Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification Author Bruneau, Anne https://orcid.org/0000-0001-5547-0796 Institut de recherche en biologie vegetale and Departement de Sciences biologiques, Universite de Montreal, 4101 Sherbrooke E., Montreal (QC) H 1 X 2 B 2, Canada anne.bruneau@umontreal.ca Author de Queiroz, Luciano Paganucci https://orcid.org/0000-0001-7436-0939 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Ringelberg, Jens J. https://orcid.org/0000-0003-0567-5210 Department of Systematic and Evolutionary Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland & School of Geosciences, University of Edinburgh, Old College, South Bridge, Edinburgh EH 8 9 YL, UK Author Borges, Leonardo M. https://orcid.org/0000-0001-9269-7316 Universidade Federal de Sao Carlos, Departamento de Botanica, Rodovia Washington Luis, Km 235, 13565 - 905, Sao Carlos, SP, Brazil Author Bortoluzzi, Roseli Lopes da Costa https://orcid.org/0000-0002-7445-7244 Programa de Pos-graduacao em Producao Vegetal, Universidade do Estado de Santa Catarina, Centro de Ciencias Agroveterinarias, Avenida Luiz de Camoes 2090, 88520 - 000, Lages, Santa Catarina, Brazil Author Brown, Gillian K. https://orcid.org/0000-0002-7940-5435 Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Toowong, Queensland, 4066, Australia Author Cardoso, Domingos B. O. S. https://orcid.org/0000-0001-7072-2656 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Programa de Pos-Graduacao em Biodiversidade e Evolucao (PPGBioEvo), Instituto de Biologia, Universidade Federal de Bahia (UFBA), Rua Barao de Jeremoabo, s. n., Ondina, 40170 - 115, Salvador, BA, Brazil Author Clark, Ruth P. https://orcid.org/0000-0001-9974-2933 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author Conceicao, Adilva de Souza https://orcid.org/0000-0002-8800-422X Programa de Pos-graduacao em Diversidade Vegetal, Universidade do Estado da Bahia, Herbario HUNEB, Campus VIII, Rua do Gangorra 503, 48608 - 240, Paulo Afonso, Bahia, Brazil Author Cota, Matheus Martins Teixeira https://orcid.org/0000-0003-0654-7501 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Demeulenaere, Else https://orcid.org/0000-0002-1815-3051 Center for Island Sustainability and Sea Grant, University of Guam, UOG Station, Mangilao, 96923, Guam Author de Stefano, Rodrigo Duno https://orcid.org/0000-0003-1707-4121 Centro de Investigacion Cientifica de Yucatan, A. C. (CICY), Calle 43 No. 130 x 32 y 34, Chuburna de Hidalgo; CP 97205, Merida, Yucatan, Mexico Author Ebinger, John E. Eastern Illinois University, Charleston, IL 61920, USA Author Ferm, Julia https://orcid.org/0000-0002-8762-3942 Department of Ecology, Environment and Plant Sciences, 10691, Stockholm University, Stockholm, Sweden Author Fonseca-Cortes, Andres https://orcid.org/0000-0001-7207-9940 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Gagnon, Edeline https://orcid.org/0000-0003-3212-9688 Department of Integrative Biology, University of Guelph, 50 Stone Road, Guelph (ON) N 1 G 2 W 1, Canada & Chair of Phytopathology, Technical University Munich, 85354 Freising, Germany & Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, UK Author Grether, Rosaura https://orcid.org/0000-0003-2673-665X Departamento de Biologia, Universidad Autonoma Metropolitana-Iztapalapa, Apdo. Postal 55 - 535, 09340 Ciudad de Mexico, Mexico Author Guerra, Ethiene https://orcid.org/0000-0002-9495-1717 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil Author Haston, Elspeth https://orcid.org/0000-0001-9144-2848 Royal Botanic Garden Edinburgh, 20 A Inverleith Row, Edinburgh, EH 3 5 LR, UK Author Herendeen, Patrick S. https://orcid.org/0000-0003-2657-8671 Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, IL 60022, USA Author Hernandez, Hector