Abstract
The aquatic and wetland plant genus Bacopa contains species that are found worldwide, with a center of diversity in the Americas. Although there have been regional taxonomic studies of the genus, a global synthesis is lacking. Variation in morphological features across species has complicated efforts to determine species boundaries and circumscribe infrageneric categories with confidence. Molecular data have enabled a more thorough understanding of phylogenetic relationships in Bacopa, and these can be used to develop a more stable taxonomy for the genus. Because there are approximately 60 species in the genus, it would be helpful to have a phylogenetically informed sectional classification and an improved understanding of ancestral biogeography and morphological evolution. We obtained novel molecular data for 28 species and produced phylogenetic hypotheses that include most species in the genus. The phylogenetic evidence supports at least four independent transitions to becoming fully aquatic species. We also reconstructed three separate ancestral dispersal events into Africa and beyond. Morphometric data were used to evaluate species boundaries and synonymy. We update sectional circumscriptions and propose a new section, Bacopa sect. Conobea, to accommodate species formerly in the genus Conobea that were found to be monophyletic with Bacopa.
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References
Ahedor AR (2019) Bacopa. In: Flora of North America editorial committee (eds) Flora of North America North of Mexico, vol. 17. Oxford University Press Inc., New York
Albach DC, Meudt HM, Oxelman B (2005) Piecing together the new Plantaginaceae. Amer J Bot 92:297–315. https://doi.org/10.3732/ajb.92.2.297
Arias Divito JC (1968) Las expediciones científicas españolas durante el signl XVIII. Instituto de Cultura Hispánica, Madrid
Atlas of Living Australia (2023) Bacopa: occurrence download. Available at: http://www.ala.org.au. Accessed 29 May 2023
Aymard CGA, Rangel-Ch JO, Minorta-Cely V, Castro-Lima F (2020) Notes on the genus Bacopa (Plantaginaceae, Gratioleae) in the Orinoquia region of Colombia and Venezuela. Harvard Pap Bot 25:195–203. https://doi.org/10.3100/hpib.v25iss2.2020.n7
Baldwin BG (1992) Phylogenetic utility of the internal transcribed spacers of nuclear ribosomal DNA in plants: an example from the Compositae. Mol Phyl Evol 1:3–16. https://doi.org/10.1016/1055-7903(92)90030-K
Barker WR (1992) Bacopa. In: Harden GJ (ed) Flora of New South Wales, vol. 3. New South Wales University Press, Sydney, pp 556–557
Barker WR, Croft R (2015) Illustrating regional differences and trends in the chronology of records of taxa. Austral Syst Bot Soc Newslett 162:16–22
Barrett SC, Strother JL (1978) Taxonomy and natural history of Bacopa (Scrophulariaceae) in California. Syst Bot 3:408–419. https://doi.org/10.2307/2418753
Barringer KA, Burger WC (2000) Flora Costaricensis: Scrophulariaceae. Fieldiana Bot 41:1–69. https://doi.org/10.5962/bhl.title.2594
Barroso GM (1952) Scrophulariaceae: indígenas e exóticas no Brasil. Rodriguésia 15:9–108
Bentham G (1836) Pentstemon staticifolius. In: Lindley J, Edwards’s botanical register, vol. 21. James Ridgway and Sons, Picadilly, t 1770
