Abstract
Portulaca is the only genus in Portulacaceae and includes ca. 115 species with worldwide distribution. 12 species (one naturalized) occur in Cuba. The taxonomic value of seeds characters for the circumscription of species, by comparing macro- and micromorphological characters, is analyzed. The seeds of Cuban purslanes do not present association patterns between types of undulations of the anticline walls. The relief of their periclinal walls allows the corroborate certain variation in the distribution range of some species. The delimitation of Portulaca species in Cuba using seeds characters, provided as an identification key and descriptions, confirms the taxonomic value of such characters and strengthens the circumscriptions supported by the rest of the plant structures. This is largely given by the standardization of macromorphological terms with a micromorphological basis, the color of the mature seeds and their dimensions.
References
Amini, M., Sajedi, S. & Domina, G. (2017) First data on the taxonomic diversity of the Portulaca oleracea aggregate (Portulacaceae) in Iran. Turkish Journal of Botany 41: 535–541. http://dx.doi.org/10.3906/bot-1611-43
Barthlott, W. (1981) Epidermal seed surface characters of plants: systematic applicability and some evolutionary aspects. Nordic Journal of Botany 1: 345–355. https://doi.org/10.1111/j.1756-1051.1981.tb00704.x
Coelho, A.A.O.P. & Giulietti, A.M. (2010) O gênero Portulaca L. (Portulacaceae) no Brasil. Acta Botanica Brasilica 24 (3): 655–670. https://doi.org/10.1590/S0102-33062010000300009
Dalavi, J.V., Chougule, R.N., Jadhav, V.D. & Yadav, S.R. (2019) Variation in Seed Micromorphology in Portulaca L. (Portulacaceae) in India. Phytomorphology 69 (1 & 2): 15–24.
Danin, A. & Anderson, L.C. (1986) Distribution of Portulaca oleracea L. (Portulacaceae) subspecies in Florida. Sida 11 (3): 318–324. [https://www.jstor.org/stable/23909826]
Danin, A. & Reyes-Betancort, J.A. (2006) The status of Portulaca oleracea L. in Tenerife, the Canary Islands. Lagascalia 26: 71–81.
Danin, A. & Raus, T. (2012) A key to 19 microspecies of the Portulaca oleracea aggregate. In: Timonin, A.K., Sukhorukov, A.P., Harper, G.H. & Nilova, M.V. (Eds.) Caryophyllales: New insights into phylogeny, systematics, and morphological evolution of the order, Proceedings of the Symposium held on 24th–27th September 2012. Lomonosov State University, Moscow, pp. 70–83.
Danin, A., Baker, I. & Baker, H.G. (1978) Cytogeography and taxonomy of the Portulaca oleracea L. polyploid complex. Israel Journal of Botany 27: 177–211.
Danin, A., Domina, G. & Raimondo, F.M. (2008) Microspecies of the Portulaca oleracea aggregate found on major Mediterranean islands (Sicily, Cyprus, Crete, Rhodes). Flora Mediterranea 18: 89–107.
Danin, A., Buldrini, F., Bandini, M. & Bosi, G. (2014) The history of the Portulaca oleracea aggregate in the Emilia-Romagna Po Plain (Italy) from the Roman Age to the present. Plant Biosystems 148 (4): 622–634. http://dx.doi.org/10.1080/11263504.2013.788098
Danin, A., Buldrini, F., Bandini Mazzanti, M., Bosi, G., Caria, M.C., Dandria, D., Lanfranco, E., Mifsud, S. & Bagella, S. (2016) Diversification of Portulaca oleracea L. complex in the Italian peninsula and adjacent islands. Botany Letters 163: 261–272. https://doi.org/10.1080/23818107.2016.1200482
Denis, D. & Ramírez-Arrieta, V.M. (2020) Si una imagen vale más que mil palabras: ¿cuánto puede decir un gráfico de cajas? Revista del Jardín Botánico Nacional 41: 57–69.
