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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<p>Johnston, I.M. (1927) Studies in the Boraginaceae VI. A revision of the South American Boraginoideae. <em>Contributions from the Gray Herbarium of Harvard University</em> 78: 1–118.</p>
<p>Johnston, I.M. (1928) Studies in the Boraginaceae–VII. 2. Notes on various Boraginoideae. <em>Contributions from the Gray Herbarium of Harvard University</em> 81: 73–83.</p>
<p>Johnston, I.M. (1939) Studies in the Boraginaceae, XIII: New or otherwise noteworthy species, chiefly from western United States. <em>Journal of the Arnold Arboretum</em> 20: 375–402. https://doi.org/10.5962/bhl.part.21105</p>
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<p>Kelley, R.B., Simpson, M.G. & Hasenstab, K.E. (2012) <em>Cryptantha</em>.<em> In: </em>Baldwin, B.G., Goldman, D.H., Keil, D.J., Patterson, R., Rosatti, T.J. & Wilken, D.H. (Eds.) <em>The Jepson Manual: Vascular Plants of California. Second Edition</em>. University of California Press, Berkeley, Los Angeles, London, pp. 455–468.</p>
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<p>Mabry, M.E. & Simpson, M.G. (2018) Evaluating the monophyly and biogeography of <em>Cryptantha</em> (Boraginaceae). <em>Systematic Botany</em> 43: 53–76. https://doi.org/10.1600/036364418X696978</p>
<p>Macbride, J.F. (1918) VI. Various American Spermatophytes, new or transferred. <em>Contributions from the Gray Herbarium of Harvard University New Series</em> 56: 50–61.</p>
<p>Nelson, A. & Macbride, J.F. (1916) Western Plant Studies III. <em>Botanical Gazette</em> 61: 30–47. https://doi.org/10.1086/331713</p>
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<p>Rebman, J.P. & Simpson, M.G. (2021) Three new species of <em>Cryptantha</em> (Boraginaceae) from the Southern Channel Islands of California. <em>Madroño.</em> [in press]</p>
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<p>Simpson, M.G. & Rebman, J.P. (2013) A new species of <em>Cryptantha</em> (Boraginaceae) from the Sierra de San Pedro Mártir, Baja California, Mexico. <em>Madroño</em> 60: 35–45. https://doi.org/10.3120/0024-9637-60.1.35</p>
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<p> </p>
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<p>Brand, A. (1925) Drei neue Gattungen der Cryptantheae. <em>Repertorium Specierum Novarum Regni Vegetabilis</em> 21: 249–254.</p>
<p>Brandegee, T.S. (1899) New species of western plants. <em>Botanical Gazette</em> 27: 444–457. https://doi.org/10.1086/327855</p>
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<p>Don, G. (1837) <em>A General History of the Dichlamydeous Plants (Original Title: A General System of Gardening and Botany)</em>. 4 (Corolliflorae): 1–908. London, UK: Gilbert & Rivington.</p>
<p>GBIF. (2020) <em>Global Biodiversity Information Facility</em>. Available from: http://www.gbif.org (accessed 26 September 2020)</p>
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<p>Gray, A. (1885) Contributions to the botany of North America: A revision of some Borragineous genera. <em>Proceedings of the American Academy of Arts and Sciences </em>20: 257–310. https://doi.org/10.2307/25138773</p>
<p>Greene, E.L. (1885) Studies in the botany of California and parts adjacent. <em>Bulletin of the California Academy of Sciences</em> 1: 179–228.</p>
<p>Greene, E.L. (1887) Some west American Asperifoliae III. <em>Pittonia</em> 1: 107–120.</p>
<p>Greene, E.L. (1889a) Some plants from the Bay of San Bartolome, Lower California. <em>Pittonia</em> 1: 287–292.</p>
