Abstract
The latest phylogenetic and taxonomic studies emphasizing Phyllanthus (Phyllanthaceae) have significantly altered the relationships between its taxa. In this work, we inferred a phylogeny for the Brazilian phyllocladiferous species of Phyllanthus, traditionally included within Phyllanthus sects. Choretropsis and Xylophylla from eastern Brazil and Caribbean, respectively. Our analysis demonstrated that Brazilian phyllocladiferous species has no relation to P. sect. Xylophylla from the Caribbean, but sect. Choretropsis, as it currently stands, does not form a monophyletic group. Thus, phylloclades arose in Phyllanthus at least in five distinct clades. Most species of sect. Choretropsis are recovered among representatives of other groups (sects. Antipodanthus and Phyllanthus subsect. Claussenianii), but no macromorphological feature could be traced for the clades we recovered. One exception is a clade containing P. dracenoides, P. gladiatus and P. pedicellatus, species with dracenoid habit and exclusive to the humid dense forests along Atlantic Forest domain, for which we describe a new section.
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Acknowledgements
We would like to thank the Universidade Estadual de Campinas, in the figure of Professor Dr. Samantha Koehler, for the infrastructure of the Molecular Biology Laboratory, used for the extractions and amplifications performed in this study. PO also thanks Raquel Moura Machado for her help in the laboratory activities. This study was financed in part by the Coordination for the Improvement of Higher Education Personnel – Brazil (CAPES) – Finance Code 001, and the National Council for Scientific and Technological Development [processes 141913/2020-6 and 442162/2020-0]. PO also thanks The Mohamed bin Zayed Species Conservation Fund [Grant 210528056] and IDEA WILD for support [Grant Number ORLABRAZ0121].
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P.O. designed and wrote the manuscript and carried out the laboratory activities. O.L.M.S. performed the bioinformatics analysis and prepared the figures. I.C. wrote and proofread the manuscript. V.C.S. wrote and proofread the manuscript.
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This study was financed in part by the Coordination for the Improvement of Higher Education Personnel – Brazil (CAPES) – Finance Code 001, and the National Council for Scientific and Technological Development [processes 141913/2020-6 and 442162/2020-0]. PO also thanks The Mohamed bin Zayed Species Conservation Fund [grant 210528056] and IDEA WILD for support [grant number ORLABRAZ0121].
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Newly generated and downloaded sequences.
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Appendix
Appendix
Supplementary material. Voucher information for the samples included in this work, represented as follows: Taxon name, origin, collector and collection number, herbarium code, accession numbers for ITS and matK. Newly generated sequences are in bold.
Astrocasia: Astrocasia neurocarpa (Müll.Arg.) I.M.Johnst. ex Standl., United States (cultivated), Wurdack 743 (DAV), AY936649, AY830261. Breynia: Breynia disticha J.R.Forst. & G.Forst., Netherlands, Utrecht Botanical Garden, Bouman and Verwijs RWB024 (L), MN915814, MN916082. Breynia androgyna (L.) Chackrab. & N.P.Balakr., Chachoengsao, VanWelzen 2006-4 (L), EU623563, EU643748. Cathetus: Cathetus beckleri (Müll.Arg.) I.Telford & J.J.Bruhl, Australia, Hosking 2680 (NE), MN915861, MN916127. Cathetus chrysanthus (Baill.) R.W.Bouman, New Caledonia, Munzinger and McPherson 796 (MO), AY936680, AY936585. Cathetus