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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Borhidi, A. (2002) Revalidación del género Solenandra Hook. F. (Rubiaceae). Acta Botanica Hungarica 44: 223–231. https://doi.org/10.1556/ABot.44.2002.3-4.3
Borhidi, A. (2003) Lorencea, gen. nov. (Condamineae, Rubiaceae) de Guatemala y México. Acta Botanica Hungarica 45: 13–21. https://doi.org/10.1556/ABot.45.2003.1-2.2
Borhidi, A. (2006) Rubiáceas de México. Académiai Kiadó, Budapest, 512 pp.
Borhidi, A. (2011) Estudios sobre Rubiáceas Mexicanas XXXV. Chiococca motleyana nomen novum. Acta Botanica Hungarica 53: 267–268. https://doi.org/10.1556/ABot.53.2011.3-4.7
Borhidi, A. (2012) Rubiáceas de México. Académiai Kiadó, Budapest, 608 pp.
Borhidi, A. (2018) Coutaportla, género endémico de México, Lorencea (Rubiaceae) endémico en Mesoamérica. Acta Botanica Hungarica 60: 31–45. https://doi.org/10.1556/034.60.2018.1-2.3
Borhidi, A. (2012) La Familia Rubiaceae en la Flora de México. Académiai Kiadó, Budapest, 704 pp.
Borhidi, A. & Diego-Pérez, N. (2001) Introducción a la taxonomía de la familia Rubiaceae en la flora de México. Acta Botanica Hungarica 60: 237–280. https://doi.org/10.1556/abot.44.2002.3-4.5
Borhidi, A. & Fernández-Zequeira, M. (1989) El género Exostema L. C. Rich. en Cuba. Acta Botanica Hungarica 35: 287–307.
Bremer, B. (1992) Phylogeny of the Rubiaceae (Chiococceae) based on molecular and morphological data-useful approaches for classification and comparative ecology. Annals of the Missouri Botanical Garden 79: 380–387. https://doi.org/10.2307/2399775
Bremer, B., Andreasen, K. & Olsson, D. (1995) Subfamilial and tribal relationships in the Rubiaceae based on rbcL sequence data. Annals of the Missouri Botanical Garden 82: 383–397. https://doi.org/10.2307/2399889
Bremer, B. & Jansen, R.K. (1991) Comparative restriction site mapping of chloroplast DNA implies new phylogenetic relationships within Rubiaceae. American Journal of Botany 78: 198–213. https://doi.org/2445243
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). Systematic Biology 48: 413–435. https://doi.org/10.1080/106351599260085
Bremer, B. & Eriksson, T. (2009) Timetree of Rubiaceae, phylogeny and dating the family, subfamilies and tribes. International Journal of Plant Science 170: 766–793. https://doi.org/10.1086/599077
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. Annals of the Missouri Botanical Garden 96: 68–78. https://doi.org/10.3417/2006205
De Jussieu, A.L. (1789) Rubiaceae. In: de Jussieu, A.L. (Ed.) Genera Plantarum: secundun ordines naturales disposita, juxta methodum in Horto regio parisiensi exaratam. Herissant, Paris, pp. 196–210.
Delprete, P.G. (1996) Evaluation of the tribes Chiococceae, Condamineeae, and Catesbaeeae (Rubiaceae) based on morphological characters. Opera Botanica Belgica 7: 165–192. https://doi.org/10.1093/botlinnean/boy029
Govaerts, R., Andersson, L., Robbrecht, E., Bridson, D., Davis, A. & Schanzer, I. (2018) World checklist of Rubiaceae. Kew, The Board of Trustees of the Royal Botanic Gardens. Available from: http://www.kew.org/wcsp (accessed 21 August 2021).
Harris, J.G. & Harris, M.W. (2001) Plant Identification Terminology: An Illustrated Glossary. Spring Lake Publishing, Spring Lake, UTAH, 206 pp.
Hooker J.D. (1873) Rubiaceae. In: Bentham, G. & Hooker, J.D. (Eds.) Genera Plantarum. Reeve & Co., London, pp. 7–151.
