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Systematics and biogeography of Bacopa (Plantaginaceae)

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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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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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Authors and Affiliations

  1. Department of Biology, University of Wisconsin-Whitewater, 800 W Main St, Whitewater, WI, 53190, USA

    Nicholas P. Tippery & Tai A. Thompson

  2. Botany Department, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, WI, 53233, USA

    Nicholas P. Tippery

  3. Department of Biology, Andrews University, Berrien Springs, MI, 49104, USA

    Daniel Gonzalez-Socoloske

  4. Meise Botanic Garden, Nieuwelaan 38, 1860, Meise, Belgium

    Frederik Leliaert

  5. Herbário Rosa Mochel-SLUI, Universidade Estadual do Maranhão, Cidade Universitária Paulo VI s/n, São Luís, Maranhão, 65054-970, Brazil

    André Vito Scatigna

  6. Departamento de Ciências Biológicas, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, Av. Pádua Dias 22, Piracicaba, São Paulo, CEP 13418-900, Brazil

    Vinicius Castro Souza

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N.P.T. wrote the main manuscript text with input from A.V.S. and F.L., and N.P.T. prepared the figures. All authors reviewed the manuscript.

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Correspondence to Nicholas P. Tippery.

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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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