Abstract
This study provides the first comprehensive checklist and analysis of species included in the Chiococceae tribe in Mexico, based on over 4000 herbarium specimens. We report 28 species in nine genera; including two endemic genera (Coutaportla and Nernstia) and 14 endemic species. It is estimated that 50 % of Chiococceae species found in Mexico are endemic. We mapped the diversity per grid cell of species using their occurrence records obtained from online databases. Distribution was concentrated in the tropical southern area of Mexico, with the most diverse states in terms of species and endemicity being Chiapas (17 species; 5 endemic), Veracruz (14 species; 4 endemic), and Oaxaca (14 species; 6 endemic). The combination of several traits in reproductive characters are useful to diagnose the generic entities of Chiococceae; for example form, size, symmetry, and merosity of the corolla; consistency, dorsiventral compression, and dehiscence of the fruit; and attachment and margin of the seed are useful for diagnosing the generic entities of Chiococceae. We also highlight taxonomic problems on the generic level (Asemnantha/Chiococca, Coutaportla/Lorencea; Solenandra/Exostema) and species level (e.g. Chiococca alba), for which further studies are needed to better comprehend the diversity of the Chiococceae tribe in Mexico.
References
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<p>Lorence, D.H. (1986) The identity of <em>Portlandia guatemalensis</em> (Rubiaceae). <em>Systematic Botany</em> 11: 209–213. https://doi.org/2418958</p>
<p>Lorence, D.H. (1990) A phylogenetic list of genera of Rubiaceae in Mexico. <em>Acta Botánica Mexicana</em> 12: 1–7. https://doi.org/10.21829/abm12.1990.599</p>
<p>Lorence, D.H. (2012a) <em>Chiococca</em>. <em>In</em>: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) <em>Flora Mesoamericana, vol. 4, part 2.</em> Missouri Botanical Garden Press, St. Louis Missouri, pp. 47–51.</p>
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<p>Mouly, A., Kainulainen, K., Person, C., Davis, A.P., Wong, K.M. & Razafimandimbison, S.G. (2014) Phylogenetic structure and clade circumscriptions in the Gardenieae complex (Rubiaceae). <em>Taxon</em> 63: 801–818. https://doi.org/10.12705/634.4</p>
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<p>Borhidi, A. (2002) Revalidación del género <em>Solenandra</em> Hook. F. (Rubiaceae). <em>Acta Botanica Hungarica</em> 44: 223–231. https://doi.org/10.1556/ABot.44.2002.3-4.3</p>
<p>Borhidi, A. (2003) <em>Lorencea</em>, gen. nov. (Condamineae, Rubiaceae) de Guatemala y México. <em>Acta Botanica Hungarica</em> 45: 13–21. https://doi.org/10.1556/ABot.45.2003.1-2.2</p>
<p>Borhidi, A. (2006) <em>Rubiáceas de México</em>. Académiai Kiadó, Budapest, 512 pp.</p>
<p>Borhidi, A. (2011) Estudios sobre Rubiáceas Mexicanas XXXV. <em>Chiococca motleyana</em> nomen novum. <em>Acta Botanica Hungarica</em> 53: 267–268. https://doi.org/10.1556/ABot.53.2011.3-4.7</p>
<p>Borhidi, A. (2012) <em>Rubiáceas de México</em>. Académiai Kiadó, Budapest, 608 pp.</p>
<p>Borhidi, A. (2018) <em>Coutaportla</em>, género endémico de México, <em>Lorencea</em> (Rubiaceae) endémico en Mesoamérica. <em>Acta Botanica Hungarica</em> 60: 31–45. https://doi.org/10.1556/034.60.2018.1-2.3</p>
<p>Borhidi, A. (2012) <em>La Familia Rubiaceae en la Flora de México</em>. Académiai Kiadó, Budapest, 704 pp.</p>
<p>Borhidi, A. & Diego-Pérez, N. (2001) Introducción a la taxonomía de la familia Rubiaceae en la flora de México. <em>Acta Botanica Hungarica</em> 60: 237–280. https://doi.org/10.1556/abot.44.2002.3-4.5</p>
<p>Borhidi, A. & Fernández-Zequeira, M. (1989) El género <em>Exostema</em> L. C. Rich. en Cuba. <em>Acta Botanica Hungarica</em> 35: 287–307.</p>
<p>Bremer, B. (1992) Phylogeny of the Rubiaceae (Chiococceae) based on molecular and morphological data-useful approaches for classification and comparative ecology. <em>Annals of the Missouri Botanical Garden</em> 79: 380–387. https://doi.org/10.2307/2399775</p>
<p>Bremer, B., Andreasen, K. & Olsson, D. (1995) Subfamilial and tribal relationships in the Rubiaceae based on <em>rbc</em>L sequence data. <em>Annals of the Missouri Botanical Garden</em> 82: 383–397. https://doi.org/10.2307/2399889</p>
<p>Bremer, B. & Jansen, R.K. (1991) Comparative restriction site mapping of chloroplast DNA implies new phylogenetic relationships within Rubiaceae. <em>American Journal of Botany</em> 78: 198–213. https://doi.org/2445243</p>
<p>Bremer, B., Jansen, R.K., Oxelman, B., Backlund, M., Lantz, H. & Kim, K.J. (1999) More characters or more taxa for a robust phylogeny in a case study from the coffee family (Rubiaceae). <em>Systematic Biology</em> 48: 413–435. https://doi.org/10.1080/106351599260085</p>
