Abstract
The genus Utricularia (Lentibulariaceae) is the most diverse and cosmopolitan among carnivorous plants. Utricularia sect. Foliosa sensu Taylor (1989) is composed of three South American species, U. amethystina, U. tricolor and U. tridentata, the first species being recognized as highly diverse. In his taxonomic revision of the genus Utricularia, Taylor (1989) circumscribed the polymorphic U. amethystina comprising 27 synonyms involving a wide variation in corolla size and colour, morphology of corolla parts, indumentum, and overall plant size. The original publications of the synonyms provide limited information disencouraging a revision study and therefore inflating U. amethystina as new variations were discovered. In order to shed light onto the problem, a taxonomic revision, a morphometric and a molecular study showed evidence of a larger complex than previously thought, and U. amethystina s.str. sister to U. tricolor. A previously published molecular phylogeny provided strong support for all sampled U. amethystina synonyms and potential new taxa found to be nested outside the U. amethystina s.str. and U. tricolor clade. Here, we incorporate these new relationships and describe and comment on the species that comprise U. sect. Foliosa, including the restablishment of six synonyms, the description of three new species and lectotypification of five names. The current study disentangles U. amethystina complex so that further studies can approach it in a more realistic way, especially studies using population genetics tools. Our study shows that more needs to be done and here U. hirtella is presented as a new complex.
References
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Barnhart, J.H. (1931) Lentibulariaceae. Botanical results of the Tyler-Duida expedition (continued). Bulletin of the Torrey Botanical Club 58 (8): 468–469. https://doi.org/10.2307/2480621
Barnhart, J.H. (1933) Pinguiculaceae. In: Small, J.K. (Ed.) Manual of Southeastern Flora. The University of North Carolina Press, Chapel Hill, pp. 1231–1238.
Beauverd, G. (1907) Plantae Damazianae Brasilienses, Determinées par différents botanistes. Utriculariaceae. Bulletin de l’Herbier Boissier, sér. 2 7: 702–703.
Benjamin, L. (1847) Lentibularieae. Linnaea 20: 488.
Benjamin, L. (1847) Utricularieae. Flora Brasiliensis 10: 222–256.
Berry, P.E. & Riina, R. (2005) Insights into the diversity of the Pantepui Flora and the biogeographic complexity of the Guayana Shield. Biologiske Skrifter 55: 145–167.
Brown, N.E. (1901) Lentibulariaceae. Transactions of the Linnean Society of London, 2nd series: Botany 6: 52–56. https://doi.org/10.1111/j.1095-8339.1901.tb00001.x
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Guedes, F.M., Coan, A.I., Araújo, G.B. & Alves, M. (2021) The identity of Utricularia trinervia (Lentibulariaceae): Taxonomic re-establishment and unpublished descriptions of trap morphology. Taxon 70 (4): 854–865. https://doi.org/10.1002/tax.12496
Hicken, C.M. (1912) Dos nuevas plantas para la Flora Uruguaya. Boletin de la Sociedad Physis 1: 179–181.
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Kelloff, C.L. & Funk, V.A. (2004) Phytogeography of the Kaieteur Falls, Potaro Plateau, Guyana: Floral distributions and affinities. Journal of Biogeography 31 (3): 501–513. https://doi.org/10.1046/j.0305-0270.2003.01038.x
Kok, P.J.R. (2010) A new species of Chironius Fitzinger, 1826 (Squamata: Colubridae) from the Pantepui region, northeastern South America. Zootaxa 2611: 31–44. https://doi.org/10.11646/zootaxa.2611.1.3
Kok, P.J.R., Ratz, S., Tegelaar, M., Aubret, F. & Means, D.B. (2015) Out of taxonomic limbo: A name for the species of Tepuihyla (Anura: Hylidae) from the Chimantá Massif, Pantepui region, northern South America. Salamandra 51: 283–314.
Kok, P.J.R., Nicolai, M.P.J., Lathrop, A. & MacCulloch, R.D. (2018) Anomaloglossus meansi sp. n., a new Pantepui species of the Anomaloglossus beebei group (Anura, Aromobatidae). ZooKeys 759: 99–116. https://doi.org/10.3897/zookeys.759.24742
Kuntze, C.E.O. (1898) Lentibulariaceae. Revisio Generum Plantarum 3 (3): 240.
Kurz, W.S. (1874) Description of Utricularia nivea Vahl. Journal of Botany, British and Foreign 12: 53–54.