<p>Greene, E.L. (1889b) A list of San Benito plants. <em>Pittonia</em> 1: 263–266.</p>
<p>Greene, E.L. (1892) New or noteworthy species, XII. <em>Pittonia</em> 2: 222–233.</p>
<p>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). <em>American Journal of Botany</em> 104: 1717–1728. https://doi.org/10.3732/ajb.1700369</p>
<p>Hasenstab-Lehman, K.E. & Simpson, M.G. (2012) Cat’s eyes and popcorn flowers: Phylogenetic systematics of the genus <em>Cryptantha</em> s.l. (Boraginaceae). <em>Systematic Botany</em> 37: 738–757. http://dx.doi.org/10.1600/036364412X648706</p>
<p>Hooker, W.J. & Arnott, G.A.W. (1840) <em>The Botany of Captain Beechy’s Voyage</em>. Henry G. Bohn, London, 444 pp.</p>
<p>Jiménez-Mejías, P., Cohen, J.I. & Naczi, R.F.C. (2017) The study of online digitized specimens revalidates <em>Andersonglossum boreale</em> as a species different from <em>A. virginianum</em> (Boraginaceae). <em>Phytotaxa</em> 295: 22–34. https://doi.org/10.11646/phytotaxa.295.1.2</p>
<p>Johnston, I.M. (1922) Undescribed plants mostly from Baja California. <em>University of California Publications in Botany</em> 7: 437–446.</p>
<p>Johnston, I.M. (1923) Studies in the Boraginaceae. Novelties and new combinations in the genus <em>Cryptantha</em>. <em>Contributions from the Gray Herbarium of Harvard University</em> 68: 52–57.</p>
<p>Johnston, I.M. (1924) Expedition of the California Academy of Sciences to the Gulf of California in 1921: The botany (the vascular plants). <em>Proceedings of the California Academy of Sciences</em> 12: 951-1218.</p>
<p>Johnston, I.M. (1925) Studies in the Boraginaceae IV. The North American species of <em>Cryptantha</em>. <em>Contributions from the Gray Herbarium of Harvard University</em> 74: 1–114.</p>
<p>Johnston, I.M. (1927) Studies in the Boraginaceae VI. A revision of the South American Boraginoideae. <em>Contributions from the Gray Herbarium of Harvard University</em> 78: 1–118.</p>
<p>Johnston, I.M. (1928) Studies in the Boraginaceae–VII. 2. Notes on various Boraginoideae. <em>Contributions from the Gray Herbarium of Harvard University</em> 81: 73–83.</p>
<p>Johnston, I.M. (1939) Studies in the Boraginaceae, XIII: New or otherwise noteworthy species, chiefly from western United States. <em>Journal of the Arnold Arboretum</em> 20: 375–402. https://doi.org/10.5962/bhl.part.21105</p>
<p>Kartesz, J.T. (2015) <em>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)].</em> Available from: http://www.bonap.org/MapSwitchboard.html (accessed 24 November 2020)</p>
<p>Kearney, T.H. & Peebles, R.H. (1960) <em>Arizona Flora</em>. Ed. 2nd, with supplement by John Thomas Howell, Elizabeth McClintock, and collaborators. University of California Press, Berkeley, California, 1085 pp.</p>
<p>Kelley, R.B., Simpson, M.G. & Hasenstab, K.E. (2012) <em>Cryptantha</em>.<em> In: </em>Baldwin, B.G., Goldman, D.H., Keil, D.J., Patterson, R., Rosatti, T.J. & Wilken, D.H. (Eds.) <em>The Jepson Manual: Vascular Plants of California. Second Edition</em>. University of California Press, Berkeley, Los Angeles, London, pp. 455–468.</p>
<p>Kelley, W.A. & Wilken, D. (1993) <em>Cryptantha</em>.<em> In: </em>Hickman, J.C. (Eds.) <em>The Jepson Manual: Higher Plants of California</em>. University of California Press, Berkeley. pp. 369–378.</p>
<p>Mabry, M.E. & Simpson, M.G. (2018) Evaluating the monophyly and biogeography of <em>Cryptantha</em> (Boraginaceae). <em>Systematic Botany</em> 43: 53–76. https://doi.org/10.1600/036364418X696978</p>
<p>Macbride, J.F. (1918) VI. Various American Spermatophytes, new or transferred. <em>Contributions from the Gray Herbarium of Harvard University New Series</em> 56: 50–61.</p>