Eggli, U. & Ford-Werntz, D. (2002) Portulacaceae. In: Eggli, U. & Ford-Werntz, D. (Eds.) Illustrated Handbook of Succulent Plants-Dicotyledons. Springer, New York, pp. 370–432. https://doi.org/10.1007/978-3-642-56316-4_48
El-Bakatoushi, R., Alframawy, A.M., Samer, M., El-Sadek, L. & Botros, W. (2013) Evolution of the Portulaca oleracea L. aggregate in Egypt on molecular and phenotypic levels revealed by morphology, inter-simple sequence repeat (ISSR) and 18S rDNA gene sequence markers. Flora 208: 464–477. https://doi.org/10.1016/j.flora.2013.07.008
García-Beltrán, J.A., Barrios, D., Toledo, S. & Testé, E. (2020) Distribución y conservación de Portulaca (Portulacaceae) en Cuba. Botanical Sciences 98 (3): 597–617. https://doi.org/10.17129/botsci.2545
García-Beltrán, J.A., Barrios, D. & Bécquer, E.R. (2021) Revisión taxonómica de Portulacaceae en Cuba. Brittonia 73 (3): 1–31. https://doi.org/10.1007/s12228-021-09653-0
Hernández-Ledesma, P., Berendsohn, W.G., Borsch, Th., Mering, S., Akhani, H., Arias, S., Castañeda-Noa, I., Eggli, U., Eriksson, R., Flores-Olvera, H., Fuentes-Bazán, S., Kadereit, G., Klak, C., Korotkova, N., Nyffeler, R., Ocampo, G., Ochoterena, H., Oxelman, B., Rabeler, R.K., Sanchez, A., Schlumpberger, B.O. & Uotila, P. (2015) A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45: 281–383. https://doi.org/10.3372/wi.45.45301
Judd, W.S. (1981) A monograph of Lyonia (Ericaceae). Journal of the Arnold Arboretum 62 (3): 315–436. https://doi.org/10.5962/bhl.part.11246
Judd, W.S. (2007) Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles. Systematic Botany Monographs 81: 1–235.
Kim, I. (2012) Anatomical and morphological features of seeds in Portulaca. Applled Microscopy 42 (4): 194–199. https://doi.org/10.9729/AM.2012.42.4.194
Legrand, D. (1962) Las especies americanas de Portulaca. Anales del Museo de Historia Natural de Montevideo 7 (3): 9–147.
León, Hno. & Alain, Hno. (1950) Novedades de la Flora Cubana (II). Contribuciones Ocasionales del Museo de Historia Natural del Colegio “De La Salle” 9: 9–424.
Matthews, J.F. (2003) Portulaca. In: Flora of North America Editorial Committee. (Ed.) Flora of North America, vol. 4. Oxford University Press, New York, pp. 496–501.
Matthews, J.F. & Levins, P.A. (1985a) The genus Portulaca in the southeastern United States. Castanea 50 (2): 96–104.
Matthews, J.F. & Levins, P.A. (1985b) Portulaca pilosa L., P. mundula I.M. Johnst. and P. parvula Gray in the southwest. Sida 11 (1): 45–61.
Matthews, J.F. & Levins, P.A. (1986) The systematic significance of seed morphology in Portulaca (Portulacaceae) under scanning electron microscopy. Systematic Botany 11 (2): 302–308. http://dx.doi.org/10.2307/2419120
Matthews, J.F., Ketron, D.W. & Zane, S.F. (1993) The biology and taxonomy of the Portulaca oleracea L. (Portulacaceae) complex in North America. Rhodora 95 (882): 166–183.
Matilla, A., Gallardo, M. and Puga-Hermida, M.I. (2005) Structural, physiological and molecular aspects of heterogeneity in seeds: a review. Seed Science Research 15: 63–76. http://dx.doi.org/10.1079/SSR2005203
Murley, M.R. (1951) Seeds of the Cruciferae of northeastern North America. American Midland Naturalist 46: 1–81. http://dx.doi.org/10.2307/2421948
Nixon, K.C. & Wheeler, Q.D. (1990) An amplification of the phylogenetics species concept. Cladistics 6: 211–223. https://doi.org/10.1111/j.1096-0031.1990.tb00541.x
Nyffeler, R. & Eggli, U. (2010) Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. https://doi.org/10.1002/tax.591021
Ocampo, G. (2013) Morphological characterization of seeds in Portulacaceae. Phytotaxa 14: 1–24. http://dx.doi.org/10.11646/phytotaxa.141.1.1
Ocampo, G. (2014) Systematic implications of seed morphological diversity in Portulacaceae (Caryophyllales). Plant Systematics and Evolution 301 (4): 1215–1226. http://dx.doi.org/10.1007/s00606-014-1146-1
Ocampo, G. & Columbus, J.T. (2012) Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae). Molecular Phylogenetics and Evolution 63: 97–112. http://dx.doi.org/10.1016/j.ympev.2011.12.017
Ortega, M.D. (1985) Aspectos fitogeográficos y ecológicos del género Portulaca L. en Cuba. Feddes Repertorium 96: 713–719. https://doi.org/10.1002/fedr.4910960735
Regalado, L., Ventosa, I. & Morejón, R. (2008) Revisión histórica de los herbarios cubanos con énfasis en las series de especímenes. Revista del Jardín Botánico Nacional 29: 101–138.