Larrañaga, D.A. (1923) Escritos de Don Damaso Antonio Larrañaga 2. Instituto histórico y geográfico del Urugay, Imprenta Nacional, Montevideo, 511 pp. https://doi.org/10.5962/bhl.title.116606
Lehmann, J.G.C. (1844) Utricularia. Botanische Zeitung (Berlin) 2: 109–110.
Linnaeus, C. (1753) Species Plantarum 1. Impensis Laurentii Salvii, Holmiae, 572 pp.
Menezes, C.G., Gasparine, E.C., Baleeiro, P.C. & de Miranda, V.F.O. (2014) Seed morphology of bladderworts: A survey on Utricularia sect. Foliosa and sect. Psyllosperma (Lentibulariaceae) with taxonomic implications. Phytotaxa 167 (2): 173–182. https://doi.org/10.11646/phytotaxa.167.2.3
Miquel, F.A.W. (1847) Lentibulariaceae. Linnaea 19: 144–145.
Pilger, R.K.F. (1914) Lentibulariaceae. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 6: 188–191. https://doi.org/10.2307/3994403
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Salerno, P.E., Ron, S.R., Señaris, J.C., Rojas-Runjaic, F.J.M., Noonam, B.P. & Cannatella, D.C. (2012) Ancient tepui summits harbor young rather than old lineages of endemic frogs. Evolution 66 (10): 3000–3013. https://doi.org/10.1111/j.1558-5646.2012.01666.x
Silva, N.G. da, Alves, R.J.V., Pereira, J.F. & Rivadavia, F. (2011) Lentibulariaceae, Serra de São José, Minas Gerais, Brazil. Check List 7 (2): 120–127. https://doi.org/10.15560/7.2.120
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Baleeiro, P.C., Sano, P.T. & Jobson, R.W. (2019) Molecular Phylogeny of the Utricularia amethystina complex (Utricularia sect. Foliosa) assessed using plastid and nuclear sequence data. Systematic Botany 44 (2): 398–404. https://doi.org/10.1600/036364419X15562052252045
Barbosa-Silva, R.G., Trovo, M., Martinelli, G. & Forzza, R.C. (2018) Up on the table mountains in Brazil: New Bromeliaceae and Eriocaulaceae (Poales) from the Pantepui in the Guayana Shield. Plant Ecology and Evolution 151 (1): 130–141. https://doi.org/10.5091/plecevo.2018.1392
Barnhart, J.H. (1931) Lentibulariaceae. Botanical results of the Tyler-Duida expedition (continued). Bulletin of the Torrey Botanical Club 58 (8): 468–469. https://doi.org/10.2307/2480621
Barnhart, J.H. (1933) Pinguiculaceae. In: Small, J.K. (Ed.) Manual of Southeastern Flora. The University of North Carolina Press, Chapel Hill, pp. 1231–1238.
Beauverd, G. (1907) Plantae Damazianae Brasilienses, Determinées par différents botanistes. Utriculariaceae. Bulletin de l’Herbier Boissier, sér. 2 7: 702–703.
Benjamin, L. (1847) Lentibularieae. Linnaea 20: 488.
Benjamin, L. (1847) Utricularieae. Flora Brasiliensis 10: 222–256.
Berry, P.E. & Riina, R. (2005) Insights into the diversity of the Pantepui Flora and the biogeographic complexity of the Guayana Shield. Biologiske Skrifter 55: 145–167.
Brown, N.E. (1901) Lentibulariaceae. Transactions of the Linnean Society of London, 2nd series: Botany 6: 52–56. https://doi.org/10.1111/j.1095-8339.1901.tb00001.x
CNC Flora (2021) Utricularia tridentata in Lista Vermelha da flora brasileira versão 2012.2 Centro Nacional de Conservação da Flora. [http://cncflora.jbrj.gov.br/portal/pt-br/profile/Utricularia tridentata]
Coelho, N., Gomes-Klein, V.L. & Dantas-Queiroz, M.V. (2017) Lentibulariaceae from Serra dos Pireneus, Goiás, Brazil. Rodriguesia 68 (4): 1347–1356. https://doi.org/10.1590/2175-7860201768416
Cross, A.T., Krueger, T.A., Gonella, P.M., Robinson, A.S. & Fleischmann, A.S. (2020) Conservation of carnivorous plants in the age of extinction. Global Ecology and Conservation 24: e01272. https://doi.org/10.1016/j.gecco.2020.e01272
De Candolle, A.L.P.P. (1844) Lentibularieae. Prodromus Systematis Naturalis Regni Vegetabilis 8. Sumptibus Sociorum Treuttel et Wu?rtz, Parisii, pp. 2–32.
de Vriese, W.H. (1848) Lentibularieae. Nederlandsch Kruidkundig Archief 1: 323–325.