<p>Nelson, A. & Macbride, J.F. (1916) Western Plant Studies III. <em>Botanical Gazette</em> 61: 30–47. https://doi.org/10.1086/331713</p>
<p>Osterhout, G.E. (1903) New plants from Colorado. <em>Bulletin of the Torrey Botanical Club</em> 30: 236–237. https://doi.org/10.2307/2478782</p>
<p>Philippe, R.A. (1860) <em>Florula Atacamensis; seu, Enumeratio plantarum, quas in itinere per desertum Atacamense</em>. Halis Saxonum, Sumptibus Eduardi Anton, 56 pp. https://doi.org/10.5962/bhl.title.162865</p>
<p>Philippe, R.A. (1891) Catalogus Plantarum in Itinere Tarapacano Lectarum. <em>Anales del Museo Nacional de Chile</em> 8: 1–92.</p>
<p>Rebman, J.P., Gibson, J. & Rich, K. (2016) Annotated checklist of the vascular plants of Baja California, Mexico. <em>Proceedings of the San Diego Society of Natural History</em> Number 45: 1–352.</p>
<p>Rebman, J.P. & Simpson, M.G. (2021) Three new species of <em>Cryptantha</em> (Boraginaceae) from the Southern Channel Islands of California. <em>Madroño.</em> [in press]</p>
<p>Reiche, K. (1907) Estudios Críticos Sobre la Flora de Chile (Continuación): Familia Borrajináceas. <em>Anales de la Universidad de Chile</em> 121: 803–835.</p>
<p>Ruiz, H. & Pavon, J. (1799) <em>Flora Peruviana et Chilensis</em>. Tomus 2: Superiorum Permissu, Typis Gabrielis de Sancha, 70 pp.</p>
<p>SEINet. (2021) <em>SEINet Portal Network</em>. Available from: http//:swbiodiversity.org/seinet/index.php (accessed 1 April 2021)</p>
<p>Simpson, M.G. & Hasenstab, K.E. (2009) <em>Cryptantha</em> of Southern California. <em>Crossosoma</em> 35: 1–59.</p>
<p>Simpson, M.G. & Rebman, J.P. (2013) A new species of <em>Cryptantha</em> (Boraginaceae) from the Sierra de San Pedro Mártir, Baja California, Mexico. <em>Madroño</em> 60: 35–45. https://doi.org/10.3120/0024-9637-60.1.35</p>
<p>Simpson, M.G., Rebman, J.P., Hasenstab-Lehman, K.E., Guilliams, C.M. & McConnell, P.O. (2013) <em>Cryptantha wigginsii</em> (Boraginaceae): A presumed extinct species rediscovered. <em>Madroño</em> 60: 24–34. https://doi.org/10.3120/0024-9637-60.1.24</p>
<p>Simpson, M.G., Dowdy, R., Rebman, J.P., Kelley, R.B. & Simpson, L.M. (2014) Recognition of two species in <em>Eremocarya</em> (Boraginaceae): Evidence from fornix bodies, nutlets, corolla size, and biogeography. <em>Madroño</em> 61: 259–275. https://doi.org/10.3120/0024-9637-61.3.259</p>
<p>Simpson, M.G., Simpson, L.M. & Rebman, J.P. (2016) A new, large-flowered variety of <em>Eremocarya micrantha</em> (Boraginaceae). <em>Madroño</em> 63: 39–54. https://doi.org/10.3120/0024-9637-63.1.39</p>
<p>Simpson, M.G. & Kelley, R.B. (2017) <em>Cryptantha nevadensis</em> var. <em>rigida</em> elevated to species with a new name, <em>C. juniperensis</em>. <em>Phytotaxa</em> 295: 227–236. https://doi.org/10.11646/phytotaxa.295.3.3</p>
<p>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). <em>Taxon</em> 66: 1406–1420. https://doi.org/10.12705/666.8</p>
<p>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. <em>American Journal of Botany</em> 104: 1600–1650. https://doi.org/10.3732/ajb.1700308</p>
<p>Simpson, M.G., Mabry, M.E. & Hasenstab-Lehman, K. (2019) Transfer of four species of <em>Cryptantha</em> to the genus <em>Johnstonella</em> (Boraginaceae). <em>Phytotaxa</em> 425: 279–289. https://doi.org/10.11646/phytotaxa.425.5.3</p>
<p>Simpson, M.G. & Rebman, J.P. (2021) A new species of <em>Cryptantha</em> restricted to dunes in northwestern Baja California, Mexico. <em>Madroño.</em> [in press]</p>
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