fasciculata Lour., China, Hong Kong, Bouman et al. 26 (L), MN915895, MN916154. Cathetus kerstingii (Jean F.Brunel) R.W.Bouman, Guinea, Darbyshire 562 (WAG), MN915950, MN916189. Cathetus myrtifolius (Wight) R.W.Bouman, Sri Lanka, Kathriarachchi et al. 12 (K), AY936712, AY936616. Cathetus petraeus (A.Chev. & Beille) R.W.Bouman, Liberia, Blyden 1037 (WAG), MN916026, MN916239. Cathetus welwitschianus (Müll.Arg.) R.W.Bouman, Tanzania, Bidgood et al. 1882 (K), AY936739, AY936640. Cathetus wheeleri (G.L.Webster) R.W.Bouman, Sri Lanka, Kathriarachchi et al. 33 (K), AY936740, AY936641. Cicca: Cicca acida (L.) Merrl., Thailand, Samuel 702-19 (K), AY936666, AY936572. Cicca ankarana (Leandri) R.W.Bouman, Ralimanana et al. 663 (K), MN915851, MN916118. Cicca chacoensis (Morong.) R.W.Bouman, Paraguay, Morong, Krapovickas et al. 45628 (K), AY936677, AY936582. Cicca physocarpa (Müll.Arg.) R.W.Bouman, Gabon, McPherson 16148 (WAG), MN916030, MN916243. Cicca pinnata (Wight) R.W.Bouman, Zimbabwe, Caster and Coates-Palgrave 2189 (K), AY936724, AY936626. Cicca polyantha (Pax) R.W.Bouman, Cameroon, Breteler 1938 (WAG), MN916033, MN916246. Dendrophyllanthus: Dendrophyllanthus cuscutiflorus (S.Moore) R.W.Bouman, Singapore, Singapore Botanical Garden, Yu 61 (L), MN915901, MN916299. Dendrophyllanthus pancherianus (Baill.) R.W.Bouman, New Caledonia, McPherson and Munzinger 18264 (MO), AY936721, AY936623. Emblica: Emblica officinallis Gaertner, India, Chase 14459 (K), AY936689, AY936594. Emblica urinaria (L.) R.W.Bouman, Philippines, Majaducon 5750 (L), MN916071, MN916273. Flueggea: Flueggea virosa (Roxb. ex Willd.) Royle, Singapore, Singapore Botanical Garden, Yu 64 (L), MN915825, MN916092. Glochidion: Glochidion carinatum (Beille) R.W.Bouman, Cambodia, Toyama et al. 3212 (FU),_, MN916138. Glochidion ellipticum Wight, China, Yunnan, Xishuangbanna Tropical Botanical Garden, Bouman and Yong 58 (HITBC), MN915826, MN916093. Heterosavia: Heterosavia bahamensis (Britton) Hoffm., Fairchild Tropical Garden (KW048), AY936749, AY830284. Kirganelia: Kirganelia dinklagei (Pax) R.W.Bouman, Gabon, Bissiengou (WAG), MN915908, MN916333. Kirganelia flexuosa (Siebold & Zuccarini) R.W.Bouman, China, Chow 132 (L), MN915927, MN916174. Kirganelia fuscolurida (Müll.Arg.) R.W.Bouman, Madagascar, Schatz 1737 (WAG), MN915934, MN916179. Kirganelia microcarpa (Benth.) Hurusawa and Tanaka, China, Yunan, Xishuangbanna Tropical Botanical Garden, Bouman and Yong RWB056 (HITBC), MN915985, MN916207. Kirganelia purpurea (Müll.Arg.) R.W.Bouman, Namibia, Ward et al. 10442 (K), MN916042, MN916254. Lingelsheimia: Lingelsheimia sp., Madagascar, Rabenantoandro et al. 1115 (MO), AY936662, AY830272. Lysiandra: Lysiandra calycina (Labill.) R.W.Bouman, Australia, Chase 2163 (K), AY936674, AY552446. Lysiandra carpentariae (Müll.Arg.) R.W.Bouman, Australia, Clarkson and Neldner 8410 (L), MN915877, MN916140. Margaritaria: Margaritaria discoidea (Baill.) G.L.Webster, Mayotte, Comoro Island, Barthelat 1092 (K), AY936664, AY830274. Moeroris: Moeroris amara (Schumach. & Thonner) R.W.Bouman, Gabon, Wieringa 8189 (WAG), MN915847, MN916114. Moeroris boehmii (Pax) R.W.Bouman, Kenya, Wieringa 8841 (WAG), MN915865, MN916129. Moeroris debilis Klein ex Willd, Sri Lanka, Kathriarachchi et al. 15 (K), AY936686, AY936591. Moeroris lokohensis (Leandri) R.W.Bouman, Madagascar, Ralimanana et al. 414 (K), AY936704, AY936606. Moeroris madagascariensis (Müll.Arg.) R.W.Bouman, Madagascar, McPherson 18925 (MO), MN915973, MN916317. Moeroris nummulariifolia (Poir.) R.W.Bouman, Madagascar, Hoffmann et al. 304 (K), AY936715, AY552444. Moeroris sepialis (Müll.Arg.) R.W.Bouman, Kenya, Luke