Funk, V., Hollowell, T., Berry, P., Kelloff, C. & Alexander, S.N. (2007) Checklist of the Plants of the Guiana Shield (Venezuela?: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana). Contributions from the United States National Herbarium 55: 1–584.
Giulietti, A.M., Pirani, J.R. & Harley, R.M. (1997) Espinhaço range region. eastern brazil. Centres of plant diversity. a guide and strategies for the conservation, Vol. 3: The Americas. WWF & IUCN, Cambridge, pp. 397–404.
Gleason, H.A. (1929) Studies on the Flora of northern South America XIII. The Tate collection from Mount Roraima and vicinity. Bulletin of the Torrey Botanical Club 56: 391–408. https://doi.org/10.2307/2480451
Gonella, P.M. & Baleeiro, P.C. (2018) Utricularia biceps (Lentibulariaceae), a new carnivorous species endemic to the campos rupestres of brazil. Phytotaxa 376 (5): 214–222. https://doi.org/10.11646/phytotaxa.376.5.4
Grace, O.M., Pérez-Escobar, O.A., Lucas, E.J., Vorontsova, M.S., Lewis, G.P., Walker, B.E., Lohman, L.G., Knapp, S., Wilkie, P., Sarkinen, T., Darbyshire, L., Lughadha, E.N., Monro, A., Woudstra, Y., Demissew, S., Muasya, A.M., Díaz, S., Baker, W.J. & Antonelli, A. (2021) Botanical monography in the Anthropocene. Trends in Plant Science 26 (5): 433–441. https://doi.org/10.1016/j.tplants.2020.12.018
Guedes, F.M., Coan, A.I., Araújo, G.B. & Alves, M. (2021) The identity of Utricularia trinervia (Lentibulariaceae): Taxonomic re-establishment and unpublished descriptions of trap morphology. Taxon 70 (4): 854–865. https://doi.org/10.1002/tax.12496
Hicken, C.M. (1912) Dos nuevas plantas para la Flora Uruguaya. Boletin de la Sociedad Physis 1: 179–181.
Hijmans, R.J., Guarino, L. & Mathur, P. (2012) DIVA-GIS Version 7.5 Manual. [http://www.diva-gis.org]
IUCN (2012) IUCN Red List Categories and Criteria. Version 3. Edited by P. by the I. S. S. Commission. Gland, Switzerland and Cambridge, UK. [https://portals.iucn.org/library/node/10315]
Jobson, R.W., Baleeiro, P.C. & Guisande, C. (2018) Systematics and evolution of Lentibulariaceae: III. Utricularia. In: Ellison, A.M. & Adamec, L. (Eds.) Carnivorous Plants: Physiology, Ecology, and Evolution. Oxford University Press, Oxford, pp. 89–104. https://doi.org/10.1093/oso/9780198779841.003.0008
Kamie?ski, F. (1891) Lentibulariaceae. In: Engler, A. & Prantl, K.A.E. (Eds.) Die natürlichen Pflanzenfamilien IV, 3b, preprint. W. Engelmann, Leipzig, 384 pp.
Kelloff, C.L. & Funk, V.A. (2004) Phytogeography of the Kaieteur Falls, Potaro Plateau, Guyana: Floral distributions and affinities. Journal of Biogeography 31 (3): 501–513. https://doi.org/10.1046/j.0305-0270.2003.01038.x
Kok, P.J.R. (2010) A new species of Chironius Fitzinger, 1826 (Squamata: Colubridae) from the Pantepui region, northeastern South America. Zootaxa 2611: 31–44. https://doi.org/10.11646/zootaxa.2611.1.3
Kok, P.J.R., Ratz, S., Tegelaar, M., Aubret, F. & Means, D.B. (2015) Out of taxonomic limbo: A name for the species of Tepuihyla (Anura: Hylidae) from the Chimantá Massif, Pantepui region, northern South America. Salamandra 51: 283–314.
