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 Tribe Sclerolobieae Benth. & Hook. f., Gen. Pl. 1: 436. 1865. Figs 78 , 79 , 80 , 81 , 82 , 83 , 84 , 85 , 86 Tachigalieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Tr. Nov.]: 252. 1943. Type: Tachigali Aubl. Type . Sclerolobium Vogel [= Tachigali Aubl.] Included genera (5). Arapatiella Rizzini & A. Mattos (2 species), Diptychandra Tul. (2), Jacqueshuberia Ducke (7), Moldenhawera Schrad. (12), Tachigali Aubl. (80-90). Description. Trees or shrubs, unarmed. Stipules present, persistent or caducous. Leaves pinnate or bipinnate, usually paripinnate, rarely imparipinnate; myrmecophilous domatia present or absent; leaflets opposite; extrafloral nectaries absent. Inflorescences terminal or lateral racemes or panicles. Flowers bisexual, bilaterally or radially symmetrical; hypanthium cupular or cylindrical; sepals 5, free or rarely connate at base; petals 5, free, equal or the dorsal, standard petal differentiated; stamens often 10 (18), rarely only one stamen and 7 or 9 staminodes, free or connate, monomorphic or dimorphic, anthers longitudinally dehiscent, rarely opening through pores; pollen in monads, rarely in tetrads or associated with viscin threads, the exine scabrate-punctate to reticulate; ovary sessile or stipitate; nectary disk absent. Fruit dehiscent along both sutures or indehiscent with flaking exocarp. Seeds ellipsoid to obovoid, laterally compressed or complanate, sometimes winged, seed coat hard or soft, endosperm scarce. Distribution. Mostly Neotropical, extending from Central to South America, in tropical rainforests, tropical cloud forests, seasonally dry forests, and fire-prone savannas. Clade-based definition. The most inclusive crown clade containing Moldenhawera floribunda Schrad. and Tachigali guianensis (Benth.) Zarucchi & Herend., but not Caesalpinia brasiliensis L., Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig. 78 ). Figure 78. Generic relationships in tribe Sclerolobieae . For description of phylogeny and support values, see Fig. 6 caption (page 63). Notes. Although the generic name Sclerolobium is currently treated under Tachigali , the tribe name Sclerolobieae ( Bentham 1865 ) has priority over Tachigalieae ( Nakai 1943 ). Tribe Sclerolobieae , as herein circumscribed based on phylogenomic analyses ( Ringelberg et al. 2022 ), includes five genera (Fig. 78 ), most of which were never closely associated with each other in traditional classifications of Caesalpinioideae ( Polhill and Vidal 1981 ; Polhill 1994 ; Lewis 2005b ; Ulibarri 2008 ). For example, Polhill and Vidal (1981) treated Tachigali and Diptychandra as part of the Sclerolobium group of the old sense tribe Caesalpinieae , whereas Arapatiella , Jacqueshuberia , and Moldenhawera were classified together with ten other Caesalpinioideae genera in the Peltophorum group of the same tribe. Not only have the relationships amongst Sclerolobieae genera now been strongly resolved for almost all nodes (Fig. 78 ), but also their monophyly is strongly supported by morphological characters and more densely sampled phylogenies based on a few plastid and nuclear loci ( Haston et al. 2003 , 2005 ; Bruneau et al. 2008 ; Chomicki et al. 2015 ; Huamantupa-Chuquimaco 2020 ). The tribe is not defined by obvious synapomorphies, although there is a tendency for the occurrence of unusually divided or foliaceous stipules (except for Diptychandra ). The genera are distinct in leaf morphology (pinnate or bipinnate leaves) (Fig. 79 ), floral symmetry (radial or bilateral), pollination syndrome (bees or birds), pollen presentation (monads or tetrads), fruit morphology, seed morphology (winged or non-winged), and dispersal syndrome (autochory, hydrochory, or anemochory). The high morphological variation found within Sclerolobieae in terms of floral architecture (Fig. 80 ) and fruit type (Fig. 81 ) may