et al. 7112 (K), AY936732, AY936633. Moeroris tenella Roxb., Comoro Island, Hoffmann et al. 372 (K), AY936733, AY936634. Moeroris stipulata Raf., Suriname, Gietling 114 (WAG), MN916062, MN916269. Nellica: Nellica maderaspatensis (L.) Raf., Australia, Hunter et al. 1532 (K), AY936707, AY936609. Nymphanthus: Nymphanthus glauscescens (Baill. Ex Miquel) R.W.Bouman, China, Yunan, Xishuangbanna Tropical Botanical Garden, Bouman and Yong 66 (HITBC), MN916040, MN916252. Nymphanthus longifolius (Miq.) R.W.Bouman, Thailand, TRP5004102 (BK), AB550090,_. Nymphanthus ruber (Lour.) Spreng., China, Hainan, Lee et al. 2441 (CUHK), AY765298,_. Nymphanthus watsonii (Airy Shaw) R.W.Bouman, Malaysia, Peninsular part, Yu 225 (L), MN916076, MN916278. Phyllanthus: Phyllanthus acuminatus Vahl, Venezuela, Breteler 4238 (WAG), MN915837, MN916109. Phyllanthus angustifolius (Sw.) Sw., Germany, Bayreuth Botanical Garden living collection, Lauerer 091479, no voucher, MN915849, MN916116. Phyllanthus angustissimus Müll.Arg., Brazil, Minas Gerais, Serranópolis de Minas, O.S. Ribas and J.M. Silva 7748 (MBM), OP740340,_. Phyllanthus arbuscula (Sw.) J.F.Gmel, Brazil, USP campus, Cordeiro 3586 (SP), OP740351, OP961939. Phyllanthus arbuscula (Sw.) J.F.Gmel, Belgium, Meisse, living collection, Reynders 19074182 (L), MN915855, MN916123. Phyllanthus arenicola Casar, Brazil, Maas and Carauta s.n. (U),_, MN916124. Phyllanthus attenuatus Miq., Venezuela, Breteler 4696 (WAG), MN915856, MN916125. Phyllanthus caparaoensis G.L.Webster, Brazil, Espírito Santo, Santa Teresa, Orlandini et al. (ESA), OP740341, OP961940. Phyllanthus caroliniensis Walter, Guyana, Jansen-Jacobs et al. 4762 (K), AY936675, AY936580. Phyllanthus chapadensis P.Orlandini & M.J.Silva, Brazil, Goiás, Alto Paraíso de Goiás, M.M. Arbo et al. 3568 (MBM!), OP740342,_. Phyllanthus choretroides Müll.Arg., Brazil, Minas Gerais, Santana de Pirapama, D.C. Zappi 763 (SPF!), OP740348, OP961941. Phyllanthus chryseus Howard, Cuba, Van ee et al. 387 (K), AY936681, AY936586. Phyllanthus chryseus R.A.Howard, Cuba, T. Borsch et al. 4280 (B, HAJB),_, LS975796. Phyllanthus cinctus Urb., Cuba, Ekman 19166, MN915892, MN916151. Phyllanthus claussenii Müll.Arg., Brazil, Minas Gerais, Hatschbach 64117 (U), MN915894, MN916153. Phyllanthus claussenii Müll.Arg., Brazil, Melo 2261 (HUEFS 29998), AY936683, AY936588. Phyllanthus comosus Urb., Cuba, T. Borsch et al. 4271 (HAJB), LS975727, LS975785. Phyllanthus dawsonii Steyerm., Brazil, da Silva 2073 (DAV), MN915902, MN916159. Phyllanthus discolor Poepp. ex Spreng., Cuba, Berazain, Rankin and Köhler 71878 (K), AY936688, AY936593. Phyllanthus dictyospermus Müll.Arg., Brazil, Santos 5712 (DAV), MN915907, MN916162. Phyllanthus dracaenoides P.Orlandini & I.Cordeiro, Brazil, Bahia, Itacaré, Orlandini et al. 284 (ESA), OP740335, OP961942. Phyllanthus dracaenoides P.Orlandini & I.Cordeiro, Brazil, Bahia, Itacaré, Orlandini et al. 285 (ESA), OP740336, OP961943. Phyllanthus epiphyllanthus L., Belgium, Meisse, living collection, Reynders 19840633 (L), MN915919, MN916169. Phyllanthus evanescens Brandegee, Nicaragua, Stevens 32461 (MO), MN915921, MN916339. Phyllanthus excisus Urban, Cuba, W.Greuter et al. 28281 (B, HAJB, P-Gr), LS975746, LS975806. Phyllanthus flagelliformis Müll.Arg., Brazil, Bahia, Boa Nova, L. Daneu 743 (CEPEC), OP740349, OP961944. Phyllanthus fluitans Benth. ex Müll.Arg., Netherlands, Kathriarachchi 70 (WU), AY936693, AY936597. Phyllanthus gladiatus Müll.Arg., Brasil, Sergipe, Areia Branca, Orlandini (ESA), OP740339, OP961945. Phyllanthus heliotropus C.Wight ex Griseb, Cuba, Maas et al. 7762 (U), MN915946, MN916186. Phyllanthus