Borhidi A, Muñiz O (1975) New plants in Cuba IV. Act Bot Acad Sci Hung 21:221–230
Bouckaert R, Vaughan TG, Barido-Sottani J, Duchêne S, Fourment M, Gavryushkina A et al (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Comput Biol 15:e1006650. https://doi.org/10.1371/journal.pcbi.1006650
Brown R (1810) Prodromus florae Novae Hollandiae, vol. 1. Richard Taylor, London. https://doi.org/10.5962/bhl.title.52309
Browne P (1756) The civil and natural history of Jamaica. Osborne and Shipton, London
Carney JA (2001) Black rice: the African origins of rice cultivation in the Americas. Harvard University Press, Cambridge
Chamisso A (1833) Scrofularineae. Linnaea 8:17–35
Chen LMJ, Lua HK, Yeo RSW, Choo LM, Ho BC, Chua KS, Koh SL (2018) Additions to the flora of Singapore—new and overlooked records of naturalised plant species. Nature Singapore 11:63–75
Cheng T, Xu C, Lei L, Li C, Zhang Y, Zhou S (2016) Barcoding the kingdom Plantae: new PCR primers for ITS regions of plants with improved universality and specificity. Molec Ecol Res 16:138–149. https://doi.org/10.1111/1755-0998.12438
Chong KY, Tan HT, Corlett RT (2009) A checklist of the total vascular plant flora of Singapore: native, naturalised and cultivated species. National University of Singapore, Singapore
Christenhusz MJM (2014) Plantaginaceae. In: Davidse G, Sousa Sánchez M, Knapp S, Chiang Cabrera F (eds) Flora Mesoamericana, vol. 5. Missouri Botanical Garden Press, St, Louis, pp 1–73
Cook CDK (1999) The number and kinds of embryo-bearing plants which have become aquatic: a survey. Perspect Pl Ecol Evol Syst 2:79–102. https://doi.org/10.1078/1433-8319-00066
D’Arcy WG (1979) Scrophulariaceae In: Woodson RE, Schery RW. Flora of Panama. Ann Missouri Bot Gard 66:173–272. https://doi.org/10.2307/2398908
DePoe CE (1969) Bacopa egensis (Poeppig) Pennell (Scrophulariaceae) in the United States. SIDA Contrib Botany 3:313–318
Díaz JF, Beiras MB (2007) Plantas invasoras de Galicia: bioloxía, distribución e métodos de control). Xunta De Galicia, Santiago de Compostela
Doyle JJ, Doyle JL (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem Bull 19:11–15
Estes D, Small RL (2008) Phylogenetic relationships of the monotypic genus Amphianthus (Plantaginaceae tribe Gratioleae) inferred from chloroplast DNA sequences. Syst Bot 33:176–182. https://doi.org/10.1600/036364408783887375
Fensham RJ, Laffineur B (2019) Defining the native and naturalized flora for the Australian continent. Austral J Bot 67:55–69. https://doi.org/10.1071/BT18168
Fritsch PW, Almeda F, Martins AB, Cruz BC, Estes D (2007) Rediscovery and phylogenetic placement of Philcoxia minensis (Plantaginaceae), with a test of carnivory. Proc Calif Acad Sci 58:447–467
GBIF (2022) Global Biodiversity Information Facility, occurrence download. Available at: https://doi.org/10.15468/dl.8x8hch. Accessed 5 Dec 2022
Ghazanfar SA, Hepper FN, Philcox D (2008) Flora of tropical east Africa: Scrophulariaceae. Royal Botanic Gardens, Kew
Gonzalez-Socoloske D, Tippery NP, Jiménez Pérez NC, Les DH (2020) New record of Bacopa egensis (Plantaginaceae) for the flora of Mexico. J Bot Res Inst Texas 14:395–403
Grayum MH, Wilson KA (1975) Bacopa monnieri (Scrophulariaceae), a new record for California. Madroño 23:161
