Abstract
A review of the species Cryptantha maritima (Boraginaceae) supports the continued recognition of the three varieties, these differing in corolla size, ovule and nutlet number, and calyx vestiture. Mapping of these taxa from verified specimens demonstrates some geographic trends, but also some overlap in ranges. In the process of our study, we discovered a new taxonomic entity. Cryptantha maritima var. vizcainensis is described as new. This variety is restricted to the Vizcaíno Desert of Baja California Sur, Mexico and should be considered rare, as it is known to date from only eleven collections. It differs from the other three varieties of the species in having a canescent, appressed-strigose stem vestiture lacking spreading trichomes, in having a relatively large corolla, and in having a fruit derived from two 1-ovuled ovary lobes, developing into two heteromorphic nutlets. We also discovered that the species Cryptantha pondii, previously treated as a synonym of C. patula, should be resurrected as a distinct taxon. Cryptantha pondii is restricted to the western Vizcaíno Desert and to Natividad Island of Baja California Sur. It is morphologically distinctive in having bracteate flowers, relatively large corollas, and four smooth nutlets heteromorphic by size. It is to date known from only three collections and should be considered extremely rare. We also reviewed the morphological and phylogenetic status of Johnstonella echinosepala. This taxon shows similarities to Cryptantha maritima and also exhibits some morphological discontinuity between Pacific and Gulf populations in Baja California Sur. Evidence from both morphological and phylogenetic studies supports the transfer of this species from the genus Johnstonella back to the genus Cryptantha. Finally, we propose that both C. pondii and C. echinosepala are likely close relatives of C. maritima, all of the “Maritimae clade,” a group distantly related to the main core of the genus Cryptantha. This study confirms the great importance of studying herbarium specimens in taxonomic research.
References
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CNPS. (2021) California Native Plant Society Rare Plant Program Inventory of Rare and Endangered Plants of California (online edition, v8-03 0.39) Available from: http://www.rareplants.cnps.org (accessed 15 January 2021)
Coville, F.V. (1895) Botany of the Death Valley Expedition. Contributions from the U. S. National Herbarium 4: 1–318.
Don, G. (1837) A General History of the Dichlamydeous Plants (Original Title: A General System of Gardening and Botany). 4 (Corolliflorae): 1–908. London, UK: Gilbert & Rivington.
GBIF. (2020) Global Biodiversity Information Facility. Available from: http://www.gbif.org (accessed 26 September 2020)
Gray, A. (1883) Contributions to North American botany. Proceedings of the American Academy of Arts and Sciences 19: 1–96. https://doi.org/10.2307/25138721
Gray, A. (1885) Contributions to the botany of North America: A revision of some Borragineous genera. Proceedings of the American Academy of Arts and Sciences 20: 257–310. https://doi.org/10.2307/25138773
Greene, E.L. (1885) Studies in the botany of California and parts adjacent. Bulletin of the California Academy of Sciences 1: 179–228.
Greene, E.L. (1887) Some west American Asperifoliae III. Pittonia 1: 107–120.
Greene, E.L. (1889a) Some plants from the Bay of San Bartolome, Lower California. Pittonia 1: 287–292.
Greene, E.L. (1889b) A list of San Benito plants. Pittonia 1: 263–266.
Greene, E.L. (1892) New or noteworthy species, XII. Pittonia 2: 222–233.
Guilliams, M.C., Hasenstab-Lehman, K., Mabry, M. & Simpson, M.G. (2017) Memoirs of a frequent flier: Phylogenomics reveals 18 long-distance dispersals between North America and South America in the popcorn flowers (Amsinckiinae, Boraginaceae). American Journal of Botany 104: 1717–1728. https://doi.org/10.3732/ajb.1700369
Hasenstab-Lehman, K.E. & Simpson, M.G. (2012) Cat’s eyes and popcorn flowers: Phylogenetic systematics of the genus Cryptantha s.l. (Boraginaceae). Systematic Botany 37: 738–757. http://dx.doi.org/10.1600/036364412X648706
Hooker, W.J. & Arnott, G.A.W. (1840) The Botany of Captain Beechy’s Voyage. Henry G. Bohn, London, 444 pp.
