Abstract
Ceratozamia rosea sp. nov., a new cycad species from the northern mountains of Chiapas, Mexico, is described and compared with its geographically proximal and morphologically similar congeners. Ceratozamia rosea is closely related to the geographically proximal species C. zoquorum, C. becerrae, and C. sancheziae, but easily distinguishable by its purplish-pink (Mexican pink) emerging leaves, a trait unique in the genus. Additionally, C. rosea presents several other qualitative and quantitative traits that are significantly differentiated from each of these species. This new species description suggests that Ceratozamia diversity in Chiapas, Mexico, might be higher than previously thought. Ceratozamia rosea is critically endangered due to over-collecting and habitat destruction.
References
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. In: Smith, V., Penev, L. (Eds.) e-Infrastructures for data publishing in biodiversity science. ZooKeys 150: 117–126. Available from: https://geocat.kew.org/ (accessed 19 April 2023). https://doi.org/10.3897/zookeys.150.2109</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Breedlove, D.E. (1981) Flora of Chiapas. Part 1: Introduction to the Flora of Chiapas. California Academy of Sciences, San Francisco, United States of America.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Brongniart, A.T. (1846) Note sur un nouveau genre de Cycadeìes du Mexique. Annales des Science Naturelles, seìr 3 5: 5–10.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Calonje, M., Stevenson, D.W. & Osborne, R. (2023) The World List of Cycads, online edition. Available from: http://www.cycadlist.org (accessed 27 March 2023).</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Donaldson, J.S. (2009) Botanic gardens science for conservation and global change. Trends in Plant Science 14 (11): 608–613. https://doi.org/10.1016/j.tplants.2009.08.008</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Ferrusquia-Villafranca, I. (1998) La Geología de México: Una sinopsis. In Ramamoorthy, T.P., Bye, R., Lot, A. & Fa, J. (Eds.) La Biodiversidad de México. UNAM, Instituto de Biología. Mexico City, Mexico. pp. 1–107.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Gaertner, J. (1791) Centuria Octava. De Fructibus et Seminibus Plantarum 2 (2): 244, pl. 133. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Grisebach, A.H.R. (1864) CXXXI. Palmae. Flora of the British West Indian Islands. Lovell Reeve, London. pp. 514. https://doi.org/10.5962/bhl.title.143</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">González, D. & Vovides, A.P. (2002) Low intralineage divergence in Ceratozamia (Zamiaceae) detected with nuclear ribosomal DNA ITS and chloroplast DNA trnL-F non-coding region. Systematic Botany 27: 654–661. https://doi.org/10.1043/0363-6445-27.4.654</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Gutiérrez-Ortega, J.S., Pérez-Farrera, M.A., Vovides, A.P., Chávez-Cortázar, A., López, S., Santos-Hernández, N.G. & Ruíz-Roblero, S.K. (2021) Ceratozamia sanchezae (Zamiaceae): a new cycad species from Chiapas Highlands (Mexico). Phytotaxa 500: 201–216. https://doi.org/10.11646/phytotaxa.500.3.4</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Habib, S., Gong, Y., Dong, S., Lindstrom, A., Stevenson, D.W., Wu, H. & Zhang, S. (2023) Phylotranscriptomics Shed Light on Intrageneric Relationships and Historical Biogeography of Ceratozamia (Cycadales). Plants 12 (3): 478. https://doi.org/10.3390/plants12030478</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hammer, Ø., Harper, D.A. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia electronica 4 (1): 9.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hemsley, W.B. (1878) Clusia guatemalensis. In: Diagnoses plantarum novarum vel minus cognitarum Mexicanarum et Centrali-Americanarum 1. pp. 2–3. https://doi.org/10.5962/bhl.title.60211</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">INEGI [Instituto Nacional de Estadística y Geografía] (2010) Compendio de información geográfica municipal 2010. Maravilla Tenejapa, Chiapas. Clave geoestadística 07115. Available from: https://www.inegi.org.mx/contenidos/app/mexicocifras/datos_geograficos/07/07115.pdf (accessed 18 April 2023).