Diversity and distribution of the Huastec Mayan medicinal plants: hotspots for bioprospecting and conservation

Biodiversity Data Journal 13: e170091 doi: 10.3897/BDJ.13.e170091 Research Article Diversity and distribution of the Huastec Mayan medicinal plants: hotspots for bioprospecting and conservation Madeleyne Cupido , José Arturo De-Nova , Virginia Gabriela Cilia-López , Francisco Javier PérezVázquez , Pablo Delgado-Sánchez ‡ Universidad Autónoma de San Luis Potosí, San Luis Potosí, Mexico Corresponding author: Madeleyne Cupido ([email protected]), José Arturo De-Nova ([email protected]) Academic editor: Gianmarco Tavilla Received: 27 Aug 2025 | Accepted: 24 Nov 2025 | Published: 08 Dec 2025 Citation: Cupido M, De-Nova JA, Cilia-López VG, Pérez-Vázquez FJ, Delgado-Sánchez P (2025) Diversity and distribution of the Huastec Mayan medicinal plants: hotspots for bioprospecting and conservation. Biodiversity Data Journal 13: e170091. https://doi.org/10.3897/BDJ.13.e170091 Abstract The multi-ethnic biocultural region Huasteca in Mexico has a variety of practices, including the use of traditional plant-based medicine. Here, we analyse the diversity and distribution of the medicinal plants used by the Tenek culture, to identify hotspots for bioprospecting and conservation. We describe their diversity, growth forms, origin and endemism, classes of diseases, parts used and conservation risk status. A total of 468 plant species used for medicinal purposes were recorded, from 113 families and 350 genera. More than 50% of the species richness is concentrated in 10 families and 11 genera. A total of 418 species are native, 23 of them are endemic, while 50 are nonnative. These species are obtained from one or more habitats, including 356 from the wild, 237 from home gardens, 111 from milpas, 20 from sugarcane plantations and 16 from disturbed habitats. The most common growth form is herbaceous and the most commonly used parts are leaves. Geographic hotspots with high species richness, disease classes diversity and high ethnomedicinal index values occur in Aquismón, San Antonio and Tantoyuca. Our results show that the biocultural region Huasteca is a reservoir of current benefits and future option values for bioprospecting and preserving of ‡ ‡ ‡ ‡ ‡ © Cupido M et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. biocultural heritage. Further studies should employ phylogenetic approaches to elucidate evolutionary patterns and processes implied selection of medicinal plants by local people of medicinal plants. This will help identify lineages with phytochemicals, improve bioprospecting and ensure benefits for local people. As knowledge holders, they contribute to the conservation of medicinal plants. Keywords biodiversity, ethnomedicine, Huasteca, risk category, traditional knowledge Introduction The use of plants has been an integral part of the traditional systems of medicine for centuries for humans around the world (Cox 1994, Roopashree et al. 2024). Plants have played an important role in traditional medicine in the world and, currently, ca. 23,842 plant species have been recorded as medicinal in the world (Pironon et al. 2024). Due to overexploitation, expansion of alien invasive species and climate change, medicinal plants are facing an increasing risk of habitat destruction and extinction (Huang et al. 2011, Huang et al. 2012, Pandey et al. 2018). Currently, identifying hotspots (critical areas for preservation) is a useful method for conservation efforts, because it helps select priority areas for conservation and improves the sustainable use of biodiversity (Myers et al. 2000). This identification of biodiversity hotspots is highly recommended in order to take into account multiple conservation indicators, such as the diversity of species found in a region, endemic species, threatened species and nationally protected species (Forest et al. 2007, Huang et al. 2012), as well as the diversity of human disease types found in a given region. However, despite being the second country in terms of medicinal plant use, Mexico has not carried out this type of study yet (Muñetón-Pérez 2009, Loraine and Mendoza-Espinoza 2010, Cruz-Pérez et al. 2021). Currently, traditional medicine in Mexico has a strong ancestral tradition in the use of medicinal plants. Between 3,000 and 4,500 species are used (Bye 1993, Argueta et al. 1994, CONABIO 2006, Muñetón-Pérez 2009) by more than 56 ethnic groups (Martin et al. 2011). Their ancestral presence and interactions with the environment and local communities have allowed the development of multiple forms of knowledge and the use of natural resources, especially plants, to address various health problems. The biocultural region Huasteca, located in northeastern Mexico (Fig. 1), has been described as multi-ethnic and a biocultural region (Alcorn 1984), where different practices have been developed over time, such as traditional medicine, which is part of the local population's lives and used to treat different diseases and symptoms (Domínguez and Alcorn 1985). The most important ethnic group is the Tenek people, a descendant lineage of the ancient Maya with a rich biocultural heritage (Alcorn 1984, De-Nova et al. 2024). This heritage is the result of the territory's complex geographical and geological configuration of their territory, in the so-called Mexican transition zone between the Nearctic and Neotropical biogeographic regions. The landscape of their plant diversity is estimated to contain ca. 1,858 species of vascular plants, mainly with a tropical affinity, that inhabit the different types of seasonally dry 2Cupido M et al tropical forests (De-Nova et al. 2024). It has recently been