Kok, P.J.R., Nicolai, M.P.J., Lathrop, A. & MacCulloch, R.D. (2018) Anomaloglossus meansi sp. n., a new Pantepui species of the Anomaloglossus beebei group (Anura, Aromobatidae). ZooKeys 759: 99–116. https://doi.org/10.3897/zookeys.759.24742
Kuntze, C.E.O. (1898) Lentibulariaceae. Revisio Generum Plantarum 3 (3): 240.
Kurz, W.S. (1874) Description of Utricularia nivea Vahl. Journal of Botany, British and Foreign 12: 53–54.
Larrañaga, D.A. (1923) Escritos de Don Damaso Antonio Larrañaga 2. Instituto histórico y geográfico del Urugay, Imprenta Nacional, Montevideo, 511 pp. https://doi.org/10.5962/bhl.title.116606
Lehmann, J.G.C. (1844) Utricularia. Botanische Zeitung (Berlin) 2: 109–110.
Linnaeus, C. (1753) Species Plantarum 1. Impensis Laurentii Salvii, Holmiae, 572 pp.
Menezes, C.G., Gasparine, E.C., Baleeiro, P.C. & de Miranda, V.F.O. (2014) Seed morphology of bladderworts: A survey on Utricularia sect. Foliosa and sect. Psyllosperma (Lentibulariaceae) with taxonomic implications. Phytotaxa 167 (2): 173–182. https://doi.org/10.11646/phytotaxa.167.2.3
Miquel, F.A.W. (1847) Lentibulariaceae. Linnaea 19: 144–145.
Pilger, R.K.F. (1914) Lentibulariaceae. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 6: 188–191. https://doi.org/10.2307/3994403
Pivello, V.R., Vieira, I., Christianini, A.V., Ribeiro, D.B., da Silva Menezes, L., Berlinck, C.N., Melo, F.P.L., Marengo, J.A., Tornquist, C.G., Tomas, W.M. & Overbeck, G.E. (2021) Understanding Brazil’s catastrophic fires: Causes, consequences and policy needed to prevent future tragedies. Perspectives in Ecology and Conservation 19 (3): 233–255. https://doi.org/10.1016/j.pecon.2021.06.005
Rando, J.G. & Pirani, R. (2011) Padrões de distribuição geográfica das espécies de Chamaecrista sect. Chamaecrista ser. Coriaceae (Benth.) H. S. Irwin & Barneby, Leguminosae – Caesalpinioideae. Brazilian Journal of Botany 34: 499–513. https://doi.org/10.1590/S0100-84042011000400004
Rapini, A., Ribeiro, P.L., Lambert, S. & Pirani, J.R. (2008) A flora dos campos rupestres da Cadeia do Espinhaço. Megadiversidade 4 (1–2): 197–217.
Rendón, J.H. & Ruiz, S.Z. (2017) Lentibulariaceae. Flora de Guerrero 77. Universidad Nacional Autónoma de México, Facultad de Ciencias. Circuito, p. 47.
Rickett, H.W. (1967) The bladderworts (Utricularia). Wild flowers of the United States, the Southeastern states 2. New York Botanical Garden , McGraw-Hill. New York, pp. 518–520.
Saint-Hilare, A. de (1833) Voyage dans le district des diamans et sur le littoral du Breìsil. Librairie-Gide, Paris, 454 pp.
Saint-Hilarie, A. de & Girard, F. (1838) Monographie des Primulaceìes et des Lentibularieìes du Breìsil meìridional. Comptes Rendus Hebdomadaires des Seìances de l’Academie des Sciences, Series. D: Sciences Naturelles 7: 868–870.
Salerno, P.E., Ron, S.R., Señaris, J.C., Rojas-Runjaic, F.J.M., Noonam, B.P. & Cannatella, D.C. (2012) Ancient tepui summits harbor young rather than old lineages of endemic frogs. Evolution 66 (10): 3000–3013. https://doi.org/10.1111/j.1558-5646.2012.01666.x
Silva, N.G. da, Alves, R.J.V., Pereira, J.F. & Rivadavia, F. (2011) Lentibulariaceae, Serra de São José, Minas Gerais, Brazil. Check List 7 (2): 120–127. https://doi.org/10.15560/7.2.120
Souza, P.C.B. & Bove, C.P. (2012) Lentibulariaceae. In: Flora dos estados de Goiás e Tocantins. Coleção Rizzo, PRPPG/UFG, Goiânia, Brasil, 69 pp.
Standley, P.C. (1936) Lentibulariaceae. Publications of the Field Museum of Natural History, Botanical Series 12: 366–367.
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