Jiménez-Mejías, P., Cohen, J.I. & Naczi, R.F.C. (2017) The study of online digitized specimens revalidates Andersonglossum boreale as a species different from A. virginianum (Boraginaceae). Phytotaxa 295: 22–34. https://doi.org/10.11646/phytotaxa.295.1.2
Johnston, I.M. (1922) Undescribed plants mostly from Baja California. University of California Publications in Botany 7: 437–446.
Johnston, I.M. (1923) Studies in the Boraginaceae. Novelties and new combinations in the genus Cryptantha. Contributions from the Gray Herbarium of Harvard University 68: 52–57.
Johnston, I.M. (1924) Expedition of the California Academy of Sciences to the Gulf of California in 1921: The botany (the vascular plants). Proceedings of the California Academy of Sciences 12: 951-1218.
Johnston, I.M. (1925) Studies in the Boraginaceae IV. The North American species of Cryptantha. Contributions from the Gray Herbarium of Harvard University 74: 1–114.
Johnston, I.M. (1927) Studies in the Boraginaceae VI. A revision of the South American Boraginoideae. Contributions from the Gray Herbarium of Harvard University 78: 1–118.
Johnston, I.M. (1928) Studies in the Boraginaceae–VII. 2. Notes on various Boraginoideae. Contributions from the Gray Herbarium of Harvard University 81: 73–83.
Johnston, I.M. (1939) Studies in the Boraginaceae, XIII: New or otherwise noteworthy species, chiefly from western United States. Journal of the Arnold Arboretum 20: 375–402. https://doi.org/10.5962/bhl.part.21105
Kartesz, J.T. (2015) The Biota of North America Program (BONAP). North American Plant Atlas. Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2010. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. Available from: http://www.bonap.org/MapSwitchboard.html (accessed 24 November 2020)
Kearney, T.H. & Peebles, R.H. (1960) Arizona Flora. Ed. 2nd, with supplement by John Thomas Howell, Elizabeth McClintock, and collaborators. University of California Press, Berkeley, California, 1085 pp.
Kelley, R.B., Simpson, M.G. & Hasenstab, K.E. (2012) Cryptantha. In: Baldwin, B.G., Goldman, D.H., Keil, D.J., Patterson, R., Rosatti, T.J. & Wilken, D.H. (Eds.) The Jepson Manual: Vascular Plants of California. Second Edition. University of California Press, Berkeley, Los Angeles, London, pp. 455–468.
Kelley, W.A. & Wilken, D. (1993) Cryptantha. In: Hickman, J.C. (Eds.) The Jepson Manual: Higher Plants of California. University of California Press, Berkeley. pp. 369–378.
Mabry, M.E. & Simpson, M.G. (2018) Evaluating the monophyly and biogeography of Cryptantha (Boraginaceae). Systematic Botany 43: 53–76. https://doi.org/10.1600/036364418X696978
Macbride, J.F. (1918) VI. Various American Spermatophytes, new or transferred. Contributions from the Gray Herbarium of Harvard University New Series 56: 50–61.
Nelson, A. & Macbride, J.F. (1916) Western Plant Studies III. Botanical Gazette 61: 30–47. https://doi.org/10.1086/331713
Osterhout, G.E. (1903) New plants from Colorado. Bulletin of the Torrey Botanical Club 30: 236–237. https://doi.org/10.2307/2478782
Philippe, R.A. (1860) Florula Atacamensis; seu, Enumeratio plantarum, quas in itinere per desertum Atacamense. Halis Saxonum, Sumptibus Eduardi Anton, 56 pp. https://doi.org/10.5962/bhl.title.162865
Philippe, R.A. (1891) Catalogus Plantarum in Itinere Tarapacano Lectarum. Anales del Museo Nacional de Chile 8: 1–92.
Rebman, J.P., Gibson, J. & Rich, K. (2016) Annotated checklist of the vascular plants of Baja California, Mexico. Proceedings of the San Diego Society of Natural History Number 45: 1–352.
Rebman, J.P. & Simpson, M.G. (2021) Three new species of Cryptantha (Boraginaceae) from the Southern Channel Islands of California. Madroño. [in press]
Reiche, K. (1907) Estudios Críticos Sobre la Flora de Chile (Continuación): Familia Borrajináceas. Anales de la Universidad de Chile 121: 803–835.