indicated from an evolutionary framework that the Huastec Mayan (Tenek) useful plants represent a vast source of current benefits and future option values (i.e. options to benefit from future uses of biodiversity) for human well-being (De-Nova et al. 2024). Traditional medicine, based on plants, contributes significantly to the health of the population, especially in indigenous communities (WHO 2013). A high proportion of information on traditional medicine has been recorded in different parts of the world, but these data are often scattered in isolated studies on a particular species or lost, as this knowledge is passed down orally from generation to generation. Our objective was to analyse the diversity and distribution of the Huastec Mayan medicinal plants used by the Tenek in the biocultural region Huasteca to spatially identify ethnomedicinal hotspots for bioprospecting and conservation. We describe taxonomic diversity, growth forms, origin and endemism, classes of diseases, parts used and risk categories. We also discuss the relevance of these plants as a source of option values, particularly in the context of the search for pharmacological compounds. Material and methods Study area The study area is the Tenek territory in the biocultural region Huasteca, located in northeastern Mexico, mainly in the States of Veracruz, Hidalgo and San Luis Potosi, including Figure 1. Tenek territory and their municipalities in States of San Luis Potosí and Veracruz, Mexico. The purple colour gradient illustrates the orography.  Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 3 high agrobiodiversity, multiple uses and option values of biodiversity for human wellbeing (De-Nova et al. 2024). The types of vegetation range from subtropical and tropical forests at lower altitudes (50-800 m) to cloud and pine-oak forests at higher altitudes (600-2,000 m). This region is located between the physiographic provinces of the coastal plain of the Gulf of Mexico and the Sierra Madre Oriental and corresponds to the hydrological region of the Panuco River (Fig. 1). The environments in the region provide important abiotic conditions that promote a high diversity of vegetation types, making it one of the priority inland regions of Mexico. Its flora includes 1,858 species (742 with some ethnobotanic use), mainly from tropical affinity that are elements of tropical and seasonally dry tropical forests, but also from temperate pine and oak forests and relicts of cloud forests (De-Nova et al. 2024). This study focuses on the Tenek people and their territory, who have inhabited the region and have managed and shaped the ecosystem for more than 3,000 years (Alcorn 1984). Over the last century, however, changes in land use caused by the commercial expansion of agriculture and livestock have significantly impacted the environment. In critical areas of the Huasteca, most of the original vegetation has disappeared and been replaced by areas of extensive agriculture and livestock (Reyes Hernández et al. 2006). Database An updated database of plant species used in traditional medicine by the Tenek culture was compiled, based on the ethnobotanical atlas published by Alcorn (1984). This atlas recorded information and data on the various uses of plants in the region. We included information on the medicinal plants listed in this atlas, including additional data on each species. This information was recently published in the catalogue of medicinal plants of the biocultural region Huasteca (Cupido et al. 2023). Nomenclature was updated using the Taxonomic Name Resolution Service (TNRS), version 5.2 (Boyle et al. 2013) available in Plants of the World Online (POWO 2025). The database was developed with the following fields: family, species, common local names, habitat, origin (native or nonnative), growth form, extinction risk category according to the Mexican NOM-059SEMARNAT-2010 (SEMARNAT 2010), as assessed by authorities and the Red List of Threatened Species (IUCN 2023). Disease classification The diseases and symptoms treated with Tenek medicinal plants correspond to a complex knowledge system that includes ancient, inherited practices, influenced by the West since the conquest. These were classified according to the ICD-11 (2024) of the World Health Organisation as follow: certain infectious or parasitic diseases (CIPD), diseases of the blood or blood-forming organs (DBBO), diseases of the circulatory system (DCS), diseases of the digestive system (DGS), diseases of the ear or mastoid process (DEMP), diseases of the visual system (DVS), diseases of the genitourinary system (DGUS), diseases of the musculoskeletal system or connective tissue (DMSC), diseases of the nervous system (DNS), diseases of the respiratory system (DRS), diseases of the skin (DS), endocrine, nutritional or metabolic diseases (ENMD), factors influencing health 4Cupido M et al status or contact with health services (FIHS), injury, poisoning or certain other consequences of external causes (IPEC), mental, behavioural or neurodevelopmental disorders (MBND), neoplasms (NE), pregnancy, childbirth or the puerperium (PCP) and symptoms, signs or clinical findings not elsewhere classified (NEC). Additionally, we included culture-bound syndromes (CBS). Geographical distribution A second database with geographical data was compiled with records of all the vascular plant species documented in the municipalities, historically occupied by the Tenek people (Tenek territory), extracted from the Herbarium SLPM database (SLPM 2025) and the GBIF platform (GBIF 2024), in order to identify hotspots that represent critical geographical areas for preserving biocultural heritage (Cámara-Leret and