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)
Kainulainen, K., Razafimandimbison, S.G. & Bremer, B. (2013) Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae (Rubiaceae). Botanical Journal of the Linnean Society 173: 387–406. https://doi.org/10.1111/boj.12038
Liogier, E.E. (1962) Flora de Cuba, vol. 5. Universidad de Puerto Rico, Rio Piedras, Puerto Rico, 324 pp.
Lorence, D.H. (1986) The identity of Portlandia guatemalensis (Rubiaceae). Systematic Botany 11: 209–213. https://doi.org/2418958
Lorence, D.H. (1990) A phylogenetic list of genera of Rubiaceae in Mexico. Acta Botánica Mexicana 12: 1–7. https://doi.org/10.21829/abm12.1990.599
Lorence, D.H. (2012a) Chiococca. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 47–51.
Lorence, D.H. (2012b) Erithalis. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 85–86.
Lorence, D.H., Van Devender, T.R. & Ferguson, G.M. (2018) Chiococca grandiflora (Rubiaceae), a new species from Northern Mexico. PhytoKeys 98: 73–83. https://doi.org/10.3897/phytokeys.98.25170
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. Botanical Sciences 95: 780–806. https://doi.org/10.17129/botsci.1871
McDowell, T. (2012) Exostema. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 167–169.
McDowell, T. & Bremer, B. (1998) Phylogeny, diversity and distribution in Exostema (Rubiaceae): implications of morphological and molecular analyses. Plant Systematics and Evolution 212: 215–246. https://doi.org/10.1007/BF01089740
McDowell, T., Volovsek, M. & Manos, P. (2003) Biogeography of Exostema (Rubiaceae) in the Caribbean region in light of molecular phylogenetic analyses. Systematic Botany 28: 431–441. https://doi.org/10.1043/0363-6445-28.2.431
Manns, U. & Bremer, B. (2010) Towards a better understanding of intertribal relationships and stable tribal delimitations within Cinchonoideae s.s. (Rubiaceae). Molecular Phylogenetics and Evolution 56: 21–39. https://doi.org/10.1016/j.ympev.2010.04.002
Morrone, J.J. (2005) Hacia una síntesis biogeográfica de México. Revista Mexicana de Biodiversidad 76: 207–252. https://doi.org/10.22201/ib.20078706e.2005.002.303
Motley, T.J, Wurdack, K.J. & Delprete, P.G. (2005) Molecular systematics of the Catesbeeae-Chiococceae complex (Rubiaceae) flower and fruit evolution and biogeographic implications. American Journal of Botany 92: 316–329. https://doi.org/10.3732/ajb.92.2.316.
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). Taxon 63: 801–818. https://doi.org/10.12705/634.4
Ochoterena, H. (2012a) Coutaportla. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 69.
Ochoterena, H. (2012b) Coutarea. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 69–70.
Ochoterena, H. (2012c) Hintonia. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 118–119.
Paudyal, S.K., Delprete, P.G. & Motley, T.J. (2014) Using molecular, morphological, and palynological evidence to transfer Strumpfia maritima to the monotypic tribe Strumpfieae (Cinchonoideae, Rubiaceae), and a re-delimitation of the tribe Chiococceae. Systematic Botany 39: 1197–1203. https://doi.org/10.1600/036364414X682580
Paudyal, S.K., Delprete, P.G, Neupane, S. & Motley, T.J. (2018) Molecular phylogenetic analysis and generic delimitations in tribe Chiococceae (Cinchonoideae, Rubiaceae). Botanical Journal of the Linnean Society 187: 365–396. https://doi.org/10.1093/botlinnean/boy029
Quantum GIS Development Team (2018) Quantum GIS Geographic Information System. Open Source Geospatial Foundation Project. [http://qgis.osgeo.org]
Robbrecht, E. (1988) Tropical Woody Rubiaceae. Opera Botanica Belgica 1: 1–272. https://doi.org/10.2307/4110534
Rova, J.H.E., Delprete, P.G., Andersson, L. & Albert, V.A. (2002) A trnL-F cpDNA sequence study of the Condamineae-Rondeletieae-Sipaneeae complex with implications on the phylogeny of Rubiaceae. American Journal of Botany 89: 145–159. https://doi.org/10.3732/ajb.89.1.145.