Ruiz, H. & Pavon, J. (1799) Flora Peruviana et Chilensis. Tomus 2: Superiorum Permissu, Typis Gabrielis de Sancha, 70 pp.
SEINet. (2021) SEINet Portal Network. Available from: http//:swbiodiversity.org/seinet/index.php (accessed 1 April 2021)
Simpson, M.G. & Hasenstab, K.E. (2009) Cryptantha of Southern California. Crossosoma 35: 1–59.
Simpson, M.G. & Rebman, J.P. (2013) A new species of Cryptantha (Boraginaceae) from the Sierra de San Pedro Mártir, Baja California, Mexico. Madroño 60: 35–45. https://doi.org/10.3120/0024-9637-60.1.35
Simpson, M.G., Rebman, J.P., Hasenstab-Lehman, K.E., Guilliams, C.M. & McConnell, P.O. (2013) Cryptantha wigginsii (Boraginaceae): A presumed extinct species rediscovered. Madroño 60: 24–34. https://doi.org/10.3120/0024-9637-60.1.24
Simpson, M.G., Dowdy, R., Rebman, J.P., Kelley, R.B. & Simpson, L.M. (2014) Recognition of two species in Eremocarya (Boraginaceae): Evidence from fornix bodies, nutlets, corolla size, and biogeography. Madroño 61: 259–275. https://doi.org/10.3120/0024-9637-61.3.259
Simpson, M.G., Simpson, L.M. & Rebman, J.P. (2016) A new, large-flowered variety of Eremocarya micrantha (Boraginaceae). Madroño 63: 39–54. https://doi.org/10.3120/0024-9637-63.1.39
Simpson, M.G. & Kelley, R.B. (2017) Cryptantha nevadensis var. rigida elevated to species with a new name, C. juniperensis. Phytotaxa 295: 227–236. https://doi.org/10.11646/phytotaxa.295.3.3
Simpson, M.G., Guilliams, C.M., Hasenstab-Lehman, K.E., Mabry, M.E. & Ripma, L. (2017a) Phylogeny of the popcorn flowers: Use of genome skimming to evaluate monophyly and interrelationships in subtribe Amsinckiinae (Boraginaceae). Taxon 66: 1406–1420. https://doi.org/10.12705/666.8
Simpson, M.G., Johnson, L.A., Villaverde, T. & Guilliams, C.M. (2017b) American amphitropical disjuncts: Perspectives from vascular plant analyses and prospects for future research. American Journal of Botany 104: 1600–1650. https://doi.org/10.3732/ajb.1700308
Simpson, M.G., Mabry, M.E. & Hasenstab-Lehman, K. (2019) Transfer of four species of Cryptantha to the genus Johnstonella (Boraginaceae). Phytotaxa 425: 279–289. https://doi.org/10.11646/phytotaxa.425.5.3
Simpson, M.G. & Rebman, J.P. (2021) A new species of Cryptantha restricted to dunes in northwestern Baja California, Mexico. Madroño. [in press]
Thiers, B. (2021) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sycamore.nybg.org/science/ih (accessed 15 January 2021)
Tomlinson, P.B. (1964) The good eye. Carolina Tips 27: 13.
Torrey, J. (1857) Appendix. Article VII. Descriptions of plants collected along the route, by W. P. Blake, and at the mouth of the Gila. In: Williamson, R.S. (Ed.) Report of Explorations in California for Railroad Routes to Connect with the Routes Near the 35th and 32d Parallels of North Latitude 5. Beverley Tucker, Washington D.C., 363 pp.
Villaseñor, J.L. (2016) Checklist of the native vascular plants of Mexico, Catálogo de las plantas vasculares nativas de México. Revista Mexicana de Biodiversidad 87: 559–902. http://dx.doi.org/10.1016/j.rmb.2016.06.017
Wiggins, I.L. (1980) Flora of Baja California. Stanford University Press, Stanford, California, 1025 pp.