<p>Bremer, B. & Eriksson, T. (2009) Timetree of Rubiaceae, phylogeny and dating the family, subfamilies and tribes. <em>International Journal of Plant Science</em> 170: 766–793. https://doi.org/10.1086/599077</p>
<p>Davis, A.P., Govaerts, R., Bridson, D.M., Ruhsam, M., Moat, J. & Brummit, N.A. (2009) A global assessment of distribution, diversity, endemism, and taxonomic effort in the Rubiaceae. <em>Annals of the Missouri Botanical Garden</em> 96: 68–78. https://doi.org/10.3417/2006205</p>
<p>De Jussieu, A.L. (1789) Rubiaceae. <em>In</em>: de Jussieu, A.L. (Ed.) <em>Genera Plantarum: secundun ordines naturales disposita, juxta methodum in Horto regio parisiensi exaratam</em>. Herissant, Paris, pp. 196–210.</p>
<p>Delprete, P.G. (1996) Evaluation of the tribes Chiococceae, Condamineeae, and Catesbaeeae (Rubiaceae) based on morphological characters. <em>Opera Botanica Belgica</em> 7: 165–192. https://doi.org/10.1093/botlinnean/boy029</p>
<p>Govaerts, R., Andersson, L., Robbrecht, E., Bridson, D., Davis, A. & Schanzer, I. (2018) <em>World checklist of Rubiaceae</em>. Kew, The Board of Trustees of the Royal Botanic Gardens. Available from: http://www.kew.org/wcsp (accessed 21 August 2021).</p>
<p>Harris, J.G. & Harris, M.W. (2001) <em>Plant Identification Terminology: An Illustrated Glossary</em>. Spring Lake Publishing, Spring Lake, UTAH, 206 pp.</p>
<p>Hooker J.D. (1873) Rubiaceae. <em>In</em>: Bentham, G. & Hooker, J.D. (Eds.) <em>Genera Plantarum</em>. Reeve & Co., London, pp. 7–151.</p>
<p>IUCN (2019) Guidelines for using the IUCN Red List Categories and Criteria, Version 14. Prepared by the Standards and Petitions Subcommittee. Available from: https://www.iucnredlist.org/resources/redlistguidelines (accessed 5 November 2021)</p>
<p>Kainulainen, K., Razafimandimbison, S.G. & Bremer, B. (2013) Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae (Rubiaceae). <em>Botanical Journal of the Linnean Society</em> 173: 387–406. https://doi.org/10.1111/boj.12038</p>
<p>Liogier, E.E. (1962) <em>Flora de Cuba, vol. 5</em>. Universidad de Puerto Rico, Rio Piedras, Puerto Rico, 324 pp.</p>
<p>Lorence, D.H. (1986) The identity of <em>Portlandia guatemalensis</em> (Rubiaceae). <em>Systematic Botany</em> 11: 209–213. https://doi.org/2418958</p>
<p>Lorence, D.H. (1990) A phylogenetic list of genera of Rubiaceae in Mexico. <em>Acta Botánica Mexicana</em> 12: 1–7. https://doi.org/10.21829/abm12.1990.599</p>
<p>Lorence, D.H. (2012a) <em>Chiococca</em>. <em>In</em>: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) <em>Flora Mesoamericana, vol. 4, part 2.</em> Missouri Botanical Garden Press, St. Louis Missouri, pp. 47–51.</p>
<p>Lorence, D.H. (2012b) <em>Erithalis</em>. <em>In</em>: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) <em>Flora Mesoamericana, vol. 4, part 2.</em> Missouri Botanical Garden Press, St. Louis Missouri, pp. 85–86.</p>
<p>Lorence, D.H., Van Devender, T.R. & Ferguson, G.M. (2018) <em>Chiococca grandiflora</em> (Rubiaceae), a new species from Northern Mexico. <em>PhytoKeys </em>98: 73–83. https://doi.org/10.3897/phytokeys.98.25170</p>
<p>Martínez-Gordillo, M., Bedolla-García, B., Cornejo-Tenorio, G., Fragoso-Martínez, I., García-Peña, M.R., González-Gallegos, J.G., Lara-Cabrera, S.I. & Zamudio, S. (2017) Lamiaceae de México. <em>Botanical Sciences</em> 95: 780–806. https://doi.org/10.17129/botsci.1871</p>
<p>McDowell, T. (2012) <em>Exostema</em>. <em>In</em>: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) <em>Flora Mesoamericana, vol. 4, part 2.</em> Missouri Botanical Garden Press, St. Louis Missouri, pp. 167–169.</p>
<p>McDowell, T. & Bremer, B. (1998) Phylogeny, diversity and distribution in <em>Exostema</em> (Rubiaceae): implications of morphological and molecular analyses. <em>Plant Systematics and Evolution</em> 212: 215–246. https://doi.org/10.1007/BF01089740</p>
<p>McDowell, T., Volovsek, M. & Manos, P. (2003) Biogeography of <em>Exostema</em> (Rubiaceae) in the Caribbean region in light of molecular phylogenetic analyses. <em>Systematic Botany</em> 28: 431–441. https://doi.org/10.1043/0363-6445-28.2.431</p>
<p>Manns, U. & Bremer, B. (2010) Towards a better understanding of intertribal relationships and stable tribal delimitations within Cinchonoideae s.s. (Rubiaceae). <em>Molecular Phylogenetics and Evolution</em> 56: 21–39. https://doi.org/10.1016/j.ympev.2010.04.002</p>
<p>Morrone, J.J. (2005) Hacia una síntesis biogeográfica de México. <em>Revista Mexicana de Biodiversidad</em> 76: 207–252. https://doi.org/10.22201/ib.20078706e.2005.002.303</p>
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