Monograph |
Corresponding author: Anne Bruneau ( anne.bruneau@umontreal.ca ) Academic editor: Sandy Knapp
© 2024 Anne Bruneau, Luciano Paganucci de Queiroz, Jens J. Ringelberg, Leonardo M. Borges, Roseli Lopes da Costa Bortoluzzi, Gillian K. Brown, Domingos B. O. S. Cardoso, Ruth P. Clark, Adilva de Souza Conceição, Matheus Martins Teixeira Cota, Else Demeulenaere, Rodrigo Duno de Stefano, John E. Ebinger, Julia Ferm, Andrés Fonseca-Cortés, Edeline Gagnon, Rosaura Grether, Ethiéne Guerra, Elspeth Haston, Patrick S. Herendeen, Héctor M. Hernández, Helen C. F. Hopkins, Isau Huamantupa-Chuquimaco, Colin E. Hughes, Stefanie M. Ickert-Bond, João Iganci, Erik J. M. Koenen, Gwilym P. Lewis, Haroldo Cavalcante de Lima, Alexandre Gibau de Lima, Melissa Luckow, Brigitte Marazzi, Bruce R. Maslin, Matías Morales, Marli Pires Morim, Daniel J. Murphy, Shawn A. O’Donnell, Filipe Gomes Oliveira, Ana Carla da Silva Oliveira, Juliana Gastaldello Rando, Pétala Gomes Ribeiro, Carolina Lima Ribeiro, Felipe da Silva Santos, David S. Seigler, Guilherme Sousa da Silva, Marcelo F. Simon, Marcos Vinícius Batista Soares, Vanessa Terra.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bruneau A, Queiroz LP, Ringelberg JJ, Borges LM, Bortoluzzi RLC, Brown GK, Cardoso DBOS, Clark RP, Conceição AS, Cota MMT, Demeulenaere E, Duno de Stefano R, Ebinger JE, Ferm J, Fonseca-Cortés A, Gagnon E, Grether R, Guerra E, Haston E, Herendeen PS, Hernández HM, Hopkins HCF, Huamantupa-Chuquimaco I, Hughes CE, Ickert-Bond SM, Iganci J, Koenen EJM, Lewis GP, Lima HC, Lima AG, Luckow M, Marazzi B, Maslin BR, Morales M, Morim MP, Murphy DJ, O’Donnell SA, Oliveira FG, Oliveira ACS, Rando JG, Ribeiro PG, Ribeiro CL, Santos FS, Seigler DS, Silva GS, Simon MF, Soares MVB, Terra V (2024) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 1-552. https://doi.org/10.3897/phytokeys.240.101716
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Caesalpinioideae is the second largest subfamily of legumes (Leguminosae) with ca. 4680 species and 163 genera. It is an ecologically and economically important group formed of mostly woody perennials that range from large canopy emergent trees to functionally herbaceous geoxyles, lianas and shrubs, and which has a global distribution, occurring on every continent except Antarctica. Following the recent re-circumscription of 15 Caesalpinioideae genera as presented in Advances in Legume Systematics 14, Part 1, and using as a basis a phylogenomic analysis of 997 nuclear gene sequences for 420 species and all but five of the genera currently recognised in the subfamily, we present a new higher-level classification for the subfamily. The new classification of Caesalpinioideae comprises eleven tribes, all of which are either new, reinstated or re-circumscribed at this rank: Caesalpinieae Rchb. (27 genera / ca. 223 species), Campsiandreae LPWG (2 / 5–22), Cassieae Bronn (7 / 695), Ceratonieae Rchb. (4 / 6), Dimorphandreae Benth. (4 / 35), Erythrophleeae LPWG (2 /13), Gleditsieae Nakai (3 / 20), Mimoseae Bronn (100 / ca. 3510), Pterogyneae LPWG (1 / 1), Schizolobieae Nakai (8 / 42–43), Sclerolobieae Benth. & Hook. f. (5 / ca. 113). Although many of these lineages have been recognised and named in the past, either as tribes or informal generic groups, their circumscriptions have varied widely and changed over the past decades, such that all the tribes described here differ in generic membership from those previously recognised. Importantly, the approximately 3500 species and 100 genera of the former subfamily Mimosoideae are now placed in the reinstated, but newly circumscribed, tribe Mimoseae. Because of the large size and ecological importance of the tribe, we also provide a clade-based classification system for Mimoseae that includes 17 named lower-level clades. Fourteen of the 100 Mimoseae genera remain unplaced in these lower-level clades: eight are resolved in two grades and six are phylogenetically isolated monogeneric lineages. In addition to the new classification, we provide a key to genera, morphological descriptions and notes for all 163 genera, all tribes, and all named clades. The diversity of growth forms, foliage, flowers and fruits are illustrated for all genera, and for each genus we also provide a distribution map, based on quality-controlled herbarium specimen localities. A glossary for specialised terms used in legume morphology is provided. This new phylogenetically based classification of Caesalpinioideae provides a solid system for communication and a framework for downstream analyses of biogeography, trait evolution and diversification, as well as for taxonomic revision of still understudied genera.
Classification, diversity, Fabaceae, Leguminosae, Mimosoideae, phylogenomics, taxonomy
Anne Bruneau1, Luciano Paganucci de Queiroz2, Jens J. Ringelberg3,4
In 1981, the first volumes of the Advances in Legume Systematics (ALS) series were published as Parts 1 and 2. Edited by Roger M. Polhill and Peter H. Raven, the intention was to disseminate new research results and information about the taxonomy, systematics and evolution of Leguminosae. It is a testimony to the success of this series, that more than four decades later, as part of the ALS series, we here present a 14th volume, published in two parts, dedicated to the classification of subfamily Caesalpinioideae.
The first part of Advances in Legume Systematics 14 (Classification of Caesalpinioideae Part 1: new generic delimitations) was published in 2022 as a special issue of PhytoKeys (https://phytokeys.pensoft.net/issue/3247/) edited by Colin E. Hughes, Luciano P. de Queiroz and Gwilym P. Lewis. ALS14 Part 1 included 16 papers focused on generic delimitation in 15 clades of Caesalpinioideae based on new phylogenomic analyses presented there. Circumscriptions of 15 genera were revised to deal with issues of non-monophyly. In this second part of Advances in Legume Systematics 14, we present a higher-level classification of Caesalpinioideae that includes a fully updated synopsis of the 163 genera now recognised (ca. 4680 species) which are placed in 11 newly described or reinstated tribes, alongside a clade-based classification for the largest of these tribes, the recircumscribed Mimoseae.
As editors of Part 2 of Advances in Legume Systematics 14, we thank the 45 contributors from 14 countries for their diligence and collaborative spirit. This ambitious project that garners the expertise from colleagues around the world could only be possible with the willingness and hard work of all, several of whom contributed to revisions of texts in addition to their own. In particular, we thank Colin Hughes, Erik Koenen and Gwilym Lewis for discussions and ideas that initiated this project, and Colin Hughes and Gwilym Lewis for their important contributions to writing, revision, and editing of texts. We thank colleagues and organisations around the world who have shared images of Caesalpinioideae. We are indebted to the PhytoKeys editorial team for their support and careful reviews, and in particular Sandy Knapp, the handling editor, and two anonymous reviewers, as well as Fabian Michelangeli and Tiina Särkinen, for their comments that helped to improve this compendium. Finally we acknowledge financial support from institutions and funding sources that supported Open Access publication costs: Université de Montréal (Canada); Universidade Estadual de Feira de Santana (Brazil); Royal Botanic Gardens, Kew (UK); Royal Botanic Gardens, Edinburgh (UK); Chicago Botanic Garden (USA); Center for Island Sustainability and Sea Grant of the University of Guam; Universidad Nacional Autónoma de México; Technical University Munich (Germany); Royal Botanic Gardens Victoria, Melbourne (Australia).
Anne Bruneau1, Luciano Paganucci de Queiroz2, Jens J. Ringelberg3,4, Gwilym P. Lewis10, Colin E. Hughes3
Citation: Bruneau A, Queiroz LP, Ringelberg JJ, Lewis GP, Hughes CE (2024) Introduction - Classification of subfamily Caesalpinioideae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 7–31. https://doi.org/10.3897/phytokeys.240.101716
With close to 800 genera and more than 22,000 species (
In 2017, the Legume Phylogeny Working Group (
Within subfamilies, new phylogenies of many legume groups have unequivocally demonstrated the non-monophyly of the tribes recognised in the classifications of
A new higher-level classification of subfamily Caesalpinioideae is therefore now both feasible and timely. Here we use the phylogenomic backbones for subfamily Caesalpinioideae from
Caesalpinioideae sensu
A Caesalpinioideae genus richness across floristic realms (according to
Map showing the global distribution of Caesalpinioideae species richness. Numbers of Caesalpinioideae species per one degree latitude / longitude grid cell. Infraspecific taxa are not counted individually but are included at the species level. All maps in this special issue are based on quality-controlled occurrence data from digitised herbarium specimens and floristic surveys (see Suppl. material
Caesalpinioideae are almost entirely woody perennials, but they are extremely diverse in stature and habit – including lianas, trees of all sizes, up to rain forest canopy emergents (e.g., Cedrelinga Ducke, Dinizia Ducke), shrubs, functionally herbaceous geoxyles, and two herbaceous aquatic species (Neptunia Lour.). Similarly, the subfamily is highly diverse in floral and fruit morphology. One of the hallmarks of Caesalpinioideae, as in many other plant groups, is repeated morphological and ecological convergences whereby similar leaf, flower and fruit morphologies, and ecological adaptations, have apparently been reinvented multiple times across lineages and through time (
Caesalpinioideae is the only legume subfamily that has bipinnate leaves, which are prevalent but not universal across the subfamily (see Glossary, Schemes 1–7). A minority of genera have species with pinnate leaves, and leaves modified into phyllodes occur in most species of the large, mainly Australian, genus Acacia Mill. and in a few species in other unrelated genera including Senna Mill. and Mimosa L. The leaves themselves, especially the bipinnate leaf, can be extremely large (e.g., Schizolobium Vogel leaves are > 1 m long) to highly reduced; aphyllous, or nearly aphyllous, species occur in some genera [e.g., Acacia, Senna, Chamaecrista (L.) Moench, Neltuma Raf., Prosopidastrum Burkart]. Across all legumes, seismonasty, i.e., leaf movements prompted by touch, is known only within subfamily Caesalpinioideae, in the genera Mimosa and Neptunia (tribe Mimoseae). Extrafloral nectaries (EFNs) are present in the majority of Caesalpinioideae (Scheme 2), are morphologically extremely diverse, and often conspicuous and abundant on the petiole or leaf rachides between pinnae or leaflet pairs, and in a few genera (e.g., Archidendron F. Muell., Macrosamanea Britton & Rose) on floral bracts (
Across the subfamily, inflorescences and flowers are morphologically highly variable. The inflorescences can be racemose, paniculate or in fascicles and the Mimoseae have characteristic capitate or spicate and frequently heteromorphic inflorescences, often with some sterile flowers, some of which develop showy staminodia (Schemes 3, 4). Although usually bisexual, flowers can also be unisexual, and in the Mimoseae inflorescences can include a mixture of both bisexual and unisexual flowers with or without sterile flowers. The flowers are generally pentamerous, but there are many variations [3–6 (8) sepals or petals], and in some species, sepals and/or petals are absent (Ceratonia L.). Flowers are generally radially symmetrical in several Caesalpinioideae tribes, including the Mimoseae, but in other clades the flowers are bilaterally symmetrical or asymmetrical. Although a majority of Caesalpinioideae flowers are bee pollinated, specialised bat, bird, butterfly and moth pollinated flowers are also common (
The subfamily is most diverse in lowland tropical and subtropical regions, only rarely occurring above 2500 m elevation, but a minority of genera have species in warm temperate zones that are not prone to severe frosts across the Americas, Europe, Asia, and Australia. More than half of Caesalpinioideae genera naturally occur in the Americas (104 of 163 genera), of which 84 are endemic. Africa (including Madagascar) has the second highest number of Caesalpinioideae genera, with 59 genera, 29 of which are endemic, followed by Asia (40 genera, 7 endemic), and Australia and the Pacific (27 genera, 6 endemic; See details in Tables
Caesalpinioideae genera richness across global floristic realms and subrealms (according to
Floristic realms/subrealms | Present genera: n (%*) | Endemic genera: n (%**) |
---|---|---|
Neotropical | 101 (61.96%) | 71 (70.30%) |
- Brazilian | 81 (49.69%) | 25 (30.86%) |
- Subtropical American | 76 (46.63%) | 17 (22.37%) |
African | 59 (36.20%) | 30 (50.85%) |
- Madagascan | 26 (15.95%) | 6 (23.08%) |
- Guineo-Congolian | 41 (25.15%) | 14 (34.15%) |
- Sudanio-Zambezian | 30 (18.40%) | 5 (17.24%) |
Indo-Malesian | 40 (24.54% hi) | 11 (27.50%) |
- Malaysian | 32 (19.63%) | 5 (15.63%) |
- Indian | 29 (17.79%) | 4 (14.29%) |
Holarctic | 23 (14.11%) | 0 (0%) |
- North American | 19 (11.66%) | 0 (0%) |
- European | 1 (0.61%) | 0 (0%) |
- Asian | 8 (4.91%) | 0 (0%) |
Australian | 21 (12.88%) | 1 (4.76%) |
Chile-Patagonian | 15 (9.20%) | 1 (6.67%) |
Saharo-Arabian | 7 (4.29%) | 0 (0%) |
- Iran-Pakistan | 4 (2.45%) | 0 (0%) |
- Saharo-Arabian | 5 (3.07%) | 0 (0%) |
Novozealandic | 2 (1.23%) | 0 (0%) |
Novozealandic | 1 (0.61%) | 0 (0%) |
Tasmanian | 1 (0.61%) | 0 (0%) |
Distribution of the genera of Caesalpinioideae across different floristic realms and subrealms.
Genus | Realms | Saharo-Arabian | Holarctic | Chile-Patagonian | Neotropical | Indo-Malesian | African | Australian | Novozealandic | ||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Sub-realms | Iran-Pakistan | Saharo-Arabian | North American | European | Asian | Brazilian | Subtropical American | Malaysian | Indian | Madagascan | Guineo-Congolian | Sudanio-Zambezian | Novozealandic | Tasmanian | |||
Total | 4 | 5 | 19 | 1 | 8 | 15 | 81 | 76 | 32 | 29 | 26 | 41 | 30 | 21 | 1 | 1 | |
% | 2.45 | 3.07 | 11.66 | 0.61 | 4.91 | 9.20 | 49.69 | 46.63 | 19.63 | 17.79 | 15.95 | 25.15 | 18.40 | 12.88 | 0.61 | 0.61 | |
Abarema | ## | ||||||||||||||||
Acacia | + | + | + | + | |||||||||||||
Acaciella | + | + | + | ||||||||||||||
Acrocarpus | # | # | |||||||||||||||
Adenanthera | + | + | + | + | |||||||||||||
Adenopodia | + | + | + | ||||||||||||||
Afrocalliandra | ## | ||||||||||||||||
Alantsilodendron | ## | ||||||||||||||||
Albizia | + | + | + | + | + | + | + | ||||||||||
Amblygonocarpus | # | # | |||||||||||||||
Anadenanthera | # | # | |||||||||||||||
Anonychium | ## | ||||||||||||||||
Arapatiella | ## | ||||||||||||||||
Archidendron | + | + | + | ||||||||||||||
Archidendropsis | ## | ||||||||||||||||
Arcoa | ## | ||||||||||||||||
Arquita | + | + | + | ||||||||||||||
Aubrevillea | ## | ||||||||||||||||
Balsamocarpon | # | ||||||||||||||||
Batesia | ## | ||||||||||||||||
Biancaea | + | + | |||||||||||||||
Blanchetiodendron | ## | ||||||||||||||||
Boliviadendron | ## | ||||||||||||||||
Burkea | # | # | |||||||||||||||
Bussea | # | # | |||||||||||||||
Caesalpinia | # | # | |||||||||||||||
Calliandra | + | + | + | + | |||||||||||||
Calliandropsis | ## | ||||||||||||||||
Calpocalyx | ## | ||||||||||||||||
Campsiandra | ## | ||||||||||||||||
Cassia | + | + | + | + | + | + | + | + | |||||||||
Cedrelinga | ## | ||||||||||||||||
Cenostigma | # | # | |||||||||||||||
Ceratonia | + | + | + | ||||||||||||||
Chamaecrista | + | + | + | + | + | + | + | + | + | + | + | ||||||
Chidlowia | ## | ||||||||||||||||
Chloroleucon | # | # | |||||||||||||||
Cojoba | # | # | |||||||||||||||
Colvillea | ## | ||||||||||||||||
Conzattia | ## | ||||||||||||||||
Cordeauxia | + | ||||||||||||||||
Coulteria | # | # | |||||||||||||||
Cylicodiscus | ## | ||||||||||||||||
Delonix | # | # | # | ||||||||||||||
Denisophytum | + | + | + | + | + | ||||||||||||
Desmanthus | + | + | + | ||||||||||||||
Dichrostachys | + | + | + | + | + | + | |||||||||||
Dimorphandra | ## | ||||||||||||||||
Dinizia | ## | ||||||||||||||||
Diptychandra | ## | ||||||||||||||||
Ebenopsis | ## | ||||||||||||||||
Entada | + | + | + | + | + | + | + | + | + | ||||||||
Enterolobium | # | # | |||||||||||||||
Erythrophleum | + | + | + | + | + | ||||||||||||
Erythrostemon | + | + | + | + | |||||||||||||
Faidherbia | + | + | + | ||||||||||||||
Falcataria | + | + | |||||||||||||||
Fillaeopsis | ## | ||||||||||||||||
Gagnebina | ## | ||||||||||||||||
Gelrebia | # | # | |||||||||||||||
Gleditsia | + | + | + | + | |||||||||||||
Gretheria | ## | ||||||||||||||||
Guilandina | + | + | + | + | + | + | + | + | |||||||||
Gwilymia | ## | ||||||||||||||||
Gymnocladus | + | + | + | ||||||||||||||
Haematoxylum | + | + | + | ||||||||||||||
Havardia | ## | ||||||||||||||||
Heliodendron | ## | ||||||||||||||||
Hererolandia | ## | ||||||||||||||||
Hesperalbizia | ## | ||||||||||||||||
Heteroflorum | ## | ||||||||||||||||
Hoffmannseggia | + | + | + | + | |||||||||||||
Hultholia | ## | ||||||||||||||||
Hydrochorea | + | + | + | ||||||||||||||
Indopiptadenia | ## | ||||||||||||||||
Inga | # | # | |||||||||||||||
Jacqueshuberia | ## | ||||||||||||||||
Jupunba | # | # | |||||||||||||||
Kanaloa | ## | ||||||||||||||||
Lachesiodendron | # | # | |||||||||||||||
Lemurodendron | ## | ||||||||||||||||
Leucaena | + | + | + | ||||||||||||||
Leucochloron | ## | ||||||||||||||||
Libidibia | # | # | |||||||||||||||
Lophocarpinia | ## | ||||||||||||||||
Lysiloma | ## | ||||||||||||||||
Macrosamanea | ## | ||||||||||||||||
Mariosousa | ## | ||||||||||||||||
Marlimorimia | ## | ||||||||||||||||
Melanoxylum | ## | ||||||||||||||||
Mezcala | ## | ||||||||||||||||
Mezoneuron | + | + | + | + | + | + | |||||||||||
Microlobius | # | # | |||||||||||||||
Mimosa | + | + | + | + | + | + | + | ||||||||||
Mimozyganthus | + | + | |||||||||||||||
Moldenhawera | ## | ||||||||||||||||
Mora | # | # | |||||||||||||||
Moullava | + | + | + | ||||||||||||||
Naiadendron | ## | ||||||||||||||||
Neltuma | + | + | + | + | |||||||||||||
Neptunia | + | + | + | + | + | + | + | + | + | ||||||||
Newtonia | # | # | |||||||||||||||
Osodendron | ## | ||||||||||||||||
Pachyelasma | ## | ||||||||||||||||
Painteria | ## | ||||||||||||||||
Parapiptadenia | ## | ||||||||||||||||
Pararchidendron | + | + | |||||||||||||||
Parasenegalia | + | + | + | ||||||||||||||
Paraserianthes | + | + | |||||||||||||||
Parkia | + | + | + | + | + | + | + | ||||||||||
Parkinsonia | + | + | + | + | + | + | |||||||||||
Paubrasilia | ## | ||||||||||||||||
Peltophorum | + | + | + | + | + | + | |||||||||||
Pentaclethra | + | + | + | ||||||||||||||
Piptadenia | # | # | |||||||||||||||
Piptadeniastrum | ## | ||||||||||||||||
Piptadeniopsis | # | # | |||||||||||||||
Pithecellobium | # | # | |||||||||||||||
Pityrocarpa | # | # | |||||||||||||||
Plathymenia | ## | ||||||||||||||||
Pomaria | + | + | + | ||||||||||||||
Prosopidastrum | ## | ||||||||||||||||
Prosopis | + | + | + | ||||||||||||||
Pseudalbizzia | # | # | |||||||||||||||
Pseudoprosopis | ## | ||||||||||||||||
Pseudosamanea | # | # | |||||||||||||||
Pseudosenegalia | ## | ||||||||||||||||
Pterogyne | # | # | |||||||||||||||
Pterolobium | + | + | + | + | + | ||||||||||||
Punjuba | # | # | |||||||||||||||
Recordoxylon | ## | ||||||||||||||||
Ricoa | ## | ||||||||||||||||
Robrichia | # | # | |||||||||||||||
Samanea | # | # | |||||||||||||||
Sanjappa | ## | ||||||||||||||||
Schizolobium | # | # | |||||||||||||||
Schleinitzia | ## | ||||||||||||||||
Senegalia | + | + | + | + | + | + | + | + | + | + | + | + | |||||
Senna | + | + | + | + | + | + | + | + | + | + | + | + | |||||
Serianthes | # | # | |||||||||||||||
Sphinga | # | # | |||||||||||||||
Stachyothyrsus | ## | ||||||||||||||||
Stenodrepanum | ## | ||||||||||||||||
Strombocarpa | + | + | + | + | |||||||||||||
Stryphnodendron | # | # | |||||||||||||||
Stuhlmannia | # | # | |||||||||||||||
Sympetalandra | ## | ||||||||||||||||
Tachigali | # | # | |||||||||||||||
Tara | # | # | |||||||||||||||
Tetrapleura | ## | ||||||||||||||||
Tetrapterocarpon | ## | ||||||||||||||||
Thailentadopsis | ## | ||||||||||||||||
Ticanto | + | + | + | + | |||||||||||||
Umtiza | ## | ||||||||||||||||
Vachellia | + | + | + | + | + | + | + | + | + | + | + | ||||||
Viguieranthus | ## | ||||||||||||||||
Vouacapoua | ## | ||||||||||||||||
Wallaceodendron | ## | ||||||||||||||||
Xerocladia | ## | ||||||||||||||||
Xylia | + | + | + | ||||||||||||||
Zapoteca | # | # | |||||||||||||||
Zuccagnia | ## | ||||||||||||||||
Zygia | # | # |
The Neotropical floristic realm (sensu
Nine genera have a pantropical distribution, occurring in all tropical floristic realms [Cassia L., Chamaecrista, Entada Adans., Guilandina L., Neptunia, Peltophorum (Vogel) Benth., Senegalia Raf., Senna, Vachellia; Table
The genera in Caesalpinioideae are geographically arranged across two major continental blocks. The American block, including the entire Neotropical realm, the Chile-Patagonian realm, and the southern part of the Holarctic realm (North American subrealm), has 83 genera restricted to it, with eight of these genera distributed across the Neotropical and Chile-Patagonian realms, some slightly extending to the southern portion of the North American subrealm (Acaciella Britton & Rose, Calliandra Benth., Desmanthus, Erythrostemon, Hoffmannseggia Cav., Mimozyganthus Burkart, Neltuma, and Strombocarpa). The second major block includes the African, Indo-Malesian, and Australian realms, with 52 genera restricted to this block and with Adenanthera L., Dichrostachys, Erythrophleum Afzel. ex R. Br., and Mezoneuron Desf. occurring in all three of these realms. Acacia weakly extends into the Madagascan (La Réunion and Mauritius) and Tasmanian subrealms.
Species richness, determined from occurrence records, indicate south-central Mexico and Central America, central-eastern Brazil, and southwestern Australia to be the most diverse regions (Fig.
The wide geographical distribution of Caesalpinioideae is matched by an equally wide ecological amplitude across the full precipitation spectrum of the tropics, spanning a 100-fold gradient in mean annual precipitation from arid deserts to seasonally dry tropical forests and savannas, and tropical rainforests (
Stability is one of the most important qualities of any taxonomic classification. It is therefore crucial that the phylogenetic framework used for assigning names to clades is robust and unlikely to change with sampling of additional taxa or genomic regions in the future. Based on the number and identity of the taxa included, the size of the genomic dataset, and the phylogenomic methods used to infer the phylogeny and assess its robustness, the phylogenetic framework employed here is currently the best available for taxonomic classification of Caesalpinioideae. This is confirmed by its overall agreement with previous smaller-scale phylogenies (see details below) and other recent independent phylogenomic studies (
The classification proposed here uses as its framework the most comprehensively sampled phylogenetic analysis of Caesalpinioideae to date (Figs
Phylogeny of Caesalpinioideae showing the tribal classification presented here. The names and phylogenetic placements of all 63 non-Mimoseae Caesalpinioideae genera are shown and known generic non-monophyly is indicated with terminal names of non-monophyletic genus in bold. The most likely placements for four unsampled genera are indicated with dashed lines; see respective treatments for details. Tribe Mimoseae has been collapsed (see Fig.
Phylogeny of tribe Mimoseae showing the clade-based classification of the tribe with two named higher-level and 17 named lower-level clades. The names and phylogenetic placements of all 100 Mimoseae genera are shown, and known generic non-monophyly is indicated with terminal names of non-monophyletic genera in bold. The most likely placement of the unsampled genus Microlobius is indicated with a dashed line; see Stryphnodendron clade treatment (page 319) for details. Branch lengths are expressed in coalescent units, and terminal branch lengths have been assigned an arbitrary uniform length for visual clarity. Monophyletic genera are represented by single branches; see Suppl. material
The phylogenomic analysis presented here includes 420 Caesalpinioideae species representing all but five of the 163 genera. The five missing genera are: Vouacapoua Aubl., which has three species and is likely a member of tribe Cassieae (e.g.,
A common feature of phylogenomic analyses employing large numbers of genes is the presence of conflict among gene trees, i.e., phylogenies based on individual genes (
The reference phylogeny used here as the basis for the new classification was inferred using ASTRAL (
Under the
Although Mimoseae have traditionally been diagnosed by a series of diagnostic features, notably valvate petal aestivation and flowers with a reduced perianth and showy androecium, mostly clustered in compact inflorescences, the morphological distinctions between the mimosoid clade and some genera of the subtending grade of caesalpinioid lineages are not always clear-cut. For example, Dinizia, once considered to be in Mimosoideae, is placed outside the mimosoid clade in molecular phylogenetic and phylogenomic analyses (
The new classification proposed here thus follows a traditional Linnean approach but is complemented by a clade-based classification of the large tribe Mimoseae. Rank-free naming of clades within subfamilies and tribes has been prevalent in the legume literature, with many examples of clade names that have become widely used and accepted, such as the dalbergioid clade (
The new classification of subfamily Caesalpinioideae comprises eleven tribes, which are either new, reinstated or re-circumscribed at this rank: Caesalpinieae Rchb., Cassieae Bronn, Campsiandreae LPWG, Ceratonieae Rchb., Dimorphandreae Benth., Erythrophleeae LPWG, Gleditsieae Nakai, Mimoseae Bronn, Pterogyneae LPWG, Schizolobieae Nakai, and Sclerolobieae Benth. & Hook. f. (Fig.
Comparison of the new phylogeny-based classification for Caesalpinioideae with classifications for these genera published in Advances in Legume Systematics, Part 1 (
Genera |
|
|
New classification |
---|---|---|---|
Arcoa Urb. | Dimorphandra group (Caesalpinieae) | Umtiza clade (Caesalpinieae) | Ceratonieae |
Tetrapterocarpon Humbert | |||
Acrocarpus Wight ex Arn. | Acrocarpus group (Caesalpinieae) | ||
Ceratonia L. | Ceratoniinae (Cassieae) | ||
Umtiza Sim | Detarieae | Gleditseae | |
Gleditsia J. Clayton | Gleditsia group (Caesalpinieae) | ||
Gymnocladus Lam. | |||
Pterogyne Tul. | Pterogyne group (Caesalpinieae) | Pterogyne group (Caesalpinieae) | Pterogyneae |
Batesia Spruce ex Benth. | Peltophorum group (Caesalpinieae) | Batesia group (Caesalpinieae) | Cassieae |
Melanoxylum Schott | |||
Recordoxylon Ducke | |||
Vouacapoua Aubl. | Unknown position | ||
Chamaecrista (L.) Moench | Cassiinae (Cassieae) | Cassiinae (Cassieae) | |
Cassia L. | |||
Senna Mill. | |||
Stuhlmannia Taub. | Caesalpinia group (Caesalpinieae) | Caesalpinia group (Caesalpinieae) | Caesalpinieae |
Cordeauxia Hemsl. | |||
Cenostigma Tul. | |||
Libidibia (DC.) Schltdl. | |||
Balsamocarpon Clos | |||
Zuccagnia Cav. | |||
Hoffmannseggia Cav. | |||
Stenodrepanum Harms | |||
Erythrostemon Klotzsch | |||
Pomaria Cav. | |||
Haematoxylum L. | |||
Lophocarpinia Burkart | |||
Caesalpinia L. | |||
Coulteria Kunth | |||
Tara Molina | |||
Guilandina L. | |||
Moullava Adans. | |||
Pterolobium R. Br. ex Wight & Arn. | |||
Mezoneuron Desf. | |||
Arquita Gagnon, G. P. Lewis & C. E. Hughes | New in 2015 | ||
Hererolandia Gagnon & G. P. Lewis | New in 2016 | ||
Denisophytum R. Vig. | Reinstated in 2016 | ||
Paubrasilia Gagnon, H. C. Lima & G. P. Lewis | New in 2016 | ||
Gelrebia Gagnon & G. P. Lewis | New in 2016 | ||
Hultholia Gagnon & G. P. Lewis | New in 2016 | ||
Biancaea Tod. | Reinstated in 2016 | ||
Ticanto Adans. | Reinstated in 2022 | ||
Schizolobium Vogel | Peltophorum group (Caesalpinieae) | Core-Peltophorum group (Caesalpinieae) | Schizolobieae |
Bussea Harms | |||
Peltophorum (Vogel) Benth. | |||
Parkinsonia L. | Caesalpinia group (Caesalpinieae) | ||
Conzattia Rose | |||
Heteroflorum M. Sousa | New in 2005 | ||
Colvillea Bojer ex Hook. | Peltophorum group (Caesalpinieae) | ||
Delonix Raf. | |||
Moldenhawera Schrad. | Moldenhawera group (Caesalp.) | Sclerolobieae | |
Arapatiella Rizzini & A. Mattos | Tachigali group (Caesalpinieae) | ||
Jacqueshuberia Ducke | |||
Tachigali Aubl. | Sclerolobium group (Caesalpinieae) | ||
Diptychandra Tul. | Unknown position | ||
Dimorphandra Schott | Dimorphandra group (Caesalpinieae) | Dimorphandra group (Caesalpinieae) | Dimorphandreae |
Mora Benth. | |||
Stachyothyrsus Harms | |||
Burkea Benth. | |||
Campsiandra Benth. | Peltophorum group (Caesalpinieae) | Unknown position | Campsiandreae |
Dinizia Ducke | Subf. Mimosoideae | Subf. Mimosoideae | |
Pachyelasma Harms | Dimorphandra group (Caesalpinieae) | Dimorphandra group (Caesalpinieae) | Erythrophleeae |
Erythrophleum Afzel. ex R. Br. | |||
Sympetalandra Stapf | Mimoseae | ||
Chidlowia Hoyle | Unknown position | ||
Other Mimoseae genera | Subf. Mimosoideae | Subf. Mimosoideae |
Caesalpinioideae as defined here includes elements from three previously recognised major groups: part of old sense tribe Caesalpinieae, part of old sense tribe Cassieae, and the nested subfamily Mimosoideae. This broad clade has been referred to as the Mimosoideae-Caesalpinieae-Cassieae or MCC clade (
In their treatment of tribe Caesalpinieae in Advances in Legume Systematics Part 1,
Below we provide a brief morphological description, overview of previous classification, and history of the phylogenetic understanding of each of the tribes proposed here. Additional details are given in the taxonomic accounts for each of these groups.
Tribe Ceratonieae comprises six species in four genera. The four genera had previously been placed in distinct generic groups and tribes: Acrocarpus Wight ex Arn. in its own generic group of Caesalpinieae by
These recent molecular analyses have highlighted several previously unsuspected morphological synapomorphies for Ceratonieae, the most striking of which is a bipinnate leaf (although mostly once pinnate in Ceratonia) terminating in a triad of pinnae arising from the same point at the apex of the rachis (
The simplest of the informal generic groups recognised by
The 20 species of the three genera of tribe Gleditsieae occur in warm temperate regions, with several disjunctions between North and South America (Gleditsia), South Africa (Umtiza), and North America and Asia (Gymnocladus). Tribe Gleditsieae is characterised by several vegetative and floral synapomorphies, such as a tubular hypanthium and sepals with trichomes on the inner surface (
Pterogyneae is here recognised as a new tribe comprising just the single species Pterogyne nitens Tul. Although not included in the phylogenomic analyses of
In Advances in Legume Systematics Part 1,
Tribe Cassieae is the largest non-mimosoid clade (in terms of species richness) in subfamily Caesalpinioideae, with 695 species, the vast majority of which are found in the genera Chamaecrista (361 species) and Senna (287 species) (Tribe Cassieae, page 83). Although broadly distributed across the tropics, most of the genera and species are found in the New World. The clade is characterised by singly pinnate or bifoliolate leaves and, in several genera, stomata on both sides of the leaflets (
The Caesalpinia group defined by
Tribe Caesalpinieae comprises ca. 223 species in 27 genera. Although two of these genera, Stenodrepanum and Hultholia, were not sampled by
Species of Caesalpinieae are highly diverse in growth forms, defence mechanisms, fruit morphologies, and pollination and seed dispersal syndromes (
Tribe Schizolobieae as here circumscribed matches the core Peltophorum group (i.e., Peltophorum group s.s.) first recovered phylogenetically by
Tribe Schizolobieae contains ca. 42 species in eight genera. It has a pantropical distribution, and a considerable portion of the clade (i.e., the Parkinsonia – Delonix subclade; Fig.
The generic composition of tribe Sclerolobieae as treated here has not been recovered previously, although its constituent genera have often been associated with each other based on morphological and molecular data. The five genera of the Sclerolobieae were placed in two generic groups of tribe Caesalpinieae by
As defined here, tribe Sclerolobieae is restricted to the Neotropics, and comprises ca. 113 species, most of which are in the genus Tachigali (80–90 species; Tribe Sclerolobieae, page 165). Several species of Tachigali are known to form close co-evolutionary associations with ants (
The four genera of the Dimorphandreae, Dimorphandra, Mora Benth., Stachyothyrsus Harms, and Burkea Benth., form a clade in most molecular phylogenetic studies (
The four genera of the Dimorphandreae contain 35 species. However, 26 are in Dimorphandra s.l., which is paraphyletic (e.g.,
It is also notable that while tribes Sclerolobieae, Schizolobieae and Dimorphandreae are each supported as monophyletic in recent phylogenomic analyses, the relationships among these three lineages are weakly supported and characterised by high gene tree conflict (Suppl. material
Dinizia and Campsiandra, previously placed in Mimosoideae and the Peltophorum group of Caesalpinieae respectively (Table
Tribe Campsiandreae comprises 5 to 22 species (the genus Campsiandra needs to be revised because several species are of dubious taxonomic status), only two of which are in Dinizia. The tribe is restricted to tropical rainforests in South America. Nodulation is reported in one species each of Dinizia and Campsiandra (
Erythrophleum and Pachyelasma have rarely been recovered as sister genera before (
The Erythrophleeae, with 12 species in Erythrophleum and one species in Pachyelasma, is restricted to the Old World tropics (Africa, Asia and Australia). Erythrophleum is reported to nodulate.
Recognising the mimosoid clade as the newly circumscribed tribe Mimoseae results in by far the largest tribe in subfamily Caesalpinioideae in terms of numbers of species (ca. 3500) and genera (100). Previous tribal classifications of the mimosoid clade (i.e., former subfamily Mimosoideae) recognised five tribes: Mimoseae, Acacieae Benth., Ingeae Benth., Parkieae (Wight & Arn.) Benth., and Mimozygantheae Burkart (
There has been ongoing debate about which genera are included in the mimosoid clade (
Tribe Mimoseae is diagnosed by valvate petal aestivation (with exceptions in Chidlowia and Sympetalandra), bipinnate leaves (except Inga and a few scattered species in other genera), flowers that are relatively small with reduced perianth and showy androecium, mostly clustered in compact inflorescences that are commonly capitate or spicate (Tribe Mimoseae, page 201), and the presence of symbiosome-type (as opposed to fixation-thread-type) root nodules (
Tribe Mimoseae, as circumscribed here, is robustly supported as monophyletic and is subtended by a relatively long branch (Fig.
The core mimosoid clade as delimited by
The ingoid clade, delimited by
Based on the phylogeny of subfamily Caesalpinioideae presented here, we recognise eleven tribes (Fig.
We provide a key to genera, as well as taxonomic descriptions and notes for tribes, named clades, and all 163 genera, and illustrate the diversity of growth forms, foliage, flowers and fruits for nearly all genera. We also provide a distribution map of the native range for each genus, based on quality-controlled herbarium specimen localities and floristic surveys. The occurrence data for Mimoseae are from
The Legume Phylogeny Working Group (LPWG)*
* Luciano Paganucci de Queiroz2, Anne Bruneau1 , Jens J. Ringelberg3,4, Leonardo M. Borges5, Roseli Lopes da Costa Bortoluzzi6, Gillian K. Brown7, Domingos B. O. S. Cardoso8,9, Ruth P. Clark10, Adilva de Souza Conceição11, Matheus Martins Teixeira Cota2, Else Demeulenaere12, Rodrigo Duno de Stefano13, John E. Ebinger14, Julia Ferm15, Andrés Fonseca-Cortés2, Edeline Gagnon16,17,18, Rosaura Grether19, Ethiéne Guerra20, Elspeth Haston18, Patrick S. Herendeen21, Héctor M. Hernández22, Helen C. F. Hopkins10, Isau Huamantupa-Chuquimaco23, Colin E. Hughes3, Stefanie M. Ickert-Bond24, João Iganci20,25, Erik J. M. Koenen26, Gwilym P. Lewis10, Haroldo Cavalcante de Lima8,27, Alexandre Gibau de Lima8,28, Melissa Luckow29, Brigitte Marazzi30, Bruce R. Maslin31,32, Matías Morales33,34, Marli Pires Morim8, Daniel J. Murphy35, Shawn A. O’Donnell36, Filipe Gomes Oliveira2, Ana Carla da Silva Oliveira2, Juliana Gastaldello Rando37, Pétala Gomes Ribeiro2, Carolina Lima Ribeiro2, Felipe da Silva Santos2, David S. Seigler38, Guilherme Sousa da Silva39, Marcelo F. Simon40, Marcos Vinícius Batista Soares20, Vanessa Terra41
Citation: Legume Phylogeny Working Group, Queiroz LP et al. (2024) 1. Subfamily Caesalpinioideae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 32–54. https://doi.org/10.3897/phytokeys.240.101716
Caesalpiniaceae R. Br., in M. Flinders, Voy. Terra Austral. 2: 551. 1814. Type: Caesalpinia L.
Caesalpinia L.
Trees, shrubs, lianas, suffruticose or functionally herbaceous, occasionally aquatic, either unarmed or commonly armed with prickles, spines, or thorns; specialised extrafloral nectaries often present on the petiole and/or on the primary and secondary leaf rachides, usually between pinnae or leaflet pairs, more rarely stipular or bracteal. Stipules in lateral position and free or absent, usually entire, less frequently divided or spinescent. Leaves usually pulvinate, bipinnate, otherwise pinnate (sometimes both types on the same plant) and then mostly paripinnate, rarely imparipinnate, less often bifoliolate, modified into phyllodes or lacking, arrangement of the pinnae and leaflets mostly opposite, rarely alternate; stipels rare and not to be confused with the more commonly present paraphyllidia. Inflorescences globose or ellipsoid capitula, spicate, paniculate, racemose or in fascicles; bracteoles commonly small or absent. Flowers usually bisexual, rarely unisexual (species dioecious or monoecious), or bisexual flowers combined with unisexual and/or sterile flowers in heteromorphic inflorescences (Mimoseae), radially, less frequently bilaterally symmetrical, or asymmetrical; hypanthium lacking or cupular, rarely tubular; sepals (3) 5 (6–8), free or fused; petals (3) 5 (6–8), free or fused, the sepals or petals or both sometimes lacking, aestivation valvate (Mimoseae) or imbricate and then the adaxial petal innermost; stamens commonly diplostemonous or haplostemonous, sometimes reduced to 3 or 4 (in some Mimosa species), frequently polystemonous (to 100+ in some Mimoseae), free or fused, sometimes heteromorphic, some or all sometimes modified or staminodial, anthers basifixed or dorsifixed, often with a stipitate or sessile apical gland, dehiscing via longitudinal slits or apical or basal poricidal slits or pores; pollen in tricolporate monads, or commonly in tetrads, bitetrads or polyads (most Mimoseae); gynoecium uni- or rarely polycarpellate, 1–many-ovulate. Fruit typically dry and dehiscent, either a legume (dehiscent along both sutures) or a follicle (dehiscent along the adaxial suture only), or dry and segmented into one-seeded articles, either without a persistent margin (a lomentum) or with a persistent margins forming a replum like a frame (a craspedium), sometimes indehiscent and somewhat fleshy (an indehiscent legume), or dry and winged (a samara), when dehiscent with papery, leathery, or woody valves, dehiscence passive (inert valves), elastic, or explosive (the valves becoming curved, spirally coiled, or arched backwards), less frequently with entire valves, but with the endocarp segmented into one-seeded articles (a cryptolomentum). Seeds usually with an open (U-shaped) or closed (O-shaped) pleurogram on both faces or lacking a pleurogram, sometimes with a fleshy aril or sarcotesta, sometimes winged, in cross section terete (with a 1:1 ratio), compressed (with more or less 2:1 ratio) or flattened (with > 4:1 ratio;
2n mostly 24, 26, 28, but also reported 2n = 14, 16, 20, 22, 36, 52, 54, 56, 72, 78, 104, 112.
Caesalpinioideae in its emended circumscription contains eleven tribes, 163 genera and ca. 4680 species. Tribes: Caesalpinieae Rchb. (27 genera / ca. 223 species), Campsiandreae LPWG (2 / 5–22), Cassieae Bronn (7 / 695), Ceratonieae Rchb. (4 / 6), Dimorphandreae Benth. (4 / 35), Erythrophleeae LPWG (2 / 13), Gleditsieae Nakai (3 / 20), Mimoseae Bronn (100 / ca. 3510), Pterogyneae LPWG (1 / 1), Schizolobieae Nakai (8 / 42–43), Sclerolobieae Benth. & Hook. f. (5 / ca. 113) (Fig.
The most inclusive crown clade containing Arcoa gonavensis Urb. and Mimosa pudica L., but not Bobgunnia fistuloides (Harms) J.H. Kirkbr. & Wiersema, Duparquetia orchidacea Baill., or Poeppigia procera C. Presl.
Pantropical, common in both wet and dry regions, with a handful of species extending to the temperate zone, less frequently frost-tolerant (Gleditsia J. Clayton, Gymnocladus Lam. and some species of Acacia Mill., Desmanthus Willd. and Senna Mill.). Caesalpinioideae species are infrequent above 2500 m in the tropics and are largely absent from mid- and high-elevation tropical montane forests. Generic diversity is highest in the Neotropics, and there are important centres of high species diversity in Mexico and Central America, central-east South America, Africa, Madagascar, parts of South East Asia and Australia (Fig.
This clade was referred to as the Mimosoideae-Caesalpinieae-Cassieae (MCC clade) (
In the following treatments, the type species of different taxa are presented. Their homotypic (nomenclatural) synonyms are indicated by the identity symbol (≡) and heterotypic (taxonomic) synonyms by the equality symbol (=), as specified in the International Code of Nomenclature for algae, fungi, and plants (
Some sections of the key, for particular groups, were developed and modified from previous publications [e.g.,
1 | Plants aphyllous or leaves reduced to phyllodes of petiolar or leaf rachis origin, lacking leaflets | 2 |
– | Plants with developed leaves | 7 |
2 | Petals imbricate in bud, the uppermost petal in the inner position; some or all anthers dehiscing by pores; flowers solitary or in racemes | 3 |
– | Petals valvate in bud; all anthers dehiscing through longitudinal slits; flowers densely clustered in capitate or spicate inflorescences | 4 |
3 | Erect herbs or subshrubs; stipules cordiform clasping the stem internodes; flowers isolated in stipule axils; pedicel with a pair of alternate bracteoles; fruits elastically dehiscent, the valves twisting upon dehiscence | * Chamaecrista |
– | Shrubs with hard woody branches; stipules small, not covering the stems; flowers in short corymbiform racemes; pedicels lacking bracteoles; fruits indehiscent or passively dehiscent, the valves not becoming twisted | * Senna |
4 | Flowers polystemonous; seeds arilate | * Acacia |
– | Flowers diplostemonous; seeds not arilate | 5 |
5 | Aphyllous trees or shrubs due to precocious leaf falling, profusely armed with spine-tipped, rigid, straight, cylindrical thorns; fruits indehiscent, compressed-turgid, pulp mealy or spongy | * Neltuma |
– | Aphyllous or phyllodial subshrubs or shrubs, either unarmed or armed with stipular spines or prickles; fruit a craspedium, a lomentum or dehiscent along the ventral suture | 6 |
6 | Subshrubs or shrubs, unarmed or armed with scattered cauline and phyllodial recurved prickles; stems not striate; leaves modified into phyllodes, not caducous | * Mimosa |
– | Small shrubs armed with stipular spines, the shoots also often spinescent, tapered to a hard point; stems green, photosynthetic, striate with longitudinal, golden corky ridges; leaves not modified in phyllodia, early caducous | * Prosopidastrum |
7 | All or some leaves bipinnate | 8 |
– | Leaves once pinnate or bifoliolate | 179 (page 51) |
8 | Petals valvate in bud; seeds often with a U-shaped or O-shaped pleurogram | 9 |
– | Petals imbricate in bud, the uppermost petal in the inner position; seeds lacking a pleurogram | 126 (page 46) |
9 | Flowers haplostemonous or diplostemonous, i.e., fertile stamens the same or exactly twice the number of petals | 10 |
– | Flowers polystemonous, rarely less than twice but always with more than the number of petals | 64 (page 40) |
10 | Flowers in globose or ellipsoid capitula | 11 |
– | Flowers in elongated spikes or spiciform racemes | 29 |
11 | Capitula with clearly marked floral differentiation, the basal flowers often but not always with elongated showy staminodes | 12 |
– | Capitula without marked floral differentiation (flowers mostly hermaphrodite or staminate) and the basal flowers always without elongated showy staminodes | 15 |
12 | Trees; calyx with imbricate lobes; capitula either with sterile staminodial flowers at base, fertile flowers at apex and modified staminate, nectar-secreting flowers between, or capitula with fertile flowers at base and modified nectar-secreting flowers at apex | * Parkia |
– | Subshrubs, small shrubs, sometimes aquatic herbs; calyx with valvate sepals, capitula with sterile staminoidal flowers at base and fertile hermaphrodite flowers at apex | 13 |
13 | Fruits stipitate, oblong, seeds transversely oriented; stipules ovate to lanceolate and striately veined | Neptunia |
– | Fruits subsessile, linear, seeds obliquely or longitudinally oriented; stipules subulate or setiform with an auriculate base | 14 |
14 | Fruits sub-cylindrical, tardily dehiscent along both sutures from the apex, valves lignified; anther glands present; these stipitate, terminal, or claviform (orbicular on a filiform stalk); pollen aggregated into tetrahedral tetrads | * Mezcala |
– | Fruits dorsi-ventrally flattened, passively dehiscent, valves chartaceous or coriaceous; anther glands absent; pollen dispersed in monads | Desmanthus |
15 | Fruits breaking up into 1-seeded articles, leaving persistent margins or not, or the valves falling entire but leaving a frame made by persistent margins | 16 |
– | Fruits indehiscent or dehiscent through one or both margins, not articulated | 18 |
16 | Fruit a craspedium, i.e., with persistent margins (like a frame) after articles dispersed or the entire valves have fallen; unarmed or branches armed with prickles (fruit rarely a lomentum, i.e. lacking persistent margins, but then the plant is always armed with prickles) | Mimosa |
– | Fruits lomentaceous, the articles falling but the fruit lacking persistent margins; plants armed with stipular spines, sometimes additionally with branches modified into thorns, but lacking prickles | 17 |
17 | Fruits stipitate with margins undulate, not thickened; seeds compressed, discoid; stipular spines ± straight; stems slightly ribbed or not and not ending in a sharp point; Paraguay and southern Bolivia | Piptadeniopsis |
– | Fruits subsessile with straight and thickened margins; seeds ± bulky; stipular spines ± curved; shoots spinescent, ribbed and green, ending in a sharp point; Mexico (Baja California) and Patagonian Argentina | * Prosopidastrum |
18 | Plants armed with stipular spines; fruits indehiscent | 19 |
– | Plants unarmed; fruits either dehiscent or indehiscent | 21 |
19 | Fruits woody, more or less tightly spirally coiled, irregularly and openly coiled or at least somewhat curved, always > 2-seeded; arid and semi-arid areas of North America (southern USA and northern Mexico) and South America (Peru to Bolivia, Argentina and Chile) | * Strombocarpa |
– | Fruits with papery valves, falcate ovate, not coiled or spiralled, 1 (2)-seeded | 20 |
20 | Calyx lobes imbricate in bud; stems not ribbed; leaves and almost sessile capitate inflorescences attached to brachyblasts arising between the straight spines; fruits narrowly winged along the upper suture; Argentina, Bolivia and Paraguay | Mimozyganthus |
– | Calyx lobes valvate in bud; stems ribbed and green; capitula pedunculate, axillary; brachyblasts absent; spines curved; fruits with a broad arched wing along the lower suture; Namaqualand in South Africa and Namibia | Xerocladia |
21 | Leaves with exactly 1 pair of pinnae and 3 leaflets per pinna; capitula pedunculate and solitary or fasciculate on axillary brachyblasts; fruits obovate, 1-seeded, small (to 3.2 cm long), dehiscent through both margins; seeds compressed, cordiform; Kaho’olawe island (Hawaii) | Kanaloa |
– | Leaves with more than one pair of pinnae or > 3 leaflets or both; brachyblasts present or absent; fruits larger (over 5 cm long) and mostly linear to oblong, dehiscent or indehiscent; seeds not cordiform | 22 |
22 | Plants from Africa, Asia and Malesia | 23 |
– | Plants from America | 25 |
23 | Fruit valves coriaceous, narrowly winged along the margins, splitting at edges but not separating over seed-chambers; floral bracts peltate, persistent; Malesia and Papuasia | Schleinitzia |
– | Fruit valves ligneous or stiffly coriaceous, elastically dehiscent from the apex through both sutures; floral bracts caducous; Africa (including Madagascar) | 24 |
24 | Leaves emerging from lateral brachyblasts; stipules joined at base and to the base of petiole, persistent; Madagascar | Alantsilodendron |
– | Leaves on branches lacking brachyblasts; stipules free and caducous; mostly continental Africa (2 species in northern Madagascar) | Xylia |
25 | Fruits held erect above foliage and elastically dehiscent from the apex along both sutures; brachyblasts present sheathed in persistent stipules; dry vegetation of central Mexico | 26 |
– | Fruits not erect, indehiscent or passively dehiscent along one or both margins; widely distributed from southern USA to South America | 27 |
26 | Shrubs to 1 m tall, profusely branched from the base; leaves and inflorescences emerging from lateral brachyblasts; stamens 5; fruits oblanceolate, plano-compressed, with thickened margins | Calliandropsis |
– | Multi-stemmed treelet or large shrub to 3.5 m; inflorescences arising from leaf axils; stamens 10; fruits linear, terete or sub-terete, the margins not noticeably thickened | * Mezcala |
27 | Fruit woody, dehiscent through one margin (a follicle), with the margins constricted between the seeds; seeds flat and orbicular with a narrowly winged margin; widespread deciduous trees in South America (naturalised in the West Indies) | Anadenanthera |
– | Fruit with straight margins and chartaceous, coriaceous or woody valves, dehiscent through one or both margins, if woody and dehiscent from one margin then with bulky seeds; seeds flat or bulky but never orbicular with a winged margin | 28 |
28 | Calyx with imbricate lobes in bud; anthers with an apical gland and glabrous; fruit indehiscent or passively dehiscent with woody valves; seeds compressed or not; evergreen trees from Amazonia | * Parkia |
– | Calyx with valvate lobes in bud; anthers eglandular, sometimes the connective with a small rounded or hooded apiculum, usually pilose, occasionally glabrous; fruit with chartaceous or coriaceous valves, inertly dehiscent along one or usually both sutures; seeds flat-compressed; deciduous shrubs or small trees from North and Central America, one species in dry vegetation of northern and north-western South America (L. leucocephala a pantropical weed) | Leucaena |
29 | Fruits breaking into 1-seeded articles resulting from fragmentation of the entire fruit wall or from just the endocarp | 30 |
– | Fruits not breaking into 1-seeded articles | 35 |
30 | Flowers with the pedicels articulated near the middle, persisting after the flowers fall; leaflets alternate; fruits dehiscing through both margins in two entire valves but the endocarp splitting into 1-seeded papery envelopes; unarmed trees | Plathymenia |
– | Flowers sessile or the pedicel not articulated; leaflets mostly opposite; fruits with the margins framing 1-seeded articles or the entire valves which break away from the persistent margin (craspedium); trees, shrubs, geoxylic subshrubs or lianas, unarmed or armed with prickles or stipular spines | 31 |
31 | Leaves lacking extrafloral nectaries on petiole and leaf rachis | 32 |
– | Leaves with extrafloral nectaries on petiole and/or leaf rachis or on shoot immediately beneath the base of stipules | 33 |
32 | Anther glands absent; plants unarmed or armed with infranodal or internodal prickles; flowers in capitula or spikes; epicarp remaining attached to endocarp on mature fruits | * Mimosa |
– | Anther glands usually present; plants unarmed, rarely (Entada spinescens) with stipular spines; flowers in spikes; epicarp separating from the endocarp on mature fruits | * Entada |
33 | Plants unarmed; epicarp separating from the endocarp on mature fruits; extrafloral nectaries mostly on the shoot immediately beneath the base of stipules, if present on petiole or leaf rachis then plant from Madagascar | * Entada |
– | Plants unarmed or armed with prickles; epicarp remaining attached to endocarp on mature fruits; extrafloral nectaries on the petiole and/or leaf rachis; plants from tropical America and continental sub-Saharan Africa | 34 |
34 | Habit variable; anther glands absent; flowers haplostemonous (in Mimosa myriadenia) or diplostemonous, pentamerous, or tetramerous, in capitula or spikes; tropical America | * Mimosa |
– | Lianas, rarely (in African Adenopodia rotundifolia) shrubs or treelets; anther glands present; flowers diplostemonous, pentamerous, in spikes; distributed in Mexico and Central America and disjunctly across sub-Saharan Africa | * Adenopodia |
35 | Plants from the Americas | 36 |
– | Plants from Africa, Asia, Malesia and Australia | 48 |
36 | Plants armed with stipular spines, solitary or paired thorns, spinescent shoots or prickles | 37 |
– | Plants unarmed | 40 |
37 | Fruits dehiscing through both margins, with papery valves | 38 |
– | Fruits indehiscent, woody or pulpy, rarely with a thin mesocarp, the endocarp fragmented into 1-seeded chambers | 39 |
38 | Branches armed with internodal or infranodal prickles; flowers with a mostly campanulate corolla | Piptadenia |
– | Branches armed with paired recurved stipular spines; flowers with a slender tubular corolla with very short teeth | Lachesiodendron |
39 | Plants armed with stipular spines | * Strombocarpa |
– | Plants armed with axillary, uninodal, solitary or paired thorns or spinescent shoots | * Neltuma |
40 | Flowers with 5 fertile stamens and 5–15 linear staminodes; fruits sickle-shaped and explosively dehiscing along both sutures, the robust woody valves twisting and becoming arched backwards | * Pentaclethra |
– | Flowers with 10 fertile stamens, staminodes absent; fruits never sickle-shaped, indehiscent or dehiscent through one or both margins, if dehiscent then passively so and the valves papery | 41 |
41 | Branches and leaves lacking ferruginous granular trichomes | 42 |
– | Young branches and leaves usually covered with ferruginous granular trichomes | 45 |
42 | Fruits compressed turgid, indehiscent, with the mesocarp thick and pulpy or spongy and the endocarp segmented in one seeded chambers | * Neltuma |
– | Fruits compressed, dehiscent through one or both margins, with a thin mesocarp and an indistinct endocarp | 43 |
43 | Fruit a legume, dehiscing along both margins; flowers mostly with reddish petals and stamens | Parapiptadenia |
– | Fruit a follicle, dehiscing along one margin only; flowers with greenish petals and whitish stamens | 44 |
44 | Extrafloral nectary between or just below the first pair of pinnae; spikes mostly solitary in axils of coevally developing leaves; fruits moniliform, with deeply constricted margins and thick coriaceous and pubescent valves | Pityrocarpa |
– | Extrafloral nectary between the base and the middle of the petiole; spikes mostly clustered in terminal efoliate pseudoracemes or below the coeval leaves; fruits with a linear or oblong body, straight or shallowly sinuous margins and thin to thick woody and glabrous valves | Marlimorimia |
45 | Branches and leaves with strong garlic smell; ferruginous granular trichomes falling early; leaves with 1–3 pairs of pinnae, each pinna comprising a single pair of leaflets, petiolar nectary absent; fruit 4–7 × 1–1.5 cm; seeds white | Microlobius |
– | Branches and leaves without evident garlic smell; ferruginous granular trichomes always present in brach tips; leaves always with more than one pair of pinnae, each pinna comprising 3 or more pairs of leaflets, petiolar nectary present; fruit 8–14 × 2–3.5 cm; seeds black, brown or ochre | 46 |
46 | Leaves with 2–4 (6) pairs of pinnae; leaflets 2.5–16 × 1.5–8 cm; spikes clustered in panicles (except in G. coriacea and G. fissurata) | Gwilymia |
– | Leaves with (3) 5–32 pairs of pinnae; leaflets 0.6–1.2 × 0.3–0.6 cm; spikes grouped in terminal pseudoracemes or in fascicles below the leaves | 47 |
47 | Branches not striate; petiolar nectary 0.5–2 mm long; leaflets alternate, abaxial surface with a tuft of trichomes at the base of the midrib; petals fused at least for ½ of their length; fruit coriaceous or woody and indehiscent or splitting along a single margin (follicle) | Stryphnodendron |
– | Branches strongly striate; petiolar nectary ca. 10 mm long; leaflets opposite, without a tuft of trichomes on the abaxial surface; petals fused only for ¹⁄3 of their length; fruit chartaceous, dehiscent along both margins (legume) | Naiadendron |
48 | Flowers of the base of the spikes sterile and with flat and showy staminodes | 49 |
– | Flowers all fertile (hermaphrodite or staminate), or if sterile, then lacking showy staminodes | 50 |
49 | Short shoots terminated by spines or composed of many persistent fused stipules; fruits with sutural ribs not greatly enlarged, either elastically dehiscent from the apex and coiling after dehiscence with the pericarp coriaceous, or indehiscent and with the pericarp woody | Dichrostachys |
– | Short shoots without terminal spines or fused stipules; fruits with sutural ribs greatly thickened or modified into flattened wings, either elastically dehiscent from the apex and recurved but not coiled after dehiscence, with the pericarp woody, or indehiscent or tardily inertly dehiscent and with the pericarp coriaceous to chartaceous | Gagnebina |
50 | Leaflets clearly alternate | 51 |
– | Leaflets opposite or subopposite | 55 |
51 | Fruit linear, curved or spirally twisted, dehiscing into 2 coriaceous or subcoriaceous valves; seeds bulky with a hard bright red or red and black testa, unwinged; India to tropical South East Asia, Malesia, Australia and Madagascar (1 species widely cultivated) | Adenanthera |
– | Fruit oblong to linear, indehiscent or dehiscent through both margins but then not spirally twisted, valves woody; seeds plano-compressed, winged or not, if bulky, then not red nor red and black; continental Africa | 52 |
52 | Fruit indehiscent, with a longitudinal crest or wing along the valves; flowers in spiciform racemes; seeds unwinged | 53 |
– | Fruit dehiscent through both margins, lacking longitudinal wings; flowers in spikes; seeds winged | 54 |
53 | Anther-glands absent; fruit bluntly tetragonal or sub-cylindrical in cross section; pedicels and calyces glabrous; bracts not apparent | Amblygonocarpus |
– | Anther-glands present, caducous; fruit with a thick longitudinal wing-like projection along each valve, thus cruciform in cross section; pedicels and calyces hairy; flowers subtended by persistent triangular bracts | Tetrapleura |
54 | Leaves lacking extrafloral nectaries; fruits oblong, 10–20 cm wide, dehiscent along both margins, with detaching endocarp; free-standing intrastaminal disk massive, clearly visible in open flower; petals lacking stellate trichomes; stamen filaments pubescent to near the middle | Fillaeopsis |
– | Leaves with small sunken extrafloral nectaries on petiole and/or leaf-rachis; fruits linear-oblong, under 5 cm wide, dehiscing along only the ventral margin, the endocarp not detaching; intrastaminal disk present but not massive; petals stellate-pubescent; stamen filaments glabrous | Cylicodiscus |
55 | Flowers with 5 fertile stamens and 5–15 staminodes | * Pentaclethra |
– | Flowers with all 10 stamens fertile, lacking staminodes | 56 |
56 | Leaves lacking extrafloral nectaries | 57 |
– | Leaves with extrafloral nectary on petiole and/or leaf-rachis | 59 |
57 | Flowers greenish; anther-glands absent; fruit laterally compressed, papyraceous, indehiscent, oblong, wind dispersed, twisted in its proximal end | Aubrevillea |
– | Flowers red or bright yellow; anther-glands present; fruit dehiscent through one or both sutures with a woody or coriaceous pericarp | 58 |
58 | Flowers bright yellow; fruit woody, dehiscing elastically from apex downwards into two recurving valves; seeds unwinged; lianas with tendrilled leaves or small trees | Pseudoprosopis |
– | Flowers red; fruit coriaceous, dehiscing along one margin only; seeds winged; big trees from rainforest canopy | Piptadeniastrum |
59 | Plants armed with internodal prickles | 60 |
– | Plants unarmed | 61 |
60 | Fruit indehiscent, cylindrical or subterete, with a pulpy or fibrous mesocarp; largest leaflets < 1.5 × 1 cm; shoots with scattered prickles | Prosopis |
– | Fruit dehiscent, plano-compressed, coriaceous, lacking a thick mesocarp; largest leaflets > 3 × 3 cm; shoots largely unarmed, occasionally with scattered prickles, mature stems with spine-tipped woody protuberances | Indopiptadenia |
61 | Fruits indehiscent with a thick spongy mesocarp; endocarp fragmented into 1-seeded chambers | Anonychium |
– | Fruits dehiscing down one or both margins, woody, with a thin mesocarp; endocarp not fragmented | 62 |
62 | Seeds unwinged; fruit clavate to dolabriform, explosively dehiscent through both margins | Calpocalyx |
– | Seeds winged; fruit dehiscing through one or both margins but passively so | 63 |
63 | Fruits opening through one margin only; flowers lacking an intrastaminal disk | Newtonia |
– | Fruits opening through both margins; flowers with a distinct intrastaminal disk | Lemurodendron |
64 | Stamens free or joined only at the very base | 65 |
– | Stamens united into a staminal tube | 73 |
65 | Plants armed with stipular spines or prickles on stem and often also on the leaf petiole and rachis (spines or prickles may be absent or rare on individual specimens) | 66 |
– | Plants unarmed | 68 |
66 | Plants armed with infranodal and/or internodal prickles; habit sometimes lianescent; peduncles lacking an obvious involucel (sometimes a small, caducous bract present) | Senegalia |
– | Plants lacking prickles, armed with stipular spines, these sometimes swollen and hollow, and always restricted to the nodal regions; habit never lianescent; peduncles usually with an involucel, rarely lacking (in the monospecific Faidherbia) | 67 |
67 | Peduncles lacking an involucel; stamens united into a tube for ca. 1 mm, anthers eglandular | Faidherbia |
– | Peduncles with involucel (usually medial or basal, rarely at base of inflorescence); stamens free to the base or occasionally shortly fused; anther glands often present | Vachellia |
68 | Funicle expanded into an aril at the point of attachment to the seed; plants native to Australia (but a few species cultivated, and sometimes invasive elsewhere) | * Acacia |
– | Seeds exarillate; plants from the Americas | 69 |
69 | Inflorescence a globose to sub-globose or ellipsoid capitulum (as broad as long or nearly so) | 70 |
– | Inflorescence a cylindrical spike more than twice as long as wide | 71 |
70 | Leaves lacking extrafloral nectaries on petiole and rachis; stamens consistently 200+, the filaments often drying yellow, orange or pink | Acaciella |
– | Leaves with extrafloral nectaries on petiole and/or rachis; stamens fewer than 150, the filaments drying straw-coloured | * Parasenegalia |
71 | Twigs commonly with brachyblasts above most nodes; anther glands absent; endemic to Bolivia | Pseudosenegalia |
– | Twigs lacking brachyblasts (except Mariosousa compacta, Mexico); anther glands present; widespread in the Americas | 72 |
72 | Fruit valves coriaceous; lenticels on twigs orange, orbicular to slightly elongated vertically, some more than 0.6 mm across | * Parasenegalia |
– | Fruit valves thinner, mostly chartaceous to cartilaginous; lenticels on twigs not orange, orbicular to slightly elongated horizontally, very rarely 0.6 mm across | Mariosousa |
73 | Leaves lacking extrafloral nectaries | 74 |
– | Leaves with extrafloral nectaries on the petiole, leaf rachis, or both | 76 |
74 | Inflorescence units in spherical capitula; fruit with membranous to coriaceous valves; pollen in 16-celled, discoid and acalymmate polyads | * Zapoteca |
– | Inflorescence units mostly obconical capitula, less frequently spherical capitula, umbels or pseudoracemes; fruit with mostly woody valves, rarely coriaceous; pollen in (7) 8 (10)-celled and flattened-ovoid polyads | 75 |
75 | Plants from the New World; pollen in 8-celled and calymmate polyads, the basal cell with a conspicuous sticky appendage | * Calliandra |
– | Plants from tropical Africa; pollen in (7) 8 (10)-celled and acalymmate polyads, the basal cell with an extremely reduced sticky appendage | Afrocalliandra |
76 | Plants from the New World | 77 |
– | Plants from continental Africa, tropical Asia, Malesia and Australia | 111 |
77 | Stems armed at all or most nodes with spines or thorns | 78 |
– | Stems unarmed | 85 |
78 | Stems armed with axillary thorns; perulate resting buds axillary to most leaves | * Chloroleucon |
– | Stems armed with stipular spines; buds not perulate, protected by the adaxial side of the petiole | 79 |
79 | Petiolar nectary below the first pair of pinnae | 80 |
– | Petiolar nectary at the point of origin of the first (or only) pair of pinnae | 82 |
80 | Trees or shrubs, generally sarmentose; flower buds flask-shaped, flowers opening at night; androecium up to 9 cm long | Sphinga |
– | Erect trees or shrubs, never sarmentose; flower buds ovoid-pyriform, flowers opening during the day; androecium usually less than 3 cm long, very rarely up to 7 cm long | 81 |
81 | Calyx shallowly campanulate, 1–2 mm long; corolla lobes recurved at anthesis; ovary disk absent | Havardia |
– | Calyx deeply campanulate, 2.8–3.4 mm long; corolla lobes erect at anthesis; ovary disk present (sometimes poorly developed) | Gretheria |
82 | Leaves with one pair of pinnae, leaflets 1 pair per pinna, if leaves with more pinnae and/or leaflets then seeds with fleshy, often brightly coloured arils | Pithecellobium |
– | Leaves with 2 or more pairs of pinnae, never 1, leaflets 2–30 pairs per pinna; seeds lacking aril | 83 |
83 | Fruit cylindrical, woody, straight or slightly curved, deeply internally septate; seeds globose; growing in lowlands of Mexico and USA (Texas) | Ebenopsis |
– | Fruit flattened or slightly subterete, subwoody and curved, without internal septa; seeds lentiform; growing in highlands of Mexico | 84 |