Zuloaga, F.O., Morrone, O. & Belgrano, M.J. (Eds.) (2008) Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monographs in Systematic Botany from the Missouri Botanical Garden 107: 1–3348. Available from: http://www2.darwin.edu.ar/Proyectos/FloraArgentina/fa.htm (accessed 28 June 2021)
Baja Flora. (2021) The Flora of Baja California. Available from: http://www.bajaflora.org (accessed 15 January 2021)
Bowdler, N. (2010) Thousands of plant species ‘undiscovered in cupboards’. BBC News Science & Environment. Available from: http://www.bbc.co.uk/news/science-environment-11913076 (accessed 27 November 2020)
Brand, A. (1925) Drei neue Gattungen der Cryptantheae. Repertorium Specierum Novarum Regni Vegetabilis 21: 249–254.
Brandegee, T.S. (1899) New species of western plants. Botanical Gazette 27: 444–457. https://doi.org/10.1086/327855
CCH1. (2021) Data provided by the Consortium of California Herbaria. Available from: http://ucjeps.berkeley.edu/consortium (accessed 1 April 2021)
CCH2. (2021) Consortium of California Herbaria CCH2 Portal. Available from: http://cch2.org/portal/index.php (accessed 15 January 2021)
Chacón, J., Luebert, F., Hilger, H.H., Ovcinnikova, S., Selvi, F., Cecchi, L., Guilliams, C.M., Hasenstab-Lehman, K., Sutorý, K., Simpson, M.G. & Weigend, M. (2016) The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera. Taxon 65: 523–546. https://doi.org/10.12705/653.6
CNPS. (2021) California Native Plant Society Rare Plant Program Inventory of Rare and Endangered Plants of California (online edition, v8-03 0.39) Available from: http://www.rareplants.cnps.org (accessed 15 January 2021)
Coville, F.V. (1895) Botany of the Death Valley Expedition. Contributions from the U. S. National Herbarium 4: 1–318.
Don, G. (1837) A General History of the Dichlamydeous Plants (Original Title: A General System of Gardening and Botany). 4 (Corolliflorae): 1–908. London, UK: Gilbert & Rivington.
GBIF. (2020) Global Biodiversity Information Facility. Available from: http://www.gbif.org (accessed 26 September 2020)
Gray, A. (1883) Contributions to North American botany. Proceedings of the American Academy of Arts and Sciences 19: 1–96. https://doi.org/10.2307/25138721
Gray, A. (1885) Contributions to the botany of North America: A revision of some Borragineous genera. Proceedings of the American Academy of Arts and Sciences 20: 257–310. https://doi.org/10.2307/25138773
Greene, E.L. (1885) Studies in the botany of California and parts adjacent. Bulletin of the California Academy of Sciences 1: 179–228.
Greene, E.L. (1887) Some west American Asperifoliae III. Pittonia 1: 107–120.
Greene, E.L. (1889a) Some plants from the Bay of San Bartolome, Lower California. Pittonia 1: 287–292.
Greene, E.L. (1889b) A list of San Benito plants. Pittonia 1: 263–266.
Greene, E.L. (1892) New or noteworthy species, XII. Pittonia 2: 222–233.
Guilliams, M.C., Hasenstab-Lehman, K., Mabry, M. & Simpson, M.G. (2017) Memoirs of a frequent flier: Phylogenomics reveals 18 long-distance dispersals between North America and South America in the popcorn flowers (Amsinckiinae, Boraginaceae). American Journal of Botany 104: 1717–1728. https://doi.org/10.3732/ajb.1700369
Hasenstab-Lehman, K.E. & Simpson, M.G. (2012) Cat’s eyes and popcorn flowers: Phylogenetic systematics of the genus Cryptantha s.l. (Boraginaceae). Systematic Botany 37: 738–757. http://dx.doi.org/10.1600/036364412X648706
Hooker, W.J. & Arnott, G.A.W. (1840) The Botany of Captain Beechy’s Voyage. Henry G. Bohn, London, 444 pp.
Jiménez-Mejías, P., Cohen, J.I. & Naczi, R.F.C. (2017) The study of online digitized specimens revalidates Andersonglossum boreale as a species different from A. virginianum (Boraginaceae). Phytotaxa 295: 22–34. https://doi.org/10.11646/phytotaxa.295.1.2
Johnston, I.M. (1922) Undescribed plants mostly from Baja California. University of California Publications in Botany 7: 437–446.
