaf constant numbers of selected Derris species in Thailand","volume":"12","author":"Phairoh","year":"2021","journal-title":"Int. J. Pharm. Sci. Res."},{"key":"ref_42","first-page":"232","article-title":"Foliar epidermal anatomy and pollen morphological studies of Mucuna pruriens (Linn.) DC. and Mucuna poggei Taub. in Jos, Nigeria","volume":"3","author":"Adeniji","year":"2024","journal-title":"Agric. Food Nat. Resour. J."},{"key":"ref_43","first-page":"9784","article-title":"Micromorphological study of some species of Papilionoideae from Nigeria","volume":"7","author":"Owolabi","year":"2018","journal-title":"Int. J. Curr. Adv. Res."},{"key":"ref_44","first-page":"126","article-title":"Leaf epidermal anatomy of medicinal plants in Lamtakhong Research Station, Nakhon Ratchasima Province","volume":"31","author":"Phromprasit","year":"2026","journal-title":"Burapha Sci. J."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1007\/s00425-026-04999-9","article-title":"Functional and evolutionary significance of the unique lime-secreting hydathodes and amphistomatic leaves in Saxifraga (Saxifragaceae)","volume":"263","author":"Tkach","year":"2026","journal-title":"Planta"},{"key":"ref_46","unstructured":"Metcalfe, C.R., and Chalk, L. (1957). Anatomy of the Dicotyledons: Leaves, Stem, and Wood in Relation to Taxonomy with Notes Economic Uses, Clarendon Press."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"151790","DOI":"10.1016\/j.flora.2021.151790","article-title":"Comparison of petiole anatomy in Flemingia and its potential for delimitation of species","volume":"278","author":"Thacker","year":"2021","journal-title":"Flora"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Pitaloka, M.K., Harrison, E.L., Hepworth, C., Wanchana, S., Toojinda, T., Phetluan, W., Brech, R.A., Narawatthana, S., Vanavichit, A., and Gray, J.E. (2021). Rice stomatal mega-papillae restrict water loss and pathogen entry. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.677839"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1186\/s12284-023-00676-7","article-title":"Small papillae regulated by SPD25 are critical for balancing photosynthetic CO2 assimilation and water loss in rice","volume":"16","author":"Zhu","year":"2023","journal-title":"Rice"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"152989","DOI":"10.1016\/j.flora.2026.152989","article-title":"Leaf anatomy and micromorphology of Otatea (Bambuseae: Guadinae): Adaptations to arid environments","volume":"340","author":"Terrazas","year":"2026","journal-title":"Flora"},{"key":"ref_51","unstructured":"Thiers, B.M. (2026, August 14). (Updated Continuously). Index Herbariorum. Available online: https:\/\/sweetgum.nybg.org\/science\/ih\/."},{"key":"ref_52","unstructured":"Metcalfe, C.R., and Chalk, L. (1979). Anatomy of the Dicotyledons: Systematic Anatomy of Leaf and Stem, with a Brief History of the Subject, Clarendon Press. [2nd ed.]."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1111\/j.1469-8137.2011.03935.x","article-title":"Photosynthetic pathway and ecological adaptation explain stomatal trait diversity amongst grasses","volume":"193","author":"Taylor","year":"2012","journal-title":"New Phytol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2605\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T04:17:18Z","timestamp":1787890638000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,26]]},"references-count":53,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15172605"],"URL":"https:\/\/doi.org\/10.3390\/plants15172605","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,26]]}}}