Bascompte 2021). The GBIF data include specimens from 40 different herbaria in Mexico and around the world, as well as direct observations curated by experts. Particular attention was paid to our institutional herbarium, SLPM, since it contains the largest collection of botanical vouchers from the region. Additionally, a copy of all specimens collected Alcorn (1984) is deposited there. An extraction was made for only the medicinal species included in the first database, including geographic coordinates and voucher information. No exclusion criteria were applied, except for specimens with gaps in relevant distribution or taxonomic information, as well as geographical inconsistencies. The years of the collection range from 1930 to 2024; however, 87% of the collection is from 1970 to 2024. Then, we assigned the classes of diseases of each species indicated in our first database (Suppl. material 1) and these were assigned to each geographical record. We used QGIS v. 2.16.3 (QGIS 2025) to visualise the geographical distribution of medicinal plant species richness (SR) and number of categories covered in grid cells of 0.045 decimal degrees (ca. 5 km ). To spatially identify ethnomedicinal hotspots highlighting the clustering of medicinal plants attending a wide number of classes of diseases, we calculated an ethnomedicinal index (EI) as: where n is the number of classes of diseases treated for the i-th species in a grid cell and N is the total number of classes of diseases treated for all species in the region (here 19 classes). Results Taxonomic richness and diversity We recorded a total of 468 plant species used in the Huastec Mayan traditional medicine in the Tenek territory, belonging to 113 families and 350 genera (Suppl. material 1 and Suppl. material 2). The families with the highest number of useful species (Fig. 2A) are 2 Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 5 Fabaceae (45 species), Asteraceae (30), Euphorbiaceae (25), Malvaceae (23), Solanaceae (19), Poaceae (15), Acanthaceae (13), Lamiaceae (13), Apocynaceae (10) and Verbenaceae (10), representing 43.37% of the total richness. The remaining 102 families include fewer than 10 species. The genera with the highest number of species (Fig. 2B) are Euphorbia L. (9), Croton L. (6) and Solanum L. (5). A total of 418 species are native (23 endemic) and 50 are non-native (Suppl. material 1 and Suppl. material 2). The most common habitats where medicinal plants are collected and obtained (Fig. 3) are in the wild (356 species), in home gardens (237) and in milpa (111). The most used growth forms (Fig. 4A) are herbaceous (228 species), trees (100) and shrubs (92). Growth forms such as rosette, creeping and epiphyte have less medicinal use. The most used parts (Fig. 4B) are leaves (201 species), followed by roots (136) and bark (69). A total of 141 species are considered in the IUCN Red List: 128 Least Concern, five Data Deficient, four Endangered, two Vulnerable, one Critically Endangered and one Near Threatened and three in the NOM-059-SEMARNAT-2010: two Threatened and one Subject to Special Protection (Suppl. material 1). Disease classification Table 1 shows the number of diseases and symptoms, as well as the number of species used for mitigation and treatment. The five disease classes with the highest values are: 1) DS, with 65 different diseases and symptoms treated by 177 species; 2) NEC with 56 Figure 2. Highest richness of medicinal plant species in the Tenek territory by families (A) and genera (B).  6Cupido M et al different diseases and symptoms and treated by 270 species; 3) DGS with 44 different diseases and symptoms treated by 161 species; 4) CBS with 40 different diseases and symptoms treated by 137 species; 5) CIPD with 30 different diseases and symptoms and 135 treated by species. A total of 259 species are used to treat more than three disease classes, 88 treat only two and 121 treat only one. Figure 3. Number of medicinal plants species recorded by habitat type in the Tenek territory.  Figure 4. Species richness of medicinal plants in the Tenek territory growth forms (A) and used parts (B).  Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 7 Disease class Diseases and symptoms No. species Certain infectious or parasitic diseases (CIPD) 30 135 Diseases of the blood or blood-forming organs (DBBO) 11 24 Diseases of the circulatory system (DCS) 14 32 Diseases of the digestive system (DGS) 44 161 Diseases of the ear or mastoid process (DEMP) 5 19 Diseases of the visual system (DVS) 11 15 Diseases of the genitourinary system (DGUS) 15 64 Diseases of the musculoskeletal system or connective tissue (DMSC) 26 52 Diseases of the nervous system (DNS) 4 16 Diseases of the respiratory system (DRS) 29 75 Diseases of the skin (DS) 65 177 Endocrine, nutritional or metabolic diseases (ENMD) 3 13 Factors influencing health status or contact with health services (FIHS) 3 10 Injury, poisoning or certain other consequences of external causes (IPEC) 19 74 Mental, behavioural or neurodevelopmental disorders (MBND) 15 26 Neoplasms (NE) 3 3 Pregnancy, childbirth or the puerperium (PCP) 23 71 Symptoms, signs or clinical findings, not elsewhere classified (NEC) 56 270 Culture-bound syndromes (CBS) 40 137 Geographical distribution The geographical database for medicinal plants in the Tenek territory includes 5,341 records for 432 species in 275 grid cells (Suppl. material 3). Species richness is concentrated mainly in five municipalities (Fig. 5A): San Antonio (304 species, with two grid cells including 92 and 274 species), Ciudad Valles (261 species, with four grid cells including 50, 58, 79 and 100 species), Aquismón (221 species, with one grid cell including 106 species) in San Luis Potosí and Tantoyuca (195 species, with two grid cells that include 93 and 108 species) and Tempoal (114 species with a grid cell with 59 species) in Veracruz. The municipalities with more classes of diseases covered by plants (Fig. 5B) are Aquismón (19 classes, with eight grid cells with more than 14 classes), Ciudad Valles (19 classes, with 28 grid cells with more than 14 classes), Huehuetlán (19 classes, with three grid cells with more than 14 classes), San Antonio (19 classes, with two grid cells with more than 14 classes), Tamuín (19 classes, with one grid cell with more than 14 classes), Tancanhuitz (19 classes, with three grid cells with more than 14 Table 1. Overview of the disease classes treated by the Huastec Mayan medicinal plants according to ICD-11 (2024) and the CBS category. 