Rzedowski, J. (1991) Diversidad y orígenes de la flora fanerogámica de México. Acta Botánica Mexicana 14: 13–21. https://doi.org/10.21829/abm14.1991.611
Sazima, M., Buzato, S. & Sazima, I. (1999) Bat-pollinated flower assemblages and bat visitors at two Atlantic forest sites in Brazil. Annals of Botany 83: 705–712. https://doi.org/10.1006/anbo.1999.0876
Schumann, K. (1891) Rubiaceae. In: Engler, A. & Prantl, K. (Eds.) Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten begründet. W. Engelmann, Leipzing, pp. 97–156.
Standley, P.C. (1921) Rubiaceae. North American Flora 32: 1–82.
Standley, P.C. (1934) Rubiales-Rubiaceae. North American Flora 32: 1–300.
Urban, I. (1923) Plantae cubenses novae. Symbolae Antillanae 9: 55–176.
Villarreal, J.A. (1987) Coutaportla pailensis (Rubiaceae), a new species from Coahuila, México. Sida 12: 223–225.
Villaseñor, J.L. (2003) Diversidad y distribución de las Magnoliophyta de México. Interciencia 28: 160–167.
Villaseñor, J.L. (2016) Checklist of the native vascular plants of Mexico. Revista Mexicana de Biodiversidad 87: 559–902. https://doi.org/10.1016/j.rmb.2016.06.017
Aiello, A. (1979) A re-examination of Portlandia (Rubiaceae) and associated taxa. Journal of the Arnold Arboretum 60: 38–126.
Alvarado-Cárdenas, L.O., Lozada-Pérez, L., Islas-Hernández, C.S., Cortez, E.B., Maya-Mandujano, K.G. & Chávez-Hernández, M.G. (2020) Apocináceas de ayer y hoy. Conocimiento histórico y reevaluación de la diversidad y distribución de Apocynaceae en México. Botanical Sciences 98: 393–416. https://doi.org/10.17129/botsci.2525
Borhidi, A. (2002) Revalidación del género Solenandra Hook. F. (Rubiaceae). Acta Botanica Hungarica 44: 223–231. https://doi.org/10.1556/ABot.44.2002.3-4.3
Borhidi, A. (2003) Lorencea, gen. nov. (Condamineae, Rubiaceae) de Guatemala y México. Acta Botanica Hungarica 45: 13–21. https://doi.org/10.1556/ABot.45.2003.1-2.2
Borhidi, A. (2006) Rubiáceas de México. Académiai Kiadó, Budapest, 512 pp.
Borhidi, A. (2011) Estudios sobre Rubiáceas Mexicanas XXXV. Chiococca motleyana nomen novum. Acta Botanica Hungarica 53: 267–268. https://doi.org/10.1556/ABot.53.2011.3-4.7
Borhidi, A. (2012) Rubiáceas de México. Académiai Kiadó, Budapest, 608 pp.
Borhidi, A. (2018) Coutaportla, género endémico de México, Lorencea (Rubiaceae) endémico en Mesoamérica. Acta Botanica Hungarica 60: 31–45. https://doi.org/10.1556/034.60.2018.1-2.3
Borhidi, A. (2012) La Familia Rubiaceae en la Flora de México. Académiai Kiadó, Budapest, 704 pp.
Borhidi, A. & Diego-Pérez, N. (2001) Introducción a la taxonomía de la familia Rubiaceae en la flora de México. Acta Botanica Hungarica 60: 237–280. https://doi.org/10.1556/abot.44.2002.3-4.5
Borhidi, A. & Fernández-Zequeira, M. (1989) El género Exostema L. C. Rich. en Cuba. Acta Botanica Hungarica 35: 287–307.