juglandifolius Willd., Ecuador, Holm-Nielsen 22881 (K), AY936699, AY936602. Phyllanthus klotzschianus Müll.Arg., Brazil, Bahia, Barra Grande, Orlandini et al. 292 (ESA), OP740343, OP961946. Phyllanthus klotzschianus Müll.Arg., Brazil, Bahia, Barra Grande, Orlandini et al. 293 (ESA),_,OP961947. Phyllanthus klotzschianus Müll.Arg., Brazil, Bahia, Mucugê, M. Groppo Jr. et al. 780 (K), AY936702, AY936605. Phyllanthus lindenianus Baill., Dominican Republic, Fuertes 345 (K),_, MN916200. Phyllanthus mimosoides Sw., Belgium, Meisse living collection, Reynders 19074179 (L), MN915989, MN916209. Phyllanthus mirificus G.L.Webster, Cuba, T. Borsch et al. 5021 (B, HAJB), LS975774, LS975835. Phyllanthus niruri L., Brazil, Espírito Santo, Santa Teresa, Orlandini et al. 264 (ESA), OP740350, OP961948. Phyllanthus niruri L., Costa Rica, Cascante 1297 (K), AY936713, AY936617. Phyllanthus orbicularis Kunth, Borneo, Eschevaria s.n. (L), MN916012, MN916225. Phyllanthus pachystylus Urb., Cuba, Van Ee et al. 402 (K), AY936720, AY936622. Phyllanthus pedicellatus P.Orlandini, I.Cordeiro & V.C.Souza, Brazil, Minas Gerais, Santa Maria do Salto, A. Campos-Rocha and N. Oleas 1698 (ESA), OP740337, OP961949. Phyllanthus pedicellatus P.Orlandini, I.Cordeiro & V.C.Souza, Brazil, Minas Gerais, Santa Maria do Salto, A. Campos-Rocha and N. Oleas 1698 (SP), OP740338, OP961950. Phyllanthus pseudocicca Grisebach, Cuba, P.A. González HFC 87681 (HAJB), LS975789, LS975820. Phyllanthus purpusii Brandegee, United States of America, Berkeley, cultivated from Mexico, Breedlove 42730, living collection Berkeley, no voucher, MN916043, MN916255. Phyllanthus ramillosus Müll.Arg., Brazil, Arbo 6945 (DAV), MN916044, MN916256. Phyllanthus salviifolius Kunth, Ecuador, Lewis et al. 3500 (K), AY936730, AY936631. Phyllanthus sarothamnoides Govaerts & Radcl.-Sm., Brazil, Bahia, Palmeiras, Orlandini et al. 296 (ESA), OP740347, OP961951. Phyllanthus sarothamnoides Govaerts & Radcl.-Sm., Brazil, Bahia, Palmeiras, Orlandini et al. 297 (ESA), OP740344, OP961952. Phyllanthus sellowianus (Klotzsch) Müll.Arg., United Kingdom, RBG Kew, living collection, Chase 14776 (K), MN916054, MN916263. Phyllanthus spartioides Pax & K.Hoffm., Brazil, Bahia, Rio de Contas, R.M. Harley and D.J. Hind 26107 (SP), OP740346, OP961953. Phyllanthus spartioides Pax & K.Hoffm., Brazil, Bahia, Rio de Contas, R.M. Harley et al. 19709 (SPF), OP740345,_. Phyllanthus spruceanus Müll.Arg., Guyana, Maas and Maas 474 (U), MN916061, MN916353. Phyllanthus subcarnosus C.Wright ex Griseb., Cuba, Mayotte, T. Borsch et al. 4707 (B, HAJB), LS975741, LS975800. Phyllanthus symphoricarpoides Kunth, Cuatrecasas 18421 (U), MN916064, MN916321. Phyllanthus tuerckheimii G.L.Webster, Guatemala, Mendez 8022 (DAV), MN916070, MN916272. Phyllanthus vacciniifolius (Müll.Arg.) Müll.Arg., Venezuela, Hokche et al. 854 (U), MN916073, MN916275. Plagiocladus: Plagiocladus diandrus (Pax & Drude) J.F.Brunel, Gabon, de Wilde and de Wilde 11641 (WAG), AY936687,_. Plagiocladus diandrus (Pax & Drude) J.F.Brunel, Gabon, Wieringa 2903 (WAG),_, AY936592. Synostemon: Synostemon albiflorus (F.Muell. ex Müll.Arg.) Airy Shaw, Australia, Forster 32329 (NE), MN916077, MN916279.
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Orlandini, P., da Silva, O.L.M., Cordeiro, I. et al. Phylogenetics of Brazilian phyllocladiferous Phyllanthus (Phyllanthaceae): a new section exclusive from the Atlantic Rain Forest, with morphological and molecular support. Plant Syst Evol 309, 36 (2023). https://doi.org/10.1007/s00606-023-01871-1
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DOI: https://doi.org/10.1007/s00606-023-01871-1