Updating the taxonomy of polymorphic plant taxa: six Atlantic Forest species segregated from the widely circumscribed Oxalis polymorpha Mart. ex Zucc. (Oxalidales, Oxalidaceae)
Abstract
Widely circumscribed taxa are good candidates for in-depth taxonomic studies. The current taxonomy of Oxalis sect. Polymorphae includes 13 species mostly endemic to the Brazilian Atlantic Forest, in eastern Brazil. Oxalis polymorpha Mart. ex Zucc., the type-species of the section, exhibits a wide morphological variability and includes at least four recognized morphotypes. We propose the segregation of six species from this widely circumscribed O. polymorpha by combining morphological and molecular systematic data from the nuclear and plastid genomes. Thus, Oxalis decipiens Progel, previously treated under O. polymorpha, is accepted as a distinct species. The same is valid for O. polymorpha subsp. tijucana Lourteig, which is recognised at species level (O. tijucana (Lourteig) Fiaschi). Four additional new species are described based on material that were formerly placed under the wide circumscription of O. polymorpha: O. amorimii Fiaschi sp. nov., O. animarum Fiaschi sp. nov., O. idimae Fiaschi sp. nov., and O. itatingae Fiaschi sp. nov. We provide evidence that one of these species, O. itatingae, is phylogenetically closer to O. sect. Holophyllum than to other species of O. sect. Polymorphae, while the other species, excluding O. tijucana, emerged as non-sister taxa within the latter. Descriptions, illustrations, geographic distribution maps, preliminary conservation assessments, taxonomic notes, and an identification key to species of Oxalis sect. Polymorphae are provided.
References
Abreu M.C. 2011. Sistemática de Oxalis L. sect. Thamnoxys (Endl.) Progel no Brasil. PhD thesis. Pernambuco Federal Rural University, Brazil.
Alencar J., Maciel J.R. & Buril M.T. 2024. Morphologically hypervariable species hinder our knowledge of biodiversity: Daustinia montana (Convolvulaceae) as a case study. Botanical Journal of the Linnean Society 204 (1): 86–101. https://doi.org/10.1093/botlinnean/boad040
Anderson B.M., Binks R.M., Byrne M., Crawford A.D. & Shepherd K.A. 2023. Using RADseq to resolve species boundaries in a morphologically complex group of yellow-flowered shrubs (Geleznowia, Rutaceae). Australian Systematic Botany 36 (4): 277–311. doi:10.1071/SB23010
Bacci L.F., Amorim A.M., Michelangeli F.A. & Goldenberg R. 2018. Increased sampling in under-collected areas sheds new light on the diversity and distribution of Bertolonia, an Atlantic Forest endemic genus. Systematic Botany 43 (3): 767–792. https://doi.org/10.1600/036364418X697490
Bachman S., Moat J., Hill A.W., de la Torre J. & Scott B. 2011. Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117–126. https://doi.org/10.3897/zookeys.150.2109
Beenje H. 2016. The Kew Plant Glossary. Royal Botanic Gardens, Kew.