84 | Leaves with 1–2 pairs of pinnae; leaflets 3–12 per pinna, blades suborbicular, broadly oblong, or elliptic (then revolute); corolla lobes ascending | Painteria |
– | Leaves with 3–7 pairs of pinnae; leaflets 10–20 per pinna, blades narrowly oblong, linear-oblong or lanceolate; corolla lobes recurving | Ricoa |
85 | Inflorescences cauliflorous, the spikes or capitula arising from efoliate nodes below the expanded leaves | 86 |
– | Inflorescence units mostly axillary or clustered in efoliate pseudoracemes or panicles, sometimes on efoliate nodes of young stems | 87 |
86 | Calyx 14–20 mm long; bracts usually with patelliform nectaries, rarely lacking nectaries | * Macrosamanea |
– | Calyx up to a maximum 14 mm long, usually much shorter; bracts lacking patelliform nectaries | * Zygia |
87 | Terminal and axillary resting buds ovoid, composed of closely imbricate, striately veined scales; flowers sessile in globose capitula | 88 |
– | Resting buds absent or, if rarely present, the scales not striately veined, or loosely imbricate; flowers sessile or not, in spikes, racemes, capitula or umbels | 91 |
88 | Capitula grouped in efoliate pseudoracemes or panicles; fruit plano-compressed, tardily dehiscent with papery valves | Blanchetiodendron |
– | Capitula solitary or fasciculate in leaf axils or at efoliate nodes below the expanding leaves; fruit dehiscent or indehiscent with thick or papery valves | 89 |
89 | Branches with brachyblasts; flowers in each capitulum dimorphic; fruits with firm valves, compressed but somewhat plump | * Chloroleucon |
– | Branches lacking brachyblasts; flowers homomorphic; fruits with stiff paper valves, plano-compressed | 90 |
90 | Lower leaflet surface with evident reticulate secondary and tertiary venation; eastern and central Brazil | Leucochloron |
– | Lower leaflet surfaces with 1–2 (3) prominent primary veins, but otherwise the venation not evident; endemic to eastern Andean valleys in Bolivia | Boliviadendron |
91 | Leaflet venation palmate or the leaflets very narrow and 1-veined | 92 |
– | Leaflet venation pinnate | 95 |
92 | Fruits indehiscent, somewhat woody and ear-shaped (reniform-auriculiform), internally septate into 1-seeded chambers | 93 |
– | Fruits dehiscent through both margins or, if indehiscent, not ear-shaped, not internally septate into 1-seeded chambers | 94 |
93 | Indumentum of leaf rachis ferruginous; leaflets in 40–80 pairs per pinna, very small (2.5–6 mm long) and 1-veined; sepals and petals ferruginous-villous externally; ovary tomentose | Robrichia |
– | Indumentum of leaf rachis pallid; leaflets either fewer pairs per pinna and/or larger, palmately-veined; sepals and petals puberulent, rarely sericeous, strigose-pilose, or glabrous; ovary at anthesis glabrous | Enterolobium |
94 | Flowers homomorphic, larger (corolla 11–63 mm long, androecium 30–70 mm long); seeds papery, lacking a pleurogram | * Macrosamanea |
– | Flowers dimorphic, smaller (corolla <6.5 mm long, androecium <20 mm long); seeds with a hard coat, presenting a U-shaped or O-shaped pleurogram | Pseudalbizzia |
95 | Capitula clustered into efoliate terminal panicles, the inflorescence units never subtended by a leaf; fruit a long plano-compressed papery loment (20–70 cm long), the articles twisted through ±90° at each isthmus | Cedrelinga |
– | Capitula solitary, fasciculate in leaf axils, or clustered in axillary efoliate pseudoracemes, or the pseudoracemes at efoliate nodes below the leaves; fruits dehiscent or indehiscent but if indehiscent never with twisting articles | 96 |
96 | Fruits indehiscent or breaking into 1-seeded articles | 97 |
– | Fruits dehiscent through one or both margins | 100 |
97 | Fruits with a strongly biconvex or sub-cylindrical body, the valves woody or pulpy and internally at least partially septate; flowers heteromorphic in each capitulum | 98 |
– | Fruits mostly plano-compressed with papery, leathery or subligneous valves; flowers either homomorphic or heteromorphic | 99 |
98 | Androecium bicoloured, the stamens basally white and distally pink or reddish | Samanea |
– | Androecium uniformly whitish | * Hydrochorea |
99 | Fruits lomentiform, the ripe leathery or subligneous valves breaking into 1-seeded articles but leaving no persistent margins; flowers heteromorphic | * Hydrochorea |
– | Fruits indehiscent or breaking into 1-seeded articles leaving a persistent margin (craspedium); flowers homomorphic | Lysiloma |
100 | Flowers within each capitulum dimorphic | 101 |
– | Flowers homomorphic | 103 |
101 | Fruit curved to spiral, chartaceous, plano-compressed, twisting at dehiscence to expose the reddish-orange endocarp | * Jupunba |
– | Fruit straight or slightly arched, the papery or woody valves not becoming spirally twisted during dehiscence | 102 |
102 | Fruit linear, the margins slender, sometimes immersed, the valves papery, barely dehiscent through one or both sutures | Pseudosamanea |
– | Fruit oblong with raised margins framing plane and woody valves, late dehiscent along one suture (follicle) | * Hydrochorea |
103 | Fruits elastically dehiscent from the apex, the chartaceous to weakly coriaceous valves becoming arched backwards; small erect or scandent shrubs | * Zapoteca |
– | Fruits dehiscent through one or both margins but the valves never recurving and becoming arched backwards; trees | 104 |
104 | Fruit falcately or spirally recurved, the valves compressed but elevated over each seed, twisting at dehiscence to reveal the reddish endocarp; seeds bicoloured (white and dark bluish) or with a translucent testa | 105 |
– | Fruit compressed with flat valves, if cylindrical and torulose with reddish valves then with black or dark brown seeds | 107 |
105 | Inflorescence a dense globose capitulum; flowers sessile; fruits with a puberulent and ferruginous epicarp; seeds with a U-shaped median pleurogram | Abarema |
– | Inflorescence in lax elongated or congested racemes or elongated spikes; flowers sessile to pedicellate; fruits without a puberulent epicarp; seeds with an apical-basal pleurogram or the pleurogram absent | 106 |
106 | Leaflets obovate, ovate, oblong, linear or rhombic, secondary veins not arched; inflorescences lax or congested racemes, rarely spikes but then in combination with 2–12 pairs of leaflets per pinna; pleurogram usually present | * Jupunba |
– | Leaflets elliptic to ovate-lanceolate, the secondary veins arched; inflorescences long-racemose or spiciform; pleurogram absent | Punjuba |
107 | Bracts with nectaries on the upper surface | * Macrosamanea |
– | Bracts lacking nectaries | 108 |
108 | Corolla slender tubular with very short teeth; fruits cylindrical, torulose or moniliform, the ripe valves externally red and coriaceous | * Cojoba |
– | Corolla campanulate, the teeth at least ¼ of the total length; fruits not as above | 109 |
109 | Leaves unijugate, each pinna with just one leaflet | * Zygia |
– | Leaves either with more pinnae or leaflets | 110 |
110 | Fruit with inconspicuously raised margins, the valves linear, plano-compressed, stiffly papery, inertly dehiscent through both sutures; capitula solitary or fasciculate in leaf axils; flowers homomorphic and sessile; tropical Mexico | Hesperalbizia |
– | Fruit with markedly raised margins framing an oblong body, the valves plane and woody and tardily dehiscing through the upper margin only (follicle); capitula clustered in pseudoracemes; flowers heteromorphic, the peripheral ones pedicellate; South America | * Hydrochorea |
111 | Fruit elastically dehiscent from apex downwards, the stiffly coriaceous valves becoming arched backwards | 112 |
– | Fruits dehiscent through one or both margins or indehiscent or lomentiform, if dehiscent through both margins the valves never become arched backwards | 113 |
112 | Branches unarmed; Madagascar | Viguieranthus |
– | Branches armed with stipular spines; north-east India and adjacent Bangladesh and Myanmar | * Calliandra |
113 | Branches armed with stipular spines | 114 |
– | Branches unarmed or with infranodal prickles | 115 |
114 | Capitula with homomorphic flowers; fruit submoniliform with leathery valves; Sri Lanka to Vietnam | Thailentadopsis |
– | Capitula usually with heteromorphic flowers; fruit plano-compressed with papery valves; Malesia | * Albizia |
115 | Leaflets alternate | 116 |
– | Leaflets opposite or subopposite | 117 |
116 | Inflorescences spikes, racemes or capitula, these clustered in pseudoracemes, panicles or umbels; fruits woody, straight, indehiscent or late dehiscent | Serianthes |
– | Inflorescences corymbiform umbels, solitary or fasciculate in leaf axils or on short-shoots; fruits chartaceous, dehiscing along one margin only, contorted to expose the reddish endocarp | Pararchidendron |
117 | Plants from Africa | 118 |
– | Plants from Asia, Australia and the Pacific | 120 |
118 | Fruits with papery valves, usually dehiscent, non-septate | * Albizia |
– | Fruits septate, the valves thin and fibrous or thick and woody, indehiscent or lomentiform | 119 |
119 | Pedicels of peripheral flowers at least 1 mm long; leaves with (1) 2–3 pairs of pinnae; fruits lomentiform with seeds dispersed as 1-seeded articles | * Hydrochorea |
– | Pedicels of peripheral flowers up to 0.5 mm long; leaves with (3) 5–35 pairs of pinnae; fruits indehiscent or if lomentiform only tardily breaking up into articles | Osodendron |
120 | Seeds lacking a pleurogram; gynoecium uni- or pluricarpellate | 121 |
– | Seeds with a pleurogram; gynoecium unicarpellate | 123 |
121 | Fruits dehiscing through one or both margins, spirally contorted, if straight then turgid, usually with the valves reddish outside and orangish-red within, sharply contrasting with the seeds; seeds black or bluish-black, unwinged | Archidendron |
– | Fruits dehiscing through both margins, straight and plano-compressed, the valves not reddish inside; seeds strongly flattened with a narrow marginal wing | 122 |
122 | Inflorescence units capitula; calyx and corolla pubescent | Heliodendron |
– | Inflorescence units spikes, spiciform racemes, racemes, rarely capitula but then with calyx and corolla glabrous | Archidendropsis |
123 | Inflorescence units capitula | * Albizia |
– | Inflorescence units racemes, spikes or spiciform racemes | 124 |
124 | Fruit plano-compressed with woody valves, the endocarp breaking into 1-seeded articles; flowers in lax axillary racemes | Wallaceodendron |
– | Fruit with papery valves, the endocarp not breaking into 1-seeded articles; flowers in dense racemes or the racemes grouped in panicles | 125 |
125 | Racemes solitary or paired in leaf axils | Paraserianthes |
– | Racemes or spikes clustered in large panicles | Falcataria |
126 | Lowermost sepal modified and boat-shaped, usually making a hood in bud | 127 |
– | Lowermost sepal not differentiated from the remaining ones | 152 (page 48) |
127 | Leaves terminating in a pair of pinnae plus a single terminal pinna | 128 |
– | Leaves terminating in a pair of pinnae or with pinnae clearly alternate | 135 |
128 | Plant armed with scattered straight conical spines, and short, lateral spinescent shoots; fruits oblong to fusiform, glabrous, dehiscing along the middle of the valves or parallel to the margin | * Haematoxylum |
– | Plant unarmed; fruits not dehiscing along the middle of the valves | 129 |
129 | Sepals persistent in fruit | Hoffmannseggia |
– | Sepals caducous in fruit | 130 |
130 | Fruits cylindrical-torulose; central and western Argentina, in subtropical wooded grassland and scrub, especially on salt flats | Stenodrepanum |
– | Fruits never cylindrical-torulose, widespread in the Americas and South Africa | 131 |
131 | Stipules linear, persistent; androecium and gynoecium cupped in the lower cucullate sepal; lower lateral sepals forming a platform at right angles to the abaxial cucullate sepal; fruits with simple trichomes, glandular-punctate trichomes, and plumose, dendritic and/or stellate trichomes | Pomaria |
– | Stipules caducous; androecium and gynoecium not cupped in the lower sepal, deflexed; lateral sepals not forming a platform; fruits glabrous or with simple and/or gland-tipped trichomes, the latter sometimes also dendritic or plumose | 132 |
132 | Fruits indehiscent, thickened and somewhat torulose; inflorescence a raceme or panicle, often corymbose; leaflets glabrescent and eglandular, or with glandular dots parallel to the midvein | * Libidibia |
– | Fruits dehiscent, often with twisting valves; inflorescence a raceme or panicle, sometimes pyramidal in shape; leaflets glabrescent to densely pubescent or with a stellate indumentum; leaflets eglandular, or with dark subepidermal glands, and/or with glandular dots sunken in the margins or parallel to the margin on the abaxial side of the leaflets | 133 |
133 | Leaflets alternate, or occasionally sub-opposite (rarely fully opposite), with dark subepidermal glands (best seen with a ×10 hand lens); stellate indumentum sometimes present on foliage and inflorescence rachis; fruit subligneous to woody, with thickened sutures | * Cenostigma |
– | Leaflets always opposite, without dark subepidermal glands; stellate indumentum never present on foliage or rachis; fruit coriaceous to subligneous, sutures not thickened | 134 |
134 | Shrubs or small to medium-sized trees varying from (0.5) 1–12 (20) m tall, occasionally functionally herbaceous subshrubs, woody at the base; flowers sometimes laterally compressed; petals yellow, red, pink or orange; ovary eglandular or covered in gland-tipped trichomes, the hairs never dendritic; widespread across low-elevation seasonally dry tropical forests and woodlands in Mexico, Central America, the Caribbean, and in Caatinga vegetation in Brazil, and in patches of dry forest, deserts, yungas-puna transition zones, and chaco-transition forests in Argentina, Bolivia, Chile and Paraguay | Erythrostemon |
– | Small to medium-sized, often decumbent shrubs, 0.3–2.5 m tall; flowers never laterally compressed; petals yellow, sometimes all five petals streaked with red markings; ovary covered in gland-tipped trichomes, which are sometimes dendritic; occurring at mid elevations in dry inter-Andean valleys, in Ecuador, Peru, Bolivia and Argentina | Arquita |
135 | Plants unarmed | 136 |
– | Plants armed | 139 |
136 | Fruit thin, plano-compressed, oblong-elliptic to elliptic, valves membranous to papyraceous, indehiscent; margin of the lower cucullate sepal pectinate-glandular; flowers unisexual; leaflets eglandular | Coulteria |
– | Fruit oblong-elliptic, elastically dehiscent with twisting valves; margin of the lower cucullate sepal entire; flowers bisexual; leaflets eglandular or with red glands | 137 |
137 | Flowers nearly actinomorphic; trees, up to 25 m tall; leaflets eglandular or with red glands; eastern Africa (Kenya and Tanzania), and northern and north-western Madagascar | * Stuhlmannia |
– | Flowers clearly zygomorphic; shrubs or small trees, up to 5 m tall; leaflets eglandular; Cuba or northern Madagascar | 138 |
138 | Fruits laterally compressed; anthers glabrous; endemic to Cuba (near Moa, in the Sierra de Nipe) | * Caesalpinia |
– | Fruits inflated and hollow; anthers pubescent; endemic to the northern tip of Madagascar (Orangea peninsula, near Antsiranana) | * Denisophytum |
139 | Trees or erect shrubs | 140 |
– | Lianas or climbing or trailing shrubs | 144 |
140 | Fruits indehiscent, somewhat fleshy, turgid and coriaceous; lower cucullate sepal with a pectinate/fimbriate or entire margin | Tara |
– | Fruits dehiscent, with valves twisting upon dehiscence, laterally-compressed and subligneous to woody; lower cucullate sepal with an entire margin | 141 |
141 | Fruits armed with woody prickles; stems with upturned thorns arising from woody protuberances; petals yellow, the standard with a conspicuous red blotch on the inner face | Paubrasilia |
– | Fruits unarmed; stems with straight to deflexed prickles; petals yellow, white, pink, red or orange | 142 |
142 | Petals pink-purple to whitish pink; bracts broadly ovate to suborbicular with an aristate apex; fruits pyriform with rounded, oblique bases; leaflets sometimes with translucent dots on lower surface | Gelrebia |
– | Petals yellow, red, orange or white; bracts lanceolate to linear with an acute to acuminate apex; fruits oblong-elliptic, short-stipitate, with cuneate base; leaflets eglandular | 143 |
143 | Petals orange, red, or white; Central America, Mexico, the Caribbean and the northern Andes (Peru to Colombia) | * Caesalpinia |
– | Petals yellow, sometimes with red markings on the standard (median petal); Somalia, Ethiopia, Argentina, Paraguay, Mexico, Florida and the Caribbean | * Denisophytum |
144 | Fruits winged, although wing sometimes very narrow | 145 |
– | Fruits without a wing | 148 |
145 | Fruit a samara with a basal 1-seeded chamber and a prolonged upper suture that is broadly winged | Pterolobium |
– | Fruit 1 or more seeded, with a longitudinal (often narrow) wing along the upper suture, if 1-seeded then with a central seed-chamber | 146 |
146 | Fruit with a prominent wing 2 mm or more wide | Mezoneuron |
– | Fruit with a less distinct wing, usually 2 mm wide or less (although occasionally up to 4 mm, or carinate) | 147 |
147 | Fruit oblong-elliptic, dehiscent, terminating in a sharp beak, 4–9-seeded | * Biancaea |
– | Fruit circular to sub-elliptic or lunate, indehiscent, 1 (rarely 2)-seeded | * Ticanto |
148 | Plants with glands on stems, leaf rachis, inflorescence, and fruits; needle-like trichomes on inflorescence rachis and pedicels | Hultholia |
– | Plants eglandular; stems with recurved prickles; pedicels and inflorescence peduncles with a few prickles near their bases | 149 |
149 | Fruit oblong to oblong-elliptic | 150 |
– | Fruit broadly elliptic to circular | 151 |
150 | Fruit oblong, indehiscent, somewhat fleshy, sub-torulose, with thickened sutures, terminating in an acute apex, exocarp and endocarp strongly adnate; seeds sub-globose | Moullava |
– | Fruit oblong to oblong-elliptic, laterally compressed, dehiscent, coriaceous to subligneous, with a smooth, regular outer surface, base often much narrower than the truncate apex which terminates in a sharp beak, exocarp and endocarp separate easily; seeds flattened to ellipsoidal | * Biancaea |
151 | Flowers unisexual, segregated into pistillate and staminate racemes; fruits usually covered in spinescent bristles; seeds globose, with parallel fracture lines concentric with the small apical hilum | Guilandina |
– | Flowers bisexual, in racemes; fruits not spinescent; seeds laterally compressed, smooth, without fracture lines | * Ticanto |
152 | Plants from the Americas | 153 |
– | Plants from Africa, Asia and Australia | 165 |
153 | Androecium strongly dimorphic including fertile stamens and staminodes | 154 |
– | Androecium homomorphic with all stamens fertile | 155 |
154 | Indumentum of rusty T-shaped trichomes; stipules pinnate; flowers showy with bright yellow clawed petals; fertile stamen 1, this clearly longer than the 9 staminodes | * Moldenhawera |
– | Indumentum lacking T-shaped trichomes; stipules linear or absent; flowers small, pale yellow to cream or dark orange to reddish, petals not clawed; fertile stamens 5; staminodes 5, spatulate, free or connate forming a dome | Dimorphandra |
155 | Stamens united basally into a tube deeply split on one side rendering the flowers slightly bilateral; fruit valves becoming recurved backwards upon dehiscence; stipules pinnate | Jacqueshuberia |
– | Stamens free; fruits indehiscent or passively dehiscent, rarely with the valves becoming longitudinally twisted; stipules entire (not pinnate) | 156 |
156 | Inflorescence units spikes with densely packed flowers | 157 |
– | Inflorescence units racemes or panicles | 159 |
157 | Sepals united into a gamosepalous calyx; fruits flat, coriaceous, indehiscent and marginally compressed or woody, elastically dehiscent along both sutures; South America in Amazonia and eastern Brazil | Dinizia |
– | Sepals free; fruit thick-walled, indehiscent, subterete or flat but then with margins not compressed | 158 |
158 | Fruit compressed with a linear outline; plants from temperate North and South America | * Gleditsia |
– | Fruit sub-terete; plants from Hispaniola (Haiti and Dominican Republic) | Arcoa |
159 | Sepals and petals similar, greenish or whitish, the sepals not covering petals in bud | 160 |
– | Sepals and petals clearly differentiated, the petals larger than sepals and yellow; petals covering petals in bud | 161 |
160 | Hypanthium elongate, sub-cylindrical; leaflet margins entire; fruit sessile | *Gymnocladus |
– | Hypanthium campanulate, discoid or turbinate; leaflet margins crenulate, rarely entire; fruit stipitate | * Gleditisia |
161 | Plants usually armed with spines or thorns, rarely unarmed; adaxial petal clearly differentiated in shape and/or colour, with a thicker claw | * Parkinsonia |
– | Plants unarmed; all petals more or less equal | 162 |
162 | Fruits dehiscent through both sutures | 163 |
– | Fruits indehiscent | 164 |
163 | Fruit 1-seeded, flattened, spatulate, oblanceolate or spoon-shaped, the valves firm-coriaceous, with the endocarp released as a thin, papery wing-like envelope; rain forests of southern Mexico to South America | Schizolobium |
– | Fruit (2) 3 (4)-seeded, plano-compressed, linear, the margins narrowly winged; dry forests of western and southern Mexico from Sonora and Baja California Sur south to Chiapas | Conzattia |
164 | Fruit flat, oblong or narrowly ellipsoid, the mesocarp thin and dry, tapering at both ends, with a firm wing-like extension on each suture, valves usually longitudinally striate; flowers bisexual; stigma peltate | * Peltophorum |
– | Fruit cylindrical, the mesocarp thickened and fibrous, valves smooth, pale orange-brown when ripe, with fibrous-spongy septae between the seeds forming marked seed cavities; flowers unisexual by reduction of the androecium or gynoecium, dioecious; stigma circular, flat or slightly concave | Heteroflorum |
165 | Flowers apetalous with a fleshy hypogynous disk wider than the calyx | * Ceratonia |
– | Flowers with sepals and petals | 166 |
166 | Flowers unisexual and tetramerous, with 4 sepals, 4 petals and 8 stamens (staminate flowers); fruit indehiscent with 4 membranous wings; Madagascar | Tetrapterocarpon |
– | Flowers bisexual or unisexual but then with 5 (6) sepals and petals; fruits unwinged or with just two straight wings along the sutures, indehiscent or dehiscent along one or both margins | 167 |
167 | Sepals and petals very similar, greenish or lavender coloured, the narrow sepals not covering petals in bud | 168 |
– | Sepals and petals clearly differentiated | 169 |
168 | Unarmed trees; hypanthium elongate, sub-cylindrical, 6–12 mm long; leaflet margins entire; fruit sessile | * Gymnocladus |
– | Trees usually armed with thorns and/or branched spines; hypanthium shortly campanulate, 1–4 mm long; leaflet margins crenulate, rarely entire; fruit stipitate | * Gleditsia |
169 | Sepals valvate in bud | 170 |
– | Sepals imbricate in bud | 171 |
170 | Flowers radial or weakly zygomorphic; all sepals free and reflexed at anthesis; petals showy and long-clawed | * Delonix |
– | Flowers strongly zygomorphic and resupinate; 4 sepals united into a lip making a spathaceous calyx; petals reduced | Colvillea |
171 | Adaxial petal differentiated in shape, colour or both, with a thicker claw; plants mostly armed with stipular spines, axillary thorns or spinescent leaf rachides, rarely unarmed | * Parkinsonia |
– | All petals similar, if clawed the claws equally thin; unarmed trees or shrubs | 172 |
172 | Flowers pedicellate in lax racemes; petals bright yellow, clawed, with wrinkled margins; style terminating in a peltate stigma | 173 |
– | Flowers sessile or shortly pedicellate in dense spikes or spiciform racemes; petals white or greenish white, not clawed and/or with entire margins; style short and stigma porate | 174 |
173 | Fruit indehiscent, flat, oblong or narrowly ellipsoid, tapering at both ends, with a wing-like extension on each suture | * Peltophorum |
– | Fruit elastically dehiscent from the apex, narrowly oblong-obovate, compressed with greatly thickened margins, the valves woody, recurving | Bussea |
174 | Leaflets alternate | 175 |
– | Leaflets opposite | 177 |
175 | Fruit 1 (2)-seeded, elliptical to oblong-elliptical, flat, thin-valved, indehiscent; petals reflexed at anthesis | Burkea |
– | Fruit 2–15-seeded, linear or oblong-linear, valves stiffly coriaceous or woody, indehiscent or late dehiscent; petals erect at anthesis | 176 |
176 | Ovary long-stipitate; petals green to greenish-yellow; fruits not septate, with thin woody or leathery valves and margins not thickened | Erythrophleum |
– | Ovary subsessile; petals red; fruits with woody resinous valves and thick, raised margins, the endocarp internally septate into 10–15 1-seeded envelopes | Pachyelasma |
177 | Petals basally united into a gamopetalous corolla; leaves terminating in a pair of pinnae | * Sympetalandra |
– | Petals free; leaves terminating in a solitary pinna | 178 |
178 | Stamens dimorphic, the 5 antepetalous shorter and with thin filaments, the 5 antesepalous longer with filaments wider towards the apex; flowers with sepals, petals and stamens whitish, in cylindrical erect spikes; West Africa | Stachyothyrsus |
– | Stamens homomorphic, all with filiform filaments; flowers with showy red hypanthium and bright orange stamens in dense drooping racemes, the pedicels twisting so that all flowers face upwards; tropical Asia | Acrocarpus |
179 | Petals valvate in bud; petals joined into a gamopetalous corolla; seeds with a U-shaped pleurogram | 180 |
– | Petals imbricate in bud, the uppermost petal in inner position; petals free; seeds lacking a pleurogram | 185 |
180 | Leaves lacking nectaries | 181 |
– | Leaf rachis with nectaries between the leaflets | 182 |
181 | Plants armed with triangular broad-based recurved prickles on the shoots; leaves densely fascicled on cushion-like brachyblasts, petiole flat and laterally expanded, leaflets small (0.2–0.6 cm long); fruit a craspedium with undulate margins, persisting after the 1-seeded articles disperse; Mexico (Coahuila) | * Mimosa |
– | Plants unarmed; leaves not fascicled, with a grooved unexpanded petiole and large leaflets (4–10 cm long); fruits elastically dehiscent from the apex, the woody valves becoming recurved backwards after dehiscence; coastal plains of the three Guianas | * Calliandra |
182 | Plants armed with straight horizontal stipular spines; fruits elastically dehiscent from the apex; India (western Ghats) | Sanjappa |
– | Plants unarmed; fruits indehiscent or passively dehiscent; Americas | 183 |
183 | Inflorescences cauliflorous | * Zygia |
– | Inflorescences solitary or fasciculate in leaf axils or on efoliate terminal shoots | 184 |
184 | Fruits indehiscent; seeds surrounded by a whitish sweet sarcotesta | Inga |
– | Fruits late dehiscent through both sutures, terete to moniliform with bright red valves; seeds lacking an aril or a sarcotesta | * Cojoba |
185 | Lowermost sepal modified and boat-shaped, usually forming a hood in bud (cucullate) | 186 |
– | Lowermost sepal not differentiated from the other four | 194 |
186 | Armed shrubs or trees, with prickles scattered along the branches or in pairs below the stipules, or plants with short shoots modified into persistent thorns | 187 |
– | Unarmed shrubs or trees | 190 |
187 | Sepals persistent in fruit; fruit cylindrical, covered with resinous hairs; pairs of needle-like prickles inserted below the stipules and leaf petiole; endemic to northern Chile, from the Coquibo and La Serena valleys | Balsamocarpon |
– | Sepals caducous; fruit flattened, non-resinous; plants armed with scattered straight spines on shoots or with curved deflexed prickles; widely distributed across Central America, Mexico, the Caribbean, South America and Namibia | 188 |
188 | Fruit a lomentum, with 4 coarsely serrate wings, breaking up into one-seeded articles; native to Paraguay and northern Argentina | Lophocarpinia |
– | Fruit unsegmented, without wings; native to Namibia and Mexico to northern South America | 189 |
189 | Fruit sub-circular to sickle-shaped, tardily dehiscent along the sutures, finely pubescent and with robust patent trichomes; endemic to Namibia | Hererolandia |
– | Fruit oblong to fusiform, dehiscent along the middle of the fruit valves or close to the fruit margin, but never along the sutures, lacking patent trichomes | * Haematoxylum |
190 | Sepals persistent; fruit indehiscent, gall-like, covered with long bristles; north-west Argentina | Zuccagnia |
– | Sepals caducous; fruits ovoid to elliptic, not gall-like, glabrous or covered in a different type of indumentum | 191 |
191 | Fruit elastically dehiscent, with valves twisting upon dehiscence, laterally-compressed and subligneous to woody, oblanceolate to oblong-elliptic | 192 |
– | Fruit indehiscent, thickened and fleshy, ovoid or elliptic | 193 |
192 | Fruit subligneous, lacking a crest; sepals valvate; stellate indumentum lacking; restricted to Africa and Madagascar | * Stuhlmannia |
– | Fruit woody, with conspicuously thickened sutures, sometimes with a crest proximally on the adaxial side; stellate indumentum often present; sepals imbricate; restricted to the Neotropics | * Cenostigma |
193 | Fruit elliptic, somewhat thick and fleshy, bright red at maturity, rounded at apex and base, 1–2-seeded; leaflets with black, sessile glands on the lower surface; seeds compressed-turgid; sepals imbricate; endemic to Hispaniola and Puerto Rico | * Libidibia |
– | Fruit ovoid, with a ligneous and brownish pericarp, apex beaked, 1–4-seeded; leaflets with red glands on the lower surface; seeds ovoid; sepals valvate; endemic to north-eastern Africa | Cordeauxia |
194 | Leaves imparipinnate and/or with alternate leaflets | 195 |
– | Leaves paripinnate with opposite leaflets | 202 |
195 | Armed trees, with simple or compound thorns on trunk and branches (sometimes thornless in cultivated plants); leaflet margins crenulate, rarely entire; temperate and subtropical North and South America, eastern Asia and Caspian region | * Gleditsia |
– | Unarmed trees or shrubs; leaflet margins entire; tropical South America, Mediterranean region and Arabian Peninsula | 196 |
196 | Plants polygamo-dioecious; flowers apetalous, with a conspicuous disk wider than the calyx; Mediterranean region of northern Africa and southern Europe to Arabian Peninsula | * Ceratonia |
– | Plants with flowers hermaphrodite, with petals and a tiny disk or the disk absent; tropical South America | 197 |
197 | Flowers pentamerous, with 5 sepals, 5 petals and 10 stamens; filaments whitish or pale yellow; fruits indehiscent or passively dehiscent; seeds various but never with a spongy wing | 198 |
– | Flowers with different numbers of sepals and/or petals and/or stamens; stamens 10–17 (25) per flower, filaments dark red and showy; fruits dehiscent with the valves loosely spiralling; seeds discoid, with a spongy wing | Campsiandra |
198 | Flowers small (ca. 3 mm diam.), entirely white or greenish-white, in spicate, axillary, catkin-like, racemes; fruit a 1-seeded samara with a style remnant laterally displaced at apex of the seed chamber | Pterogyne |
– | Flowers > 7 mm diam with yellow petals; flowers in terminal panicles, petals pale yellow or bright yellow; fruit dehiscing through one or both margins | 199 |
199 | Leaves with a nectary on the petiole or leaf rachis, between leaflets; fruits dehiscing only through one suture; flowers small (to 15 mm diam); petals pale yellow, obovate, not clawed | 200 |
– | Leaves lacking nectaries; fruits indehiscent or dehiscent along both sutures; flowers larger (> 20 mm diam.); petals with bright yellow blades widely expanded from the claws | 201 |
200 | Ovary 1-ovulate; fruit 1-seeded, globose to pyriform, subligneous, late dehiscent, the surface not ribbed | Vouacapoua |
– | Ovary with more than 1 ovule; fruit inflated, oblong, woody, longitudinally ribbed, with 2–3 red seeds | Batesia |
201 | Fruit linear, narrowly winged along the upper suture, the wing up to 3 mm wide, late dehiscent along both margins, the endocarp not septate | Recordoxylon |
– | Fruit oblong, dehiscent along both margins to release winged 1-seeded envelopes resulting from endocarp fragmentation | Melanoxylum |
202 | Anthers of some stamens or staminodes dehiscing through pores | 203 |
– | All anthers dehiscing through longitudinal slits | 206 |
203 | Androecium with just one fertile stamen and 7 or 9 staminodes, the fertile stamen much longer and equalling the length of the style, the anther with a villous connective; staminodes short, barely extending beyond the ovary length; stipules pinnately compound; indumentum of rusty T-shaped trichomes | * Moldenhawera |
– | Androecium of 7–10 fertile stamens, staminodes up to 3, usually much shorter than the fertile ones; stipule blades never compound; indumentum various but never including T-shaped trichomes | 204 |
204 | Pedicels with a pair of alternate bracteoles near the middle; fruits elastically dehiscent, the valves twisting after dehiscence; anthers with pubescent sutures; extrafloral nectaries, when present, with a concave or flat head | * Chamaecrista |
– | Bracteoles when present at the base of the pedicels; fruits indehiscent or passively dehiscent, valves remaining straight after dehiscence; anther sutures glabrous; extrafloral nectaries, when present, with a convex head | 205 |
205 | Foliar nectaries absent; hypanthium solid, turbinate or conical; 3 abaxial stamens with sigmoidal filaments and longitudinally dehiscent anthers; adaxial anthers with basal pores; fruits woody to coriaceous, indehiscent, commonly cylindrical or less often compressed, rarely fragmenting into 1-seeded segments | Cassia |
– | Foliar nectaries often present on the petiole and/or leaf rachis; hypanthium absent; all stamen filaments straight or curved (but not sigmoidal); abaxial and median stamens with apical pores, the 3 adaxial stamens sterile (staminodes); fruits dehiscent with chartaceous or papery valves or indehiscent, with a flat compressed to cylindrical body | * Senna |
206 | Plants polygamo-dioecious; flowers apetalous, with a conspicuous disk wider than the calyx | * Ceratonia |
– | Flowers hermaphrodite; flowers with 5 petals and a tiny disk or a disk absent | 207 |
207 | Plants from tropical Africa and Malesia | 208 |
– | Plants from tropical America | 211 |
208 | Plants armed with thorny branches or spinescent shoots | 209 |
– | Plants unarmed | 210 |
209 | Leaflets 2–3 pairs, symmetrical and petiolulate; flowers showy, zygomorphic, petals clawed and white but the upper petal slightly larger and with a yellow blotch; sepals valvate; Madagascar | * Delonix |
– | Leaflets 5 or more pairs per leaf, markedly asymmetrical, sessile; flowers small, greenish, actinomorphic; calyx open in bud, not covering the petals; South Africa | Umtiza |
210 | Flowers wine-red in loose pendulous panicles; petals and stamens free to the hypanthium rim; fruits elastically dehiscent, the valves twisting; West African rainforests | Chidlowia |
– | Flowers white to greenish-white in dense spiciform racemes; petals and stamens joined at the base; fruits indehiscent, longitudinally striate; Malesia | * Sympetalandra |
211 | Fruits samaroid, oblong-elliptical, tapered at each end, with a central seed-nucleus and a thin marginal wing, the exocarp flaking when ripe; hypanthium cupular, either symmetrical (the ovary attached in the middle of the hypanthium) or asymmetrical (ovary laterally attached) | Tachigali |
– | Fruits dehiscent, oblong to linear with woody valves; hypanthium funnel-shaped or disk-shaped and symmetrical | 212 |
212 | Stipules foliaceous, suborbicular and coriaceous, persistent or late caducous; flowers > 2 cm long in terminal corymbose panicles; fruits elastically dehiscent from apex, the valves rolling up backwards | Arapatiella |
– | Stipules membranous and caducous, lanceolate; flowers < 1 cm long, in axillary racemes or spikes | 213 |
213 | Flowers distinctly pedicellate in racemes; sepals and petals reflexed; all 10 stamens fertile; fruits with thin ligneous valves, passively dehiscent; seeds plano-compressed with a wide marginal wing | Diptychandra |
– | Flowers sessile or subsessile in dense spikes or spiciform racemes grouped in woody terminal panicles; sepals and petals erect or spreading; androecium of 5 fertile stamens alternating with 5 staminodes; fruits woody, the valves twisting after dehiscence; seeds large (mostly > 10 cm long and weighing about 1 kg), bulky, unwinged | Mora |
Illustrated glossary (Schemes 1–7) of morphological terms used in the key and in the descriptions of Caesalpinioideae.
Gwilym P. Lewis10
Citation: Lewis GP (2024) 2. Tribe Ceratonieae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 62–69. https://doi.org/10.3897/phytokeys.240.101716
Ceratoniinae H.S. Irwin & Barneby in R.M. Polhill & P.H. Raven, Adv. Legume System. 1: 98. 1981.
Ceratonia L.
(4). Acrocarpus Wight ex Arn. (1 species), Arcoa Urb. (1), Ceratonia L. (2), Tetrapterocarpon Humbert (2).
Unarmed (rarely the stipules of juvenile leaves spinescent) shrubs or trees. Stipules spinescent (Arcoa), minute, caducous or lacking. Leaves pinnate or bipinnate with a terminal pinna. Inflorescences erect or pendent racemes or panicles, sometimes clustered on short shoots or ramiflorous. Flowers unisexual or bisexual, a short hypanthium usually present, and sometimes a prominent central pulviniform disk; sepals valvate to slightly imbricate; petals absent or 4–5 (6) per flower; androecium haplo- to diplostemonous, or occasionally stamens more than 2× petal number. Fruits dehiscent or indehiscent, linear to linear-oblong with an adaxial narrow wing, or oblong-ellipsoid, or laterally compressed and 4-winged (the wings in two unequal pairs), 1–several-seeded. Seeds compressed, usually separated by areas of pulpy mesocarp, pleurogram lacking.
Highly disjunct in Hispaniola (Arcoa), Madagascar (Tetrapterocarpon), tropical (South-)East Asia (Acrocarpus), and north-eastern Africa, the Mediterranean, Oman, Yemen and Somalia (Ceratonia).
The most inclusive crown clade containing Arcoa gonavensis Urb. and Ceratonia siliqua L., but not Umtiza listeriana Sim, Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig.
The tribal name Ceratonieae was first published by
Generic relationships in tribe Ceratonieae. Left part of figure shows complete genus-level Caesalpinioideae phylogeny with the Ceratonieae indicated with a red rectangle. Branch lengths are expressed in coalescent units and terminal branches were assigned an arbitrary uniform length for visual clarity. Support for relationships is based on fractions of supporting and conflicting gene trees: pie charts show gene tree support and conflict per node (blue representing supporting gene trees, green gene trees supporting the most common alternative topology, red gene trees supporting further alternative topologies, grey gene trees uninformative for this node), and numbers above pie charts are Internode Certainty All support values [both calculated with PhyParts (
Only two characters have been found to be shared by members of tribe Ceratonieae: bipinnate leaves with a single terminal pinna (Fig.
Flower, fruit, and vegetative characters of tribe Ceratonieae A Arcoa gonavensis Urb., foliage and fruits, Dominican Republic B Tetrapterocarpon geayi Humbert, fruits and part of bipinnate leaf, Madagascar (Du Puy M410) C–E Acrocarpus fraxinifolius Wight & Arn., India, cultivated tree C inflorescence D dehisced fruits and new flush foliage E part of a bipinnate leaf F–J Ceratonia siliqua L. F tree, Jordan, semi-desert near Petra G fruits, Israel, Mt. Scopus Botanical Gardens, Jerusalem H pistillate flowers, Crete I staminate flowers, Crete J staminate inflorescences, Israel, Jerusalem Botanical Gardens. Photo credits A F Jimenez R. B D Du Puy C–E https://efloraofindia.com/2011/02/01/acrocarpus-fraxinifolius/ E D Valke F, G, J O Fragman-Sapir H, I G Lewis.
Arcoa gonavensis Urb.
Shrub or small tree, with well-developed brachylasts. Stipules spinescent, caducous. Leaves pinnate when juvenile, bipinnate with a terminal pinna when mature, leaflets opposite to subopposite, sessile. Inflorescence a sparsely branching panicle of spikes arising from a thickened woody brachyblast. Flowers unisexual, staminate flowers with a prominent pistillode, pistillate flowers with staminoidia; sepals free to a very short hypanthium; petals 5 (6); a short cupuliform disk present centrally; stamens (or staminodes) number more than twice sepal number (12 or more per flower); pollen markedly irregular and coarsely reticulate, porate with prominent pores; ovary with an appressed rust-coloured indumentum, stigma terminal and capitate. Fruits oblong-ellipsoid, subterete, thick-walled, indehiscent, 1–few-seeded. Seeds ovate, compressed, surrounded by copious pulp, pleurogram lacking (Fig.
Unknown.
Arid tropical vegetation on limestone hills and cliffs.
Named for Count George von Arco (ca. 1903).
Unknown.
Originally placed by
Tetrapterocarpon geayi Humbert
Unarmed trees or shrubs, dioecious; brachyblasts absent. Stipules inconspicuous, minute and caducous. Leaves bipinnate, ending in a terminal pinna; leaflets alternate. Inflorescences axillary, spike-like racemes of small, subsessile flowers, usually aggregated into panicles. Flowers unisexual, actinomorphic, 4-merous (sepals and petals 4 per flower), greenish; sepals equal, petals equal, both whorls imbricate in young bud; androecium diplostemonous in staminate flowers, with one whorl of 4 fertile stamens, their filaments with an apical tuft of hairs behind the anthers, and one whorl of 4 hairy staminodes, lacking anthers; pollen with a scabrate-punctate sculpture pattern; pistillate flowers with a stipitate, compressed-fusiform ovary, stigma capitate and bilobed. Fruits membranous, indehiscent, compressed, 4-winged (in two unequal pairs), 1-seeded (Fig.
Unknown.
Seasonally dry tropical to xerophytic forest and thicket, on limestone, basalt, or sand.
From Greek, tetra- (= four), ptero- (= winged) and carpos (= fruit), the fruits have four dry, papery wings in unequal pairs.
Tetrapterocarpon geayi is used locally for carpentry, cart construction and to make charcoal (Du Puy and Rabevohitra 2002).
Although placed in the Dimorphandra group of the Caesalpinieae by
Du Puy and Rabevohitra (2002);
Acrocarpus fraxinifolius Wight & Arn.
Unarmed evergreen tree, bark pale grey, smooth, brachyblasts absent. Stipules not seen, presumed lacking, at least on mature leaves. Leaves large, bipinnate with a single terminal pinna, leaflets ovate-lanceolate, acuminate (Fig.
2n = 24 (
Tropical and subtropical broad-leaved rainforest and evergreen gallery forest.
From Greek, acro- (= summit or top) and carpos (= fruit), most probably alluding to the long-stipitate ovaries and fruits.
Timber of A. fraxinifolius (pink cedar tree) is used to make tea boxes, furniture, and plywood; the species is widely grown as an ornamental and is also used for fodder, gums, and bee forage (for honey) (
Acrocarpus was earlier placed in its own Acrocarpus group of tribe Caesalpinieae (
Siliqua Duhamel, Traité Arbr. Arbust. 2: 261. 1755, nom. superfl.
Ceratia Adans., Fam. Pl. 2: 319. 1763. Type not designated.
Ceratonia siliqua L.
Long-lived, evergreen, small to medium-sized trees (to ca. 12 m) (Fig.
2n = 24 (
Two species, one (C. siliqua) native to north-eastern Africa and the eastern Mediterranean (its native range uncertain due to its long history of cultivation), and Ceratonia oreothauma, with two distinct subspecies, one in Oman and Yemen and the other in the Somali Republic (Fig.
Distribution of Ceratonia based on quality-controlled digitised herbarium records. Note that many distribution points observed here are from cultivated Ceratonia specimens. The true native range of the genus Ceratonia is likely to be more restricted than depicted on the map. See Suppl. material
Mediterranean scrubland (dry hillsides in garigue and coastal and submaritime maquis) (C. siliqua); rocky limestone slopes and gullies (C. oreothauma); 0–2000 m.
From ‘Ceratonia’, ‘ceronia’ or ‘ceratea’ (Greek names for C. siliqua), or possibly from ceras (Greek = horn) referring to the long, curved pods of C. siliqua.
Ceratonia siliqua is widely cultivated in the Mediterranean for forest-forage and its nutritious fruits; the Romans were harvesting the species as early as 79 AD. Carob seeds are said to be the original carat used as a standard weight by jewellers. Ceratonia siliqua is also used for wood, as a chocolate and coffee substitute, and occasionally to make alcohol. Carob seed gum is used in foods, cosmetics, medicines, photographic film emulsions, adhesives, paints, inks, and polishes (
The two Ceratonia species are differentiated, amongst other characters, by the pollen type, tetracolporate in C. siliqua and tricolporate in C. oreothauma (
Patrick S. Herendeen21, Anne Bruneau1
Citation: Herendeen PS, Bruneau A (2024) 3. Tribe Gleditsieae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 70–77. https://doi.org/10.3897/phytokeys.240.101716
Gleditsia J. Clayton
(3). Gleditsia J. Clayton (13 species), Gymnocladus Lam. (6), Umtiza Sim (1).
Deciduous or evergreen trees, with simple or branched thorns or unarmed; branches with or without lateral brachyblasts. Stipules inconspicuous (Gleditsia), foliaceous, caducous (Gymnocladus), or absent (Umtiza). Leaves bipinnate, pinnate, or intermediate with one or more pinnae replaced by a leaflet. Inflorescences panicles or racemes. Flowers regular, bisexual (Umtiza), or unisexual and then androdioecious or dioecious (Gleditsia, Gymnocladus), small, white or greenish to violet, mostly 5-merous; hypanthium short to elongate; calyx gamosepalous; petals small; stamens (6) 10, free; pollen in tricolporate monads, exine perforate, reticulate; ovary sessile or stipitate. Fruit compressed or turgid, papery, leathery or woody, dehiscent, tardily dehiscent or indehiscent, sometimes with pulpy interior, with few to 25 (40) seeds. Seeds compressed to subterete, orbicular or ovoid-elliptic.
Temperate to subtropical North America, South America, Asia, and South Africa.
The most inclusive crown clade containing Umtiza listeriana Sim and Gymnocladus dioicus (L.) K. Koch, but not Ceratonia siliqua L., Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig.
Tribe Gleditsieae was first named by
Umtiza listeriana Sim
Small evergreen trees to 12 m or shrubs, branches armed with stout spines that are frequently branched; branches with lateral brachyblasts (Fig.
Gleditsieae, diversity of vegetative morphology and fruits A–C Umtiza listeriana Sim A small tree B branch with pinnately compound leaves and inflorescence C branch with thorn at node D Gymnocladus assamicus Kanjilal ex P.C. Kanjilal branch with bipinnate leaves and inflorescence E Gleditsia japonica Miq., bipinnate leaf with several pinnae replaced by single leaflets, cultivated at the Royal Botanic Gardens, Kew F Gleditsia amorphoides (Griseb.) Taub., branched thorns G Umtiza listeriana dehisced fruits H Gymnocladus dioicus L., fruit I Gleditsia triacanthos L., fruits. Photo credits A–C, G SAplants, Wikimedia Commons (CC-BY-SA 4.0) D B Choudhury E, F GP Lewis H Chicago Botanic Garden (photographer unknown) I A Schnabel.
Gleditsieae, floral diversity A, B Umtiza listeriana Sim A branch with inflorescence of hermaphrodite flowers B close-up of flowers C, D Gymnocladus assamicus Kanjilal ex P.C. Kanjilal C inflorescence of hermaphrodite flowers D dissected hermaphrodite flower E Gymnocladus chinensis Baill., inflorescence, cultivated at the US National Arboretum F Gymnocladus dioicus L., cultivated at Chicago Botanic Garden, upright inflorescence of hermaphrodite flowers G branch with leaves and inflorescence H Gleditsia triacanthos L., inflorescence of staminate flowers I Gleditsia caspica Desf., inflorescence, cultivated at the Royal Botanic Gardens, Kew. Photo credits A, B SAplants, Wikimedia Commons (CC-BY-SA 4.0) C, D Murata E PS Herendeen F, G R Carlson H Royal Botanic Gardens, Kew (photographer unknown) I M Svanderlik.
Unknown.
Monospecific (U. listeriana), from Eastern Cape region, South Africa, primarily along the Buffalo River between East London and King William’s Town (Fig.
Subtropical dry forest, bushland and thicket.
Derived from ‘umthiza’, the African vernacular name for the single species, U. listeriana.
Used locally for medicine and has alleged magical properties, witchdoctors have used sticks of the tree as healing wands; the wood was once used to house propeller shafts in small boats because its oiliness provided constant lubrication (
The genus is monospecific with Umtiza listeriana endemic to the Eastern Cape region of South Africa where the species is endangered and protected. The Umtiza Nature Reserve is named for this species.
Gymnocladus canadensis Lam., nom. illeg. [= Gymnocladus dioicus (L.) Koch (≡ Guilandina dioica L.; vide
Large to medium size deciduous trees, unarmed, stems often stout, androdioecious (individuals with either staminate flowers or hermaphrodite flowers) or dioecious. Stipules small, inconspicuous. Leaves even- or irregularly-bipinnate, pinnae 3–10 pairs, proximal-most sometimes reduced to single leaflets, opposite, subopposite or alternate; leaflets alternate, 6–30 per pinna, lamina margin entire (Figs
2n = 28 (
Six species in eastern and central North America, China, Vietnam, India, Burma, Thailand (Fig.
Temperate bottomlands and riparian woodlands, tropical and subtropical montane woodlands and wooded hillsides (
The genus name derives from the Greek (= naked branch) because G. dioicus is one of the latest trees to leaf out in the spring.
Multiple species of Gymnocladus are used by people in different parts of the world (
The taxonomy of the species of Gymnocladus is well established, but the monophyly of the genus has not been adequately tested. Only G. dioicus and/or G. chinensis have been included in phylogenetic analyses (
Melilobus Mitch., Diss. Gen. Pl.: 37. 1769. Type: Melilobus heterophyla Raf. [= Gleditsia triacanthos L.]
Asacara Raf., Neogenyton: 2. 1825. Type: Asacara aquatica (Marshall) Raf. [≡ Gleditsia aquatica Marshall]
Garugandra Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 96. 1879. Type: Garugandra amorphoides Griseb. [≡ Gleditsia amorphoides (Griseb.) Taub.]
Caesalpiniodes Kuntze, Rev. Gen. 1: 166. 1891. Type: Caesalpiniodes triacanthum (L.) Kuntze [≡ Gleditsia triacanthos L.]
Pogocybe Pierre, Fl. Cochinch.: t. 392B. 1899. Type: Pogocybe entadoides Pierre. [= Gleditsia australis Hemsl.]
Gleditsia triacanthos L.
Large to medium size deciduous trees (G. saxatilis was described as evergreen), usually well-armed with simple or compound thorns on trunk and branches (Fig.
2n = 28 (
Thirteen species and 1 hybrid taxon. Eastern and central North America to northern Mexico; Bolivia to northern Argentina; China, Tibet, Vietnam, Korea, Japan, Malaysia, Philippines, Azerbaijan (Fig.
Species are generally found in temperate and subtropical woodlands and thickets on sandy and rocky slopes, and lowland wet forest and swamp forest (
The genus name commemorates Johann Gottlieb Gleditsch, director of the Berlin Botanical Garden, who died in 1786.
Multiple species of Gleditsia are used by people in different parts of the world (
Phylogenetic analyses indicate that the genus is monophyletic (
Luciano Paganucci de Queiroz2, Filipe Gomes Oliveira2
Citation: Queiroz LP, Oliveira FG (2024) I4. Tribe Pterogyneae In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 78–82. https://doi.org/10.3897/phytokeys.240.101716
Differing from all other tribes of subfamily Caesalpinioideae by the combination of imparipinnate leaves with a well-formed rachis extension, alternate leaflets, compact catkin-like racemes, flowers small, polygamous and regular, the ovary with a marginal wing, style subterminal, fruit a samara with the basal seed chamber with a small subterminal beak (style remnant), and chromosome number 2n = 20.
Pterogyne Tul. (designated here).
Tribe Pterogyneae includes only the genus Pterogyne with one species.
The most inclusive crown clade containing Pterogyne nitens Tul., but not Ceratonia siliqua L., Umtiza listeriana Sim or Cassia fistula L.
The tribe Pterogyneae is being proposed here to include the single genus Pterogyne. This genus was previously ascribed to the tribe Cynometreae (currently subfamily Detarioideae) by
Contrasting phylogenetic positions for Pterogyne as supported by different analyses A sister to all Caesalpinioideae, except Ceratonieae and Gleditsiae (ca. 1500 nuclear genes;
The uncertain phylogenetic position of Pterogyne probably reflects the short diversification time between the divergence of the four distinct lineages, Pterogyne, Cassieae, Caesalpinieae, and the remaining Caesalpinioideae, in the Paleocene, with Pterogyne representing a long branch diverging from those other lineages between 57 Ma and 60 Ma (
The recognition of the monogeneric (and monospecific) tribe Pterogyneae is additionally supported by its unique combination of morphological features, as highlighted in the diagnosis above. This isolated evolutionary position is also corroborated by a chromosome number of 2n = 20 for Pterogyne nitens, which is cytologically unique in subfamily Caesalpinioideae (
Pterogyne nitens Tul.
Medium to tall trees, mostly 4–8 m but sometimes to 20 m, unarmed (Fig.
Pterogyne nitens Tul., the only species of tribe Pterogyneae A cultivated tree in Brasília (Brazil) B flowering branch showing the foliage and inflorescences; note the alternate leaflets; the insets show an expanding leaf (left) highlighting the tiny caducous stipules (arrow) and a stipel at the leaflet attachment (right) C inflorescences with a visiting Syrphidae fly (Diptera) D part of the spicate raceme with some flowers removed E close-up of flowers F samaras, the inset showing the subterminal style remnant at the top of the seed chamber (arrows) G samara seed chamber opened to show the seed. Photo credits A–G RT Queiroz https://rubens-plantasdobrasil.blogspot.com/.
2n = 20 (
Monospecific (P. nitens), distributed across north-eastern, eastern and central Brazil, northern Argentina and Paraguay and south-eastern Bolivia (Fig.
Pterogyne nitens is a tree of the South American tropical and subtropical seasonally dry forests of north-eastern Brazil (Caatinga), central Brazil (Cerrado) and chaquean forests (northern Argentina, Paraguay and southern Bolivia), also occurring in eastern Brazilian semi-deciduous forests of the Mata Atlântica. It occurs mostly as a pioneer tree colonizing degraded areas. It flowers in a very short period and the flowers are visited by a wide range of small insects (
From pteros (Greek: πτέρυξ = wing) and gynos- (Greek: γυνή = woman, gynoecium) referring to the ovary provided with a narrow wing along one margin.
Pterogyne nitens is planted as an urban tree (Fig.
The phylogenetic position of the genus is still uncertain and conflicting among different analyses (see Fig.
The compact catkin-like racemes with rather small flowers and the samara fruits are similar to those found in the African-Asiatic genus Pterolobium R. Br. ex Wight & Arn. of tribe Caesalpinieae (
Juliana Gastaldello Rando37, Matheus Martins Teixeira Cota2, Alexandre Gibau de Lima8,28, Roseli Lopes da Costa Bortoluzzi6, Brigitte Marazzi30, Adilva de Souza Conceição11
Citation: Rando JG, Cota MMT, Lima AG, Bortolozzi RLC, Marazzi B, Conceição AS (2024) 5. Tribe Cassieae In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 83–102. https://doi.org/10.3897/phytokeys.240.101716
Cassiaceae Vest, Anleit. Stud. Bot.: 270, 291. 1818. Type: Cassia L.
Irregulares Bronn, Form. Pl. Legumin.: 13. 1822. Type: Cassia L.
Cassiinae Wight & Arn., Prodr. Fl. Ind. Orient.: 280. 1834. Type: Cassia L.
Cassioideae Burmeist., Handb. Naturgesch.: 319. 1837. Type: Cassia L.
Melanoxyleae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 254. 1943. Type: Melanoxylum Schott
Cassia L.
(7). Batesia Spruce ex Benth. (1 species), Cassia L. (39), Chamaecrista (L.) Moench (361), Melanoxylum Schott (1), Recordoxylon Ducke (3), Senna Mill. (287), Vouacapoua Aubl. (3).
Trees, shrubs, subshrubs or vines. Stipules diverse in shape and size, persistent or caducous. Leaves bifoliolate, paripinnate and/or imparipinnate, extrafloral nectaries present in Batesia and Vouacapoua, and in several species of Chamaecrista and Senna, on the petiole and/or leaf rachis, and/or peduncles of inflorescences; leaflets opposite, subopposite or alternate. Inflorescences racemes or panicles, terminal, axillary and sometimes ramiflorous. Flowers hypogynous or perigynous, bilaterally symmetrical, radially symmetrical or asymmetrical; sepals 5, free; petals 5, free, equal, subequal or differentiated; stamens 2–10, free, homomorphic or heteromorphic, anthers poricidal (Cassia, Chamaecrista and Senna) or longitudinally dehiscent; pollen unknown for most genera, 3-colporate with long apertures in Chamaecrista and Senna; ovary sessile or stipitate. Fruit a legume, follicle, indehiscent or dehiscent, with valves opening elastically or not. Seeds mostly compressed, exarillate, variable in shape and colour.
The tribe has a pantropical distribution in wet forests, seasonally dry forests and woodlands, savannas and deserts, a few species extending to temperate areas. The highest diversity of the richest genera (Chamaecrista and Senna) occurs in the Neotropical region.
The most inclusive crown clade containing Cassia fistula L. and Melanoxylum brauna Schott, but not Ceratonia siliqua L., Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig.
Tribe Cassieae sensu
Cassia, Chamaecrista and Senna are the largest genera in the tribe. Because of similarity in floral morphology, they were all treated under the single genus Cassia until they were segregated by
Generic relationships in tribe Cassieae. Two most likely positions of the unsampled genus Vouacapoua are indicated with dashed lines: 1) as sister to all Cassieae [following
Examples of variation in stamens and pistil among Cassieae genera A Cassia moschata Kunth (Rando et al. 1184) B Senna macranthera (DC. ex Collad.) H.S. Irwin & Barneby (Rando et al. 272) C Chamaecrista bahiae (H.S. Irwin) H.S. Irwin & Barneby (Rando et al. 1214) D Melanoxylum brauna Schott (Cardoso et al. 2439) E Recordoxylon speciosum (Benoist) Gazel ex Barneby (Pereira-Silva et al. 15631) F Vouacapoua americana Aubl (Nascimento 245). G Batesia floribunda Spruce ex Benth (based on photographs of Projeto Flora Reserva Ducke, INPA/DFID, comm. Mike Hopkins). Scale: 1 cm. Drawn by Najla M.B. Scheidegger / @ilustre.nt.
The position of Vouacapoua has proved more difficult to resolve. Even though not sampled by
Cassieae is a morphologically heterogeneous group, with only a few features shared among all or nearly all genera. All Cassieae have once-pinnate or bifoliolate leaves, most species have yellow petals, with only a few having red, orange, pink or white ones. Cassia, Senna and Chamaecrista are characterised by poricidal anthers (Fig.
Vouacapoua americana Aubl.
Unarmed trees. Stipules not observed. Leaves spiral, imparipinnate; petiole terete; extrafloral nectaries on pulvinus or between all pairs of leaflets, sessile, secretory surface convex, sometimes absent; leaflets 7–11, opposite. Inflorescence a panicle; bract 1, caducous, bracteoles 2, caducous. Flowers perigynous, radially symmetrical; hypanthium campanulate; sepals 5, free; petals 5, yellow, free; stamens 10, free, filaments glabrous, anthers longitudinally dehiscent; pollen unknown; ovary shortly stipitate, attached to the base of the hypanthium. Fruit obovoid to ellipsoid drupaceous legume, rugose and velutinous, one (rarely 2–3)-seeded, swollen over seed, dehiscent. Seeds globose to obovoid, with a brownish, smooth and glossy testa.
Unknown.
Three species, V. americana, V. macropetala Sandwith and V. pallidior Ducke, restricted to northern South America in the wet Amazonian forests of Brazil, British Guiana, French Guiana and Suriname (
Examples of Batesia, Melanoxylum, Recordoxylon and Vouacapoua diversity A Batesia floribunda Spruce ex Benth. extrafloral nectary, petiolules and part of leaflets (Cota 1158) B, C Vouacapoua americana Aubl. (Cardoso et al. 3450) B leaves and immature fruit C mature fruits and seeds on the ground D, E Melanoxylum brauna Schott (Cardoso et al. 2439) D inflorescence, buds and flowers E flower F–H Recordoxylon speciosum (Benoist) Gazel ex Barneby (Pereira-Silva et al. 15631) F trunk of mature individual G flowers and immature fruit H flower. Photo credits A–E D Cardoso F–H G Pereira-Silva.
The genus is known only on well-drained soils from lowland tropical rainforests of the Amazon basin (“terra firme”).
The timber of V. americana is used in civil and naval constructions and furniture (
The name originated with the Galibi (also called Kalinã), indigenous people from northern South America. “Vouacapoua” or “Voicapou” are Galibi names for Vouacapoua americana (
Vouacapoua is characterised by a terete petiole, extrafloral nectaries on the pulvinus or between the pairs of leaflets, glabrous stamen filaments (Fig.
Seeds of V. americana are dispersed by small rodents (Myoprocta exilis and Dasyprocta leporina) that bury the fruits at short distances from the source tree (
Batesia floribunda Spruce ex Benth.
Unarmed trees. Stipules not observed. Leaves spiral, imparipinnate or, rarely, paripinnate; extrafloral nectaries present between the proximal pair of leaflets, sometimes also between the distal ones, secretory surface flat and disc shaped; petiole narrowly winged; leaflets 9–13, opposite. Inflorescence a panicle; bract 1, caducous, bracteoles 2, caducous. Flowers perigynous, radially symmetrical; hypanthium campanulate; sepals 5, free; petals 5, yellow, free; stamens 10, homomorphic, filaments villous at the base, anthers longitudinally dehiscent; pollen unknown; ovary shortly stipitate, attached to the base of the hypanthium. Fruit an ellipsoid to oblong-obovate follicle, turgid and slightly compressed, the valves fleshy-coriaceus with strongly raised veins, dehiscent. Seeds globose with a reddish and smooth testa.
Unknown.
Monospecific (B. floribunda), occurring in northern and north-western South America in the wet Amazonian forests of Brazil, French Guiana, Colombia and Peru (Fig.
The genus is known only on well-drained soils from lowland tropical rainforests of the Amazon basin (‘terra firme”).
The timber of B. floribunda is used in the construction of fine furniture (
Named after the English naturalist Henry Walter Bates, who explored the Amazon rainforests with A.R. Wallace in the 19th century (
Batesia is characterised by the presence of winged leaf petioles, extrafloral nectaries between the leaflet pairs, and mainly by the fruit follicle with bright red seeds. In the original description, Batesia was described with three bracteoles on the pedicels, but observation of herbarium specimens lead us to the conclusion that there are two bracteoles and one bract. Herbarium specimens (vegetative or with fruits and seeds) of B. floribunda are often misidentified as Ormosia Jacks. (Papilionoideae). However, Ormosia displays terete leaf petioles, lacks extrafloral nectaries, and has a distinct fruit. The relationship of Batesia, in particular to Chamaecrista and a clade that groups Recordoxylon and Melanoxylum, remains poorly understood and requires further study (Fig.
Melanoxylon Schott, Syst. Veg., ed. 16 [Sprengel] 4(2): Cur. Post. 406. 1827, orth. var.
Perittium Vogel, 1837. Linnaea 11: 408. 1837. Type: Perittium ferrugineum Vogel [= Melanoxylum brauna Schott]
Melanoxylum brauna Schott
Unarmed trees, bark thick. Stipules caducous. Leaves spiral, imparipinnate; extrafloral nectaries absent; leaflets 11–21, opposite to subopposite. Inflorescence a panicle; bracts and bracteoles caducous. Flowers perigynous, bilaterally symmetrical; hypanthium infundibuliform; sepals 5, free; petals 5, yellow, free, clawed, glabrous; stamens 10, slightly heteromorphic, filaments ferruginous tomentose at the base, anthers longitudinally dehiscent; pollen unknown; ovary 11–13-ovulate, ferruginous tomentose. Fruit an oblong, slightly curved, compressed legume, dehiscing through both margins, valves with raised transverse ribs, tomentose, endocarp breaking up into one seeded transversely oblong envelopes. Seeds oblong-depressed, with smooth, opaque and dark reddish testa.
Unknown.
Monospecific (M. brauna), restricted to Brazil, occurring predominantly along the eastern Brazilian coast, but entering the interior in drier vegetations (Fig.
Melanoxylum brauna occurs preferentially in wet habitats, mostly in tropical rainforests; its occurrence in drier Brazilian vegetation (seasonally deciduous and semi-deciduous Forests) is apparently associated with wetter areas within these ecosystems.
The timber of M. brauna is largely used in the construction of fine furniture, and the bark is a source of tannin for medicinal purposes (
Melano + xylon from Greek meaning “black” and “wood”, respectively. The name is related to the black heartwood of the plant (
Melanoxylum is characterised by its imparipinnate leaves, and by its dense inflorescence and characteristic fruit, an oblong legume, slightly curved with articulate endocarp, breaking up into transversely oblong parts.
Recordoxylon amazonicum (Ducke) Ducke [≡ Melanoxylon amazonicum Ducke (= Recordoxylon speciosum (Benoist) Gazel ex Barneby)]
Unarmed trees. Stipules not observed. Leaves spiral, imparipinnate; leaflets 7–15, alternate to opposite; extrafloral nectaries absent. Inflorescence a panicle; bract 1, caducous, bracteoles 2, caducous. Flowers perigynous, bilaterally symmetrical, hypanthium campanulate; sepals 5, free; petals 5, the innermost petal yellow with white spot at the base, the others yellow, free, clawed; stamens 10, slightly heteromorphic, filaments glabrous, anthers longitudinally dehiscent; pollen unknown; ovary shortly stipitate, attached to the base of the hypanthium. Fruit oblong, straight, compressed legume, with a longitudinal rib close to the superior margin, dehiscent. Seeds elliptic-depressed, with a brownish and rugose testa.
Unknown.
Three species, R. pulcherrimum Barneby, R. speciosum, and R. stenopetalum Ducke, restricted to northern South America in the wet Amazonian forests of Brazil, Guyana, French Guiana and Venezuela (
The genus is known from lowland tropical rainforests of the Amazon basin, on well-drained soils (“terra firme”) to poorly drained soils (“igapó” forest).
Named after the American botanist Samuel James Record, an important wood anatomist who observed some wood structures in R. speciosum (as Melanoxylum amazonicum;
Recordoxylon is characterised by its bilaterally symmetrical flowers, clawed petals, glabrous stamen filaments, longitudinally dehiscent anthers (Fig.