M. https://orcid.org/0000-0002-1741-5515 Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Cd. Universitaria, 04510 Ciudad de Mexico, Mexico Author Hopkins, Helen C. F. https://orcid.org/0000-0003-4984-8224 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author Huamantupa-Chuquimaco, Isau https://orcid.org/0000-0002-4153-5875 Herbario Alwyn Gentry (HAG), Universidad Nacional Amazonica de Madre de Dios (UNAMAD), AV. Jorge Chavez N ° 1160, Madre de Dios, Peru Author Hughes, Colin E. https://orcid.org/0000-0002-9701-0699 Department of Systematic and Evolutionary Botany, University of Zurich, Zollikerstrasse 107, 8008 Zurich, Switzerland Author Ickert-Bond, Stefanie M. https://orcid.org/0000-0001-8198-8898 Department of Biology & Wildlife & Herbarium (ALA) at the University of Alaska Museum of the North, University of Alaska Fairbanks, P. O. Box 756960, Fairbanks AK 99775 - 6960, USA Author Iganci, Joao https://orcid.org/0000-0002-5740-3666 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil & Programa de Pos-Graduacao em Fisiologia Vegetal, Universidade Federal de Pelotas, Instituto de Biologia, Campus Universitario Capao do Leao, Passeio Andre Dreyfus, Departamento de Botanica, Predio 21, Pelotas, Rio Grande do Sul, 96010 - 900, Brazil Author Koenen, Erik J. M. https://orcid.org/0000-0002-4825-4339 Evolutionary Biology & Ecology, Universite Libre de Bruxelles, Faculte des Sciences, Campus du Solbosch - CP 160 / 12, Avenue F. D. Roosevelt, 50, 1050 Bruxelles, Belgium Author Lewis, Gwilym P. https://orcid.org/0000-0003-2599-4577 Accelerated Taxonomy Department, Royal Botanic Gardens, Kew, Richmond, TW 9 3 AE, UK Author de Lima, Haroldo Cavalcante https://orcid.org/0000-0003-2154-670X Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Instituto Nacional da Mata Atlantica / INMA-MCTI, Av. Jose Ruschi, 4, Centro, 29650 - 000, Santa Teresa, Espirito Santo, Brazil Author de Lima, Alexandre Gibau https://orcid.org/0000-0002-9168-2507 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil & Department of Biological and Environmental Sciences, University of Gothenburg, Gothenburg, Sweden Author Luckow, Melissa https://orcid.org/0009-0007-2543-0516 School of Integrative Plant Science, Plant Biology Section, Cornell University, 215 Garden Avenue, Roberts Hall 260, Ithaca, NY 14853, USA Author Marazzi, Brigitte https://orcid.org/0000-0003-3252-5816 Natural History Museum of Canton Ticino, Viale C. Cattaneo 4, 6900 Lugano, Switzerland Author Maslin, Bruce R. https://orcid.org/0000-0002-3039-0973 Western Australian Herbarium, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia, 6983, Australia & Singapore Herbarium, 1 Cluny Road, Singapore, Singapore Author Morales, Matias https://orcid.org/0000-0001-5540-9725 Instituto de Recursos Biologicos, CIRN-CNIA, INTA. N. Repetto & Los Reseros s. n., Hurlingham, Buenos Aires, Argentina & Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), Godoy Cruz 2290 (C 1425 FQB), Ciudad Autonoma de Buenos Aires, Argentina Author Morim, Marli Pires https://orcid.org/0000-0003-0872-8429 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro, Pacheco Leao 915, 22460 - 030, Rio de Janeiro, RJ, Brazil Author Murphy, Daniel J. https://orcid.org/0000-0002-8358-363X Royal Botanic Gardens Victoria, Melbourne, Victoria, 3004, Australia Author O'Donnell, Shawn A. https://orcid.org/0000-0003-0731-7425 Geography and Environmental Sciences, Northumbria University, Ellison Place, Newcastle upon Tyne, NE 1 8 ST, UK Author Oliveira, Filipe Gomes https://orcid.org/0000-0003-0244-3262 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Oliveira, Ana Carla da Silva https://orcid.org/0000-0001-7042-5360 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Rando, Juliana Gastaldello https://orcid.org/0000-0002-3714-8231 Programa