Bouckaert R., Vaughan T.G., Barido-Sottani J., Duchêne S., Fourment M., Gavryushkina A., Heled J., Jones G., Kühnert D., Maio N., Matschiner M., Mendes F.K., Müller N.F., Ogilvie H.A., du Plessis L., Popinga A., Rambaut A., Rasmussen D., Siveroni I., Suchard M.A., Wu C.-H., Xie D., Zhang C., Stadler T. & Drummond A.J. 2019. BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology 15 (4): e1006650. https://doi.org/10.1371/journal.pcbi.1006650
Cabral F.S., Lima D.F., Vaio M. & Fiaschi P. 2024. Molecular phylogenetics of Oxalis subg. Thamnoxys (Oxalidaceae) reveals artificial arrangements of traditional sections. Botanical Journal of the Linnean Society 204 (2): 174–186. https://doi.org/10.1093/botlinnean/boad044
Chautems A., Lopes T.C.C., Peixoto M. & Rossini J. 2010. Taxonomic revision of Sinningia Nees (Gesneriaceae) IV: six new species from Brazil and a long overlooked taxon. Candollea 65: 241–266. https://doi.org/10.15553/c2010v652a6
Costa T.S., Sakuragui C.M. & Fiaschi P. 2019. Flora do Rio de Janeiro: Oxalidaceae. Rodriguésia 70: e02382017. https://doi.org/10.1590/2175-7860201970043
Darriba D., Posada D., Kozlov A.M., Stamatakis A., Morel B. & Flouri T. 2020. ModelTest-NG: a new and scalable tool for the selection of DNA and protein evolutionary models. Molecular Biology and Evolution 37 (1): 291–294. https://doi.org/10.1093/molbev/msz189
Dias I.R., Medeiros T.T., Nova M.F.V. & Solé M. 2014. Amphibians of Serra Bonita, southern Bahia: a new hotpoint within Brazilʼs Atlantic Forest hotspot. ZooKeys 449: 105–130. https://doi.org/10.3897/zookeys.449.7494
Doyle J.J. & Doyle J.L. 1990. Isolation of plant DNA from fresh tissue. Focus 12: 13–15.
Drummond A.J., Suchard M.A., Xie D. & Rambaut A. 2012. Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution 29 (8): 1969–1973. https://doi.org/10.1093/molbev/mss075
Eiten G. 1963. Taxonomy and regional variation of Oxalis section Corniculatae L. Introduction, keys and synopsis of the species. American Midland Naturalist 69 (2): 257–309. https://doi.org/10.2307/2422912
Ellis B., Daly D.C., Hickey L.J., Johnson K.R., Mitchell J.D., Wilf P. & Wing S.L. 2009. Manual of Leaf Architecture. Cornell University Press, Ithaca. https://doi.org/10.1079/9781845935849.0000
Ferreira G.E., Costa I.G., Araujo A.O., Hopkins M.G. & Chautems A. 2016. Three new species of Besleria (Gesneriaceae) from the southeastern Brazilian Atlantic rainforest. Phytotaxa 263 (3): 270–278. https://doi.org/10.11646/phytotaxa.263.3.7
Fiaschi P. 2012. Two new species of Oxalis sect. Polymorphae (Oxalidaceae) from the Brazilian Atlantic Forest. Kew Bulletin 67: 33–38. https://doi.org/10.1007/s12225-012-9330-3
Fiaschi P. 2014. Three new species and a revised key to species of Oxalis section Polymorphae (Oxalidaceae). Brittonia 66: 134–150. https://doi.org/10.1007/s12228-013-9315-z
Fiaschi P. & Pirani J.R. 2005. Four new species of Schefflera (Araliaceae) from Espírito Santo state, Brazil. Kew Bulletin 60: 77–85. Available from https://www.jstor.org/stable/4110886 [accessed 21 Mar. 2025].
Fiaschi P., Bilk K.D., Sartor M.F., Costa-Lima J.L. da & Lima D.F. 2024. Systematics of Oxalis subgenus Thamnoxys section Holophyllum (Oxalidales, Oxalidaceae). Systematic Botany 49 (1): 154–191. https://doi.org/10.1600/036364424X17110456120758
Fiaschi P., Cabral, F.S., Caballero, L.G.& Lima D.F. 2024. Updating the taxonomy of polymorphic plant taxa: six Atlantic Forest species segregated from the widely circumscribed Oxalis polymorpha Mart. ex Zucc. (Oxalidaceae). Figshare. Dataset. https://doi.org/10.6084/m9.figshare.27324762.v1
Groppo M., Costa I.G., Bruniera C.P., Ferreira C., Afonso L., Maeoka H.M., Dutra F.V., Gasparino E.C. & Ferreira P.L. 2019. Rock star flowers: Conchocarpus hendrixii (Galipeinae, Rutaceae), a new species from Eastern Brazil with notes on its phylogenetic position in the genus. Phytotaxa 422 (1): 75–92. https://doi.org/10.11646/phytotaxa.422.1.5
Heibl C. 2005. Studies on the Systematics, Evolution, and Biogeography of Oxalis Sections Caesiae, Carnosae, and Giganteae, Endemic to the Atacama Desert of Northern Chile. Master thesis. Ludwig-Maximilians-Universität, München, Germany.