8Cupido M et al classes) and Tanlajás (19 classes, with one grid cell with more than 14 classes) in San Luis Potosí and Tancocó (19 classes, with one grid cell with more than 14 classes), Tantoyuca (19 classes, with two grid cells with more than 14 classes) and Tempoal (19 classes, with four grid cells with more than 14 classes) in Veracruz. Four ethnomedicinal hotspots (Fig. 6) are located in municipalities San Antonio (two grid cells; ethnomedicinal index of 46.7 and 15.3) Tantoyuca (two grid cells; ethnomedicinal index of 21.8 and 15.9), Ciudad Valles (two grid cells; ethnomedicinal index of 17.4 and 15.3) and Aquismón (one grid cell; ethnomedicinal index of 17.5). Figure 5. Medicinal plants in the Tenek territory by species richness (A) and classes of diseases (B). Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 9 Conclusions The relevance of cartography in exploring hotspots for bioprospecting and conserving natural ethnomedicinal resources that have known benefits and value for human wellbeing has not been widely studied. Our synthesis of the existing information on diversity and distribution of medicinal plant species in the Tenek territory reveals geographical zones with a high number of medicinal plant species, classes of diseases treated and ethnomedicinal indices. They represent hotspots for biodiversity conservation and nature option values for future benefits (Futuyama 1995, Faith et al. 2010). The municipalities with the main diversity of ethnomedicinal plant species and classes of diseases (San Antonio, Ciudad Valles, Aquismón, Tantoyuca and Tempoal) could be considered as a natural drugstore with a high variety of valuable options that represent the future benefits provided by biodiversity (Futuyama 1995, Faith et al. 2010). The EI (ethnomedicinal index) allows identifying particular hotspots with the greatest potential to develop bioprospecting activities in Aquismón, San Antonio, Ciudad Valles and Tantoyuca, where medicinal plants are assets highlighting the economic value of biodiversity (Costanza et al. 1997). Public policies and biodiversity conservation strategies are more likely to succeed in the long term if adequate methodologies are used to identify and allocate economic resources to benefit local inhabitants (Wang et al. 2022). The presence of natural protected areas in the biocultural region Huasteca, such as the Sierra del Abra Tanchipa Biosphere Reserve in Ciudad Valles, ensures the conservation of ethnomedicinal resources, where disturbance levels have been previously noted as low (De-Nova et al. 2018) and the participation of locals, government and enterprises have strengths and opportunities to combine bioprospecting and conserving biodiversity and traditional knowledge. Additionally, the future marketing and commercial trade of medicinal plants provide significant opportunities for local communities to profit through initiatives, based on ethical, equitable and sustainable principles. This profit stems from their traditional ethnobotanical knowledge and proximity to the resources, both of which are considered forms of biocultural heritage. Acknowledgements This research was funded by the Secretaría de Ciencia, Humanidades, Tecnología e Innovación (SECIHTI) (grant CF-320270 to J.A.D.N. and Doctoral Scholarship 1007054 to M.C.). The first author would like to thank the IIZD and PMPCA of the UASLP for their facilities and financial support for this research. Author contributions MC, JADN and VGCL conceived and designed the study. FJPV and PDS supervised, revised and reviewed the manuscript. All authors read and approved the final version of the manuscript. 16 Cupido M et al Conflicts of interest The authors have declared that no competing interests exist. References • Alcorn J (1984) Huastec Mayan ethnobotany. American Ethnologist • Alonso-Castro A, Maldonado-Miranda JJ, Zarate-Martinez A, del Rosario Jacobo-Salcedo M, et al. (2012) Medicinal plants used in the Huasteca Potosina, México. Journal of ethnopharmacology 143 (1): 292‑298. https://doi.org/10.1016/j.jep.2012.06.035 • Argueta A, Cano A, Rodarate ME (1994) Atlas de las Plantas Medicinales de la Medicina Tradicional Mexicana. Instituto Nacional Indigenista, Mexico, DF1786. • Barthas B (1996) De la selva al naranjal (transformaciones en la agricultura indígena en la Huasteca potosina). 