Cassia [infragen. unranked] Chamaecrista L., Sp. Pl. 1: 379. 1753. Type: Cassia chamaecrista L., nom. utique rejic. [≡ Cassia fasciculata Michx. (≡ Chamaecrista fasciculata (Michx.) Greene)]
Cassia sect. Chamaecrista
(L.) DC., Hist. Nat. Méd. Casses 24, 118.1816. Lectotype (designated by
Sooja Siebold, Verh. Batav. Genootsch. Kunst. 12: 56. 1830. Type: Sooja nomame Siebold, nom. inval. (nom. nud.) [≡ Cassia mimosoides var. nomame Makino (≡ Chamaecrista nomame (Makino) H. Ohashi)]
Disterepta Raf., Sylva Tellur.: 126. 1838. Type: Disterepta pilosa (L.) Raf. [≡ Cassia pilosa L. (≡ Chamaecrista pilosa (L.) Greene)]
Hepteireca Raf., Sylva Tellur.: 126. 1838. Type: Hepteireca glandulosa (L.) Raf. [≡ Cassia glandulosa L. (≡ Chamaecrista glandulosa (L.) Greene)]
Dialanthera Raf., Sylva Tellur.: 127. 1838. Type: Dialanthera glandulosa (L.) Raf. [≡ Cassia glandulosa L. (≡ Chamaecrista glandulosa (L.) Greene)]
Xamacrista Raf., Sylva Tellur.: 127. 1838. Type: Xamacrista trifolia Raf. [= Cassia chamaecrista L. (≡ Chamaecrista fasciculata (Michx.) Greene)]
Nictitella Raf., Sylva Tellur.: 128. 1838. Lectotype (designated by Irwin & Barneby, 1982): Nictitella amena Raf. [= Cassia nictitans L. (≡ Chamaecrista nictitans (L.) Moench)]
Ophiocaulon Raf., Sylva Tellur.: 129. 1838. Lectotype (designated by Irwin & Barneby, 1982): Ophiocaulon serpens (L.) Raf. [≡ Cassia serpens L. (≡ Chamaecrista serpens (L.) Greene)]
Cassia subg. Lasiorhegma Vogel ex Benth., Fl. Bras. 15(2): 129. 1870. Type not designated.
Chamaecrista nictitans (L.) Moench [≡ Cassia nictitans L.]
Trees, treelets, shrubs and subshrubs, lacking spines or prickles. Stipules diverse in shape and size, persistent or caducous. Leaves distichous or spiral, bifoliolate or paripinnate; extrafloral nectaries when present on petiole, generally on the rachis between the pairs of leaflets or in the axis of the inflorescence, sessile or stipitate, the secretory surface concave, rarely convex; leaflets 1–65 pairs. Inflorescence a fascicle, raceme or panicle; bract 1, caducous or persistent, bracteoles 2, alternate, located at mid-length or slightly above the pedicels, persistent. Flowers hypogynous, asymmetrical, hypanthium absent; sepals 5, free; petals 5, free, yellow, or yellow with red base, sometimes red, orange or pink; stamens 5–10, homomorphic, filaments glabrous, anthers dehiscent by apical pores, pubescent laterally, rarely with the indumentum covering the entire anther; pollen subprolate to prolate, syncopate, fused at the poles; ovary stipitate. Fruit a compressed legume, valves papyraceous or coriaceous, elastically dehiscent through both margins, becoming twisted after dehiscence. Seeds variable in shape and colour.
Haploid numbers n = 7, 8, 14, 16, 24 (
Chamaecrista species typically occur in open environments. Although several species are widespread, such as C. rotundifolia (Pers.) Greene, C. mimosoides (L.) Greene and C. flexuosa (L.) Greene, a high diversity is concentrated in Brazilian savannas and in the “campos rupestres’’ vegetation (
Examples of Chamaecrista diversity A Chamaecrista xinguensis (Ducke) H.S. Irwin & Barneby trunk of mature individual (Rando et al. 1208) B C. compitalis (H.S. Irwin & Barneby) H.S. Irwin & Barneby base of trunk of mature individual (Rando et al. 1364) C C. ensiformis (Vell.) H.S. Irwin & Barneby flowering branch (Rando & Cota 1366) D C. flexuosa (L.) Greene flower, leaves in background E C. desvauxii var. latistipula (Benth.) G.P. Lewis, branch with flowers and fruit F C. ramosa var. curvifolia (Vogel) G.P. Lewis branches and flower G C. distichoclada (Benth.) H.S. Irwin & Barneby flowering branch (Rando et al. 1230) H C. lineata (Sw.) Greene leaves and flower (Rando 964) I C. andromedea (Mart. ex Benth.) H.S. Irwin & Barneby branch with leaves and flowers (Rando et al. 1251) J C. vauthieri (Benth.) H.S. Irwin & Barneby flowering branches (Cardoso et al. 4096) K C. ochnacea var. purpurascens (Benth.) H.S. Irwin & Barneby inflorescence and leaves L C. scabra (Pohl ex Benth.) H.S. Irwin & Barneby leaves and inflorescence (Rando et al. 1266). Photo credits A, C, G–I JG Rando B JG Jardim D–F, K H Moreira J D Cardoso L MF Simon.
Some species are used in traditional African medicine. For example, C. absus (L.) H.S. Irwin & Barneby is used as a purgative, for treating wounds and sores, and also against syphilis (
A composite name from the Greek Chamae (= small, of little growth), and the Latin crista, referring to the crest (
The largest genus of the tribe can be easily recognised by a set of features: the presence of two bracteoles on the pedicels, stamens generally homomorphic, poricidal anthers (Fig.
Cathartocarpus Pers., Syn. Pl. 1: 459. 1805. Type: Cathartocarpus fistula (L.) Pers. [≡ Cassia fistula L.]
Bactyrilobium Willd., Enum. Pl.: 439. 1809. Type: Bactyrilobium fistula (L.) Willd. [≡ Cassia fistula L.]
Cassiana Raf., Amer. Monthly Mag. & Crit. Rev. 1: 266. 1818. Type not designated.
Mac-leayiaMontrouz., Mém. Acad. Imp. Sci. Lyon, Sect. Sci., sér. 2, 10: 198. 1860. Type: Mac-leayia multiflora Montrouz. [= Cassia artensis Beauvis.]
Cassia fistula L.
Trees or shrubs, lacking spines or prickles. Stipules 0.1–1.0 cm, in general caducous. Leaves distichous or spiral, paripinnate; extrafloral nectaries absent; leaflets 2–25 pairs, opposite. Inflorescence an axillary raceme; bract 1, caducous or persistent, bracteoles 2, at the base of pedicels, usually caducous. Flowers hypogynous, bilaterally symmetrical, hypanthium solid, turbinate or conic; sepals 5, free, reflexed at anthesis; petals 5, yellow or pink, less often red, white or mixed, the median petal a different colour from the rest; stamens 10, heteromorphic, 3 with long sigmoidal filaments and longitudinally dehiscent anthers, 7 adaxial ones varying in length, organised in groups of 5+2 or 4+3, anthers with basal poricidal dehiscence; pollen unknown; ovary shortly stipitate, attached to the base of the hypanthium. Fruit a linear-oblong, often long, cylindrical or quadrangular, woody, indehiscent legume (except in C. hintoni Sandwith). Seeds obovoid to ellipsoid, compressed, smooth and glossy with castaneous or brownish testa.
Examples of Cassia diversity A Cassia moschata Kunth inflorescence B C. ferruginea (Schrad.) Schrad. ex DC. branch showing the spiraled leaves and an inflorescence (Rando et al. 125) C C. fistula L. branch with leaves and an inflorescence D C. moschata branches with leaves, an inflorescence and fruits (left corner) in the background (Rando et al. 1184) E C. javanica L. branches with leaves and an inflorescence F C. grandis L.f. flowers. Photo credits A–D JG Rando E D Gissi F D Cardoso.
Haploid numbers n = 12, 13, 14 (
Thirty-nine species (
Cassia occurs preferentially in tropical rainforests; a few species extend to or occur exclusively in temperate areas, such as C. ferruginea Schrad. ex DC. and C. leptophylla Vogel (
Many uses are reported for several Cassia species as medicinal plants (
Derived from the ancient Greek name casia for the aromatic and fragrant plants (
Cassia is characterised by showy dense inflorescences, flowers with an androecium with 3 long sigmoidal stamens with longitudinally dehiscent anthers and 7 adaxial stamens varying in length, organised in groups of 5+2 or 4+3, and these poricidal at the base of the thecae (instead of apically as in Chamaecrista and Senna) (Fig.
Chamaecassia Link, Handbuch 2: 139. 1831. Type: Chamaecassia laevigata (Willd.) Link [≡ Cassia laevigata Willd. (= Senna septemtrionalis (Viv.) H.S. Irwin & Barneby)]
Chamaefistula (DC. ex Collad.) G. Don, Gen. Hist. 2: 106. 1832. Type: Chamaefistula corymbosa (Lam.) G. Don [≡ Cassia corymbosa Lam. (≡ Senna corymbosa (Lam.) H.S. Irwin & Barneby)]
Adipera Raf., Sylva Tellur.: 129. 1838. Type: Adipera herbertiana (Lindl.) Raf. [= Senna × floribunda (Cav.) H.S. Irwin & Barneby]
Diallobus Raf., Sylva Tellur.: 128. 1838. Type: Diallobus tora (L.) Raf. [≡ Cassia tora L. (≡ Senna tora (L.) Roxb.)]
Ditremexa Raf., Sylva Tellur.: 127. 1838. Lectotype (designated by Britton and Wilson 1924): Ditremexa occidentalis (L.) Britton & Rose [≡ Cassia occidentalis L. (≡ Senna occidentalis (L.) Link)]
Emelista Raf., Sylva Tellur.: 127. 1838. Type: Emelista obtusifolia (L.) Raf. [≡ Cassia obtusifolia L. (≡ Senna obtusifolia (L.) H.S. Irwin & Barneby)]
Herpetica Raf., Sylva Tellur.: 123. 1838. Type: Herpetica alata (L.) Raf. [≡ Cassia alata L. (≡ Senna alata (L.) Roxb.)]
Isandrina Raf., Sylva Tellur.: 126. 1838. Type: Isandrina arborescens Raf. [= Senna atomaria (L.) H.S. Irwin & Barneby]
Panisia Raf., Sylva Tellur.: 128. 1838. Type: Panisia biflora (L.) Raf. [≡ Cassiabiflora L., but typus not established, probably either = Senna pallida (Vahl) H.S. Irwin & Barneby sensu lato or S. angustisiliqua (Lam.) H.S. Irwin & Barneby]
Peiranisia Raf., Sylva Tellur.: 127. 1838. Type: Peiranisia aversiflora (Herb.) Raf. [≡ Cassia aversiflora Herb. (≡ Senna aversiflora (Herb.) H.S. Irwin & Barneby)]
Scolodia Raf., Sylva Tellur.: 128. 1838. Type: Scolodia viminea (L.) Raf. [≡ Cassia viminea L. (≡ Senna viminea (L.) H.S. Irwin & Barneby)]
Cassia subg. Senna (Mill.) Benth., Fl. Bras. 15(2): 83, 96. 1870. Type not cited, but inferred as Senna alexandrina Mill.
Chamaesenna
Raf. ex Pittier, Arb. Arbus. Orden Legum.: 130. 1928. Lectotype (designated by
Cowellocassia Britton, N. Amer. Fl. 23: 251. 1930. Type: Cowellocassia scleroxyla (Britton) Britton [≡ Cassia scleroxyla Britton (= Senna domingensis (Spreng.) H.S. Irwin & Barneby)]
Earleocassia Britton, N. Amer. Fl. 23: 247. 1930. Type: Earleocassia roemeriana (Scheele) Britton [≡ Cassia roemeriana Scheele (≡ Senna roemeriana (Scheele) H.S. Irwin & Barneby)]
Echinocassia Britton & Rose, N. Amer. Fl. 23: 251. 1930. Type: Echinocassia aculeata (Pohl ex Benth.) Britton & Rose [≡ Cassia aculeata Pohl ex Benth. (≡ Senna aculeata (Pohl ex Benth.) H.S. Irwin & Barneby)]
Desmodiocassia Britton & Rose, N. Amer. Fl. 23: 244. 1930. Type: Desmodiocassia villosa (Mill.) Britton & Rose [≡ Cassia villosa Mill. (≡ Senna villosa (Mill.) H.S. Irwin & Barneby]
Gaumerocassia Britton, N. Amer. Fl. 23: 252. 1930. Type: Gaumerocassia peralteana (Kunth) Britton [≡ Cassia peralteana Kunth (≡ Senna peralteana (Kunth) H.S. Irwin & Barneby)]
Leonocassia Britton, N. Amer. Fl. 23: 268. 1930. Type: Leonocassia stenophylla (Benth.) Britton [≡ Cassia stenophylla Benth. (≡ Senna stenophylla (Benth.) H.S. Irwin & Barneby)]
Palmerocassia Britton, N. Amer. Fl. 23: 253. 1930. Type: Palmerocassia wislizeni (A. Gray) Britton [≡ Cassia wislizeni A. Gray (≡ Senna wislizeni (A. Gray) H.S. Irwin & Barneby)]
Phragmocassia Britton & Rose, N. Amer. Fl. 23: 245. 1930. Type: Phragmocassia skinneri (Benth.) Britton & Rose [≡ Cassia skinneri Benth. (≡ Senna skinneri (Benth.) H.S. Irwin & Barneby)]
Pseudocassia
Britton & Rose, N. Amer. Fl. 23: 230. 1930. Lectotype (designated by
Pterocassia Britton & Rose, N. Amer. Fl. 23: 243. 1930. Type: Pterocassia galeottiana (M. Martens) Britton & Rose [≡ Cassia galeottiana M. Martens (≡ Senna galeottiana (M. Martens) H.S. Irwin & Barneby)]
Tharpia Britton & Rose, N. Amer. Fl. 23: 246. 1930. Type: Tharpia pumilio (A. Gray) Britton & Rose [≡ Cassia pumilio A. Gray (≡ Senna pumilio (A. Gray) H.S. Irwin & Barneby)]
Vogelocassia Britton, N. Amer. Fl. 23: 258. 1930. Type: Vogelocassia leiophylla (Vogel) Britton [≡ Cassia leiophylla Vogel (≡ Senna leiophylla (Vogel) H.S. Irwin & Barneby)]
Xerocassia Britton & Rose, N. Amer. Fl. 23: 246. 1930. Type: Xerocassia armata (S. Watson) Britton & Rose [≡ Cassia armata S.Watson (≡ Senna armata (S. Watson) H.S. Irwin & Barneby)]
Senna alexandrina Mill.
Trees, treelets, erect or scandent shrubs, subshrubs, vines, rarely aphyllous shrubs with cladode-like branches, unarmed or rarely with spines or prickles. Stipules diverse, caducous or persistent, rarely with embedded extrafloral nectary tissue. Leaves distichous or spiral, bifoliolate or paripinnate, rarely absent or reduced to a petiolar phyllode; extrafloral nectaries when present, on the petiole and/or on the leaf rachis, sessile or stipitate, the secretory surface convex; leaflets 1–many pairs, opposite. Inflorescences racemes or panicles; bract 1, persistent or caducous, bracteoles absent. Flowers hypogynous, bilaterally symmetrical or asymmetrical; hypanthium absent; sepals 5, free; petals 5, free, yellow; androecium comprising 3 adaxial staminodes and (4) 6–7 heteromorphic stamens, grouped in 2 sets of 4 median and (0) 2–3 abaxial stamens, rarely all 10 stamens fertile and homomorphic, filaments glabrous, anthers apically poricidal; pollen prolate-spheroidal to prolate, not syncopate; ovary stipitate. Fruit an indehiscent legume, a follicle or a legume dehiscing through both margins, linear or oblong, cylindrical, laterally compressed or tetragonal, fleshy or dry. Seeds variable in shape and colour.
Haploid chromosome numbers n = 11, 12, 13, 14 and 28 (
287 species (
Examples of Senna diversity A Senna macranthera (DC. ex Collad.) H.S. Irwin & Barneby flowering branch (Lima et al. 422) B Senna alata (L.) Roxb. flowering branch C Senna corifolia (Benth.) H.S. Irwin & Barneby flowering branch (Rando et al. 936) D Senna angulata (Vogel) H.S. Irwin & Barneby fruits E Senna multijuga (Rich.) H.S. Irwin & Barneby mature individual (Lima 408) F Senna pendula (Humb.& Bonpl. ex Willd.) H.S. Irwin & Barneby flower (Lima et al. 435) G Senna spectabilis (DC.) H.S. Irwin & Barneby flower (unvouchered) H Senna rugosa (G. Don) H.S. Irwin & Barneby leaf with extrafloral nectaries between leaflets (Lima et al. 538). Photo credits A, B, E, F, H A Lima C JG Rando D F Logan G RT Queiroz https://rubens-plantasdobrasil.blogspot.com/.
Senna occurs in a wide range of habitats, including wet forests, seasonally deciduous forests, seasonally semi-deciduous forests, savannas, deserts and also anthropised areas (
Species of Senna are used in traditional and conventional medicine, as bee forage for honey, ornamentals and for timber. Seeds are roasted and ground as a coffee substitute (
The name Senna derives from the Arabic “sana” or “sanna” which refers to plants with cathartic properties (
Senna is morphologically characterised by the paripinnate leaves, extrafloral nectaries present in many species (on leaves, stipules, bracts, and sepals; leaf nectaries with convex secretory surface, the others are embedded and externally not clearly visible), yellow petals, heteromorphic androecium usually with 6–7 fertile stamens (Fig.
Phylogenies have consistently retrieved seven strongly supported clades supported by floral morphological features and presence or absence of extrafloral nectaries (
Senna is outstanding within legumes with respect to its specialisation in pollination mode. This specialisation is expressed especially in the androecium in which the diversity pertains to patterns of heteranthery and anther elaborations, including dehiscence patterns, pointing direction of the pores, and extension of the lateral furrow (
Molecular dating analyses (
Edeline Gagnon16,17,18, Ruth P. Clark10, Jens J. Ringelberg3,4, Gwilym P. Lewis10
Citation: Gagnon E, Clark RP, Ringelberg JJ, Lewis GP (2024) 6. Tribe Caesalpinieae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 103–145. https://doi.org/10.3897/phytokeys.240.101716
Poincianeae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 253. 1943. Type: Poinciana L. [= Caesalpinia L.]
Caesalpinia L.
(27). Arquita Gagnon, G.P. Lewis & C.E. Hughes (5 species), Balsamocarpon Clos (1), Biancaea Tod. (6), Caesalpinia L. (9), Cenostigma Tul. (15), Cordeauxia Hemsl. (1), Coulteria Kunth (11), Denisophytum R. Vig. (8), Erythrostemon Klotzsch (31), Gelrebia Gagnon & G.P. Lewis (8), Guilandina L. (up to 20), Haematoxylum L. (5), Hererolandia Gagnon & G.P. Lewis (1), Hoffmannseggia Cav. (23), Hultholia Gagnon & G.P. Lewis (1), Libidibia (DC.) Schltdl. (7), Lophocarpinia Burkart (1), Mezoneuron Desf. (24), Moullava Adans. (4), Paubrasilia Gagnon, H.C. Lima & G.P. Lewis (1), Pomaria Cav. (16), Pterolobium R. Br. ex Wight & Arn. (10), Stenodrepanum Harms (1), Stuhlmannia Taub. (1), Tara Molina (3), Ticanto Adans. (9), Zuccagnia Cav. (1).
Trees, shrubs, subshrubs, or herbs, sometimes scandent, often with prickles, thorns, glands, or glandular hairs. Stipules (best seen on young flush foliage and on seedlings) variable across the tribe, ranging from minute, lanceolate-deltate to triangular, ovate, or orbicular, sometimes foliaceous, the margins sometimes ciliate-fimbriate, persistent, caducous, or apparently lacking (at least on mature leaves). Leaves pinnate or bipinnate. Inflorescences terminal and/or axillary racemes or panicles; bracteoles absent; pedicels often jointed. Flowers zygomorphic, or rarely almost actinomorphic; hypanthium usually present (rarely a short calyx tube); sepals generally free to hypanthium-rim, the lowermost sepal modified, often forming a hood (cucullate) over the other four sepals in bud, imbricate to valvate; petals 5, the median (innermost) petal usually clearly differentiated; stamens 10, all similar, the filaments usually hairy, especially basally, and sometimes glandular, anthers mostly dorsifixed, introrse; pollen tricolporate monads, mostly spherical, surface reticulate and with granular-membraned margos surrounding weakly developed colpi (a margocolpus); ovary subsessile to short-stipitate, stigma usually crateriform. Fruits diverse, 1–several-seeded. Seeds, flattened, or globose.
The tribe is pantropical, found predominantly in seasonally dry tropical forests and shrublands, but extending in a subset of clades into tropical and warm temperate savannas, tropical wet forests, and tropical coastal habitats (
The most inclusive crown clade containing Caesalpinia brasiliensis L. and Erythrostemon gilliesii (Hook.) Klotzsch, but not Cassia fistula L., Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig.
The tribe comprises ca. 223 species (the number of species of Guilandina is unresolved) in 27 genera. The tribe Caesalpinieae was first described by
Although there are no unique diagnostic morphological synapomorphies for the Caesalpinieae, it can be recognised by a combination of features, including the presence of glandular trichomes, prickles or spines, bilaterally symmetrical flowers with a somewhat modified lower sepal, and free stamens crowded around the pistil, although none of these characters are ubiquitous within the tribe. Flowers vary greatly (Fig.
Flowers of Caesalpinieae A Tara spinosa Britton & Rose, Peru, Ancash (Hughes et al. 3043) B Cenostigma pluviosum var. pluviosum, Bolivia, Santa Cruz (Wood et al. 26552) C Caesalpinia bahamensis Lam., Cuba (Lewis 1853) D Hultholia mimosoides (Lam.) Gagnon & G.P. Lewis, India E Pomaria burchellii (DC.) B.B. Simpson & G.P. Lewis, Botswana Ghanzi district F Erythrostemon coccineus (G.P. Lewis & J.L. Contr.) Gagnon & G.P. Lewis, Mexico, Oaxaca (Lewis et al. 1802) G Caesalpinia pulcherrima (L.) Sw., Honduras H Erythrostemon melanadenius (Rose) Gagnon & G.P. Lewis, Mexico, Oaxaca (Hughes et al. 2091) I Pterolobium stellatum (Forssk.) Brenan, Africa J Guilandina bonduc L., India K Gelrebia trothaei (Harms) Gagnon & G.P. Lewis, Tanzania. Photo credits A E Gagnon B, F–H CE Hughes C GP Lewis D VR Vinayaraj, India Biodiversity Portal (https://indiabiodiversity.org/group/wild_orchids_of_india/observation/show/335155), the basionym of Hultholia mimosoides E O Bourquin, Flora of Zimbabwe (https://www.zimbabweflora.co.zw/speciesdata/image-display.php?species_id=127200&image_id=6) I P Van Wyk J M Sanjappa K PJ Cribb.
Fruits of Caesalpinieae A Hererolandia pearsonii (L. Bolus) Gagnon & G.P. Lewis, Namibia, Sesriem Canyon B Haematoxylum brasiletto H. Karst., Mexico (Lewis 2057) C Lophocarpinia aculeatifolia (Burkart) Burkart, Paraguay, (Fortunato 8650) D Hoffmannseggia arequipensis Ulibarri, Peru, Arequipa, (Hughes et al. 2342) E Balsamocarpon brevifolium Clos, Chile (Baxter et al. DCI 1859) F Zuccagnia punctata Cav., Argentina, Mendoza G Cordeauxia edulis Hemsl., Somalia H Erythrostemon coccineus (G.P. Lewis & J.L. Contr.) Gagnon & G.P. Lewis, Mexico, Oaxaca (Lewis et al. 1802) I Paubrasilia echinata (Lam.) Gagnon, H.C. Lima & G.P. Lewis, Brazil J Libidibia paraguariensis (Parodi) G.P. Lewis, Bolivia, Santa Cruz (Hughes 2475) K Coulteria mollis Kunth, Guatemala (Lewis et al. 1714) L Cenostigma pluviosum var. cabralianum (G.P. Lewis) Gagnon & G.P. Lewis, Brazil (Lewis et al. 2019) M Pomaria jamesii (Torr. & A. Gray) Walp., USA. Photo credits A AA Dreyer B, I, K, L GP Lewis C RH Fortunato D, H, J CE Hughes E P Baxter F Italo Specogna, Flora mendocina (http://www.floramendocina.com.ar/clase_3/zuccagnia_punctata_p9558.html) G M Thulin M P Alexander, SEINet Arizona Chapter (https://swbiodiversity.org/seinet/imagelib/imgdetails.php?imgid=253949).
Fruits of the Caesalpinieae A Gelrebia rubra (Engl.) Gagnon & G.P. Lewis, Namibia B Moullava spicata (Dalzell ex Wight) Nicolson, India, Maharashtra C Hultholia mimosoides (Lam.) Gagnon & G.P. Lewis, India D Guilandina bonduc L., Madagascar E Pterolobium stellatum (Forssk.) Brenan, Zimbabwe F Ticanto sinensis (Hemsley) R. Clark & Gagnon, China (Clark 415) G Biancaea decapetala (Roth) O. Deg., Peru, Ancash (Hughes et al. 2227) H Mezoneuron kauaiense (H. Mann) Hillebr. I Mezoneuron andamanicum Prain, Thailand (Clark 251). Photo credits A Dave U, iNaturalist (https://www.inaturalist.org/photos/41085094) B P Awale, Flowers of India (http://www.flowersofindia.net/) C VR Vinayaraj, India Biodiversity Portal (https://indiabiodiversity.org/observation/show/335158), the basionym of Hultholia mimosoides D GP Lewis E BT Wursten, Flora of Zimbabwe (https://www.zimbabweflora.co.zw/speciesdata/image-display.php?species_id=127190&image_id=1) F, I P Suksathan G CE Hughes H D Eickhoff, https://www.flickr.com/photos/dweickhoff/4822012867/in/photostream/.
Vegetative traits of the Caesalpinieae A Hoffmannseggia minor (Phil.) Ulibarri, Bolivia B Hererolandia pearsonii (L. Bolus) Gagnon & G.P. Lewis, Namibia, Sesriem Canyon C Moullava spicata (Dalzell ex Wight) Nicolson, India, Maharastra D Cordeauxia edulis Hemsl., undersurface of leaflets showing glands, Somalia E Pomaria austrotexana B.B. Simpson, USA, Texas F Small tree of Erythrostemon nicaraguensis (G.P. Lewis) Gagnon & G.P. Lewis, Nicaragua, Esteli (Hawkins et al. 4) G Leopard bark of Libidibia paraguariensis (D. Parodi) G.P. Lewis, Bolivia, Santa Cruz (Hughes 2475) H Lenticelled bark of Erythrostemon nicaraguensis (G.P. Lewis) Gagnon & G.P. Lewis, Nicaragua, Esteli (Hawkins et al. 4) I Young flush of leaves of Cenostigma pluviosum var. intermedium (G.P. Lewis) Gagnon & G.P. Lewis, Brazil, Bahia (Lima et al. 7901) J Fluted trunk of Haematoxylum brasiletto H. Karst., Mexico, Oaxaca, (Hughes 1947) K Prickles on woody protuberances on a young trunk of Paubrasilia echinata (Lam.) Gagnon, H.C. Lima & G.P. Lewis, Brazil, Bahia (Lima et al. 7909) L Recurved spines of Biancaea decapetala (Roth) O. Deg., Peru, Ancash, (Hughes et al. 3055). Photo credits A GP Lewis B AA Dreyer C Shivaprakash, iNaturalist (https://inaturalist.ca/observations/81864757) D M Thulin E WR Carr F–H, J CE Hughes I, K, L E Gagnon.
At the generic level, fruits are highly variable and taxonomically more useful than flowers. Several of the genera can be differentiated based on fruit characteristics. For example, the fruits of Balsamocarpon, Cordeauxia, Coulteria, Cenostigma, Guilandina, Haematoxylum, Hererolandia, Hultholia, Libidibia, Lophocarpinia, Moullava, Paubrasilia, Pterolobium and Zuccagnia are all distinctive and provide useful diagnostic synapomorphies for these genera (Figs
The pollen details of individual genera are not easily extracted from the literature because so many generic names have been reinstated from within Caesalpinia s.l., or new genera described, since the palynological study of the Caesalpinioideae by
For an exhaustive list of accepted species names and synonyms in tribe Caesalpinieae, together with their currently accepted equivalents see
Stuhlmannia moavii Taub.
Unarmed trees. Stipules minute conical projections, caducous. Leaves pinnate or bipinnate and then ending in a pair of pinnae, pinnae in (1) 2–10 opposite pairs, with reddish glands; leaflets in 3–12 opposite to subopposite pairs per pinna, eglandular or with red glands on the lower surface. Inflorescence a terminal or axillary raceme. Flowers bisexual, sub-actinomorphic; hypanthium persisting as a shallow cup at the pedicel apex as the fruit matures; sepals 5, caducous, valvate in bud, lowermost sepal not conspicuously differentiated in bud; petals 5, free, yellow, the median petal with red markings, slightly smaller than the other 4; stamens 10, filaments pubescent; ovary stipitate, with red sessile glands, glabrous to pubescent. Fruit a flattened, oblong, woody, elliptic legume, dehiscing along both sutures, valves twisting. Seeds flattened, sub-circular to ovate, brown.
Unknown.
Seasonally dry tropical forest, woodland on limestone and in riverine forest.
Named by Taubert for the German naturalist Franz Ludwig Stuhlmann (1863–1928).
Unknown.
The genus is phylogenetically and morphologically closely related to Cordeauxia (Fig.
Cordeauxia edulis Hemsl.
Multi-stemmed, unarmed, evergreen shrubs, red gland dots on stems. Stipules caducous or lacking (not seen). Leaves pinnate; leaflets in (1) 2–4 (6) pairs, coriaceous, with conspicuous red glands on the lower surface (Fig.
2n = 24 (
Monospecific (C. edulis), in north-eastern Africa (Somalia and Ethiopia). Introduced in Israel, Kenya, Sudan, Tanzania, and Yemen (
Seasonally dry tropical (semi-desert) bushland and thicket on sand.
Named by Hemsley for Captain H. E. S. Cordeaux (1870–1943), one time H. M. Commissioner in Somalia.
The seeds of C. edulis (yeheb nut) are used as human food and have potential as an arid-land food; also used as livestock fodder, production of a red dye, as medicine, wood, an insecticide, and a soap substitute (
Cordeauxia is closely related to the genus Stuhlmannia but is easily distinguished by its distinct habit: a shrub with a large tap-root (vs. medium-sized tree), and large, cornute, inertly dehiscent fruit with ovoid seeds (vs. non-cornute, explosively dehiscent fruit with compressed seeds).
Cenostigma macrophyllum Tul.
Unarmed multi-stemmed shrubs, small compact trees, or large trees to 35 m, the larger trees with fluted trunks at maturity. Stipules filiform, spathulate-cucullate or lanceolate, caducous or sub-persistent, unknown for some species. Leaves pinnate or bipinnate, sometimes with stellate hairs or various types of sessile or stalked glands; species with pinnate leaves either with three leaflets or 2–9 pairs of opposite leaflets; species with bipinnate leaves with 1–11 pairs of opposite to alternate pinnae, plus a terminal pinna, each pinna with 3–29 alternate to subopposite (occasionally opposite), eglandular leaflets, or with black subepidermal glands on the undersurface, and/or with conspicuous, sessile or punctate glands on the undersurface or along the margins, in addition to stipitate glands. Inflorescence an axillary or terminal raceme, sometimes pyramidal in shape, sometimes aggregated into large showy panicles, pedicels articulated. Flowers bisexual, zygomorphic; hypanthium persisting as a small cup or wide shallow cup, or abscising as a ring around the pedicel apex or fruit stipe as the fruit matures; sepals 5, caducous, the lower cucullate sepal generally slightly longer than the other four; petals 5, free, bright yellow, the median petal with red or orange markings on the inner surface of the blade, the outer surface of the petal claw with short-stalked glands; stamens 10, free, filaments pubescent on lower portion, usually with short-stipitate glands along entire length; ovary pubescent with glands intermixed. Fruits laterally compressed, coriaceous to woody legumes with conspicuously thickened margins, dehiscent, sometimes explosively so, 2–6 (8)-seeded. Seeds ochre, brown, or mottled, shiny.
2n = 24 [C. microphyllum (Mart ex G. Don) Gagnon & G.P. Lewis, C. pluviosum (DC.) Gagnon & G.P. Lewis, C. pyramidale (Tul.) Gagnon & G.P. Lewis], 2n = 48 [C. bracteosum (Tul.) Gagnon & G.P. Lewis] (
Twenty-three taxa in 15 species confined to the Neotropics. The genus extends around the Amazonian arc of dry forests and adjacent cerrado vegetation, as well as throughout Central America, and extending to the Caribbean, with endemics in Cuba and Hispaniola (Fig.
Seasonally dry tropical forest, bushland, and thicket (restinga, caatinga, semi-arid thorn scrub), wooded grassland (cerrado and cerradão) and terra firme forest.
From ceno- (Greek = empty) and stigma, presumably alluding to the chambered stigma (a character of many species of the Caesalpinieae, and not restricted to Cenostigma).
Cenostigma pluviosum is often planted as an ornamental street tree in South America. Other species are used for their timber and production of charcoal, as well as for local medicine (
Based on phylogenetic and morphological evidence, Gagnon and Lewis in
Caesalpinia sect. Libidibia DC., Prodr. [A.P. de Candolle] 2: 483. 1825. Type: Caesalpinia coriaria (Jacq.) Willd. [≡ Poinciana coriaria Jacq. (≡ Libidibia coriaria (Jacq.) Schltdl.)]
Stahlia Bello, Anal. Soc. Esp. Hist. Nat. 10: 255. 1881. Type: Stahlia maritima Bello [= Libidibia monosperma (Tul.) Gagnon & G.P. Lewis]
Libidibia coriaria (Jacq.) Schltdl. [≡ Poinciana coriaria Jacq.]
Small to medium-sized or large unarmed trees; bark hard, smooth, with a patchwork of shades of grey, white, and pale green, often referred to as snake bark [except in L. coriaria and L. monosperma (Tul.) Gagnon & G.P. Lewis, where it is rough and fissured]. Stipules caducous or lacking (not seen). Leaves bipinnate, rarely pinnate (L. monosperma); bipinnate leaves with 2–10 pairs of opposite pinnae plus a single terminal pinna and 3–28(30) pairs of opposite leaflets per pinna; pinnate leaves with 4–6 pairs of opposite to subopposite leaflets; leaflets eglandular or with subsessile gland dots on the undersurface of the blades, on either side of the midvein. Inflorescence a terminal or axillary raceme or panicle, sometimes corymbose. Flowers bisexual, zygomorphic; hypanthium usually not persistent as the fruit matures; sepals 5, caducous, the lower sepal slightly longer and cucullate in bud; petals 5, free, yellow, or white, the median petal sometimes flecked or blotched orange or red; stamens 10, free, pubescent on the lower half of the filaments, eglandular [except for L. ferrea (Mart. ex Tul.) L.P. Queiroz, which has stipitate glands]; ovary eglandular. Fruit coriaceous to woody, straight (contorted in L. coriaria), indehiscent, eglandular, glabrous, black (red and somewhat fleshy in L. monosperma). Seeds somewhat laterally compressed.
2n = 24 [L. coriaria, L. ferrea, L. paraguariensis (D. Parodi) G.P. Lewis, L. punctata (Willd.) Britton], and 2n = 48 (L. ferrea) (
Ten taxa in seven species in the Neotropics. One species (L. monosperma, previously in the monospecific genus Stahlia) is endemic to Puerto Rico and the Dominican Republic. The other species are found across a circum–Amazonian arc of dry forests and adjacent cerrado vegetation, across the Andes, as well as throughout Central America (Fig.
Seasonally dry tropical forests and thorn scrub (including Brazilian Caatinga) and savanna woodlands. Libidibia monosperma occurs along the margins of mangrove swamps and in marshy deltas, in drier edaphic conditions.
The name Libidibia is derived from the vernacular name ‘libi-dibi’ or ‘divi-divi’ used for some species.
Libidibia species are widely used as ornamental park and street trees. Their fruits are rich in tannin and used commercially in the tanning industry and sometimes used for animal fodder, ink and local medicines. The wood and timber are prized in turnery and for parts of guitars and violins, as well as for decorative inlay and cabinet work. Some species are used in heavy construction (railway sleepers, beams, bridge supports), for tool handles and as firewood (
The genus needs revising; other species are perhaps waiting to be discovered and described, both in the field and in herbaria (
Barreto Valdés (2013);
Balsamocarpon brevifolium Clos
Shrub to 2 m tall, with 3–5 mm long, deflexed or patent, woody, nodal, often paired, sometimes caducous, woody spines. Stipules deltoid, glandular, caducous. Leaves pinnate, in fascicles on short brachyblasts; leaflets in 3–4 pairs, glabrous, fleshy. Inflorescence a short raceme. Flowers bisexual, sub-zygomorphic; a short hypanthium persisting (sometimes with the sepals still attached) and tightly adhering to the base of the fruit as it matures; sepals 5, fimbriate; petals 5, free, yellow, with glandular trichomes on the dorsal surface; stamens 10, free, filaments pubescent; ovary glandular and finely pubescent. Fruit thick, turgid, resinous, glandular, and indehiscent, 3–4-seeded. Seeds round, orange-brown.
Unknown.
Desert scrub, rocky hillsides.
From balsamo- (Greek = balsam) and carpos (Greek = fruit), the fruits yield a sticky resin traditionally used for tanning.
Fruit resin used in the tanning industry, and wood locally used for charcoal production and as firewood (
Over-exploitation and increased fragmentation of the remaining populations of B. brevifolium mean that the species is vulnerable to extinction (
Zuccagnia punctata Cav.
Shrubs. Stipules caducous (not seen). Leaves pinnate; leaflets 5–13 pairs, subopposite, with glandular dots on both surfaces of the leaflet blades. Inflorescence a terminal, erect raceme. Flowers bisexual, zygomorphic; the calyx (hypanthium and sepals) persistent at fruit maturity; sepals 5, glabrous; the lower sepal cucullate and covering the other four in bud; petals 5, free, yellow, glandular trichomes on the dorsal surface of the petal blades; stamens 10, free, pubescent; ovary pilose. Fruit ovoid-acute, oblique, laterally compressed, indehiscent, 1-seeded, gall-like, on a short stipe and covered with long reddish-brown bristles at maturity. Seeds laterally compressed.
2n = 24 (
Dry temperate upland and montane bushlands and thickets on sandy plains.
Named by Cavanilles for the Italian physician, traveller and plant collector, Attilio Zuccagni (1754–1807).
Minor local medicinal uses; the leaves yield a yellow dye (
Although recorded and described from Chile in the 19th Century, the genus has been cited as doubtful for the flora of Chile (
Larrea Ortega, Nov. Rar. Pl. Descr. Dec.: 15. t. 2. 1797, nom. rej., non Larrea Cav., Anales Hist. Nat. 2(4): 119. 1800 [Zygophyllaceae]. Type: Larrea glauca Ortega [≡ Hoffmannseggia glauca (Ortega) Eifert]
Moparia Britton & Rose, N. Amer. Fl. 23(5): 317. 1930. Type: Moparia repens (Eastw.) Britton & Rose [≡ Caesalpinia repens Eastw. (≡ Hoffmannseggia repens (Eastw.) Cockerell)]
Hoffmannseggia falcaria Cav., nom. superfl. [≡ Hoffmannseggia glauca (Ortega) Eifert (≡ Larrea glauca Ortega)]
Perennial woody herbs, most species forming a basal rosette, or subshrubs, unarmed, often arising from bud-bearing and tuberous roots, shoots pubescent and with gland-tipped trichomes. Stipules lanceolate, ovate or deltate, acuminate, caducous or persistent. Leaves bipinnate, ending in a pair of pinnae plus a single terminal pinna (except for Hoffmannseggia aphylla (Phil.) G.P. Lewis & Sotuyo); pinnae in 1–13 opposite pairs; leaflets small and numerous, in 2–15 (18) opposite pairs per pinna. Inflorescence a terminal or axillary raceme. Flowers bisexual, zygomorphic; calyx (hypanthium and sepals) usually persistent as the fruit matures; sepals 5, weakly imbricate; petals 5, free, yellow to orange, the median petal often with red markings; stamens 10, free, filaments pubescent; ovary glabrous to pubescent, eglandular to glandular. Fruit laterally compressed, straight or sometimes falcate, the sutures almost parallel, papery to leathery, glabrous to pubescent, eglandular or with glandular trichomes, indehiscent or dehiscent, with twisting valves. Seeds compressed, ovoid.
2n = 24 [H. drepanocarpa A. Gray, H. eremophila (Phil.) Burkart ex Ulibarri, H. glauca (Ortega) Eifert, H. microphylla Torr., H. oxycarpa Benth., H. viscosa (Ruiz & Pav.) Hook. & Arn.] (
Twenty-five taxa in 23 species, with an amphitropical distribution in the Americas: 10 species restricted to North America (southern USA and Mexico), 12 in South America (mainly Andean), and one species (H. glauca) widespread throughout the range of the genus (Fig.
Subtropical desert and semi-desert grassland, often in open areas and on disturbed sites, on sandy, rocky, or calcareous soils.
Named by Antonio José Cavanilles for the German botanist, entomologist and ornithologist, Johann Centurius Graf von Hoffmannsegg (1766–1849).
Hoffmannseggia glauca produces tubers once eaten by indigenous groups in North America (but sometimes becomes a noxious weed); the roots of H. intricata Brandegee produce a reddish-brown dye (
A complete synopsis and key to species [except H. aphylla which was transferred to the genus by
Stenodrepanum bergii Harms
Suffrutescent shrub, or perennial herb, (10) 20–40 cm, with bud-bearing and occasionally tuber-forming roots; glabrous, with globose sessile glands scattered along the branches. Stipules ovate. Leaves bipinnate, pinnae in 1–3 opposite pairs plus a single terminal pinna; leaflets in 5–9 opposite to subopposite pairs per pinna, embedded glands on the lower surface. Inflorescence a lax, terminal raceme. Flowers bisexual, zygomorphic; hypanthium persistent as a small cup at the apex of the pedicel as the fruit matures; sepals 5, caducous, glandular, the lower cucullate sepal covering the other four in bud; petals 5, free, yellow, the median petal with red markings, stipitate glands on the dorsal surface; stamens 10, free, filaments pubescent and glandular; ovary glandular. Fruit linear to slightly falcate, cylindrical, torulose, 1–5-seeded. Seeds ovoid.
2n = 24, 36 (
Subtropical wooded grassland and scrub, especially close to salt pans.
From Greek, steno- (= narrow) and drepano- (= sickle), in allusion to the narrow sickle-shaped fruit.
Unknown.
Morphologically similar in appearance to the genus Hoffmannseggia but with a distinctive linear to slightly falcate, cylindrical, torulose fruit. Resolved as sister to Hoffmannseggia in
Poincianella Britton & Rose, N. Amer. Fl. 23(5): 327. 1930. Type: Poincianella mexicana (A. Gray) Britton & Rose [≡ Caesalpinia mexicana A. Gray (≡ Erythrostemon mexicanus (Rose) Gagnon & G.P. Lewis)]
Schrammia Britton & Rose, N. Amer. Fl. 23(5): 317. 1930. Type: Schrammia caudata (A. Gray) Britton & Rose [≡ Hoffmannseggia caudata A. Gray (≡ Erythrostemon caudatus (A. Gray) Gagnon & G.P. Lewis)]
Erythrostemon gilliesii (Hook.) Klotzsch [≡ Poinciana gilliesii Hook.]
Shrubs or small to medium-sized trees, occasionally suffrutices, unarmed (except E. glandulosus (Bertero ex DC.) Gagnon & G.P. Lewis). Stipules ovate-lanceolate, ovate, or orbicular, acute to acuminate, sometimes foliaceous, cordate and auriculate at the base, caducous or less often persistent. Leaves bipinnate, usually ending in a single terminal pinna; pinnae in 1–6 (15), opposite pairs; leaflets in 2–13 (20) opposite pairs per pinna, leaflet blades eglandular or with conspicuous black sessile glands along the margin, these sometimes sunken in the sinuses of the crenulated margin. Inflorescence an axillary or terminal raceme. Flowers bisexual, zygomorphic; hypanthium persistent as a wide or narrow, shallow or deep cup, or sometimes abscising to form a free ring around the pedicel apex as the fruit matures; sepals 5, lower sepal cucullate in bud, all sepals caducous; petals 5, free, bright golden yellow to creamish yellow, salmon pink or pink-scarlet, the median petal often with red-orange markings, the corolla diverse in form; stamens 10, free, filaments pubescent, eglandular or with stipitate glands; ovary pubescent, eglandular or with sessile or stipitate glands. Fruit a chartaceous to coriaceous or slightly woody, laterally compressed legume, elastically dehiscent with twisting valves, eglandular or with stipitate glands, (1) 2–7 (8)-seeded. Seeds yellow to ochre-brown or mottled with grey and black.
2n = 24 [E. exostemma (Moc. & Sessé ex DC.) Gagnon & G.P. Lewis, E. gilliesii, E. hughesii (G.P. Lewis) Gagnon & G.P. Lewis, E. melanadenius (Rose) Gagnon & G.P. Lewis, E. mexicanus (Rose) Gagnon & G.P. Lewis, E. nelsonii (Britton & Rose) Gagnon & G.P. Lewis, E. yucatanensis (Greenm.) Gagnon & G.P. Lewis] (
Thirty-four taxa in 31 species: 22 species are found across the southern USA, Mexico, and Central America, one occurs in the Caribbean (Cuba and Hispaniola), and eight occur in South America (Fig.
Seasonally dry tropical forests across the Neotropics; also occurring in deserts, yungas-puna transition zones, and chaco-transition forests (Argentina, Bolivia, Chile, Paraguay).
From Greek, erythro- (= red) and stemon (= stamen), the type species E. gilliesii has long red exserted stamens, but this is unusual in the genus as circumscribed here.
Erythrostemon gilliesii is widely cultivated as a garden ornamental and is hardy in Mediterranean and temperate regions (
Originally described as a monospecific genus, its circumscription was recently emended to include many species previously placed in Central American and Mexican Poincianella (
Melanosticta DC., Prodr. [A.P. de Candolle] 2: 485. 1825. Type: Melanosticta burchellii DC. [≡ Pomaria burchellii (DC.) B.B. Simpson & G.P. Lewis]
Cladotrichium
Vogel, Linnea 11: 401. 1837. Lectotype (designated by
Pomaria glandulosa Cav.
Small shrubs, subshrubs, or perennial herbs, with a moderate to dense indumentum of simple curled hairs, sometimes also scattered plumose trichomes, intermixed with sessile, oblate glands (drying black) on stems. Stipules mostly laciniate, glandular, persistent. Leaves bipinnate, pinnae in 1–8 (11) opposite pairs, plus a terminal pinna; leaflets small, 2–16 (27) opposite pairs per pinna, always with multiple sessile glands on their lower surface (these orange in the field, drying black). Inflorescence a terminal or axillary raceme. Flowers bisexual, zygomorphic; hypanthium persistent as a small shallow cup as the fruit matures; sepals 5, caducous, lanceolate, the lower sepal cucullate, covering the other 4 in bud, and closely embracing the androecium and gynoecium at anthesis; petals 5, free, yellow, white, red, or pink; stamens 10, filaments pubescent; ovary sparsely to densely hairy and glandular. Fruit a linear or sickle-shaped, laterally compressed legume, with a sparse to dense covering of plumose/dendritic or stellate trichomes (sometimes obscure and restricted to fruit margin) intermixed with sessile oblate glands (drying black), elastically dehiscent, with twisting valves. Seeds laterally compressed.
2n = 24 P. rubicunda (Vogel) B.B. Simpson & G.P. Lewis, P. stipularis (Vogel) B.B. Simpson & G.P. Lewis) (
Mainly in subtropical dry grassland and in degraded sites, many on limestone.
Named by Cavanilles for Dominic Pomar, botanist from Valencia, and doctor to Philip III (1598–1621), King of Spain.
Unknown.
Revisions of the species of Pomaria are available for North America (
Arquita mimosifolia (Griseb.) Gagnon, G.P. Lewis & C.E. Hughes [≡ Caesalpinia mimosifolia Griseb.]
Small to medium-sized, often decumbent, shrubs, usually with glandular trichomes on various parts of the plant. Stipules ovate-obovate to deltoid, usually with a fimbriate-glandular margin, caducous. Leaves bipinnate; pinnae 1–5 pairs, usually with a single terminal pinna; leaflets in 4–12 opposite pairs per pinna, often with maroon/black glands in depressions on crenulated leaflet margins, and sometimes with occasional sessile black glands on the undersurface of leaflet blades. Inflorescence a leaf-opposed raceme. Flowers bisexual, zygomorphic; hypanthium persistent as a small shallow cup at the pedicel apex as the fruit matures; sepals 5, caducous, the lower sepal cucullate; petals 5, free, yellow to orange, median petal sometimes streaked red; stamens 10, free; ovary usually covered with gland-tipped trichomes. Fruits laterally compressed, lunate-falcate legumes, covered sparsely to densely with gland-tipped trichomes, these sometimes dendritic. Seeds laterally compressed, ovate-orbicular, the testa shiny olive-grey, sometimes mottled or streaked black.
2n = 24 (A. mimosifolia) (
Six taxa in five species restricted to the Andes in South America, in disjunct inter-Andean valleys, in Ecuador, Peru, Bolivia and Argentina (Fig.
Tropical and subtropical seasonally dry, montane, and rupestral habitats.
Arquita is the vernacular name for A. trichocarpa (Griseb.) Gagnon, G.P. Lewis & C.E. Hughes in Argentina (
Unknown.
A revision of Arquita with a key to species is available in
Hererolandia pearsonii (L. Bolus) Gagnon & G.P. Lewis [≡ Caesalpinia pearsonii L. Bolus]
Multi-stemmed shrubs armed with curved, deflexed prickles. Stipules not seen. Leaves pinnate, borne in fascicles on short woody brachyblasts that are usually subtended by a pair of tiny (sometimes obscure) prickles; leaflets (4) 5–7 (9) pairs, opposite, eglandular. Inflorescence a short raceme. Flowers zygomorphic, bisexual; hypanthium short, persistent as a ring around the stipe of the fruit; sepals 5, free, the lower sepal cucullate and covering the other 4 sepals in bud, all sepals caducous; petals 5, yellow, free; stamens 10, free, pubescent on the lower half; ovary pubescent. Fruit a thinly woody, laterally compressed, almost circular to strongly sickle-shaped legume, dehiscing along the sutures, finely pubescent and covered in robust trichomes, usually 1-seeded. Seeds laterally compressed.
Unknown.
Semi-desert and desert areas, on stony, sandy soils.
The type locality of H. pearsonii is in the semiarid Hereroland, a region of eastern Namibia inhabited by the Herero people, who are nomadic cattle herders.
Unknown.
The genus was described by
Haematoxylon L., Philosophia Botanica: 34. 1764, orth. var.
Cymbosepalum Baker, Bull. Misc. Inform. Kew 1895 (100–101): 103. 1895. Type: Cymbosepalum baronii Baker [= Haematoxylum campechianum L.]
Haematoxylum campechianum L.
Multi-stemmed shrubs to medium-sized trees, armed with scattered straight conical spines, and short, lateral spinescent shoots; mature trees with conspicuously fluted trunks, shrubs often with ribbed branches. Stipules minute, acuminate, caducous. Leaves pinnate or bipinnate (both can be present on the same individual in some species), eglandular; pinnate leaves with 2–6 pairs of leaflets; bipinnate leaves with 1–3 pairs of pinnae plus a terminal pinna, each pinna with 2–5 (6) pairs of opposite leaflets. Inflorescence a terminal or axillary raceme or panicle. Flowers bisexual, actinomorphic to zygomorphic; the short hypanthium persisting in fruit as a small cup; sepals 5, free, the lower sepal cucullate and slightly covering the other 4 in bud, sepals caducous; petals 5, yellow to pale yellow or white, free; stamens 10, free, filaments pubescent, particularly on the lower half; ovary glabrous to pubescent. Fruit laterally flattened, membranous to chartaceous, dehiscing along the middle of the valves, or near the margin of the fruit, but never along the sutures, 1–3-seeded. Seeds oblong to reniform, flattened.
2n = 24 (H. campechianum) (
Five species: two in Central America (Salvador to Costa Rica), Mexico, South America (Colombia and Venezuela) and the Caribbean (perhaps introduced), two endemic to Mexico, and one in Southern Africa (Namibia; Fig.
Deserts, seasonally dry tropical semi-deciduous scrub and thorn scrub, sandy riverbeds, and dry rocky hillsides. One species (H. campechianum) is known to grow in frequently inundated marshy areas by rivers.
From Greek, haemato- (= bloody) and xylon (= wood), alluding to the blood-red heartwood of H. campechianum which produces a brilliant red dye.
The heartwood of H. campechianum is the source of a colourless chemical, haematoxylin, which upon oxidation turns to haematein, a commercial dark violet dye used for wool, silk, cotton, fur, leather, bone and synthetic fibre dying, and with iron chromium mordants to obtain red and black; also used as a stain in microscopical preparations (particularly to show up cell nuclei), and ink for writing and painting and the rich red colour has been used to adulterate wine. Species are also used medicinally, as ornamentals and living hedges and the bee flowers yield a high-quality honey (
This genus is easily diagnosable by the ascending secondary veins of its leaflets, which form a sharp angle with the primary vein. There is a key to species by
Barreto Valdés (2013);
Lophocarpinia aculeatifolia (Burkart) Burkart [≡ Cenostigma aculeatifolium Burkart]
Shrubs, armed with scattered straight, conical, 2–5 mm long spines on shoots; leaves and inflorescences crowded on brachyblasts. Stipules acuminate, caducous. Leaves pinnate, leaflets in 2 (3) opposite pairs, eglandular, with a pair of small prickles at the insertions of the leaflets. Inflorescence a short, corymbiform, pubescent raceme, each with 3–6 flowers. Flowers zygomorphic, bisexual; hypanthium turbinate, fleshy, persistent at the apex of the pedicel as the fruit matures; sepals 5 caducous, lower sepal cucullate and covering the other 4 sepals in bud, embracing the androecium and gynoecium at anthesis; petals 5, yellow to yellow-orange, free, the median petal differentiated from the rest by a fleshy claw and wavy blade margins, pubescent; stamens 10, free, filaments pubescent; ovary glabrous. Fruit a lomentum, with 1–5 segments, falcate, with 4 coarsely serrate wings. Seeds ellipsoid to reniform, smooth.
Unknown.
Chaco woodlands and seasonally dry tropical to subtropical forests.
From Greek, lopho- (= combed or crested) and carpos (= fruit), the fruit has 4 crested wings, the ending -inia signifies a close relationship with Caesalpinia.
Unknown.
Lophocarpinia is closely related to the genus Haematoxylum but has a distinctive lomentaceous fruit with coarsely serrated wings.
Poinciana L., Sp. Pl. 1: 380. 1753. Type: Poinciana pulcherrima L. [≡ Caesalpinia pulcherrima (L.) Sw.]
Caesalpinia sect. Brasilettia DC., Prodr. [A.P. de Candolle] 2: 481. 1825. Type: Caesalpinia brasiliensis L.
Brasilettia (DC.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Type: Brasilettia brasiliensis (L.) Kuntze [≡ Caesalpinia brasiliensis L.]
Caesalpinia brasiliensis L.
Shrubs or small trees, usually armed with curved deflexed prickles. Stipules minute, caducous or apparently lacking. Leaves bipinnate, pinnae (1) 2–6 pairs, opposite; leaflets 3–13 pairs per pinna, alternate to opposite. Inflorescence a terminal or axillary raceme or panicle. Flowers pedicellate, bisexual, zygomorphic; hypanthium persistent as a cup at the apex of the pedicel as the fruit matures, if the fruit stipitate then the stipe exerted from the hypanthial cup, or the whole calyx persistent (e.g., in Caesalpinia pulcherrima (L.) Sw.); sepals 5, caducous or persistent, eglandular, the lower sepal strongly cucullate and covering the other 4 sepals in bud; petals 5, variable in colour (yellow, white, red, orange or green), the corolla also variable in shape; stamens 10, free, the filaments pubescent; ovary glabrous and eglandular. Fruit a wingless, unarmed, coriaceous, glabrous, eglandular, explosively dehiscent legume, with twisting valves, 3–7-seeded. Seeds laterally compressed.
2n = 24 (C. bahamensis Lam., C. pulcherrima) (
Nine species restricted to the Neotropics (apart from the pantropically cultivated C. pulcherrima). One species (C. cassioides Willd.) occurs in the northern Andes from Peru to Colombia, one (C. pulcherrima) is likely native in Guatemala and the state of Sonora in Mexico (but is widely cultivated), C. nipensis Urb. is endemic to Cuba, and all other species are also Caribbean in distribution (Fig.
Distribution of Caesalpinia based on quality-controlled digitised herbarium records. The distribution of Caesalpinia pulcherrima in Sonora and Guatemala is based on specimen records that morphologically appear to be non-cultivated, but the exact native distribution of the species remains difficult to ascertain because it has been widely cultivated for a long time. See Suppl. material
Seasonally dry tropical forests, coastal thicket, bushlands and thorn scrubs, dry plains, and riparian woodlands, on soils derived from limestone or sandstone.
Named by Linnaeus for Andrea Cesalpino (1519–1603), Italian naturalist, botanical collector, systematist and philosopher, physician to Pope Clement VIII, professor of medicine and botany in Pisa and Rome.
Caesalpinia pulcherrima is widely cultivated pantropically as a garden and park ornamental and has various medicinal properties (
Caesalpinia, as recently re-circumscribed (
Barreto Valdés (2013);
Denisophytum madagascariense R. Vig.
Shrubs or small trees, armed with straight or curved, deflexed prickles, scattered along shoots and in pairs at the petiole base (except D. madagascariense which is unarmed). Stipules minute, or foliaceous and conspicuous, caducous or persistent. Leaves bipinnate, pinnae in 1–6 opposite pairs; leaflets 2–10 (11) opposite pairs per pinna. Inflorescence a terminal or axillary raceme. Flowers bisexual, zygomorphic; a short hypanthium persistent at the pedicel apex as the fruit matures; sepals 5, caducous, lower sepal cucullate and covering the other 4 sepals in bud; petals 5, free, yellow, the median petal sometimes with red markings on the inner face of the blade; stamens 10, free, filaments pubescent and eglandular; ovary glabrous. Fruits coriaceous, laterally compressed (inflated in D. madagascariense), glabrous, eglandular, stipitate legumes, elastically dehiscent, with twisting valves. Seeds laterally compressed.
2n = 24 [D. pauciflorum (Griseb.) Gagnon & G.P. Lewis] (
Nine taxa in eight species. Three species are distributed in Mexico, Florida, and the Caribbean, one species is endemic to Paraguay, Bolivia, and Argentina, one is endemic to northern Madagascar, and the other three occur in northern Kenya, Somalia, and Arabia (Fig.
Low deciduous seasonally dry tropical woodlands or scrublands, also in open pine woodlands, coastal plains and foothills. Species in Madagascar and Africa grow in limestone soils.
It has been hypothesised that Denisophytum honours Marcel Denis, a botanist with expertise in the genus Euphorbia L. in Madagascar, and a friend and collaborator of René Viguier, the genus author (
Unknown.
An evaluation of species limits is needed for this genus. It has a highly disjunct trans-continental distribution typical of lineages occupying the succulent biome sensu
Barreto Valdés (2013);
Paubrasilia echinata (Lam.) Gagnon, H.C. Lima & G.P. Lewis [≡ Caesalpinia echinata Lam.]
Medium sized to large tree, armed with small to large, upturned prickles, these usually arising from woody protuberances, bark flaking in large woody plates; heartwood red, with the trunk exuding a red sap when injured. Stipules caducous (on seedlings lanceolate, acute to acuminate). Leaves bipinnate, ending with a pair of pinnae; pinnae in (2) 3–20 alternate pairs; leaflets alternate, (2) 3–19 (21) leaflets per pinna (generally the number of leaflets is inversely proportional to their size). Inflorescence a terminal, or occasionally axillary, raceme or panicle, with 15–40 flowers. Flowers bisexual, zygomorphic; hypanthium persistent as a shallow cup or abscising to form a small free ring around the pedicel apex as the fruit matures; sepals 5, caducous, the lowest sepal cucullate, covering the other 4 in bud; petals 5, free, bright yellow, eglandular, the median petal with a blood-red blotch on the inner face; stamens 10, free, eglandular, the filaments densely pubescent on lower half; ovary pubescent with bristles intermixed. Fruit a spiny, finely pubescent, sub-lunate, woody, elastically dehiscent legume with twisting valves, 1–2-seeded. Seeds laterally compressed.
2n = 24 (
Dry coastal cactus scrub often on rocky outcrops, inland in Mata Atlântica, and in tall restinga on well-drained sandy soil.
“Pau-brasil” is the national tree of Brazil and has long been associated with the country. The Latinization of its well-known and much used common name recognises the importance of the species to Brazil.
Widely cultivated in Brazil as an ornamental street or park tree, and sometimes in plantations. The tree’s red sap was once used for dying cotton and cloth and its wood is much prized for the manufacture of high-quality violin bows (
Originally described as Caesalpinia echinata by
Guaymasia Britton & Rose, N. Amer. Fl. 23(5): 322. 1930. Type: Guaymasia pumila Britton & Rose [≡ Coulteria pumila (Britton & Rose) Sotuyo & G.P. Lewis]
(designated by
Trees or shrubs, unarmed, dioecious. Stipules minute, caducous or lacking. Leaves bipinnate, pinnae in 2–6 opposite to subopposite pairs; leaflets in (2) 4–12 (14) opposite to alternate pairs per pinna, eglandular, glabrous to velvety pubescent. Inflorescence an axillary or terminal raceme. Flowers unisexual, zygomorphic; hypanthium either persisting as a shallow cup or not persistent as the fruit matures; sepals 5, caducous, with a lower glandular-pectinate, cucullate sepal, covering the other 4 in bud; petals 5, yellow, free; staminate flowers with 10 free stamens, filaments pubescent, eglandular; pistillate flower with ovary pubescent or glabrous. Fruit chartaceous to papyraceous, laterally-compressed, oblong to elliptic (occasionally suborbicular), indehiscent (or sometimes opening along one suture), wingless, often persisting to next flowering season, eglandular, glabrous to densely velutinous, 1–6-seeded. Seeds ovate, orbicular, or sub-quadrate, compressed.
2n = 24 (
Eleven species in Mexico and Central America, one species extending to Belize, Aruba, Cuba, Jamaica and Curaçao, one to Venezuela (including Isla Margarita) and Colombia (Fig.
Seasonally dry tropical forest, semi-deciduous and deciduous woodlands, xerophytic scrub forests and thorn scrub, on sandy, calcareous, or metamorphic substrates, some species endemic on limestone.
Named by Kunth for the Irish botanist Thomas Coulter (1793–1846) who collected in central Mexico (1825–1834) and was curator of the herbarium at Trinity College, Dublin, Ireland.
A preferred local firewood in its native range; some species are used as ornamentals in living fences (
Nicarago Britton & Rose, N. Amer. Fl. 23(5): 319. 1930. Type: Nicarago vesicaria (L.) Britton & Rose [≡ Caesalpinia vesicaria L. (≡ Tara vesicaria (L.) Molinari, Sánchez Och. & Mayta)]
Russellodendron Britton & Rose, N. Amer. Fl. 23(5): 320. 1930. Type: Russellodedron cacalaco (Bonpl.) Britton & Rose [≡ Caesalpinia cacalaco Bonpl. (≡ Tara cacalaco (Bonpl.) Molinari & Sánchez Och.)]
Tara tinctoria Molina [= Tara spinosa (Molina) Britton & Rose]
Shrubs or trees, armed with deflexed prickles. Stipules minute, deltate-lanceolate, caducous, or lacking. Leaves bipinnate, ending with a pair of pinnae, sometimes armed with prickles at the base of the pinnae and leaflets; pinnae in 2–5 opposite pairs; leaflets 1–8 opposite pairs per pinna. Inflorescence a terminal or axillary raceme or panicle. Flowers bisexual, zygomorphic; hypanthium persistent as a shallow cup at the pedicel apex or abscising as a narrow free ring as the fruit matures; sepals 5, eglandular, caducous, lower sepal cucullate covering the other 4 sepals in bud, with a pectinate, fimbriate or entire margin; petals 5, free, yellow, the median petal with red markings; stamens 10, free, the filaments pubescent, eglandular; ovary puberulent to glabrescent. Fruit an indehiscent legume, straight, oblong, laterally compressed, slightly turgid and somewhat fleshy, coriaceous. Seeds ellipsoid, brown, shiny.
2n = 24 (all three species) (
Three species, one in South America (T. spinosa, which occurs from central and south-western Colombia through Ecuador and Peru into Bolivia), one in Mexico [T. cacalaco (Bonpl.) Molinari & Sánchez Och.] and one in Mexico, Guatemala, Nicaragua and extending into the Caribbean [T. vesicaria (L.) Molinari, Sánchez Och. & Mayta] (Fig.
Seasonally dry tropical forests to semi-arid thorn scrubs.
Derived from the vernacular name ‘tara’ in Peru, Bolivia, and Chile.
Tara spinosa is widely cultivated across the tropics and subtropics (including in the Canary Islands) as a source of tannins, gums and firewood, and occasionally as an ornamental (
As originally described by Molina, the genus Tara contained a single binomial. The genus Coulteria, as described by
Barreto Valdés (2013);
Gelrebia rubra (Engl.) Gagnon & G.P. Lewis [≡ Hoffmannseggia rubra Engl.]
Erect to scrambling shrubs, armed with scattered, straight, or curved, deflexed prickles. Stipules minutes, caducous or lacking. Leaves bipinnate, ending in a pair of pinnae; pinnae 1–17 opposite pairs; leaflets 1–33 opposite pairs per pinna [except in G. glandulosopedicellata (R.Wilczek) Gagnon & G.P. Lewis], lower surface of the blades with numerous subepidermal glands or translucent dots. Inflorescence a terminal or axillary raceme. Flowers bisexual, zygomorphic; hypanthium persisting as a wide shallow cup at the pedicel apex as the fruit matures; sepals 5, caducous, lower sepal strongly cucullate (occasionally with a beaked apex), covering the other 4 sepals in bud before anthesis; petals 5, free, dark pinkish mauve to light pinkish white, eglandular; stamens 10, free, filaments pubescent and eglandular; ovary glabrous. Fruit a coriaceous, broadly oblong-ovate to obliquely pyriform legume. Seeds laterally compressed.
Unknown.
Nine taxa in eight species, restricted to Africa, in Namibia, Angola, Botswana, South Africa, Mozambique, northern Kenya, Ethiopia, Somalia, and the Democratic Republic of the Congo (Zaire, Katanga) (Fig.
Deciduous bushlands, dry woodlands, on rocky ridges, often along dry riverbeds, or on sandy valley floors. One species also found in degraded savannas.
Gelreb or gelrib is the Somali name for Gelrebia trothaei subsp. erlangeri (Harms) Gagnon & G.P. Lewis, meaning ‘camel trap’ and clearly alluding to the robust deflexed prickles characteristic of the species, and indeed the whole genus, which can hinder the passage of camels.
Unknown.
Bonduc Mill., Gard. Dict. Arb. Ed. 4: 28. 1754.
Caesalpinia subg. Guilandina (L.) Gillis & Proctor, J. Arnold. Arbor. 55(3): 426. 1974. Type: Caesalpinia bonduc (L.) Roxb. [≡ Guilandina bonduc L.]
Guilandina bonduc L.
Lianas or scrambling shrubs, generally densely armed with robust recurved prickles. Stipules subulate to compound-foliaceous, caducous to persistent. Leaves bipinnate, with recurved prickles on leaf and pinnae rachides; pinnae in 3–11 opposite pairs; leaflets 3–24 opposite to alternate pairs per pinna, eglandular. Inflorescence a terminal or axillary raceme, often branched near the base. Flowers unisexual, segregated on separate staminate and pistillate racemes (sometimes the staminate flowers cryptically hermaphrodite but with the anthers lacking pollen); hypanthium either not persistent in fruit or persisting as a narrow tube closely adhering to the fruit stipe; sepals 5, caducous, valvate in bud, the lower sepal slightly cucullate; petals 5, free, yellow, barely exceeding the sepals; stamens 10, free, pubescent near the filament base; ovary usually covered in bristly trichomes. Fruits oblong elliptic, inflated legumes, usually armed with 5–10 mm long slender spinescent bristles. Seeds obovoid to globular, ca. 2 cm in diameter, smooth, grey, pale to dark brown, or orange, with parallel fracture lines concentric with the small apical hilum.
2n = 24 (G. bonduc) (
Up to 20 species (see notes), widely distributed as far north as some of the southern and eastern Japanese islands, south to South Africa, Madagascar, and Australia. Also occurring in South America and the Caribbean, China, India, Myanmar (Burma), Indo China, Hong Kong, and Taiwan (Fig.
Coastal thickets on sand, in secondary forests, and lowland rainforests, occasionally on limestone.
Named by Linnaeus for Melchior Wieland (1515–1589), Prussian naturalist, traveller, and scholar from Königsberg, who settled in Italy and italianised his name to ‘Guilandini’, or Guilandinus in Latin; he was sent to the Levant, Asia, and Africa (1559–1560), was captured by pirates and finally ransomed by Gabriele Falloppio.