</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">IUCN (2023) The IUCN Red List of Threatened Species. Version 2022-2. Available from: https://www.iucnredlist.org (accessed 29 March 2023). </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Jacquin, N.J. (1781) Selectarum Stirpium Americanarum Historia. Ex Officina Krausiana, Vindobonæ. pp. 284. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Koch, K. (1855) Monstera acuminata. Index Seminum (Berlin): 4.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Liebmann (1849) Chamaedorea tepejilote. Historia Naturalis Palmarum 3 (9): 308</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus, C.V. (1759) Cedrela odorata. Systema Naturae, Editio Decima 2: 940. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Marris, E. (2006) Plant science: Gardens in full bloom. Nature 440: 860–863. https://doi.org/10.1038/440860a</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martínez-Domínguez, L., Nicolalde-Morejón, F., Vergara-Silva, F. & Stevenson, D.W. (2022) Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1–102. https://doi.org/10.3897/phytokeys.208.80382</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martínez-Domínguez, L., Nicolalde-Morejón, F., Vergara-Silva, F., Stevenson, D.W. & del Callejo, E. (2017) Cryptic diversity, sympatry, and other integrative taxonomy scenarios in the Mexican Ceratozamia miqueliana complex (Zamiaceae). Organisms Diversity & Evolution 17 (4): 727–752. https://doi.org/10.1007/s13127-017-0341-7</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1849) Synopsis Palmarum. In systema receptarum, indicata simul patria et adjectis passim characteribus. Historia Naturalis Palmarum 3 (9): 307–344.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1853) Ordo Palmarum. Historia Naturalis Palmarum 3 (10): 323. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1838) Chamaedorea oblongata. Historia Naturalis Palmarum 3 (7): 160.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Medina-Villarreal, A., González-Astorga, J. & de los Monteros, A.E. (2019) Evolution of Ceratozamia cycads: A proximate-ultimate approach. Molecular Phylogenetics and Evolution 139: 106530. https://doi.org/10.1016/j.ympev.2019.106530</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miquel, F.A.W. (1847) Over eenige nieuwe of zeldame Cycadeën in den Hortus Botanicus Amsterdam. Eerste gedeelte. Tijdschrift voor de wisen natuurkundige wetenschappen, uitgegeven door de eerste klasse van het Kon. Ned. Instituut van Wetenschappen, Letteren en Schoone Kunsten 1: 33–43. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Otto, F., Dietrich, A. (1852) Chamaedorea ernesti-augusti. Allgemeine Gartenzeitung 20 (10): 73–74. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P. & Iglesias, C. (2001) A new species of Ceratozamia (Zamiaceae) from Chiapas, Mexico. Botanical Journal of the Linnean Society 137: 77–80. https://doi.org/10.1006/bojl.2001.0448</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A. (2005) Estudio sistemático de los complejos Ceratozamia norstogii D. W. Stev. y Ceratozamia miqueliana H. Wendl. (Cycadales, Zamiaceae). Universidad Autónoma de Querétaro. Santiago de Queretaro, Mexico. pp. 185</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Gregory, T.J., Chemnick, J., Salas-Morales, S., Calonje, M. & Díaz-Jiménez, P. (2022) Ceratozamia schiblii (Zamiaceae): A new cycad species from the eastern mountains of Oaxaca, Mexico. Taxonomy 2 (3): 324–338. https://doi.org/10.3390/taxonomy2030025</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Haynes, J.L., Chemnick, J., Salas-Morales, S.H., Calonje, M. & Vovides, A.P. (2021a) Ceratozamia aurantiaca (Zamiaceae): A new cycad species from the northern rainforests of Oaxaca, Mexico. Taxonomy 1 (3): 243–255. https://doi.org/10.3390/taxonomy1030018</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Vovides, A.P., Calonje, M. & Díaz-Jiménez, P. (2021b) Ceratozamia dominguezii (Zamiaceae): A new cycad species from southeastern Mexico. Taxonomy 1 (4): 345–359. https://doi.org/10.3390/taxonomy1040026</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., González, D., Hernández-Sandoval, L. & Martínez, M. (2007) Variación morfológica y genética del complejo Ceratozamia miqueliana H.A.Wendl. (Cycadales, Zamiaceae). Memoirs of New York Botanical Garden 97: 519–541.