Hammel BE, Grayum MH, Herrera C, Zamora N (2015) Manual de plantas de Costa Rica, vol. 8. Missouri Botanical Garden Press, Saint Louis
Heberling JM (2022) Herbaria as big data sources of plant traits. Int J Pl Sci 183:87–118. https://doi.org/10.1086/717623
Hepper FN (1963) Scrophulariaceae. In: Hutchinson J, Dalziel JM, Hepper FN (eds) Flora of west tropical Africa, ed. 2. Crown Agents for Overseas Governments, London, pp 352–374
Hong DY, Yang HB, Jin CL, Holmgren NH (1998) Scrophulariaceae. In: Wu ZY, Raven PH (eds) Flora of China, vol. 18. Missouri Botanical Garden Press, St. Louis, pp 1–212
Kadono Y (2004) Alien aquatic plants naturalized in Japan: history and present status. Global Environm Res 8:163–169
Kuzmina ML, Braukmann TW, Fazekas AJ, Graham SW, Dewaard SL, Rodrigues A, Bennett BA, Dickinson TA, Saarela JM, Catling PM, Newmaster SG (2017) Using herbarium-derived DNAs to assemble a large-scale DNA barcode library for the vascular plants of Canada. Appl Pl Sci 5:1700079. https://doi.org/10.3732/apps.1700079
Lansdown RV, Knees SG, Patzelt A (2013) Bacopa monnieri. The IUCN Red List of threatened species 2013:e.T164168A17722668. https://doi.org/10.2305/IUCN.UK.2013-1.RLTS.T164168A17722668.en
Linnaeus C (1759) Systema naturae, vol. 2. Lars Salvius, Stockholm
Maddison WP, Maddison DR (2023) Mesquite: a modular system for evolutionary analysis, ver. 3.81. Available at: http://www.mesquiteproject.org/. Accessed 2 Jun 2023
Mielcarek R (1994) Un nouveau Bacopa (Scrophulariaceae) de Guinée (Afrique de l’Ouest). Fragm Florist Geobot 39:25–27
Nguyen LT, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molec Biol Evol 32:268–274. https://doi.org/10.1093/molbev/msu300
Olmstead RG, Sweere JA (1994) Combining data in phylogenetic systematics: an empirical approach using three molecular data sets in the Solanaceae. Syst Biol 43:467–481. https://doi.org/10.1093/sysbio/43.4.467
Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisens WJ, Reeves PA (2001) Disintegration of the Scrophulariaceae. Amer J Bot 88:348–361. https://doi.org/10.2307/2657024
Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND, Powell GVN, Underwood EC, D’Amico JA, Itoua I, Strand HE, Morrison JC et al (2001) Terrestrial ecoregions of the world: a new map of life on earth: e new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. Bioscience 51:933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Pennell FW (1935) The Scrophulariaceae of eastern temperate North America. Acad Nat Sci Philadelphia Monogr 1:1–650
Pennell FW (1946) Reconsideration of the Bacopa-Herpestis problem of the Scrophulariaceae. Proc Acad Phil 98:83–98
Philcox D (1972) Bacopa Aublet. In: Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, Webb DA (eds) Flora Europaea, vol. 3. Diapensiaceae to Myoporaceae. Cambridge University Press, Cambridge
Philcox D (1990) Scrophulariaceae. In: Launert E, Pope GV (eds) Flora Zambesiaca, vol. 8., part 2. Flora Zambesiaca Managing Committee, London, pp 1–179
POWO (2022) Plants of the world online. Royal botanic gardens, Kew. Available at: http://www.plantsoftheworldonline.org/. Accessed 14 Jul 2023
R Core Team (2023). R: A language and environment for statistical computing, version 4.3.0. R Foundation for Statistical Computing, Vienna. Available at: https://www.R-project.org/.. Accessed 4 May 2023