There are reports of the genus being poisonous, although some species are used medicinally. The large globose seeds of G. bonduc and G. major are used as marbles, necklace beads and buttons (
Guilandina lacks a recent global taxonomic account and there are doubts about the number of species, with estimates ranging from seven to 20. The genus needs to be revised with particular attention given to species delimitation, synonymy, and geographical distribution. Six Caesalpinia binomials included as belonging to Guilandina in
Hultholia mimosoides (Lam.) Gagnon & G. P. Lewis [≡ Caesalpinia mimosoides Lam.]
Climbing woody shrub; branches densely armed with short, robust, needle-like trichomes; young stems pubescent, with rust-coloured, hyaline hairs, and dome-shaped glands, topped with a few hairs. Stipules subulate, caducous. Leaves bipinnate; pinnae in 10–30 opposite pairs, with a pair of deflexed prickles at the insertion of the pinnae on the leaf rachis and at the insertion of leaflets on the pinnae rachides; leaflets 7–20 pairs per pinna, opposite, glabrous, eglandular. Inflorescence a terminal or leaf-opposed, lax raceme, with 50 or more flowers. Flowers bisexual, zygomorphic; hypanthium persisting as a wide shallow cup at the pedicel apex as fruit matures; sepals 5, caducous, the lower sepal strongly cucullate; petals 5, free, bright yellow, dark glands present on the blade, median petal smaller than the 4 lateral petals; stamens 10, free, filaments pubescent at least on the lower half; ovary densely pubescent and with glandular dots (often obscured by the dense pubescence). Fruit an obovoid, falcate, vesicular, unarmed, dehiscent legume, sparsely pubescent, particularly along the margin, and with a few obscure stellate hairs, and covered in gland dots, 1–3-seeded. Seeds sub-globose, grey.
2n = 24 (
In secondary thickets and clearings, often on roadsides.
The name Hultholia honours the Cambodian botanist Dr. Sovanmoly Hul Thol (born 1946), whose doctoral thesis, “Contribution à la révision de quelques genres de Caesalpiniaceae, representés en Asie” (Hul Thol 1976), is an important revision of the Asian species and genera of the Caesalpinieae, and particularly the genus Pterolobium.
Although Hultholia mimosoides is not known to be cultivated, the young, pungent, flowering shoots are sold as a vegetable in markets in Vientiane (Laos) (Vidal and Hul Thol 1976).
This monospecific genus was described by
Almeloveenia Dennst., Schlüssel Hortus Malab.: 32. 1818. Type: Almeloveenia spinosa Dennst. [= Moullava spicata (Dalzell ex Wight) Nicolson]
Cinclidocarpus Zoll. & Moritzi, Natuur-Geneesk. Arch. Ned.-Indië iii: 81. 1846. Type: Cinclidocarpus nitidus Zoll. & Moritzi [= Moullava tortuosa (Roxb.) Gagnon & G.P. Lewis]
Wagatea Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 3: 90. 1851. Type: Wagatea spicata Dalzell ex Wight [≡ Moullava spicata (Dalzell ex Wight) Nicolson]
Caesalpinia sect. Cinclidocarpus (Zoll. & Moritzi) Benth. & Hook., Gen. Pl. 6.: 565–567. 1865. Type not designated.
Moullava spicata (Dalzell ex Wight) Nicolson [≡ Wagatea spicata Dalzell ex Wight]
Lianas and scrambling shrubs, armed with deflexed prickles. Leaves bipinnate, ending with a pair of pinnae; pinnae in 7–20 opposite pairs; leaflets in 5–40 opposite pairs per pinna. Stipules caducous or lacking (not seen). Inflorescence an elongated terminal or axillary raceme, the racemes sometimes aggregated into panicles. Flowers bisexual, sub-actinomorphic or zygomorphic; hypanthium persisting either as a distinct cup or as a wide shallow calyx remnant at the pedicel apex as the fruit matures; sepals 5, caducous, eglandular, glabrous, the lower sepal strongly cucullate, covering the other 4 sepals in bud; petals 5, free, yellow, the median (innermost upper) and lateral petals sometimes streaked red, eglandular; stamens 10, free, barely exserted beyond the corolla, densely pubescent on lower half of filaments; ovary glabrous or pubescent. Fruit fleshy, oblong-elliptic, unarmed, indehiscent, sub-torulose, with thickened sutures, drying black (immature fruits of M. spicata red-tomentose), exocarp and endocarp strongly adnate, glabrous, 1–4-seeded. Seeds sub-globular, olive-brown to black.
2n = 24 [M. digyna (Rottler) Gagnon & G.P. Lewis, M. spicata] (
The Asian species are found in seasonally dry tropical semi-evergreen forest margins, secondary thickets, and on mountain slopes, up to 1200 m elevation. The African species occurs mostly in riverine habitats in lowland rainforests.
Derived from the vernacular name of Moullava spicata, “mulu” (Malayalam: spiny), a spiny climber.
Moullava spicata is used for medicine (
First described as a monospecific genus from India, its description was emended in
Campecia Adans., Fam. Pl. (Adanson) 2: 318. 1763 (no type species designated, and no species names ever published in this genus. It is thus not possible to apply this name which is rejected against Biancaea).
Caesalpinia sect. Sappania DC., Prodr. [A. P. de Candolle] 2: 484. 1825. Type not designated.
Biancaea scandens Tod. [= Biancaea decapetala (Roth) Deg.]
Lianas, climbing or trailing shrubs or small trees, armed with short, slightly recurved prickles. Stipules lanceolate-oblong to broadly ovate, sometimes amplexicaul at base, caducous or persistent. Leaves bipinnate, ending with a pair of pinnae, rachis armed with pairs of prickles at the base of each pinna, sometimes also scattered on the rachis; pinnae in 4–19 opposite or alternate pairs; leaflets in 5–20 opposite or alternate pairs per pinna. Inflorescence a terminal or axillary raceme or panicle. Flowers bisexual, zygomorphic; hypanthium persisting as a small cup or wider shallow cup or occasionally as an abscised free ring around the pedicel apex as the fruit matures; sepals 5, caducous, usually pubescent, the lower sepal cucullate and covering the other 4 in bud; petals 5, free, yellow to white, eglandular, the median petal smaller than the other 4, and inrolled towards the centre; stamens 10, filaments densely pubescent especially at the base; ovary densely velutinous. Fruit a coriaceous, pubescent, glabrescent or glabrous, eglandular, dehiscent, wingless, laterally compressed (but somewhat inflated and often with a narrow wing along the upper suture in B. decaptala), 1–9-seeded legume. Seeds flat, black, or brown.
2n = 24 (B. decapetala, B. sappan) (
Six species widespread across southern Asia. Biancaea decapetala, native to Asia, has been widely introduced across the tropics as a hedge plant or ornamental and is invasive in South Africa and Hawaii (Fig.
Primary forest and forest margins, grasslands, scrub vegetation, riverine habitats, secondary thickets, and clearings. From the coast to mountain slopes.
It is presumed that the Italian botanist Agostino Todaro (1818–1892) named Biancaea for his fellow countryman and botanist Giuseppe Bianca (1801–1883).
Biancaea decapetala often grown as a living fence (
Gagnon and Lewis in
Cantuffa J.F. Gmel., Syst. Nat., ed 13[bis]. 2(1): 677. 1791, nom. rej. vs. Pterolobium R. Br. ex Wight & Arn. Type: Cantuffa exosa J.F. Gmel. [≡ Pterolobium exosum (J.F. Gmel.) Baker f. (= Pterolobium stellatum (Forssk.) Brenan)]
Reichardia Roth, Nov. Pl. Sp.: 210. 1821, nom. illeg., non Roth, Bot. Abh. Beobacht. 35. 1787, nec Roth, Catal. Bot. 2: 64. 1800. Lectotype: Reichardia hexapetala Roth [≡ Pterolobium hexapetalum (Roth) Santapau & Wagh]
Pterolobium lacerans R. Br. ex Wight & Arn., nom. illeg. [Pterolobium exosum (J.F. Gmel.) Baker f. [≡ Cantuffa exosa J.F. Gmel. (= Pterolobium stellatum (Forssk.) Brenan)]
Lianas or scrambling / trailing shrubs, armed with prickles. Stipules small, inconspicuous, subulate to triangular-subulate, caducous. Leaves bipinnate; pinnae in 5–20 opposite pairs; leaflets in 6–25 opposite pairs per pinna. Inflorescence a terminal or axillary raceme, often aggregated into panicles. Flowers bisexual, sub-actinomorphic to zygomorphic; hypanthium persisting as a minute cup or ridge at the pedicel apex as the fruit matures; sepals 5, caducous, the lower sepal cucullate, covering the other 4 sepals in bud; petals 5, free, yellow to white, equal to slightly differentiated, the median petal sometimes in-rolled; stamens 10, free, filaments pubescent (occasionally glabrous); ovary pubescent. Fruit a red to brown samara, 1 (2)-seeded. Seeds ovate-oblong, sub-compressed.
2n = 24 (P. stellatum) (
Seasonally dry tropical upland evergreen forests, riverine and humid forests, woodlands, and wooded grasslands.
From Greek, ptero- (= wing) and lobion (= fruit), in reference to the fruit which resembles a samara.
The leaves of P. stellatum are used in parts of Africa as a leather dye, for ink and as medicine; plants are grown as living fences (
Vidal and Hul Thol (1974) published a revision of Pterolobium, with a key to species.
Mezonevron Desf., Mém. Mus. Hist. Nat. 4: 245. 1818, orth. var.
Mezoneurum DC., Prodr. [A.P. de Candolle] 2: 484. 1825, orth. var.
Caesalpinia subg. Mezoneuron (Desf.) Vidal ex Herend. & Zarucchi, Ann. Missouri Bot. Gard. 77(4): 854. 1990.
Mezoneuron glabrum Desf. [= Mezoneuron pubescens Desf.]
Scrambling shrubs or lianas, occasionally medium-sized trees, usually armed with recurved prickles on stem and leaves, rarely unarmed. Stipules triangular, often caducous. Leaves bipinnate, ending in a pair of pinnae; pinnae in (1) 2–18 opposite to sub-opposite pairs; leaflets in 1–15 opposite to alternate pairs per pinna. Inflorescence a terminal or axillary raceme (often aggregated into panicles). Flowers bisexual, zygomorphic; hypanthium persisting as a small cup or wide shallow cup as the fruit matures; sepals 5, imbricate, the lower sepal cucullate, and overlapping the other 4 in bud, caducous; petals 5, free, usually yellow with red markings on the median petal, or occasionally red, pink or cream, the median petal somewhat modified (either with a fleshy ligule or a patch of hairs on the inner surface between the blade and claw, or the petal bilobed); stamens 10, free, usually all pubescent or villose on lower half, or one or all glabrous; ovary glabrous to hairy. Fruit laterally compressed, indehiscent, chartaceous, coriaceous or woody, venose, longitudinally (and often broadly) winged along the upper suture, the wing 2–20 mm wide, 1–13-seeded. Seeds ± transversely arranged in seed chamber, compressed.
2n = 22 (M. cucullatum, M. kauaiense) (
Tropical and subtropical riverine forests, lowland rainforests, swamp forests, seasonally dry forests, thickets, vine forests and wooded grasslands, especially along forest and river margins.
From Greek, meso- (= middle) or meizon (= greater) and neuron (= nerve), the upper suture of the fruit is bordered by a usually broad longitudinal wing so that the suture appears as a prominent sub-central nerve or vein.
Used for medicine; the wood of M. kauaiense (Mann.) Hillebr. (“uhiuhi”) from Hawaii was once used locally for spears and in house construction (
Mezoneuron is absent from the American tropics and subtropics today, but fossil records show that the genus was widespread across North America, as well as in Europe, during the middle Eocene, about 45 Ma (
Caesalpinia sect. Nugaria DC., Prodr. [A.P. de Candolle] 2: 481. 1825. Type not designated.
Guilandina paniculata Lam. [= Ticanto crista (L.) R. Clark & Gagnon]
Scandent shrubs or lianas to 15 m. Stems usually with scattered, recurved prickles. Stipules triangular, caducous. Leaves bipinnate, leaf rachis with recurved prickles at base of pinnae and usually scattered in between, pinnae in 1–16 opposite pairs; leaflets in 2–15 opposite pairs per pinna. Inflorescence a terminal or axillary raceme or panicle, bracts (at base of racemes) and bracteoles (at base of pedicels) caducous; pedicels articulated. Flowers bisexual, zygomorphic; hypanthium persisting as a small cup as the fruit matures; sepals 5, caducous, the lower lobe cucullate over the others in bud; petals 5, the median petal distinct from the others in shape, usually with a ± circular patch of hairs on the inner surface, the lateral petals glabrous or with few hairs; stamens 10, free, the basal half tomentose; ovary 1–2-ovuled, glabrous or hairy, stigma funnel-shaped and more or less papillate, or truncate. Fruit coriaceous or ligneous, dehiscent or indehiscent, elliptic, lunate, or suborbicular, the upper suture lacking a wing, or with a narrow wing up to 4 mm wide, one species with a carinate wing 5–6 mm deep, fruit usually glabrous, otherwise tomentose, 1 (2)-seeded. Seeds circular to reniform, compressed.
2n = 24 (T. crista) (
Nine species, mainly in southern and central China, with a few records in Laos, Myanmar, northern Thailand, and northern Vietnam, and one species (T. crista) throughout South East Asia, extending into Australia and Oceania (Fig.
Scrub or (riverine) forests, on limestone or in sandy parts of mangroves, up to 1800 m elevation.
Ticanto was a vernacular name used for these plants by the Brachmanes (Brahmans or Brahmins), a sector of Hinduism.
Numerous studies report pharmacological uses of Caesalpinia crista but appear to concern the species Guilandina bonduc to which the name Caesalpinia crista is often erroneously applied. Confirmed uses of true C. crista are not known to the present authors.
Ticanto is a genus recently re-instated by
Colin E. Hughes3, Elspeth Haston18
Citation: Hughes CE, Haston E (2024) 7. Tribe Schizolobieae In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 146–164. https://doi.org/10.3897/phytokeys.240.101716
Schizolobium Vogel, Linnaea 11: 399. 1837.
(8). Bussea Harms (7 species), Colvillea Bojer ex Hook. (1), Conzattia Rose (1), Delonix Raf. (12), Heteroflorum M. Sousa (1), Parkinsonia L. (12), Peltophorum (Vogel) Benth. (6–7), Schizolobium Vogel (2).
Trees or occasionally shrubs, (3) 5–40 m and up to 1.5 m stem diameter, typically branching low down with spreading, flat-topped crowns, some species of Delonix with swollen trunks; outer bark generally thin, pale silvery grey, or green, inner bark green; mainly unarmed, occasionally armed with axillary thorns, stipular spines and/or spinescent leaf rachides and spinescent side shoots; brachyblasts usually absent, rarely present. Stipules absent, minute, setiform, acicular and caducous, or lobed, pinnatifid or bipinnatifid, and persistent, or spinescent and persistent. Leaves bipinnate, variable in leaf formula, sometimes large, rarely paripinnate and much reduced. Inflorescences axillary racemes or terminal panicles, pedicels usually jointed. Flowers usually showy, almost always yellow, occasionally orange, whitish-pink or red, usually bisexual, but sometimes unisexual (Heteroflorum, Conzattia, and Parkinsonia anacantha Brenan are at least partially dioecious); a distinct hypanthium always present, this generally short, discoid or shallowly campanulate, but larger and obliquely turbinate in Schizolobium; sepals 5, subequal, either imbricate or valvate, free or partially fused, usually strongly reflexed; petals usually 5, occasionally 1, subequal or variable, longer or shorter clawed, usually crinkled, often with erose margins; stamens 10, usually spreading, rarely clustered around the ovary; pollen in oblate tricolporate monads with moderately to coarsely reticulate surface ornamentation. Fruits diverse, 1–many-seeded, usually flat, but sometimes terete or thickened, generally linear or linear-oblong or spathulate oblanceolate, the valves papery, coriaceous or woody, sutures sometimes thickened or with a narrow wing, indehiscent or dehiscent along both sutures, sometimes tardily so, usually inertly so, or rarely elastically from the apex. Seeds oblong-ellipsoid or ovate, discoid, lenticular and flattened to sub-spherical or globose, albuminous or ex-albuminous, integument hard.
Tribe Schizolobieae occupies a strikingly disjunct pantropical distribution spanning dry and wet tropical forests and savannas.
The most inclusive crown clade containing Schizolobium parahyba (Vell.) S.F. Blake and Delonix decaryi (R. Vig.) Capuron, but not Caesalpinia brasiliensis L., Dimorphandra conjugata (Splitg.) Sandwith, or Mimosa sensitiva L. (Fig.
Tribe Schizolobieae is here delimited to include eight genera and ca. 45 species. The monophyly and generic contents of this clade were first established using molecular data by
Tribe Schizolobieae is equivalent to the Peltophorum group s.s. of
Although varying markedly in stature, from shrubs and small trees to gigantic, buttressed trees up to 40 m tall in Schizolobium, all species of tribe Schizolobieae share a characteristic tree form and bark (Fig.
Variation in tree form and bark among genera of tribe Schizolobieae A, B Schizolobium amazonicum Huber ex Ducke, tree to 20 m, Veracruz, Mexico (Hughes 1880) C young sapling of Schizolobium amazonicum showing typical ‘tree fern habit’ with large leaves and green bark D Peltophorum dubium (Spreng.) Taub., cultivated as an ornamental, central plaza in Saipina, Santa Cruz, Bolivia (Hughes 2461) E bark of Peltophorum africanum Sond. F Parkinsonia peruviana C.E. Hughes, Daza & Hawkins, small tree in dry thorn scrub in the upper Marañon valley, Peru (Hughes 2213) G green shoot and axillary thorns, Parkinsonia andicola (Griseb.) Varjão & Mansano, Chuquisaca, Bolivia (Hughes 2313) H green bark of Parkinsonia praecox (Ruiz & Pavon ex Hook.) Hawkins, Oaxaca, Mexico (Hughes 1298) I tree of Parkinsonia africana Sond., South Africa J, K small tree of Heteroflorum sclerocarpum M. Sousa showing typical low forked branching habit, spreading crown and pale metallic grey bark with storied protuberences, Guerrero, Mexico (Hughes 1845) L, M Conzattia multiflora Standl., showing typical umbrella crown habit and pale grey metallic bark in succulent-rich seasonally dry forest, Puebla, Mexico (Hughes 1824) N small tree of Colvillea racemosa Bojer, Madagascar (Du Puy 102) O bottle tree of Delonix floribunda (Baill.) Capuron, south-western Madagascar (Bruneau 1405) P twisted and stunted growth form of Delonix pumila Du Puy, Phillipson & R. Rabev. in south-western Madagascar (Du Puy 456) Q small tree of Delonix regia (Bojer) Raf. with a spreading crown, flowering preceding or coinciding with leaf flush, N Madagascar (Du Puy 578). Photo credits A, B, D, F, G, H, J, K, L, M CE Hughes C A Bayer Tamayo File:Tambor (Schizolobium parahyba) (14101065740).jpg - Wikimedia Commons E W Coville (https://plantnet.org/) I Wi Boshoff http://hdl.handle.net/11660/5647 N, P, Q D Du Puy O A Bruneau.
Leaf formula of the mainly bipinnate leaves (only Delonix edule has once-pinnate leaves) is highly variable among genera of Schizolobieae, ranging from highly reduced leaves with very short rachides and very reduced leaflets in some species of Parkinsonia, to what are some of the largest leaves of any genus in subfamily Caesalpinioideae, at 1–2 m long on young saplings of Schizolobium (Fig.
The petals of flowers of all genera are showy, mainly bright yellow, but in some species of Delonix and Peltophorum orange, whitish-pink or red (Fig.
While Schizolobieae can be found in all tropical vegetation types, the greatest diversity of genera and species occurs in the trans-continental succulent biome sensu
Schizolobium excelsum Vogel [= Schizolobium parahyba (Vell.) S.F. Blake]
Unarmed trees, 25–40 m, the trunk frequently 50 cm and up to 150 cm diameter, often buttressed, saplings typically unbranched to 2–3 m bearing at the apex a cluster of huge, tree fern-like leaves (Fig.
2n = 26 (
Two species, one widespread across the Neotropics from south-central Mexico, Central America, and Amazonia and the second in coastal Brazil from Santa Catarina north to Bahia (Fig.
Flowers of tribe Schizolobieae A Schizolobium amazonicum Huber ex Ducke, Veracruz, Mexico (Hughes 1880) B Peltophorum dubium (Spreng.) Taub., Santa Cruz, Bolivia (Hughes 2461) C Bussea perrieri R. Vig., western Madagascar D Parkinsonia aculeata L., Zacapa, Guatemala (Hughes 1222) E Parkinsonia florida (Benth. ex A. Gray) S. Watson, Sonora, Mexico (Hughes 1562) F Conzattia multiflora Standl., Puebla, Mexico (Hughes 931) G Colvillea racemosa Bojer, south-western Madagascar (Bruneau 1403) H Delonix regia (Bojer) Raf., northern Madagascar (Du Puy 578) I Delonix leucantha subsp. gracilis Du Puy, Phillipson & R. Rabev., Madagascar (Du Puy 87). Photo credits A, B, D–F CE Hughes C feno, iNaturalist (https://www.inaturalist.org/photos/103942022) G A Bruneau H, I D Du Puy.
Fruits of tribe Schizolobieae A, B dehisced fruits of Schizolobium amazonicum Huber ex Ducke showing the papery endocarp surrounding the single seed, Veracruz, Mexico (Hughes 1880) C Peltophorum dubium (Spreng.) Taub., Chiapas, Mexico (Hughes 1685) D Peltophorum pterocarpum (DC.) Backer ex K. Heyne, India E Bussea sakalava Du Puy & R. Rabev., Madagascar (Du Puy 248) F dehisced fruit of Bussea perrieri R. Vig., north-west Madagascar G ripe, tardily dehiscent fruits of Parkinsonia andicola (Griseb.) Varjão & Mansano, Chuquisaca, Bolivia (Hughes 2619) H Parkinsonia florida (Benth. ex A. Gray) S. Watson, Sonora, Mexico (Hughes 1562) I ripe indehiscent fruits of Parkinsonia peruviana C.E. Hughes, Daza & Hawkins, La Libertad, Peru (Eastwood 82) J Conzattia multiflora Standl., Puebla, Mexico (Hughes 1815) K–M indehiscent ripe fruits of Heteroflorum sclerocarpum M. Sousa, Guerrero, Mexico (Hughes 1845) L Heteroflorum sclerocarpum Guerrero, Mexico (Hughes 1845) showing a legume that has been broken apart manually N Delonix boiviniana (Baill.) Capuron, Madagascar (D Du Puy 301) O Delonix edule (H. Perrier) Babineau & Bruneau, ripe fruits, Madagascar (Du Puy 449) P Colvillea racemosa Bojer, ripe dehiscing fruits, Madagascar (Du Puy 304). Photo credits A–C, G–M CE Hughes D D Valke File:Peltophorum pterocarpum (2745008096).jpg - Wikimedia Commons E, N–P D Du Puy F Andry.A.R, iNaturalist (https://www.inaturalist.org/photos/28896062).
Mainly confined to terra firme in tropical moist forest and often common in secondary forest. Deciduous, at least briefly, usually flowering when leafless just before leaf flush. Seeds wind-dispersed.
From Greek, schizo- (= split or divided) and -lobion (= fruit), in reference to the splitting of the exocarp from the endocarp at maturity.
Renowned for its fast growth rate and very light soft wood. Widely cultivated as an ornamental and sometimes as coffee or cacao shade; young saplings, with their up to 2m-long leaves resemble tree ferns (Fig.
Schizolobium is sister to the clade comprising the rest of the tribe (Fig.
Bussea massaiensis (Taub.) Harms [≡ Peltophorum massaiense Taub.]
Unarmed shrubs or more usually treelets or trees, (2) 6–25 m, with an erect branching habit, the trunk 70–100 cm in diameter, bark smooth, pale whitish-grey, young shoots with a brown or rusty indumentum. Stipules small, subulate, caducous. Leaves bipinnate, petiolar and rachis glands absent, (1) 8–14 pairs of pinnae and (5) 8–11 (26) pairs of opposite leaflets, these rhombic and unequal-sided, the base truncate, the midvein oblique, dark green and glossy above. Inflorescences terminal or subterminal in the upper leaf axils, paniculate or racemose, often combined into a compound terminal inflorescence, the axes minutely velvety brown, pedicels not jointed, bracts mostly caducous (persistent in B. massaiensis subsp. rhodesica). Flowers bisexual, weakly zygomorphic; hypanthium shallowly campanulate; sepals 5, imbricate in bud, becoming reflexed, the 3 inner sepals with membranous erose hyaline margins; petals 5, the lateral pairs sub-equal, the upper petal reduced, all petals crinkled, the margins erose, tapering at the base to a short robust claw, bright yellow (Fig.
2n = 22 (
Seven species, five from Africa (one endemic to Mozambique, one in West Africa, two in Tanzania and one in Angola – Congo), plus two species endemic to western and northern Madagascar (Fig.
Seasonally dry tropical forest, thickets and deciduous bushland, often on sandy soils, moist semi-deciduous forest and rainforest.
Named in honour of Walter Carl Otto Busse (1865–1933), German botanist and ecologist who collected material of the type species in East Africa in 1901.
The wood is hard and used for construction and firewood.
Robustly supported as sister to Peltophorum (Fig.
Baryxylum Lour., Fl. Cochinch. 1: 266–267. 1790, nom rej. against Peltophorum (Vogel) Benth. Type: Baryxylum rufum Lour. [= Peltophorum dasyrhachis var. dasyrhachis]
Caesalpinia sect. Peltophorum Vogel, Linnaea 11(3): 406. 1837. Type: Caesalpinia dubia Spreng. [≡ Peltophorum dubium (Spreng.) Taub.]
Peltophorum vogelianum Walpers, nom. illeg. [= Peltophorum dubium (Spreng.) Taub.]
Unarmed trees (Fig.
2n = 26 (
Six (or seven) species. One (or two) in the Neotropics, one in southern Africa, four in South East Asia, one of which, P. pterocarpum (DC.) Backer ex K. Heyne, reaching N Australia (Fig.
In lowland deciduous and evergreen moist forests, coastal vegetation along beaches and mangrove edges, frequent in forest clearings and secondary vegetation. Evergreen or often deciduous. Flowering often immediately preceding leaf flush, indehiscent narrowly-winged fruits probably wind-dispersed. Some introduced and cultivated species weakly naturalised in places, the true native distribution of P. dubium in some doubt in some parts of the Neotropics, e.g., in Bolivia (
From Greek, pelte- (= shield) and -phoros (= bearing), in reference to the large peltate shield-like centrally-attached stigma.
Several species are introduced and cultivated as ornamentals (Fig.
The genus Peltophorum resembles Schizolobium in potentially gigantic stature, abruptly bipinnate leaves, massive terminal racemose-paniculate inflorescence of handsome yellow flowers, and almost equal calyx-lobes, but differs in the grossly peltate stigma which Peltophorum shares with Bussea which is its sister genus (Fig.
Cercidium Tul., Arch. Mus. Hist. Nat. 4: 133. 1844. Type: Cercidium spinosum Tul. [= Parkinsonia praecox (Ruiz & Pav.) Hawkins]
Rhetinophloeum H. Karst., Fl. Columb. 2: 25. 1862. Type: Rhetinophloeum viride H. Karst. [= Parkinsonia praecox (Ruiz & Pav.) Hawkins]
Peltophoropsis Chiov., Ann. Bot. (Rome) 13(3): 385. 1915. Type: Peltophoropsis scioana Chiov. [≡ Parkinsonia scioana (Chiov.) Brenan]
Cercidiopsis Britton & Rose, N. Amer. Fl. 23(5): 306. 1930. Type: Cercidiopsis microphylla (Torr.) Britton & Rose [≡ Parkinsonia microphylla Torr.]
Parkinsonia aculeata L.
Highly branched shrubs or small trees (Fig.
2n = 28 (
Twelve species, eight confined to the New World (including two named hybrid species), and four in Africa. Parkinsonia occupies a striking disjunct amphi-Atlantic distribution across arid and semi-arid parts of the New World from the southern USA to Argentina and disjunctly in arid parts of north-eastern and south-western Africa (Fig.
Distribution of Parkinsonia based on quality-controlled digitised herbarium records [excluding P. aculeata whose native range remains uncertain, and which is weedy and invasive in many areas of Neotropics and elsewhere – see
Parkinsonia is confined to seasonally dry, semi-arid and arid climates, growing in Chaco woodlands, seasonally dry tropical forests, deserts and semi-deserts, occupying most enclaves of the disjunct trans-continental succulent biome distribution (sensu
Named in honour of John Parkinson (1567–1650), the British apothecary and herbalist to King James I of England.
Parkinsonia aculeata is widely cultivated as an ornamental street tree in arid zones.
Parkinsonia is robustly supported as sister to the clade comprising Heteroflorum + Conzattia + Delonix + Colvillea (Fig.
Conzattia arborea Rose [= Conzattia multiflora (B.L. Rob.) Standl.]
Unarmed, small to medium-sized tree, 3–10 (15) m, generally forking low down with a large spreading flat-topped ‘umbrella’ crown (Fig.
Unknown.
Confined to seasonally dry tropical forest. Strongly deciduous, fruits ripening when leafless (Fig.
Named in honour of the Italian-born Mexican botanist Cassiano Conzatti (1862–1951), a prodigious plant collector in Mexico and Chile and director of the Escuela Normal in Oaxaca, Mexico.
Conzattia has been listed as an important source of medicine and wood for construction and fuel, and immature fruits are used as a minor food in Mexico (
Conzattia is robustly supported as sister to Heteroflorum (Fig.
Heteroflorum sclerocarpum M. Sousa
Unarmed small trees to 15 m, forked near the base, forming a wide spreading umbrella crown (Fig.
Unknown.
Monospecific. A narrowly restricted endemic of southern Mexico, disjunctly distributed close to the Pacific coast in the lower reaches of the Río Balsas watershed around El Infiernillo (Michoacán and Guerrero) and in coastal Oaxaca (Fig.
Confined to seasonally dry tropical forest, occasionally forming dominant stands (
The name Heteroflorum is derived from hetero- (Greek = different, uneven) and -florum (plural of flos; Latin = flower) in reference to the different sized functionally male and female flower forms associated with dioecy.
Unknown.
Heteroflorum is robustly supported as sister to a second monospecific Mexican endemic genus, Conzattia (Fig.
Colvillea racemosa Bojer ex Hook.
Unarmed, small to medium-sized trees, 8–15 (20) m and 50–70 (90) cm stem diameter (Fig.
2n = 28 (
Seasonally dry forests and spiny thickets. Trees are deciduous. The flowers are nectariferous and although often eaten by lemurs, they are pollinated by Siouimanga sunbirds (Du Puy and Rabevohitra 2002).
Named in honour of Sir Charles Colville (1770–1843), distinguished Scottish officer under Wellington in the Napoleonic wars and governor of Mauritius, 1828–1834.
Sometimes planted as an ornamental in villages (Fig.
The status of Colvillea as a genus distinct from Delonix remains questionable as its placement either as sister to Delonix, or nested within it is unstable in phylogenies that have densely sampled Delonix (
Aprevalia Baill., Bull. Mens. Soc. Linn. Paris 1: 428. 1884. Type: Aprevalia floribunda Baill. [≡ Delonix floribunda (Baill.) Capuron]
Lemuropisum H. Perrier, Bull. Soc. Bot. France 85: 494. 1939. Type: Lemuropisum edule H. Perrier [≡ Delonix edulis (H. Perrier) Babineau & Bruneau]
Delonix sect. Aprevalia (Baill.) Capuron, Adansonia, n.s. 8(1): 12. 1968. Type: Delonix floribunda (Baill.) Capuron [≡ Aprevalia floribunda Baill.]
Delonix regia (Bojer ex Hook.) Raf. [≡ Poinciana regia Bojer ex Hook.]
Shrubs (D. edule) or small to medium-sized trees to 30 m, often forking low down and forming spreading, flat-topped umbrella crowns (Fig.
2n = 26 or 28 (
Twelve species, ten endemic to south-western, western and northern Madagascar, one widespread in eastern and north-eastern Africa extending to adjacent Arabia, and one restricted to northern Kenya and Somalia (Fig.
Seasonally dry and arid thickets including the spiny dry forests of south-western Madagascar, Acacia-Commiphora woodland in north-eastern Africa, and semi-desert scrub. A few species have distinctive swollen trunks and ‘bottle tree’ habits (Fig.
From Greek, delo- (= evident, conspicuous) and -onyx (= claw or nail) in reference to the fact that the petals have very long claws.
The well-known Flamboyant or Flame tree, Delonix regia with its magnificent showy racemes of scarlet flowers is a favourite in cultivation as an ornamental street and garden tree throughout the tropics (
The genus Delonix is sister to the morphologically similar Colvillea. These two genera together are robustly supported as sister to the clade comprising two monospecific endemic Mexican genera Conzattia and Heteroflorum (Fig.
Haroldo Cavalcante de Lima8,27, Isau Huamantupa-Chuquimaco23, Domingos B. O. S. Cardoso8,9
Citation: Lima HC, Huamantupa-Chuquimaco I, Cardoso DBOS (2024) 8. Tribe Sclerolobieae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 165–176. https://doi.org/10.3897/phytokeys.240.101716
Tachigalieae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Tr. Nov.]: 252. 1943. Type: Tachigali Aubl.
Sclerolobium Vogel [= Tachigali Aubl.]
(5). Arapatiella Rizzini & A. Mattos (2 species), Diptychandra Tul. (2), Jacqueshuberia Ducke (7), Moldenhawera Schrad. (12), Tachigali Aubl. (80–90).
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.
Mostly Neotropical, extending from Central to South America, in tropical rainforests, tropical cloud forests, seasonally dry forests, and fire-prone savannas.
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.
Although the generic name Sclerolobium is currently treated under Tachigali, the tribe name Sclerolobieae (
Not only have the relationships amongst Sclerolobieae genera now been strongly resolved for almost all nodes (Fig.
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.
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.
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.
Dolichonemia Nees, Flora 4 (19): 303. 1821. Type: Dolichonemia speciosa Nees [= Moldenhawera floribunda Schrad.]
Moldenhawera floribunda Schrad.
Trees or shrubs, the young branches often rust-coloured with T-shaped trichomes. Stipules often pinnate. Leaves paripinnate; bipinnate, partially bipinnate; leaflets opposite. Inflorescences in terminal corymbiform racemes, fasciculate or solitary; bracts and bracteoles setaceous, caducous. Flowers apparently radially symmetrical, but essentially bilaterally symmetrical because of the single large fertile stamen; hypanthium infilled; sepals (4) 5, free; petals (4) 5, yellow, rarely pink (M. acuminata Afr. Fern. & P. Bezerra), clawed, the margins crimped; androecium dimorphic, fertile stamen 1, filament elongated, connective hairy, anther dehiscing through longitudinal slits, staminodes 7 or 9, filaments short or elongated, anthers dehiscing by longitudinal slits or pores, or indehiscent; pollen grains in monads, perforate to an almost vermiculate-reticulate tectum, scabrate-punctate; ovary sessile, stigma truncate. Fruit a dehiscent, linear, compressed legume, the two valves woody and coiling at dehiscence, 4–8-seeded. Seeds ovate to oblong, compressed.
Unknown.
Twelve species and one variety, exclusively in Brazil, mostly along the eastern Atlantic coast (Bahia, Espírito Santo and Rio de Janeiro states), but two species are in montane areas of inland Bahia and Minas Gerais states and one species is in northern Brazil (Maranhão and Piauí states) (Fig.
Most species occur in the Atlantic Forest, commonly in moist coastal forest or associated with coastal vegetation on sand soil (restinga). Three species occur in savanna vegetation, mainly in rocky grasslands.
The genus is named after Johann Jakob Moldenhawer (1766–1827), German professor of botany at Kiel and one of the founders of plant anatomy.
The yellow densely flowered inflorescences make the genus potentially of interest as an ornamental for gardens and as street trees. The red timber from M. intermedia G.P. Lewis & L.P. Queiroz is used for furniture (
Moldenhawera is characterised by the Malpighiaceae-like bilaterally symmetrical flowers, hypanthium infilled and the unusual androecium with a single fertile stamen with bearded connective and seven or nine short staminodes. The genus presents variation in the leaf division, with pinnate and/or bipinnate leaves, sometimes in the same species. The last taxonomic revision of the genus, including a key to all species, was provided by
Diptychandra aurantiaca Tul.
Trees or shrubs, unarmed. Stipules scale-like. Leaves paripinnate; leaflets 2–4, opposite. Inflorescences in terminal unbranched racemes, 3.5–20 cm long; bracts and bracteoles minute, caducous. Flowers radially symmetrical; hypanthium cupulate; sepals 5, free, often with red dots; petals 5, yellow, clawed; androecium monomorphic, stamens 10, free; pollen in tetrads, moderately reticulate; ovary stipitate, stigma punctiform. Fruit a dehiscent, linear-oblong, compressed legume, the two valves ligneous with resinous dots, 1–2 (4)-seeded. Seeds ellipsoid-complanate, winged.
Unknown.
Three species, one of which is widespread throughout most of western Brazil extending to Bolivia and Paraguay; and the remaining two are each endemic to north-eastern Brazil and Colombia (Fig.
Mostly in savannas and seasonally dry forests, except for D. granadillo C. Romero & Arbeláez which occurs in Andean mountain forests.
The generic name is derived from Greek, diptycho- (= twice folded) and -andro (= stamen), the stamen filaments are twice folded in bud.
Species of the genus are used as timber for construction and charcoal. The wood of D. granadillo has been reported to be very hard and difficult to saw (
Diptychandra is characterised by the paripinnate leaves and the flowers radially symmetrical with cupulate hypanthium and sepals often with red dots. The fruits are dehiscent, linear-oblong, bearing valves with resinous dots and 1–2(4) winged seeds. The last taxonomic revision of the genus, including a key to all species, was provided by
Arapatiella trepocarpa Rizzini & A. Mattos [= Arapatiella psilophylla (Harms) R.S. Cowan]
Trees, unarmed. Stipules foliaceous, orbiculate to reniform. Leaves paripinnate; leaflets 2–4, opposite. Inflorescences in terminal paniculate racemes, 4–10 cm long; bracts and bracteoles small, caducous. Flowers radially symmetrical; hypanthium turbinate; sepals 5, free; petals 5, white, clawed; androecium monomorphic, stamens 10, free; pollen grains in monads, associated with viscin threads, coarsely reticulate; ovary stipitate, stigma disciform-peltate. Fruit a linear-oblanceolate, laterally compressed legume, elastically dehiscent, the woody valves coiling backwards from the apex at dehiscence, 4–5 (8)-seeded. Seeds ellipsoid-compressed.
Unknown.
The genus is only known from lowland Atlantic moist coastal forests on “tabuleiro terciário” of southern Bahia and northern Espírito Santo states in Brazil.
The generic name is derived from the local vernacular name “arapati”.
The species of the genus are used as timber for construction.
Arapatiella is characterised by the foliaceous stipules, orbiculate to reniform, leaves paripinnate, and flowers radially symmetrical with turbinate hypanthium and ten exserted stamens. The fruits are elastically dehiscent from the apex, the woody valves coiling backwards at dehiscence. A taxonomic study of the genus, including a key to all species, was provided by Lima and Kuntz (2020).
Jacqueshuberia quinquangulata Ducke
Small trees or shrubs, unarmed. Stipules pinnate. Leaves bipinnate; pinnae and leaflets opposite, (2) 4–30 pairs of pinnae, 7–80 pairs of leaflets per pinna. Inflorescences terminal racemes, elongated or corymbiform; bracts and bracteoles setaceous, caducous. Flowers bilaterally symmetrical; hypanthium cupulate or campanulate, slightly ribbed; sepals 5, free; petals 5, yellow or dark purplish red, ovate, slightly unequal, sessile, lacking a claw; androecium monomorphic, stamens 10, joined in lower part; pollen in monads, associated with viscin threads, coarsely reticulate; ovary sessile, stigma oblique capitate. Fruit a dehiscent, linear, compressed legume, ribbed on the edges, the two ligneous valves coiling backwards from the apex at dehiscence, 4–8-seeded. Seeds oblong-ellipsoid, compressed.
Unknown.
Seven species across north-western South America, where two are endemic in Amazonian Brazil, two in Venezuelan Guayana, one in Guyana, one in Colombia and one in Peru (Fig.
White sand forests of the Amazon basin, montane forests on sandstone and savannas in the Guiana Shield.
The generic name honours Jacques E. Huber (1867–1914), a Swiss botanist who explored the Amazon region.
Potentially of interest as an ornamental because of its wide range of flower colour, including red, purple, and yellow.
Jacqueshuberia is characterised by the combination of foliaceous and pinnate stipules, bipinnate leaves, the pinnae and leaflets opposite, flowers bilaterally symmetrical with a cupular hypanthium and ten exserted stamens. The fruits are dehiscent, linear, with 4–8 seeds. The genus was last revised by
Cuba Scop., Intr. Hist. Nat.: 300. 1777. Type not designated.
Cubaea Schreb., Gen. Pl.: 278. 1789. Type not designated.
Valentinia Neck., Elem. Bot. 2: 450. 1790, opus utique oppr.
Tachia Pers., Syn. Pl. 1: 459. 1805, non Tachia Aublet, Hist. Pl. Guiane: 75. 1775. (Gentianaceae). Type: Tachia paniculata Pers. [≡ Tachigali paniculata Aubl.]
Sclerolobium Vogel, Linnaea 11: 395. 1837. Type: Sclerolobium denudatum Vogel [≡ Tachigali denudata (Vogel) Oliveira-Filho]
Tachigali paniculata Aubl.
Trees, unarmed. Stipules foliaceous, pinnate or pectinate; persistent to caducous. Leaves paripinnate; leaflets 2–20 pairs, opposite and inversely symmetrical; petiole and/or rachis usually with myrmecophilous domatia. Inflorescences in paniculate terminal racemes or in leaf axils of terminal branches; bracts equal in shape to, but smaller than, the stipules; bracteoles minute, lanceolate or subulate. Flowers radially or bilaterally symmetrical; hypanthium cupulate or obliquely cylindrical; sepals 5, free; petals 5, yellow or orange, lineate, lanceolate or spathulate, sometimes clawed; stamens 10, rarely 15–16, monomorphic with equal filaments, or dimorphic with 7 filaments longer, subulate, and 3 shorter, falcate or sigmoidal; pollen in monads, finely reticulate; ovary stipitate, stigma truncate. Fruit an indehiscent, compressed, oblong-elliptic or oblong, 1–3-seeded cryptosamara; exocarp flaking at maturity; mesocarp surrounded by a subligneous and thin wing; endocarp hyaline and membranous. Seeds oblong-ellipsoid, compressed.
2n = 24 (
Seventy-eight formally described species, but recent taxonomic estimates suggest the genus may include more than 90 species (
Three species occur in evergreen and semi-deciduous lowland forests of Central America, T. costaricensis (N. Zamora & Pùveda) N. Zamora & van der Werff, T. panamensis van der Werff & N. Zamora, and T. versicolor Standl. & L.O. Williams (
The generic name is derived from the vernacular name “tachi” for stinging ants.
Used as timber for construction and charcoal. Tachigali vulgaris L.F. Gomes da Silva & H.C. Lima is planted in forest restoration (
Tachigali is recognisable by the combination of paripinnate leaves, the leaflets inversely symmetrical, stipules foliaceous and mostly pinnate, the petiole and/or rachis usually with myrmecophilous domatia, and very distinct strongly laterally compressed wind-dispersed fruits (cryptosamaras). Floral symmetry marks a major subdivision within Tachigali, and was once used to define two genera (
Guilherme Sousa da Silva39, Marcelo F. Simon40
Citation: Silva GS, Simon MF (2024) 9. Tribe Dimorphandeae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 177–186. https://doi.org/10.3897/phytokeys.240.101716
Dimorphandrinae Walp., Repert. Bot. Syst. 1: 854. 1843. Type: Dimorphandra Schott
Moreae Britton & Rose, N. Amer. Fl. 23: 201, 217. 1930. Type: Mora Benth.
Dimorphandra Schott
(4). Burkea Hook. (1 species), Dimorphandra Schott (26), Mora Benth. (6), Stachyothyrsus Harms (2).
Unarmed shrubs, treelets to canopy trees up to 40 m high; trunk buttressed or not; brachyblasts absent; branches glabrous or pilose. Stipules present or absent. Leaves pinnate or bipinnate; extrafloral nectaries on the petiole occurring only in Stachyothyrsus; pinnae (in bipinnate leaves) 1-many pairs; leaflets 1-many pairs, opposite or alternate, variable in size and shape. Inflorescences short or elongated spiciform racemes or spikes, often arranged in corymbose or paniculate synflorescences; bracteoles small or absent. Flowers 5-merous, diplostemonous; stamens alternate, sometimes with 5 fertile and 5 staminodes; anther glands absent or present (Burkea); pollen tricolpate monads. Fruit a typical legume or samara (Burkea), dehiscent or indehiscent, variable in size and shape, 1-multiseeded. Seeds flat-compressed to ovoid, with a hard or thin testa, areolas absent, present on both sides only in Burkea.
Rainforests, seasonally dry forests, and savannas, in tropical regions of the Americas and Africa.
The most inclusive crown clade containing Dimorphandra gardneriana Tul. and Burkea africana Hook., but not Campsiandra comosa Benth., Tachigali guianensis (Benth.) Zarucchi & Herend. or Schizolobium parahyba (Vell.) S.F. Blake (Fig.
The informal Dimorphandra group of the old sense subfamily Caesalpinioideae, which originally included 10 genera (
Members of the Dimorphandreae (Figs
Leaves and inflorescences of tribe Dimorphandreae A Mora paraensis (Ducke) Ducke (Simon 1663) B Stachyothyrsus staudtii Harms C Burkea africana Hook. D Dimorphandra mediocris Ducke (Simon 4209) E Dimorphandra cuprea Sprague & Sandwith (Farroñay 1804) F Dimorphandra parviflora Spruce ex Benth. (Simon 1176) G Dimorphandra mollis Benth. H Dimorphandra pennigera Tul. I Dimorphandra ignea Ducke J Dimorphandra vernicosa Spreng. ex Benth. (Cardoso 3279) K Burkea africana L Dimorphandra gardneriana Tul. (Silva 21). Photo credits A, D, F, G MF Simon B Nicolas Texier (CC-BY-NC-ND-3.0) C AR Lecuona (CC-BY-NC-4.0) E, H, I F Javier Farroñay Pacaya J D Cardoso K AE van Wyk and S Malan L G Sousa da Silva.
Flowers and fruits of the tribe Dimorphandreae A Dimorphandra cuprea Sprague & Sandwith (Farroñay 1804) B Dimorphandra pennigera Tul. C Dimorphandra vernicosa Spreng. ex Benth. (Cardoso 3279) D Dimorphandra ignea Ducke E Stachyothyrsus staudtii Harms F Dimorphandra gardneriana Tul. (Simon 2715) G Dimorphandra macrostachya Benth. (Silva 540) H Dimorphandra cuprea Sprague & Sandwith (Farroñay 1804) I Stachyothyrsus staudtii Harms J Burkea africana Hook. K–M Mora paraensis (Ducke) Ducke (K, L Costa 28). Photo credits A, B, D, H F Javier Farroñay Pacaya C D Cardoso E Flora of the world F MF Simon G G Sousa da Silva I N Texier (CC-BY-NC-ND-3.0) J C Sydes (CC-BY-NC-4.0) K, L J Barbosa Pedrosa Costa M A Rocha Dantas.
Although Dimorphandreae forms a well-supported clade, Dimorphandra, the largest genus in the tribe, was recovered as non-monophyletic (Fig.
Dimorphandra exaltata Schott
Unarmed small or medium trees 3–5 (7) m, to canopy trees 40 (50) m; trunk buttressed or not; brachyblasts absent; usually pubescent in all parts with reddish to dark or light brown hairs. Stipules present or absent, caducous. Leaves bipinnate, rachis 5–90 cm long; pinnae 1–40 or more pairs; leaflets 1–50 pairs, opposite or alternate, commonly glabrous above and pubescent below, venation brochidodromous. Inflorescences short or elongated spiciform racemes, often arranged in paniculate or corymbose synflorescences; bracteoles caducous or absent. Flowers small, pale yellow to cream or dark orange to reddish, fragrant, 5-merous, diplostemonous; calyx cupuliform, tubular or campanulate; petals obovate, oblong or spatulate, glabrous or pilose; fertile stamens 5, filaments thin, with oblong, dorsifixed anthers, anther glands absent; staminodes 5, spatulate, free or connate forming a dome, with or without a rudimentary anther at the apex, usually deciduous at anthesis; pollen tricolpate monads; ovary sessile, subsessile, or stipitate, multi-ovulate, glabrous or densely hairy, style short to absent, stigma conical, terminal. Fruit dehiscent or indehiscent legumes, linear-oblong, curved or suborbicular, flat, valves leathery or woody. Seeds orbicular, flat, cylindrical or oblong.
2n = 28 (
26 species, including four subspecies (
Predominates in tropical rainforests, including both terra firme and seasonally flooded (igapó) forests, white-sand forests (campinarana), savannas, and less often in seasonally dry forests. Fruits of D. mollis Benth. are important food resources for tapirs (
Dimorphandra refers to the androecium that has five stamens alternating with five staminodes, that is, two morphological forms of the stamens.
Dimorphandra species have the ability to fix nitrogen (
The genus is classified into three subgenera: Dimorphandra (11 species), Phaneropsia Tul. (5 species) and Pocillum Tul. (10 species), which can be differentiated by leaf morphology, inflorescence architecture and fruit shape (
Mora excelsa Benth.
Unarmed large trees, 15–45 m; trunk buttressed; brachyblasts absent; glabrous. Stipules small, caducous. Leaves paripinnate, alternate; petiole 2–6 cm long, rachis 5–10 cm long; leaflets (1) 2–6 (7) pairs, opposite, large, long-acuminate, glabrous and smooth, secondary veins inconspicuous. Inflorescences spiciform racemes arranged in paniculate synflorescences. Flowers small, white or yellow, 5-merous, diplostemonous; bracteoles small, caducous; calyx with a very short tube and short ciliated lobes; petals oblong or ovate, finely ciliated at the apex; fertile stamens 5, filaments thick, anthers covered with caducous white hairs, anther glands absent; staminodes 5; pollen tricolpate monads; ovary sessile or nearly so, few-ovuled, style compressed with a thin terminal stigma. Fruit a dehiscent legume, flat, elliptic or oblong, coriaceous to woody, the valves twist after dehiscence. Seeds large, flattened or suborbicular, with a membranous testa.
Unknown.
The genus comprises six species occurring in Central America (Panama, Costa Rica), northern South America (Brazil, Colombia, Ecuador, Guyana, Suriname, Venezuela) and Greater Antilles (Haiti, Dominican Republic and Trinidad Tobago) (Fig.
Mora species generally occur in periodically flooded forests, swamps and mangroves. Mora excelsa forms monodominant forests in the Guianas (
Derived from the widely used Arawak vernacular name ‘mora’.
Mora species are valuable timber in the Guianas, being used in construction, industrial flooring and for charcoal (
Mora has a complex taxonomic history, having been treated under Dimorphandra by several authors. Mora can be readily distinguished from Dimorphandra by the paripinnate leaves, which are always glabrous (vs. bipinnate, often pubescent); anthers with conspicuous caducous white hairs (vs. glabrous); style longer than the ovary (vs. shorter); large, soft seeds, with a membranous testa (vs. small seeds with a hard testa) (
Kaoue Pellegr., Bull. Soc. Bot. France 80: 464. 1933. Type: Kaoue stapfiana (A. Chev.) Pellegr. [≡ Oxystigma stapfiana A. Chev. (≡ Stachyothyrsus stapfiana (A. Chev.) J. Léonard & Voorh.)]
Stachyothyrsus staudtii Harms
Unarmed trees 20–25 (30) m; trunk not buttressed; brachyblasts absent; glabrous. Stipules small, caducous. Leaves bipinnate, rachis 4–20 cm; pinnae 2 pairs; leaflets 3–5 pairs per pinna, opposite, glabrous, venation reticulate. Inflorescences spiciform racemes, sometimes arranged in paniculate synflorescences; bracts triangular-rounded, persistent. Flowers small, whitish, fragrant, 5-merous, diplostemonous; calyx short, cupuliform; petals obovate-oblong; stamens dimorphic, the 5 antesepalous slightly longer than the 5 antepetalous (shorter), anther glands absent; pollen tricolpate monads; ovary short, 2–3-ovulate, style short, stigma slightly bilobed. Fruit dehiscent legume, curved, flat, 1–2-seeded. Seeds irregular shaped, dark.
Unknown.
Two species, restricted to the central-west coast of Africa in Ivory Coast, Liberia and Sierra Leone (S. stapfiana J. Leonard & Voorhoeve), and Cameroon, Gabon, Equatorial Guinea and Congo (S. staudtii) (Fig.
Stachyothyrsus occurs in tropical rainforests, including swamps, along river banks, and secondary forests.
From Greek, Stachys (= spike) and thyrsus (= wand, panicle), in reference to the spicate inflorescences aggregated into showy panicles.
Stachyothyrsus stapfiana leaves are used for thatching, while its wood has only limited importance (
Stachyothyrsus is most closely related to Burkea (the genera have stamens arising at the insertion or just slightly above the ovary insertion), but they are easily distinguished by the opposite, less numerous and larger leaflets in Stachyothyrsus. In addition, Stachyothyrsus species occur only in rainforests along the west coast of Africa, whereas Burkea is widely distributed in African savannas and dry forests.
Burkea africana Hook.
Unarmed shrubs or small trees, exceptionally reaching 20–30 (35) m; trunk not buttressed; brachyblasts absent; indumentum on leaves and inflorescences composed by simple ferrugineous trichomes. Stipules minute, caducous. Leaves bipinnate, clustered at the tip of the branches; petiole and rachis together 7–32 cm long; pinnae (1) 2–5 (7) pairs; leaflets 5–15 (18) per pinna, alternate. Inflorescences elongate spiciform racemes 5–30 cm long, crowded at the tip of branches. Flowers small, whitish, sweet-scented, 5-merous, diplostemonous; calyx campanulate; petals obovate or elliptic-obtuse, glabrous; stamens 10, homomorphic; anthers oblong, anther glands present; pollen tricolpate monads; ovary subsessile, hirsute, 1–2-ovulate, style very short, thick, stigma conspicuous, capitate. Fruit samaroid, oblong or elliptical, flat, indehiscent, 1 (2)-seeded. Seed obovate, compressed, albuminous, cotyledon thin, flat, radicle short.
2n = 28 (
Monospecific (B. africana), widespread in Africa (except for the rainforest regions), occurring mainly in the west, centre and south of the continent, extending into Senegal, Sudan and Uganda, south to Namibia, Botswana and northern South Africa (Fig.
The species inhabits savannas and seasonally dry forests at elevations from 40–1740 m. Trees with a bole diameter above 12.5 cm are fire resistant, being sufficiently protected by their corky bark. It is frequent and abundant in many regions (e.g., southern Africa), but often occurs in a dispersed and non-aggregated pattern.
Named after the British botanist Joseph Burke (1812–1873), who collected plants and animals (especially in South Africa) for the Earl of Derby.
An edible gum is produced from the stem, tender young leaves are cooked and eaten as a vegetable while young flowers are eaten in sauces. The bark, roots and leaves are commonly used in traditional medicine (
Burkea is the most unusual genus within the tribe, with young leaves and inflorescences clustered at the tip of shoots, capitate stigma, and samaroid, monospermic fruits, whereas the other genera have leaves and inflorescence which are not clustered at the tip of shoots, small stigma, and multi-seeded legumes. The dispersal of the flat and dry fruits of Burkea is possibly by wind since the trees occur in dry and open environments (
Gwilym P. Lewis10
Citation: Lewis GP (2024) 10. Tribe Campsiandreae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 187–192. https://doi.org/10.3897/phytokeys.240.101716
Campsiandreae is characterised by a combination of flowers with a cupular hypanthium, imbricate petals and stamens long-exserted from the corolla. The genus Dinizia is similar in appearance to genera of tribes Dimorphandreae and Mimoseae by its bipinnate leaves and small flowers in dense spicate racemes but differs by the combination of alternate leaflets, imbricate petal aestivation and all stamens fertile (in Mimoseae petals are valvate and genera of Dimorphandreae with bipinnate leaves and spicate inflorescences have opposite leaflets and an androecium comprising 5 fertile stamens and 5 staminodes). The genus Campsiandra is similar in appearance to Arapatiella Rizzini & A. Mattos (tribe Sclerolobieae) in having paripinnate leaves and regular perigynous flowers with a turbinate hypanthium and exserted stamens but differs by the variable number of stamens (vs. stamens 10 in Arapatiella), fruits indehiscent or inertly dehiscent, the valves not strongly twisting after dehiscence (vs. fruits elastically dehiscent from the apex, the valves rolling inwards in Arapatiella).
Campsiandra Benth.
(2). Campsiandra Benth. (3 to ca. 20 species), Dinizia Ducke (2).
Medium to large trees. Stipules caducous or lacking. Leaves pinnate with opposite, often gland-dotted leaflets, or bipinnate with alternate, eglandular leaflets; extrafloral nectaries absent. Inflorescence a compound spiciform raceme, or showy multi-flowered panicle. Flowers hermaphrodite or functionally staminate, pedicellate (the pedicels up to 3 cm long in Campsiandra); calyx with 5 imbricate lobes or a tube with 5 broadly triangular lobes; petals 5, imbricate, whitish (sometimes with rose-reddish markings), cream coloured or yellow; stamens 10–17 (25) per flower, exserted from the corolla, anthers eglandular; pollen in monads or tetrads; ovary stipitate. Fruit laterally compressed, coriaceous or woody, inertly dehiscent or indehiscent. Seeds discoid with a marginal spongy wing, or elliptic to obovate, hard, and wingless.
South America, mainly in the Amazon and Orinoco basins in flooded forests and swamp forests (Campsiandra) or in non-flooded Amazonian forests (Dinizia excelsa Ducke), or semi-deciduous Atlantic rainforest (Dinizia jueirana-facao G.P. Lewis & G.S. Siqueira).
The most inclusive crown clade containing Campsiandra laurifolia Benth. and Dinizia excelsa Ducke, but not Delonix decaryi (R. Vig.) Capuron, Dimorphandra conjugata (Splitg.) Sandwith or Mimosa sensitiva L. (Fig.
Phylogenetic analyses of a few molecular markers had suggested that the genus Dinizia could be more closely related to the Dimorphandra group of the old sense subfamily Caesalpinioideae, but with low support (
(here designated). Campsiandra laurifolia Benth.
Medium to large trees (6–25 m tall). Stipules inconspicuous and caducous. Leaves imparipinnate, 7–13-foliolate, leaflets in opposite pairs plus a single terminal leaflet, often gland-dotted although these frequently obscured by a waxy epidermal layer; extrafloral nectaries absent. Inflorescence a multi-flowered, often showy, terminal panicle (Fig.
Flower, fruit and vegetative characters of tribe Campsiandreae A foliage and fruits, Campsiandra comosa Benth., Brazil B compound inflorescence, Campsiandra angustifolia Spruce ex Benth., Peru C fruit and seed, Campsiandra laurifolia Benth., cultivated Rio de Janeiro Botanic Gardens, Brazil D inflorescence and foliage, Campsiandra sp., Amazonas, Brazil E–I Dinizia jueirana facao G.P. Lewis & G.S. Siqueira, Reserva Natural Vale, Espírito Santo, Brazil E foliage and fruits F trunk and crown of mature tree G inflorescences H mature fruits (hand for scale) I rough bark of trunk. Photo credits A D Cardoso B T Pennington D M Falção E, G, I D Folli C, F, H GP Lewis.
Unknown.
The number of species in the genus varies greatly depending on the treatment consulted.
Mainly in the Amazon and Orinoco basins in flooded forests, swamp forests (of both black and white-water rivers), on alluvial plains, white sand riverine beaches and embankments, with seeds being mostly water dispersed.
The seeds of a few species are used locally to make a flour and medicinally (
From Greek, campso, campsis (= bending, a bend) and andro- (= man, anther) referring to the long wavy stamens of especially the first described species.
Campsiandra was originally placed in the informal Peltophorum group of tribe Caesalpinieae sensu
Dinizia excelsa Ducke
Large forest canopy-emergent unarmed trees (Fig.
2n = 26 (28) (D. excelsa) (
Two species, one widespread in northern and central-western Amazonian Brazil, Guyana, and Suriname (D. excelsa), the other narrowly restricted to a small area of Eastern Brazil in Espirito Santo state (D. jueirana-facao) (Fig.
Non-flooded Amazonian forests (D. excelsa), or semi-deciduous Atlantic rainforest (D. jueirana-facao).
Named by Ducke for his friend José Picanço Diniz, doctor-in-law and philanthropist.
The wood of D. excelsa is very resistant and has been widely used in civil and naval construction, for railway sleepers, cabinetwork and joinery, as well as for battens, props, beams, girders, posts, stakes, door and window frames, floor-boards, carts, wagons and bridges (
The genus was placed in its own Dinizia group of tribe Mimoseae by
Luciano Paganucci de Queiroz2, Anne Bruneau1
Citation: Queiroz LP, Bruneau A (2024) 11. Tribe Erythrophleeae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 193–200. https://doi.org/10.3897/phytokeys.240.101716
Unarmed trees or treelets with macrophyllidious bipinnate leaves, alternate leaflets, flowers shortly pedicellate, densely packed in elongate spicate racemes, small, regular, with a short cupular hypanthium, sepals and petals ascending, almost erect. Similar to the genera Adenanthera L., Amblygonocarpus Harms and Tetrapleura Benth. (Adenanthera clade, tribe Mimoseae) in habit of unarmed trees with ample bipinnate leaves with alternate leaflets and small pedicellate flowers in spicate racemes, but differing by the ascending perianth giving a closed aspect to the flowers (vs. flowers open because sepals and petals are reflexed backwards) and seeds lacking a pleurogram. Also differentiated from Adenanthera by fruits straight or slightly curved with thick woody valves (vs. valves thin coriaceous and twisted after dehiscence) and from Amblygonocarpus and Tetrapleura by the fruits with flat valves [vs. tetragonal with a median rib (Amblygonocarpus) or wing (Tetrapleura) on each valve].
(designated here). Erythrophleum Afzel. ex R. Br.
(2). Erythrophleum Afzel. ex R. Br. (12 species), Pachyelasma Harms (1).
Unarmed trees or treelets; trunk with rough bark and a reddish sap when cut, brachyblasts absent. Stipules inconspicuous, mostly caducous. Leaves bipinnate, ample, macrophyllidious, with few pinnae and few leaflets per pinna, leaflets alternate, elliptical to oblong, frequently asymmetrical, pinnately veined. Inflorescences spicate racemes clustered in terminal or axillary panicles. Flowers perigynous, shortly pedicellate, 5-merous, bisexual, sepals and petals ascending, almost erect, the perianth almost cylindrical; stamens 10, free, the filaments glabrous or pubescent, anthers dehiscing through longitudinal slits; pollen in tricolporate monads; ovary stipitate, pluriovulate, style conical to cylindrical. Fruit dehiscent or, rarely, indehiscent, valves stiffly coriaceous or resinous, endocarp not septate nor breaking into one-seeded envelopes. Seeds slightly compressed, without pleurogram.
The most inclusive crown clade containing Erythrophleum suaveolens (Guill. & Perr.) Brenan and Pachyelasma tessmannii (Harms) Harms, but not Campsiandra laurifolia Benth., Dimorphandra conjugata (Splitg.) Sandwith or Pentaclethra macrophylla Benth. (Fig.
Tropical Africa (including Madagascar), eastern and south-eastern Asia and Australia.
The new tribe Erythrophleeae is here proposed to include Erythrophleum and Pachyelasma, two genera which were previously included in the informal Dimorphandra group of old sense tribe Caesalpinieae (sensu
Erythrophleum and Pachyelasma share a combination of morphological traits only rarely found in non-Mimoseae Caesalpinioideae, such as bipinnate leaves and small pedicellate perigynous flowers clustered in dense spicate racemes. Structural extrafloral nectaries that were characterised as parenchymatous and elevated with a small domed structure with a central pore (
Erythrophleum Afzel. ex G. Don, Gen. Hist. 2: 424. 1832. Type: Erythrophleum guineense G. Don [= Erythrophleum suaveolens (Guill. & Perr.) Brenan], nom. superfl.
Fillaea Guill. & Perr., Fl. Seneg. Tent. 1: 242, pl. 55. 1832. Type: Fillaea suaveolens Guill. & Perr. [≡ Erythrophleum suaveolens (Guill. & Perr.) Brenan]
Mavia G. Bertol., Mem. Reale Accad. Sci. Ist. Bologna 2: 570. 1850. Type: Mavia judicialis G. Bertol. [= Erythrophleum suaveolens (Guill. & Perr.) Brenan]
Laboucheria F. Muell., J. Proc. Linn. Soc., Bot. 3: 158. 1859. Type: Laboucheria chlorostachya F. Muell. [≡ Erythrophleum chlorostachys (F. Muell.) Baill.]
Erythrophleum suaveolens (Guill. & Perr.) Brenan
Unarmed trees to 30 m, treelets, rarely shrubs; trunk with rough bark; short shoots absent. Stipules very small, caducous. Leaves bipinnate; extrafloral nectaries present (7 species) or absent (3), with the secretory surface sunken in a pit capped by a small round pore (
2n = 24, 28 (
Twelve species, five in tropical sub-Saharan Africa and Madagascar, four in eastern and south-eastern Asia (south-eastern China, Cambodia, Laos, Vietnam, Thailand, Taiwan) and three in northern Australia (Fig.
Habit, bark and foliage of Erythrophleeae A Erythrophleum fordii Oliv. at Hùng Temple, Vietnam B bark of Erythrophleum couminga Baill. in Madagascar C Pachyelasma tessmannii (Harms) Harms in Congolian rainforest D, E, H Erythrophleum chlorostachys Baill. D rugous bark E tree in northern Australian savannas H foliage F Erythrophleum lasianthum Corbishley foliage G Erythrophleum suaveolens (Guill. & Perr.) Brenan foliage and terminal inflorescences, Ntumbachusi falls, Zambia. Photo credits A Hungda (https://tropical.theferns.info/image.php?id=Erythrophleum+fordii) B Solofo Eric Rakotoarisoa, iNaturalist (https://www.inaturalist.org/photos/42581094) C Bart Wursten (https://www.flickr.com/photos/zimbart/8212743587/in/photolist-dvJsxn-dvJnwi) D, E, H G Mahajan (https://alchetron.com/Erythrophleum-chlorostachys) F JMK (https://wikiwand.com/en/Erythrophleum_lasianthum) G MG Bingham (https://malawiflora.com/speciesdata/image-display.php?species_id=126540&image_id=8).
Tropical lowland wet forests in western Africa and south-eastern Asia, seasonally dry forests, woodlands and savannas in central-southern Africa and Australia.
From Greek, erythros (= red) and phloio (= bark), in reference to the red juice which flows from the trunk when cut.
Almost all parts of the plants of species of Erythrophleum are highly toxic for humans and livestock, due mostly to alkaloids, which have a Digitalis-like action on the heart, and to saponins (
The crushed bark of some species (E. lasianthum and E. suaveolens) is used as a fish and rat poison and crushed seeds are used as a component of arrow poison (
Timber of some species, usually named as ‘Tali’, is used for railway sleepers, boat building and canoes, heavy construction and joinery, firewood, and charcoal (
Together the twelve species of Erythrophleum comprise a morphologically cohesive genus, diagnosed by the combination of macrophyllidious bipinnate leaves, pedicellate flowers with straight and almost erect sepals and petals, and a long stipitate ovary tapering to a short conical style. The genus can be differentiated from Pachyelasma by the length of the ovary stipe (long in Erythrophleum vs. short in Pachyelasma), flower colour (green to greenish-yellow vs. reddish) and fruits (thin woody or leathery valves and margins not thick vs. resinous valves and thick, raised margins).
There is some controversy regarding the date and place of publication and the type species of the genus.
Seven of the twelve species have been included in various molecular phylogenetic and genetic studies, all of which support the monophyly of the genus as sampled (
Extrafloral nectaries were first reported in the 1980’s and they are now known to be present on the leaf rachis of at least seven of the twelve species (
Pachyelasma tessmannii (Harms) Harms [≡ Stachyothyrsus tessmannii Harms]
Unarmed trees, frequently emergent above forest canopies, to 60 m and 2.5 m diameter (Fig.