IUCN 2012. The IUCN Red List Categories and Criteria, version 3.1, Second Edition. IUCN Red List Unit, Cambridge (UK). Available from https://www.iucnredlist.org [accessed 15 Jul. 2023].
Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
Lopez A., Panseri A.F. & Urtubey E. 2013. Revision of Oxalis section Palmatifoliae DC. (Oxalidaceae). Phytotaxa 138 (1): 1–14. https://doi.org/10.11646/phytotaxa.138.1.1
Lourteig A. 1994. Oxalis L. subgénero Thamnoxys (Endl.) Reiche emend Lourt. Bradea 7 (1): 1–199.
Lourteig A. 2000. Oxalis L. subgéneros Monoxalis (Small) Lourt., Oxalis y Trifidus Lourt. Bradea 7 (2): 1–629.
Mello-Silva R. & Cabral A. 2022. Taxonomic revision of Barbacenia (Velloziaceae) Atlantic Forest Inselberg group, with two new species. Annals of the Missouri Botanical Garden 107: 32–63. https://doi.org/10.3417/2022685
Miller M.A., Pfeiffer W. & Schwartz T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE): 1–8. Gateway Computing, New Orleans. https://doi.org/10.1109/GCE.2010.5676129
Moraes P.L.R. 2009. The Brazilian Herbarium of Maximilian, Prince of Wied. Neodiversity 4: 16–51.
Mori S.A. 2011. Chapter 4. From the Field. In: Mori S.A., Berkov A., Gracie C.A. & Hecklau E.F. (eds) Tropical Plant Collecting: from the Field to the Internet: 131–190. TECC Editora, Florianópolis.
Mori S.A., Boom B.M., Carvalho A.M. & Santos T.S. 1983. Southern Bahian moist forests. The Botanical Review 49: 155–232. https://doi.org/10.1007/BF02861011
Nery E.K. & Fiaschi P. 2019. Geometric morphometrics dismiss the polymorphic Hydrocotyle quinqueloba (Araliaceae) from the Neotropics. Systematic Botany 44: 451–469. https://doi.org/10.1600/036364419X15561132273558
Nery E.K., Matchin-Viera M.E., Camacho O., Caddah M.K. & Fiaschi P. 2020. Delimiting a constellation: integrative taxonomy of a star-shaped Hydrocotyle species complex (Araliaceae) from the Brazilian Atlantic Forest. Plant Systematics and Evolution 306: 57. https://doi.org/10.1007/s00606-020-01682-8
Nuernberg-Silva A. & Fiaschi P. 2021. Taxonomic revision and morphological delimitation of Oxalis sect. Ripariae (Oxalidaceae). Phytotaxa 529: 125–159. https://doi.org/10.11646/phytotaxa.529.1.11
Oliveira R.S., Urdampilleta J.D. & Dutilh J.H.A. 2017. A new Hippeastrum (Amaryllidaceae) species from Brazil. Phytotaxa 307 (2): 147–152. https://doi.org/10.11646/phytotaxa.307.2.6
Oliveira-Filho A.T. 2017. NeoTropTree, Flora arbórea da Região Neotropical: Um banco de dados envolvendo biogeografia, diversidade e conservação. Universidade Federal de Minas Gerais. Available from http://www.neotroptree.info [accessed 21 Mar. 2025].
Pessoa E., Sader M., Pedrosa-Harand A. & Alves M. 2020. A natural hybrid, an autopolyploid, or a new species? An integrative case study of a distinctive Costus species (Costaceae) from the Atlantic Forest of Brazil. Systematics and Biodiversity 18 (1): 42–56. https://doi.org/10.1080/14772000.2020.1729890
Progel A. 1877. Oxalideae. In: Martius C.F.P. & Eichler A.G. (eds) Flora brasiliensis, Vol. 12, Part II: 474–520, plates 102–116. Frid. Fleischer, Lipsiae.