Robinson BL (1909) Diagnoses and transfers of tropical American phanerogams. Contrib Gray Herb Harvard Univ 613–626
Ratnasingham S, Hebert PD (2007) BOLD: the barcode of life data system. Molec Ecol Notes 3:355–364
Raus T (2003) Bacopa (Scrophulariaceae) in Greece and adjacent countries. Bocconea 16:745–749
Romanowski N (2011) Wetland weeds: causes, cures and compromises. CSIRO Publishing, Collingwood
Scatigna AV, Fritsch PW, Souza VC, Simões AO (2018) Phylogenetic relationships and morphological evolution in the carnivorous genus Philcoxia (Plantaginaceae, Gratioleae). Syst Bot 43:910–919. https://doi.org/10.1600/036364418X697814
Scatigna AV, Souza VC, Sosa MM, Colletta DG, Machado RM, Simões AO (2022) Phylogenetics of Gratioleae (Plantaginaceae): paraphyly of Stemodia and its implications for generic circumscriptions, with insights from floral evolution. Bot J Linn Soc 200:194–217. https://doi.org/10.1093/botlinnean/boac013
Scatigna AV (2023) Looks can be deceiving: recently published Bacopa llanorum (Plantaginaceae) is synonymous with Lindernia brachyphylla (Linderniaceae) and B. debilis is an unnecessary combination. Phytotaxa 613:64–68. https://doi.org/10.11646/phytotaxa.613.1.6
Schrank F (1828) Plantae novae aut minus cognitae. Syll Pl Ratisb 2:55–82
Schwarz G (1978) Estimating the dimension of a model. Ann Stat 6:461–464
Sloane H (1696) Catalogus plantarum quae in insula Jamaica sponte proveniunt. D. Brown, London
Sloane H (1707) A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica, vol. 1. Bentley and Magnes, London
Sosa MM, Florentín JE, Scatigna AV (2021) Sinopsis de las especies del género Bacopa (Plantaginaceae, Gratioleae) de Paraguay y sus estados de conservación. Bol Soc Argent Bot 56:71–80
Sosa MM, Moroni P, O’Leary N (2018) A taxonomic revision of the genus Bacopa (Gratioleae, Plantaginaceae) in Argentina. Phytotaxa 336:1–27. https://doi.org/10.11646/phytotaxa.336.1.1
Sosa MM (2009) Estudios morfo-anatómicos, cromosómicos y palinológicos en especies de Stemodia (Scrophulariaceae). PhD Thesis, Instituto de Botánica del Nordeste, Universidad Nacional del Nordeste, Corrientes
Souza VC, Giulietti AM (2009) Levantamento das espécies de Scrophulariaceae sensu lato nativas do Brasil. Pesquisas Bot 60:7–288
Souza VC (2003) Scrophulariaceae. In: Wanderley MGL, Shepherd GJ, Melhem TS, Giulietti AM, Kirizawa M (eds) Flora Fanerogâmica do Estado de São Paulo Online, vol. 3. Fundação de Amparo à Pesquisa do Estado de São Paulo, São Paulo, pp 297–321
Standley PC, Williams LO (1973) Scrophulariaceae. Flora of Guatemala, part 9. Fieldiana Bot 24:319–416
Sutton D, Hampshire R (2001) Scrophulariaceae. In: Stevens WD, Ulloa Ulloa C, Pool A, Montiel OM (eds) Flora de Nicaragua. Monogr Syst Bot Missouri Bot Gard 85: 2354–2368
Taberlet P, Gielly L, Pautou G, Bouvet J (1991) Universal primers for amplification of three non-coding regions of chloroplast DNA. Pl Molec Biol 17:1105–1109. https://doi.org/10.1007/BF00037152
Tamjidi Eramsadati SS, Mohammadpour P, Ramzi S (2021) New distribution of Bacopa monnieri from North of Iran. Rostaniha 22:150–154. https://doi.org/10.22092/botany.2021.354988.1252