1. Centro Francés de Estudios Mexicanos y Centroamericanos: Institut français de recherche scientifique pour le développement en coopération, México, D.F. • Beroni S, Goldson-Barnaby A, Petrea F (2023) A mini review of Desmodium incanum - An underutilized herb in Jamaica. Caribbean Journal of Science 11 (1): 08‑14. https:// doi.org/10.55434/CBI.2023.10102 • Bessada SF, Barreira JM, Oliveira MBPP (2015) Asteraceae species with most prominent bioactivity and their potential applications: A review. Industrial Crops and Products 76: 604‑615. https://doi.org/10.1016/j.indcrop.2015.07.073 • Blanckaert I, Paredes-Flores M, Espinosa-García F, Piñero D, Lira R (2012) Ethnobotanical, morphological, phytochemical and molecular evidence for the incipient domestication of Epazote (Chenopodium ambrosioides L.: Chenopodiaceae) in a semiarid region of Mexico. Genetic Resources and Crop Evolution 59 (4): 557‑573. https:// doi.org/10.1007/s10722-011-9704-7 • Boyle B, Hopkins N, Lu Z, Raygoza Garay JA, Mozzherin D, Rees T, Matasci N, Narro M, Piel W, Mckay S, Lowry S, Freeland C, Peet R, Enquist B (2013) The taxonomic name resolution service: an online tool for automated standardization of plant names. BMC Bioinformatics 14 (16). https://doi.org/10.1186/1471-2105-14-16 • Buenz E, Verpoorte R, and Bauer BA (2018) The ethnopharmacologic contribution to bioprospecting natural products. Annual Review of Pharmacology and Toxicology 58: 509‑530. https://doi.org/10.1146/annurev-pharmtox-010617-052703 • Bye R (1993) The role of humans in the diversification of plants in Mexico. In: Ramamorthy TP, Bye R, Lot A, Fa J (Eds) Biological diversity of Mexico. Origins and distribution. Oxford University Press. London, 707. pp. [ISBN 019506674X]. • Bye R, Linares E, Estrada E (1995) Biological diversity of medicinal plants in México. In: Springer (Ed.) Phytochemistry of Medicinal Plants. [ISBN 978-1-4899-1778-2]. https:// doi.org/10.1007/978-1-4899-1778-2_4 • Caballero-George C, Gupta M (2011) A quarter century of pharmacognostic research on Panamanian flora: a review. Planta Medica 77 (11): 1189‑1202. https://doi.org/10.1055/ s-0030-1271187 • Caballero J, Cortés L, Martínez-Alfaro M, Lira-Saade R (2004) Uso y manejo tradicional de la diversidad vegetal, In: Biodiversidad de Oaxaca. A J García-Mendoza, MJ Ordóñez, Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 17 & M Briones-Salas. Instituto de Biología, UNAM-Fondo Oaxaqueño para la Conservación de la Naturaleza-World Wildlife Fund, México • Cámara-Leret R, Bascompte J (2021) Language extinction triggers the loss of unique medicinal knowledge. Proceedings of the National Academy of Sciences 118 (24): e2103683118. https://doi.org/10.1073/pnas.2103683118 • Casanova-Pérez C, Delgado-Caballero CE, Cruz-Bautista P, Casanova-Pérez L (2022) Plantas medicinales usadas por los Tének en la Huasteca, México. CienciaUAT 16 (2): 40‑58. https://doi.org/10.29059/cienciauat.v16i2.1576 • Chamkhi I, Hnini M, Aurag J (2022) Conventional medicinal uses, phytoconstituents, and biological activities of Euphorbia officinarum L.: A systematic review. Advances in Pharmacological and Pharmaceutical Sciences 2022 https://doi.org/ 10.1155/2022/9971085 • Chidambaram K, Alqahtani T, Alghazwani Y, Aldahish A, Annadurai S, Venkatesan K, Dhandapani K, Thilagam E, Venkatesan K, Paulsamy P, Vasudevan R, Kandasamy G (2022) Medicinal plants of Solanum species: The promising sources of phyto-insecticidal compounds. Journal of Tropical Medicine 2022: 1‑22. https://doi.org/ 10.1155/2022/4952221 • Cilia-López VG, Cariño-Cortés R, Zurita-Salinas LR, Cilia-López VG, Cariño-Cortés R, Zurita-Salinas LR (2021) Ethnopharmacology of the Asteraceae family in Mexico. Botanical Sciences 99 (3): 455‑486. https://doi.org/10.17129/botsci.2715 • Coe F, Anderson G (2005) Snakebite ethnopharmacopoeia of eastern Nicaragua. Journal of Ethnopharmacology 96 (1-2): 303-323. https://doi.org/10.1016/j.jep.2004.09.026 • CONABIO (2006) Capital natural y bienestar social. Redacta, S.A. de C.V. URL: http:// www.conabio.gob.mx/2ep/images/3/37/capital_natural_2EP.pdf • Costanza R, d'Arge R, de Groot R, Farber S, Grasso M, Hannon B, Limburg K, Naeem S, O'Neill R, Paruelo J, Raskin R, Sutton P, van den Belt M (1997) The value of the world's ecosystem services and natural capital. Nature 387 (6630): 253‑260. https://doi.org/ 10.1038/387253a0 • Cox PA (1994) The ethnobotanical approach to drug discovery: strengths and limitations. In: Ciba Foundation Symposium, et al. (Ed.) Ciba Foundation Symposium 185 - Ethnobotany and the Search for New Drugs. URL: https://onlinelibrary.wiley.com/doi/abs/ 10.1002/9780470514634.ch3 [ISBN 978-0-470-51463-4]. • Cruz-Pérez A, Barrera-Ramos J, Bernal-Ramírez L, Bravo-Avilez D, et al. (2021) Actualized inventory of medicinal plants used in traditional medicine in Oaxaca, Mexico. Journal of Ethnobiology and Ethnomedicine 17 (7): 1‑15. https://doi.org/10.1186/ s13002-020-00431-y • Cupido M, De-Nova A, Guerrero-González M, Pérez-Vázquez FJ, Méndez-Rodríguez KB, Delgado-Sánchez P (2022) GC-MS analysis of phytochemical compounds of Opuntia megarrhiza (Cactaceae), an endangered plant of Mexico. PeerJ Organic Chemistry 4: e5. https://doi.org/10.7717/peerj-ochem.5 • Cupido M, De-Nova JA, Cilia-Lopez VG (2023) Catálogo de plantas medicinales de la región biocultural Huasteca. URL: https://biocultural.uaslp.mx/ • Cupido M, De-Nova JA, Cilia-López VG (2024) Aproximaciones evolutivas en etnobotánica de plantas medicinales y bioprospección. Botanical Sciences 102 (1): 26‑38. https://doi.org/10.17129/botsci.3325 18 Cupido M et al • Cupido M, De-Nova JA (2025) Catálogo de plantas medicinales de la región biocultural Huasteca. Revista Universitarios Potosinos 21 (281): 9‑13. URL: https://leka.uaslp.mx/ index.php/universitarios-potosinos/article/view/772 • Davis C, Choisy P (2024) Medicinal plants meet modern biodiversity science. Current biology 34 (4): R158‑R173. https://doi.org/10.1016/j.cub.2023.12.038 • De Lima LF, Andrade-Pinheiro JC, Freitas MA, da Silva AI, Fonseca VJA, da Silva TG, da Silva JCP, de Lima RH, Sales DL, Neves RP, de Brito ES, Ribeiro PRV, Canuto KM, Coutinho HDM, Siyadatpanah A, Kim B, Morais-Braga MFB (2022) Anti-Candida properties of Gossypium hirsutum L.: Enhancement of fungal growth, biofilm production and antifungal resistance. Pharmaceutics 14 (4). https://doi.org/10.3390/ pharmaceutics14040698 • De-Nova J, De-Nova E, Flores Rivas J, Reyes-Hernández H, et al. (2025) San Luis Potosí. Historia, cultura y paisaje. El Paisaje. In: Guevara Sada S, Moreno-Casasola P (Eds) La deforestación, la fragmentación y la conectividad. Primera edición. El Colegio de San Luis, Universidad Veracruzana, Mexico, 379-398 pp. URL: https:// librosdigitales.colsan.edu.mx/ebook/San_Luis_Potosi_El_paisaje.pdf [ISBN 978-607-2627-38-3/978-607-2621-64-0)]. • De-Nova JA, Castillo-Lara P, Salinas Rodríguez MM, Fortanelli-Martínez J, Mora-Olivo A (2018) Los Bosques Tropicales Estacionales>. In: Reyes-Hernández H, et al. (Ed.) Reserva de la Biosfera Sierra del Abra Tanchipa. Biodiversidad y acciones para su conservación. Primera edición. México, San Luis Potosí, 59 pp. URL: https:// www.researchgate.net/profile/J-Arturo-De-Nova/publication/ 326799033_Reserva_de_la_Biosfera_Sierra_del_Abra_Tanchipa_Biodiversidad_y_Acciones_para_su_Conservacion/ links/5b69d703299bf14c6d951c11/Reserva-de-la-Biosfera-Sierra-del-Abra-TanchipaBiodiversidad-y-Acciones-para-su-Conservacion.pdf#page=60 [ISBN 978-607-535-054-7]. • De-Nova JA, Villegas-Ortega DS, Cupido M, Cilia-López VG (2024) Evolutionary clustering in neotropical biocultural heritage: the Huastec Mayan useful plants. Botanical Journal of the Linnean Society 204 (4): 316‑326. https://doi.org/10.1093/botlinnean/ boad061 • Domínguez X, Alcorn J (1985) Screening of medicinal plants used by Huastec Mayans of northeastern Mexico. Journal of Ethnopharmacology 13 (2): 139‑156. https://doi.org/ 10.1016/0378-8741(85)90002-9 • Elizalde-Romero CA, Montoya-Inzunza LA, Contreras-Angulo LA, Heredia JB, GutiérrezGrijalva EP (2021) Solanum fruits: phytochemicals, bioaccessibility and bioavailability, and their relationship with their health-promoting effects. Frontiers in Nutrition 8 https:// doi.org/10.3389/fnut.2021.790582 • Ernst M, Grace O, Saslis-Lagoudakis CH, Nilsson N, Simonsen HT, Rønsted N (2015) Global medicinal uses of Euphorbia L. (Euphorbiaceae). Journal of Ethnopharmacology 176: 90‑101. https://doi.org/10.1016/j.jep.2015.10.025 • Faith DP, Magallón S, Hendry AP, Conti E, Yahara T, Donoghue MJ (2010) Evosystem services: an evolutionary perspective on the links between biodiversity and human wellbeing. Current Opinion in Environmental Sustainability 2 (1): 66‑74. https://doi.org/ 10.1016/j.cosust.2010.04.002 • Fidler B, Goldberg T (2014) Ingenol Mebutate Gel (Picato). Pharmacy and therapeutics 39 (1): 40‑46. URL: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3956386/ Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 19 • Fongnzossie-Fedoung E, Biwole AB, Nyangono Biyegue CF, Ngansop Tounkam M, Akono Ntonga P, Nguiamba VP, Essono DM, Forbi Funwi P, Tonga C, Nguenang GM, Kemeuze V, Sonwa DJ, Tsabang N, Bouelet IS, Tize Z, Boum AT, Momo Solefack MC, Betti JL, Nouga Bissoue A, Lehman LG, Mapongmetsem PM, Nneme Nneme L, Ngono Ngane RA, Ngogang Yonkeu J (2023) A review of Cameroonian medicinal plants with potentials for the management of the COVID-19 pandemic. Advances in Traditional Medicine 23 (1): 59‑84. https://doi.org/10.1007/s13596-021-00567-6 • Forest F, Grenyer R, Rouget M, Davies TJ, Cowling R, Faith D, Balmford A, Manning J, Procheş Ş, van der Bank M, Reeves G, Hedderson TJ, Savolainen V (2007) Preserving the evolutionary potential of floras in biodiversity hotspots. Nature 445 (7129): 757‑760. https://doi.org/10.1038/nature05587 • Futuyama DJ (1995) The uses of evolutionary biology. Science 267 (5194): 41‑42. https:// doi.org/10.1126/science.7809608 • García-Willis C, Vela-Ortega R, Maya-Leal ME (2009) Epidemiología de la mordedura por ofidio en pacientes pediátricos. Boletín Médico del Hospital Infantil de México 66 (3): 254‑259. URL: https://www.medigraphic.com/pdfs/bmhim/hi-2009/hii093f.pdf • Garnatje T, Peñuelas J, Vallès J (2017) Ethnobotany, Phylogeny, and 'Omics' for Human Health and Food Security. Trends in Plant Science 22 (3): 187‑191. https://doi.org/ 10.1016/j.tplants.2017.01.001 • GBIF (2024) Global Biodiversity Information. URL: https://www.gbif.org/ • Gitima G, Gebre A, Berhanu Y, Wato T (2025) Exploring indigenous wisdom: Ethnobotanical documentation and conservation of medicinal plants in Goba District, Southwest Ethiopia. Scientific African 27 (e02571). https://doi.org/10.1016/j.sciaf. 2025.e02571 • Hamada FA, Hamed AI, Sheded MG, Shaheen ASM (2010) Macro, micro-morphological and bioactivity aspects of naturalized exotic Solanum diphyllum L. Al-Azhar Bulletin of Science175‑206. • Hasan M, Azam MNK, Ahmed MN, Hirashima A (2015) A randomized ethnomedicinal survey of snakebite treatment in southwestern parts of Bangladesh. Journal of Traditional and Complementary Medicine 6 (4): 337‑342. https://doi.org/10.1016/j.jtcme.2015.03.007 • Hernandez M, Avila-Bello CH, Morales-Mavil JE (2007) Etnobotánica y ecología de plantas utilizadas por tres curanderos contra la mordedura de serpiente en la región de Acayucan, Veracruz, México. Boletin de la Sociedad Botanica de Mexico 81: 89‑100. URL: https://www.redalyc.org/pdf/577/57708106.pdf • Huang J, Chen B, Liu C, Lai J, Zhang J, Ma K (2012) Identifying hotspots of endemic woody seed plant diversity in China. Diversity and Distributions 18 (7): 673‑688. https:// doi.org/10.1111/j.1472-4642.2011.00845.x • Huang L, Peng H, Xiao P (2011) Development trend of traditional Chinese medicine resources. Zhongguo Zhong Yao Za Zhi 36 (1): 1‑4. https://doi.org/10.1155/2015/218901 • ICD-11 (2024) International Classification of Diseases (ICD-11) of the World Health Organization. URL: https://www.who.int/standards/classifications/classification-ofdiseases • Ijatuyi E, Lamm A, Yessoufou K, Suinyuy T, et al. (2025) Integration of indigenous knowledge with scientific knowledge: A systematic review. Environmental Science & Policy. Environmental Science & Policy 170: 104119. https://doi.org/10.1016/j.envsci. 