Unknown.
Monospecific (P. tessmannii), distributed in west tropical Africa (Nigeria, Cameroon, Central African Republic, Equatorial Guinea, Gabon, Congo, Democratic Republic of Congo; Fig.
Tribe Erythrophleeae inflorescences, fruits and seeds A–D Spicate racemes of species of Erythrophleum with small flowers, erect sepals and petals and hairy stamen filaments A E. suaveolens (Guill. & Perr.) Brenan with a visiting fly B E. chlorostachys (F. Muell.) Baill. (Ironwood) C, D E. lasianthum Corbishley E–I Fruits and seeds E dehisced fruits of Erythrophleum suaveolens F, G ripe fruits of Erythrophleum lasianthum H, I dispersed fruits of Pachyelasma tessmannii (Harms) Harms on the forest floor, showing the flat valves and raised margins. Photo credits: A AlkalIn (https://commons.wikimedia.org/wiki/File:Flowers_of_Erythrophleum_suaveolens.jpg) B T Harley (https://territorynativeplants.com.au/erythrophleum-chlorostachys-ironwood) C, D SAplants E Oliver Haumann, iNaturalist (https://www.inaturalist.org/photos/60428121) F, G JMK (https://commons.wikimedia.org/wiki/File:Erythrophleum_lasianthum,_loof_en_peule,_Manie_van_der_Schijff_BT,_a.jpg) H B Wursten (https://flickr.com/photos/zimbart/8212726705) I T Stévart (https://tropicos.org/ImageDownload.aspx?imageid=100336252).
Tropical lowlands rainforests of the Guinean and Congolian ecoregions.
From the Greek, pachy- (= thick) and elasmos (= plate), in reference to the thick pod.
Pachyelasma tessmannii, locally known as Mekogho and Mundumbula in Gabon, is used medicinally in various capacities. Fruits are used in traditional folk medicine to cure diarrhoea and abdominal pain (
Pachyelasma tessmannii is a dominant tree in West African rainforests, where it is one of the tallest trees, frequently emergent beyond the forest canopy. Its flowers are described as having a very unpleasant odour at night, which can be detected even from a distance of 300 m (Breteler 1026, P).
Luciano Paganucci de Queiroz2, Erik J. M. Koenen26, Colin E. Hughes3, Melissa Luckow29, Gwilym P. Lewis10, Jens J. Ringelberg3,4, Anne Bruneau1
Citation: Queiroz LP, Koenen EJM, Hughes CE, Luckow M, Lewis GP, Ringelberg JJ, Bruneau A (2024) 9. Tribe Mimoseae. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 201–206. https://doi.org/10.3897/phytokeys.240.101716
MimosaceaeR. Br., in M. Flinders, Voy. Terra Austral. 2: 551. 1814, nom. cons. Type: Mimosa L.
MimosoideaeDC., Prodr. [A.P. de Candolle] 2: 424. 1825. Type: Mimosa L.
AcacieaeDumort., Anal. Fam. Pl.: 40. 1829. Type: Acacia Mill., nom. cons.
AcaciinaeWight & Arn., Prodr. Fl. Ind. Orient.: 267. 1834. Type: Acacia Mill., nom. cons.
ParkiinaeWight & Arn., Prodr. Fl. Ind. Orient.: 279. 1834. Type: Parkia R. Br.
AcaciaceaeE. Mey., Comm. Pl. Afr. Austr. 1: 164. 1836. Type: Acacia Mill., nom. cons.
DesmanthinaeBenth., J. Bot. (Hooker) 2: 128. 1840. Type: Desmanthus Willd.
ParkieaeEndl., Gen. Pl.: 1323. 1840. Type: Parkia R. Br.
AdenantherinaeBenth., J. Bot. (Hooker) 4: 331. 1841. Type: Adenanthera L.
MimosineaeJ. Presl, Nowočeská Bibl. [Wšobecný Rostl.] 7: 346: 421. 1846. Type: Mimosa L.
AdenanthereaeBenth. & Hook.f., Gen. Pl. 1: 437. 1865. Type: Adenanthera L.
Ingeae Benth. & Hook.f., Gen. Pl. 1: 437. 1865. Type: Inga Mill.
PiptadenieaeBenth., Trans. Linn. Soc. London 30: 343, 358. 1875. Type: Piptadenia Benth.
DesmantheaeKuntze, in von Post & Kuntze, Lex. Gen. Phan.: 646. 1903. Type: Desmanthus Willd.
Mimozygantheae Burkart, Darwiniana 3: 447. 1939. Type: Mimozyganthus Burkart
AlbizieaeNakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 251. 1943. Type: Albizia Durazz.
Affonseeae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 251. 1943. Type: Affonsea A. St.-Hil. [= Inga Mill.]
Mimosa L.
Trees, shrubs, lianas, suffruticose or herbs, occasionally aquatic; unarmed or armed with prickles, spines or thorns. Stipules lateral and free or absent. Leaves bipinnate, less frequently paripinnate or modified into phyllodes (many Acacia, some Mimosa), rarely absent; pinnae and leaflets mostly opposite, rarely alternate; paraphyllidia (reduced basal leaflet pair on the pinnae) present or absent; specialised extrafloral nectaries often present on the petiole and/or on the primary and secondary rachides. Inflorescence globose, ellipsoid, umbelliform or corymbiform capitula, spikes or spiciform racemes; arising singly, paired or many from axillary fascicles, more frequently clustered in diversely arranged synflorescences. Flowers bisexual or frequently bisexual flowers combined with unisexual and/or sterile flowers in heteromorphic inflorescences, radially symmetrical; hypanthium mostly lacking; sepals and petals (3) 5 (6–8), mostly fused, sepals valvate in bud, rarely imbricate (Mimozyganthus, Parkia, Pentaclethra), petal aestivation valvate, rarely imbricate (Chidlowia, Sympetalandra), frequently the base of petals and stamens joined into a tube (stemonozone); stamens diplostemonous, haplostemonous or polystemonous, sometimes modified into showy staminodia, free or the filaments fused, anthers basifixed or dorsifixed, dehiscing via longitudinal slits, often with a stipitate or sessile apical gland; pollen commonly in tetrads, bitetrads or polyads, rarely in monads; gynoecium uni- or rarely polycarpellate, 1–many ovulate. Fruits 1–many-seeded, indehiscent or dehiscent along one or both sutures, often explosively or elastically dehiscent, also often lomentum or craspedium, the endocarp indistinct or separate and fragmented into 1-seeded envelopes. Seeds usually with an open (U-shaped) or closed (O-shaped) pleurogram on both faces, sometimes with a fleshy aril or sarcotesta; sometimes winged, hilum usually apical, lens usually inconspicuous; embryo straight. Root nodules present, indeterminate, and always symbiosome-type, or absent (at least 7 genera).
(100). Abarema Pittier (2 species), Acacia Mill. (1082), Acaciella Britton & Rose (15), Adenanthera L. (12), Adenopodia C. Presl (7), Afrocalliandra E.R. Souza & L.P. Queiroz (2), Alantsilodendron Villiers (11), Albizia Durazz. (ca. 90), Amblygonocarpus Harms (1), Anadenanthera Speg. (2–4), Anonychium (Benth.) Schweinf. (1), Archidendron F. Muell. (ca. 120), Archidendropsis I.C. Nielsen (11), Aubrevillea Pellegr. (2), Blanchetiodendron Barneby & J.W. Grimes (1), Boliviadendron E.R. Souza & C.E. Hughes (1), Calliandra Benth. (140), Calliandropsis H.M. Hern. & P. Guinet (1), Calpocalyx Harms (11), Cedrelinga Ducke (1), Chidlowia Hoyle (1), Chloroleucon (Benth.) Britton & Rose (10), Cojoba Britton & Rose (13–19), Cylicodiscus Harms (1), Desmanthus Willd. (23), Dichrostachys (DC.) Wight & Arn. (13–14), Ebenopsis Britton & Rose (3), Entada Adans. (40), Enterolobium Mart. (8), Faidherbia A. Chev. (1), Falcataria (I.C. Nielsen) Barneby & J.W. Grimes (3), Fillaeopsis Harms (1), Gagnebina Neck. ex DC. (7), Gretheria R. Duno & Torke (2), Gwilymia A.G. Lima, Paula-Souza & Scalon (7), Havardia Small (3), Heliodendron Gill.K. Br. & Bayly (3), Hesperalbizia Barneby & J.W. Grimes (1), Hydrochorea Barneby & J.W. Grimes (10), Indopiptadenia Brenan (1), Inga Mill. (ca. 300), Jupunba Britton & Rose (37), Kanaloa Lorence & K.R. Wood (1), Lachesiodendron P.G. Ribeiro, L.P. Queiroz & Luckow (1), Lemurodendron Villiers & Guinet (1), Leucaena Benth. (24), Leucochloron Barneby & J.W. Grimes (4), Lysiloma Benth. (8), Macrosamanea Britton & Rose ex Britton & Killip (12), Mariosousa Seigler & Ebinger (14), Marlimorimia L.P. Queiroz, L.M. Borges, Marc.F. Simon & P.G. Ribeiro (6), Mezcala C.E. Hughes & J.L. Contr. (1), Microlobius C. Presl (1), Mimosa L. (615), Mimozyganthus Burkart (1), Naiadendron A.G. Lima, Paula-Souza & Scalon (1), Neltuma Raf. (30), Neptunia Lour. (22), Newtonia Baill. (11), Osodendron E.J.M. Koenen (3), Painteria Britton & Rose (2), Parapiptadenia Brenan (6), Pararchidendron I.C. Nielsen (1), Parasenegalia Seigler & Ebinger (11), Paraserianthes I.C. Nielsen (1), Parkia R. Br. (ca. 35), Pentaclethra Benth. (3), Piptadenia Benth. (28), Piptadeniastrum Brenan (1), Piptadeniopsis Burkart (1), Pithecellobium Mart. (19), Pityrocarpa (Benth.) Britton & Rose (7), Plathymenia Benth. (1), Prosopidastrum Burkart (ca. 6), Prosopis L. (3), Pseudalbizzia Britton & Rose (17), Pseudoprosopis Harms (7), Pseudosamanea Harms (3), Pseudosenegalia Seigler & Ebinger (2), Punjuba Britton & Rose (5), Ricoa R. Duno & Torke (1), Robrichia (Barneby & J.W. Grimes) A.R.M. Luz & E.R. Souza (3), Samanea (Benth.) Merr. (3), Sanjappa E.R. Souza & M.V. Krishnaraj (1), Schleinitzia Warb. ex J.C. Willis (4), Senegalia Raf. (219), Serianthes Benth. (18), Sphinga Barneby & J.W. Grimes (3), Strombocarpa Engelm. & A. Gray (10), Stryphnodendron Mart. (28), Sympetalandra Stapf (5), Tetrapleura Benth. (2), Thailentadopsis Kosterm. (3), Vachellia Wight & Arn. (164), Viguieranthus Villiers (18), Wallaceodendron Koord. (1), Xerocladia Harv. (1), Xylia Benth. (9), Zapoteca H.M. Hern. (22), Zygia P. Browne (ca. 60).
Pantropical, with a few species extending marginally into warm temperate regions in North America and Asia, and extratropical South America, southern Africa and Australia.
The most inclusive crown clade containing Mimosa sensitiva L. and Pentaclethra macrophylla Benth., but not Pachyelasma tessmannii (Harms) Harms, Dimorphandra conjugata (Splitg.) Sandwith or Delonix decaryi (R. Vig.) Capuron (Fig.
Tribe Mimoseae as circumscribed here broadly coincides with the limits of the old sense subfamily Mimosoideae as adopted in several classical works (e.g.,
Phylogenetic studies have since shown that most of the genera included in the Mimosoideae comprise a monophyletic group, but nested in a paraphyletic old-sense subfamily Caesalpinioideae (
When revising the subfamilial classification for the Leguminosae, the Legume Phylogeny Working Group (
In Advances in Legume Systematics Part 1, five tribes were recognised in subfamily Mimosoideae: Parkieae, Mimoseae, Mimozygantheae, Acacieae and Ingeae (
Despite this putative hard polytomy along the backbone of the ingoid clade, the phylogenomic backbone of the Mimoseae of
The alternative solution for classification of this group, that of recognising multiple tribes within the mimosoid clade, is untenable given the imbalanced, “ladder-like” phylogenomic backbone of the mimosoid legumes (Fig.
Thus, in the following taxonomic arrangement, 25 treatments are presented for tribe Mimoseae (the numbers between brackets refer to the number of genera):
Tribe Mimoseae
13. Adenanthera clade (7 genera)
14. Sympetalandra (1)
15. Chidlowia (1)
16. Entada clade (3)
17. Newtonia grade (4)
Core mimosoid clade
18. Cylicodiscus (1)
19. Prosopis clade (2)
20. Neltuma clade (3)
21. Dichrostachys clade (14)
22. Parkia clade (3)
23. Lachesiodendron (1)
24. Stryphnodendron clade (7)
25. Mimosa clade (3)
Ingoid clade
26. Senegalia grade (4)
27. Calliandra clade (3)
28. Zapoteca clade (5)
29. Cojoba clade (3)
30. Pithecellobium clade (7)
31. Archidendron clade (9)
32. Cedrelinga (1)
33. Pseudosamanea (1)
34. Jupunba clade (4)
35. Samanea clade (2)
36. Albizia clade (3)
37. Inga clade (8)
Melissa Luckow29
Citation: Luckow M (2024) 13. Adenanthera clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 207–224. https://doi.org/10.3897/phytokeys.240.101716
Adenanthera clade
Figs
Included genera (7). Amblygonocarpus Harms (1 species), Adenanthera L. (12), Calpocalyx Harms (11), Pentaclethra Benth. (3), Pseudoprosopis Harms (7), Tetrapleura Benth. (2), Xylia Benth. (9).
Description. Predominantly large trees, occasionally lianas and small trees, unarmed except for uncinate lignified tendrils in climbing lianas of Pseudoprosopis; brachyblasts absent. Stipules small, linear to triangular, caducous. Leaves bipinnate, macrophyllous to microphyllous; pinnae opposite, subopposite, or alternate, few to many pairs per leaf; foliar nectaries absent except in Xylia and Calpocalyx, where they are present between the proximal pair of pinnae and sometimes between additional pinnae pairs, mounded or sunken into the petiole; leaflets opposite, subopposite, or alternate, usually petiolulate but sessile in Pentaclethra. Inflorescence usually racemose but sometimes spicate (Pentaclethra, Calpocalyx), and capitate or umbellate in Xylia; the primary inflorescences either axillary and solitary or paired, and immersed in the foliage, but more often grouped into terminal paniculiform secondary inflorescences exserted from the foliage; pedicels sometimes jointed where they join the calyx and remaining as peg-like structures when the flowers abscise. Flowers 5-merous, usually all hermaphrodite and appearing bisexual, functionally pistillate flowers reported in some species of Calpocalyx; hypanthium absent; sepals valvate in bud, imbricate in one genus (Pentaclethra), connate, often attenuate proximally and forming a pseudopedicel; petals valvate in bud, free or loosely joined at base, sometimes adnate to the stamens and forming a stemonozone; stamens 10, free, anthers bearing stipitate glands (absent in Amblygonocarpus); pollen usually in 8–32-grained calymmate polyads (tricolporate monads in Pentaclethra); ovary sessile or stipitate, stigma porate. Fruits variable, the valves woody and explosively dehiscent through both sutures in Pentaclethra, Xylia, Pseudoprosopis, and Calpocalyx, coriaceous to chartaceous and the valves curling post-dehiscence in Adenanthera; indehiscent in Tetrapleura and Amblygonocarpus. Seeds unwinged, the testa hard and bearing a pleurogram in most genera, recalcitrant with a papery testa and lacking a pleurogram in Pentaclethra and Calpocalyx.
Distribution. Humid forests of Africa and Asia, with only one species (Pentaclethra macroloba (Willd.) Kuntze) in the New World.
Clade-based definition. The most inclusive crown clade containing the most recent common ancestor of Pentaclethra macrophylla Benth. and Xylia torreana Brenan, but not Sympetalandra schmutzii Steenis, Entada pervillei (Vatke) R. Vig. or Pachyelsama tessmannii (Harms) Harms (Fig.
Notes. The Adenanthera clade includes all genera from the Adenanthera group of
The transitional nature of the Adenanthera clade is evidenced by the presence of characters that are very unusual among other members of the Mimoseae such as imbricate sepals and alternate leaflets. Flowers of Pentaclethra are distinct from other genera in the group having imbricate sepals, five staminodia alternating with fertile stamens within each flower, and a unique hooded anther gland unlike any other in Mimoseae. What Pentaclethra shares with most other members of the Adenanthera clade are woody, explosively dehiscent pods, another character uncommon outside this clade, and which may be the ancestral fruit state of the Adenanthera clade.
The relationship between Tetrapleura and Amblygonocarpus is morphologically strong. The fruits are quite similar, being indehiscent and four-angled in cross-section, with the angles elaborated into wings in Tetrapleura (Fig.
The Pseudoprosopis, Xylia and Calpocalyx clade shares the character of leaflets and pinnae opposite one another. Pseudoprosopis stands out as the only genus in the Adenanthera clade to have a liana habit, and some species have unique lignified tendrils for climbing (Fig.
Habit and growth form in the Adenanthera clade A Xylia xylocarpa var. kerrii (Craib & Hutch.) I.C. Nielsen scaly bark B Xylia torreana Brenan canopy C Pentaclethra macroloba (Willd.) Kuntze trunk showing buttresses D Tetrapleura tetraptera (Schumach. & Thonn.) Taub. canopy (Harris 9659) E Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre pale grey, scaly bark (Catarino 2067) F Pseudoprosopis gilletii (De Wild.) Villiers liana with leaves and lignified tendrils (Texier 1558) G Adenanthera pavonina L. thin scaly bark H habit I Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre growing in savanna (Coates Palgrave M806). Photo credits A Dechaphaetkrathok B F du Randt C D DeMelo D DJ Harris E L Catarino F N Texier G, H Shelbyfarmer I M Coates Palgrave.
Flowers and inflorescences in the Adenanthera clade A Adenanthera pavonina L. inflorescence B, C Tetrapleura tetraptera (Schumach. & Thonn.) Taub. (Harris 9663) B reflexed petals and white filaments on the flowers C racemose inflorescence D Xylia hoffmannii (Vatke) Drake umbellate inflorescence (Ratovoson 838) E Calpocalyx klainei Pierre ex Harms inflorescence (Texia 976) F Pseudoprosopis bampsiana Lisowski flower detail showing pseudopedicels (Bidault 2515) G, H Pentaclethra macroloba (Willd.) Kuntze G inflorescences congested at the ends of branches H close-up of the inflorescence showing the long filamentous staminodes. Photo credits A CheongWeei Gan B, C DJ Harris D F Ratovoson E N Texia F E Bidault G, H D Cardoso.
Fruits and seeds in the Adenanthera clade A Adenanthera pavonina L. dehisced fruits with exposed persistent red seeds B Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre Indehiscent four-angled fruits (Catarino 2067) C Pseudoprosopis fischeri Harms young fruit (Bingham 8055) D Xylia xylocarpa var. kerrii (Craib & Hutch.) I.C.Nielsen young dolabriform pod E Tetrapleura tetraptera (Schumach. & Thonn.) Taub. immature winged, indehiscent pod (Harris 9659) F Pentaclethra macrophylla Benth. pod and recalcitrant seeds G, H Calpocalyx ngouniensis Pellegr. woody valves from dehisced pod (Texier 695). Photo credits A Shelbyfarmer B L Catarino C MG Bingham D B Peroth E DJ Harris 9659 F CJ Porto, Fundación Tierra Ibérica G H N Texier.
Most Adenanthera clade species inhabit humid forests, all in Africa and Asia, except for one New World species of Pentaclethra. Some species tolerate littoral and gallery evergreen forests, but Xylia xylocarpa (Roxb.) W. Theob. is the only species to grow in drier semi-deciduous forests and Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre is the only one that has moved into the savannas of Africa (Fig.
Pentaclethra filamentosa Benth. [≡ Pentaclethra macroloba (Willd.) Kuntze (≡ Acacia macroloba Willd.)]
Large, unarmed, often buttressed trees 10–35 m tall, (Fig.
2n = 26 (
Three species, one in Latin America, widespread from Central America (Guatemala, Costa Rica, and Panama) into South America (Venezuela, Colombia, and Brazil), and the islands of Trinidad and Tobago; two species in West Africa, from Senegal to Angola and the Congo, also the islands of Principe and San José (Fig.
Lowland humid and sub-humid forests, often on riverine or waterlogged soils, sometimes the dominant tree species particularly in the Americas. Seeds are demonstrated to be hydrochorous in Pentaclethra macroloba (
Claimed to be from the Greek, pente (= five) and kleithro (= bolt), alluding to five imbricate sepals and five petals joined at their bases (
Pentaclethra macrophylla has many traditional uses, and is often planted around homes to shade gardens, improve soil fertility, and provide food, medicine, and timber. Known as the African oil bean tree, Owala oil tree, or Atta bean, the seeds are either roasted for up to 12 hours or fermented for several days to make an edible paste called ugba, high in protein and oil (
Pentaclethra is often considered a transitional genus between the core mimosoids and the rest of the Caesalpinioideae. It shares a number of characters with the genus Dimorphandra Schott such as woody, clavate pods, flowers bearing staminodia alternating with fertile stamens, and imbricate sepals; however recent studies have shown the floral characters to be independently derived (
The genus has been considered a classic example of a Gondwanan distribution, being disjunct between western Africa and Central and South America. The very short-lived (recalcitrant) seeds with a thin, papery testa coupled with a heavy, elastically dehiscent fruit seems to preclude a dispersal event across the Atlantic, although the much younger age now estimated for the legume family argues against a vicariant distribution. It is intriguing that P. macrophylla occurs on islands off the west coast of Africa and P. macroloba in Trinidad and Tobago. Seeds of P. macroloba are hydrochorous in freshwater (
Tetrapleura thonningii Benth., nom. illeg. [= Tetrapleura tetraptera (Schumach. & Thonn.) Taub. (≡ Adenanthera tetraptera Schumach. & Thonn.)]
Unarmed trees or shrubs 8–25 m tall (Fig.
2n = 26 (
Common in secondary forest but growing best in undisturbed rainforest; high forest zones, riverine forests, southern savanna woodland, and as forest outliers in the African plains.
From the Greek, tetra (= four) and pleura (= ribs), referring to the four ribs on the fruits.
A valued forest species for its fragrant fruits and seeds which are used to season food. The fruits, seeds, leaves, and bark are all used in folk medicine to treat a wide variety of ailments. The chemistry of the fruits has been studied and found to be a potential treatment for diabetes and to reduce inflammation (
Tetrapleura differs from Amblygonocarpus in minor characters as described below under Amblygonocarpus. Although they might be treated as a single genus, the two genera tend to be ecologically distinct. Tetrapleura is a rainforest tree that only occasionally moves into the savanna as a forest outlier whereas Amblygonocarpus is a true inhabitant of savannas and deciduous forests.
Amblygonocarpus schweinfurthii Harms [= Amblygonocarpus andongensis (Welw. ex Oliv.) Exell & Torre (= Tetrapleura andongensis Welw. ex Oliv.)]
Unarmed trees 8–20 m tall, 25–75 cm diameter, with a broad, umbrella-shaped crown (Fig.
2n = 28 (
One species (A. andongensis), widespread in savannas from northern Ghana east to Sudan, in the south from Angola through Zambia and Botswana to Tanzania and Mozambique (Fig.
Savannas and deciduous woodlands, frequently on sandy soils, often associated with Senegalia Raf., Burkea Benth. and Albizia Durazz.
From the Greek, ambly- (= blunt), gonia- (= angle), -carpus (= fruit), in reference to the angled fruits.
The seeds of Amblygonocarpus are harvested in the wild and eaten roasted. The roots, bark, and leaves are used in folk medicine to treat a wide variety of ailments. The wood is extremely hard and is used to make furniture, small implements, and heavy-duty flooring; also used as a fuel and to make charcoal (
Amblygonocarpus is closely related to Tetrapleura and Adenanthera in the recent phylogeny of
Gonsii Adans., Fam. Pl. 2: 318. 1763. Type not designated.
Adenanthera pavonina L.
Unarmed trees or shrubs (A. marina Nielsen), (3) 20–45 m tall, to 1 m diameter (Fig.
2n = 26, 28 (
Twelve species in tropical Asia, Australasia, Melanesia, Solomon Islands, and Madagascar (Fig.
Primary and secondary rainforests 100–700 m elevation, as a canopy tree in dipterocarp forests (one species an understory tree), peat swamps and swampy forests, forest margins, savannahs, alluvial forests, and two species coastal.
Greek Aden- (= gland) and anthera, referring to the gland present at the apex of the anther.
Planted in villages as an ornamental, used as shade trees for coffee and as a nitrogen-fixer. The seeds are toxic fresh but can be cooked and eaten, as can the leaves. The seeds are also used for making jewelry and in India to make a red dye. Leaves, bark, and seeds have been used in traditional medicine, and recent studies of seed extracts have shown A. pavonina to have antibacterial and anti-inflammatory effects (
Adenanthera pavonina is widely naturalised in Africa and parts of Asia, and in the New World in the Caribbean, Florida, and northern South America. The red or bicoloured seeds are persistent after the pods dehisce and are bird-dispersed.
Pseudoprosopis fischeri (Taub.) Harms [≡ Prosopis fischeri Taub.]
Large woody lianas or scandent shrubs or small trees, 3–6 m, coppicing in one species; unarmed but sometimes having large, hooked, lignified tendrils for climbing (Fig.
Unknown.
Lowland humid and sub-humid forests in Guineo-Congolian region, where they may occur in primary forests and also in secondary forests and riparian areas. In Tanzania and Mozambique, they are found on the edges of evergreen and deciduous gallery forests, or in thickets in cutover regions.
Pseudo (false) and Prosopis, referring to its similarity to the genus Prosopis L.
Used as a fish poison in the Democratic Republic of Congo (
Pseudoprosopis is the only genus in the Adenanthera clade having the liana habit. The uncinate lignified tendrils on the stems, although not universal in the genus, are also distinctive. Two species, Ps. bampsiana Lisowski, and Ps. euryphylla Harms are listed as vulnerable and near-threatened respectively by IUCN.
Esclerona Raf., Sylva Tellur.: 120. 1838. Type: Esclerona montana Raf. [= Xylia xylocarpa (Roxb.) Taub. (≡ Mimosa xylocarpa Roxb.)]
Xylolobus Kuntze, Lex. Gen. Phan.: 598. 1903, nom. superfl. Type not designated.
Xylia dolabriformis Benth. [= Xylia xylocarpa var. xylocarpa (≡ Mimosa xylocarpa Roxb.)]
Shrubs or trees, (6) 15–30 m tall, to 40 cm diameter, unarmed, evergreen or deciduous (Fig.
Nine species, West Africa from Guinea to Ghana; central and southern Africa from Democratic Republic of the Congo to Tanzania, Mozambique, and South Africa, Madagascar, India, and South East Asia (Fig.
n = 12 (
Evergreen, semi-deciduous and gallery forests.
From the Greek xylon, referring to the very hard wood in this genus.
Valued as a timber tree. The wood is very hard and is used in heavy construction, for houses, bridges, ship building, and tools as well as for fuel and charcoal. Used in reforestation in South East Asia. Seeds are eaten as a vegetable, and the bark and seeds are used in folk medicine to treat a wide variety of ailments. Xylia xylocarpa is grown as a shade tree in India (
Xylia has distinctive woody, dolabriform legumes, seen elsewhere only in Pentaclethra, Calypocalyx, Pseudoprosopis and Dimorphandra. The inflorescences vary from 1–2 heads borne in the axils of coeval, well-developed leaves in some species to complex paniculiform inflorescences with numerous aggregated heads and suppressed leaves in other species. In a few species, simple, linear bracts are present in the inflorescence instead of leaves and the peduncles may bear 1-several bracts below the developing inflorescences. Generally, these bracts are caducous. Inflorescences are also characterised by having precocious development of the foliar glands which reach their full size on very small leaves. The glands are conspicuous in the inflorescence and possibly serve either to distract potential floral predators from the developing flowers or to attract protectors. Likewise, the terminus of the rachis is precociously developed, overtopping the pinnae and leaflets, and appearing bract-like. As discussed in more detail in the Adenanthera clade notes, Calpocalyx is nested within Xylia, rendering the latter non-monophyletic (Fig.
Calpocalyx dinklagei (Taub.) Harms [≡ Erythrophloeum dinklagei Taub.]
Small trees or shrubs 5–20 m to tall forest trees to 50 m or more, the latter often with buttresses or water roots, unarmed, glabrous or pubescent; brachyblasts absent. Stipules small, linear, caducous and absent from most specimens. Leaves bipinnate, petiole usually terete, occasionally slightly sulcate, petiolar and rachis glands present, usually sunken into the petiole; pinnae 1 pair, 10–30 (50) cm long, opposite, articulated to the petiole; leaflets opposite, 1–9 pairs per pinna, the proximal pair of leaflets usually reduced to a single leaflet, macrophyllous, obovate to elliptic, petiolulate, the petiolules articulate to the rachis. Inflorescences of spikes, 2.5–11 cm long, oblong, subtended by triangular bracts, either solitary and axillary or more often aggregated into terminal complex-branched paniculiform secondary inflorescences with the spikes arranged in fascicles of 1–5, these subtended by three-parted bracteoles which bear an enlarged circular gland on the center bract, panicle immersed or exserted above the foliage; entire inflorescence usually fuscous to golden pubescent, anthesis often with centrifugal maturation (Fig.
n = 12 (
Littoral and coastal forests, and rainforest. Shrubs and treelets are generally in the understory of undisturbed forest but also flourish in older secondary forests. Larger buttressed trees occur in evergreen lowland forests, two species have hollow branchlets and are inhabited by ants, and cauliflory is reported in one species.
From Greek, calpo (= urn) and kylix (= drinking cup), referring to the urn-shaped calyx.
The wood is a valuable source of lumber and is used in construction for flooring, shipbuilding, furniture, and agricultural implements. The bark is used in traditional medicine to treat wounds and C. dinklagei has recently been found to contain powerful anti-inflammatory compounds (
Species boundaries in Calpocalyx are difficult and more sampling is necessary to properly delimit species. The genus has a number of unique features in the Adenanthera clade. Ant-associations, while common among the members of the Adenanthera clade, have not resulted in the formation of domatia except in C. cauliflorus Hoyle and C. winkleri (Harms) Harms. Mimoseae usually have synchronous flowering within an inflorescence, but some species of Calpocalyx demonstrate remarkable centrifugal maturation within a spike. Most species in the genus are listed as vulnerable by the IUCN, including C. atlanticus Villiers, C. brevifolius Villiers, C. cauliflorus Hoyle, C. heitzii Pellegr., C. klainei Pierre ex Harms, C. letestui Pellegr., and C. ngouniensis Pellegr. Calpocalyx is here recovered as nested within Xylia (Fig.
Anne Bruneau1
Citation: Bruneau A (2024) 14. Sympetalandra. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 225–227. https://doi.org/10.3897/phytokeys.240.101716
Sympetalandra borneensis Stapf
Sympetalandra is here considered a monogeneric lineage, whose relationship to other Mimoseae remains difficult to determine. Although historically considered a member of non-mimosoid Caesalpinioideae (
Unarmed, small to medium-sized tree (to 30 m), buttressed. Stipules minute or caducous. Leaves bipinnate (Fig.
Sympetalandra unijuga (Airy Shaw) Steenis A bipinnate leaves with opposite pinnae and opposite leaflets B fruit, Malaysia (Madani 83436) C flowers in bud and early anthesis D slender, branched inflorescences E close-up of flower. Photo credits A, C–E Digital Flora of the Philippines (
Unknown, but recent phylogenomic analyses suggest the genus may be polyploid (
Five species [S. borneensis, S. densiflora (Elmer) Steenis, S. hildebrandii Steenis, S. schmutzii Steenis, S. unijuga (Airy Shaw) Steenis] restricted to Malesia (Fig.
Lowland tropical forests. Sympetalandra densiflora, known as Kamatog, is considered near-threatened under the IUCN Red List in the Philippines.
The bark of S. schmutzii is used as a fish poison (
The generic name refers to the petals that are shortly connate at their base.
Sympetalandra was placed in the Dimorphandra group of tribe Caesalpinieae by
Sympetalandra is subtended by a relatively long branch in the phylogenomic studies of
Anne Bruneau1
Citation: Bruneau A (2024) 15. Chidlowia. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 228–230. https://doi.org/10.3897/phytokeys.240.101716
Chidlowia sanguinea Hoyle
Chidlowia is here considered a monospecific lineage, whose relationship to other Mimoseae remains difficult to determine. In all phylogenetic analyses in which the genus is included, it is resolved as a distinct lineage, generally on a long branch, but strongly supported within Mimoseae (
Unarmed tree, small to medium sized, up to 25 (30) m (Fig.
Chidlowia sanguinea Hoyle A tree with steep buttresses and twisted trunk B bark greyish to brownish with many lenticels, inner bark pink to reddish brown C parapinnate leaves with 4–6 pairs of leaflets D inflorescence, long and pendulous panicle E tip of inflorescence showing multiple buds F deep red flowers with long stamens and style G fruit, with coriaceous woody valves H dehisced fruit showing under-developed seed remnants. Photo credits A–C C Jongkind (WAG) D–H X van der Burgt (K).
Unknown.
Trees of evergreen and moist semi-deciduous Guineo-Congolian forest.
Named in honour of English silviculturist, Chidlow Vigne, who worked at the Gold Coast Forest Service, and was the first collector to recognise the distinctiveness of specimens of this species.
In Ivory Coast the wood, which is very hard and known as ‘bala’, is locally used for joinery, stakes and rifle butts (
Chidlowia was placed in the Dimorphandra group of tribe Caesalpinieae by
In Chidlowia, the singly pinnate leaves, relatively large flowers with showy red petals which are strongly imbricate in bud, the large explosively dehiscent woody fruits, and seeds lacking a pleurogram are all more suggestive of placement outside the mimosoids.
Chidlowia has flowers similar to those found in Sympetalandra with a fleshy and thick floral disc joined to the base of the calyx, petals and stamens simulating a hypanthium (
Shawn A. O’Donnell36, Gwilym P. Lewis10
Citation: O’Donnell SA, Lewis GP (2024) 16. Entada clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 231–240. https://doi.org/10.3897/phytokeys.240.101716
Entada clade
Figs
Included genera (3). Aubrevillea Pellegr. (2 species), Entada Adans. (40), Piptadeniastrum Brenan (1).
Description. Trees, shrubs, lianas and geoxylic suffrutices, unarmed (except for Entada spinescens Brenan with spinescent stipules). Stipules inconspicuous, setaceous, or in E. spinescens rigid, subconical, divaricate, spinescent, or in Piptadeniastrum africanum (Hook. f.) Brenan linear, densely pubescent, caducous. Leaves bipinnate, lacking extrafloral nectaries (except in a few species of Entada). Inflorescences spiciform racemes or spikes, axillary or terminal, sometimes clustered into fascicles or panicles. Flowers 5-merous, bisexual or staminate, sessile or pedicellate; calyx gamosepalous, campanulate or cupuliform; petals free or slightly united at their bases, stemonozone present; stamens (8) 10, anthers with a sessile or stipitate gland, or eglandular; pollen tricolporate, infratectum columellate, exine perforate to finely reticulate, released as monads; ovary multi-ovulate. Fruit papyraceous, with a basal twist, indehiscent, or coriaceous and dehiscent along one suture, or a craspedium breaking up to leave the sutures as a persistent replum. Seeds globular or laterally compressed, winged or not, with or without a pleurogram.
Distribution. Widespread across the tropics, with highest species diversity in sub-Saharan Africa (Aubrevillea and Piptadeniastrum are confined to tropical Africa), reaching subtropical latitudes in southern Africa and eastern Asia.
Clade-based definition. The most inclusive crown clade containing Entada phaseoloides (L.) Merr. and Piptadeniastrum africanum (Hook. f.) Brenan, but not Chidlowia sanguinea Hoyle, Pentaclethra macroloba (Willd.) Kuntze or Prosopis africana (Guill. & Perr.) Taub. (Fig.
Notes. As then known, species of the Entada clade had been distributed by
A sister relationship between Piptadeniastrum africanum and the Entada group, rather than with the Newtonia group as
Gigalobium P. Browne, Civ. Nat. Hist. Jamaica: 362. 1756, nom. rej. vs. Entada Adans.. Lectotype (designated by Panigrahi in Taxon 34 : 714. 1985): Entada gigas (L.) Fawc. & Rendle [≡ Mimosa gigas L.]
Perima Raf., Sylva Tellur.: 118. 1838. Type: Perima odorata Raf., nom. illeg. [= Mimosa scandens L. (= Entada phaseoloides (L.) Merr.)]
Strepsilobus Raf., Sylva Tellur.: 117. 1838. Type: Strepsilobus scandens (L.) Raf. [≡ Mimosa scandens L. (= Entada phaseoloides (L.) Merr.)]
Elephantorrhiza Benth., J. Bot. (Hooker) 4: 344. 1841. Type: Elephantorrhiza burchellii Benth., nom. illeg. [≡ Acacia elephantorrhiza Burch. ex DC., nom. illeg. (= Entada elephantina (Burch.) S.A. O’Donnell & G.P. Lewis)]
Pusaetha L. ex Kuntze, Revis. Gen. Pl. 1: 204. 1891. Type: Pusaetha scandens (L.) Kuntze [≡ Mimosa scandens L. (= Entada phaseoloides (L.) Merr.)]
Entadopsis Britton, N. Amer. Fl. 23: 191. 1928. Type: Entadopsis polystachya (L.) Britton [≡ Mimosa polystachya L. (≡ Entada polystachya (L.) DC.)]
Entada monostachya DC., nom. illeg. [≡ Mimosa entada L. (≡ Entada rheedei subsp. rheedei)]
Lianas (Fig.
Entada clade variation in habit A Entada rheedei Spreng., huge woody liana, India B Entada wahlbergii Harv., thin woody climber in fruit, Benin C Entada leptostachya Harms, scandent shrub in fruit, Madagascar D Entada burkei (Benth.) S.A. O’Donnell & G.P. Lewis, erect shrub, South Africa E Entada abyssinica Steud. ex A. Rich., small tree, Rwanda F Entada elephantina (Burch.) S.A. O’Donnell & G.P. Lewis, geoxylic suffrutex, South Africa G Aubrevillea platycarpa Pellegr., large tree with plank buttresses, Guinea H Piptadeniastrum africanum (Hook.f.) Brenan, large tree with aliform buttresses, Guinea I P. africanum, large tree in forest canopy, Sierra Leone. Photo credits A Shiwali Samant, iNaturalist (https://www.inaturalist.org/photos/58810243) B M Schmidt,
Entada clade variation in foliage and flowers A Entada rheedei Spreng., young bipinnate leaf terminating in a bifurcate tendril (modified terminal pinnae pair), India B Entada tuberosa R. Vig., bipinnate leaf terminating in a bifurcate, thickened tendril (modified terminal pinnae pair), and rachis and rachillae terminating in a yellowish glandular mucro, Madagascar C Entada mannii (Oliv.) Tisser., oblong leaflets with rounded apices and midvein positioned centrally, Republic of Congo D Entada obliqua (Burtt Davy) S.A. O’Donnell & G.P. Lewis, asymmetric leaflets with acute apices and midvein positioned closer to distal margin, South Africa E Aubrevillea platycarpa Pellegr., oblong-obovate leaflets with emarginate apices, Guinea F Piptadeniastrum africanum (Hook.f.) Brenan, finely divided bipinnate leaves, Benin G E. rheedei, subsessile flowers with greenish-yellow corollas and cream-coloured stamen filaments, India H Entada stuhlmannii (Taub.) Harms, spiciform raceme of flowers with deep red corolla and stamen filaments, Mozambique I P. africanum, spiciform raceme of flowers with yellowish-white corolla (base of petals tinged pinkish) and stamen filaments, and red anthers with a white apical anther gland, Republic of Congo J P. africanum, terminal panicles of spiciform racemes, Republic of Congo. Photo credits A Shiwalee Sawant, iNaturalist (https://www.inaturalist.org/photos/58810126) B Andry.A.R, iNaturalist (https://www.inaturalist.org/photos/30264619) C DJ Harris,
Entada clade variation in fruits and seeds A Entada rheedei Spreng., mature, gigantic, torulose, slightly curved, segmented craspedium with woody endocarp, India B Entada gigas (L.) Fawc. & Rendle, mature, laxly spirally twisted, segmented craspedium, Costa Rica C Entada spiralis Ridl., immature, tightly spirally twisted, segmented craspedium, Singapore D Entada africana Guill. & Perr., mature, segmented craspedia, distinctly umbonate over seeds, with exocarp peeling away, Togo E Entada polystachya (L.) DC., mature, segmented craspedia, slightly umbonate over seeds, with exocarp already shed, Costa Rica F Entada dolichorrhachis Brenan, mature, small, torulose, slightly curved, segmented craspedium, Zambia G Entada burkei (Benth.) S.A. O’Donnell & G.P. Lewis, immature craspedia, not segmented, South Africa H E. burkei, mature craspedia, not segmented, entire valves breaking away from replum, exocarp peeling away, South Africa I Entada goetzei (Harms) Harms, immature, elongate craspedia, not segmented, distinctly umbonate over seeds, Mozambique J Aubrevillea platycarpa Pellegr., papery, indehiscent fruits with twisted bases (holotype A Aubreville 990, MNHN-P-P00418246), Côte d’Ivoire K Piptadeniastrum africanum (Hook.f.) Brenan, coriaceous pods, dehiscent along single suture, Democratic Republic of Congo L P. africanum, flattened, oblong seeds surrounded by broad, membranous wing, with funicle attached at middle of long axis of seed, Uganda M E. gigas, large, laterally compressed, cordate seed without pleurogram, collected in beach wrack, USA N E. rheedei, large, laterally compressed, globular seeds without pleurogram, South Africa O E. africana, laterally compressed, elliptic seeds with closed pleurogram, Togo P E. burkei, globular seed without pleurogram, South Africa. Scale bars: 1 cm (M); 2 cm (N); 5 mm (O, P). Photo credits A Dinesh Valke, iNaturalist (https://www.inaturalist.org/photos/159405372) B Pedro Blanco, iNaturalist (https://www.inaturalist.org/photos/181372721) C Cerlin Ng CC BY-NC-SA 2.0 D B Eichhorn,
2n = 28 (
Forty species, widespread, primarily tropical, but reaching subtropical latitudes in southern Africa and eastern Asia (Fig.
Frequently in riparian and littoral vegetation including at the landward fringes of mangroves, though also in savanna, open woodland, thickets and open, dry to dense, humid forest, often on sandy substrates.
Likely from the indigenous name for the plant in Malabar, India (
Seeds and bark of several species are ground and used in traditional medicines, as soap, and as fish poison; leaves and detoxified seeds are eaten as famine food (
The study of
Aubrevillea kerstingii (Harms) Pellegr. [≡ Piptadenia kerstingii Harms]
Tall trees (Fig.
Unknown.
Two species (A. kerstingii and A. platycarpa Pellegr.) from tropical west and central Africa, from Liberia and Guinea east to eastern Central African Republic and south to the Democratic Republic of Congo (Fig.
Rainforest, with outliers in seasonally dry Sudanian woodland and wooded grassland.
Named for Prof. A. Aubreville (1897–1982), a noted French forester, ecologist and taxonomist (
Used in traditional medicines, for timber, and as a shade tree (
Piptadeniastrum africanum (Hook. f.) Brenan [≡ Piptadenia africana Hook. f.]
Tall trees, unarmed, with ramified, aliform buttresses often more than 3 m high (Fig.
2n = 26, diploid (
Rainforest and in riparian vegetation within more seasonally dry tropical forest.
Used for commercial timber, charcoal, fish and rodent poison, in traditional medicines and in rituals (
In his revision of the African members of Piptadenia (sensu
Colin E. Hughes3, Melissa Luckow29, Gwilym P. Lewis10
Citation: Hughes CE, Luckow M, Lewis GP (2024) 17. Newtonia grade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 241–249. https://doi.org/10.3897/phytokeys.240.101716
Newtonia grade
Figs
Included genera (4). Anonychium (Benth.) Schweinf. (1 species), Fillaeopsis Harms (1), Newtonia Baill. (11), Plathymenia Benth. (1).
The genera Anonychium, Plathymenia, Fillaeopsis and Newtonia, the first three monospecific, and the four together comprising a total of 14 species, form a grade with respect to the core mimosoid clade (Figs
All four of these genera comprise unarmed trees, the plesiomorphic condition in Mimoseae (
Prosopis sect. Anonychium Benth., J. Bot. (Hooker) 4: 347. 1841. Type: Prosopis oblonga Benth. [= Anonychium africanum (Guill. & Perr.) C.E. Hughes & G.P. Lewis]
Anonychium lanceolatum (Benth.) Schweinf. [≡ Prosopis lanceolata Benth. (= Anonychium africanum (Guill. & Perr.) C.E. Hughes & G.P. Lewis)]
Unarmed trees 4–20 m high (Fig.
Habit, habitats, inflorescences, and fruits of Newtonia and allies A small tree of Anonychium africanum (Guill. & Perr.) C.E. Hughes & G.P. Lewis in Senegal, West Africa B small twisted treelet of Plathymenia reticulata Benth. in savanna grasslands (Cerrado) in eastern Bolivia C large, buttressed tree base of Fillaeopsis discophora Harms in tropical wet forest, Ebo, Cameroon D large buttressed tree base of Newtonia buchananii (Baker f.) G.C.C. Gilbert & Boutique in semi-evergreen forest in Mozambique E spicate inflorescence of Anonychium africanum F spicate inflorescences of Plathymenia reticulata, eastern Bolivia G spicate inflorescences of Fillaeopsis discophora, Gabon H spicate inflorescences of Newtonia buchananii in Malawi I indehiscent pods of Anonychium africanum collected as livestock feed in Burkina Faso, West Africa J tardily dehiscent pods with chartaceous or weakly coriaceous valves of Plathymenia reticulata, eastern Bolivia K pod valves and papery endocarp packets surrounding seeds of Plathymenia reticulata, Bahia, Brazil L tardily dehiscent pods with chartaceous valves of Fillaeopsis discophora in tropical wet forest in Mayombe, Congo (Brazzaville) M large winged seeds of Fillaeopsis discophora in tropical wet forest in Ambam, Cameroon N fruits and winged seeds of Newtonia buchananii in cultivation in South Africa. Photo credits A S Christensen www.westafricanplants.senckenberg.de B, F, J, K, N CE Hughes C, L, M X van der Burgt D Stefaan Dondeyne E M Arbonnier https://agritrop.cirad.fr/ G E Bidault, Missouri Botanical Garden, http://legacy.tropicos.org/Image/100618759 H G Baumann www.westafricanplants.senckenberg.de I M Schmidt www.westafricanplants.senckenberg.de.
2n = 28 (
Native across the whole Sahelian savanna belt. Seeds dispersed by herbivores.
Anonychium from the Latin or Greek onych = ónyx (= nail or claw), meaning the absence of nails or claws, in reference to the lack of armature of this genus.
Trees of Anonychium are maintained and managed by farming and pastoralist communities in traditional silvo-pastoral and agroforestry systems throughout the African Sahel (Figs
The genus Anonychium was resurrected by
Pirottantha Speg., Anales Soc. Ci. Argent. 82: 226. 1916 (publ. 1917). Type: Pirottantha modesta Speg. [= Plathymenia reticulata Benth.]
Plathymenia reticulata Benth.
Unarmed deciduous trees (Fig.
2n = 26 (
Monospecific (P. reticulata), restricted to South America, primarily in central and eastern Brazil but extending to east-central Paraguay, Bolivia, and southern Suriname (Fig.
Mainly in deciduous rocky cerrado (savanna) (Fig.
From Greek, platy (= flat) and hymen (= membrane), in reference to the membranous endocarp breaking up into papery square envelopes in which the seeds are dispersed (Fig.
The timber of P. reticulata is used for furniture and fence posts. The bark has been used medicinally (
Plathymenia forms its own monogeneric (and monospecific) lineage embedded in the grade that subtends the core mimosoid clade (Fig.
Fillaeopsis discophora Harms
Large, often buttressed, trees 24–40 m, 40–80 (100) cm diameter (Fig.
Unknown.
Monospecific (F. discophora), restricted to central Africa in Cameroon, Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Gabon, Nigeria, Republic of Congo (Fig.
Large canopy-emergent trees growing in closed-canopy evergreen lowland tropical rainforests (Fig.
From Greek Fillaea- (a legume genus name, now a synonym of Erythrophleum), and -opsis (= appearance) and hence named for its resemblance to the genus Fillaea.
The wood is light and mainly used for veneer and plywood (
Fillaeopsis forms its own monogeneric (and monospecific) lineage embedded in the grade that subtends the core mimosoid clade (Fig.
The leaves of Fillaeopsis are very similar to those of Cylicodiscus, although flowers and fruits of the two genera are quite different. When sterile the two genera can be distinguished by the following characters: Fillaeopsis lacks a gland at the apex of the petiole, whereas Cylicodiscus has a small, sunken gland; leaflets of Fillaeopsis have a conspicuous marginal vein and are elliptical in shape, whereas leaflets of Cylicodiscus have an inconspicuous marginal vein and are ovate.
Newtonia insignis Baill. [= N. duparquetiana (Baill.) Keay]
Unarmed large trees, often with buttresses (Fig.
2n = 26 (
Newtonia species are (with the exception of N. scandens Villiers) large trees, often with buttresses (Fig.
Named by Baillon in honour of the English philosopher and mathematician Isaac Newton.
Various species are used for timber (
Newtonia is one of three genera of large, unarmed African trees with winged seeds that form part of the grade subtending the core mimosoid clade (Fig.
In the past, several New World species were included in Newtonia. These are now placed in the genus Marlimorimia, a genus of the Stryphnodendron clade segregated from Pseudopiptadenia Rauschert by
Colin E. Hughes3, Melissa Luckow29, Gwilym P. Lewis10
Citation: Hughes CE, Luckow M, Lewis GP (2024) 18. Cylicodiscus. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 250–253. https://doi.org/10.3897/phytokeys.240.101716
Cyrtoxiphus Harms, Nat. Pflanzenfam. Nachtr. II–IV 1: 203. 1897. Type: Cyrtoxiphus staudtii Harms [= Cylicodiscus gabunensis Harms]
Cylicodiscus gabunensis Harms
Although Cylicodiscus is clearly a member of the core mimosoid clade, its phylogenetic relationship to the Prosopis clade, from which it is separated by a very short branch, and to the Neltuma clade, are not well-resolved (Fig.
Phylogeny showing the relationships among Cylicodiscus and the genera of the Prosopis clade and the Neltuma clade, the first-branching lineages of the core mimosoid clade of tribe Mimoseae. Substantial gene tree conflict is associated with the relationships of Cylicodiscus which is separated from the Prosopis clade by a very short branch (see inset for exact branch lengths printed above branches). For description of phylogeny and support values, see Fig.
Very large trees 25–65 m, to 70+ cm trunk diameter, bole straight cylindrical, thick wandering buttresses with knee-like outgrowths (Fig.
Habit, leaves, flowers and fruits of Cylicodiscus in wet tropical forest in west Africa A, B Cylicodiscus gabunensis Harms, large canopy emergent tree C mature tree trunk with large knee-like buttresses D immature understory sapling showing sharp-pointed woody protuberances on stem E leaves showing leaflets with conspicuous drip tips F inflorescences G strap-like legumes (ca. 1 m long) showing follicular dehiscence H canopy-emergent tree crown with long strap-like pendulous fruits I 12 cm long winged seeds. Photo credits A, B, D, E, G W Hawthorne C X van der Burgt F, H, I R Ndonda Makemba.
Unknown.
Monospecific (C. gabunensis), west and central Africa in Cameroon, Côte d’Ivoire, Equatorial Guinea, Gabon, Ghana, and Nigeria, widespread but apparently uncommon (or perhaps just difficult to collect!) (Fig.
Very large canopy-emergent trees with massive crowns in well-drained evergreen lowland tropical Guineo-Congolan rainforest (Fig.
From Greek cylico (= cup-shaped) and Latin discus (= disk), referring to the cup-shaped floral disk to which the stamens are attached.
The timber of C. gabunensis is very dense and used in heavy construction, for railway sleepers and flooring (Ndonda Makemba et al. 2019).
Phylogenetically, Cylicodiscus is separated by a very short branch from the Prosopis clade (Fig.
Habit, armature, inflorescences, and fruits of genera of the Prosopis clade A Small tree of Indopiptadenia oudhensis (Brandis) Brenan in disturbed seasonal monsoon vegetation in Uttar Pradesh, northern India B, C scattered internodal sharply tipped woody protuberances on trunk and branches of young Indopiptadenia oudhensis trees in Uttar Pradesh, northern India D trees of Prosopis cineraria (L.) Druce lopped for animal fodder on the arid fringes of the Thar desert, Rajasthan, India E internodal prickles on young shoot of Prosopis farcta (Banks & Sol.) J.F. Macbride F inflorescences of Indipiptadenia oudhensis in Uttar Pradesh, northern India G inflorescences of Prosopis farcta H unripe, plano-compressed fruits of Indopiptadenia oudhensis in Uttar Pradesh, northern India I indehiscent fruits with a thickened mesocarp of Prosopis farcta. Photo credits A–C, F, H O Bajpai and L Babu Chaudhary D CE Hughes E Zeynel Cebeci https://commons.wikimedia.org/wiki/File:Prosopis_farcta_-_Syrian_mesquite_01.JPG G Eitan Ferman https://de.wikipedia.org/wiki/Prosopis_farcta#/media/Datei:Prosopis_farcta,_flower.jpg I Zeynel Cebeci https://en.wikipedia.org/wiki/Prosopis_farcta#/media/File:Prosopis_farcta_01.JPG.
Colin E. Hughes3, Melissa Luckow29, Gwilym P. Lewis10
Citation: Hughes CE, Luckow M, Lewis GP (2024) 19. Prosopis clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 254–259. https://doi.org/10.3897/phytokeys.240.101716
Prosopis clade
Figs
Included genera (2). Indopiptadenia Brenan (1 species), Prosopis L. (3).
Description. Small trees, shrubs or subshrubs (Fig.
Distribution. Restricted to the Old World in northern Africa, the Middle East, Pakistan, northern India and Nepal in dry and arid thorn scrub and seasonally dry deciduous or semi-evergreen monsoon forests.
Clade-based definition. The most inclusive crown clade containing Indopiptadenia oudhensis (Brandis) Brenan and Prosopis cineraria (L.) Druce, but not Cylicodiscus gabunensis Harms, Xerocladia viridiramis (Burch.) Taub. or Newtonia hildebrandtii (Vatke) Torre (Figs
Notes. The non-monophyly of the former Prosopis s.l. prompted its division into four segregate genera, Anonychium (Benth.) Schweinf., Prosopis, Neltuma Raf. and Strombocarpa Engelm. & A. Gray by
The sister group relationship between Prosopis and the Indo-Nepalese monospecific genus Indopiptadenia, i.e., the Prosopis clade as circumscribed here, is robustly supported (Fig.
Indopiptadenia oudhensis (Brandis) Brenan [≡ Piptadenia oudhensis Brandis]
Trees 3–12 m tall (Fig.
Unknown.
Monospecific (I. oudhensis), occupying a narrow band restricted to the Terai region of the outlying lowest foothills of the Himalayas at scattered localities almost entirely within Nepal and just barely traversing the border into northern India in a few places (Fig.
Seasonal deciduous or semi-evergreen monsoon forests and disturbed riverine vegetation (Fig.
Indopiptadenia refers to the superficial resemblance to the genus Piptadenia and Indo- (from India).
The leaves are used locally for fodder and the wood as fuel and timber. The wood is very hard, strong, and durable (
Indopiptadenia remained poorly known, based on just a handful of collections, until
Lagonychium M. Bieb., Fl. Taur.-Caucas. 3: 288. 1819. Type: Lagonychium stephanianum M. Bieb. [= Prosopis farcta (Banks & Sol.) J.F. Macbr.]
Prosopis sect. Adenopis DC., Prodr. [A.P. de Candolle] 2: 446. 1825. Type: Prosopis spicigera L. [= Prosopis cineraria (L.) Druce]
Pleuromenes Raf., Sylva Tellur.: 144. 1838. Type: Pleuromenes heterocarpa (Delile) Raf. [≡ Acacia heterocarpa Delile (= Prosopis farcta (Banks & Sol.) J.F. Macbr.)]
Prosopis spicigera L. [= Prosopis cineraria (L.) Druce]
Prickly subshrubs, shrubs, small trees (Fig.
2n = 28 (
Three species distributed across arid parts of north Africa (but apparently the genus rare at its western limits in Libya and Tunisia), the Middle East, Pakistan and north-western India (especially Punjab and Rajasthan) and reaching its northern limits in Azerbaijan (Fig.
Abundant in dry and arid parts of north-western India, where P. cineraria is sometimes the most common tree in parts of Punjab and Rajasthan (Fig.
Prosopis cineraria is a highly valued tree, protected and cultivated in silvopastoral and other agroforestry systems as a source of high-quality durable wood, leaves for fodder, fruits for livestock feed and flowers as bee forage (
Although P. cineraria and P. farcta are well-known, easily distinguished and widely distributed species, the third species of the genus, P. koelziana Burkart, is poorly-known from just a handful of collections from Iran. It has fruits similar to P. farcta and leaves similar to P. cineraria, and was suggested by
Colin E. Hughes3, Melissa Luckow29, Gwilym P. Lewis10
Citation: Hughes CE, Luckow M, Lewis GP (2024) 20. Neltuma clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 260–268. https://doi.org/10.3897/phytokeys.240.101716
Neltuma clade
Figs
Included genera (3). Xerocladia Harv. (1 species), Strombocarpa Engelm. & A. Gray (10), Neltuma Raf. (ca. 30).
Description. Intricately much-branched erect to prostrate shrubs and small trees (Fig.
Habit and armature of the genera of the Neltuma clade A shrub of Xerocladia viridiramis Taub. in the Namib desert, Namibia B small tree of Strombocarpa tamarugo (Phil.) C.E Hughes & G.P. Lewis, on the Pampa del Tamarugal in the Atacama desert, northern Chile C shrubby treelet of Strombocarpa ferox (Griseb.) C.E. Hughes & G.P. Lewis in seasonally dry tropical scrubland in southern Bolivia D small tree of Neltuma kuntzei (Harms ex C.E.O. Kuntze) C.E. Hughes & G.P. Lewis in seasonally dry tropical forest in southern Bolivia E small tree of Neltuma juliflora (Sw.) Raf. in seasonally dry tropical scrubland on deep black vertisol soil in central Nicaragua F straight cylindrical spinescent shoots of the sub-aphyllous Neltuma kuntzei G short recurved stipular spines and unijugate leaves of Xerocladia viridiramis H slender pale whitish stipular spines, unijugate leaves, and globose to ovoid-elliptic capitula of Strombocarpa strombulifera (Lam.) A. Gray I axillary nodal spine of Neltuma juliflora, central Nicaragua. Photo credits A H Kolberg, Plants of Namibia https://herbaria.plants.ox.ac.uk/bol/Namibia B O Whaley C–F, I CE Hughes G GN Dreber https://www.southernafricanplants.net/plantdata_sub.php?Mspec_ID=6570 H Guillermo Debandi, iNaturalist (https://guatemala.inaturalist.org/photos/1502506).
Inflorescences and fruits of the genera of the Neltuma clade A capitulum of Xerocladia viridiramis Taub. B spike of Strombocarpa palmeri (S. Watson) C.E. Hughes & G.P. Lewis C spike of Strombocarpa tamarugo (Phil.) C.E Hughes & G.P. Lewis D erect spike of Neltuma rubriflora (Hassl.) C.E. Hughes & G.P. Lewis E pendulous spikes of Neltuma flexuosa (DC.) C.E. Hughes & G.P. Lewis, Antofagasta, Chile F spikes of Neltuma laevigata (Humb. & Bonpl. ex Willd.) Britton & Rose, south-central Mexico G indehiscent pods of Neltuma laevigata, south-central Mexico H unripe winged monospermous fruits of Xerocladia viridiramis in the Namib desert, Namibia I ripe fruits of Strombocarpa tamarugo, Atacama desert, northern Chile J spirally coiled indehiscent ‘screw-bean’ legumes of Strombocarpa strombulifera (Lam.) A. Gray. Photo credits A, H H Kolberg, Plants of Namibia https://herbaria.plants.ox.ac.uk/bol/namibia) B, F, G CE Hughes C, I O Whaley D X Enboc- Encontro de Botânicos do Centro Oeste E P Hechenleitner J Dick Culbert.
Distribution. Amphi-Atlantic, widespread in seasonally dry tropical and arid regions of the Americas, there with a somewhat bicentric predominantly amphitropical distribution of species diversity in the Mexican-Texan and the Argentinian-Chilean-Paraguayan regions, and in ecologically similar arid regions of Namibia and South Africa.
Clade-based definition. The most inclusive crown clade containing Xerocladia viridiramis (Burch.) Taub. and Neltuma laevigata (Humb. & Bonpl. ex Willd.) Britton & Rose, but not Cylicodiscus gabunensis Harms, Neptunia oleracea Lour. or Newtonia hildebrandtii (Vatke) Torre (Figs
Notes. The Neltuma clade and the relationships among its three genera are robustly supported (
Species of the Neltuma clade are almost entirely confined to arid and semi-arid succulent-rich, fire-free scrubland vegetation spanning three highly disjunct centres of species diversity in North America, South America, plus the outlying monospecific Xerocladia in southern Africa, providing another striking example of phylogenetic conservatism across the trans-continental succulent biome (
Xerocladia zeyheri Harv. [= Xerocladia viridiramis (Burch.) Taub.]
Small, rigid, densely and intricately much-branched shrub to 1 m (Fig.
Unknown.
Arid scrubland in sandy river soils, on riverbanks, alluvium and saline flats.
From Greek, xero- (= dry) and -cladion (= branch), in reference to the dense branching habit typical of small shrubs in arid climates.
Unknown.
Xerocladia has long been thought to be closely related to the former Prosopis s.l. (
Material referred to under the name Xerocladia pampeana Speg. from Argentina, shows clear affinities to the genus Prosopidastrum Burkart, as suggested by
Spirolobium A.D. Orb., Voy. Amér. Mér. 8 (Atlas, Bot): t. 13. 1839, nom. rej. vs. Spirolobium Baill., Bull. Mens. Soc. Linn. Paris 1: 773. 1889 (Apocynaceae). Type: Spirolobium australe A.D. Orb. [= Strombocarpa strombulifera (Lam.) A. Gray]
Sopropis Britton & Rose, N. Amer. Fl. 23: 182. 1928. Type: Sopropis palmeri (S. Watson) Britton & Rose [≡ Prosopis palmeri S. Watson (≡ Strombocarpa palmeri (S. Watson) C.E. Hughes & G.P. Lewis)]
Strombocarpa strombulifera (Lam.) A. Gray [≡ Mimosa strombulifera Lam.]
Low spiny, sometimes creeping, shrubs or small trees (Fig.
2n = 28 (
Ten species. Restricted to the New World and there occupying a markedly bicentric amphitropical distribution in arid and semi-arid regions of North America [southern USA, especially in the Sonoran Desert, Baja California, and northern Mexico (Coahuila)] and South America (south-central Peru to Argentina, Bolivia, and Chile) (Fig.
Often abundant in cactus-rich, semi-desert Monte vegetation, deserts and arid mesetas, dry river-beds and washes (Fig.
Strombo- (Italian = conch) and -carpa (Greek = fruit), in reference to the resemblance of the fruits to the spiral shells of some tropical marine molluscs (Fig.
Fruits browsed by cattle and sheep and much valued in arid deserts for that purpose (
Strombocarpa is one of three genera segregated from Prosopis s.l. (
Prosopis sect. Algarobia DC., Prodr. [A.P. de Candolle] 2: 446. 1825. Type: Prosopis algarobilla Griseb. [= Neltuma affinis (Spreng.) C.E. Hughes & G.P. Lewis]
Mitostax Raf., Sylva Tellur.: 120. 1838. Type: Mitostax pallida (Humb. & Bonpl. ex Willd.) Raf. [≡ Acacia pallida Humb. & Bonpl. ex Willd. (≡ Neltuma pallida (Humb. & Bonpl. ex Willd.) C.E. Hughes & G.P. Lewis)]
Algarobia (DC.) Benth., Pl. Hartw.: 13. 1839. Type: Algarobia dulcis (Kunth) Benth. [= Neltuma laevigata (Humb. & Bonpl. ex Willd.) Britton & Rose]
Prosopis sect. Monilicarpa Ruiz Leal ex Burkart, J. Arnold Arbor. 57(3): 230. 1976. Type: Prosopis argentina Burkart [≡ Neltuma argentina (Burkart) C.E. Hughes & G.P. Lewis]
Neltuma juliflora (Sw.) Raf. [≡ Mimosa juliflora Sw.]
Spiny, erect to prostrate subshrubs, shrubs and small trees, (0.1) 4–10 (20) m tall, usually with a short trunk to 40–60 (>100) cm diameter (Fig.
2n = 26 or 28 (
Probably around 30 species. Widespread across seasonally dry tropical and arid regions of the Americas with a pseudo-amphitropical bicentric pattern of greatest species diversity in the Mexican-Texan and Argentinian-Chilean-Paraguayan regions, especially diverse and abundant in the Chaco, with an outlying disjunct occurrence of Neltuma ruscifolia (Griseb.) C.E. Hughes & G.P. Lewis of questionable nativity in the Caatinga in north-east Brazil, also extending into warm and some colder temperate areas in Texas and Nevada in the north and Patagonia in the south, where N. denudans Benth. reaches 48°S (Fig.
Dominant across large tracts of the Gran Chaco in mixed sub-xerophyllous woodland, also in Monte vegetation, open desert forests in dry quebradas (Fig.
Possibly derived from the common name Mulla Thumma in the Dravidian language Teluga in the Indian states of Andhra Pradesh and Telangana, where N. juliflora is introduced.
The wood generally hard, dense, durable and flexible, and widely used for fence posts, parquet flooring, barrels, firewood and charcoal and the dependable provision of large quantities of protein- and sugar-rich, non-toxic, highly palatable and nutritious fruits (
Neltuma is one of three genera segregated from Prosopis s.l. (
Thirteen species of ‘Prosopis’ have been described since the publication of
Colin E. Hughes3, Melissa Luckow29
Citation: Hughes CE, Luckow M (2024) 19. Dichrostachys clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 269–298. https://doi.org/10.3897/phytokeys.240.101716
Dichrostachys clade
Figs
Included genera (14). Alantsilodendron Villiers (11 species), Calliandropsis H.M. Hern. & Guinet (1), Desmanthus Willd. (23), Dichrostachys (DC.) Wight & Arn. (13–14), Gagnebina Neck. ex DC. (7), Kanaloa Lorence & K.R. Wood (1), Lemurodendron Villiers & Guinet (1), Leucaena Benth. (24), Mezcala C.E. Hughes & J.L. Contr. (1), Mimozyganthus Burkart (1), Neptunia Lour. (ca. 22), Piptadeniopsis Burkart (1), Prosopidastrum Burkart (ca. 6), Schleinitzia Warb. ex J.C. Willis (4).
Description. Predominantly small trees and shrubs 1–15 m, occasionally larger trees to 30 m and 1 m stem diameter (Lemurodendron), or functionally herbaceous subshrubs from a woody taproot (some Desmanthus and Neptunia) and, in two species of Neptunia, floating aquatic herbs; mostly unarmed, or in Mimozyganthus, Piptadeniopsis and Prosopidastrum, armed with spinescent stipules, and occasionally spinescent shoots (Dichrostachys). Stipules small and caducous, or more often persistent and foliaceous or setiform, spinose in three genera. Leaves bipinnate, a nectary almost always present on the petiole or at point of insertion of first, and sometimes multiple pinnae pairs, these variably sessile or stipitate, ellipsoid, round, and crateriform or raised; pinnae 1–40 (60) pairs, leaflets (1) 2–60 pairs per pinna. Inflorescences capitate or spicate, solitary or in fascicles in axils of coevally developing leaves or forming compound panicles exserted beyond the foliage. Flowers variably homomorphic to strongly heteromorphic, usually so to some degree, with variable proportions of sterile, functionally staminate and hermaphrodite flowers in most genera, or subsets of species within genera, sterile flowers often with showy white, pink or yellow filiform or petaloid staminodia, or staminodia absent; sepals always valvate in bud except in Mimozyganthus, where imbricate; petals valvate; spherical claviform anther glands present or absent, or sometimes the connective of the anther with a minute sub-cylindrical or hooded apiculum; pollen unit highly variable including tricolporate monads, tetrahedral tetrads and calymmate or acalymmate polyads; ovary sessile or stipitate, glabrous or pubescent; stigma porate to funnelform, broadly peltate in two genera. Fruits generally linear or linear-oblong and straight, occasionally coiled, in two genera obovate or suborbicular, usually plano-compressed, the sutures more or less raised and in one genus (Gagnebina) the fruits usually winged, generally inertly dehiscent along one, or more usually both sutures, or in four genera elastically dehiscent from the apex, the valves recurving, or indehiscent and lomentiform (Prosopidastrum and Piptadeniopsis), or in one genus (Schleinitzia) margins separating but the valves remaining attached. Seeds generally unwinged, but winged in Lemurodendron and narrowly so in Mimozyganthus, pleurogram generally present, but absent in Lemurodendron, Mimozyganthus and Piptadeniopsis.