Projeto MapBiomas 2024. Projeto MapBiomas – Coleção 8 da Série Anual de Mapas de Cobertura e Uso de Solo do Brasil. Available from https://plataforma.brasil.mapbiomas.org [accessed 20 Jun. 2024].
Radford A.E., Dickson W.C., Massey J.R. & Bell C.R. 1974. Vascular Plant Systematics. Harper and Row, New York.
Rambaut A., Drummond A.J., Xie D., Baele G. & Suchard M.A. 2018. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67 (5): 901–904. https://doi.org/10.1093/sysbio/syy032
Richetti E., Costa T.S., Cabral W.S., Fiaschi P., Sakuragui C.M. & Lusa M.G. 2022. Morphoanatomy and phylogenetics reveals a distinct species of Oxalis sect. Polymorphae (Oxalidaceae) from the Brazilian Atlantic Forest. Plant Systematics and Evolution 308: 16. https://doi.org/10.1007/s00606-022-01807-1
Richetti E., Lima D.F., Fiaschi P. & Lusa M.G. 2023. Macro- and micromorphology reveal four entities in the highly variable Oxalis polymorpha Mart. ex Zucc. (Oxalidaceae). Botany 101: 43–60. https://doi.org/10.1139/cjb-2022-0073
Sambuichi R.H.R. & Haridasan M. 2007. Recovery of species richness and conservation of native Atlantic forest trees in the cacao plantations of southern Bahia in Brazil. Biodiversity and Conservation 16: 3681–3701. https://doi.org/10.1007/s10531-006-9017-x
Schroth G., Faria D., Araujo M., Bede L., Van Bael S.A., Cassano C.R., Oliveira L.C. & Delabie J.H.C. 2011. Conservation in tropical landscape mosaics: the case of the cacao landscape of southern Bahia, Brazil. Biodiversity and Conservation 20: 1635–1654. https://doi.org/10.1007/s10531-011-0052-x
Sobral M., Costa I.G., Souza M.C. & Zorzanelli J.P.F. 2017. Five new species and one new combination in Brazilian Myrtaceae. Phytotaxa 307 (4): 233–244. https://doi.org/10.11646/phytotaxa.307.4.1
Stamatakis A. 2014. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
Stoughton T.R., Jolles D.D., OʼQuinn R.L. 2017. The western springbeauties, Claytonia lanceolata (Montiaceae): a review and revised taxonomy for California. Systematic Botany 42 (2): 283–300. https://doi.org/10.1600/036364417X695475
Thiers B. continuously updated. Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Available from https://sweetgum.nybg.org/science/ih/ [accessed 21 Mar. 2025].
Thomas W.W. & Barbosa M.R.V. 2008. Natural vegetation types in the Atlantic Coastal Forest of Northeastern Brazil. In: Thomas W.W. (ed.) The Atlantic Coastal Forest of Northeastern Brazil: 6–20. Memoirs of the New York Botanical Garden, Vol. 100. The New York Botanical Garden Press, Bronx.
Vaio M., Nascimento J., Mendes S., Ibiapino A., Felix L.P., Gardner A., Emshwiller E., Fiaschi P. & Guerra M. 2018. Multiple karyotype changes distinguish two closely related species of Oxalis (O. psoraleoides and O. rhombeo-ovata) and suggest an artificial grouping of section Polymorphae (Oxalidaceae). Botanical Journal of the Linnean Society 188: 269–280. https://doi.org/10.1093/botlinnean/boy054
Valadares R.T. & Coelho M.A.N. 2017. Anthurium idimae, a new species of Araceae from Southeastern Brazil. Phytotaxa 316 (1): 73–78. https://doi.org/10.11646/phytotaxa.316.1.7
Zuccarini J.G. 1825. Monographie der amerikanischen Oxalis-Arten. Königliche Akademie der Wissenschaften, München 9: 125–184. https://doi.org/10.5962/bhl.title.161310
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