Ricceri, C. & Arrigoni, P.V. (2000) L’aggregato di Portulaca oleracea L. (Portulacaceae). Parlatorea 4: 91–97.
Santos, T.A.V. (2016) Estudos micromorfológicos em Portulaca L. (Portulacaceae) do Brasil. Universidade Estadual de Feira de Santana, Feira de Santana, 92 pp.
Santos, T.A.V., Coelho, A.A.O.P., Dórea, M.C., Santos, F.A.R., Leite, K.R.B. & Oliveira, R.P. (2016) Micromorphological features revealing two new species of Portulaca (Portulacaceae) from Brazil, segregated from P. hirsutissima. Phytotaxa 270 (2): 103–115. http://dx.doi.org/10.11646/phytotaxa.270.2.3
StatSoft. (2011) Statistica: data analysis software system, v.10. StatSoft, Tulsa. Available from: http://www.statsoft.com (accessed 17 June 2021)
Stearn, W.T. (2004) Botanical Latin 4<sup>th</sup> ed. Timber Press, Portland, 546 pp.
Thiers, B.M. (2021 onwards) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih. (accessed 17 June 2021)
Walter, J., Vekslyarska, T. & Dobeš, C. (2015) Flow cytometric, chromosomal and morphometric analyses challenge current taxonomic concepts in the Portulaca oleracea complex (Portulacaeae, Caryophyllales). Botanical Journal of the Linnean Society 179: 144–156. https://doi.org/10.1111/boj.12309
Wheeler, Q.D. & Platnick, N.I. (2000) The phylogenetic species concept (sensu Wheeler and Platnick). In: Wheeler, Q.D. & Meier, R. (Eds.) Species concepts and phylogenetic theory: A debate. Columbia University Press, New York, pp. 55–69.
Wilson, P. (1932) Portulaca. In: Britton, N.L., Murrill, W.A. & Barnhart, J.H. (Ed.) North American Flora, vol. 21. The New York Botanical Garden, New York, pp. 328–336.
Barthlott, W. (1981) Epidermal seed surface characters of plants: systematic applicability and some evolutionary aspects. Nordic Journal of Botany 1: 345–355. https://doi.org/10.1111/j.1756-1051.1981.tb00704.x
Coelho, A.A.O.P. & Giulietti, A.M. (2010) O gênero Portulaca L. (Portulacaceae) no Brasil. Acta Botanica Brasilica 24 (3): 655–670. https://doi.org/10.1590/S0102-33062010000300009
Dalavi, J.V., Chougule, R.N., Jadhav, V.D. & Yadav, S.R. (2019) Variation in Seed Micromorphology in Portulaca L. (Portulacaceae) in India. Phytomorphology 69 (1 & 2): 15–24.
Danin, A. & Anderson, L.C. (1986) Distribution of Portulaca oleracea L. (Portulacaceae) subspecies in Florida. Sida 11 (3): 318–324. [https://www.jstor.org/stable/23909826]
Danin, A. & Reyes-Betancort, J.A. (2006) The status of Portulaca oleracea L. in Tenerife, the Canary Islands. Lagascalia 26: 71–81.
Danin, A. & Raus, T. (2012) A key to 19 microspecies of the Portulaca oleracea aggregate. In: Timonin, A.K., Sukhorukov, A.P., Harper, G.H. & Nilova, M.V. (Eds.) Caryophyllales: New insights into phylogeny, systematics, and morphological evolution of the order, Proceedings of the Symposium held on 24th–27th September 2012. Lomonosov State University, Moscow, pp. 70–83.
Danin, A., Baker, I. & Baker, H.G. (1978) Cytogeography and taxonomy of the Portulaca oleracea L. polyploid complex. Israel Journal of Botany 27: 177–211.