Distribution. Pantropical including Africa, North and South America, Asia, Australia and the Pacific, with the core species and generic diversity concentrated in seasonally dry tropical vegetation in Madagascar, Mexico and the Chaco region of South America. Outside this core amphi-Atlantic dry tropical range are a number of notable outliers: in the Pacific with Kanaloa (endemic to Hawaii) and Schleinitzia (western Pacific), in wet forests of north-eastern Madagascar with the narrowly endemic monospecific Lemurodendron, pantropically in aquatic and seasonally flooded habitats with Neptunia, the only known truly aquatic member of subfamily Caesalpinioideae, and extending weakly into temperate North America (Desmanthus).
Clade-based definition. The most inclusive crown clade containing Lemurodendron capuronii Villiers & Guinet and Dichrostachys unijuga Baker, but not Prosopis spicigera L., Vachellia tortilis (Forssk.) Galasso & Banfi or Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow (Fig.
Notes. The Dichrostachys clade sensu
While a close relationship among the genera of the Dichrostachys and Leucaena subclades has long been recognised, the Mimozyganthus subclade was identified and placed with the Dichrostachys and Leucaena groups by
Habit and growth forms of genera of the Dichrostachys clade A herbaceous to weakly woody thicket of shrubby Neptunia plena (L.) Benth. in seasonally inundated ruderal habitat, Bolivia (Särkinen 2159) B, C medium-sized to large trees of Lemurodendron capuronii Villiers & P. Guinet, north-western Madagascar (Koenen 429) D small tree of Leucaena matudae (Zárate) C.E. Hughes in seasonally dry tropical forest, Guerrero, Mexico (Hughes 2153) E shrub of Kanaloa kahoolawensis Lorence & K.R. Wood cultivated, National Tropical Botanical Garden, Kaua’i, Hawaii F small treelet of Desmanthus fruticosus Rose, Baja California, Mexico (Hughes 1532) G prostrate woody suffrutex of Desmanthus acuminatus Benth., Potosi, Bolivia (Hughes 2314) H shrub of Mimozyganthus carinatus (Griseb.) Burkart, Santa Cruz, Bolivia (Hughes 2476) I shrub of Prosopidastrum globosum (Gillies ex Hook. & Arn.) Burkart, Mendoza, Argentina J shrublet of Calliandropsis nervosa (Britton & Rose) H.M. Hern. & P. Guinet, Puebla, Mexico (Hughes 1784) K small treelet of Dichrostachys cinerea (L.) Wight & Arn., Kruger National Park, South Africa L small treelet of Alantsilodendron pilosum Villiers, S. Madagascar (Luckow 4162) M treelet of Gagnebina pterocarpa (Lam.) Baill., Madagascar N shrub of Dichrostachys venosa Villiers, in dry spiny forest, southern Madagascar. Photo credits A, D–H, J CE Hughes B, C E Koenen I Estepa Patagónica K B Dupont L M Luckow M C Commenge naturetropicale N D Du Puy.
Inflorescences and flowers of genera of the Dichrostachys clade A Lemurodendron capuronii Villiers & P. Guinet (Koenen 435) B inflorescence of Neptunia plena (L.) Benth., basal flowers sterile with showy yellow petaloid staminodia (Wood 26650) C inflorescence of Leucaena esculenta (Sessé & Mociňo ex DC.) Benth. (Hughes 1326) D inflorescences and tergeminate leaves of Kanaloa kahoolawensis Lorence & K.R. Wood E fascicles of capitate inflorescences in axils of coevally developing leaves, Schleinitzia fosbergii Nevling & Niezgoda F inflorescences of Desmanthus bicornutus S. Watson showing basal sterile flowers with exserted white showy staminodia (Hughes 1526) G capitate inflorescences of Prosopidastrum globosum (Gillies ex Hook. & Arn.) Burkart H semi-spherical capitate inflorescences of Calliandropsis nervosa (Britton & Rose) H.M. Hern. & P. Guinet (Hughes 1784) I inflorescence of Alantsilodendron ramosum Villiers J spicate inflorescences of Gagnebina pterocarpa (Lam.) Baill., the basal flowers sterile with showy white staminodia K spicate inflorescence of Dichrostachys cinerea (L.) Wight & Arn., basal flowers sterile with showy pink staminodia L compact spikes of Dichrostachys akataensis Villiers the basal flowers sterile with showy pale pink staminodia. Photo credits A E Koenen B, C, F, H, J, K CE Hughes D A Palomino E Lauren Gutierrez G D Cabral, Flora Mondocina I, L D Du Puy.
Fruits of genera of the Dichrostachys clade A leaves and pod of Lemurodendron capuronii Villiers & P. Guinet, north-eastern Madagascar (Koenen 429) B Neptunia plena (L.) Benth. fruits dehiscing along the dorsal suture (Wood 26650) C ripe pods of Leucaena cruziana Britton & Rose, Oaxaca, Mexico (Hughes 1300) D cluster of unripe pods on a single inflorescence of Leucaena leucocephala subsp. glabrata (Rose) S. Zárate, Honduras (Hughes 1557) E ripe pods of Desmanthus leptophyllus Kunth dehiscing along both sutures (Hughes 3098) F unripe fruit of Mezcala balsensis (J.L. Contr.) C.E. Hughes & J.L. Contr., held erect, Guerrero, Mexico G ripe fruits of Mezcala balsensis dehiscing from the apex along both sutures H, I unripe and ripe fruits of Kanaloa kahoolawensis Lorence & K.R. Wood, cultivated at the Olinda rare plant propagation facility, Maui, Hawaii, USA J ripe fruits of Schleinitzia insularum (Guilemin) Burkart, the valves tardily splitting along both sutures but indehiscent K winged fruits of Gagnebina pterocarpa (Lam.) Baill., Madagascar L unripe, lomentiform fruits of Prosopidastrum globosum (Gillies ex Hook. & Arn.) Burkart which break up into one- or few-seeded fragments when ripe M ripe fruits of Mimozyganthus carinatus (Griseb.) Burkart, S Bolivia (Hughes 2476) N ripe fruits, elastically dehisced from the apex, Calliandropsis nervosa (Britton & Rose) H.M. Hern. & P. Guinet, Puebla, Mexico (Hughes 1784) O unripe coiled fruits of Dichrostachys cinerea (L.) Wight & Arn. P ripe fruits dehiscing elastically from the apex, Dichrostachys venosa Villiers, southern Madagascar Q ripe fruits of Alantsilodendron pilosum Villiers, Madagascar. Photo credits A E Koenen B–E, M, N C Hughes F, G J Luis Contreras H, I A Palomino J G McCormack, Cook Islands Biodiversity K, P, Q D Du Puy L I Specogno, Flora Mendocina O Atamari (CC-BY-SA-3.0).
The presence of heteromorphic inflorescences (Fig.
Fruits and seed dispersal mechanisms are diverse and apparently also evolutionarily labile, including pods inertly dehiscent along one or both sutures, sometimes tardily so and the valves remaining attached (in Schleinitzia; Fig.
These apparently complex and highly labile trajectories of fruit and flower trait evolution are likely attributable, at least in part, to reticulation given that polyploidy appears to be prevalent across the Dichrostachys clade. At least two ancient whole genome duplications are likely associated with this clade, one subtending the genus Leucaena (
Recent re-delimitation to ensure generic monophyly has led to segregation of the monospecific Mezcala to deal with the non-monophyly of Desmanthus (
None of the genera of the Dichrostachys clade are species-rich; only Leucaena and Desmanthus have more than 20 species; six genera are monospecific. The majority of genera and species are concentrated in seasonally dry tropical forests of Madagascar and the Neotropics: Alantsilodendron, Gagnebina and Dichrostachys mainly in drier parts of Madagascar, and Calliandropsis, Desmanthus, Leucaena, Mezcala, Mimozyganthus, Piptadeniopsis and Prosopidastrum mainly in the seasonally dry Neotropics. This amphi-Atlantic distribution largely confined to highly disjunct areas of seasonally dry tropical forests and scrublands has been interpreted as the result of trans-continental phylogenetic biome conservatism across the succulent biome sensu
Lemurodendron capuronii Villiers & P. Guinet
Unarmed trees to 30 m and 1 m stem diameter (Fig.
Unknown, but many gene duplications suggestive of a polyploid (Ringelberg et al. unpubl. data).
Monospecific (L. capuronii), a very narrowly restricted endemic in north-eastern Madagascar, known from just a handful of collections from the 1950s and 1960s, but recollected in 2014 from south of Vohemar, Antsiranana (Fig.
Mixed lowland evergreen and deciduous forest on igneous rock (gabbro), to 200 m elevation. Seeds likely wind-dispersed.
From Greek, lemur and -dendron (= tree), in reference to the endemic distribution of the genus in Madagascar where extant lemurs are also endemic.
The wood is locally harvested for making charcoal, potentially threatening the survival of this globally very rare genus (Erik Koenen, field observation).
Lemurodendron is sister to Neptunia in recent phylogenomic analyses (
Hemidesmas Raf., Sylva Tellur.: 119. 1838. Type: Hemidesmas plenus (L.) Raf. [≡ Neptunia plena (L.) Benth. (≡ Mimosa plena L.)]
Neptunia oleracea Lour.
Unarmed herbaceous perennials from a woody taproot (Fig.
Neptunia species comprise a polyploid series with diploids, tetraploids and hexaploids: 2n = 26, 28, 52, 56, 72, 78 (
Ca. 22 species (10 new Australian species added by
Neptunia is unique among Caesalpinioideae in having two species with an aquatic habit, occurring by dams, or in lagoons and swampy areas with little water movement. The remaining species are terrestrial but sometimes grow in seasonally inundated riverine or other ruderal habitats. It is also one of relatively few functionally herbaceous perennial Mimoseae (a habit found elsewhere in Desmanthus, Mimosa, and Entada). Although the majority of species are tropical, the genus also extends to warm temperate regions in the southern USA. In Australia, where the genus is most diverse, Neptunia species are generally found on heavy clay vertisols in grassland and open eucalypt woodland (
Named after the Roman god of fresh water and the sea, Neptune, in reference to the unusual aquatic habit of two species, including the type, in the genus.
Neptunia oleracea is a nutritious vegetable rich in vitamins and is cultivated as a food plant in Asia and Africa, where young shoots and fruits are eaten raw or cooked as a green vegetable (
Neptunia has long been considered closely allied to members of the Dichrostachys Group (
The ten new species described by
Ryncholeucaena Britton & Rose, Fl. N. Amer. 23: 130. 1928. Type: Ryncholeucaena greggii (S. Watson) Britton & Rose [≡ Leucaena greggii S. Watson]
Caudoleucaena Britton & Rose, Fl. N. Amer. 23: 130. 1928. Type: Caudoleucaena retusa (Benth.) Britton & Rose [≡ Leucaena retusa Benth.]
Leucaena diversifolia (Schlecht.) Benth. [≡ Acacia diversifolia Schlecht.]
Unarmed small or medium-sized trees, 2–15 (20) m (Fig.
2n = 52, 56, 104, 112 (
Twenty-four species plus two named hybrids (
Distribution of Leucaena based on quality-controlled digitised herbarium records. Note that the pantropically cultivated and widely introduced species Leucaena leucocephala (Lam.) de Wit, whose true native distribution remains unknown (
Seasonally dry tropical forest and semi-arid thorn scrub; two species extending to warm temperate scrublands and one species to rainforests. Seed dispersal passive. All species deciduous. Nodulating, symbiosome present. Bee-pollinated (Xylocopa and other smaller bees).
From Leucaino (Latin = becoming white), probably referring to the predominantly white stamen filaments.
Unripe seeds of 13 species are used as minor foods in Mexico and are widely cultivated and incipiently domesticated in backyards in south-central Mexico (
The Leucaena subclade (Leucaena, Schleinitzia, Mezcala, Kanaloa plus Desmanthus) is equivalent to the informal Leucaena group of
Schleinitzia novoguineensis (Warb.) L.I. Nevling & C.J. Niezgoda [≡ Piptadenia novoguineensis Warb.]
Trees or shrubs, 2–20 (–25) m, unarmed, young stems angled with corky ridges, brachyblasts absent. Stipules acicular to triangular, persistent. Leaves bipinnate; petiole ventrally sulcate, nectary at apex of petiole or mid-petiole, additional nectaries between distal-most and sometimes all pinnae, crateriform to urceolate; pinnae 4–30 pairs, opposite to subopposite; leaflets 20–60 pairs per pinna, opposite, linear to oblong; venation obscure except central midvein. Inflorescence capitate, globose, clustered 2–7 per node in terminal, largely efoliate panicles (Fig.
2n = 52 or 54 (
Four species disjunctly distributed across the western Pacific basin in New Guinea, the Philippines, Melanesia, Micronesia and Polynesia (Fig.
In lowland rainforests, especially common in secondary vegetation near the coast, and in littoral habitats above high-tide limits on calcareous substrates, especially coral limestone and coral sand, and often forming thickets on coastal beach strands (
Named in honour of Vice Admiral George Schleinitz (1834–1910), governor of German New Guinea (now part of Papua New Guinea).
The wood is favoured for cremations, handicrafts and frames for fish nets (
Schleinitzia is characterised by a fused whorl of bracts subtending the capitate inflorescence, similar to those of Kanaloa and Leucaena. Pollen in calymmate tetrahedral tetrads is also seen in Lemurodendron, Leucaena and Mezcala. Schleinitzia differs from related genera in having the flowers borne on pseudopedicels instead of being sessile.
Mezcala balsensis (J.L. Contr.) C.E. Hughes & J.L. Contr. [≡ Desmanthus balsensis J.L. Contr.]
Unarmed multi-stemmed treelet or large shrub to 3.5 m, brachyblasts present, sheathed in persistent stipules. Stipules setiform with striate membranous wings. Leaves bipinnate, a stipitate nectary between basal pinnae pair; pinnae 2–4 (5) pairs, opposite; leaflets 8–14 pairs per pinna, oblong, opposite. Inflorescences capitate, 1–2 per leaf axil, composed of 30–50 functionally sterile, functionally staminate and hermaphrodite flowers, proportions of each flower type variable; bracteoles subtending each flower peltate or deltate setiform. Flowers with sepals and petals valvate in bud; each inflorescence with 0–5 sterile flowers proximally, staminodia filamentous, white; similar to the hermaphrodite flowers but smaller and lacking anthers and ovary; functionally staminate flowers 12–30 per inflorescence, lacking an ovary, borne above the sterile flowers; hermaphrodite flowers apical, 5–25 per inflorescence, the calyx obconic, petals 5, free, white or pale green, stamens 10, cream-white, anthers with a minute orbicular gland on a filiform stalk; pollen in tetrahedral tetrads with striate exine ornamentation; ovary sessile, glabrous, stigma narrow funnelform. Fruits held erect above shoots, linear-oblong, straight or weakly arcuate, terete or sub-terete, 3–10 cm long, 5–13-seeded, seeds longitudinally inserted, valves initially fleshy becoming woody or sub-woody when ripe, elastically but tardily dehiscent from the apex along both sutures (Fig.
Unknown.
Seasonally dry tropical forest and semi-arid thorn scrub on shallow calcareous rocky soils. Seed dispersal passive. Strongly deciduous. Nodulation status unknown. Bee-pollinated (Xylocopa and other generalist bee species).
Named for the Mezcala culture which blossomed from 700 to 200 BC in the central Balsas Depression in Guerrero Mexico where the genus is endemic.
Unknown.
Mezcala was established as a monospecific genus to account for the non-monophyly of Desmanthus (
Kanaloa kahoolawensis Lorence & K.R. Wood
Unarmed shrub, 0.75–1 m, branches dense, decumbent (Fig.
2n = 28 (
Monospecific (K. kahoolawensis), narrowly restricted to the island of Kaho’olawe, Hawaii (Fig.
Steep rocky cliffs and screes derived from basaltic lava flows. Seed dispersal passive.
Kanaloa is the name of a Hawaiian deity who according to legend used the island of Kaho’olawe to rest and recoup his energies. According to
Unknown.
The affinities of Kanaloa have been in doubt due to lack of definitive support in previous phylogenetic analyses (
Acuan Medik., Theodora: 63. 1786, nom. rej. vs. Desmanthus Willd. Type: Acuan virgatum (L.) Medikus [(≡ Desmanthus virgatus (L.) Willd.)]
Darlingtonia DC., Ann. Sci. Nat. (Paris) 4: 97. 1825. Type: Darlingtonia brachyloba (Willd.) DC. [≡ Acacia brachyloba Willd. (≡ Desmanthus illinoensis (Michx.) MacMill. ex B.L. Rob. & Fernald)]
Desmanthus virgatus (L.) Willd. [≡ Mimosa virgata L.]
Unarmed, mostly woody shrubs or small trees to 4 m, or functionally herbaceous perennials much-branched from a woody base (Fig.
2n = 28 (
23 species. Three endemic species in the USA, with six extending to Mexico, six endemic species in Mexico, with three species extending to South and Central America, one endemic to Argentina, two bicentric in the USA and Argentina, one in the Antilles and north-western South America, one widespread USA to Argentina (Fig.
Seasonally dry tropical forest and semi-arid thorn scrub (ca. 10 species); warm temperate scrubland and oak woodland (seven species), three species extending to cool temperate grasslands and woodlands, two species in wet tropics (but absent from the Amazon basin), very often in disturbed open habitats, abandoned pastures and coastal thickets. Seed dispersal passive. Usually deciduous. Nodulating, symbiosome present. Bee-pollinated (Xylocopa and generalist bee species).
From Greek, desme (= bundle) and anthos (= flower), in reference to dense capitate inflorescences.
Used as ornamentals, livestock fodder and in erosion control; Desmanthus illinoensis (Michx.) MacMill. ex B.L. Rob. & Fernald is used for food (leaves, cooked seeds), medicine and is a potential pulse crop (
The delimitation of Desmanthus was altered by
Mimozyganthus carinatus (Griseb.) Burkart [≡ Mimosa carinata Griseb.]
Shrub or small treelet, 3–5 m (Fig.
2n = 28 (
Monospecific (M. carinatus), centred on the Chaco phytogeographic region in north-west and west-central Argentina, north-west Paraguay and south-east Bolivia (Fig.
Tropical and subtropical arid and semi-arid Chaco woodlands and adjacent Piedmont seasonally dry scrub and forest, 150–1200 m elevation. The papery marginally-winged fruits suggest that the fruit is likely wind-dispersed.
From Greek, zygo- (= paired) and -anthos (= flower), referring to the inflorescences which arise in the yoke between a pair of spinescent stipules, plus the Greek prefix mimo- (= mime or mimic) indicating placement of the genus in what was then subfamily Mimosoideae.
Used locally for firewood and lumber; apparently a dye can be extracted from the stems (
Until recently, Mimozyganthus was sometimes considered to be ‘transitional’ between subfamily Caesalpinioideae and the former Mimosoideae and was placed in its own tribe Mimozygantheae (
Prosopidastrum mexicanum (Dressler) Burkart [≡ Prosopis globosa var. mexicana Dressler]
Small often aphyllous shrubs, 0.5–1.5 (3) m (Fig.
2n = 14, 28 (
Six to eight species occupying a highly disjunct amphitropical distribution in Baja California and Isla Socorro, Mexico and central Argentina (Fig.
In arid subtropical scrub and semi-desert vegetation. Often aphyllous. Fruits breaking up into one- or few-seeded segments.
From -astrum (Latin = partial resemblance to) and Prosopis, another mimosoid genus which it resembles.
Unknown.
Prosopidastrum shares with Piptadeniopsis and Mimozyganthus, the two genera with which it is resolved in a robustly supported subclade of the Dichrostachys clade (Fig.
In the past, only 1–2 species were recognised until
It is difficult to reliably distinguish P. angusticarpum R.A. Palacios & Hoc from P. striatum (Benth.) R.A. Palacios & Hoc as the diameter of the inflorescence does not correlate with other characters, all of which overlap except petal colour and pubescence. The status of the segregates P. gracile R.A. Palacios & Hoc and P. benthami (Chodat & Wilczlek) R.A. Palacios & Hoc is also doubtful as they appear to differ from one another only in the colour of the petals (white vs. green-tipped). Thus, pending more detailed taxonomic revisionary work, there is uncertainty about whether six or eight species should be recognised in the genus.
Piptadeniopsis lomentifera Burkart
Shrub or small branched treelet 3–7 m, shoots striate, green for first two years, terminal shoots often slender and somewhat pendulous, armed with 1–3 mm long stipular spines at nodes. Leaves bipinnate, small, borne on short axillary tuberculous brachyblasts or alternate on older growth; petiole 3–10 mm long, puberulent, terminating in a caducous spicule; sessile, circular nectaries at point of insertion of pinnae and 1–2 minute sessile nectaries at insertion of distal leaflet pairs; pinnae 1 pair, leaflets (2) 3 pairs, elliptic-obovate, obtuse and emarginate at apex, weakly asymmetric at base, glabrous or sparsely pubescent, 3–4-veined. Inflorescence capitate, solitary, axillary on 1 cm peduncles; bracteoles subtending each flower rhomboid, acuminate, incurved, caducous. Flowers uniformly hermaphrodite; sepals valvate, fused to form a cupular calyx, villous; petals valvate, almost free at anthesis, villous; stamens 10, free, anthers with a minute stipitate spherical gland apical on the connective; pollen in columellate 8–12 (16)-grained polyads, the individual grains with porate apertures; ovary stipitate, pubescent, stigma punctate, inconspicuous. Fruits stipitate, true loments, breaking up into 3–9 one-seeded articles with no persistent replum, linear, straight, compressed, margins raised, to 8 cm long × 0.8–1 cm wide, valves sub-coriaceous, glabrous. Seeds very compressed, ovate, 7 × 7 × 1 mm, wingless, testa thin, pleurogram absent.
2n = 28 (
Tropical and subtropical seasonally dry thorn forest and Chaco woodland. Usually deciduous.
From -opsis (Greek = appearance) and Piptadenia, another genus of mimosoid legume.
Unknown.
Piptadeniopsis is robustly supported as sister to Prosopidastrum and these two genera together with Mimozyganthus form a subclade (Fig.
Calliandropsis nervosa (Britton & Rose) H.M. Hern. & P. Guinet [≡ Anneslia nervosa Britton & Rose]
Small, unarmed shrubs to 1 m, profusely and intricately branched from the base (Fig.
Unknown.
Seasonally dry thorn scrub on dry rocky calcareous soils, 1450–2000 m elevation. Seed dispersal mechanical via elastically dehiscent pods. Deciduous.
From Greek, -opsis (= appearance), referring to the resemblance to (the fruits of) the genus Calliandra.
Unknown.
Calliandropsis is robustly supported as sister to the rest of the Dichrostachys subclade (Fig.
Cailliea Guill. & Perr., Fl. Seneg. Tent.: 239. 1832, nom. rej. vs. Dichrostachys Wight & Arn. Type: Cailliea dichrostachys Guill. & Perr., nom. illeg. [≡ Mimosa nutans Pers. (= Dichrostachys cinerea subsp. africana Brenan & Brummitt)]
Dichrostachys cinerea (L.) Wight & Arnott [≡ Mimosa cinerea L.]
Shrubs or small trees to 7 m (Fig.
2n = 28 (36, 54, 56, 78) in D. cinerea (
ca. 13–14 species, D. cinerea with many varieties. One species native and widespread in Africa, India, and Australia, and widely introduced and weedy elsewhere; one species restricted to the Horn of Africa; one endemic to Socotra; one endemic to Australia; 10 species in Madagascar (Fig.
Many diverse habitats, from open savannas and seasonally dry forests to arid tropical scrub. Mostly deciduous. Dichrostachys cinerea is a problematic weed in many parts of the tropics, where it can form pure stands and outcompete native vegetation.
From Greek, di- (= two), chroma (= colour) and -stachys (= spike), in reference to the bi-coloured spicate inflorescence found in the genus (Fig.
Wood used for fuel and for small tools, leaves and fruits are important sources of livestock forage. Dichrostachys cinerea has been used in soil reclamation projects and as an ornamental (
As currently circumscribed, Dichrostachys is non-monophyletic (Fig.
Gagnebina tamarascina (Lam.) DC. [≡ Mimosa tamariscina Lam. (= Gagnebina pterocarpa (Lam.) Baill.)]
Unarmed shrubs or small trees, branched from the base, 1–6 (10) m (Fig.
2n = 26 (
Usually along rivers in evergreen or deciduous woodlands; one species [G. commersoniana (Baill.) R. Vig.] weedy and found in a wide array of habitats. The winged pods of G. pterocarpa have allowed it to colonise islands in the Indian Ocean; G. commersonia also has lightweight pods and in addition to being a widespread weed in Madagascar, it is found on Aldabra.
Named in honour of Françoise Gagnepain, French botanist (1866–1952).
Wood used locally for building houses, reportedly used in making Maintirano paper (Bosch et al. 2011).
Closely related to Dichrostachys and Alantsilodendron (Fig.
Alantsilodendron villosum (R. Vig.) Villiers [≡ Dichrostachys villosa R. Vig.]
Unarmed (weak thorns in one species) shrubs or small treelets (Fig.
Unknown.
Restricted to seasonal xerophytic deciduous woodlands and scrub, especially in the spiny forests of south-western Madagascar, deciduous in long dry season.
From alantsili-, the Madagascan name for dry forest, and -dendron (Greek = tree) in reference to the distribution of the genus restricted to dry forest.
Unknown.
Alantsilodendron is closely related to Dichrostachys and Gagnebina (Fig.
Helen C. F. Hopkins10, David S. Seigler38, John E. Ebinger14, Vanessa Terra41
Citation: Hopkins HCF, Seigler DS, Ebinger JE, Terra V (2024) 22. Parkia clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 299–315. https://doi.org/10.3897/phytokeys.240.101716
Parkia clade
Figs
Included genera (3). Anadenanthera Speg. (2–ca. 4 species), Parkia R. Br. (ca. 35), Vachellia Wight & Arn. (164).
Description. Trees or shrubs. Stipules small and caducous (Anadenanthera, Parkia) or persistent and spinose (Vachellia). Leaves bipinnate, usually alternate (some exceptions in Parkia); extrafloral nectaries present on the petiole, and commonly also distally on the rachis (Vachellia, Parkia); leaflets opposite (rarely alternate in Parkia), oblong, narrowly oblong, cultrate, elliptic, linear or rarely otherwise, main-vein typically centric and parallel to the margins in oblong and narrowly oblong leaflets, petiolule or point of attachment typically centric or subcentric (a few exceptions). Inflorescences globose (all 3 genera), or clavate, oblate, subglobose or biglobose (Parkia p.p.), or spicate or oblongoid (Vachellia p.p.). Flowers homomorphic (all 3 genera) or heteromorphic (most species of Parkia); perianth 5-merous; stamens 10 (Anadenanthera, Parkia) or many (Vachellia), anther glands present or absent; pollen in polyads; styles elongate-filiform, stigmas cup-shaped to poriform. Fruits dehiscent or not, containing pulp, or gum, or neither. Seeds in 1 or occasionally in 2 series, with a pleurogram (possible exceptions in Parkia).
Geographic distribution. Pantropical; Vachellia and Parkia are both pantropical whereas Anadenanthera is confined to the Americas.
Clade-based definition. The most inclusive crown clade containing Vachellia tortilis (Forssk.) Galasso & Banfi and Parkia bicolor A. Chev., but not Dichrostachys unijuga Baker, Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow, or Stryphnodendron adstringens (Mart.) Coville (Fig.
Notes. The Parkia clade sensu
Many characters show a patchwork distribution within the clade. These include: stipular spines (present in Vachellia, sometimes enlarged in Africa and the Americas and inhabited by ants; absent in Anadenanthera and Parkia); brachyblasts (present in Vachellia; absent in Anadenanthera and Parkia); involucre of bracts on the peduncle (present in Vachellia and Anadenanthera, absent in Parkia); calyx lobes (equal and valvate in Vachellia and Anadenanthera; unequal and imbricate in Parkia); stamen number (10 in Parkia and Anadenanthera, 20–100 in Vachellia); stemonozone (absent in Vachellia, present in Anadenanthera and often in Parkia); and pollen grains (colporate in Vachellia, porate in Parkia and Anadenanthera). Evidence of polyploidy is seen in Vachellia but not the other two genera. Root nodulation is apparently common in Vachellia and Anadenanthera and absent in Parkia (
Most, though not all, species of Parkia are bat-pollinated and many of their unusual morphological characters can be related to chiropterophily (see Parkia Notes). In contrast, the smaller inflorescences in Anadenanthera and Vachellia commonly exhibit characters typical of melittophily or generalised pollination by diurnal insects (short, slender peduncles, fragrant floral odours, diurnal anthesis).
Aldina E. Mey., Comment. Pl. Africae Austr. 171. 1836, nom. illeg., non Aldina Adans. (1763) nom. rej., nec Aldina Endl. 1840. nom. cons. Type not designated.
Farnesia Gasp., Deser. Nuov. Gen. 1838, non Farnesia Heist. ex Fabr., Enum. Pl. Hort. Helmstad. ed. 2, 400. 1763. Type: Farnesia odora Gasp., nom. illeg. [= Mimosa farnesiana L. (≡ Vachellia farnesiana (L.) Wight & Arn.)]
Gumifera Raf., Sylva Tellur.: 118. 1838. Lectotype: Gumifera nilotica (L.) Raf. [≡ Mimosa nilotica L. (≡ Vachellia nilotica (L.) P.J.H. Hurter & Mabb.)]
Poponax Raf., Sylva Tellur.: 118. 1838. Type: Poponax tortuosa (L.) Raf. [≡ Mimosa tortuosa L. (≡ Vachellia tortuosa (L.) Seigler & Ebinger)]
Delaportea Thorel ex Gagnep., Notul. Syst. (Paris) 2: 118. 1911. Type: Delaportea armata Thorel ex Gagnep. [≡ Pithecellobium harmandianum Pierre (≡ Vachellia harmandiana (Pierre) Maslin, Seigler & Ebinger)]
Pithecodendron Speg., Physis (Buenos Aires) 6: 313. 1923. Type: Pithecodendron argentinense Speg. [= Mimosa horrida L. (≡ Vachellia horrida (L.) Kyal. & Boatwr.)]
Nimiria Prain ex Craib, Bull. Misc. Inform. Kew 1927: 393. 1927. Type: Nimiria siamensis Craib [≡ Vachellia siamensis (Craib) Maslin, Seigler & Ebinger]
Acaciopsis Britton & Rose, N. Amer. Fl. 23: 93. 1928. Type: Acaciopsis pringlei (Rose) Britton & Rose [≡ Acacia pringlei Rose (≡ Vachellia pringlei (Rose) Seigler & Ebinger)]
Bahamia Britton & Rose, N. Amer. Fl. 23: 86. 1928. Type: Bahamia acuifera (Benth.) Britton & Rose [≡ Acacia acuifera Benth. (≡ Vachellia acuifera (Benth.) Seigler & Ebinger)]
Feracacia Britton & Rose, N. Amer. Fl. 23: 86. 1928. Type: Feracacia daemon (Ekman & Urb.) Britton & Léon [≡ Acacia daemon Ekman & Urb. (≡ Vachellia daemon (Ekman & Urb.) Seigler & Ebinger)]
Fishlockia Britton & Rose, N. Amer. Fl. 23: 91. 1928. Type: Fishlockia anegadensis (Britton) Britton & Rose [≡ Acacia anegadensis Britton (≡ Vachellia anegadensis (Britton) Seigler & Ebinger)]
Lucaya Britton & Rose, N. Amer. Fl. 23: 87. 1928. Type: Lucaya choriophylla (Benth.) Britton & Rose [≡ Acacia choriophylla Benth. (≡ Vachellia choriophylla (Benth.) Seigler & Ebinger)]
Myrmecodendron Britton & Rose, N. Amer. Fl. 23: 91. 1928. Type: Myrmecodendron hindsii (Rose) Britton & Rose [≡ Acacia hindsii Benth. (≡ Vachellia hindsii (Benth.) Seigler & Ebinger)]
Tauroceras Britton & Rose, N. Amer. Fl. 23: 85. 1928. Type: Tauroceras spadicigerum (Schltdl. & Cham.) Britton & Rose [≡ Acacia spadicigera Schltdl. & Cham. (= Vachellia cornigera (L.) Seigler & Ebinger)]
Acacia subg. Acacia sensu Vassal, Bull. Soc. Hist. Nat. Toulouse 108: 139. 1972.
(designated by
Shrubs or trees, 0.5–30 m, rarely a prostrate shrub; bark mostly dark to light brown to black or grey, rarely whitish-reddish to yellowish, and papery, peeling or corky, with most species commonly rough to furrowed; brachyblasts usually present; prickles absent. Stipules spinescent, woody, paired at the nodes, straight to curved, sometimes asymmetrical, in some species enlarged and inhabited by ants, occasionally exceeding 200 mm long; a number of African species with galls subtending the spines. Leaves caducous (sometimes evergreen), alternate and also commonly clustered (2–8) on short shoots, these brachyblast leaves usually smaller and with fewer pinna pairs and leaflets than the alternately arranged leaves on fast growing branches; petioles with one or more extrafloral nectaries, often with one to several on the rachis, most commonly between the pinna pairs; pinnae 1–many (60+) pairs, mostly opposite; leaflets 1-many (70+) pairs per pinna, mostly opposite, sessile to subsessile, the apex in most American myrmecophytes bearing a small yellow detachable Beltian body. Inflorescences capitula to cylindrical spikes, solitary or clustered in leaf axils or on short shoots, rarely in pseudo-racemes or pseudo-panicles, sometimes andromonoecious; peduncles with a small involucel, usually medial or basal, rarely at the base of the inflorescence. Flowers bracteate, sessile to subsessile, actinomorphic; calyx 4–5 (6)-lobed; corolla 4–5 (6)-lobed; stamens 20–100, yellow to gold or creamy white, filaments usually separate to the base or occasionally shortly fused (or rarely and irregularly a greater degree of fusion), exserted, anthers small, dorsifixed, apical glands often present; pollen in polyads of 16 (8, 12, 24, 32, 48) grains, colporate with the exine surface psilate and nexine with columellae (
Morphological features of Vachellia A branches with spicate inflorescences of V. rigidula (Benth.) Seigler & Ebinger B habit of V. bidwillii (Benth.) Kodela (M. Simmons 3191) C inflorescences and leaves of V. farnesiana (L.) Wight & Arn. D habit of V. farnesiana E gall and spines of V. drepanolobium (Harms ex Y. Sjöstedt) P.J.H. Hurter F spines and inflorescences of V. cornigera (L.) Seigler & Ebinger (Seigler 16051) G partial leaf of V. cornigera with Beltian Bodies (Tamaulipas, Mexico) (Seigler 15962) H fruit of V. caven (Molina) Seigler & Ebinger I habit of V. schaffneri (S. Watson) Seigler & Ebinger J leaves and stems of V. karroo (Hayne) Banfi & Galasso K inflorescences and leaves of V. nilotica subsp. indica (Benth.) Kyal. & Boatwr. (Simmons 1032) L fruits of V. nilotica subsp. kraussiana (Benth.) Kyal. & Boatwr. Photo credits A D Seigler B, K J Simmons C, J S Navie D, F, G, I B Maslin E T Nicholls, Nature Education Center H RT Queiroz http://rubens-plantasdobrasil.blogspot.com/ L D Kunjithapatham.
x = 13. Many American species of Vachellia are diploids with 2n = 26 (
Vachellia (164 species in total) is represented by 61 species (some with forms and varieties) in the Americas, 75 in Africa and Madagascar, nine in Australia and the Pacific, and 33 in Asia (including about 15 also found in Africa) (
Distribution of Vachellia based on quality-controlled digitised herbarium records. The map includes records for introduced/naturalised taxa as well as native ones. For instance, most records in eastern Brazil are due to V. farnesiana, which is naturalised there. The Indian subcontinent may be under-sampled (see
Vachellia is widespread in most tropical and many subtropical areas of the world. A few species enter warm temperate regions but their distribution is generally limited to areas that lack killing frosts. Vachellia nilotica and V. farnesiana have been widely introduced into the Old and New World, respectively. The latter has a pantropical distribution and, including probable introductions, is the most widely distributed of all members of the genus.
Many members of Vachellia are tenacious, spiny and invasive. Although most species favour disturbed, arid sites from sea level to ca. 2800 m, others (e.g., V. mayana (Lundell) Seigler & Ebinger) are limited to relatively undisturbed rainforests. Species are known from many forest types including primary and secondary tropical evergreen forests, rainforests, semi-deciduous forests, dry and wet secondary forests, montane, gallery, riverine forests, and are frequently found in Quercus-Juniperus woodlands, but uncommonly in secondary vegetation of cloud forests and ground water forests in Africa (
They are associated with many soil types including sands, clay, serpentine soils, volcanic soils, limestone and rocky limestone soils, karst limestone, coastal dunes, granitic washes, loam, saline soils, heavy black soils, gravelly soils, Kalahari sands, seasonally flooded alluvium, black cotton soils, hard-pan grey soils, dry water courses, margins of seasonal swamps, and they sometimes grow on termite mounds (
Vachellia was named in honour of the Reverend George Harvey Vachell (1799–1839), a collector of the flora of China.
Vachellia species are often used for firewood and for making charcoal and are the major source of fuel in many areas of the world. They often serve as forage for livestock especially in times of drought (
A number of especially spiny species are grown for living fences. In Africa these are used to surround kraals. Others, such as Vachellia leucophloea and V. nilotica, are planted along roadsides and in gardens. Some are grown as shade trees for coffee, cacao and other crops (
Vachellia farnesiana and V. caven (Molina) Seigler & Ebinger were early introduced into Spain, France and Italy (
The genus Vachellia was once part of a widely circumscribed Acacia Mill., and as first described by
No intrageneric classification has yet been published for Vachellia. Major works that present descriptions and/or keys include:
Vachellia species can readily be distinguished from other members of Acacia s.l., and especially the large segregate genus Senegalia that is also common in both Old and New World tropics, by the presence of stipular spines, the absence of prickles, the presence of an involucre on the peduncle, the absence of a torus-shaped nectary at the base of the ovary or ovary stalk, pollen polyads with colporate apertures and nexine possessing columellae, and different seedling development (
Most Vachellia species have sessile petiolar glands, usually near the lowermost pair of pinnae, and smaller rachis glands are commonly found between the upper one or more pairs of pinnae. These glands, at least during the rapid growth associated with flower development, appear to produce a food reward for pollinators and various insects, especially ants.
The flower-heads of Vachellia species are visited by large numbers of different insects, especially bees, occasionally birds, and rarely by bats; however, pollination biology has only been examined in detail for a few species (
Seeds of the swollen-thorn American Vachellia species are dispersed by birds (
Twelve Neotropical myrmecophytes share many adaptive ecological and morphological traits, most of which appear to be related to their mutualistic association with acacia-ants of the genus Pseudomyrmex Lund, 1831 (
In approximately 11 species of African Vachellia, the stipular spines are distinctly and characteristically swollen into structures commonly known as ant-galls. These species are sometimes called Whistle Thorn acacias because of the noise caused by wind blowing over the ant entry holes. These galls are often inhabited by ants, commonly those of the genus Crematogaster Lund, 1831.
A number of hybrids have been identified in both Africa (
Piptadenia sect. Niopa Benth., J. Bot. (Hooker) 4: 340. 1841. Type: Piptadenia peregrina (L.) Benth. [≡ Mimosa peregrina L. (≡ Anadenanthera peregrina (L.) Speg.)]
Niopa (Benth.) Britton & Rose, Addisonia 12: 37, t. 403. 1927. Type: Niopa peregrina (L.) Britton & Rose [≡ Mimosa peregrina L. (≡ Anadenanthera peregrina (L.) Speg.)]
(designated by
Unarmed trees or shrubs, 3–30 m high; trunk ± smooth or with mammillate projections; bark often suberose, sometimes thick. Stipules small, bristly, caducous; bracts enclosing new shoots broad, commonly persistent. Leaves feathery; extrafloral nectaries on petiole above base, sometimes on rachis between ultimate pairs of pinnae, small, round; pinnae 7–35 pairs, opposite or almost so; leaflets 20–80 pairs per pinna, opposite or almost so, ± narrowly oblong, lanceolate, cultrate to slightly falcate, sometimes imbricate, main-vein central and straight. Compound inflorescences of pedunculate capitula, in fascicles of up to 7 peduncles inserted in series in successive leaf axils or forming terminal paniculate groups by suppression of leaves; peduncles each with 2 membranous bracts united to form an annular involucre; capitula spherical, 1–2 cm diameter including stamens, each with 35–60 flowers, greenish white to creamy yellow or rarely orange, fragrant. Flowers hermaphroditic or sometimes staminate and hermaphroditic [at least in A. colubrina (Vell.) Brenan], small, sessile; calyx campanulate, loosely gamosepalous, to 3 mm long; corolla tubular-campanulate, to 4 mm long, lobes loosely cohering to the level of the calyx mouth; stamens 10, far-exserted, free distally, adnate to corolla proximally, anther gland present (at least in bud, A. colubrina) or absent (A. peregrina); pollen in polyads of 8, 12 or commonly 16 grains, porate, exine granular, columellae absent (
Morphological features of Anadenanthera (A–I) and Parkia sect. Platyparkia (J–L) A A. colubrina (Vell.) Brenan capitulum cut in half, at anthesis B capitula at anthesis and in bud C shrub D Anadenanthera sp. foliage and petiolar nectary E A. colubrina mature pods F mammillate projections on trunk G dehisced pods H A. peregrina var. falcata (Benth.) Altschul mammillate trunks I A. peregrina var. peregrina trees with smooth trunks J Parkia pendula (Willd.) Benth. ex Walp. (Hopkins & Hopkins 273), Neotropics, tree crown K capitulum at anthesis, nectar exuding from apical flowers L pods with enlarged adaxial suture secreting gum into which the seeds have fallen. Photo credits A–I RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ J–L MJG Hopkins and HCF Hopkins.
2n = 26 (24) (
2–4 species, possibly more (
Tropical and subtropical seasonally dry forests, along rivers and at forest margins, in woodland, thickets and wooded grassland (cerrado, savanna) and caatinga, often planted near villages, sometimes weedy and found along roadsides and on wasteland; growing on a range of substrates, sometimes dominant; to 2000 (–2700) m. Trees are partly deciduous.
an- (Gr., lacking), aden- (Gr., gland), anthera (L., anther), indicating a lack of anther glands, although A. colubrina has them in bud.
In pre-Columbian times, ground seeds of both species were used by Amerindians as a source of hallucinogenic snuff (A. peregrina – cohoba, niopo, yopo; A. colubrina var. cebil – cebil, curupay, vilca), probably for 3000+ years. The northern species, A. peregrina, is still used for magical, medicinal, religious and stimulative purposes (
The long-established treatment by
In addition to its capitula and dehiscent pods, distinctive characters of Anadenanthera include the thin, rimmed or very narrowly winged seeds that are circular in outline and lack endosperm. The species recognised by
The use of Anadenanthera as an hallucinogen is dependent on the presence in the seeds, fruits and bark of many populations of a range of indole alkaloids derived from tryptamine and related to serotonin; among these, bufotenin is often especially abundant and psychoactive (
Plants of Anadenanthera colubrina are partially self-incompatible and its main pollinators are diurnal eusocial bees; eusocial honey wasps are nectar thieves or sometimes pollinators, and other insects are largely thieves of nectar and/or pollen (Kiill and da Silva 2016;
Trees are possibly mast fruiting and seed dispersal is by gravity and probably wind and leaf-cutter ants (
Paryphosphaera H. Karst., Fl. Columb. 2: 7, tab. 104. 1862. Type: Paryphosphera arborea H. Karst. [= Parkia nitida Miq.]
Parkia africana R. Br., nom. superfl. [≡ Parkia biglobosa (Jacq.) R. Br. ex G. Don (≡ Mimosa biglobosa Jacq.)]
Unarmed trees or rarely shrubs, 3–40 m, evergreen or rarely deciduous; trunk sometimes buttressed, bark variable. Stipules small, caducous. Leaves alternate, (sub)opposite or clustered at the ends of twigs; pinnae 1–55 pairs, opposite, subopposite or rarely alternate; leaflets 3–110 pairs per pinna, opposite or rarely alternate (P. biglobosa), linear to oblong or slightly sigmoid or rarely elliptic and 3–45 × 1–13 mm, or rarely ovate (P. singularis Miq.) and then to 120 × 75 mm; main-vein central, straight or slightly sigmoid; extrafloral nectaries often present on petiole near the base, elliptic, single or double (or heart-shaped), and sometimes on the rachis between the pinnae, especially in seedlings, small, round. Compound inflorescences of pedunculate capitula arranged in axillary or terminal, short to very long racemes or panicles; principal axis 0.15–5 m long, erect, horizontal, pendent or projecting at all angles, within or beneath the crown to far-extending beyond it; peduncles alternate or (sub)opposite, 1–115 cm long, pendent, erect or projecting at all angles, tough, sometimes thick and robust; 4 caducous bracts enclosing the capitulum in young bud stage. Capitula of 3 types: in sect. Sphaeroparkia: globose, 1–5 cm diameter, with 120–650 flowers, all fertile, lacking specialised nectar-secreting flowers, red or yellow at anthesis; in sect. Platyparkia: oblate, 2.7–3.5 × 4–5 cm, with 1060–1325 flowers, these of 2 sorts, those in the middle and at the base fertile, those at the apex modified and nectar-secreting, capitula red; in sect. Parkia: clavate, subglobose or biglobose, 4–21.5 × 3–8 cm, with 1090–3240 flowers, these of 3 main sorts: fertile ones forming an apical ball, below this a constricted cylinder or depressed ring of nectar-secreting flowers, at the base a zone of staminodial flowers in which the filaments are short to far-projecting and then forming a wide fringe, capitula yellow (sometimes the fringe white), reddish (bright to dull red, pink, orange or purplish), or occasionally bicoloured (red at the base, apical ball yellow). Flowers tubular, each subtended by an obdeltate-spathulate bract, slightly longer than the calyx. Fertile flowers hermaphroditic, functionally staminate, or a mixture; calyx almost bilabiate with 2 large lobes and 3 smaller ones, lobes imbricate in bud (or sub-equal and sub-imbricate in P. ulei (Harms) Kuhlm.); corolla lobes with lower parts variously connate and often adnate to the filament-tube; stamens 10, shortly exserted, filaments usually connate proximally and free distally, anthers basifixed (most species) or dorsifixed (sect. Sphaeroparkia), with or without an apical gland; pollen in polyads of 16, 28 or 32 grains, porate, exine granular or with columellae, variously ornamented (
Morphological features of Parkia sect. Parkia (A–L) and sect. Sphaeroparkia (M, N) A P. bicolor A. Chev. pendent capitulum approaching anthesis, Korup National Park, Cameroon B P. biglobosa (Jacq.) R. Br. ex G. Don pendent capitulum cut in half, Ibadan, Nigeria C P. decussata Ducke erect capitulum at anthesis, Neotropics (Hopkins & Hopkins 237) D P. timoriana (DC.) Merr. pendent capitulum, South East Asia (H.C.F. Hopkins 634) E P. gigantocarpa Ducke pendent capitulum and another cut in half, shortly post-anthesis, finger ring gives scale, Neotropics (Hopkins & Hopkins 298) F P. igneiflora Ducke pendent capitulum near anthesis, Neotropics (Hopkins & Hopkins 230) G P. speciosa Hassk., capitula at anthesis, Temburong, Brunei H P. discolor Spruce ex Benth. indehiscent pods nearing maturity, Neotropics (Hopkins & Hopkins 264) I, J P. bicolor I ripe indehiscent pod with yellow valves containing orange pulp, Bero Mts, Guinea-Conakry J immature pods, Korup National Park, Cameroon K P. cachimboensis H.C. Hopkins dehiscent pods lacking gum, the seeds attached by their funicles, Serra do Cachimbo, Brazil L P. igneiflora var. aurea Ducke vel aff. erect compound inflorescence axes projecting above the tree crown bearing pendent yellow capitula on short pendent peduncles, Cachoeira Berro d’Agua, AM, Brazil M, N P. multijuga Benth. M capitula at anthesis and in bud, Trombetas, Brazil N old pod from ground plus seeds, INPA, Manaus, Brazil. Photo credits A R Grünmeier B HCF Hopkins C–F, H, K, N MJG Hopkins and HCF Hopkins G I Nielsen I M Cheek J X van der Burgt L L Mello M unknown.
2n = 26 (22, 24) (
Currently ca. 35 species but more are likely to be recognised as a result of genetic studies (e.g.,
The genus is pantropical (Fig.
Tropical, predominantly occurring in moist habitats; most species are found in lowland rainforest (occasionally to 1500 m elevation), others grow in riparian forest, fresh-water flooded forest (várzea and igapó), woodland and wooded grassland (cerrado, savanna), and campinarana. Less common habitats in South America include coastal restinga (P. bahiae H.C. Hopkins), rocky savanna (cerrado rupestre; P. cachimboensis H.C. Hopkins) and sub-Andean dwarf forest (P. nana D.A. Neill), and in South East Asia and Malesia, peat swamp forest (P. paya H.C. Hopkins), tidal streams and Nypa swamp (P. sherfeseei Merr.), and dry evergreen and/or deciduous forest (P. leiophylla Kurz, P. sumatrana Miq.).
Named for the Scottish explorer Mungo Park (1771–1806), who investigated the course of the Niger River in West Africa and mentioned what became Parkia biglobosa as the nitta tree in the account of his first expedition to the region (
In West Africa, the seeds of Parkia biglobosa (African locust bean, néré, nété) are fermented into a widely used pungent condiment (dawadawa, soumbala, iru); the sweet mealy pulp around the seeds is also consumed (
Following
The generic limits of Parkia are unchanged from those of
Parkia is one of the most variable genera in the Mimoseae. However, despite variation in the structure of the capitula, in the morphology of the flowers, fruits and seeds, in the type of germination (phaneroepigeal, phanerogeal or cryptohypogeal) and in pollen sculpturing (
This genus has a number of unusual characters compared with others in this and closely related clades. Some appear idiosyncratic, such as the opposite leaves in a few Neotropical and one Asian species, and the lack of root nodulation. However, many of its distinctive features can be related to reproductive biology, including the sometimes very elongated compound inflorescence axes, tough and sometimes long, pendent or erect peduncles, capitula commonly composed of very numerous, relatively large flowers, foetid floral odours, and crepuscular/nocturnal anthesis (diurnal only in P. ulei). Sections Parkia and Platyparkia are pollinated by bats that typically land on the capitula to lap nectar (rather than by hovering to feed), belonging to the Phyllostomidae in the Americas and the Pteropodidae in Africa, Asia and the Pacific; various non-volant mammals, insects including bees, and birds, are occasional pollen vectors and nectar and/or pollen thieves (e.g.,
The wide range in fruit characters is reflected in a variety of dispersal mechanisms. Seed-dispersers include chimpanzees, various Neotropical and Paleotropical monkeys and perhaps birds, and for fruits that fall to the ground readily at maturity, large rodents (Parkia multijuga, Fig.
Pétala Gomes Ribeiro2, Leonardo M. Borges5
Citation: Ribeiro PG, Borges LM (2024) 23. Lachesiodendron. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 316–318. https://doi.org/10.3897/phytokeys.240.101716
Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow [≡ Acacia viridiflora Kunth]
Lachesiodendron has an isolated position in the Mimoseae phylogeny, between the Parkia clade and the node including the Mimosa, Stryphnodendron and Ingoid clades (
Trees (2) 3–20 m; indumentum puberulent to rarely glabrous; brachyblasts absent; branches armed with stipules modified into lignified spines, down-curved, paired at branch nodes, prickles absent, lenticels present. Stipules spinescent. Leaves bipinnate, unarmed; extrafloral nectaries on the petiole, on the leaf rachis between distal pairs of pinnae, and on the pinnae between distal pairs of leaflets; pinnae 5–15 pairs, opposite or sub-opposite; leaflets 20–50 pairs, opposite. Inflorescences 1–2 (3) axillary spikes. Flowers 5-merous, yellowish green; calyx gamosepalous, campanulate; corolla gamopetalous, cylindrical; stamens 10, anthers with a short-stipitate caducous apical gland; pollen in 8-grained polyads; ovary glabrous, long stipitate, exserted from the corolla, stigma in a terminal pore. Fruit a flat-compressed legume with thick margins, straight, not constricted between the 8–10 seeds, valves thin, coriaceous. Seeds ovate to obovate with a U-shaped pleurogram on both faces (Fig.
Unknown.
Confined to seasonally dry tropical forests and woodlands.
From ‘Lachesis’, in reference to the bushmaster viper [Lachesis muta (Linnaeus, 1766)] whose vernacular name (surucucu) is also applied to the tree in Brazil, probably because the pair of nodal spines resembles the viper’s fangs (Fig.
Lachesiodendron viridiflorum is used as fodder, for the timber, for environmental restoration, and provides good quality firewood and charcoal (
Lachesiodendron was recently described by
Leonardo M. Borges5, Marcelo F. Simon40, Pétala Gomes Ribeiro2, Melissa Luckow29, Alexandre Gibau de Lima8,28
Citation: Borges LM, Simon MF, Ribeiro PG, Luckow M, Lima AG (2024) 24. Stryphnodendron clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 319–331. https://doi.org/10.3897/phytokeys.240.101716
Stryphnodendron clade
Figs
Included genera (7). Gwilymia A.G. Lima, Paula-Souza & Scalon (7 species), Marlimorimia L.P. Queiroz, L.M. Borges, Marc.F. Simon & P.G. Ribeiro (6), Microlobius C. Presl (1), Naiadendron A.G. Lima, Paula-Souza & Scalon (1), Parapiptadenia Brenan (6), Pityrocarpa (Benth.) Britton & Rose (7), Stryphnodendron Mart. (28).
Description. Trees, rarely shrubs or subshrubs; indumentum composed of simple trichomes and sometimes also reddish granular trichomes; brachyblasts absent or present; branches and leaves unarmed, with a garlic smell in one genus. Stipules absent or present, usually caducous. Leaves bipinnate, extrafloral nectaries present on the petiole, rachis and pinnae; pinnae 2–many pairs, opposite or subopposite, variable in size and shape; leaflets 1–many pairs, opposite or alternate, variable in size and shape. Inflorescences cylindrical spikes, solitary or in groups, in the axils or supra-axillary to coevally developing leaves, or in efoliate nodes, sometimes further arranged in complex synflorescences. Flowers 5-merous; calyx gamosepalous; corolla gamopetalous; stamens 10, anthers with an apical gland; pollen usually in 16-grained polyads, but sometimes in groups of 4–32 grains. Fruit a dehiscent or indehiscent legume or a follicle, variable in shape and size. Seeds ellipsoid or flat-compressed, winged or not, brown, dark or white, pleurogram absent or present.
Geographic distribution. Tropical America, from Mexico to Argentina and Uruguay.
Clade-based definition. The most inclusive crown clade including Stryphnodendron adstringens (Mart.) Coville and Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson, but not Vachellia tortilis (Forssk.) Galasso & Banfi, Piptadenia adiantoides (Spreng.) J.F. Macbr. or Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow (Fig.
Generic relationships in the Stryphnodendron clade (tribe Mimoseae). The most likely position of unsampled genus Microlobius is indicated with a dashed line [following
Notes. Members of the Stryphnodendron clade are mostly trees, always unarmed, with spicate inflorescences, and pentamerous, diplostemonous flowers with glands at the apex of the anthers (Fig.
Flowering branches in the Stryphnodendron clade A Gwilymia coriacea (Benth.) A.G. Lima, Paula-Souza & Scalon (Simon 3482) B Microlobius foetidus (Jacq.) M. Sousa & G. Andrade C Stryphnodendron adstringens (Mart.) Coville D S. gracile Rizzini & Heringer E Marlimorimia bahiana (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borges F Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson G Parapiptadenia blanchetii (Benth.) Vaz & M.P. Lima H Naiadendron duckeanum (Occhioni) A.G. Lima, Paula-Souza & Scalon (Simon 1457). Photo credits A, H MF Simon B T Iwane C, D H Moreira E LP Queiroz F D Cardoso G PG Ribeiro.
Habit and fruits in the Stryphnodendron clade A Marlimorimia psilostachya (DC.) L.P. Queiroz & Marc.F. Simon showing the tree habit B Stryphnodendron platyspicum Rizzini & Heringer (Simon 2017) showing the geoxyle subshrub habit C Gwilymia coriacea (Benth.) A.G. Lima, Paula-Souza & Scalon (Simon 3730) D Microlobius foetidus (Jacq.) M. Sousa & G. Andrade E Stryphnodendron rotundifolium Mart. fruit, manually opened revealing internal septa and seeds F Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro G Naiadendron duckeanum (Occhioni) A.G. Lima, Paula-Souza & Scalon (Pereira-Silva 15692) H Parapiptadenia rigida (Benth.) Brenan showing a yellow inflorescence and dehiscing fruit I Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro. Photo credits A R Aguilar B, C, E MF Simon D H Hulsberg F G Carvalho-Sobrinho G G Pereira-Silva H RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ I LM Borges.
Similarly to the Mimosa clade (page 332), genera of the Stryphnodendron clade have diplostemonous flowers, pollen in tetrads or polyads and a narrowing style with a small porate stigma at the tip and, thus, were included in the informal Piptadenia group of Mimoseae (
Gwilymia paniculata (Poepp. & Endl.) A.G. Lima, Paula-Souza & Scalon [≡ Stryphnodendron paniculatum Poepp. & Endl.]
Trees; indumentum composed of simple and granular trichomes; brachyblasts absent; branches unarmed, young shoots and leaves covered with reddish granular trichomes. Stipules caducous. Leaves bipinnate; extrafloral nectaries on the petiole, between pinnae and between apical leaflets; pinnae 2–4 (6) pairs, opposite or subopposite; leaflets 3–5 pairs, opposite, variable in shape. Inflorescence units cylindrical spikes, arranged in fascicles of 2–5 spikes in pseudoracemes or panicles. Flowers 5-merous, whitish, yellowish or reddish; calyx gamosepalous, cupulate; corolla gamopetalous, cupulate to tubular; stamens 10, anthers with an apical gland; pollen in 16-grained polyads, but also in groups of 4–28 grains; ovary included in the corolla. Fruit an indehiscent nucoid legume, curved, falcate or spiralled; valves coriaceous or woody. Seeds lenticular, wingless, pleurogram present.
Unknown.
Seven species of which six occur in Brazil, with one extending to Bolivia and another to French Guiana, Guyana, Suriname and Venezuela. One species is narrowly endemic in French Guiana (Fig.
Five species occur in the Amazon rainforest, while two are found in Cerrado savannas, one of which also extends to the seasonally dry tropical forests and woodlands of north-eastern Brazil.
Gwilymia is in homage to the Royal Botanic Gardens, Kew botanist, Dr. Gwilym P. Lewis.
Unknown.
The genus was recently described to accommodate species previously assigned to Stryphnodendron, but that were both morphologically and phylogenetically distinct (
Goldmania Rose ex Micheli, Mém. Soc. Phys. Genève 34: 274. 1903. Type: Goldmania platycarpa Rose [= Microlobius foetidus (Jacq.) M. Sousa & G. Andrade]
Microlobius mimosoides C. Presl [= Microlobius foetidus (Jacq.) M. Sousa & G. Andrade]
Trees or shrubs, indumentum composed of simple and granular trichomes, brachyblasts present; branches and leaves unarmed, with a strong garlic odour. Stipules caducous. Leaves bipinnate, extrafloral nectaries between pinnae pairs, and sometimes also between leaflets, but never on the petiole; pinnae 1–2 (3) pairs, opposite; leaflets 1–2 pairs, opposite, obovate or elliptic, with or without a tuft of trichomes at the base of the abaxial surface. Inflorescence units cylindrical spikes, arranged in fascicles of 2–5 spikes in pseudoracemes. Flowers 5-merous, white to yellow; calyx gamosepalous, campanulate; corolla gamopetalous, narrowly campanulate; stamens 10, anthers with an apical gland; pollen in 8-grained polyads; ovary included in the corolla. Fruit a follicle, subfalcate; valves coriaceous. Seeds obovate, wingless, white, pleurogram present.
2n = 26 (
Monospecific (M. foetidus), with two varieties. Microlobius foetidus var. foetidus occurs in Mexico, Honduras and Venezuela, and M. foetidus var. paraguensis (Benth.) M. Sousa & G. Andrade is restricted to Bolivia, Paraguay, Brazil and Argentina (Fig.
Seasonally dry tropical forests and woodlands.
From Greek, micro (= small) and lobos (= pods), in reference to the small lignified ovaries present in the original material, which were thought to be mature fruits (
Unknown.
The status of Microlobius as a distinct genus has been questioned since its description (as Goldmania), particularly in comparison to the original, more ample circumscription of Piptadenia (
Among members of the Stryphnodendron clade, Microlobius is diagnosed by the following combination of characters: presence of garlic odour in the wood and leaves, petioles without extrafloral nectaries, leaves with a single pair of pinnae, opposite leaflets, and fruits follicles with white seeds (
Folianthera Raf., Sylva Tellur.: 120. 1838. Type: Folianthera guianensis (Aubl.) Raf. [≡ Mimosa guianensis Aubl. (≡ Stryphnodendron guianense (Aubl.) Benth.)]
Stryphnodendron barbadetiman (Vell.) Mart. [= Stryphnodendron adstringens (Mart.) Coville]
Trees, shrubs, or geoxyle subshrubs; indumentum composed of simple and granular trichomes; brachyblasts absent; branches unarmed, young shoots and leaves ferruginous, covered with reddish granular trichomes, not odoriferous. Stipules usually caducous. Leaves bipinnate; extrafloral nectaries present on the petiole, between or just below pinnae pairs and between or just below the distal pairs of leaflets; pinnae (3) 5–32 pairs, subopposite, opposite or rarely alternate; leaflets 8–20 pairs, alternate. Inflorescence units cylindrical spikes, arranged in fascicles of 2–6 spikes in pseudoracemes. Flowers 5-merous, whitish, yellowish or reddish; calyx gamosepalous, campanulate; corolla gamopetalous, narrowly campanulate; stamens 10, anthers with an apical gland; pollen usually in 16-grained polyads, but also in groups of 4–32 grains; ovary included in the corolla. Fruit an indehiscent legume or a follicle, oblong, linear or slightly curved; valves woody or coriaceous. Seeds obovoid or ellipsoid, wingless, black, brown, or ochre, pleurogram present.
2n = 26 (
The majority of species occur either in rainforests (most in the Brazilian Amazon), or in savannas (Brazilian Cerrado), and four extend into seasonally dry tropical forests and woodlands (
From Greek, stryphno (= sour, adstringent) and dendron (= tree), in reference to the astringent properties of its bark.
Stryphnodendron adstringens is widely used due to its tannin-rich bark with astringent properties (
Phylogenetic and morphological evidence together with diagnosability supported the segregation of part of Stryphnodendron species into two new genera (Gwilymia and Naiadendron;
Among other members of the Stryphnodendron clade, the genus Stryphnodendron can be recognised by joint occurrence of ferruginous indumentum (densely covered with reddish granular trichomes) on young branches and leaves, relatively small (0.6–1.2 × 0.3–0.6 cm) and alternate leaflets, spikes arranged in pseudoracemes, and the fruit an indehiscent nucoid legume or a follicle.
Newtonia sect. Neonewtonia Burkart, Fl. Il. Catarin. fasc. LEGU: 285. 1979. Type: Newtonia nitida (Benth.) Brenan [≡ Piptadenia nitida Benth. (= Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro)]
Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro [≡ Acacia contorta DC.]
Trees; indumentum composed of simple trichomes; brachyblasts absent; branches and leaves unarmed, not odoriferous. Stipules caducous. Leaves bipinnate; extrafloral nectaries on the lower half of the petiole; pinnae (2) 5–many pairs, opposite; leaflets (6) 10–many pairs, opposite, mostly oblong to linear, rarely rhomboid. Inflorescence units cylindrical spikes, arranged in terminal pseudoracemes or clustered in efoliate nodes below the leaves. Flowers 5-merous, white to yellowish or greenish; calyx gamosepalous; corolla gamopetalous; stamens 10, anthers with an apical gland; pollen in polyads with 8, 12 or 16 grains; ovary included or exserted from the corolla. Fruit a follicle; margins straight, rarely sinuous and constricted where the seeds abort. Seeds flat compressed, dark, narrowly winged, pleurogram absent.
Unknown.
Six species, three in eastern Brazil, one in northern South America and Costa Rica, one endemic to Colombia, and one endemic to Venezuela (Fig.
All species of Marlimorimia occur in rainforests.
Marlimorimia honours the Rio de Janeiro Botanical Garden botanist, Dr. Marli Pires Morim.
Marlimorimia warmingii (Benth.) L.P. Queiroz & P.G. Ribeiro is used as firewood, timber, to make agricultural tools, and could be used as an ornamental and for ecological restoration (
The genus was described to accommodate species that could not retain the name Pseudopiptadenia Rauschert, as it had to be synonymised under Pityrocarpa based on phylogenetic evidence (see below and
Two species of the genus, Marlimorimia colombiana (Britton & Killip) L.P. Queiroz & Marc.F. Simon, from Colombia, and M. pittieri (Harms) L.P. Queiroz & L.M. Borges, from Venezuela, are poorly known and have been assigned to the genus based on the morphology of type specimens only (
Naiadendron duckeanum (Occhioni) A.G. Lima, Paula-Souza & Scalon [≡ Stryphnodendron duckeanum Occhioni]
Trees; indumentum composed of simple and granular trichomes; brachyblasts absent; branches unarmed, strongly striate, young shoots and leaves ferruginous with reddish granular trichomes, not odoriferous. Stipules caducous. Leaves bipinnate, not odoriferous; extrafloral nectaries on the petiole, rachis and pinnae; pinnae 10–22 pairs, subopposite to opposite; leaflets 15–23 pairs, opposite. Inflorescence units cylindrical spikes grouped in fascicles of 3–5 in pseudoracemes. Flowers 5-merous, white to yellowish; calyx gamosepalous; corolla gamopetalous; stamens 10, anthers with an apical gland; pollen in (12) 16-grained polyads; ovary included. Fruit a legume, dehiscent along both margins, linear to narrow-oblong, laterally-compressed; valves chartaceous. Seeds obovate or elliptic, wingless, ochre, pleurogram present.
Unknown.
Rainforests (terra firme, often disturbed), on clay or sandy soil.
From naiades, Greek mythology’s nymphs of freshwater, and dendron (Greek = tree) in reference to the name given to the Brazilian Amazon (Naiades) by Carl Friedrich Philipp von Martius, where the single species of the genus occurs.
Unknown.
Analysis of the extrafloral nectaries and fruits of Stryphnodendron duckeanum, initially described based on flowering material only, showed the species did not fit the limits of Stryphnodendron and that it could belong to Piptadenia (Rupert Barneby unpublished note;
Naiadendron and Parapiptadenia are the only members of the Stryphnodendron clade with typical legume fruits, i.e., dehiscing along both margins. However, Naiadendron is readily set apart by the ferruginous indumentum covering both its young branches and leaves. Strongly striate branches and petiolar nectaries 8–12 mm long also differentiate Naiadendron from all other members of the clade.
Parapiptadenia rigida (Benth.) Brenan [≡ Piptadenia rigida Benth.]