Johnston, I.M. (1923) Studies in the Boraginaceae. Novelties and new combinations in the genus Cryptantha. Contributions from the Gray Herbarium of Harvard University 68: 52–57.
Johnston, I.M. (1924) Expedition of the California Academy of Sciences to the Gulf of California in 1921: The botany (the vascular plants). Proceedings of the California Academy of Sciences 12: 951-1218.
Johnston, I.M. (1925) Studies in the Boraginaceae IV. The North American species of Cryptantha. Contributions from the Gray Herbarium of Harvard University 74: 1–114.
Johnston, I.M. (1927) Studies in the Boraginaceae VI. A revision of the South American Boraginoideae. Contributions from the Gray Herbarium of Harvard University 78: 1–118.
Johnston, I.M. (1928) Studies in the Boraginaceae–VII. 2. Notes on various Boraginoideae. Contributions from the Gray Herbarium of Harvard University 81: 73–83.
Johnston, I.M. (1939) Studies in the Boraginaceae, XIII: New or otherwise noteworthy species, chiefly from western United States. Journal of the Arnold Arboretum 20: 375–402. https://doi.org/10.5962/bhl.part.21105
Kartesz, J.T. (2015) The Biota of North America Program (BONAP). North American Plant Atlas. Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2010. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. Available from: http://www.bonap.org/MapSwitchboard.html (accessed 24 November 2020)
Kearney, T.H. & Peebles, R.H. (1960) Arizona Flora. Ed. 2nd, with supplement by John Thomas Howell, Elizabeth McClintock, and collaborators. University of California Press, Berkeley, California, 1085 pp.
Kelley, R.B., Simpson, M.G. & Hasenstab, K.E. (2012) Cryptantha. In: Baldwin, B.G., Goldman, D.H., Keil, D.J., Patterson, R., Rosatti, T.J. & Wilken, D.H. (Eds.) The Jepson Manual: Vascular Plants of California. Second Edition. University of California Press, Berkeley, Los Angeles, London, pp. 455–468.
Kelley, W.A. & Wilken, D. (1993) Cryptantha. In: Hickman, J.C. (Eds.) The Jepson Manual: Higher Plants of California. University of California Press, Berkeley. pp. 369–378.
Mabry, M.E. & Simpson, M.G. (2018) Evaluating the monophyly and biogeography of Cryptantha (Boraginaceae). Systematic Botany 43: 53–76. https://doi.org/10.1600/036364418X696978
Macbride, J.F. (1918) VI. Various American Spermatophytes, new or transferred. Contributions from the Gray Herbarium of Harvard University New Series 56: 50–61.
Nelson, A. & Macbride, J.F. (1916) Western Plant Studies III. Botanical Gazette 61: 30–47. https://doi.org/10.1086/331713
Osterhout, G.E. (1903) New plants from Colorado. Bulletin of the Torrey Botanical Club 30: 236–237. https://doi.org/10.2307/2478782
Philippe, R.A. (1860) Florula Atacamensis; seu, Enumeratio plantarum, quas in itinere per desertum Atacamense. Halis Saxonum, Sumptibus Eduardi Anton, 56 pp. https://doi.org/10.5962/bhl.title.162865
Philippe, R.A. (1891) Catalogus Plantarum in Itinere Tarapacano Lectarum. Anales del Museo Nacional de Chile 8: 1–92.
Rebman, J.P., Gibson, J. & Rich, K. (2016) Annotated checklist of the vascular plants of Baja California, Mexico. Proceedings of the San Diego Society of Natural History Number 45: 1–352.
Rebman, J.P. & Simpson, M.G. (2021) Three new species of Cryptantha (Boraginaceae) from the Southern Channel Islands of California. Madroño. [in press]
Reiche, K. (1907) Estudios Críticos Sobre la Flora de Chile (Continuación): Familia Borrajináceas. Anales de la Universidad de Chile 121: 803–835.