explain why its constituent genera have long remained unnoticed as belonging to the same natural group. For example, Tachigali and Diptychandra have similarly small flowers, which are mostly radially symmetrical, but ontogenetically the apparently morphologically homogeneous flowers of Tachigali display considerable variation ( Casanova et al. 2020 ) and are quite different from those of Diptychandra . Many Tachigali species also possess bilaterally symmetrical flowers, with a curved hypanthium similar to the much bigger flowers of Arapatiella and Jacqueshuberia . The Moldenhawera flowers are striking in their more or less radially symmetrical Malpighiaceae -like architecture, where the often-yellow petals are marginally crimped and long-clawed, and the cluster of staminodes are much smaller than the single bearded fertile stamen and resemble the oil glands (elaiophores) typical of the Malpighiaceae flowers ( Queiroz et al. 2023 ). The fruits are wind-dispersed, compressed cryptosamaras in Tachigali , whereas all other genera in the tribe have elastically dehiscent, laterally compressed pods. Within Sclerolobieae there is also a high degree of variation in pollen ( Graham and Barker 1981 ; Banks et al. 2003 ; Banks and Lewis 2018 ), which can be colporate, in monads or tetrads, and with scabrate-punctate to reticulate ornamentation; pollen connecting viscin threads are present in Arapatiella and Jacqueshuberia . Faria et al. (2022) report nodulation with a fixation thread type of nodule anatomy in at least one species each of Moldenhawera , Tachigali , Jacqueshuberia , and a confirmed absence of nodulation in Arapatiella (unknown in Diptychandra ). Figure 79. Vegetative morphology of Sclerolobieae A Moldenhawera nutans L.P. Queiroz, G.P. Lewis & Allkin shrubby individual in a Restinga sand dune field B Diptychandra aurantiaca (Mart.) Tul. tree in a Caatinga seasonally dry setting C Arapatiella psilophylla (Harms) R.S. Cowan flowering tree D Tachigali amplifolia (Ducke) Barneby large, buttressed tree E Jacqueshuberia purpurea Ducke pinnate stipule F leaf G detail of a leaf H Tachigali rugosa (Mart. ex Benth.) Zarucchi & Pipoly leaf with characteristic inversely symmetrical leaflets I, J Tachigali sp. details of leaf domatia K, L variation in stipule shape. Photo credits A, C, E-L D Cardoso B RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ D I Huamantupa. Figure 80. Floral morphology of Sclerolobieae A Moldenhawera lushnathiana Yakovlev highly branched inflorescence B Moldenhawera nutans L.P. Queiroz, G.P. Lewis & Allkin flowering branch, showing the typical navicular leaves with rust-coloured leaflet abaxial surface C flowers D Diptychandra aurantiaca (Mart.) Tul. flowering branch E inflorescence F Arapatiella psilophylla (Harms) R.S. Cowan inflorescences G flowers H Jacqueshuberia purpurea Ducke flower I Tachigali macrostachya Huber candelabrum-like inflorescences J Tachigali amarumayu Huamantupa, H.C. Lima & D.B.O.S. Cardoso densely paniculate inflorescences K Tachigali paraensis (Huber) Barneby radially symmetrical flowers L Tachigali paniculata Aubl. bilaterally symmetrical flowers M Tachigali macrostachya bilaterally symmetrical flowers. Photo credits A LP Queiroz B, C, F, G, I, K-M D Cardoso J I Huamantupa D, E RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ H M Cohn-Haft. Figure 81. Fruit morphology of Sclerolobieae A Moldenhawera polysperma (Vell.) Stellfeld old fruit valves B Diptychandra aurantiaca (Mart.) Tul. fruits C fruit opened showing the characteristic winged seed D Arapatiella psilophylla (Harms) R.S. Cowan immature fruit E Jacqueshuberia purpurea Ducke immature fruits F Tachigali macrostachya Huber immature fruits G Tachigali rugosa (Mart. ex Benth.) Zarucchi & Pipoly mature fruits. Photo credits A C Vivas B, C RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ D-F D Cardoso G LP Queiroz.