2025.104119 20 Cupido M et al • Islam M, Ara H, Ahmad K, Uddin M (2019) A review on medicinal uses of different plants of Euphorbiaceae family. Universal Journal of Pharmaceutical Research 4 (1): 45‑49. https://doi.org/10.22270/ujpr.v4i1.236 • IUCN (2023) The IUCN Red List of Threatened Species. URL: https://www.iucnredlist.org/ en • Kadir M, Karmoker JR, Alam MR, Jahan SR, et al. (2015) Ethnopharmacological survey of medicinal plants used by traditional healers and indigenous people in Chittagong Hill Tracts, Bangladesh, for the treatment of snakebite. Evidence-Based Complementary and Alternative Medicine (23). https://doi.org/10.1155/2015/871675.871675 • Kumar A, Kumar S, Ramchiary N, Singh P, et al. (2021) Role of traditional ethnobotanical knowledge and indigenous communities in achieving sustainable development goals. Sustainability 13: 3062. https://doi.org/10.3390/su13063062 • Kurian G, Paddikkala J (2010) Oral delivery of insulin with Desmodium gangeticum root aqueous extract protects rat hearts against ischemia reperfusion injury in streptozotocin induced diabetic rats. Asian Pacific Journal of Tropical Medicine 3 (2): 94‑100. https:// doi.org/10.1016/s1995-7645(10)60043-0 • Leija-Loredo E, Reyes-Hernández H, Fotanelli-Martínez J, Palacio-Aponte G (2011) Situación actual del bosque de niebla en el estado de San Luis Potosí, México. Investigación y Ciencia de la Universidad Autónoma de Aguascalientes 53: 3‑11. https:// doi.org/10.33064/iycuaa2011534487 • Leonti M (2011) The future is written: Impact of scripts on the cognition, selection, knowledge and transmission of medicinal plant use and its implications for ethnobotany and ethnopharmacology. Journal of Ethnopharmacology 134 (3): 542‑555. https://doi.org/ 10.1016/j.jep.2011.01.017 • Loraine S, Mendoza-Espinoza J (2010) Las plantas medicinales en la lucha contra el cáncer, relevancia para México. Revista Mexicana de Ciencias Farmacéutica 41 (4): 18‑27. URL: https://www.redalyc.org/articulo.oa?id=57916060003 • Martin G, Camacho-Benavides CI, Del Campo-García CA, Anta-Fonseca S, et al. (2011) Indigenous and community conserved areas in Oaxaca, Mexico. J Environ Manage. Journal of Environmental Management 22 (2): 250‑266. https://doi.org/ 10.1108/14777831111113419 • Moremi M, Makolo F, Viljoen A, Kamatou G (2021) A review of biological activities and phytochemistry of six ethnomedicinally important South African Croton species. Journal of Ethnopharmacology 280 https://doi.org/10.1016/j.jep.2021.114416 • Muanda FN, Bouayed J, Djilani A, Yao C, Soulimani R, Dicko A (2011) Chemical composition and, cellular evaluation of the antioxidant activity of Desmodium adscendens eaves. Evidence-Based Complementary and Alternative Medicine 2011 (1). https:// doi.org/10.1155/2011/620862 • Muñetón-Pérez P (2009) Plantas medicinales: un complemento vital para la salud de los mexicanos. Entrevista con el Dr. Erick Estrada Lugo. Revista Digital Universitaria 10 (9): 1‑9. URL: http://www.revista.unam.mx/vol.10/num9/art58/int58.htm • Myers N, Mittermeier R, Mittermeier C, da Fonseca GB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403 (6772): 853‑858. https://doi.org/ 10.1038/35002501 • Nelson L, Shih MJ, Balick MJ (2007) Handbook of Poisonous and Injurious Plants. Second Edition. Springer-Verlag, New York, NY, USA. URL: https://scispace.com/pdf/ handbook-of-poisonous-and-injurious-plants-4jyqf2tuci.pdf [ISBN 10: 0-387-31268-4] Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 21 • Omale J, Friday ET (2010) Phytochemical composition, bioactivity and wound healing potential of Euphorbia heterophylla (Euphorbiaceae) leaf extract. International Journal on Pharmaceutical and Biomedical Research 1 (1): 54‑63. • Pandey A, Chandra Sekar K, Joshi B, Rawal RS (2018) Threat assessment of high-value medicinal plants of cold desert areas in Johar valley, Kailash Sacred Landscape, India. Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology 153 (1): 39‑47. https://doi.org/10.1080/11263504.2018.1448010 • Pironon S, Ondo I, Diazgranados M, Allkin R, Baquero A, Cámara-Leret R, Canteiro C, Dennehy-Carr Z, Govaerts R, Hargreaves S, Hudson A (2024) The global distribution of plants used by humans. Science 383: 293‑297. https://doi.org/10.1126/science.adg802 • POWO (2025) Plants of the World Online | Kew Science. URL: https:// powo.science.kew.org/ • QGIS (2025) Quantum GIS development team. version 3.42 Münster. URL: https:// qgis.org/community/organisation/ • Reyes Hernández H, Aguilar Robledo M, Aguirre Rivera JR, Trejo Vázquez I (2006) Cambios en la cubierta vegetal y uso del suelo en el área del proyecto Pujal-Coy, San Luis Potosí, México, 1973-2000. Investigaciones Geográficas (59)26‑42. URL: http:// www.scielo.org.mx/scielo.php? script=sci_abstract&pid=S0188-46112006000100003&lng=es&nrm=iso&tlng=es • Ríos-Oviedo Á, Cetzal-Ix W, López-Castilla HJ, Basu SK, Tamayo-Cen I, De la Torre Espinosa Z (2025) Ethnomedicinal uses of the flora as an alternative during the treatment of SARS-CoV-2 in the Mayan communities of the Yucatan Peninsula, Mexico. Journal of Herbs, Spices & Medicinal Plants 31 (3): 362‑380. https://doi.org/ 10.1080/10496475.2025.2498921 • Rolnik A, Olas B (2021) The plants of the Asteraceae family as agents in the protection of human health. International Journal of Molecular Sciences 16 (22): 3009. https://doi.org/ 10.3390/ijms22063009 • Roopashree S, Anitha J, Challa