Bremer, B. (1992) Phylogeny of the Rubiaceae (Chiococceae) based on molecular and morphological data-useful approaches for classification and comparative ecology. Annals of the Missouri Botanical Garden 79: 380–387. https://doi.org/10.2307/2399775
Bremer, B., Andreasen, K. & Olsson, D. (1995) Subfamilial and tribal relationships in the Rubiaceae based on rbcL sequence data. Annals of the Missouri Botanical Garden 82: 383–397. https://doi.org/10.2307/2399889
Bremer, B. & Jansen, R.K. (1991) Comparative restriction site mapping of chloroplast DNA implies new phylogenetic relationships within Rubiaceae. American Journal of Botany 78: 198–213. https://doi.org/2445243
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). Systematic Biology 48: 413–435. https://doi.org/10.1080/106351599260085
Bremer, B. & Eriksson, T. (2009) Timetree of Rubiaceae, phylogeny and dating the family, subfamilies and tribes. International Journal of Plant Science 170: 766–793. https://doi.org/10.1086/599077
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. Annals of the Missouri Botanical Garden 96: 68–78. https://doi.org/10.3417/2006205
De Jussieu, A.L. (1789) Rubiaceae. In: de Jussieu, A.L. (Ed.) Genera Plantarum: secundun ordines naturales disposita, juxta methodum in Horto regio parisiensi exaratam. Herissant, Paris, pp. 196–210.
Delprete, P.G. (1996) Evaluation of the tribes Chiococceae, Condamineeae, and Catesbaeeae (Rubiaceae) based on morphological characters. Opera Botanica Belgica 7: 165–192. https://doi.org/10.1093/botlinnean/boy029
Govaerts, R., Andersson, L., Robbrecht, E., Bridson, D., Davis, A. & Schanzer, I. (2018) World checklist of Rubiaceae. Kew, The Board of Trustees of the Royal Botanic Gardens. Available from: http://www.kew.org/wcsp (accessed 21 August 2021).
Harris, J.G. & Harris, M.W. (2001) Plant Identification Terminology: An Illustrated Glossary. Spring Lake Publishing, Spring Lake, UTAH, 206 pp.
Hooker J.D. (1873) Rubiaceae. In: Bentham, G. & Hooker, J.D. (Eds.) Genera Plantarum. Reeve & Co., London, pp. 7–151.
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)
Kainulainen, K., Razafimandimbison, S.G. & Bremer, B. (2013) Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae (Rubiaceae). Botanical Journal of the Linnean Society 173: 387–406. https://doi.org/10.1111/boj.12038
Liogier, E.E. (1962) Flora de Cuba, vol. 5. Universidad de Puerto Rico, Rio Piedras, Puerto Rico, 324 pp.
Lorence, D.H. (1986) The identity of Portlandia guatemalensis (Rubiaceae). Systematic Botany 11: 209–213. https://doi.org/2418958
Lorence, D.H. (1990) A phylogenetic list of genera of Rubiaceae in Mexico. Acta Botánica Mexicana 12: 1–7. https://doi.org/10.21829/abm12.1990.599
Lorence, D.H. (2012a) Chiococca. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 47–51.
Lorence, D.H. (2012b) Erithalis. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 85–86.
Lorence, D.H., Van Devender, T.R. & Ferguson, G.M. (2018) Chiococca grandiflora (Rubiaceae), a new species from Northern Mexico. PhytoKeys 98: 73–83. https://doi.org/10.3897/phytokeys.98.25170
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. Botanical Sciences 95: 780–806. https://doi.org/10.17129/botsci.1871
McDowell, T. (2012) Exostema. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 167–169.