Trees; indumentum composed of simple trichomes; brachyblasts absent; branches and leaves unarmed, not odoriferous. Stipules present or absent. Leaves bipinnate; extrafloral nectaries on the petiole, and usually between the distal pair of pinnae and leaflets; pinnae 1–8 pairs, opposite; leaflets (1) 2–26 pairs, opposite, elliptic to oblong. Inflorescence units cylindrical spikes, solitary, axillary or supra-axillary to coevally developing leaves. Flowers 5-merous, reddish, rarely yellowish; calyx gamosepalous, campanulate; corolla gamopetalous, campanulate to tubular; stamens 10, anthers with an apical gland; pollen in 8, 12 or 16-grained polyads; ovary included in the corolla. Fruit a legume, flat-compressed, valves undulate above the seeds (rarely plane). Seeds flat-compressed, dark, winged, pleurogram absent.
2n = 26 (
Six species, four from north-eastern Brazil, one extending also to south-eastern Brazil; and two occurring in southern Brazil, Argentina, Paraguay, and Uruguay, one of which also reaches Bolivia, western Brazil and Peru (Fig.
Sub-tropical forests, rainforests, seasonally dry tropical forests and woodlands.
From para (Greek = next to) and piptadenia, in reference to the close taxonomic relationship to the genus Piptadenia (
Parapiptadenia rigida has medicinal properties, is used as fodder, timber, firewood, an ornamental, as a source of saponins, gum, tannins, cellulose, for paper production, and for ecological restoration (
In addition to Anadenanthera Speg., Parapiptadenia is the only segregate from Piptadenia that was confirmed as monophyletic (
Piptadenia sect. Pityrocarpa Benth., J. Bot. (Hooker) 4: 339. 1841. Type: Piptadenia moniliformis Benth. [≡ Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson]
Monoschisma Brenan, Kew Bull. 10(2): 179. 1955, nom. inval., non Monoschisma Duby, Mém. Soc. Phys. Genève 19: 294. 1868 (Musci, Meteoriaceae). Type: Monoschisma leptostachyum (Benth.) Brenan [≡ Piptadenia leptostachya Benth. (≡ Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro)]
Pseudopiptadenia Rauschert, Taxon 31(3): 559. 1982. Type: Pseudopiptadenia leptostachya (Benth.) Rauschert [≡ Piptadenia leptostachya Benth. (≡ Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro)]
(designated by
Trees or shrubs; indumentum composed of simple trichomes; brachyblasts absent; branches and leaves unarmed, not odoriferous. Stipules present, sometimes caducous. Leaves bipinnate; extrafloral nectaries between or just below the first pair of pinnae; pinnae 1–4 (10) pairs, opposite; leaflets 1–10 (20) pairs per pinna, opposite, rhomboid, sometimes asymmetrically elliptical or lanceolate. Inflorescences cylindrical spikes, white to yellowish or greenish, solitary (rarely 2) in the axils of coevally developing leaves. Flowers 5-merous; calyx cupulate; petals free (sometimes joined in one species); stamens 10, anthers with an apical gland; pollen in polyads with 8 or 16 grains; ovary included or exserted from the corolla; stigma in a terminal pore. Fruit a follicle, flat compressed, margins deeply and regularly constricted, rarely sinuous and shallowly constricted. Seeds flat compressed, dark, narrowly winged, or rarely ovoid or discoid, white and wingless, pleurogram absent from dark seeds, but present in the white ones.
2n = 26 (
Seven species occurring in three major areas in tropical America: Mexico and Central America, northern South America (Guyana, Venezuela), and eastern Brazil (Fig.
Rainforests (Brazil), seasonally dry tropical forests and woodlands (Brazil and Mexico), savannas (Venezuela) and Chaco (Paraguay).
From Greek, pityron (= scurf; husks of bran) and carpus (= fruit), in reference to the leprose fruits of some species (e.g., Pityrocarpa moniliformis).
Pityrocarpa moniliformis is used as fodder, timber for light constructions, firewood, and for soil enrichment, ecological restoration and honey production; it is also rich in tannins (
Originally described as a section of Piptadenia (
The recent updates to the circumscription of Pityrocarpa rendered the genus more variable with respect to seed morphology —a character previously used to distinguish it from Parapiptadenia and Pseudopiptadenia, for example— but also highlighted the taxonomic relevance of particular traits for recognition of the genus, such as extrafloral nectaries between or just below the first pair of pinnae and inflorescences in general solitary and axillary to coevally developing leaves (
Leonardo M. Borges5, Marcelo F. Simon40, Matías Morales33,34, Melissa Luckow29, Pétala Gomes Ribeiro2, Rosaura Grether19
Citation: Borges LM, Simon MF, Morales M, Luckow M, Ribeiro PG, Grether R (2024) 25. Mimosa clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 332–342. https://doi.org/10.3897/phytokeys.240.101716
Mimosa clade
Figs
Included genera (3). Adenopodia C. Presl (7 species), Mimosa L. (615), Piptadenia Benth. (28).
Description. Trees, shrubs, herbs, geoxyles, and lianas; brachyblasts absent or sometimes present; indumentum composed of either simple, glandular, or complex multicellular trichomes, or a combination of them; branches armed or not with scattered, serial, or nodal prickles. Stipules present, in general caducous. Leaves bipinnate (rarely once pinnate or phyllodinous with transversely dilated leafstalks), rachis armed or not; extrafloral nectaries absent (most Mimosa) or present on the petiole, sometimes also on the rachis and pinnae; pinnae 1–many pairs, opposite, armed or not; leaflets 1–many pairs per pinna, sessile, mostly opposite, variable in size and shape. Inflorescence globose or ellipsoid capitula, or cylindrical racemes or spikes, solitary, fasciculate or organised in complex synflorescences, sometimes developing from short shoots on efoliate branches. Flowers 3–5 (6)-merous, haplo- or diplostemonous; anther glands absent (Mimosa) or present; pollen in tetrads or in polyads with 8, 12 or 16 grains; ovary linear to oblong or elliptic, indumentum variable, stigma generally in a terminal pore. Fruit a legume with entire valves, a craspedium (valves breaking up in monospermic articles leaving persistent margins), or an unjointed craspedium (valves remain entire after separating from the margins), variable in size, shape, indumentum and seed number. Seeds lenticular, elliptic to subspherical or rhomboid, wingless; pleurogram present.
Geographic distribution. Species of the Mimosa clade occur in almost all tropical and subtropical vegetation formations from the USA to Argentina, but are most diverse in the dry areas of the American tropics. Approximately 40 species of Mimosa are native to continental Africa, Madagascar and Asia, while four species of Adenopodia occur in Africa.
Clade-based definition. The most inclusive crown clade containing Piptadenia adiantoides (Spreng.) J.F. Macbr. and Mimosa pudica L., but not Senegalia nigrescens (Oliv.) P.J.H. Hurter, Stryphnodendron adstringens (Mart.) Coville or Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow (Fig.
Notes. Members of the Mimosa clade were previously treated as part of the Piptadenia group, an informal assemblage of genera with haplo- or diplostemonous flowers, pollen in tetrads or polyads, and a narrowing style with a small porate stigma at the tip (
All three genera of the Mimosa clade include plants armed with internodal prickles (although unarmed plants are common in Mimosa) with leaflets varying greatly in size and number and flowers arranged in spiciform racemes or spikes, even though globose capitula are common in Mimosa. Mimosa stands out from the other two genera by its anthers lacking apical glands; Adenopodia and Mimosa share fruits with valves separating from the persistent margins and breaking up into monospermic segments (a craspedium; in a few species of Mimosa the valves remain entire forming an unjointed craspedium, and in one species the fruits are lomentiform), while those of Piptadenia are typical legumes dehiscing along both sutures; some species also have non-pentamerous flowers (3-, 4- or more rarely 6-merous in Mimosa), and extrafloral nectaries are present across all three genera of the clade including in Mimosa section Mimadenia Barneby, but subsequently lost across the vast majority of species of Mimosa (
Phylogenomic analyses show that Adenopodia is robustly supported as sister to Mimosa (Fig.
Piptadenia latifolia Benth. [= Piptadenia adiantoides (Spreng.) J.F. Macbr.]
Trees 3–30 m, shrubs or lianas; indumentum composed of simple trichomes, commonly absent; brachyblasts absent; branches armed with recurved or straight prickles, these scattered or aligned in raised ribs, rarely in groups of three at the leaf nodes. Stipules present, caducous. Leaves bipinnate, commonly with prickles on petioles and pinnae; extrafloral nectaries on the petiole, on the rachis between distal pairs of pinnae, and on the pinnae between distal pairs of leaflets; pinnae 1–14 pairs, opposite; leaflets 1–many pairs, opposite. Inflorescences cylindrical spikes, white, cream, yellow or red, isolated, in fascicles, or arranged in complex racemose or paniculate synflorescences. Flowers 5-merous, diplostemonous; calyx gamosepalous; corolla gamopetalous; stamens 10, anthers with a caducous apical claviform gland; pollen in tetrads or in polyads with 8, 12 or 16 grains; ovary glabrous to pubescent; stigma in a terminal pore. Fruit a flat compressed legume with thick margins, straight, usually papery. Seeds compressed, obovate to orbicular in outline, pleurogram present.
2n = 26 [Piptadenia retusa (Jacq.) P.G. Ribeiro, Seigler & Ebinger;
Twenty-eight species in tropical America, from western and southern Mexico to southern Brazil and perhaps also at the northern limits of Argentina (Fig.
Adenopodia and Piptadenia diversity A Adenopodia gymnantha Brenan fruiting branch B fruits C Piptadenia ramosissima Benth., branch with prickles aligned along raised ribs D P. retusa (Jacq.) P.G. Ribeiro, Seigler & Ebinger, vegetative branch E, F P. micracantha Benth. E flowering branch F fruits. Photo credits A, B EOA Pérez C–F LP Queiroz.
Rainforests, seasonally dry tropical forests and woodlands, and riparian forests within Neotropical savannas.
From Greek, pipto (= to fall) and aden (= gland), in reference to the caducous anther glands.
Some species are used as a fodder, a source of tannins, as ornamentals, for timber, woodwork, firewood, paper and honey production, soil enrichment and ecological restoration (
Piptadenia was originally broadly circumscribed to accommodate mimosoid legumes with glandular anthers, and flat, dehiscent fruits with thin valves (
Although a densely sampled phylogeny of Piptadenia is still pending, the genus can be diagnosed by the stems armed with recurved prickles, extrafloral nectaries on the petioles, cylindrical spikes, presence of anther glands, and flat, papery, straight legumes. Three species in the genus are known to nodulate (
Pseudoentada Britton & Rose, N. Amer. Fl. 23: 191. 1928. Type: Pseudoentada patens (Hook. & Arn.) Britton & Rose [≡ Inga patens Hook. & Arn. (≡ Adenopodia patens (Hook. & Arn.) J.R. Dixon ex Brenan)]
Entada subg. Acanthentada Brenan, Kew Bull. 20: 366. 1966. Type: Entada spicata (E. Mey.) Druce [≡ Mimosa spicata E. Mey. (≡ Adenopodia spicata (E. Mey.) C. Presl)]
Adenopodia spicata (E. Mey.) C. Presl [≡ Mimosa spicata E. Mey.]
Lianas, rarely shrubs or treelets; indumentum composed of simple trichomes; brachyblasts absent; branches armed with scattered, sometimes paired, recurved prickles. Stipules present, caducous or persistent. Leaves bipinnate, often with prickles on petioles and pinnae; extrafloral nectaries on the petiole and sometimes also between each pinnae pair; pinnae 1–many pairs, opposite; leaflets 1–many pairs, opposite. Inflorescences cylindrical spikes, white, yellow, pink or purple, often organised in panicles. Flowers 5-merous, diplostemonous; calyx gamosepalous; corolla polypetalous or gamopetalous; stamens 10, anthers with an apical gland; pollen in 16-grained polyads; ovary pubescent. Fruit a craspedium, straight or curved, sometimes armed along the margins. Seeds ellipsoid to spheroid; pleurogram present.
Unknown, indicated as possibly 2n = 28 by
Seven species (see notes below). Three species are restricted to Mexico and Central America, and four occur disjunctly across sub-Saharan Africa (Fig.
In the New World confined to seasonally dry tropical forest, in Africa, on the margins of rain and deciduous forests, thickets and disturbed habitats.
From Greek, adeno- (= gland) and -podia (= foot), likely a reference to the shortly pedicellate anther glands.
Adenopodia spicata has anti-hypertensive properties (
Schrankia Willd., Sp. Pl., ed. 4(2): 888, 1041. 1806, nom. cons. Lectotype: Schrankia aculeata Willd. [= Mimosa quadrivalvis L.]
Eburnax Raf., New Fl. [Rafinesque] 1: 42. 1836. Type: Eburnax pudica (L.) Raf. [≡ Mimosa pudica L.]
Sensitiva Raf., Sylva Tellur.: 119. 1838. Type: Sensitiva pudica Raf. [≡ Mimosa pudica L. ?]
Lomoplis
Raf., Sylva Tellur.: 118. 1838. Lectotype (designated by
Leptoglottis DC. ex Torr. & A. Gray, Fl. N. Amer. (Torr. & A. Gray) 1: 695. 1840. Type: Leptoglottis nuttallii DC. ex Torr. & A. Gray [≡ Mimosa quadrivalvis var. nuttallii (DC. ex Torr. & A. Gray) Beard ex Barneby]
Morongia Britton, Mem. Torrey Bot. Club 5: 191. 1894. Lectotype (designated here): Morongia angustata (Torr. & A. Gray) Britton [≡ Mimosa quadrivalvis var. angustata (Torr. & A. Gray) Barneby]
Schranckiastrum Hassl., Repert. Spec. Nov. Regni Veg. 16: 151. 1919. Type: Schranckiastrum insigne Hassl. [≡ Mimosa insignis (Hassl.) Barneby]
Acanthopteron Britton, N. Amer. Fl. 23: 179. 1928. Type: Acanthopteron laceratum (Rose) Britton [≡ Mimosa lacerata Rose]
Haitimimosa Britton, N. Amer. Fl. 23: 179. 1928. Type: Haitimimosa extranea (Benth.) Britton [≡ Mimosa extranea Benth.]
Mimosopsis Britton & Rose, N. Amer. Fl. 23: 174. 1928. Type: Mimosopsis prolifica (S. Watson) Britton & Rose [≡ Mimosa prolifica S. Watson]
Neomimosa Britton & Rose, N. Amer. Fl. 23: 172. 1928. Type: Neomimosa eurycarpa (B.L. Rob.) Britton & Rose [≡ Mimosa eurycarpa B.L. Rob.]
Pteromimosa Britton, N. Amer. Fl. 23: 171. 1928. Type: Pteromimosa bahamensis (Benth.) Britton [≡ Mimosa bahamensis Benth.]
Mimosa sensitiva L.
Perennial shrubs, herbs, geoxyle subshrubs, lianas, treelets and trees, occasionally monocarpic; indumentum composed of numerous different combinations of simple, glandular, and complex multicellular trichomes and sharp-pointed setae; brachyblasts sometimes present; branches armed or not with scattered, serial, or nodal prickles. Stipules present, caducous or persistent. Leaves bipinnate, rarely once-pinnate (M. unipinnata B.D. Parfitt & Pinkava), sometimes reduced to phyllodes bearing ephemeral pinnae (M. ephedroides (Gillies ex Hook. & Arn.) Benth., M. extranea Benth., M. phyllodinea Benth.), rachis armed or not, sometimes with a spicular projection between pinnae pairs; extrafloral nectaries absent in the vast majority of species, if present, on the petiole and sometimes also on the rachis and pinnae; pinnae 1–many pairs, opposite, armed or not; leaflets 1–many pairs per pinna, sessile, mostly opposite, variable in size and shape; paraphyllidia mostly present. Inflorescences globose or ellipsoid capitula, cylindrical spikes or racemes, white, pink, yellow or red, solitary or fasciculate in the axils of coevally developing leaves, or organised in complex efoliate synflorescences, sometimes developing from short shoots on efoliate branches. Flowers 3–5 (6)-merous, haplo- or diplostemonous; bisexual or staminate, the latter located at the lower parts of the inflorescence, rarely all flowers bisexual; calyx gamosepalous; corolla gamopetalous; anther glands absent; pollen in tetrads or polyads with 8, 12, or rarely 16 grains; ovary glabrous to lanose. Fruit a typical craspedium or an unjointed craspedium, rarely lomentiform or follicular, variable in size, shape, indumentum, armature and seed number. Seeds lenticular, ellipsoid to spheroidal or rhombic.
The basic chromosome number in Mimosa is reported to be x = 13 (
615 species. Most are local endemics distributed right across the Americas from the USA to Argentina, with ca. 40 occurring in Africa, Madagascar (32 endemics) and Asia (Fig.
Mimosa vegetative diversity A M. dasilvae A.S.L. Silva & Secco, a shrub B M. pumilio Barneby, a geoxyle subshrub (Simon 3187) C M. dutrae Malme, a trailing subshrub D M. texana var. filipes (Britton & Rose) Barneby, a tree (Simon 845) E M. oedoclada Barneby showing the indumentum composed by simple, glandular and setiform trichomes (Simon 2588) F M. supravisa Barneby showing a branch armed with prickles (Jordão 442). Photo credits A CO Andrino B, D–F MF Simon C N Dahmer.
Mimosa leaf, inflorescence and fruit diversity A M. tequilana S. Watson showing bipinnate leaves with one pair of pinnae and two pairs of broad leaflets per pinna (Simon 813) B M. splendida Barneby showing multi-pinnate leaves with many small leaflets (Simon 739) C M. bimucronata (DC.) Kuntze showing capitate inflorescences (Simon 872) D M. benthamii J.F. Macbr. showing cylindrical inflorescences E M. irrigua Barneby showing a craspedium F M. foliolosa Benth. showing many unjointed craspedia. Photo credits A–C, F MF Simon D R Grether E D Cardoso.
Mimosa is one of the most ecologically adaptable, widespread and ubiquitous genera of Mimoseae, occurring across the full tropical rainfall gradient and in almost all tropical and subtropical vegetation types in the Americas, including seasonally dry tropical and subtropical forests and semi-arid thorn scrub, desert, tropical and subtropical savannas and grasslands, lowland tropical rainforest, mid-elevation moist subtropical forest, wetlands, temperate forests, rarely on sand dunes, and often in disturbed open habitats. Some species become weedy outside their native range. This ecological adaptability is closely correlated with the diversity of growth forms - lianas in wet forests, stiffly-branched xerophytic shrubs in arid and semi-arid ecologies, and geoxyles in savannas (
Either from Spanish, mimoso (= sensitive;
Mimosa species are used as living fences (e.g., M. caesalpiniifolia Benth. in Brazil), timber (e.g., M. scabrella Benth. in southern Brazil and Argentina), fodder (e.g., M. strigillosa Torr. & A. Gray, in Argentina and the USA;
Mimosa is classified into a complex infrageneric and infraspecific hierarchy (five sections, 41 series, 39 subseries, 18 subspecies and 268 varieties;
Most species nodulate, particularly with beta-rhizobia (Reis Junior et al. 2010;
Rapid movement of leaves and leaf parts in response to touch is a trademark of Mimosa. Although leaf movement evolved in many lineages within the genus, it is restricted to less than 5% of the species (
The highly disjunct amphi-Atlantic distribution of Mimosa (Fig.
We could not confirm the type species of Sensitiva, one of the many generic synonyms of Mimosa. Although Sensitiva was validly published and circumscribed to include species close to M. pudica and M. sensitiva,
Bruce R. Maslin31,32, Vanessa Terra41, David S. Seigler38, John E. Ebinger14, Colin E. Hughes3
Citation: Maslin BR, Terra V, Seigler DS, Ebinger JE, Hughes CE (2024) 26. Senegalia grade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 343–357. https://doi.org/10.3897/phytokeys.240.101716
Senegalia grade
Figs
Included genera (4). Mariosousa Seigler & Ebinger (14 species), Parasenegalia Seigler & Ebinger (11), Pseudosenegalia Seigler & Ebinger (2), Senegalia Raf. (219).
Distribution. Pantropical; all four genera occur in the New World, but Senegalia extends to Africa, Asia and Australia.
Notes. Species of the four Senegalia grade genera, together with species of Acacia Mill., Acaciella Britton & Rose and Vachellia Wight & Arn., were formerly included in what was previously called Acacia s.l. Recent phylogenomic analyses have confirmed that, as broadly circumscribed, Acacia s.l. is polyphyletic (
A number of morphological characters, typical of all (or most) species of the four Senegalia grade genera, help distinguish these genera from the other three that were originally included in Acacia s.l. These include the absence of stipular spines, the bipinnate leaves having glands on the petiole and/or rachis (although occasionally absent from some individual plants), peduncles lacking a multi-bracteate involucre, flowers commonly sessile or subsessile (but if pedicellate, the pedicel not persistent on receptacle after flowers have dropped), and pollen (unknown for Pseudosenegalia) comprising 16-grained polyads, the grains porate, and lacking pseudocolpi and having a granular exine (i.e., lacking columellae). For further discussion on the morphological characters of Acacia, Acaciella, and Vachellia see their respective treatments in the Archidendron clade (page 404), Calliandra clade (page 358), and Parkia clade (page 299).
For several years prior to fragmentation Acacia s.l., many species that are now included in genera of the Senegalia grade had been assigned to Acacia subg. Aculeiferum Vassal by various authors (e.g.,
Over the following two decades, compelling evidence, especially from phylogenetic studies, confirmed that Acacia s.l. was polyphyletic and could not be sustained as a single genus (e.g.,
Pseudosenegalia was shown to be monophyletic and sister to a subclade containing Mariosousa and Parasenegalia by
Despite the segregation of these four genera, the large, pantropical genus Senegalia is not always supported as monophyletic in recent phylogenomic studies (
A key to the four genera of the Senegalia grade, plus Acaciella, is provided in
Manganaroa
Speg., Bol. Acad. Nac. Ci. [Córdoba] 26(2): 227. [12 Oct.] 1922. Lectotype (designated by
Dugandia Britton & Killip, Ann. New York Acad. Sci. 35(3): 137. 1936. Type: Dugandia rostrata (Humb. & Bonpl. ex Willd.) Britton & Killip [≡ Acacia rostrata Humb. & Bonpl. ex Willd. (≡ Senegalia rostrata (Humb. & Bonpl. ex Willd.) Seigler & Ebinger)]
(designated by Britton and Rose, N. Amer. Fl. 23: 106. 1928). Senegalia triacantha Raf., nom. illeg. [≡ Mimosa senegal L. (≡ Senegalia senegal (L.) Britton)]. Note:
Two sections are currently recognised:
Senegalia sect. Senegalia
Acacia subg. Aculeiferum Vassal, Bull. Soc. Nat. Hist. Toulouse 108: 138. 1972; Vassal, Trav. Lab. Forest. Toulouse Tome 1, Vol. 8, Art. 17: 15. 1972. Type: Acacia senegal (L.) Willd. [≡ Mimosa senegal L. (≡ Senegalia senegal (L.) Britton]. Note: The subgenus was based on Acacia ser. Vulgares Benth., London J. Bot. 1: 322. 1842, not on Acacia ser. Vulgares and ser. Filicinae Benth. as given by
Senegalia sect. Monacanthea (Vassal) Maslin, Pl. Diversity 41: 371. 2019.
Acacia subg. Aculeiferum sect. Monacanthea Vassal, Bull. Soc. Nat. Hist. Toulouse 108: 139. 1972; Vassal, Trav. Lab. Forest. Toulouse Tome 1, Vol. 8, Art. 17: 15. 1972. Type: Acacia ataxacantha DC. [≡ Senegalia ataxacantha (DC.) Kyal. & Boatwr.]
Armed (with prickles) lianas, shrubs or trees (rarely taller than ca. 12 m); bark usually grey to brown and hard, sometimes yellowish and papery or corky when young; brachyblasts sometimes present; prickles present on branchlets (1, 2 or 3 at leaf nodes, or scattered irregularly or in rows along internodes; sometimes absent from some individuals and/or herbarium specimens) and often on underside of petiole and/or rachis. Stipules usually small and caducous, infrequently large and ± persistent, not spinescent. Leaves bipinnate, not sensitive; extrafloral nectaries present on petiole and/or rachis (sessile or sometimes stipitate) and normally on the pinnae (sessile); pinnae 1–40 (60) pairs; paraphyllidia present or absent; leaflets opposite or rarely alternate [e.g., S. comosa (Gagnep.) Maslin, Seigler & Ebinger, sect. Monacanthea s.s.], petiolulate or sometimes sessile, (1) 3–80 (100) pairs per pinna. Inflorescences comprising pedunculate heads, spikes or rarely spiciform racemes that are solitary or in fascicles in leaf axils or arranged in compound racemes or panicles; peduncles lacking a multi-bracteate involucre. Flowers all hermaphrodite or sometimes staminate and hermaphrodite within the one inflorescence, uniform, 5-merous, with a basal nectariferous disk, sessile to sub-sessile or sometimes pedicellate, commonly white to cream, sometimes yellow or red; perianth connate, valvate, not scarious; stamens numerous (ca. 30+), free; anther glands present or absent; pollen normally comprising 16-grained polyads, normally porate and lacking pseudocolpi, exine surface psilate (often with circular depressions) or variously rugulate, exine lacking columellae (or rarely columellae very short); ovary sessile to stipitate. Fruits normally dehiscent, rarely indehiscent [e.g., S. pentagona (Schumach. & Thonn.) Kyal. & Boatwr., sect. Monacanthea p.p.] or breaking into 1-seeded articles (e.g., S. rostrata, S. monacantha, sect. Monacanthea p.p.), clearly flattened or rarely elliptic in cross-section, valves normally chartaceous to coriaceous, infrequently crustaceous to sub-woody [e.g., S. rugata (Lam.) Britton & Rose, sect. Monacanthea p.p.]. Seeds not winged, exarillate; pleurogram U-shaped (open at hilar end), occasionally circular or elliptic, rarely absent [e.g., S. pedicellata (Benth.) Seigler & Ebinger, sect. Monacanthea p.p.].
2n = 26 (known for 25 species, including those of sect. Senegalia and sect. Monacanthea p.p.), 2n = 39, 52 [S. laeta (R. Br. ex Benth.) Seigler & Ebinger, sect. Senegalia], 2n = 40 [S. galpinii (Burt Davy) Seigler & Ebinger, sect. Senegalia] and 2n = 26, 52 and 104 (S. ataxacantha, sect. Monacanthea s.s.) (
A total of 219 species accommodated in three infrageneric groups (see Notes below): sect. Senegalia (51 species), sect. Monacanthea s.s. (four species) and sect. Monacanthea p.p. (164 species). The genus is distributed pantropically in the Americas (99 species), the African region (Africa and Madagascar, 68 species), Asia (Arabian Peninsula to East and South East Asia, 57 species) and north-east Australia (two species) (Fig.
The majority of Senegalia species occur in seasonally dry tropical habitats, especially seasonally dry tropical forests, scrublands and savannas. Despite this predilection for seasonally dry biomes, the genus shows wide adaptability, with some species occurring in wetter lowland tropical vegetation types, which accounts for its almost cosmopolitan distribution across the tropics. Details of habitats for many species of Senegalia are provided in
The genus name refers to Senegal, a country in West Africa where the lectotype S. senegal, was collected.
The most valuable commercial species of Senegalia is the widespread African S. senegal (sect. Senegalia), which is the classical source of Gum Arabic. This water-soluble gum is also derived from a variety of other sources, including Vachellia seyal (Delile) P.J.H. Hurter, where it is harvested from trees both in the wild and in plantations. Gum Arabic is used primarily in the food and soft-drink industries as a stabilizer but also has applications in the production of paint, glue and cosmetics, in textile industries and for viscosity control in inks. Also in Africa, the two sect. Senegalia species S. nigrescens and S. caffra (Thunb.) P.J.H. Hurter & Mabb. are sources of tannin (
In Asia, the most commonly utilised species is S. catechu (L. f.) P.J.H. Hurter & Mabb. (sect. Senegalia). In Pakistan for example, its very durable and termite-resistant wood is used for house construction and for making agricultural implements; it is also an excellent source of firewood and is regarded as one of the best woods for charcoal production; tannin extracted from the wood is used for tanning (
In the Americas many species of Senegalia are used locally for firewood; the wood is often converted to charcoal, but this is not usually economically important. Furthermore, almost all species of Senegalia that are large enough are used for making small tools, small instruments, furniture, and fence posts. The larger species are sometimes used for lumber [e.g., Senegalia aristeguietana (L. Cárdenas) Seigler & Ebinger], but most do not have significant commercial value.
Prior to the recent fragmentation of the former, broadly defined genus Acacia, species now regarded as Senegalia had been referred to Acacia subg. Aculeiferum as defined by
In summary, the present concept of Senegalia is based on a combination of genetic and morphological evidence as detailed by
Senegalia sect. Senegalia: cauline prickles 1, 2 or 3 at the nodes, flowers almost always in spikes; 51 species in Africa and Asia (Fig.
Morphological features of A–C Senegalia sect. Monacanthea s.s. and D–J Senegalia sect. Senegalia A–C Senegalia ataxacantha (DC.) Kyal. & Boatwr. A lianescent shrub habit B branch showing internodal prickles C spicate inflorescence, Pretoria National Botanical Garden, South Africa D, E S. modesta (Wall.) P.J.H. Hurter D papery peeling bark on branch, living collection at Singapore Botanic Gardens E branch showing two prickles at nodes and leaves with few pinnae, Asia F S. catechu (L.f.) P.J.H. Hurter & Mabb. spicate inflorescence, South China Botanic Garden, Guangzhou G S. laeta (R. Br. ex Benth.) Seigler & Ebinger habit H S. senegal (L.) Britton three cauline prickles at nodes I S. goetzei (Harms) Kyal. & Boatwr. two cauline prickles at nodes and spicate inflorescence J S. polyacantha subsp. campylacantha (Hochst. ex A. Rich.) Kyal. and Boatwr. habit. Photo credits A P Birnbaum B S Piry C E Koenen D, E B Maslin F Y Chen G M Schmidt H A Dreyer I C Boucher Chisale J E Faust. A, B, G–J from African plants – A Photo Guide (www.africanplants.senckenberg.de).
Senegalia sect. Monacanthea s.s.: cauline prickles internodal, flowers in spikes; four species in Africa (Fig.
Senegalia sect. Monacanthea p.p.: cauline prickles internodal but sometimes also at the nodes in some American species; flowers in heads or spikes; 164 species in Australia, Asia, Africa and the Americas. The name Manganaroa is available if this group is ever treated as a distinct genus (Fig.
Morphological features of Senegalia sect. Monacanthea p.p. A Senegalia × emoryana (Benth.) Britton & Rose habit, New World B S. micrantha (Benth.) Britton & Rose leaf, New World C S. grandistipula (Benth.) Seigler & Ebinger large foliaceous stipules (such stipules not especially common in Senegalia) (Terra 715), New World D Senegalia × emoryana internodal prickles, New World E S. subsessilis Britton & Rose thin-textured fruits, New World F S. sakalava (Drake) Boatwr. globose scarlet inflorescence (scarlet flowers are rare in Senegalia but are found in several species from Madagascar) (Koenen 215), Madagascar G S. pruinescens (Kurz) Maslin, Seigler & Ebinger lianescent shrub habit and terminal paniculate inflorescences (insert of head showing calyx red in upper part) (Maslin 11023), Asia H S. pennata subsp. insuavis (Lace) Maslin, Seigler & Ebinger liana habit (Maslin 11016), Asia I S. menabeensis (Villiers & Du Puy) Boatwr. spicate inflorescences (Du Puy M359), Madagascar J S. rugata (Lam.) Britton & Rose hard-textured pods, Macau, China K S. tonkinensis (I.C. Nielsen) Maslin, Seigler & Ebinger branch showing internodal prickles and two glands on petiole (uncommon in Senegalia) (Maslin 11041), Asia. Photo credits A, B, D, E, H B Maslin C V Terra F E Koenen G, K L Bai I D Du Puy J L-x Yuan.
The pollen description above is based on
Distribution of Senegalia based on quality-controlled digitised herbarium records. Note that the Indian subcontinent was only sparsely sampled for this map. Details of species distribution, based on states of India and other countries of the subcontinent, are provided in
Parasenegalia skleroxyla (Tussac) Seigler & Ebinger [≡ Acacia skleroxyla Tussac]
Unarmed trees (some to 25 m), shrubs (often lianescent) or lianas; bark (known for only few species) grey to brown, smooth to shallowly flat-ridged or shallowly fissured; brachyblasts absent. Stipules small, often falling tardily, rarely absent (P. lundellii Seigler & Ebinger), not spinescent. Leaves bipinnate, not sensitive; extrafloral nectaries present on petiole (variably positioned) and rachis (below uppermost 1–7 pairs or sometimes all pairs of pinnae), sessile or occasionally stipitate; pinnae normally 3–16 (–20) pairs [except 1–2 pairs in P. miersii (Benth.) Seigler & Ebinger]; paraphyllidia present or absent; leaflets opposite, normally 6–46 (65) pairs per pinna (except 1 or rarely 2 pairs in P. miersii). Inflorescences comprising pedunculate, loosely flowered spikes (1–4 in leaf axils) or densely flowered heads (1–4 in leaf axils or arranged in large, axillary and/or terminal racemes or panicles); peduncles lacking a multi-bracteate involucre. Flowers hermaphrodite, uniform, 5-merous, with a basal nectariferous disk, sessile, white to cream (sometimes aging yellowish); perianth connate, valvate, not scarious; stamens numerous (40–140), free; anther gland normally present; pollen comprising 16-grained polyads, the grains porate and lacking pseudocolpi, exine surface psilate with circular depressions or rugulate-perforate, exine lacking columellae; ovary sessile to subsessile (except stipitate with stipe to 1.6 mm long in P. vogeliana). Fruits dehiscent along both sutures (sometimes tardily so), flattened, valves chartaceous or coriaceous. Seeds sometimes with a narrow marginal wing, exarillate; pleurogram U-shaped (open at hilar end) or sometimes circular, rarely absent (e.g., P. miersii) (Fig.
Morphology of Mariosousa, Parasenegalia and Pseudosenegalia A Mariosousa acatlensis (Benth.) Seigler & Ebinger spicate inflorescences (Seigler 16002) B M. coulteri (Benth.) Seigler & Ebinger bark not papery C M. dolichostachya (S.F. Blake) Seigler & Ebinger branchlet unarmed (Seigler 16035) D–F M. salazari (Britton & Rose) Seigler & Ebinger D exfoliating papery bark (Seigler 16057) E habit F fruits (Seigler 15978) G M. usumacintensis (Lundell) Seigler & Ebinger leaf (Seigler 16027) H Pseudosenegalia feddeana (Harms) Seigler & Ebinger fruits (Atahuachi 1146) I, J Parasenegalia miersii (Benth.) Seigler & Ebinger (Terra, Coutinho & Dalvi 667) I branch showing unusual leaves (i.e., pinnae few and leaflets large) J inflorescences K Pseudosenegalia feddeana contorted tree habit. Photo credits A–G B Maslin H, K CE Hughes I, J V Terra.
2n = 26 (known only for P. visco), fide
Eleven species, widely but discontinuously distributed in the Neotropics from the Caribbean (three endemic species) through Central America (one species in Belize and Guatemala) to South America (two species in Argentina, Bolivia, Chile and Peru, and five endemic to Brazil) (Fig.
Commonly found in evergreen or semi-evergreen tropical, often riparian, forests or disturbed second growth forests and thickets, 0–800 m. (except P. visco from northern Argentina, Bolivia, northern Chile and Peru, which grows in seasonally wet montane regions between 1000–3000 m), as well as seasonally dry deciduous or semi-deciduous forests, savannas and desert scrub.
The genus name Parasenegalia is from the Greek para (= beside, near) + Senegalia, in reference to the close relationship to that genus.
The wood of P. muricata (L.) Seigler & Ebinger is sometimes used for construction and P. visco is commonly cultivated in Peru where it is economically important as a source of cabinet wood (
Prior to the recognition of Parasenegalia, most species now assigned to this genus had been included in a broadly circumscribed Senegalia (e.g.,
When Parasenegalia was first recognised it included seven species that were comprehensively described and illustrated (see Seigler and Ebinger in
Parasenegalia and Pseudosenegalia were simultaneously described by
The principal morphological characters that distinguish Parasenegalia from Pseudosenegalia are noted below under the latter genus. Note that pollen data are only available for three species of Parasenegalia, namely, P. miersii, P. santosii (G.P. Lewis) Seigler & Ebinger and P. visco (
Pseudosenegalia feddeana (Harms) Seigler & Ebinger [≡ Acacia feddeana Harms]
Unarmed trees (some to 12 m) or shrubs; bark white to grey-white and smooth; brachyblasts present at some or all nodes. Stipules small, persistent or falling late, not spinescent. Leaves bipinnate, not sensitive; pinnae 1–7 pairs; paraphyllidia absent; leaflets opposite, 11–26 pairs per pinna; gland present at apex of petiole and (when leaves multijugate) on rachis below the uppermost 1–2 pinna pairs, sessile. Inflorescences comprising pedunculate, loosely flowered spikes, 1 or 2 in leaf axils; peduncles lacking a multi-bracteate involucre. Flowers hermaphrodite, uniform, 5-merous, with a basal nectariferous disk, sessile, white or pale creamy white; perianth connate, valvate, scarious; stamens numerous (100–150), free; anther glands absent; pollen not studied; ovary stipitate. Fruits dehiscent along both sutures, flattened or elliptic in cross-section, valves coriaceous, twisting after dehiscence. Seeds not winged, exarillate; pleurogram U-shaped (open at hilar end) or absent in P. feddeana (Fig.
Unknown.
In seasonally dry tropical forest, dry scrub and thorn-scrub between 1300 and 3300 m elevation.
The generic name Pseudosenegalia is from Greek pseudo (= false) + Senegalia, in reference to a superficial resemblance to that genus.
Pseudosenegalia feddeana is important locally as a forage plant and its wood is used as a fuel; also, its seeds are edible and used as a coffee substitute (
Both species of Pseudosenegalia were originally described under Acacia and subsequently transferred to Senegalia by
Species of Pseudosenegalia are readily distinguished from those of the more speciose and widespread Parasenegalia by their extremely small leaflets (1–3 × 0.5–1 mm compared with normally (3) 5–18 × 0.5–11 mm in Parasenegalia), the presence of brachyblasts on branchlets, the absence of anther glands and by a scarious-textured perianth. The latter trait is otherwise unknown in the Senegalia grade. Even in the very large genus Acacia (1082 species), only one species, A. unifissilis Court, is described as having a scarious calyx (
Mariosousa coulteri (Benth.) Seigler & Ebinger [≡ Acacia coulteri Benth.]
Unarmed shrubs and trees; bark hard and fissured or scaly, sometimes papery and exfoliating; brachyblasts normally absent (except commonly present in M. compacta (Rose) Seigler & Ebinger). Stipules small, mostly persistent, not spinescent. Leaves bipinnate, not sensitive; extrafloral nectaries present on petiole and/or rachis (petiole glands commonly absent in M. gentryi Seigler & Ebinger, M. millefolia (S. Watson) Seigler & Ebinger and M. salazarii (Britton & Rose) Seigler & Ebinger), sessile or occasionally stipitate; pinnae 1–20 (30) pairs; paraphyllidia present or absent; leaflets opposite, (4) 8–65 pairs per pinna. Inflorescences comprising pedunculate, loosely flowered spikes, 1–4 in leaf axils or arranged in short, normally terminal, racemose clusters; peduncles lacking a multi-bracteate involucre. Flowers hermaphrodite, uniform, 5-merous, with a basal nectariferous disk, sessile, creamy white; perianth connate, valvate, not scarious; stamens numerous (50+), free; anther glands normally present; pollen comprising 16-grained polyads, porate and lacking pseudocolpi, exine surface rugulate, exine lacking columellae; ovary stipitate. Fruits dehiscent along both sutures, strongly flattened, valves chartaceous or coriaceous. Seeds exarillate, not winged, pleurogram U-shaped (open at hilar end) and areole usually large (covering 50%–70% of the seed) (Fig.
Unknown.
Fourteen species distributed from Arizona and New Mexico (one species), south through Mexico (where all species of the genus occur) to Belize, Guatemala, El Salvador, Honduras, Nicaragua and Costa Rica (where three species occur, the most widespread being M. centralis (Britton & Rose) Seigler & Ebinger) (Fig.
Mainly in seasonally dry tropical forests, thorn-scrub and thickets, weakly extending into moister semi-deciduous tropical forests; on arid hills or sometimes rocky deserts or desert grasslands, 0–2200 m elevation.
The genus Mariosousa honours Mario Sousa (1940–2017), legume specialist and former Director of the Herbarium of the Instituto de Biología (MEXU), Universidad Nacional Autónoma de México.
Mariosousa coulteri is one of the few trees of this genus with enough size and quality of wood to be of some local importance and useful for lumber (
Immediately prior to the description of Mariosousa, the 13 species that were subsequently assigned to this genus were included in the informal Acacia coulteri species-group (
Phylogenetic analyses of plastid and/or nuclear sequence data have supported the genus as monophyletic (
Mariosousa heterophylla (Benth.) Seigler & Ebinger [= Acacia willardiana Rose, ≡ Mariosousa willardiana (Rose) Seigler & Ebinger], which is endemic to the state of Sonora, Mexico, is morphologically somewhat unusual, but is clearly nested within Mariosousa (fide
Héctor M. Hernández22
Citation: Hernández HM (2024) 27. Calliandra clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 358–370. https://doi.org/10.3897/phytokeys.240.101716
Calliandra clade
Figs
Included genera (3). Acaciella Britton & Rose (15 species), Afrocalliandra E.R. Souza & L.P. Queiroz (2), Calliandra Benth. (140).
Description. Shrubs or small trees, rarely perennial herbs or geoxylic subshrubs, the branches often provided with brachyblasts from which leaves and inflorescences arise, usually unarmed but in a few cases armed with long shoots tapering at apex into a stout thorn or with spinescent stipules. Stipules persistent or caducous, rarely spinescent. Leaves bipinnate, once pinnate in a single species; petioles and rachis without extrafloral nectaries; pinnae 1–many pairs; leaflets (1) 2–many pairs per pinna, small (ca. 5 mm) to up to 12 cm long. Inflorescences capitate, umbelliform or sometimes elongated into short racemes, usually pedunculate, solitary or fasciculate, arising from leaf axils or from brachyblasts, sometimes with the capitula or umbels forming long, terminal pseudoracemes through suppression of distal leaves; capitula either homomorphic or heteromorphic. Flowers sessile, subsessile or pedicellate; calyx often cup-shaped, less frequently turbinate, hemispherical or inflated, (3) 5 (6)-merous; corolla cup-shaped to tubular, (3) 5 (6)-merous; stamens (8) 10 to moderately or extremely numerous (up to 300 in Acaciella) in a single flower, exserted or long exserted from the corolla, free from the base or basally fused to form a conspicuous tube, this disproportionately long in the central flowers of heteromorphic capitula, red, pink or white, or white in the basal half and pink or red in the distal half, rarely bright yellow; anthers eglandular; pollen in 8-grained polyads, rarely with 7 or up to 10 grains in Afrocalliandra, the polyads calymmate or acalymmate, with or without a basal grain acutely narrowed and bearing a sticky appendage. Fruits linear, oblanceolate, narrowly oblong or linear-oblanceolate, flattened, straight or slightly curved, with thickened margins, the valves membranous, coriaceous, chartaceous or ligneous, dehiscing passively or elastically along both margins, recurving from apex downwards. Seeds without an aril, discoid, ovoid or rhomboid, usually compressed, with or without U-shaped pleurogram.
Geographic distribution. The Calliandra clade is primarily a Neotropical group, with the exception of Afrocalliandra whose two species are allopatrically distributed in restricted arid areas of Kenya and adjacent Somalia, and in the northern Cape Province of South Africa.
Clade-based definition. The most inclusive crown clade containing Calliandra houstoniana (Mill.) Standl. and Acaciella angustissima (Mill.) Britton, but not Zapoteca caracasana (Jacq.) H.M. Hern., Senegalia bahiensis (Benth.) Seigler & Ebinger or Pithecellobium dulce (Roxb.) Benth. (Fig.
Notes. In the phylogenomic analyses of
The phylogenetic affinity of the three genera (ca. 157 species) of the Calliandra clade is reflected by the presence of several shared morphological characters: predominantly unarmed shrubs or small trees, bipinnate leaves [except C. hymenaeoides (Rich.) Benth.], lack of extrafloral nectaries, 8-grained polyads, dehiscent, flat pods with thickened margins, and exarillate, usually compressed seeds. Acaciella has the most disparate morphological characters compared with the other two genera, especially with respect to flower traits (e.g., lack of staminal tube, short and extremely numerous stamens), and pods with passive, rather than active (elastic) dehiscence.
All species of the clade are unarmed and lack spinescent stipules, except two species of Calliandra [C. haematomma (DC.) Benth. and C. pauciflora (A. Rich.) Griseb.] and Afrocalliandra redacta (J.H. Ross) E.R. Souza & L.P. Queiroz. In addition, Afrocalliandra gilbertii (Thulin & Hunde) E.R. Souza & L.P. Queiroz and some Brazilian species of Calliandra (e.g., C. spinosa Ducke) are armed with tapering branches becoming spinescent at the tips.
Eight-grained polyads are common to all members of the clade, although those of Calliandra are characteristically calymmate, all grains being covered by a common exine. In contrast, the polyads of all species of Acaciella and Afrocalliandra are acalymmate; in Afrocalliandra redacta the number of grains per polyad has been reported to vary from 7 to 10, but 8-grained polyads are the most common (Robbertse and von Teichman 1979). However, it is important to emphasise that Calliandra and Afrocalliandra share a unique combination of polyad characters within Mimoseae. The polyads in these two genera have an ellipsoid, tear-shaped outline displaying a highly modified “basal” grain with an acute apex, this provided with a mucilaginous appendage. This sticky appendage, which is associated with the transfer of polyads by pollinators, is extremely large and conspicuous in Calliandra (see fig. 2 in Hernández 1986), and appears to be rather rudimentary in A. gilbertii (see fig. 3G in
All species, with the exception of the two Afrocalliandra species, occur in tropical and subtropical regions of the Americas. Mexico appears to have been an important region for the evolution of the clade with almost all species of Acaciella occurring in this country (14 of 15 species), and 39 species of Calliandra, 19 of which are endemic. However, Brazil is by far the main centre of distribution of Calliandra, with ca. 71 species, most of them endemic. The group as a whole is absent (Acaciella) or poorly represented (Calliandra) in the Amazon basin. The vast majority of species occur in areas dominated by seasonally dry tropical climates and even in semi-desert regions, and less frequently in more mesic Quercus or Pinus-Quercus forests, grasslands or savannas. Although not known for Afrocalliandra, the two other genera of the Calliandra clade are reported to fix nitrogen with a symbiosome-type nodule anatomy (
Acacia ser. Filicinae Benth., London J. Bot. 1: 322. 1842. Type not designated.
Acacia sect. Filicinae (Benth.) Taub., Nat. Pflanzenfam. 3(3): 113. 1894. Type not designated.
Senegalia sect. Filicinae (Benth.) Pedley, Bot. J. Linn. Soc. 92(3): 238. 1986. Type: Senegalia angustissima (Mill.) Pedley [≡ Mimosa angustissima Mill. (≡ Acaciella angustissima (Mill.) Britton & Rose)]
Acaciella villosa (Sw.) Britton & Rose [≡ Mimosa villosa Sw.]
Shrubs or small trees to 12 m (Fig.
Diversity of plant growth forms of the genera of the Calliandra clade A shrubby habit of Acaciella angustissima var. angustissima, Texas, USA B treelet of Acaciella angustissima var. angustissima, Chiquimula, Guatemala (Hughes 1487) C shrub of Calliandra mollissima (Humb. & Bonpl. ex Willd.) Benth., Marañón Valley, Peru (Särkinen 2198) D stunted xerophytic shrublet of Afrocalliandra redacta (J.H. Ross) E.R. Souza & L.P. Queiroz, Kuboes, South Africa E stunted xerophytic shrublet of Calliandra chilensis Benth., northern Chile F small treelet of Calliandra calothyrsus Meisn., Siguatepeque, Honduras G shrub of Calliandra californica Benth., Baja California Sur, Mexico (Hughes 1546) H treelet of Calliandra fuscipila Harms, Serra do Espinhaço, Bahia, Brazil (Queiroz 15626) I, J functionally herbaceous geoxyle of Calliandra longipes Benth. arising from a stout lignotuber and resprouting and fruiting after fire in savanna woodland, Santa Cruz, Bolivia (Wood 26548) K geoxylic shrublet of Calliandra mucugeana Renvoize, arising from a stout woody underground stem and forming carpet-like thickets, Serra do Espinhaço, Bahia, Brazil (Queiroz 15540). Photo credits A Ron Stephens, iNaturalist (https://www.inaturalist.org/photos/199321699) B, C, F–K CE Hughes D Pietermier, iNaturalist (https://www.inaturalist.org/photos/152398478) E J Jiménez Castillo.
Variation in inflorescences of the genera of the Calliandra clade. A compound terminal panicle of capitula of Acaciella villosa (Sw.) Britton & Rose, Oaxaca, Mexico (Hughes 1333) B capitula of Acaciella villosa, Piura, Peru (Hughes 2635) C flowers of Afrocalliandra redacta (J.H. Ross) E.R. Souza & L.P. Queiroz, Kuboes, South Africa D erect terminal racemose inflorescence of Calliandra calothyrsus Meisn. flowering at night, Siguatepeque, Honduras (Macqueen 3) E erect terminal racemose inflorescence of Calliandra juzepczukii Standl., Oaxaca, Mexico (Hughes 1675) F erect terminal inflorescence with buds and flowers opening acropetally, Calliandra grandiflora (L’Her.) Benth., Mexico City, Mexico G inflorescence of Calliandra taxifolia (Kunth) Benth., Mollendo, Arequipa, Peru (Hughes 2357) H heteromorphic inflorescence of Calliandra mollissima (Humb. & Bonpl. ex Willd) Benth., Marañón Valley, Peru, showing central flowers with enlarged staminal tubes (Särkinen 2198) I Calliandra nebulosa Barneby, Serra do Espinhaço, Bahia, Brazil (Queiroz 15624) J flowers of Calliandra longipinna Benth., Serra do Espinhaço, Bahia, Brazil (Queiroz 15603) K long pedicellate flowers of Calliandra leptopoda Benth., Serra do Espinhaço, Bahia, Brazil L leaves, foliaceous stipules and terminal inflorescences of Calliandra lanata Benth., Serra do Espinhaço, Bahia, Brazil. Photo credits A, B, D, E, G–J CE Hughes C Pietermier, iNaturalist (https://www.inaturalist.org/photos/15308160) F Mvz-juangonzalezromero, iNaturalist (https://www.inaturalist.org/photos/195966667) K, L E de Souza.
Variation in fruits across genera of the Calliandra clade A pendulous fruit of Acaciella angustissima var. angustissima, Chiapas, Mexico B erect fruit and spinescent stipules of Afrocalliandra redacta (J.H. Ross) E.R. Souza & L.P. Queiroz, Kuboes, South Africa C, D fruits of Calliandra taxifolia (Kunth) Benth., Mollendo, Arequipa, Peru (Hughes 2357) C unripe green erect fruits D ripe fruits elastically dehiscent from the apex, the valves recurved backwards E ripe and unripe fruits of Calliandra luetzelburgii Harms, Serra do Espinhaço, Bahia, Brazil (Queiroz 15618) F unripe, green, erect fruits of Calliandra viscidula Benth., Serra do Espinhaço, Bahia, Brazil (Queiroz 15541) G, H fruits of Calliandra houstoniana (Mill.) Standl., Chiapas, Mexico (Hughes 1271 & 1287) I unripe fruits of Calliandra chilensis Benth., northern Chile J unripe, erect fruits of Calliandra bahiana var. erythematosa Barneby, Serra do Espinhaço, Bahia, Brazil (Queiroz 15622). Photo credits A Neptalí Ramírez Marcial, iNaturalist (https://www.inaturalist.org/photos/199788369) B Pietermier, iNaturalist (https://www.inaturalist.org/photos/152398411) C–H, J CE Hughes I J Jiménez Castillo.
2n = 26 [A. texensis (Torr. & A.Gray) Britton & Rose] (
Fifteen species are currently recognised (Rico Arce and Bachman 2006), all restricted to the Neotropics. With the exception of A. glauca (L.) L. Rico, all species of Acaciella occur in Mexico, and 11 of them are endemic to this country. Species have been recorded from south-eastern United States, throughout Mexico and south to Costa Rica and western Panama in Central America, with scattered records in the Antilles (Cuba, Jamaica and Dominican Republic), and South America (Venezuela, Colombia, Ecuador, Peru, Bolivia and Argentina). The genus is absent in the Amazonian basin (Fig.
Sea level to 2500 m elevation, primarily in areas covered by seasonally dry tropical deciduous forests, thorn scrub, semi-desert vegetation, and mixed Quercus-Pinus forest.
Small Acacia, from -ellus (suffix) used to form diminutives.
In its native range, the leaves of A. angustissima are reported to be used as forage for livestock, the roots for tanning, and different parts of the plants are utilised in traditional medicine and for the production of fermented beverages (Rico Arce and Bachman 2006). In addition, due to its high growth-rate and its ability to fix atmospheric nitrogen and accumulate tannins, A. angustissima is being tested as a multi-use, agroforestry species (Rico Arce and Bachman 2006;
Acaciella has traditionally been related to Acacia Mill. s.l., and species were grouped into Acacia ser. Filicinae Benth. (
Despite the resemblance of the flowers of Acaciella to those of Acacia, Rico Arce and Bachman (2006) resurrected Acaciella as a distinct genus, based on a combination of morphological traits, essentially the lack of spines and extrafloral nectaries, and 8-grained polyads (Rico Arce and Bachman 2006). Gómez-Acevedo et al. (2010), using morphological and molecular evidence, established Acaciella [excl. A. chamelensis (L. Rico) L. Rico] as monophyletic and phylogenetically related to Calliandra rather than to Acacia.
Afrocalliandra redacta (J.H. Ross) E.R. Souza & L.P. Queiroz [≡ Acacia redacta J.H. Ross]
Shrubs to 2.5 m, densely branched (Fig.
Unknown.
Two species distributed allopatrically in distant arid areas of Africa. Afrocalliandra redacta is endemic to a restricted area of the Northern Cape Province, across the mountains of the Richtersveld, South Africa, and A. gilbertii is found in the Mandera District, Kenya and in adjacent Somalia (Fig.
Both species of Afrocalliandra occur in arid zones, in areas surrounded by open, shrubby vegetation. Afrocalliandra redacta has been reported to grow on rocky slopes of schistoid granite, at 700 m elevation, whereas A. gilbertii has been found on sandstone ridges at 450–500 m.
African Calliandra, from Greek, calli- (= beautiful) and -andrus (= male), pertaining to the highly attractive stamens.
Unknown.
Based on a nuclear and plastid DNA sequence phylogeny,
Contrasting with the current view to maintain Calliandra and Afrocalliandra as separate genera,
Anneslia Salisb., Parad. Lond.: pl. 64. 1807, nom. ut. rej. vs. Calliandra Benth. Type: Anneslia falcifolia Salisb. [≡ Calliandra houstoniana (Mill.) Standl.]
Clelia Casar., Nov. Stirp. Brasil. Dec. 10: 83. 1845. Type: Clelia ornata Casar. [≡ Calliandra harrisii (Lindl.) Benth.]
Codonandra H. Karst., Fl. Columb. 2: 43, t. 122. 1863. Type: Codonandra purpurea H. Karst. [≡ Calliandra magdalenae var. magdalenae]
Guinetia L. Rico & M. Sousa, Kew Bull. 54(4): 977. 1999 (publ. 2000). Type: Guinetia tehuantepecensis L. Rico & M. Sousa [≡ Calliandra tehuantepecensis (L. Rico & M. Sousa) E.R. Souza & L.P. Queiroz]
Calliandra houstoniana (Mill.) Standl. [≡ Mimosa houstoniana Mill.]
Shrubs or small trees (Fig.
n = 8, 11 (
Approximately 140 species restricted to the Neotropics, occurring from southern United States (southern Arizona, New Mexico and Texas), throughout Mexico, Central America and South America, south to Uruguay, and northern Argentina and Chile (Fig.
Most species occur at low or moderate elevations, in seasonally dry tropical deciduous forests, and less frequently in wet forests, riparian vegetation, tropical savannas, campos rupestres (rupestrian grasslands) from Espinhaço range in eastern Brazil, temperate grasslands, and even in semi-desert or desert vegetation. Seeds disperse over short distances through the explosive dehiscing mechanism. Most Mesoamerican and Mexican species have nocturnal anthesis and are legitimately pollinated by sphingid moths or bats; however, hummingbird pollination in day-flowering species appears to be also common in the genus.
From Greek, calli- (= beautiful) and -andrus (= male), pertaining to the highly attractive stamens.
Calliandra calothyrsus Meissn. has proved to be an excellent fast-growing, multi-use shrub or small tree as a source of fuelwood and fodder, as well as for soil improvement (
For over a century, after publication of
Phylogenetic analyses using morphological and molecular sequence data have shown that all the American species of Calliandra, plus the monotypic Guinetia L. Rico & M. Sousa, later transferred to Calliandra [C. tehuantepecensis (L. Rico & M. Sousa) E.R. Souza & L.P. Queiroz], constitute a monophyletic group (
The generic placement of one Asian species, C. umbrosa (Wall.) Benth. from India, Bangladesh and Myanmar, which is still included in Calliandra, remains to be evaluated
Generic relationships in the Zapoteca clade (tribe Mimoseae). Note that Calliandra p.p. represents a single taxon, which is currently unplaced [and is not the type species of Calliandra (see tribe notes and footnote on page 382)]. For description of phylogeny and support values, see Fig.
Héctor M. Hernández22
Citation: Hernández HM (2024) 28. Zapoteca clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 371–383. https://doi.org/10.3897/phytokeys.240.101716
Zapoteca clade
Figs
Included genera (5). Faidherbia A. Chev. (1 species), Sanjappa E.R. Souza & M.V. Krishnaraj (1), Thailentadopsis Kosterm. (3), Viguieranthus Villiers (18), Zapoteca H.M. Hern. (22).
Description. Shrubs or trees, armed with spinescent stipules or unarmed. Stipules leafy or spinescent, persistent or caducous. Leaves bipinnate, rarely pinnate, with or without extrafloral nectaries on the petiole, leaf rachis or pinnae; pinnae 1–numerous pairs; leaflets 1–numerous pairs per pinna, opposite or rarely alternate, small (ca. 5 mm) to up to 22 cm long. Inflorescences spherical heads, umbels, or elongated spikes or racemes, pedunculate, usually densely-flowered, solitary or fasciculate, arising from leaf axils, sometimes forming long, terminal pseudopanicles, the capitula always homomorphic. Flowers hermaphroditic or functionally staminate; calyx cup-shaped, less frequently campanulate, obovoid, obconical or hemispherical, (3–4) 5-lobed; corolla campanulate or infundibuliform, (3–4) 5-lobed; stamens numerous, long exserted from the corolla, to 4.3 cm long, always with the base fused forming a conspicuous tube, this rarely as short as 1 mm, the staminal tube always included in the corolla, usually white, rarely pink or red-purple, or white in the basal half and pink or red in the distal half, anthers eglandular; pollen in 16-grained polyads (32-grained in Faidherbia), always acalymmate, discoid, heteromorphic; ovary sessile or shortly stipitate, style filiform, the stigma cup-shaped, capitate or discoid. Fruits linear to elliptic, oblanceolate or obovate, flattened or thickened, straight, slightly curved or curled and twisted, with or without thickened margins, the valves membranous, chartaceous, coriaceous or ligneous, brownish or orange when mature, usually but not always dehiscent elastically along both margins, recurving from apex downwards, or indehiscent. Seeds ovoid, oblong or rhomboid, with or without a pleurogram.
Geographic distribution. The Zapoteca clade displays a pantropical distribution, with Zapoteca restricted to tropical and subtropical America, Viguieranthus endemic to Madagascar and the Comoro Islands, Faidherbia widespread throughout Africa, Thailentadopsis restricted to parts of Indochina (Thailand and Vietnam) and Sri Lanka, and Sanjappa limited to south-west India.
Clade-based definition. The most inclusive crown clade containing Zapoteca caracasana (Jacq.) H.M. Hern. and Sanjappa cynometroides (Bedd.) E.R. Souza & Krishnaraj, but not Calliandra houstoniana (Mill.) Standl., Hesperalbizia occidentalis (Brandegee) Barneby & J.W. Grimes or Pithecellobium dulce (Roxb.) Benth. (Fig.
Notes. The Zapoteca clade comprises five genera with 45 known species. Faidherbia, Thailentadopsis, Sanjappa and Viguieranthus were shown to form a monophyletic group in phylogenetic analyses of Ingeae (
The individual genera may be easily distinguished. Three genera (Faidherbia, Thailentadopsis and Sanjappa) are armed with paired, persistent or caducous, straight, lignescent stipules, whereas Zapoteca (excl. Z. aculeata Spencer ex Benth.) and Viguieranthus are unarmed. The leaves are usually bipinnate, and have a single pair of pinnae in most species of Viguieranthus, 1–2 pairs in Thailentadopsis, and 1–10 pairs in Zapoteca and Faidherbia. In contrast, the leaves of Sanjappa and two species of Viguieranthus (V. brevipennatus Villiers and V. unifoliolatus Villiers) may be interpreted as pinnate, and are reduced to a single pair of leaflets per leaf; the highly reduced pinnate leaves of these two genera may have been derived from bipinnate-leaved ancestors. The number of leaflets per pinna is also highly variable, and leaflet size is usually inversely proportional to the number of leaflet pairs per pinna.
Fruits of Zapoteca and Sanjappa are essentially flat and straight, have thickened margins, and are elastically dehiscent, a dehiscing mechanism typical of Calliandra species, for example. The fruits of Viguieranthus are morphologically similar to those of the two previous genera, but the dehiscing mechanism appears to be passive, rather than active (elastically dehiscent). The submoniliform fruits of Thailentadopsis are dehiscent, but also lack the explosive mechanism of Zapoteca and Sanjappa. Contrasting with the previous four genera, the fruits of Faidherbia are indehiscent and markedly different morphologically. In fact, Faidherbia, when compared with the other genera in the clade, has the most divergent morphological features (i.e., large trees, spicate inflorescences, short staminal tube, 32-grained polyads, thick, twisted, indehiscent fruits), although sharing the straight stipular spines with Thailentadopsis and Sanjappa.
As noted in the Calliandra clade (page 358), two Asian species of Calliandra, C. umbrosa (Wall.) Benth. and a yet undescribed species, are the last remaining of the species excluded from Calliandra by
Calliandra ser. Laetevirentes Benth., London J. Bot. 3: 97. 1844. Type not designated.
Zapoteca tetragona (Willd.) H.M. Hern. [≡ Acacia tetragona Willd.]
Erect, suberect or scandent shrubs, rarely small trees, usually unarmed, very rarely armed with spinescent stipules. Stipules leafy or rarely spinescent, usually persistent. Leaves bipinnate, usually without extrafloral nectaries, but 3 (4) species with cup-shaped or cylindrical glands on the rachis or pinnae or near the base of the petiole; pinnae 1–numerous pairs; leaflets (1) 2–numerous pairs per pinna, opposite, small (ca. 5 mm) to up to 22 cm long. Inflorescences capitate, spherical in bud and at anthesis, homomorphic, pedunculate, densely-flowered, solitary or fasciculate, arising from leaf axils, sometimes forming long, terminal pseudopanicles. Flowers bracteate, hermaphroditic or functionally staminate; calyx cup-shaped, dentate or denticulate; corolla campanulate or infundibuliform, the petals valvate in bud, usually revolute at anthesis; stamens ca. 30–60, long exserted from the corolla, 1.9–4.3 cm, always with the base fused forming a conspicuous tube, the staminal tube always included in the corolla, white, pink or red-purple, or white in the basal half and pink or red in the distal half, anthers eglandular; pollen in 16-grained polyads, acalymmate, discoid, heteromorphic, nearly always with eccentric lens-shaped thickenings on the central cells on one side of the polyad; ovary sessile or shortly stipitate, usually 10–15-ovulate, style filiform, with the stigma cup-shaped. Fruits usually pendent, linear, flattened, straight or rarely slightly curved, with thickened margins, the valves membranous or rarely coriaceous, dehiscing elastically along both margins, recurving from apex downwards. Seeds ovoid or rhomboid, usually with irregular or U-shaped pleurogram.
Morphological features of the Zapoteca clade A, B Faidherbia albida (Delile) A. Chev. A habit B leaves C, D Sanjappa cynometroides (Bedd.) E.R. Souza & M.V. Krishnaraj C habit D pinnate leaves and inflorescence in bud E Zapoteca aculeata (Spruce ex Benth.) H.M. Hern. foliage and inflorescences at anthesis (Neill 18437) F Zapoteca formosa subsp. schottii (Torr. ex S. Watson) H.M. Hern. habit and dehisced fruits G Viguieranthus glaber Villiers leaves and inflorescences post-anthesis. Photo credits A Gobabeb - Namib Research Institute B Dave U C, D G Suresh E D Neill F D Beckman G C Davidson.
Inflorescence and fruit morphology in the Zapoteca clade A, B Faidherbia albida (Delile) A. Chev. A inflorescences at anthesis B indehiscent fruit C, D Sanjappa cynometroides (Bedd.) E.R. Souza & M.V. Krishnaraj C inflorescence post-anthesis D pinnate leaves and immature fruit E Thailentadopsis tenuis (Craib) Kosterm. inflorescence F Viguieranthus perrieri (R. Vig.) Villiers inflorescence at anthesis G Viguieranthus alternans (Benth.) Villiers foliage and fruits before dehiscence H Viguieranthus brevipennatus Villiers inflorescence at anthesis (Randrianarivony 175) I Zapoteca tetragona (Willd.) H.M. Hern. inflorescence at anthesis J Zapoteca portoricencis subsp. portoricensis inflorescence at anthesis (Amith 74438) K Zapoteca formosa subsp. schottii (Torr. ex S. Watson) H.M. Hern. dehisced fruit. Photo credits A S Piry B B Adkins C, D G Suresh E https://marketingoemoffice.com/Data_Pithecellobium.html F F Ratovoson G F Rakotoarivony H T Randrianarivony I FJ Gómez Marín J J Amith K D Beckman.
n = 13 (Hernández 1986,
Twenty-two described species, all of which are restricted to the Neotropics. Its distribution extends from southern United States (southern Arizona and Texas), throughout Mexico, Central America and South America, south to Paraguay and northern Argentina, including the West Indies and the Revillagigedo Islands (Fig.
Populations occur from sea-level to 2850 m, although most species are found at low or moderate elevations, in areas covered by seasonally dry tropical deciduous forests, and less frequently in wet forests, and even in desert or semi-desert vegetation. Seed dispersal covers short distances through the explosive dehiscing mechanism. Settling moths of the families Noctuidae, Pyralidae and Geometridae are confirmed to be the primary pollinators (
Named after the Zapotec ethnic group of Oaxaca, Mexico, where a significant number of taxa occur.
No significant uses are known, although Z. formosa subsp. formosa is reported to be potentially important as a forage resource for sheep (
Detailed morphological and cytological observations supported the recognition of Zapoteca as a genus distinct from Calliandra by consistently having spherical, homomorphic, capituliform inflorescences; 16-grained, acalymmate, discoid polyads, with lens-shaped thickenings in the central grains [excluding Z. nervosa (Urban) H.M. Hern.]; cup-shaped stigmas; and a basic chromosome number of x = 13. In contrast, the Neotropical species of Calliandra are characterised by having spherical, obconic, capituliform, umbelliform or racemose, homomorphic or heteromorphic inflorescences; capitate or fungiform stigmas; 8-grained, bisymmetric, calymmate polyads with a mucilaginous basal appendage; and atypical (n = 8 and 11, rather than the x = 13 of closely related genera) chromosome numbers (
The pattern of differentiation within Calliandra s.l. sensu Bentam (1875) provided the basis for the segregation of the species in ser. Laetevirentes, together with two other species placed by Bentham in his ser. Macrophyllae (C. aculeata and C. amazonica Benth.), into the new genus Zapoteca (Hernández 1986,
Viguieranthus alternans (Benth.) Villiers [≡ Calliandra alternans Benth.]