Tippery NP, Pesch JD, Murphy BJ, Bautzmann RL (2020) Genetic diversity of native and introduced Phragmites (common reed) in Wisconsin. Genetica 148:165–172. https://doi.org/10.1007/s10709-020-00098-z
Tippery NP, Pawinski KC, Jeninga AJ (2021) Taxonomic evaluation of Nymphoides (Menyanthaceae) in eastern Asia. Blumea 66:249–262. https://doi.org/10.3767/blumea.2021.66.03.08
Tippery NP, Gonzalez-Socoloske D, Leliaert F, Scatigna AV, Souza VC (2023) Data for Systematics and biogeography of Bacopa (Plantaginaceae). Available at. https://doi.org/10.6084/m9.figshare.23739456
Tungphatthong C, Somnuek J, Phadungcharoen T, Ingkaninan K, Denduangboripant KJ, Sukrong S (2018) DNA barcoding of species of Bacopa coupled with high-resolution melting analysis. Genome 61:867–877. https://doi.org/10.1139/gen-2018-0059
Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber WH, Li DZ, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ, Smith GF (eds) (2018) International code of nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, Jul 2017. [Regnum Vegetabile 159]. Koeltz Botanical Books, Glashütten. https://doi.org/10.12705/Code.2018
Villaseñor JL (2016) Checklist of the native vascular plants of Mexico. Revista Mex Biodivers 87:559–902. https://doi.org/10.1016/j.rmb.2016.06.017
Weakley AS (2023) Flora of the southeastern United States. University of North Carolina Herbarium, North Carolina Botanical Garden, Chapel Hill
Wettstein R (1891) Scrophulariaceae. In: Engler A, Prantl K (eds) Die natürlichen Pflanzenfamilien, Teil 4, Abt. 3b, pp 39–107 https://doi.org/10.5962/bhl.title.4635
Wickham H (2016) ggplot2: elegant graphics for data analysis. Springer-Verlag, New York
Yamazaki T (1990) Scrophulariaceae. Flora of Thailand, vol. 5, part 2. Forest Herbarium, Royal Forest Department, Bangkok
Yu Y, Harris AJ, Blair C, He XJ (2015) RASP (reconstruct ancestral state in phylogenies): a tool for historical biogeography. Molec Phylogen Evol 87:46–49. https://doi.org/10.1016/j.ympev.2015.03.008
Yu G, Smith D, Zhu H, Guan Y, Lam TTY (2017) ggtree: an R package for visualization and annotation of phylogenetic trees with their covariates and other associated data. Methods Ecol Evol 8:28–36. https://doi.org/10.1111/2041-210X.12628
Yu Y, Blair C, He XJ (2020) RASP 4: ancestral state reconstruction tool for multiple genes and characters. Molec Biol Evol 37:604–606. https://doi.org/10.1093/molbev/msz257
Acknowledgements
We are grateful for the curators and staff of herbaria who provided access to material for examination and DNA sampling: Ann Bogaerts (BR), Rick Ree (F), and Matthew Pace (NY). Russell Barrett and Bill Barker provided helpful insights about B. monnieri in Australia. Research was funded in part by the University of Wisconsin—Whitewater Undergraduate Research Program. AVS gratefully acknowledges research fellowship support from Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) (PDCTR-00123/20) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (301691/2021-5).
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Appendix 1
Appendix 1
Voucher information and GenBank accession numbers for newly obtained DNA sequences (nuclear ribosomal ITS / ndhF / trnL-trnF region). Sequences not obtained for a given specimen are indicated with a dash (–).