Ruiz, H. & Pavon, J. (1799) Flora Peruviana et Chilensis. Tomus 2: Superiorum Permissu, Typis Gabrielis de Sancha, 70 pp.
SEINet. (2021) SEINet Portal Network. Available from: http//:swbiodiversity.org/seinet/index.php (accessed 1 April 2021)
Simpson, M.G. & Hasenstab, K.E. (2009) Cryptantha of Southern California. Crossosoma 35: 1–59.
Simpson, M.G. & Rebman, J.P. (2013) A new species of Cryptantha (Boraginaceae) from the Sierra de San Pedro Mártir, Baja California, Mexico. Madroño 60: 35–45. https://doi.org/10.3120/0024-9637-60.1.35
Simpson, M.G., Rebman, J.P., Hasenstab-Lehman, K.E., Guilliams, C.M. & McConnell, P.O. (2013) Cryptantha wigginsii (Boraginaceae): A presumed extinct species rediscovered. Madroño 60: 24–34. https://doi.org/10.3120/0024-9637-60.1.24
Simpson, M.G., Dowdy, R., Rebman, J.P., Kelley, R.B. & Simpson, L.M. (2014) Recognition of two species in Eremocarya (Boraginaceae): Evidence from fornix bodies, nutlets, corolla size, and biogeography. Madroño 61: 259–275. https://doi.org/10.3120/0024-9637-61.3.259
Simpson, M.G., Simpson, L.M. & Rebman, J.P. (2016) A new, large-flowered variety of Eremocarya micrantha (Boraginaceae). Madroño 63: 39–54. https://doi.org/10.3120/0024-9637-63.1.39
Simpson, M.G. & Kelley, R.B. (2017) Cryptantha nevadensis var. rigida elevated to species with a new name, C. juniperensis. Phytotaxa 295: 227–236. https://doi.org/10.11646/phytotaxa.295.3.3
Simpson, M.G., Guilliams, C.M., Hasenstab-Lehman, K.E., Mabry, M.E. & Ripma, L. (2017a) Phylogeny of the popcorn flowers: Use of genome skimming to evaluate monophyly and interrelationships in subtribe Amsinckiinae (Boraginaceae). Taxon 66: 1406–1420. https://doi.org/10.12705/666.8
Simpson, M.G., Johnson, L.A., Villaverde, T. & Guilliams, C.M. (2017b) American amphitropical disjuncts: Perspectives from vascular plant analyses and prospects for future research. American Journal of Botany 104: 1600–1650. https://doi.org/10.3732/ajb.1700308
Simpson, M.G., Mabry, M.E. & Hasenstab-Lehman, K. (2019) Transfer of four species of Cryptantha to the genus Johnstonella (Boraginaceae). Phytotaxa 425: 279–289. https://doi.org/10.11646/phytotaxa.425.5.3
Simpson, M.G. & Rebman, J.P. (2021) A new species of Cryptantha restricted to dunes in northwestern Baja California, Mexico. Madroño. [in press]
Thiers, B. (2021) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sycamore.nybg.org/science/ih (accessed 15 January 2021)
Tomlinson, P.B. (1964) The good eye. Carolina Tips 27: 13.
Torrey, J. (1857) Appendix. Article VII. Descriptions of plants collected along the route, by W. P. Blake, and at the mouth of the Gila. In: Williamson, R.S. (Ed.) Report of Explorations in California for Railroad Routes to Connect with the Routes Near the 35th and 32d Parallels of North Latitude 5. Beverley Tucker, Washington D.C., 363 pp.
Villaseñor, J.L. (2016) Checklist of the native vascular plants of Mexico, Catálogo de las plantas vasculares nativas de México. Revista Mexicana de Biodiversidad 87: 559–902. http://dx.doi.org/10.1016/j.rmb.2016.06.017
Wiggins, I.L. (1980) Flora of Baja California. Stanford University Press, Stanford, California, 1025 pp.
Zuloaga, F.O., Morrone, O. & Belgrano, M.J. (Eds.) (2008) Catálogo de las Plantas Vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monographs in Systematic Botany from the Missouri Botanical Garden 107: 1–3348. Available from: http://www2.darwin.edu.ar/Proyectos/FloraArgentina/fa.htm (accessed 28 June 2021)