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., González, D., López, S., Hernandez-Sandoval, L. & Martínez, M. (2017) Estimation of genetic variation in closely related cycad species in Ceratozamia (Zamiaceae: Cycadales) using RAPDs markers. Revista de Biología Tropical 65 (1): 303–319.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., Martinez-Camilo, R., Melendez, N.M. & Iglesias, C. (2009) A reassessment of the Ceratozamia miqueliana species complex (Zamiaceae) of southeastern Mexico, with comments on species relationships. Systematics and Biodiversity 7 433–443. https://doi.org/10.1017/S1477200009990211</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Reed, C.F. (1968) Index Thelypteridis. Phytologia 17 (4): 292.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Rose, J.N. (1901) Undescribed plants from Guatemala and other Central American Republics XXII. Botanical Gazette 31 (2): 109–125. https://doi.org/10.1086/328081 </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sargent, C.S. (1890) Notes on North American Trees.- XVII. Garden & Forest 3 (118): 260. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Schutzman, B. (1984) A new species of Zamia L. (Zamiaceae, Cycadales) from Chiapas, Mexico. Phytologia 55 (5): 299.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Schutzman, B. & Vovides, A.P. (1998) A new species of Zamia (Zamiaceae, Cycadales) from Eastern, Chiapas, Mexico. Novon 8 (4): 441–446. https://doi.org/10.2307/3391871</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Stevenson, D.W., Sabato, S. & Torres, M.V. (1986) A new species of Ceratozamia (Zamiaceae) from Veracruz, Mexico with comments on species relationships, habitats, and vegetative morphology in Ceratozamia. Brittonia 38 (1): 17–26. https://doi.org/10.2307/2807413</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.H., Li, D.Z., Marhold, K., May, T.W., McNeil, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Available from: https://www.iapt-taxon.org/nomen/main.php (accessed 18 April 2023) https://doi.org/10.12705/Code.2018</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Schutzman, B., Iglesias, C. & Hernández-Sandoval, L. (2004) A new species of Ceratozamia (Zamiaceae) from Tabasco and Chiapas, Mexico. Botanical Journal of the Linnean Society 146 (1): 123–128. https://doi.org/10.1111/j.1095-8339.2004.00312.x</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., González, D., Pérez-Farrera, M.A., Avendaño, S. & Bárcenas, C. (2004a) A review of research on the cycad genus Ceratozamia Brongn. (Zamiaceae) in Mexico. Taxon 53: 291–297. https://doi.org/10.2307/4135609</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., González, D. & Avendaño, S. (2004b) Relationships and phytogeography in Ceratozamia (Zamiaceae). In: Walters, T. & Osborne, R. (Eds.) Cycad classification: concepts and recommendations. CABI Publishing, Wallingford. pp. 109–125. https://doi.org/10.1079/9780851997414.0109</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Avendaño, S., Medina-Villarreal, A., González-Astorga, J. & Galicia, S. (2020) A revision of the Ceratozamia miqueliana (Zamiaceae) species complex based on analyses of leaflet anatomical characters. Flora 270: 151649. https://doi.org/10.1016/j.flora.2020.151649</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Salinas-Rodríguez, M.M., Galicia, S., Díaz-Jiménez, P., Calonje, M. & Gutiérrez-Ortega, J.S. (2022) Morphological and anatomical analyses clarify the species definition of Ceratozamia latifolia Miq. (Zamiaceae) and lead to the description of a new species: Ceratozamia reesii. Phytotaxa 575: 224–252. https://doi.org/10.11646/phytotaxa.575.3.2</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Stevenson, D.W., Pérez-Farrera, M.A., López, S. & Avendaño, S. (2016) What is Ceratozamia mexicana (Zamiaceae)? Botanical Sciences 94 (2): 419–429. https://doi.org/10.17129/botsci.449</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Wendland, H. (1854) Descriptio speciei novae e genere Ceratozamiae. In: Index palmarum, cyclanthearum, pandanearum, cycadearum, quae in hortis europaeis coluntur, synonymis gravioribus interpositis. Hannoverae. pp. 68.