Danin, A., Domina, G. & Raimondo, F.M. (2008) Microspecies of the Portulaca oleracea aggregate found on major Mediterranean islands (Sicily, Cyprus, Crete, Rhodes). Flora Mediterranea 18: 89–107.
Danin, A., Buldrini, F., Bandini, M. & Bosi, G. (2014) The history of the Portulaca oleracea aggregate in the Emilia-Romagna Po Plain (Italy) from the Roman Age to the present. Plant Biosystems 148 (4): 622–634. http://dx.doi.org/10.1080/11263504.2013.788098
Danin, A., Buldrini, F., Bandini Mazzanti, M., Bosi, G., Caria, M.C., Dandria, D., Lanfranco, E., Mifsud, S. & Bagella, S. (2016) Diversification of Portulaca oleracea L. complex in the Italian peninsula and adjacent islands. Botany Letters 163: 261–272. https://doi.org/10.1080/23818107.2016.1200482
Denis, D. & Ramírez-Arrieta, V.M. (2020) Si una imagen vale más que mil palabras: ¿cuánto puede decir un gráfico de cajas? Revista del Jardín Botánico Nacional 41: 57–69.
Eggli, U. & Ford-Werntz, D. (2002) Portulacaceae. In: Eggli, U. & Ford-Werntz, D. (Eds.) Illustrated Handbook of Succulent Plants-Dicotyledons. Springer, New York, pp. 370–432. https://doi.org/10.1007/978-3-642-56316-4_48
El-Bakatoushi, R., Alframawy, A.M., Samer, M., El-Sadek, L. & Botros, W. (2013) Evolution of the Portulaca oleracea L. aggregate in Egypt on molecular and phenotypic levels revealed by morphology, inter-simple sequence repeat (ISSR) and 18S rDNA gene sequence markers. Flora 208: 464–477. https://doi.org/10.1016/j.flora.2013.07.008
García-Beltrán, J.A., Barrios, D., Toledo, S. & Testé, E. (2020) Distribución y conservación de Portulaca (Portulacaceae) en Cuba. Botanical Sciences 98 (3): 597–617. https://doi.org/10.17129/botsci.2545
García-Beltrán, J.A., Barrios, D. & Bécquer, E.R. (2021) Revisión taxonómica de Portulacaceae en Cuba. Brittonia 73 (3): 1–31. https://doi.org/10.1007/s12228-021-09653-0
Hernández-Ledesma, P., Berendsohn, W.G., Borsch, Th., Mering, S., Akhani, H., Arias, S., Castañeda-Noa, I., Eggli, U., Eriksson, R., Flores-Olvera, H., Fuentes-Bazán, S., Kadereit, G., Klak, C., Korotkova, N., Nyffeler, R., Ocampo, G., Ochoterena, H., Oxelman, B., Rabeler, R.K., Sanchez, A., Schlumpberger, B.O. & Uotila, P. (2015) A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45: 281–383. https://doi.org/10.3372/wi.45.45301
Judd, W.S. (1981) A monograph of Lyonia (Ericaceae). Journal of the Arnold Arboretum 62 (3): 315–436. https://doi.org/10.5962/bhl.part.11246
Judd, W.S. (2007) Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles. Systematic Botany Monographs 81: 1–235.
Kim, I. (2012) Anatomical and morphological features of seeds in Portulaca. Applled Microscopy 42 (4): 194–199. https://doi.org/10.9729/AM.2012.42.4.194
Legrand, D. (1962) Las especies americanas de Portulaca. Anales del Museo de Historia Natural de Montevideo 7 (3): 9–147.
León, Hno. & Alain, Hno. (1950) Novedades de la Flora Cubana (II). Contribuciones Ocasionales del Museo de Historia Natural del Colegio “De La Salle” 9: 9–424.
Matthews, J.F. (2003) Portulaca. In: Flora of North America Editorial Committee. (Ed.) Flora of North America, vol. 4. Oxford University Press, New York, pp. 496–501.
Matthews, J.F. & Levins, P.A. (1985a) The genus Portulaca in the southeastern United States. Castanea 50 (2): 96–104.
Matthews, J.F. & Levins, P.A. (1985b) Portulaca pilosa L., P. mundula I.M. Johnst. and P. parvula Gray in the southwest. Sida 11 (1): 45–61.