Bacopa angulata—(Baco18) Brazil: Ceará: Fortaleza, Drouet 2358 (F0878008) (OR484593 / OR474274 /–); B. aquatica—(Baco82) Venezuela: Bolivar: Cerro Baraguan, Wurdack and Morachino 41218 (NY00499249) (OR484594 / OR474255 / OR474304); B. aubletiana (Baco100)—French Guiana: Commune de Roura, Gonzalez 1895 (NY01340905) (OR484595 / OR474297 / OR474331); (Baco98) Brazil: Minas Gerais: Serra do Cipó, Irwin et al. 20292a (NY00911572) (OR484596 / OR474298 /–); (Baco99) Venezuela: Bolívar, Maguire et al. 36024 (NY01340904) (OR484597 / OR474299 /–); B. bacopoides—(Baco36) Belize: Belize Distr., Dwyer 11464 (F1808606) (OR484598 / OR474279 /–); B. caroliniana—(Baco24) USA: South Carolina: Barnwell Co.: Williston, Nelson and Kennemore 14913 (F2163741) (OR484599 / OR474281 / OR474316); B. congesta (Baco25) Argentina: Apóstoles: San José, Cabrera and Sáenz, 29149 (F1866565) (–/ OR474256 /–); B. crenata–(Baco09) Liberia: Grand Cape Mount, Jongkind 11707 (BR0000019040271) (OR484600 / OR474275 / OR474311); (Baco66) Senegal: Basse-Casamance: Koubalan, Vanden Berghen 9328 (BR0000019038070) (OR484603 / OR474276 / OR474312); (Baco12) Liberia: Bong, north-east of Gbarnga, Jongkind 10,978 (BR0000019038087) (OR484601 /–/–); (Baco62) Democratic Republic of the Congo: Kinshasa: Mbudi, Nsimundele 2284 (BR0000020362683) (OR484602 /–/–); B. dubia—(Baco32) Argentina: Corrientes: Santo Tomé, Krapovickas et al. 17067 (F1708053) (OR484604 / OR474290 / OR474325); B. egensis–(Baco01) México: Tabasco: Balancán: Laguna Leona Vicario, Gonzalez-Socoloske 325 (F) (MT676828 / MT677654 / OR474326); (Baco68) Democratic Republic of the Congo: Kwilu: Kikwit, Masens 477 (BR0000019038766) (OR484606 / OR474292 / OR474327); (Baco71) Democratic Republic of the Congo: Odzala, Lejoly 96/995 (BR0000019039299) (OR484607 / OR474293 / OR474328); (Baco04) Gabon: Moyen-Ogooue: Lac Onangué, Lachenaud 2028 (BR0000016176270) (OR484605 / OR474291 /–); B. erecta—(Baco69) Senegal: Basse-Casamance: Abéné, Vanden Berghen 9512 (BR0000019039176) (OR484608 / OR474278 / OR474314); (Baco14) Senegal: Basse-Casamance: Abéné, Vanden Berghen 9501 (BR0000019039169) (–/–/ OR474313); (Baco13) Senegal: Basse-Casamance: environs de Ziguinchor, Vanden Berghen s.n. (BR0000019039022) (–/ OR474277 /–); B. gracilis—(Baco34) Brazil: Roraima: Serra Tepequem, Prance et al. 4301 (F1814598) (–/ OR474257 /–); B. gratioloides—(Baco86) Guyana: Rupununi: Manari, Jansen-Jacobs et al. 4623 (NY01340911) (OR484617 / OR474260 / OR474305); (Baco83) Brazil: Bahia: Camaçari: Arembepe, Cardoso et al. 1004 (NY01019220) (OR484615 / OR474258 /–); (Baco84) Brazil: Bahia: Caetité, Souza et al. 5401 (NY01016160) (OR484616 / OR474259 /–); B. hamiltoniana—(Baco08) Ghana: Brong-Ahafo, Jongkind 2412 (BR0000005151837) (OR484618 / OR474272 /–); (Baco73) Benin: Atakora: Natitingou, Kotopounga, Houngnon 7562 (BR0000016136335) (OR484619 / OR474273 /–); B. innominata—(Baco27) USA: Florida: Duval Co.: Jacksonville, Curtiss 5743 (F0097241) (OR484620 / OR474294 / OR474329); (Baco55) USA: Virginia: King William Co.: Mattaponi River, Fernald and Long 12799 (F1489437) (OR484621 / OR474295 / OR474330); B. micromonnieria—(Baco38) Cuba: Camagüey: Minas, Dahlgren and Macbride 