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Breedlove, D.E. (1981) Flora of Chiapas. Part 1: Introduction to the Flora of Chiapas. California Academy of Sciences, San Francisco, United States of America.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Brongniart, A.T. (1846) Note sur un nouveau genre de Cycadeìes du Mexique. Annales des Science Naturelles, seìr 3 5: 5–10.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Calonje, M., Stevenson, D.W. & Osborne, R. (2023) The World List of Cycads, online edition. Available from: http://www.cycadlist.org (accessed 27 March 2023).</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Donaldson, J.S. (2009) Botanic gardens science for conservation and global change. Trends in Plant Science 14 (11): 608–613. https://doi.org/10.1016/j.tplants.2009.08.008</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Ferrusquia-Villafranca, I. (1998) La Geología de México: Una sinopsis. In Ramamoorthy, T.P., Bye, R., Lot, A. & Fa, J. (Eds.) La Biodiversidad de México. UNAM, Instituto de Biología. Mexico City, Mexico. pp. 1–107.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Gaertner, J. (1791) Centuria Octava. De Fructibus et Seminibus Plantarum 2 (2): 244, pl. 133. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Grisebach, A.H.R. (1864) CXXXI. Palmae. Flora of the British West Indian Islands. Lovell Reeve, London. pp. 514. https://doi.org/10.5962/bhl.title.143</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">González, D. & Vovides, A.P. (2002) Low intralineage divergence in Ceratozamia (Zamiaceae) detected with nuclear ribosomal DNA ITS and chloroplast DNA trnL-F non-coding region. Systematic Botany 27: 654–661. https://doi.org/10.1043/0363-6445-27.4.654</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Gutiérrez-Ortega, J.S., Pérez-Farrera, M.A., Vovides, A.P., Chávez-Cortázar, A., López, S., Santos-Hernández, N.G. & Ruíz-Roblero, S.K. (2021) Ceratozamia sanchezae (Zamiaceae): a new cycad species from Chiapas Highlands (Mexico). Phytotaxa 500: 201–216. https://doi.org/10.11646/phytotaxa.500.3.4</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Habib, S., Gong, Y., Dong, S., Lindstrom, A., Stevenson, D.W., Wu, H. & Zhang, S. (2023) Phylotranscriptomics Shed Light on Intrageneric Relationships and Historical Biogeography of Ceratozamia (Cycadales). Plants 12 (3): 478. https://doi.org/10.3390/plants12030478</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hammer, Ø., Harper, D.A. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia electronica 4 (1): 9.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Hemsley, W.B. (1878) Clusia guatemalensis. In: Diagnoses plantarum novarum vel minus cognitarum Mexicanarum et Centrali-Americanarum 1. pp. 2–3. https://doi.org/10.5962/bhl.title.60211</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">INEGI [Instituto Nacional de Estadística y Geografía] (2010) Compendio de información geográfica municipal 2010. Maravilla Tenejapa, Chiapas. Clave geoestadística 07115. Available from: https://www.inegi.org.mx/contenidos/app/mexicocifras/datos_geograficos/07/07115.pdf (accessed 18 April 2023).</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">IUCN (2023) The IUCN Red List of Threatened Species. Version 2022-2. Available from: https://www.iucnredlist.org (accessed 29 March 2023). </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Jacquin, N.J. (1781) Selectarum Stirpium Americanarum Historia. Ex Officina Krausiana, Vindobonæ. pp. 284. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Koch, K. (1855) Monstera acuminata. Index Seminum (Berlin): 4.