Matthews, J.F. & Levins, P.A. (1986) The systematic significance of seed morphology in Portulaca (Portulacaceae) under scanning electron microscopy. Systematic Botany 11 (2): 302–308. http://dx.doi.org/10.2307/2419120
Matthews, J.F., Ketron, D.W. & Zane, S.F. (1993) The biology and taxonomy of the Portulaca oleracea L. (Portulacaceae) complex in North America. Rhodora 95 (882): 166–183.
Matilla, A., Gallardo, M. and Puga-Hermida, M.I. (2005) Structural, physiological and molecular aspects of heterogeneity in seeds: a review. Seed Science Research 15: 63–76. http://dx.doi.org/10.1079/SSR2005203
Murley, M.R. (1951) Seeds of the Cruciferae of northeastern North America. American Midland Naturalist 46: 1–81. http://dx.doi.org/10.2307/2421948
Nixon, K.C. & Wheeler, Q.D. (1990) An amplification of the phylogenetics species concept. Cladistics 6: 211–223. https://doi.org/10.1111/j.1096-0031.1990.tb00541.x
Nyffeler, R. & Eggli, U. (2010) Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227–240. https://doi.org/10.1002/tax.591021
Ocampo, G. (2013) Morphological characterization of seeds in Portulacaceae. Phytotaxa 14: 1–24. http://dx.doi.org/10.11646/phytotaxa.141.1.1
Ocampo, G. (2014) Systematic implications of seed morphological diversity in Portulacaceae (Caryophyllales). Plant Systematics and Evolution 301 (4): 1215–1226. http://dx.doi.org/10.1007/s00606-014-1146-1
Ocampo, G. & Columbus, J.T. (2012) Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae). Molecular Phylogenetics and Evolution 63: 97–112. http://dx.doi.org/10.1016/j.ympev.2011.12.017
Ortega, M.D. (1985) Aspectos fitogeográficos y ecológicos del género Portulaca L. en Cuba. Feddes Repertorium 96: 713–719. https://doi.org/10.1002/fedr.4910960735
Regalado, L., Ventosa, I. & Morejón, R. (2008) Revisión histórica de los herbarios cubanos con énfasis en las series de especímenes. Revista del Jardín Botánico Nacional 29: 101–138.
Ricceri, C. & Arrigoni, P.V. (2000) L’aggregato di Portulaca oleracea L. (Portulacaceae). Parlatorea 4: 91–97.
Santos, T.A.V. (2016) Estudos micromorfológicos em Portulaca L. (Portulacaceae) do Brasil. Universidade Estadual de Feira de Santana, Feira de Santana, 92 pp.
Santos, T.A.V., Coelho, A.A.O.P., Dórea, M.C., Santos, F.A.R., Leite, K.R.B. & Oliveira, R.P. (2016) Micromorphological features revealing two new species of Portulaca (Portulacaceae) from Brazil, segregated from P. hirsutissima. Phytotaxa 270 (2): 103–115. http://dx.doi.org/10.11646/phytotaxa.270.2.3
StatSoft. (2011) Statistica: data analysis software system, v.10. StatSoft, Tulsa. Available from: http://www.statsoft.com (accessed 17 June 2021)
Stearn, W.T. (2004) Botanical Latin 4<sup>th</sup> ed. Timber Press, Portland, 546 pp.
Thiers, B.M. (2021 onwards) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih. (accessed 17 June 2021)
Walter, J., Vekslyarska, T. & Dobeš, C. (2015) Flow cytometric, chromosomal and morphometric analyses challenge current taxonomic concepts in the Portulaca oleracea complex (Portulacaeae, Caryophyllales). Botanical Journal of the Linnean Society 179: 144–156. https://doi.org/10.1111/boj.12309
Wheeler, Q.D. & Platnick, N.I. (2000) The phylogenetic species concept (sensu Wheeler and Platnick). In: Wheeler, Q.D. & Meier, R. (Eds.) Species concepts and phylogenetic theory: A debate. Columbia University Press, New York, pp. 55–69.
Wilson, P. (1932) Portulaca. In: Britton, N.L., Murrill, W.A. & Barnhart, J.H. (Ed.) North American Flora, vol. 21. The New York Botanical Garden, New York, pp. 328–336.