136 (F1414142) (OR484622 / OR474261 /–); B. monnierioides—(Baco44) Honduras: El Paraíso: Suyate, Standley 28053 (F1514743) (OR484623 / OR474282 / OR474317); (Baco46) Brazil: Goiás: Chapada dos Veadeiros, Irwin et al. 24528 (F1798588) (OR484624 / OR474283 / OR474318); B. myriophylloides—(Baco41) Colombia: Boyacá: Los Llanos, Haught 2705 (F1067137) (OR484625 / OR474284 / OR474319); B. occultans—(Baco79) Niger: Niamey: Saga, Leman 122 (BR0000019040332) (OR484626 / OR474285 / OR474320); B. pubescens—(Baco02) Benin: Collines: Abira (Dassa), Ayichedehou 1921 (BR0000005548644) (OR484609 / OR474264 / OR474307); (Baco03) Benin: Borgou: Pèrèrè, Binassi, Adjakidjé 2283 (BR0000019039862) (OR484610 / OR474265 / OR474308); (Baco06) Benin: Borgou: Tchaourou, Goro, Sokpon 602 (BR0000019039855) (OR484611 / OR474267 / OR474309); (Baco15) Benin: Borgou: Bétérou, Ayichedehou 1419 (BR0000005548842) (OR484613 / OR474269 / OR474310); (Baco74) Benin: Donga: Aoro, sud de Bassila, Lejoly 98/290B (BR0000005548941) (OR484614 / OR474270 /–); (Baco05) Benin: Borgou: Wari Maro (Bétérou), Ayichedehou 1353 (BR0000005548743) (–/ OR474266 /–); (Baco11) Benin: Atakora: Matéri, Porga, Houngnon 7400 (BR0000019039831) (OR484612 / OR474268 /–); (Baco75) Benin: Borgou: Parakou, Houngnon 6267 (BR0000019040196) (–/ OR474271 /–); B. reflexa—(Baco 47) Venezuela: Guárico: Calabozo, Aristeguieta 4822 (F1568610) (OR484627 / OR474286 / OR474321); B. reptans—(Baco90) Brazil: Mato Grosso: Barra do Garças, Hunt and Ramos 6088 (NY00911440) (OR484628 / OR474289 / OR474324); B. salzmannii—(Baco51) México: Chiapas: Pueblo Nuevo Solistachuacán, Shilom Ton 2531 (F1749314) (–/ OR474296 /–); B. scabra—(Baco21) Guatemala: Jutiapa: Jutiapa, Standley 76003 (F1148600) (OR484629 /–/–); (Baco52) Paraguay: Guairá: Colonia Independencia, Pedersen 10104 (F2286507) (OR484630 / OR474262 / OR474306); B. scoparioides—(Baco101) Brazil: Bahia: Una, Amorim et al. 2382 (NY00823192) (OR484631 / OR474300 / OR474332); (Baco103) Brazil: Amazonas: Manaus: Perto da Cidade Nova, Nelson 1353 (NY01173160) (OR484632 / OR474301 / OR474333); B. serpyllifolia—(Baco94) Guyana: Rupununi: Dadanawa, Jansen-Jacobs et al. 3207 (NY01340908) (OR484633 / OR474287 / OR474322); (Baco96) Guyana: Rupununi: Shea Rock, Jansen-Jacobs et al. 4990 (NY01340907) (OR484634 / OR474288 / OR474323); B. sessiliflora—(Baco54) Costa Rica: Puntarenas: Garabito, Hammel 19996 (F2212974) (OR484635 / OR474280 / OR474315); B. stricta—(Baco19) Brazil: Bahia: Piritiba, Noblick 1846 (F1925792) (OR484636 /–/–); (Baco31) Dominican Republic: San José de Ocoa, Liogier 18488 (F1775988) (OR484637 / OR474263 /–); (Baco56) Puerto Rico, Heller and Heller 716 (F0119827) (OR484638 /–/–); Mecardonia procumbens (Mill.) Small—(Baco61) Bahamas; Grand Bahama, Correll 44789 (F1776836) (OR484639 / OR474303 /–); (Baco45) Guatemala: Petén: Sayaxché, Tún Ortíz 1141 (F1683234) (–/ OR474302 /–).
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Tippery, N.P., Gonzalez-Socoloske, D., Leliaert, F. et al. Systematics and biogeography of Bacopa (Plantaginaceae). Plant Syst Evol 310, 7 (2024). https://doi.org/10.1007/s00606-023-01884-w
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DOI: https://doi.org/10.1007/s00606-023-01884-w