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Liebmann (1849) Chamaedorea tepejilote. Historia Naturalis Palmarum 3 (9): 308</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Linnaeus, C.V. (1759) Cedrela odorata. Systema Naturae, Editio Decima 2: 940. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Marris, E. (2006) Plant science: Gardens in full bloom. Nature 440: 860–863. https://doi.org/10.1038/440860a</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martínez-Domínguez, L., Nicolalde-Morejón, F., Vergara-Silva, F. & Stevenson, D.W. (2022) Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1–102. https://doi.org/10.3897/phytokeys.208.80382</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martínez-Domínguez, L., Nicolalde-Morejón, F., Vergara-Silva, F., Stevenson, D.W. & del Callejo, E. (2017) Cryptic diversity, sympatry, and other integrative taxonomy scenarios in the Mexican Ceratozamia miqueliana complex (Zamiaceae). Organisms Diversity & Evolution 17 (4): 727–752. https://doi.org/10.1007/s13127-017-0341-7</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1849) Synopsis Palmarum. In systema receptarum, indicata simul patria et adjectis passim characteribus. Historia Naturalis Palmarum 3 (9): 307–344.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1853) Ordo Palmarum. Historia Naturalis Palmarum 3 (10): 323. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Martius, C.F.P.V. (1838) Chamaedorea oblongata. Historia Naturalis Palmarum 3 (7): 160.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Medina-Villarreal, A., González-Astorga, J. & de los Monteros, A.E. (2019) Evolution of Ceratozamia cycads: A proximate-ultimate approach. Molecular Phylogenetics and Evolution 139: 106530. https://doi.org/10.1016/j.ympev.2019.106530</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Miquel, F.A.W. (1847) Over eenige nieuwe of zeldame Cycadeën in den Hortus Botanicus Amsterdam. Eerste gedeelte. Tijdschrift voor de wisen natuurkundige wetenschappen, uitgegeven door de eerste klasse van het Kon. Ned. Instituut van Wetenschappen, Letteren en Schoone Kunsten 1: 33–43. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Otto, F., Dietrich, A. (1852) Chamaedorea ernesti-augusti. Allgemeine Gartenzeitung 20 (10): 73–74. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P. & Iglesias, C. (2001) A new species of Ceratozamia (Zamiaceae) from Chiapas, Mexico. Botanical Journal of the Linnean Society 137: 77–80. https://doi.org/10.1006/bojl.2001.0448</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A. (2005) Estudio sistemático de los complejos Ceratozamia norstogii D. W. Stev. y Ceratozamia miqueliana H. Wendl. (Cycadales, Zamiaceae). Universidad Autónoma de Querétaro. Santiago de Queretaro, Mexico. pp. 185</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Gregory, T.J., Chemnick, J., Salas-Morales, S., Calonje, M. & Díaz-Jiménez, P. (2022) Ceratozamia schiblii (Zamiaceae): A new cycad species from the eastern mountains of Oaxaca, Mexico. Taxonomy 2 (3): 324–338. https://doi.org/10.3390/taxonomy2030025</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Haynes, J.L., Chemnick, J., Salas-Morales, S.H., Calonje, M. & Vovides, A.P. (2021a) Ceratozamia aurantiaca (Zamiaceae): A new cycad species from the northern rainforests of Oaxaca, Mexico. Taxonomy 1 (3): 243–255. https://doi.org/10.3390/taxonomy1030018</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Vovides, A.P., Calonje, M. & Díaz-Jiménez, P. (2021b) Ceratozamia dominguezii (Zamiaceae): A new cycad species from southeastern Mexico. Taxonomy 1 (4): 345–359. https://doi.org/10.3390/taxonomy1040026</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., González, D., Hernández-Sandoval, L. & Martínez, M. (2007) Variación morfológica y genética del complejo Ceratozamia miqueliana H.A.Wendl. (Cycadales, Zamiaceae). Memoirs of New York Botanical Garden 97: 519–541.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., González, D., López, S., Hernandez-Sandoval, L. & Martínez, M. (2017) Estimation of genetic variation in closely related cycad species in Ceratozamia (Zamiaceae: Cycadales) using RAPDs markers. Revista de Biología Tropical 65 (1): 303–319.