de Pos-graduacao em Ciencias Ambientais, Universidade Federal do Oeste da Bahia, Rua Professor Jose Seabra Lemos 316, 47800 - 021, Barreiras, Bahia, Brazil Author Ribeiro, Petala Gomes https://orcid.org/0000-0002-0070-9971 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Ribeiro, Carolina Lima https://orcid.org/0000-0001-9508-2894 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Santos, Felipe da Silva https://orcid.org/0000-0002-1068-0578 Universidade Estadual de Feira de Santana, Departamento de Ciencias Biologicas, Av. Transnordestina s / n, Campus, Novo Horizonte. 44036 - 900, Feira de Santana, BA, Brazil Author Seigler, David S. https://orcid.org/0009-0003-5177-5893 Department of Plant Biology, University of Illinois, Urbana, IL 61801, USA Author da Silva, Guilherme Sousa https://orcid.org/0000-0002-4250-0017 Instituto de Biologia, Universidade Estadual de Campinas, Campinas, 13083 - 876, Sao Paulo / SP, Brazil Author Simon, Marcelo F. https://orcid.org/0000-0002-5732-1716 Empresa Brasileira de Pesquisa Agropecuaria (Embrapa) Recursos Geneticos e Biotecnologia, Parque Estacao Biologica, Caixa Postal 02372, 70770 - 917, Brasilia / DF, Brazil Author Soares, Marcos Vinicius Batista https://orcid.org/0000-0003-2660-1771 Universidade Federal do Rio Grande do Sul, Programa de Pos-Graduacao em Botanica, Av. Bento Goncalves 9500, Bloco IV - Predio 43433, Porto Alegre, RS, 91501 - 970, Brazil Author Terra, Vanessa https://orcid.org/0000-0001-5669-1304 Instituto de Biologia, Universidade Federal de Santa Maria, 97105 - 900, Santa Maria / RS, Brazil text PhytoKeys 2024 2024-04-03 240 1 552 http://dx.doi.org/10.3897/phytokeys.240.101716 journal article http://dx.doi.org/10.3897/phytokeys.240.101716 1314-2003-240-1 B699D9DE2B435B1093DE3C38C703D430 Pseudosamanea Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11(101): 54. 1930. Figs 241 , 244 , 245 Type . Pseudosamanea guachapele (Kunth) Harms [≡ Acacia guachapele Kunth] The phylogenetic position of Pseudosamanea is not well resolved in the Mimoseae phylogeny ( Koenen et al. 2020a ; Ringelberg et al. 2022 ). The genus is placed in a large, putative hard polytomy in the ingoid clade (Fig. 241 ), but its exact position is difficult to ascertain, and differs across phylogenetic analyses ( Koenen et al. 2020a ). Pseudosamanea is therefore not treated as part of any clade within Mimoseae and is here recognised as a single genus lineage. Description. Unarmed trees 6-30 m, with wide spreading crown; bark fissured and exfoliating in irregular rectangular layers. Stipules triangular to lanceolate, sericeous, caducous. Leaves bipinnate, extrafloral nectaries usually present near the petiole base, another between the terminal pair of pinnae and less frequently along the rachides; pinnae 3-13 pairs, paraphyllidia present, caducous; leaflets 5-30 pairs, obovate, elliptic or lanceolate. Inflorescence units heteromorphic umbelliform capitula, 2-6-fascicled in leaf axil, the terminal flower sessile and stouter. Flowers 5 (6-7)-merous; calyx gamosepalous, ferruginous villose; corolla gamopetalous, campanulate, lobes triangular, canescent to yellowish villose; stamens numerous, the filaments basally united into a tube; pollen in 24-32-celled polyads; ovary ovoid. Fruits indehiscent or dehiscent through one margin, oblong, laterally compressed, valves papery, puberulent, margins slightly thicker. Seeds ovoid, laterally compressed, pleurogram present (Fig. 244 ). Figure 244. General morphology in Pseudosamanea species A, B, E, G Pseudosamanea carbonaria (Britton) E.J.M. Koenen (left) A habit B exfoliating bark E bipinnate microphyllidious leaves and umbelliform capitula G papery oblong laterally compressed pods C, D, F, H Pseudosamanea guachapele (Kunth) Harms (right) C habit D exfoliating