S, Mahesh TR, Venkatesan VK, Guluwadi S (2024) Mapping of soil suitability for medicinal plants using machine learning methods. Scientific Reports 14: 3741. https://doi.org/10.1038/s41598-024-54465-3 • Samy R, Thwin MM, Gopalakrishnakone P, Ignacimuthu S (2008) Ethnobotanical survey of folk plants for the treatment of snakebites in Southern part of Tamilnadu, India. Journal of Ethnopharmacology 115 (2): 302‑312. https://doi.org/10.1016/j.jep.2007.10.006 • Sanz-Biset J, Campos-de-la-Cruz J, Epiquién-Rivera MA, Cañigueral S (2009) A first survey on the medicinal plants of the Chazuta valley (Peruvian Amazon). Journal of Ethnopharmacology 122 (2): 333‑362. https://doi.org/10.1016/j.jep.2008.12.009 • SEMARNAT (2010) Norma Oficial Mexicana NOM-059-SEMARNAT-2010. Secretaría del Medio Ambientey Recursos Naturales. Protección ambiental-Especies nativas de México de flora y fauna silvestres-Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio-Lista de especies en riesgo. URL: https://www.dof.gob.mx/ normasOficiales/4254/semarnat/semarnat.htm • SLPM (2025) Herbario Isidro Palacios - SLPM - UASLP. URL: https://slpm.uaslp.mx/ • Tangjitman K, Wongsawad C, Kamwong K, Sukkho T, Trisonthi C (2015) Ethnomedicinal plants used for digestive system disorders by the Karen of northern Thailand. Journal of Ethnobiology and Ethnomedicine 11 (27). https://doi.org/10.1186/s13002-015-0011-9 22 Cupido M et al • Tay Z, Sánchez JGD, Vega JTS, Sánchez DR, et al. (2002) Serpientes y reptiles de importancia médica en México. Revista de la Facultad de Medicina UNAM 45 (5): 212‑219. URL: https://www.medigraphic.com/pdfs/facmed/un-2002/un025e.pdf • Todd W, Towne CE, Clarke JB (2023) Importance of centering traditional knowledge and Indigenous culture in geoscience education. Journal of Geoscience Education 71 (3): 403‑414. https://doi.org/10.1080/10899995.2023.2172976 • Toledo V, Barrera-Bassols N (2008) La memoria biocultural: la importancia ecológica de la sabidurías tradicionales. Icaria Editorial, Barcelona. URL: https:// books.google.com.mx/books? id=5LAJ8kp0BjUC&lpg=PA11&dq=La%20memoria%20biocultural. %20La%20importancia%20ecol%C3%B3gica%20de%20las%20sabidur%C3%ADas%20tradicionales%2C&lr&pg=PP1#v=one • Tsioutsiou EE, Amountzias V, Vontzalidou A, Dina E, Stevanović ZD, Cheilari A, Aligiannis N (2022) Medicinal plants used traditionally for skin related problems in the South Balkan and East Mediterranean region—A review. Frontiers in Pharmacology 13: 936047. https://doi.org/10.3389/fphar.2022.936047 • Vásquez J, Jiménez SL, Gómez IC, Rey JP, et al. (2013) Snakebites and ethnobotany in the eastern region of Antioquia, Colombia—the traditional use of plants. Journal of Ethnopharmacology 146 (2): 449‑455. https://doi.org/10.1016/j.jep.2012.12.043 • Wang Y, Fahad S, Wei L, Luo B, Luo J (2022) Assessing the role of financial development and financial inclusion to enhance environmental sustainability: Do financial inclusion and eco-innovation promote sustainable development? Frontiers in Environmental Science 10: 1056478. https://doi.org/10.3389/fenvs.2022.1056478 • WHO (2013) Traditional medicine. World Health Organization. Executive Board 134th Session Provisional Agenda Item 9.1. Geneva, Switzerland. URL: https://apps.who.int/gb/ ebwha/pdf_files/EB134/B134_24-en.pdf • Zhu Z, Ma K, Ran X, Zhang H, Zheng C, Han T, Zhang Q, Qin L (2011) Analgesic, antiinflammatory and antipyretic activities of the petroleum ether fraction from the ethanol extract of Desmodium podocarpum. Journal of Ethnopharmacology 133 (3): 1126‑1131. https://doi.org/10.1016/j.jep.2010.11.042 Diversity and distribution of the Huastec Mayan medicinal plants: hotspots ... 23 Supplementary materials Suppl. material 1: Table S1 Authors: Madeleyne Cupido & Arturo De-Nova Data type: Species and classes of disease Brief description: Table S1. Plant species used in the Huastec Mayan traditional medicine, the classes of disease they treat according to ICD-11 and the CBS category and their risk status according to NOM-059 and IUCN. Certain infectious or parasitic diseases (CIPD), diseases of the blood or blood-forming organs (DBBO), diseases of the circulatory system (DCS), diseases of the digestive system (DGS), diseases of the ear or mastoid process (DEMP), diseases of the visual system (DVS), diseases of the genitourinary system (DGUS), diseases of the musculoskeletal system or connective tissue (DMSC), diseases of the nervous system (DNS), diseases of the respiratory system (DRS), diseases of the skin (DS), endocrine, nutritional or metabolic diseases (ENMD), factors influencing health status or contact with health services (FIHS), injury, poisoning or certain other consequences of external causes (IPEC), mental, behavioural or neurodevelopmental disorders (MBND), neoplasms (NE), pregnancy, childbirth or the puerperium (PCP) and symptoms, signs or clinical findings not elsewhere classified (NEC). Additionally, we included culture-bound syndromes (CBS). Download file (480.74 kb) Suppl. material 2: Table S2 Authors: Madeleyne Cupido & Arturo De-Nova Data type: Plant species Brief description: Table S2. Database of the plant species used in the Huastec Mayan traditional medicine, taxonomy, common names, origin, growth forms, habitat, risk status according to NOM-059 and IUCN, classes of disease they treat according to ICD-11 and the CBS category. Download file (707.13 kb) Suppl. material 3: Table S3 Authors: Madeleyne Cupido & Arturo De-Nova Data type: Occurrences Brief description: Geographical data from records of the vascular plant species with ethnomedicinal information documented in the Tenek territory (https://doi.org/10.5281/zenodo. 15798528). Download file (789.70 kb) 24 Cupido M et al