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Pérez-Farrera, M.A., Vovides, A.P., Martinez-Camilo, R., Melendez, N.M. & Iglesias, C. (2009) A reassessment of the Ceratozamia miqueliana species complex (Zamiaceae) of southeastern Mexico, with comments on species relationships. Systematics and Biodiversity 7 433–443. https://doi.org/10.1017/S1477200009990211</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Reed, C.F. (1968) Index Thelypteridis. Phytologia 17 (4): 292.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Rose, J.N. (1901) Undescribed plants from Guatemala and other Central American Republics XXII. Botanical Gazette 31 (2): 109–125. https://doi.org/10.1086/328081 </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Sargent, C.S. (1890) Notes on North American Trees.- XVII. Garden & Forest 3 (118): 260. </span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Schutzman, B. (1984) A new species of Zamia L. (Zamiaceae, Cycadales) from Chiapas, Mexico. Phytologia 55 (5): 299.</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Schutzman, B. & Vovides, A.P. (1998) A new species of Zamia (Zamiaceae, Cycadales) from Eastern, Chiapas, Mexico. Novon 8 (4): 441–446. https://doi.org/10.2307/3391871</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Stevenson, D.W., Sabato, S. & Torres, M.V. (1986) A new species of Ceratozamia (Zamiaceae) from Veracruz, Mexico with comments on species relationships, habitats, and vegetative morphology in Ceratozamia. Brittonia 38 (1): 17–26. https://doi.org/10.2307/2807413</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.H., Li, D.Z., Marhold, K., May, T.W., McNeil, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Available from: https://www.iapt-taxon.org/nomen/main.php (accessed 18 April 2023) https://doi.org/10.12705/Code.2018</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Schutzman, B., Iglesias, C. & Hernández-Sandoval, L. (2004) A new species of Ceratozamia (Zamiaceae) from Tabasco and Chiapas, Mexico. Botanical Journal of the Linnean Society 146 (1): 123–128. https://doi.org/10.1111/j.1095-8339.2004.00312.x</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., González, D., Pérez-Farrera, M.A., Avendaño, S. & Bárcenas, C. (2004a) A review of research on the cycad genus Ceratozamia Brongn. (Zamiaceae) in Mexico. Taxon 53: 291–297. https://doi.org/10.2307/4135609</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., González, D. & Avendaño, S. (2004b) Relationships and phytogeography in Ceratozamia (Zamiaceae). In: Walters, T. & Osborne, R. (Eds.) Cycad classification: concepts and recommendations. CABI Publishing, Wallingford. pp. 109–125. https://doi.org/10.1079/9780851997414.0109</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Gutiérrez-Ortega, J.S., Avendaño, S., Medina-Villarreal, A., González-Astorga, J. & Galicia, S. (2020) A revision of the Ceratozamia miqueliana (Zamiaceae) species complex based on analyses of leaflet anatomical characters. Flora 270: 151649. https://doi.org/10.1016/j.flora.2020.151649</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Pérez-Farrera, M.A., Salinas-Rodríguez, M.M., Galicia, S., Díaz-Jiménez, P., Calonje, M. & Gutiérrez-Ortega, J.S. (2022) Morphological and anatomical analyses clarify the species definition of Ceratozamia latifolia Miq. (Zamiaceae) and lead to the description of a new species: Ceratozamia reesii. Phytotaxa 575: 224–252. https://doi.org/10.11646/phytotaxa.575.3.2</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Vovides, A.P., Stevenson, D.W., Pérez-Farrera, M.A., López, S. & Avendaño, S. (2016) What is Ceratozamia mexicana (Zamiaceae)? Botanical Sciences 94 (2): 419–429. https://doi.org/10.17129/botsci.449</span></span></span>
<p align="justify"><span style="color: #000000;"><span style="font-family: Times New Roman, serif;"><span style="font-size: small;">Wendland, H. (1854) Descriptio speciei novae e genere Ceratozamiae. In: Index palmarum, cyclanthearum, pandanearum, cycadearum, quae in hortis europaeis coluntur, synonymis gravioribus interpositis. Hannoverae. pp. 68.</span></span></span>