bark F bipinnate macrophyllidious leaves and umbelliform capitula H papery oblong laterally compressed fruits. Scale bars: 2 cm ( E ); 4 cm ( F ); 3 cm ( G ); 2.5 cm ( H ). Photo credits A Bioexploradores Farallones, iNaturalist (https://www.inaturalist.org/photos/30575576) B, D, H A Fonseca-Cortes C Juan Manuel de Roux, iNaturalist (https://www.inaturalist.org/photos/112891215) E Juan Manuel de Roux, iNaturalist (https://www.inaturalist.org/photos/54043095) F Cynthia Tercero, iNaturalist (https://www.inaturalist.org/photos/59667691) G Juan Carlos Delgado Madrid, iNaturalist (https://www.inaturalist.org/photos/64538395). Chromosome number. 2 n = 26 [ P. carbonaria (Britton) E.J.M. Koenen and P. guachapele ] (Rico Arce 1992). Included species and geographic distribution. Three species known to date: P. carbonaria , P. cubana (Britton & P. Wilson) Barneby & J.W. Grimes and P. guachapele . Pseudosamanea species occur from the south of Mexico to the north of Peru, with one species in Cuba (Fig. 245 ; Barneby and Grimes 1996 ). The native range of P. carbonaria is not known with certainty, but it is presumed to be native from Colombia, Panama and Venezuela from 700-1800 m elevation and introduced in Indonesia (Koenen 2022), Central America, the Greater Antilles, Peru and Brazil ( Barneby and Grimes 1996 ; under Albizia carbonaria Britton). Pseudosamanea cubana is endemic to the south coast of Cuba to 50 m elevation ( Barneby and Grimes 1996 ). Pseudosamanea guachapele is present from the south of Mexico to the north of Peru between 0-1000 m and introduced in Brazil and Cameroon and probably elsewhere ( Barneby and Grimes 1996 ). Figure 245. Distribution of Pseudosamanea based on quality-controlled digitised herbarium records. See Suppl. material 1 for the source of occurrence data. Ecology. Pseudosamanea carbonaria grows in dry and humid forests ( Barneby and Grimes 1996 ), usually near water courses, its seeds are usually dispersed with the fruit by barochory and hydrochory (pers. obs.). Pseudosamanea cubana grows in riverine forests and savannas with palms ( Barneby and Grimes 1996 ). Pseudosamanea guachapele grows in dry forests ( Barneby and Grimes 1996 ). Little is known about the interactions with ants, floral visitors and pollinators. Etymology. From the Greek pseudo (= false) and Samanea , due to the similarity to that genus (Lewis and Rico Arce 2005). Human uses. The three species can be used as timber. Pseudosamanea carbonaria is cultivated for shading coffee crops, P. guachapele is used as living fencing in Colombia, and both are used as ornamentals ( Barneby and Grimes 1996 ). Notes. As defined by Barneby and Grimes (1996) , Pseudosamanea included two species of macrophyllidious trees (Fig. 244F ), P. cubana and P. guachapele . The phylogenomic analyses of Ringelberg et al. (2022) showed Albizia carbonaria to be sister to Pseudosamanea , rather than grouping with species of Albizia section Arthrosamanea (Britton & Rose) Barneby & J.W. Grimes (now the genus Pseudalbizzia Britton & Rose; Peraza et al. 2022 ), where it had previously been classified. Albizia carbonaria was combined in Pseudosamanea based on molecular and morphological evidence ( Koenen 2022a ). Although P. carbonaria is a microphyllidious tree (Fig. 244E ), it shares with the other two species of Pseudosamanea exfoliating bark (Fig. 244B, D ), heteromorphic umbelliform inflorescences (Fig. 244E, F ) in which the central flower is bigger and sessile, and oblong laterally compressed papery legumes (Fig. 244G, H ) ( Koenen 2022a ). Pollen also supports this recircumscription, P. carbonaria having pollen with 32-celled polyads as in P. guachapele ( Koenen 2022a ). Taxonomic references. Barneby and Grimes (1996) ; Koenen (2022a) ; Lewis and Rico Arce (2005).