McDowell, T. & Bremer, B. (1998) Phylogeny, diversity and distribution in Exostema (Rubiaceae): implications of morphological and molecular analyses. Plant Systematics and Evolution 212: 215–246. https://doi.org/10.1007/BF01089740
McDowell, T., Volovsek, M. & Manos, P. (2003) Biogeography of Exostema (Rubiaceae) in the Caribbean region in light of molecular phylogenetic analyses. Systematic Botany 28: 431–441. https://doi.org/10.1043/0363-6445-28.2.431
Manns, U. & Bremer, B. (2010) Towards a better understanding of intertribal relationships and stable tribal delimitations within Cinchonoideae s.s. (Rubiaceae). Molecular Phylogenetics and Evolution 56: 21–39. https://doi.org/10.1016/j.ympev.2010.04.002
Morrone, J.J. (2005) Hacia una síntesis biogeográfica de México. Revista Mexicana de Biodiversidad 76: 207–252. https://doi.org/10.22201/ib.20078706e.2005.002.303
Motley, T.J, Wurdack, K.J. & Delprete, P.G. (2005) Molecular systematics of the Catesbeeae-Chiococceae complex (Rubiaceae) flower and fruit evolution and biogeographic implications. American Journal of Botany 92: 316–329. https://doi.org/10.3732/ajb.92.2.316.
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). Taxon 63: 801–818. https://doi.org/10.12705/634.4
Ochoterena, H. (2012a) Coutaportla. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 69.
Ochoterena, H. (2012b) Coutarea. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 69–70.
Ochoterena, H. (2012c) Hintonia. In: Davidse, G., Sousa, M., Knapp, S. & Chiang, F. (Eds.) Flora Mesoamericana, vol. 4, part 2. Missouri Botanical Garden Press, St. Louis Missouri, pp. 118–119.
Paudyal, S.K., Delprete, P.G. & Motley, T.J. (2014) Using molecular, morphological, and palynological evidence to transfer Strumpfia maritima to the monotypic tribe Strumpfieae (Cinchonoideae, Rubiaceae), and a re-delimitation of the tribe Chiococceae. Systematic Botany 39: 1197–1203. https://doi.org/10.1600/036364414X682580
Paudyal, S.K., Delprete, P.G, Neupane, S. & Motley, T.J. (2018) Molecular phylogenetic analysis and generic delimitations in tribe Chiococceae (Cinchonoideae, Rubiaceae). Botanical Journal of the Linnean Society 187: 365–396. https://doi.org/10.1093/botlinnean/boy029
Quantum GIS Development Team (2018) Quantum GIS Geographic Information System. Open Source Geospatial Foundation Project. [http://qgis.osgeo.org]
Robbrecht, E. (1988) Tropical Woody Rubiaceae. Opera Botanica Belgica 1: 1–272. https://doi.org/10.2307/4110534
Rova, J.H.E., Delprete, P.G., Andersson, L. & Albert, V.A. (2002) A trnL-F cpDNA sequence study of the Condamineae-Rondeletieae-Sipaneeae complex with implications on the phylogeny of Rubiaceae. American Journal of Botany 89: 145–159. https://doi.org/10.3732/ajb.89.1.145.
Rzedowski, J. (1991) Diversidad y orígenes de la flora fanerogámica de México. Acta Botánica Mexicana 14: 13–21. https://doi.org/10.21829/abm14.1991.611
Sazima, M., Buzato, S. & Sazima, I. (1999) Bat-pollinated flower assemblages and bat visitors at two Atlantic forest sites in Brazil. Annals of Botany 83: 705–712. https://doi.org/10.1006/anbo.1999.0876
Schumann, K. (1891) Rubiaceae. In: Engler, A. & Prantl, K. (Eds.) Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten begründet. W. Engelmann, Leipzing, pp. 97–156.
Standley, P.C. (1921) Rubiaceae. North American Flora 32: 1–82.
Standley, P.C. (1934) Rubiales-Rubiaceae. North American Flora 32: 1–300.
Urban, I. (1923) Plantae cubenses novae. Symbolae Antillanae 9: 55–176.
Villarreal, J.A. (1987) Coutaportla pailensis (Rubiaceae), a new species from Coahuila, México. Sida 12: 223–225.
Villaseñor, J.L. (2003) Diversidad y distribución de las Magnoliophyta de México. Interciencia 28: 160–167.
Villaseñor, J.L. (2016) Checklist of the native vascular plants of Mexico. Revista Mexicana de Biodiversidad 87: 559–902. https://doi.org/10.1016/j.rmb.2016.06.017