Unarmed shrubs or trees to 25 m, with trunks up to 80 cm diameter. Stipules coriaceous, not spinescent, more or less persistent. Leaves bipinnate, rarely pinnate; petioles with an extrafloral nectary at the apex, sometimes also present on the rachis tips, sometimes the petioles and rachis with narrowly winged margins; pinnae of bipinnate leaves 1 pair; leaflets alternate to opposite, sometimes 1 or 3 per pinna, 1 pair per leaf in pinnate leaves, up to 7.7 × 4.6 cm, but frequently more numerous (to 34 per pinna), and smaller (9–35 × 2–15 mm). Inflorescences spherical heads, elongate spikes or racemes, pedunculate, solitary or fasciculate, arising from leaf axils, sometimes organised in pseudopanicles, homomorphic. Flowers hermaphrodite; calyx usually cup-shaped, less frequently broadly obovoid, obconical or hemispherical, 4–5-lobed; corolla obconical, with the petals connate into a tube, 4–5-lobed; stamens numerous, long exserted from the corolla, white, always with the base fused forming a tube inserted in the corolla, anthers eglandular; pollen in 16-grained polyads, acalymmate, discoid, heteromorphic, without eccentric lens-shaped thickenings on the central cells; ovary sessile or short-pedicellate; stigma funnel-shaped or capitate. Fruits linear-oblanceolate, linear-elliptic, or linear-obovate to narrowly obovate, flattened, straight or slightly curved, with thickened margins, the valves chartaceous, coriaceous or ligneous, dehiscing along both margins. Seeds ovoid, oblong or rhomboid, without a pleurogram.
Unknown.
Most species are reported from the eastern, humid, evergreen forests in Madagascar. However, some species grow in areas covered by dry deciduous woodland or xerophytic scrubland. Populations are reported from sea level to 1600 m (
Named after René Viguier (1880–1931), French botanist.
The wood of some species is reported to be used as firewood, and for house construction and joinery (
In the original description of Viguieranthus, published in The Leguminosae of Madagascar one year after J.F. Villiers’ death (
In light of the previous considerations, Viguieranthus is a Madagascan genus, characterised by constantly having leafy (not spinescent) stipules; bipinnate leaves with only 1 pair of pinnae; a variable number of opposite or alternate pairs of leaflets per pinna, sometimes reduced to one leaflet per pinna (V. unifoliolatus Villiers and V. brevipennatus Villiers); and homomorphic inflorescences (
Faidherbia albida (Dalile) A. Chev. [≡ Acacia albida Dalile]
Trees to 30 m, with trunks up to 2 m diameter, the spreading branches forming a rounded crown, armed with spinescent stipules. Stipules spinescent, paired, persistent, straight or slightly curved, to 2 (3.3) cm long. Leaves bipinnate, cup-shaped extrafloral nectaries on the leaf rachis at the junction of each pair of pinnae, absent on the petiole; pinnae (2) 3–10 pairs; leaflets 6–23 pairs per pinna, opposite, small (3.5–9 mm long), linear or linear-oblong to slightly obovate-oblong. Inflorescences spicate, pedunculate, solitary, arising from leaf axils, collectively forming a terminal pseudopanicle. Flowers hermaphrodite; calyx cup-shaped, 5-lobed; corolla campanulate, pale pink inside, the 5 lobes divided almost to the base; stamens 35–55, long exserted from the corolla, with the base fused for ± 1 mm forming a short tube, yellowish-white to pale cream, anthers eglandular; pollen organised in 32-grained polyads, these acalymmate, subcircular, flattened; ovary short-stipitate. Fruits thick, flattened, falcate or conspicuously curled and twisted, bright orange when mature, indehiscent. Seeds elliptic-lenticular, compressed, with an elliptic pleurogram.
2n = 26 (
Monospecific (F. albida), widespread in Africa, especially across eastern Africa, from Egypt southwards into Sudan, Ethiopia, Somalia, Kenya, Uganda, Tanzania, Mozambique, Zambia and Zimbabwe, to the Transvaal, South Africa. Also reported from Mauritania, Senegal and The Gambia, and in Angola southwards to Botswana (Fig.
Found predominantly in seasonally dry tropical climates, in forests, woodlands and especially in savannas where it is often abundant, on alluvial soils, along seasonal watercourses and around lakes and swamps, from near sea level to 2600 m elevation. It shows an unusual reverse phenology being characteristically leafless during the rainy season. Faidherbia albida is reported to have a rather generalist reproductive system (
Named after the French Major Louis Léon César Faidherbe (1818–1889), Governor of Senegal and founder of Dakar.
The trunks are used to make canoes, and the wood is employed as firewood, and for handicrafts; various medicinal uses are reported. The leaves and fruits are used as fodder for domestic animals, and are also consumed by camels and elephants (
Originally described as Acacia albida Dalile, this species has several unusual characters as compared with the other African species of Acacia Mill., such as the lack of petiolar extrafloral nectaries, these being present only at the junction of each pair of pinnae, the basally connate filaments forming a short tube (in line with most members of the ingoid clade and all members of the Zapoteca clade), the eglandular anthers, and the 32-grained polyads (
Thailentadopsis tenuis (Craib) Kosterm. [≡ Pithecellobium tenue Craib]
Shrubs or small trees up to 7 m tall, the trunk to 15 cm diameter, armed with spinescent stipules. Stipules spinescent, paired, persistent, lignescent. Leaves bipinnate; stalked extrafloral nectaries between each pair of pinnae and usually between each pair of leaflets; petioles, leaf and pinnae rachides winged or not; pinnae 1–2 pairs; leaflets 1–6 pairs per pinna, opposite, increasing in size from base to apex, small to medium sized. Inflorescences capitate or umbellate, pedunculate, few-flowered, fasciculate, arising from leaf axils, sometimes forming terminal pseudopanicles, homomorphic. Flowers hermaphroditic; calyx campanulate; corolla campanulate or infundibuliform; stamens numerous, long exserted from the corolla, always with the base fused forming a conspicuous tube, the staminal tube always included in and equal in length to the corolla, white, anthers eglandular; pollen in 16-grained polyads, acalymmate, discoid, heteromorphic. Fruits submoniliform, flattened, usually with the interseminal spaces constricted, straight or markedly curved, with thickened margins, the valves coriaceous, dehiscent. Seeds ovoid, with ovate pleurogram.
Unknown.
Populations are reported from 100 to 900 m elevation, in evergreen and open savanna forest, in shaded areas by streams or along rivers, on soils derived from limestone or granitic bedrock.
From Thailand (home of the type species), Entada (a mimosoid genus, for the superficial similarity of the fruits), and -opsis (Greek = similar to).
Unknown.
The three species currently recognised within Thailentadopsis have been placed in different genera, including Acacia, Painteria Britton & Rose and Pithecellobium Mart., and
Sanjappa cynometroides (Bedd.) E.R. Souza & M.V. Krishnaraj [≡ Calliandra cynometroides Bedd.]
Small trees to 6 m, the trunk to 35 cm diameter, armed with spinescent stipules. Stipules spinescent, paired, unequal, straight, lignescent, caducous. Leaves pinnate; extrafloral nectaries circular, slightly raised, at the tip of petiole near the junction of the leaflets; leaflets 1 pair per leaf, opposite, relatively large (ca. 5–12 cm long). Inflorescences capitate, pedunculate, 7–15-flowered, solitary, arising from leaf axils, homomorphic. Flowers hermaphroditic; calyx cup-shaped, dentate, 3-lobed; corolla tubular-campanulate, the 3 lobes usually revolute at anthesis; stamens long-exserted from the corolla, to 2.4 cm long, with their bases fused forming a conspicuous tube, the staminal tube always included in the corolla, white, anthers eglandular; pollen in 16-grained polyads, acalymmate, discoid, heteromorphic; ovary sessile, 5–7-ovulate, style filiform, with the stigma discoid. Fruits oblanceolate, flattened, straight or slightly undulate marginally, with thickened margins, rigidly coriaceous, dehiscing elastically along both margins, recurving from apex downwards. Seeds rhomboid, with a rhomboid pleurogram.
Unknown.
Sanjappa cynometroides has been found growing in evergreen and sub-evergreen forest areas, near streams, at 300–1100 m elevation. Individuals are reported to occur at very low densities in natural populations (
Named after Dr. Munivenkatappa Sanjappa, Senior Scientist at the Botanical Garden, University of Agricultural Sciences, Bengaluru, India.
Unknown.
Sanjappa was described to accommodate one Old World species of Calliandra sensu
Sanjappa is nested with high support in a clade with Faidherbia and Thailentadopsis, characterised by the conspicuous spinescent stipules (
Rodrigo Duno de Stefano13
Citation: Duno de Stefano R (2024) 29. Cojoba clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 384–390. https://doi.org/10.3897/phytokeys.240.101716
Cojoba clade
Figs
Included genera (3). Cojoba Britton & Rose (13–19 species), Hesperalbizia Barneby & J.W. Grimes (1), Lysiloma Benth. (8).
Description. Unarmed trees and arborescent shrubs, either micro- or macrophyllidious. Stipules small, ephemeral, caducous, rarely persistent or lignescent. Leaves bipinnate, rarely paripinnate; extrafloral nectaries sessile; leaflet venation simple, weakly pinnate, pinnate, or palmate-pinnate. Inflorescences capitula arising either singly or fasciculate, spherical, receptacle subglobose, rarely spicate-racemose. Flowers sessile, homomorphic or nearly so, the central flower sometimes a little stouter, but not longer than the others, 5-merous; calyx campanulate, 5-veined or nearly without veins, very shortly toothed; corolla tubular or very narrowly trumpet-shaped; internally lacking callous and nectar disk; stamens numerous, joined at the base, exserted from corolla; ovary sessile or subsessile, linear-ellipsoid. Fruits variable, more or less pendulous, generally linear to broad-linear, plano-compressed, or cylindrical, straight, retrofalcate or spirally contorted, scarcely or deeply constricted between seeds, dehiscent through one or both sutures, but craspedial or indehiscent in Lysiloma, and framed by prominent sutures, and the exocarp exfoliating, valves fleshy, leathery, stiffly papery to papery-crustaceous. Seeds variable, compressed to flattened, subglobose, discoid, oblong-ellipsoid to ellipsoid.
Distribution. Neotropics: Mexico, Central America, South America, and Greater Antilles.
Clade based definition. The most inclusive crown clade containing Hesperalbizia occidentalis (Brandegee) Barneby & J.W. Grimes and Cojoba arborea (L.) Britton & Rose, but not Zapoteca caracasana (Jacq.) H.M. Hern., Pithecellobium dulce (Roxb.) Benth. or Calliandra houstoniana (Mill.) Standl. (Fig.
Notes. The Cojoba clade was first recognised in the phylogenetic analyses of Iganci et al. (2015), and the relationships were later supported in the analyses of
Within the Cojoba clade, Lysiloma and Hesperalbizia are supported as sister genera which share plano-compressed fruits, as opposed to the cylindrical fruits of Cojoba, and these two sister genera are apparently genetically weakly divergent as evidenced by lack of robust support in species-level phylogenies (
Hesperalbizia occidentalis (Brandegee) Barneby & J.W. Grimes [≡ Albizia occidentalis Brandegee]
Unarmed trees 20 (30) m, bark pale silvery grey. Stipules small, membranous, caducous, cordate at base. Leaves bipinnate; extrafloral nectaries below mid-petiole, sometimes close to the base, sessile, round or elliptical, shallowly concave, commonly smaller ones at the tip of the petiole and the rachis of the pinnae; pinnae 3–8 pairs; leaflets 5–10 pairs per pinna, venation pinnate. Inflorescence 5–20-flowered capitula, receptacle subglobose, spherical, arising either singly or fasciculate. Flowers sessile, homomorphic, 5-merous; calyx campanulate, lobes ovate or short-deltate; corolla narrowly vase-shaped; stamens 52–76, 20–24 mm long, joined at base into a well exserted, 5–11 mm long tube; pollen in 16-celled polyads, more or less isodiametric; ovary linear-elliptic, glabrous, stipitate, style shortly exserted, slightly dilated at tip. Fruits solitary or paired, broad-linear plano-compressed, long-stipitate, 8–12 (13)-seeded, inertly dehiscent through both sutures. Seeds transverse, compressed discoid or oblong-ellipsoid, areolate, funicle filiform; testa crustaceous, brown, and lustrous; pleurogram closed.
Morphology of Cojoba clade A–C Cojoba arborea (L.) Britton & Rose A habit B leaves C fruit D Cojoba graciliflora (S.F. Blake) Britton & Rose in Mexico, fruit E–G Hesperalbizia occidentalis (Brandegee) Barneby & J.W. Grimes E habit F trunk G leaves and fruits H Lysiloma candidum Brandegee, habit l Lysiloma acapulcense (Kunth) Benth. fruit J–N Lysiloma latisiliquum (L.) Benth. J habit K trunk L leaves and stipule M flowers N fruit O, P Lysiloma tergeminum Benth., fruits. Photo credits A–C GA Romero González D, J–N R Duno de Stefano E–I, O, P CE Hughes.
2n = 26 (Rico Arce 1992).
Largely confined to seasonally dry tropical deciduous forests and woodlands, from sea level to 1500 m elevation, locally abundant in tropical and marginally subtropical western Mexico. Strongly deciduous, fruits long-retained on leafless trees, flowering preceding or coinciding with new leaf flush at the end of the dry season. Seed dispersal passive.
Greek = hesperos, the evening star, and by extension the West, plus the generic name Albizia in reference to the westerly distribution of H. occidentalis within Mexico and its previous placement in the genus Albizia.
The plano-compressed, stiffly papery fruits of H. occidentalis are very similar to the fruits of some species of Pseudalbizzia and Albizia, from which Hesperalbizia was segregated by
Lysiloma schiedeanum Benth. [= Lysiloma divaricatum (Jacq.) J.F. Macbr.]
Unarmed small trees or shrubs. Stipules commonly submembranous, linear to obliquely dilated, semicordate, or flabellate, absent from all fruiting and most mature flowering specimens. Leaves bipinnate, extrafloral nectaries near or well above mid-petiole, sessile, either cupular or mounded, narrow-pored, and smaller nectaries often between some distal pinnae-pairs and toward tips of pinnae; pinnae 1–30 pairs; leaflets 1.5–50 pairs per pinna, venation simple, or weakly pinnate, or palmate-pinnate. Inflorescence either spicate-racemose, or spherical capitate, or hemispherical umbelliform. Flowers homomorphic or almost so, the central flower sometimes a little stouter, but never longer than the others; calyx campanulate, short-toothed; corolla tubular; stamens 12–32, joined into a tube at the base, exserted; pollen in 16-grained polyads, more or less isodiametric; ovary sessile or almost so, glabrous at anthesis. Fruit either a craspedium or an indehiscent legume, long persistent on tree, stipitate, in profile linear or oblong-elliptic, acute or obtuse, framed by prominent sutures, the papery-crustaceous valves plano-compressed except where crumpled or low-bullate over seeds, straight but sometimes twisted through 180° either above or below middle, the exocarp exfoliating to reveal the stramineous endocarp, less often not exfoliating in craspedial species, the valves separating from the persistent sutures and falling together out of the marginal frame, the seeds then released on the ground. Seeds transverse, compressed ellipsoid, funicle filiform, lustrous brown, pleurogram commonly U-shaped, sometimes semicircular, or closed.
2n = 26 (
Eight species, in Mexico, Central America as far south as Costa Rica, and the Greater Antilles, one species reaching subtropical south Florida, and one species extending north into south-eastern Arizona in the USA (Fig.
Predominantly seasonally dry tropical forests and adjacent arid and semi-arid thorn scrub in north-western Mexico and the southern USA, extending into mid-elevation pine-oak formations and weakly into wetter less seasonal lowland tropical rainforest. Generally on freely-drained soils, coastal sand dunes and shallow limestone soils. Most species are deciduous, flowering often coinciding with leaf flush at the end of the dry season. Fruits are often long persistent on the tree.
Derived from the Greek, Lysis (= to loose) and loma (= edge), referring to the shedding of the legume sutural frame (or edge) at fruit maturity (i.e., craspedial dehiscence).
The carpological character that distinguishes most Lysiloma species from other New World ingoid genera is the craspedial fruit with sutures which usually persist as a frame after the valves have separated (
Pithecellobium sect. Cojoba (Britton & Rose) Mohlenbr., Reinwardtia 6: 446. 1963. Type: Pithecellobium arboreum (L.) Urb. [≡ Mimosa arborea L. (≡ Cojoba arborea (L.) Britton & Rose)]
Obolinga Barneby, Brittonia 41: 170. 1989. Type: Obolinga zanonii Barneby [≡ Cojoba zanonii (Barneby) Barneby & Grimes]
Cojoba arborea (L.) Britton & Rose [≡ Mimosa arborea L.]
Unarmed medium-sized to sometimes large canopy-emergent trees to 60 m tall and 1 m stem diameter, and arborescent shrubs, either micro- or macrophyllidious. Stipules small, ephemeral, or obsolescent, rarely persistent, sometimes lignescent. Leaves bipinnate, in one species [C. rufescens (Benth.) Britton & Rose] paripinnate, extrafloral nectaries sessile, cupular thick-rimmed on petiole at, or close, below pinnae-pairs, exceptionally 1–2 on petiole; pinnae 1–22 pairs; leaflets 2–50 pairs per pinna, variable in shape, sometimes rhombic, asymmetric, venation palmate, pinnate, or indistinct. Inflorescence spherical capitula, receptacle subglobose, arising either singly or fasciculate. Flowers sessile, homomorphic, 5-merous; calyx campanulate or deeply campanulate, 5-veins or almost veinless, very shortly toothed; corolla tubular or very narrowly trumpet-shaped; stamens 20–66, joined at the base, shortly exceeding or as long as the corolla; pollen in 16-grained polyads, isodiametric; intrastaminal disc absent; ovary sessile or subsessile, ellipsoid to linear-ellipsoid. Fruits dehiscence through one or both margins, pendulous, linear or broad-linear in profile and simply retrofalcate or randomly to spirally contorted, shallowly to deeply constricted between seeds (moniliform), the sutures immersed; valves leathery-fleshy and glossy red when fresh, after dehiscence withering brown, crumpling and contracting to expose the seeds; endocarp smooth tan internally. Seeds ellipsoid to subglobose, funicle straight or simply bent, subfiliform and wiry, or narrowly dilated and stiff after dehiscence, black or dark brown, lacking pleurogram.
Unknown.
Twelve species (
Most species of Cojoba are adapted to wet tropical lowland or submontane to montane evergreen (cloud) forest, and riparian forest, up to ca. 2600 m elevation, but the four West Indian species grow in semi-deciduous lowland scrub, woodland, or chaparral. Flowering apparently at any time of year and in some species more or less continuously so. Seeds likely bird-dispersed.
Cojoba is a vernacular name for C. arborea in Puerto Rico.
The current concept of the genus Cojoba (
Rodrigo Duno de Stefano13
Citation: Duno de Stefano R (2024) 30. Pithecellobium clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 391–403. https://doi.org/10.3897/phytokeys.240.101716
Pithecellobium clade
Figs
Included genera (7). Ebenopsis Britton & Rose (3 species), Gretheria Duno & Torke (2), Havardia Small (3), Painteria Britton & Rose (2), Pithecellobium Mart. (19), Ricoa Duno & Torke (1), Sphinga Barneby & J.W. Grimes (3).
Description. Small trees, shrubs, sometimes sarmentose, growth sympodial or monopodial; branches armed with stipular spines, rarely absent, proleptic, dimorphic, with vegetative and reproductive brachyblasts; buds protected by adaxial side of petiole. Stipules spinescent. Leaves bipinnate, coeval or late suppressed; extrafloral nectaries cupular, either sessile or shortly stipitate, below mid-petiole or below proximal pinna-pair, between each pinna pair, at tip of all pinnae; leaflets macrophyllous or microphyllous, opposite, rarely alternate, venation pinnate, reticulate, subpalmate, palmate, or simple. Inflorescences spikes or globose capitula that arise singly or in fascicles. Flowers uniform; sepals 5, rarely 4 or 6; petals 5, rarely 4 or 6, connate, equal, radially symmetrical; stamens numerous, connate at the base forming a short tube, as long as the corolla or long exerted, anthers elliptic in outline, distinctly wider than long, dehiscence longitudinal; pollen in 16-grained polyads, (more or less) isodiametric; intrastaminal disc present or poorly developed; ovary sessile or shorty stipitate, body compressed-ellipsoid, style about as long as androecium, the stigma poriform. Fruits oblong or broad-linear in profile, plano-compressed turgid, recurved or coiled and sometimes also twisted, in some species cylindrically terete; valves stiffly papery, chartaceous, thinly coriaceous, fleshy, leathery, or woody, cavity continuous or septate, dehiscence inert, through both sutures; funicle straight, sinuous, sigmoid or not, spongy arilliform, red, pink, or whitish. Seeds lentiform or plumply obese, with pleurogram.
Distribution. The Pithecellobium clade is restricted to the New World and strongly centred in Mexico (two genera endemic there and two others nearly so), more sparsely extending to Central America, northern South America (two genera) and the Caribbean (two genera). Mainly below 1000 m elevation but two genera and three species in the highlands of Mexico between 1400–2360 m, predominantly in seasonally dry and arid thickets and forests, less frequent in coastal vegetation and humid forests.
Clade based definition. The most inclusive crown clade containing Pithecellobium dulce (Roxb.) Benth. and Sphinga platyloba (DC.) Barneby & Grimes, but not Cojoba arborea (L.) Britton & Rose, Zapoteca caracasana (Jacq.) H.M. Hern. or Cedrelinga cateniformis (Ducke) Ducke (Fig.
Notes. The Pithecellobium clade was first recognised as the informal Pithecellobium alliance of tribe Ingeae by
The Pithecellobium clade is characterised by stipular spines and brachyblasts that are either strictly vegetative or vegetative and reproductive. Although stipular spines occur in seven ingoid genera worldwide, in the New World the only other ingoid clade taxa known to have stipular spines are Zapoteca aculeata (Spruce ex Benth.) H.M. Hern. and Calliandra pauciflora (A. Rich.) Griseb. but Zapoteca and Calliandra differ by having fruits with elastic dehiscence from the apex and in their pollen morphology (e.g., pollen 16-grained vs. 8-grained; Hernández 1986). Some other ingoid clade taxa are armed with spiny peduncles [some species of Chloroleucon (Benth.) Britton & Rose], or prickles [Albizia corniculata (Lour.) Druce, A. myriophylla Benth., A. umbellata (Vahl) E.J.M. Koenen, and Senegalia species], but none have stipular spines.
In
Spiroloba Raf., Sylva Tellur.: 119. 1838. Type not designated.
Pithecellobium unguis-cati (L.) Benth. [≡ Mimosa unguis-cati L.]
Shrubs and trees, 1–15 m. Stipules spinescent, sometimes absent on some flowering branches, minute or obsolete in P. keyense Britton. Leaves bipinnate, the rachis occasionally winged, extrafloral nectaries cupular, either sessile or shortly stipitate, between each pair of pinna, and also at tip of all pinnae; pinnae 1–2 (11) pairs; leaflets 4–16 (23) pairs per pinna, variously obovate, elliptic, oblong, obovate and suborbicular, venation pinnate and usually also reticulate. Inflorescence units either capitula or relatively short dense spikes, either axillary to the leaves, on efoliate brachyblasts, or paniculately pseudoracemose. Flowers sessile, homomorphic, 5-merous; calyx hemispherical, campanulate or sub-cylindrical, shortly toothed; corolla tubular or trumpet-shaped, rarely turbinate, lobes erect or reflexed; stamens 16–76, the tube as long or longer than corolla; intrastaminal disc with small callosities, rarely developed into lobes; pollen in 16-grained polyads, isodiametric; ovary either oblong or ellipsoid, either sessile or long-stipitate. Fruits oblong or linear pods, reflexed or coiled and sometimes also twisted, the shallowly undulate or evenly curved sutures broad but not prominent, valves fleshy leathery or woody, red or fuscous, biconvex over seeds, the cavity continuous, incipiently septate or compartmentalised into locules; passively or elastically dehiscent either through both sutures or through the ventral suture only. Seeds following dehiscence, dangling on and invested by a red, pink, or whitish, spongy, biconvex arilliform funicle, the testa typically hard, lustrous brown or black, pleurogram present or absent.
2n = 26 (Yeh et al. 1986; Tapia-Pastrana and Gómez-Acevedo 2005).
Twenty species, 18 recognised by
Largely confined to seasonally dry and arid vegetation including coastal dunes, thickets, thorn-scrub, and spiny chaparral, savanna margins, brush-savanna, pine woodland, pine-palmetto savanna, and deciduous or semi-deciduous forests. Several species are adapted to shallow rocky coastal limestone soils sometimes derived from coral reefs, e.g., in Belize and the Bahamas. The conspicuous spongy arils clasping the seeds (Fig.
Morphology of selected species of the Pithecellobium clade A Ebenopsis ebano (Berland.) Barneby & J.W. Grimes, fruit B, C Havardia albicans (Kunth) Britton & Rose B leaves C inflorescence D Gretheria campylacantha (L. Rico & M. Sousa) Duno & Torke, fruit E Ricoa leptophylla (DC.) Duno & Torke, inflorescence and young flowers F Painteria elachistophylla (A. Gray ex S. Watson) Britton & Rose, fruit G Pithecellobium dulce (Roxb.) Benth. habit H Pithecellobium unguis-cati (L.) Benth. inflorescence I Pithecellobium lanceolatum (Humb. & Bonpl. ex Willd.) Benth. pod and seeds J Pithecellobium winzerlingii Britton & Rose pod and seed K Sphinga acatlensis (Benth.) Barneby & J.W. Grimes flowers L Sphinga platyloba (Bertero ex DC.) Barneby & J.W. Grimes flowers. Photo credits A, E, F, H R Duno de Stefano B, C, J GA Romero González D, I, K CE Hughes G D Pedersen L C Ramírez.
Greek = pithecos, monkey + ellobion, earring; in allusion to the pendulous, contorted fruit, the spelling and meaning often modified from Pithecolobium, as though from pithecos + lobos, fruit, and later to Pithecollobium.
Pithecellobium dulce has edible seed-arils (
Pithecellobium was long considered the main “dustbin” genus of the old sense tribe Ingeae. The genus has gradually changed its circumscription from the late nineteenth century to its current much reduced form, defined for the first time by
Pithecellobium can be distinguished from all other genera of former tribe Ingeae by the presence of a seed funicle modified into a usually conspicuous spongy white, pink or crimson aril that cups the lower half of the seed (Fig.
One species, P. × bahamense Northrop is hypothesised by
Sphinga platyloba (DC.) Barneby & J.W. Grimes [≡ Acacia platyloba DC.]
Shrubs and small trees, sometimes sarmentose. Stipules subulate, spinescent. Leaves bipinnate, extrafloral nectaries below mid-petiole; pinnae 1–4 pairs; leaflets 1–14 pairs per pinna, opposite, obovate or oblong-obovate, venation pinnate or subpalmate. Inflorescences capitula, all or almost all arising from brachyblasts. Flowers sessile or nearly so, homomorphic, 5-merous, the perianth greatly elongated, flower-buds flask-shaped; calyx cylindrical-campanulate; corolla narrowly trumpet-shaped; stamens 34–176, the tube greatly elongated and far exserted; pollen in 16-celled polyads, more or less isodiametric; intrastaminal disc clasping stipe of ovary; ovary cylindrical. Fruits broad-linear, plano-compressed legumes, 6–10-seeded, the stiffly papery valves framed by sutures, the cavity continuous, dehiscent through both sutures. Seeds transverse on a dilated, contorted or sigmoid funicle.
2n = 26 (Rico Arce 1992).
Three species, south-central Mexico, sporadically in Central America (Belize, Guatemala and Nicaragua), to northern South America (Colombia, Venezuela), and Aruba, Curaçao (Dutch West Indies), and one species endemic to Cuba (Fig.
Arid or seasonally dry tropical forest, mattoral and thorn scrub, to 1600 m elevation in south-central Mexico. Night-flowering and pollinated by Sphingid moths.
Sphinx (Sphingidae), the putative pollinator + Tupi inga, vernacular for several mimosoid legumes (
Unknown.
The three species of Sphinga were all known to
Pithecolobium sect. Ortholobium Benth., Trans. Linn. Soc. London 30: 592. 1875. Type: Pithecellobium albicans (Kunth) Benth. [≡ Acacia albicans Kunth (≡ Havardia albicans (Kunth) Britton & Rose)]
Havardia brevifolia (Benth.) Small [≡ Pithecellobium brevifolium Benth. (= Havardia pallens (Benth.) Britton & Rose)]
Trees or bushy treelets 2–9(–12) m. Stipules spinescent. Leaves bipinnate, extrafloral nectaries sessile, shallowly cupular, thick-rimmed, inserted near or below mid-petiole, rarely 2, rarely at very base, sometimes a little above mid-petiole, sometimes rudimentary; pinnae 1–11 (13) pairs; leaflets 7–36 pairs per pinna, alternate, rarely opposite, linear, linear-oblong, to narrowly oblong-obovate, venation simple or pinnate. Inflorescence 6–37-flowered capitula or capituliform racemes. Flowers sessile or very shortly pedicellate, homomorphic, 5-merous; calyx hemispherical, shallowly campanulate to deeply campanulate, teeth deltoid; corolla campanulate, narrowly vase-shaped to subcylindrical, lobes recurving to erect; stamens (28) 30–52; pollen in 16-celled polyads, more or less isodiametric; intrastaminal disc 5-lobed or simple callosities; ovary subsessile or shortly stipitate, slenderly ellipsoid, style about as long as stamens, the stigma poriform. Fruits oblong to broad-linear, straight, plano-compressed legumes, 8–13-seeded, valves framed by a bluntly 3-angulate suture, valves chartaceous or thinly coriaceous, stiffly papery, low-convex over each seed on one face or on both, externally veinless, the cavity continuous; dehiscence tardy, inert, through both sutures. Seeds transverse, orbicular or oblong-elliptic, compressed-disciform and flattened around the periphery, funicle distally sigmoid, pleurogram U-shaped to closed.
Unknown.
Three species, almost entirely distributed in Mexico mainly in the north and the Yucatan peninsula, extending marginally north into the USA (south Texas), and south into northern Guatemala and Belize (Fig.
Largely confined to seasonally dry, drought deciduous woodland and arid thorn scrub and arid mattoral in the Sonoran desert, Tamaulipan plains and coastal thickets on sand, secondary forest, and extending weakly into lowland rainforest, occasionally on limestone. In parts of Mexico, abundant. For example, H. albicans (Kunth) Britton & Rose is very common in the peninsula of Yucatán, and H. pallens is abundant in parts of northern Mexico.
Named after Dr. Valery Havard (1846–1927), U.S. Army, a diligent student of the North American flora.
The boiled wood of H. albicans is used in the chemical industry to colour cement (e.g., for swimming pools;
Gretheria sonorae (S. Watson) Duno & Torke [≡ Pithecellobium sonorae S. Watson]
Arborescent shrubs and small, often multi-stemmed trees, 2–14 m. Stipules stout, recurved, early lignescent, spinescent, these persistent on shoots and trunks long after leaves are shed. Leaves bipinnate, extrafloral nectaries at or below the midpoint of the petiole, sessile, shallow-cupular, thick-rimmed or plane and dimpled; pinnae 1–6 (13) pairs; leaflets 10–31 pairs per pinna, opposite, venation pinnate, the midrib slightly displaced, giving rise on each side to 2–5 weak secondary veins. Inflorescences capituliform racemes arising from brachyblasts. Flowers sessile, homomorphic, 5-merous; calyx deeply campanulate; corolla subcylindrical, lobes erect, white-silky strigose dorsally; stamens 40–52; pollen in 16-celled polyads, more or less isodiametric; intrastaminal disc 5-lobed or simple callosities; ovary ellipsoid, shortly stipitate. Fruits oblong, contracted, body straight or almost straight, plano-compressed legumes, the valves bluntly framed by longitudinally 3-ridged sutures, stiff, somewhat brittle, brownish-green, externally veinless, glabrous, red-granular or both granular and puberulent outside, the cavity continuous. Seeds transverse, compressed, disciform to orbicular in outline, funicle dilated, sigmoid, pleurogram U-shaped.
Unknown.
Two species, disjunctly scattered along the Pacific coast of Mexico, from Sonora south to Central America (Honduras, Nicaragua, and northwestern Costa Rica) (Fig.
Tropical dry forests, thorn scrub and brush-woodlands, from sea level to 400 m elevation, occasionally to 700 m.
The generic name honours Rosaura Grether González, a Mexican botanist.
Gretheria was segregated from Havardia by
Ricoa leptophylla (DC.) Duno & Torke [≡ Acacia leptophylla DC.]
Xerophytic shrubs 0.2–1.5 m, often growing in patches several meters in diameter. Stipules straight to recurved spines, persistent. Leaves bipinnate, extrafloral nectaries subsessile or shortly stipitate, circular, between the first pinnae pair (sometimes also between the second pair), absent on the pinnae; pinnae 3–7 (9) pairs; leaflets 8–25 pairs, opposite, venation weakly developed, nearly simple or 1-branched, subcentric. Inflorescences capitula, arising from brachyblasts. Flowers sessile, mostly homomorphic but some functionally staminate, 5-merous; calyx campanulate, teeth ovate or deltate; corolla tubular, lobes ovate, recurving; stamens 40–76; intrastaminal callosities developed, sometimes obscure or wanting in staminate flowers; polyads 16-celled, more or less isodiametric; ovary slenderly ellipsoid, compressed, shortly stipitate. Fruits falcately or subcircinnately broadly linear fruits, the valves stiffly leathery, at first plano-compressed, becoming turgid and low-convex (on both faces of legume) over each seed, indistinctly venulose, the cavity continuous, dehiscence inert through both sutures. Seeds obliquely descending, compressed-lentiform, funicle straight or sinuous (but not sigmoid), testa smooth, hard, moderately lustrous, dark castaneous, the pleurogram incomplete.
Unknown.
Ricoa leptophylla grows in dry or semi-arid grasslands and thornscrub, extending into the lower edge of the pine-oak belt, on both basaltic and calcareous substrates, at 1600–2800 m.
The generic name honours María Lourdes Rico, whose profound dedication and unrelenting commitment to botanical research over decades has deeply enhanced knowledge and understanding of the Leguminosae, especially of the ingoid clade.
Ricoa was segregated from Painteria to account for the non-monophyly of that genus (
The common name for R. leptophylla is Huisache, a name that is also applied to Vachellia farnesiana (L.) Wight & Arn. and other related species with spinescent stipules (
Painteria revoluta (Rose) Britton & Rose [≡ Pithecellobium revolutum Rose]
Shrubs 0.3–15 cm. Stipules of long-shoots lignescent, of short-shoots acicular or subulate, rather closely imbricate. Leaves bipinnate, extrafloral nectaries between proximal pinna-pair, rarely between 2 pairs; pinnae 1–7 pairs; leaflets 3–20 pairs per pinna, suborbicular, elliptic, and broadly oblong, completely revolute in P. revoluta, venation palmate or almost simple. Inflorescences shortly spiciform capitula arising from brachyblasts. Flowers sessile or almost so, homomorphic, 5-merous; calyx campanulate or hemispherical, lobes deltoid; corolla sub-cylindrical, the lobes either ascending or recurving; stamens 28–76, the tube either included or shortly exserted; pollen in 16-celled polyads, more or less isodiametric; intrastaminal small callosities, sometimes obscure or wanting; ovary slenderly ellipsoid, compressed, on a short stipe. Fruits compressed legumes, retrofalcate or retrocircinnate, the leathery valves biconvex over each seed, the cavity continuous; dehiscence tardy through both sutures; funicle straight or sinuous, not sigmoid. Seeds compressed-lentiform.
Unknown.
Plains and hillsides in desert grassland and brush communities, mainly between 1400–2750 m.
The genus is named after Joseph Hannum Painter (1879–1908), botanist and assistant curator in the division of plants of the United States National Museum, who collected the type species in Querétaro.
Unknown.
Hoopesia Buckley, Proc. Acad. Nat. Sci. Philadelphia 13: 453. 1862. Type: Hoopesia arborea Buckley [= Ebenopsis ebano (Berland.) Barneby & J.W. Grimes]
Siderocarpos Small, Bull. New York Bot. Gard. 2: 91. 1901, nom. illeg., non Siderocarpus Pierre, Not. Bot. Sapot.: 31. 1890 (Sapotaceae). Type: Siderocarpos flexicaulis (Benth.) Small [≡ Acacia flexicaulis Benth. (= Ebenopsis ebano (Berland.) Barneby & J.W. Grimes)]
Ebenopsis ebano (Berland.) Barneby & Grimes [≡ Mimosa ebano Berland.]
Trees and shrubs, 2–8 (20) m. Stipules early lignescent, persistent. Leaves bipinnate, extrafloral nectaries interpinnal, cupular, short-stipitate; pinnae 1–4 pairs; leaflets 2–7 pairs per pinna, opposite, oblong, oblong-obovate rhombic-oblong, obovate-elliptic to suborbicular, venation palmate. Inflorescences spikes or capitula arising from brachyblasts. Flowers sessile, homomorphic, 5-merous; calyx campanulate; corolla tubular, lobes erect; stamens 32–66, the tube included or barely exserted from the corolla; pollen in 16-celled polyads, more or less isodiametric; intrastaminal disc lobed or small callosities. Fruits massive, compressed sausage-like, erect or slightly curved legumes, the woody valves produced inwardly as pithy interseminal septa, the exocarp in age breaking into polygons; dehiscence tardy, inert, the sutures at first separating at each end of pod but not gaping, ultimately separating through their whole length. Seeds transverse on straight, subterete funicle, irregularly globose, resembling chickpeas, red-castaneous in colour.
Unknown.
Three species. Markedly disjunct across Mexico in Baja California, Sonora-Sinaloa, north-eastern Mexico, and the northern half of the Yucatán Peninsula, the genus almost endemic to Mexico, but extending into south Texas (Fig.
In tropical and subtropical arid thickets and dry forests and adjacent desert hillsides and desertic fringes in Baja California and Sonora, mostly below 500 m, cultivated in Florida. Often locally abundant as a shrubby treelet forming thickets in thorn scrub and chaparral on impermeable caliche substrates in parts of Tamaulipas and Baja California. The fruits ripen and open slowly and are sometimes described as indehiscent, but after falling, often entire, the valves are eventually fully dehiscent shedding their seeds passively on the ground.
From the words ebano (Diospyros crassiflora Hiern, Ebeneaceae) and ópsis (“aspect”, “appearance”). The wood resembling African ebony in appearance.
In Mexico, the hard wood is used for fence posts and provides high quality charcoal, perhaps accounting for the scarcity of larger trees.
Else Demeulenaere12, Stefanie M. Ickert-Bond24, Daniel J. Murphy35, Bruce R. Maslin31,32, Gillian K. Brown7
Citation: Demeulenaere E, Ickert-Bond SM, Murphy DJ, Maslin BR, Brown GK (2024) 31. Archidendron clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 404–429. https://doi.org/10.3897/phytokeys.240.101716
Archidendron clade
Figs
Included genera (9). Acacia Mill. (1082 species), Archidendron F. Muell. (ca. 120), Archidendropsis I.C. Nielsen (11), Falcataria (I.C. Nielsen) Barneby & J.W. Grimes (3), Heliodendron Gill.K. Br. & Bayly (3), Pararchidendron I.C. Nielsen (1), Paraserianthes I.C. Nielsen (1), Serianthes Benth. (18), Wallaceodendron Koord. (1).
Description. Trees or shrubs. Stipules mostly present, caducous, variable in shape, rarely spinescent (some Acacia and Heliodendron) or absent (some Archidendron). Leaves mostly bipinnate, rarely unifoliolate (some Archidendron), reduced to scales or polymorphic phyllodes or absent (many Acacia); extrafloral nectaries present or absent, variable in shape; leaflets (in bipinnate leaves) opposite or alternate. Inflorescences either simple or compound racemes, panicles, spikes, capitula or umbels. Flowers uniform, bisexual, rarely unisexual (some Acacia and Archidendron), 5- or 4-merous (some Archidendron, Archidendropsis); calyx gamosepalous, less frequently free or absent (some Acacia), valvate; corolla gamopetalous, valvate; stamens numerous, united into a tube at the base, or free (most species of Acacia), staminal tube and corolla tube shortly untied at the base; pollen in 4, 8, 12, 16 or 32-celled polyads; ovary usually 1, less frequently 2 to several in some genera, sessile or stipitate. Fruit dehiscent through one or both sutures or indehiscent (Serianthes, rarely Acacia), endocarp indistinct or bright coloured or sometimes separating from the exocarp and breaking into 1-seeded envelopes (Wallaceodendron). Seeds flattened or swollen, unwinged or narrowly winged, seed coat hard or thin, funiculate or not, pleurogram present, U-shaped (Falcataria, Pararchidendron, Serianthes, Wallaceodendron, some Acacia), closed (some Acacia) or absent (Archidendron, Archidendropsis, Heliodendron).
Distribution. The members of the Archidendron clade are largely restricted to the Indomalayan and Australasian regions, from southern India and Sri Lanka in the west, to New Caledonia in the east; and from Taiwan and the Ryukyu Islands in the north, to Australia in the south.
Clade-based definition. The most inclusive crown clade including Archidendron lucidum (Benth.) I.C. Nielsen and Wallaceodendron celebicum Koord., but not Pithecellobium dulce (Roxb.) Benth., Cedrelinga cateniformis (Ducke) Ducke or Punjuba racemiflora (Donn. Sm.) Britton & Rose (Fig.
Habit and leaf morphology diversity in genera of the Archidendron clade A Archidendron clypearia (Jack) I.C. Nielsen B Heliodendron thozetianum (F. Muell.) Gill.K. Br. & Bayly C Falcataria falcata (L.) Greuter & R. Rankin D Heliodendron basalticum (F. Muell.) Gill.K. Br. & Bayly E Serianthes kanehirae var. kanehirae F Serianthes kanehirae var. yapensis Fosberg G Pararchidendron pruinosum (Benth.) I.C. Nielsen H Paraserianthes lophantha subsp. montana (Jungh.) Benth. with tomentose stems and leaves I Wallaceodendron celebicum Koord. Photo credits A C Ng B R Cumming C J Teo D D Richter E, F JB Friday G G Brown H F and K Starr I Plantoholic Sheila.
Inflorescence and flower diversity of genera in the Archidendron clade A Archidendron vaillantii (F. Muell.) F.Muell. B Archidendropsis paivana (E. Fourn.) I.C. Nielsen C Falcataria falcata (L.) Greuter & R. Rankin D Heliodendron thozetianum (F. Muell.) Gill.K. Br. & Bayly E Serianthes sp. F Pararchidendron pruinosum (Benth.) I.C. Nielsen G Paraserianthes lophantha (Willd.) I.C. Nielsen H Wallaceodendron celebicum Koord. Photo credits A, F A Furhmann B B Henry C JB Friday D S Worboys E T Rodd G AndyBonsai H Plantaholic Sheila.
Fruit and seed diversity of genera in the Archidendron clade A fruit of Archidendron lucyi F. Muell. (Brown 169) B open pod of Archidendropsis streptocarpa (E. Fourn.) I.C. Nielsen showing winged seeds C opened pod of Falcataria falcata (L.) Greuter & R. Rankin D Heliodendron xanthoxylon (C.T. White & W.D. Francis) Gill.K. Br. & Bayly opened pod (Hyland 9229) E Heliodendron basalticum (F. Muell.) Gill.K. Br. & Bayly winged seeds (Canning & B. Rimes 6173) F–H Serianthes dilmyi Fosberg showing the indehiscent (to very tardily dehiscent) (rarely) woody pods with transverse seeds which are each isolated in a chamber I Pararchidendron pruinosum (Benth.) I.C. Nielsen fruit and seedling J Paraserianthes lophantha (Willd.) I.C. Nielsen immature pods (Brown 204A) K Wallaceodendron celebicum Koord. pod. Photo credits A, I, J G Brown B B Henry C JB Friday D, E Queensland Herbarium F–H R CJ Lim K Plantaholic Sheila.
Diversity in habit and foliage of Acacia A arborescent habit of A. cyperophylla var. omearana Maslin B ‘Minni Ritchi’ bark of A. grasbyi Maiden C prostrate habit of A. pulviniformis Maiden D shrub habit of A. brachybotrya Benth. E A. argyrophylla Hook. showing 1-veined phyllodes and globose inflorescences F bipinnate foliage of A. leucoclada subsp. argentifolia Tindale G A. spondylophylla F. Muell. showing phyllodes arranged in regular whorls along branches H A. alata var. biglandulosa Meisn. showing decurrent phyllodes that form bifarious wings along branches I A. tetraptera Maslin showing unusually small phyllodes (B.R. Maslin 5792). Photo credits A, C, E, H, I B Maslin B A George D D Murphy F L Jessup G K Brennan.
Diversity in inflorescences and flowers of Acacia A heads in terminal panicle and racemes of A. pyrifolia var. pyrifolia (B.R. Maslin 8425) B heads in racemes of A. neriifolia A. Cunn. ex Benth. C axillary heads of A. heterochroa subsp. heterochroa (B.R. Maslin 4766) D axillary spikes with densely arranged flowers of A. fecunda Maslin (B.R. Maslin 8762) E spikes with loosely arranged flowers and arranged in short racemes of A. leptostachya Benth. Photo credits A, C, D B Maslin B unknown E IB Armitage.
Diversity in fruits and seeds of Acacia A pods of A. catenulata subsp. occidentalis Maslin that readily break into 1-seeded loments (B.R. Maslin 8880) B fruits of the ‘Mulga’ species A. incurvaneura Maslin & J.E.Reid (B.R. Maslin 9304B) C A. ancistrocarpa Maiden & Blakely fruits are superficially similar to those of Calliandra D A. ampliceps Maslin showing brittle fruits with seeds having bright red aril (bird dispersed) (B.R. Maslin 8660) E A. coolgardiensis Maiden showing terete fruits F irregularly coiled fruits of A. auriculiformis A. Cunn ex Benth. showing seeds encircled by showy aril. Photo credits A–E B Maslin F K Brennan.
Notes. The Archidendron clade was first identified by
Within the Archidendron clade, only the Serianthes clade is consistently resolved, grouping Falcataria, Serianthes, and Wallaceodendron, with Serianthes and Falcataria strongly supported as sister genera (
Wallaceodendron celebicum Koord.
Unarmed medium-sized to large trees up to 45 m. Stipules caducous. Leaves bipinnate, extrafloral nectaries between pinnae pairs; pinnae 2–3 pairs; leaflets 3–6 pairs per pinna, opposite. Inflorescence a solitary or paired raceme, peduncle 5–16 cm long. Flowers uniform, bisexual, 5-merous, subtended by ca. 2 mm long, triangular, caducous bracts; calyx gamosepalous, valvate; corolla gamopetalous, valvate; stamens numerous, united into a tube at the base, staminal tube and corolla tube shortly united at the base; pollen in 16-celled polyads with a perforated tectum; ovary solitary. Fruit a woody legume, flat, straight to slightly curved, tardily dehiscent, not segmented, not reddish inside; exocarp thin, crustaceous, mesocarp woody, endocarp chartaceous loosening and at dehiscence forming small, closed envelopes around each seed. Seeds strongly flattened, circular, unwinged, with U-shaped pleurogram, without aril; testa with a thick sclerotesta.
2n = 26 (
Monospecific (W. celebicum), endemic to Malesia [North Sulawesi (Celebes) and the Philippines] (Fig.
Wallaceodendron can be found along seashores and inland in primary rainforest to 850 m elevation.
Named after the British botanist Alfred Russel Wallace (1823–1913), explorer, zoologist, and plant collector (
Wallaceodendron celebicum [banúyo (=ironwood), derham mahogany] is used for furniture, flooring, light construction, telegraph poles, music instruments, decorative veneers, carvings and sculptures (
Paraserianthes sect. Falcataria I.C. Nielsen, Bull. Mus. Natl. Hist. Nat., B, Adansonia Sér. 4, 5(3): 327. 1983 (publ. 1984). Type: Paraserianthes falcataria (L.) Nielsen [≡ Adenanthera falcataria L.]
Falcataria falcata (L.) Greuter & R. Rankin [≡ Falcataria moluccana (Miq.) Barneby & Grimes (≡ Albizia moluccana Miq.)]
Unarmed medium-sized to large trees up to 40 m. Stipules caducous. Leaves bipinnate, extrafloral nectaries disk-shaped; pinnae 8–24 pairs; leaflets (4) 8–27 (33) pairs per pinna, opposite, subsessile. Inflorescences 2–3 times branched, efoliate panicles of few-flowered spikes, each panicle axillary to a fully expanded leaf, the terminal meristem of each annual branch-complement continuing beyond the fertile axes. Flowers bisexual, 5-merous, homomorphic and sessile; calyx gamosepalous, campanulate or hemispherical; corolla gamopetalous, sericeous; stamens numerous; pollen in 16-celled polyads with a thin exine and a thick nexine, without costae; intrastaminal disc around the base of the ovary. Fruits broad-linear, straight, plano-compressed, narrowly winged along the ventral suture, inertly dehiscent through both sutures. Seeds compressed ellipsoid, coat hard, testa brown, U-shaped pleurogram.
2n = 26 [Albizia falcata (L.) Baker (= Falcataria falcata)] (
Three species [F. falcata, F. pullenii (Verdc.) Gill.K. Br., D.J. Murphy & Ladiges, F. toona (F.M. Bailey) Gill.K. Br., D.J. Murphy & Ladiges], native to the Moluccas, New Guinea, Solomon Islands, and Queensland (
Tropical rainforest and coastal dry rainforests, spanning elevations up to 2300 m (Lewis and Rico Arce 2005).
The name Falcataria is derived from the Latin word falcatus which means “sickle-shaped, hooked” (
The wood is commercially used for pulpwood, veneers, light construction, crafts and furniture, fuel and charcoal (
The molecular phylogenetic analyses of
The name applied to the type species of the genus has a complicated history (see
Albizia sect. Serianthes (Benth.) F. Muell., Fragm. 8: 165. 1874. Type: Albizia grandiflora (Benth.) F. Muell. [≡ Serianthes dilmyi Fosberg]
Serianthes grandiflora Benth.
Unarmed shrubs, 6–10 m, and trees to 27 m. Stipules linear to filiform, observed only in the seedling stage. Leaves bipinnate, extrafloral nectary circular or elliptical usually on the lower half of the petiole, additional nectaries usually on the leaf rachis and on the pinnae between the leaflets insertion; pinnae (4) 6–22 pairs, opposite or subopposite; leaflets (6) 8–25 (30) pairs per pinna, alternate except for the distal pair, sessile, mostly asymmetrical with a diagonal midvein. Inflorescence an umbel, raceme, or panicle composed of pedunculate spikes, pedunculate racemes or 1–4-flowered capitula. Flowers uniform, bisexual, 5-merous; calyx gamosepalous, valvate, usually circumscissile at the base; corolla gamopetalous, valvate, tube untied with the staminal tube in the lower part; stamens numerous, united into a tube at the base; pollen in 16-celled polyads with a perforated tectum in each locule; ovaries 1, less frequently 2, sessile. Fruits indehiscent to very tardily dehiscent (rarely), woody with transverse seeds which are isolated in a chamber each. Seeds flattened with a hard, black testa and a U-shaped pleurogram, lack a wing.
2n = 26 (only two counts of Serianthes kanehirae Fosberg are known) (
ca. 18 species, five subspecies, and two varieties, widely distributed in the Indo-Pacific region (Fig.
The more widespread Serianthes dilmyi Fosberg is adapted to supralittoral habitats. The other Serianthes taxa occur from coastal shrub, moist forest to lowland wet tropical rainforest on calcareous or volcanic soils, but in New Caledonia Serianthes is also found in maquis on ultramafic (serpentine) soils (
Serianthes is derived from the Greek serikon (= silk) and anthos (= flower) because the flowers of the tree have a silky appearance (
The wood of Serianthes was used traditionally mostly for canoes, paddles, traditional boats, traditional and ceremonial houses, and for storyboards (
Three main revisionary treatments (
The distribution of certain Serianthes taxa across different islands (and a few mainland areas) results in vernacular names being expressed in either different languages or even the same language, but with a different name. For example, the Palauan name of Serianthes kanehirae var. kanehirae Fosberg is ukall or kumer, while the Yapese name of Serianthes kanehirae var. yapensis Fosberg is gumor. Serianthes nelsonii Merr. occurs on two islands of the Mariana Islands archipelago. The CHamoru name for the tree is håyun lågu on Guam and tronkon guåfi on Rota.
Albizia sect. Spiciflorae Benth., London J. Bot. 3: 85. 1844. Type: Albizia fulgens (Labill.) Benth. [≡ Acacia fulgens Labill. (≡ Archidendropsis fulgens (Labill.) I.C. Nielsen)]
Albizia sect. Spiciflorae ser. Platyspermae Benth., Trans. Linn. Soc., London 30: 558. 1875. Type not designated.
Archidendropsis fulgens (Labill.) I.C. Nielsen [≡ Acacia fulgens Labill.]
Unarmed trees or shrubs. Stipules small ovate or filiform and often caducous, or large auriculate, orbicular, or cordate and persistent, never thorny. Leaves bipinnate, pinnae 1–14 pairs; leaflets 1–25 pairs per pinna, opposite or alternate; extrafloral nectaries circular, elliptic or obtriangular, on leaf rachis and sometimes also on the pinnae. Inflorescences in axillary or terminal spikes, spiciform racemes, racemes or in one species [A. fournieri (Vieill.) I.C. Nielsen] capitate, but when capitate the calyx and corolla are glabrous. Flowers uniform, bisexual, 4- or 5-merous, red, white, green, yellow or pink, usually subtended by very small tetrameric or pentameric bracts; calyx gamosepalous, cupular to campanulate; corolla gamopetalous, funnel shaped; stamens numerous, filaments united into a tube at the base; pollen in 16 or 32-celled polyads, tectum with non-isometric channels; ovaries mostly one, several in A. oblonga (Hemsl.) I.C. Nielsen. Fruits usually flat and dehiscent along both sutures, not partitioned inside and endocarp not separating from the exocarp. Seeds flattened or swollen, narrowly winged when flattened, thin-walled and lacking a pleurogram.
Unknown.
Eleven species from New Caledonia, New Britain, the Solomon Islands and on the island of New Guinea (Fig.
Most species with a restricted area of distribution, found in rainforests, gallery forest, wooded ravines, or mesophyllous forest.
From the generic name Archidendron with the Latin suffix -opsis, referring to the similarity to that genus.
Unknown.
Archidendropsis was published as “Gen. B” by
Pararchidendron pruinosum (Benth.) I.C. Nielsen [≡ Pithecellobium pruinosum Benth.]
Unarmed trees or shrubs. Stipules linear not spinescent, usually inconspicuous and caducous. Leaves bipinnate, extrafloral nectaries flat to concave on leaf rachis and pinnae; pinnae 1–4 pairs; leaflets 3–12 pairs per pinnae, alternate, petiolulate. Inflorescence units globose, umbellate, clustered in axillary, pedunculate racemes. Flowers uniform, bisexual, 5-merous, pedicellate; calyx gamosepalous, cupular to hemispherical; corolla gamopetalous, tubular to slightly funnel-shaped, valvate; stamens numerous, united into a tube at the base and shortly united with the corolla tube; pollen in 16-celled polyads, non-isometric channels crossing +/- entirely the tectum, non-radially oriented; ovary solitary, stipitate. Fruit curved into a circle to contorted, chartaceous, flattened, not segmented, dehiscent first along ventral suture, reddish inside, the endocarp not forming envelopes around each seed. Seeds ellipsoid, obovoid or subglobose, with U-shaped pleurogram, without aril, with a thick, black sclerotesta, unwinged.
Unknown.
Monospecific (P. pruinosum), with four varieties, in Java, Lesser Sunda Islands (Sumbawa, West Flores), Timor-Leste, the island of New Guinea, and Australia (Queensland and New South Wales) (Fig.
In Australia this species is reported from the rainforest, coastal scrub and semi-deciduous forest (sea level to 800 m); in the Malesian area it is found in montane rainforest, both primary and secondary (400–2250 m).
The Greek prefix para (= close by) alludes to the similarity of this genus to Archidendron (
Pararchidendron is sold in Australia as a horticultural decorative plant that can be a screen break or shade and produces a decorative general purpose timber (
Pararchidendron was identified as “Gen. C” by
Pithecellobium sect. Clyplearia Benth., London J. Bot. 3: 206. 1844. Type: Pithecellobium clypearia (Jack) Benth. [≡ Inga clypearia Jack (≡ Archidendron clypearia (Jack) I.C. Nielsen)]
Ortholobium Gagnep., Bull. Soc. Bot. Fr. 99: 36. 1952, nom. inval. (no Latin descr.)
Cylindrokelupha Kosterm., Bull. Org. natuurw. Onderz. 20: 20. 1954. Type: Cylindrokelupha bubalina (Jack) Kosterm. [≡ Inga bubalina Jack (≡ Archidendron bubalinum (Jack) I.C. Nielsen)]
Morolobium Kosterm., Bull. Org. natuurw. Onderz. 20: 20. 1954. Type: Morolobium monopterum (Kosterm.) Kosterm. [≡ Pithecellobium monopterum Kosterm. (≡ Archidendron monopterum (Kosterm.) I.C. Nielsen]
Paralbizzia Kosterm., Bull. Org. natuurw. Onderz. 20: 23. 1954. Type: Paralbizzia turgida (Merr.) Kosterm. [≡ Pithecellobium turgidum Merr. (≡ Archidendron turgidum (Merr.) I.C. Nielsen)]
Archidendron vaillantii (F. Muell.) F. Muell. [≡ Pithecellobium vaillantii F. Muell.]
Unarmed trees or shrubs, small to medium sized. Stipules present, sometimes glandular, or absent. Leaves bipinnate, rarely unifoliolate; pinnae 1–14 pairs; leaflets mostly opposite, rarely alternate (3 species), variable in number, shape and size; extrafloral nectaries sessile, sunken, raised or stipitate, round, boat-shaped, triangular or irregular shaped present on the rachis, additional glands present (various combinations and shapes) or absent on the pinnae. Inflorescences simple or compound in pedunculate capitula, umbels, corymbs or racemes, if compound may be arranged in cauliflorous, ramiflorous, axillary or terminal panicles; extrafloral nectaries sometimes present on floral bracts and capitula. Flowers uniform, bisexual or unisexual, 4- or 5-merous, yellow, green, white or cream; calyx gamosepalous; corolla gamopetalous; stamens numerous, united into a tube at the base and joined with the corolla in the lower part; pollen in (4, 12) 16 (32)-celled polyads; ovary 1–several per flower, sessile or stipitate. Fruits chartaceous, coriaceous, fleshy or woody, straight, curved or spirally twisted, flat to terete, sometimes internally segmented, dehiscing either along the dorsal or ventral suture, sometimes along both sutures, most often reddish outside and orange-reddish inside. Seeds may be funiculate, ellipsoid, or flattened, with a black or bluish black testa, pleurogram lacking, unwinged.
2n = 26. Only three species [A. clypearia, A. jiringa (Jack) I.C. Nielsen, A. turgidum] have published chromosome numbers (
Ninety-nine species described in this Indomalayan-Australasian genus and an additional 20 putative species that are poorly known due to limited collections or destroyed types. Archidendron is distributed from Kerala (southern India) and Sri Lanka in the west, to the Solomon Islands in the east; and from Taiwan and the Ryukyu Islands in the north, to Australia in the south (Fig.
Lowland and montane tropical and subtropical rainforests.
According to
The seeds of A. jiringa are eaten in Thailand, Malaysia and Indonesia in a dish known as “jenkol”; the young shoots are also eaten. Archidendron jiringa has also been used as timber, dye (from the pods) and the leaves (and those of A. lucidum) have been used for traditional medicine (e.g., for the treatment of diabetes, inflammatory diseases and cancer) (
Recent evidence from molecular phylogenies suggests that Archidendron is non-monophyletic (
Archidendropsis subg. Basaltica I.C. Nielsen, Bull. Mus. Natl. Hist. Nat., B, Adansonia Sér. 4, 5(3): 325. 1984. Type: Archidendropsis basaltica (F. Muell.) I.C. Nielsen [≡ Acacia basaltica F. Muell. (≡ Heliodendron basalticum (F. Muell.) Gill.K. Br. & Bayly)]
Heliodendron basalticum (F. Muell.) Gill.K. Br. & Bayly [≡ Acacia basaltica F. Muell.]
Trees or shrubs to 37 m. Stipules either resembling small thorns to 1.2 mm long that are caducous, or persistent circular-ovate glands 1–3 mm in diameter. Leaves bipinnate; extrafloral nectaries at junction of pinnae circular or triangular to rhombic, circular glands sometimes at junction of leaflet petiolules; pinnae 1–2 pairs; leaflets 1.5–11 pairs per pinna, opposite, subsessile or long (3.5–7 mm) petiolulate, elliptic to elliptic-lanceolate or oblong. Inflorescence capitula, either simple or arranged into a panicle to 35 cm long. Flowers uniform, bisexual, 4- or 5-merous, yellow to cream, hairy, sessile; calyx gamosepalous, tubular to subcampanulate, symmetrical; corolla gamopetalous, tubular to narrowly campanulate; stamens numerous, united basally into a tube that equals or slightly exceeds the corolla tube; pollen in 16-celled polyads, tectum with isometric channels; ovary solitary and shortly stipitate. Fruits brown, valves chartaceous, oblong, flat and dehiscing along both sutures. Seeds lacking a pleurogram, flat, circular to ovate or obliquely ovate, with a narrow 0.2–1 mm peripheral, membranous wing.
Unknown.
Three species restricted to the state of Queensland in Australia (Fig.
One species [H. xanthoxylon (C.T. White & W.D. Francis) Gill.K. Br. & Bayly] is found in rainforest remnants on granitic soils whereas the other two species [H. basalticum and H. thozetianum (F. Muell.) Gill.K. Br. & Bayly] are found in drier habitats on sandy to sandy-clay loam soils, either in eucalypt woodland, acacia shrubland or semi-deciduous vine thickets.
From helios (Greek = sun) in reference to the endemic distribution in the Australian state of Queensland, widely known as the “sunshine state”, and to the capitate, sun-like inflorescences of yellow flowers, and dendron (Greek = tree) to the tree habit.
Heliodendron has been used as timber for general building and the manufacture of window frames, cabinets, barrels and boat building (
Heliodendron was newly described for the morphologically distinct Archidendropsis subg. Basaltica after phylogenetic studies found the two subgenera of Archidendropsis to be non-monophyletic (
Albizia sect. Lophantha ser. Pachyspermae Benth., Trans. Linn. Soc., London 30: 559. 1875, nom. illeg. Type: Albizia lophantha (Willd.) Benth. [≡ Acacia lophantha Willd. (≡ Paraserianthes lophantha (Willd.) I.C. Nielsen)]
Albizia sect. Pachyspermae (Benth.) Fosberg, Reinwardtia 7: 74. 1965. Type: Albizia lophantha (Willd.) Benth. [≡ Acacia lophantha Willd. (≡ Paraserianthes lophantha (Willd.) I.C. Nielsen)]
Paraserianthes lophantha (Willd.) I.C. Nielsen [≡ Acacia lophantha Willd.]
Unarmed shrub or tree to 10 m. Stipules to 2 mm long, either linear, puberulous and caducous (P. lophantha subsp. lophantha), or subcordate-triangular to ovate-lanceolate and densely tomentose [P. lophantha subsp. montana (Jungh.) I.C. Nielsen]. Leaves bipinnate, extrafloral nectaries present, elliptic to oblong on the petiole, smaller circular glands sometimes between the terminal pairs of pinnae and/or leaflets; pinnae 7–14 pairs, opposite; leaflets 15–40 pairs, sessile, opposite, inequilaterally narrowly oblong to lanceolate to oblong. Inflorescences solitary or paired axillary racemes, 5.2–18 cm long. Flowers uniform, bisexual, 5-merous, yellowish-green; calyx gamosepalous, narrowly cup-shaped; corolla gamopetalous, funnel-shaped; stamens numerous and united into a tube at the base; pollen in 16-celled polyads, with costae (pores surrounded by distinct thickenings), surface with few tectal channels; ovary sessile (P. lophantha subsp. montana) or shortly stipitate to subsessile (P. lophantha subsp. lophantha). Fruits chartaceous, flat, straight-edged and dehiscent along both sutures. Seeds subcircular-elliptic or oblong, black, hard testa, flat or convex and wingless, with U-shaped pleurogram.
2n = 26 (
Monospecific (P. lophantha), with two recognised subspecies: P. lophantha subsp. lophantha, native to south-west Western Australia, and P. lophantha subsp. montana, native to Sumatra and Java, and the Lesser Sunda islands of Bali and Flores (Fig.
In Australia, Paraserianthes occurs in coastal forests, and coastal or near-coastal open eucalypt forest, thicket, shrubland and grassland (
The Greek prefix para (= close by) refers to the similarity of this genus to Serianthes (
Paraserianthes lophantha is an ecologically, horticulturally and economically important species across the tropics. It has been planted as part of reforestation programs for its rapid growth, as an ornamental, and also as a shade tree in cocoa and coffee plantations (
Paraserianthes was originally described with two sections (sect. Paraserianthes and sect. Falcataria;
Acacia sect. Phyllodineae DC., Prodr. [A.P. de Candolle] 2: 448. 1825. Type not designated.
Racosperma Mart., Hort. Reg. Monac.: 188. 1829, nom. inval. (name not accepted by author)
Phyllodoce Link, Handbuch 2: 132. 1829, non Phyllodoce Salisb., Parad. Lond. ad t. 36. 1806 (Ericaceae). Type not designated.
Racosperma
Mart., Index Seminum [München (Monacensis)]: 4. 1835. Lectotype (designated by
Cuparilla
Raf., Sylva Tellur.: 120. 1838. Lectotype (designated by
Drepaphyla
Raf., Sylva Tellur.: 120. 1838. Lectotype (designated by
Hectandra
Raf., Sylva Tellur.: 120. 1838. Lectotype (designated by
Zigmaloba Raf., Sylva Tellur.: 120. 1838. Type: Zigmaloba sulcata (R. Br.) Raf. [≡ Acacia sulcata R. Br.]
Chithonanthus Lehm., Pl. Preiss. 2: 368. 1848. Type: Chithonanthus restiaceus (Benth.) Lehm. [≡ Acacia restiacea Benth.]
Tetracheilos Lehm., Pl. Preiss. 2: 368. 1848. Type: Tetracheilos tetragonocarpa Meisn., nom. illeg.
Acacia sect. Phyllodoce (Link) Kuntze, in T.E. von Post & C.E.O. Kuntze, Lex. Gen. Phan.: 2. 1903. Type not designated.
Phytomorula
Kofoid, Univ. Calif. Publ. Bot. 6: 38. 1914. Type: Phytomorula regularis Kofoid. [described as an alga but shown to be pollen of Acacia sp., vide
Acacia penninervis Sieber ex DC.
Trees, shrubs or subshrubs, never lianas; prickles absent. Stipules normally present and caducous, rarely spinose. Leaves bipinnate or modified to polymorphic phyllodes, rarely reduced to scales or absent; extrafloral nectaries normally present. Inflorescences globose or oblongoid capitula or spikes, axillary or aggregated in racemes or infrequently panicles, pedunculate or sometimes sessile. Flowers bisexual or staminate and bisexual within a single inflorescence, 5-merous or sometimes 4-merous, uniform, white to golden, rarely mauve-pink or red; sepals free to united, very rarely absent; corolla connate, valvate; stamens numerous, normally free, rarely basally united; pollen in polyads normally 8, 12 or 16 grains, the grains extraporate or sometimes porate, surface with pseudocolpi, exine granular (i.e., lacking columellae that occur in Vachellia Wight & Arn.); ovary solitary or very rarely 2–5 (e.g., A. celastrifolia Benth.). Fruits variable, dehiscent, rarely indehiscent. Seeds normally with a pleurogram and without endosperm; funicle arillate or sometimes exarillate.
Mostly 2n = 26 with ca. 100 species examined (
1082 species, the majority in Australia. These species are distributed throughout the continent with the major centre of species-richness located in south-west Western Australia, and secondary centres of richness in eastern Australia south of the Tropic of Capricorn associated with the Great Dividing Range, and in northern and north-eastern Australia. Although species of Acacia are a conspicuous component of the central Australian arid zone, this is a relatively species-poor region (
Species of Acacia within Australia grow in a wide range of habitats, from coastal to subalpine, tropical to arid ecological vegetation classes. They are particularly conspicuous and common in many semi-arid and sub-tropical shrublands and woodlands, and while they dominate much of the arid zone the species numbers are, relatively speaking, not especially high. The diverse shrublands of Western Australia and Eucalyptus woodlands of eastern Australia associated with the Great Dividing Range, located between the arid and temperate zones, are especially species-rich, but Acacia is often not a dominant element of the vegetation (
It is normally regarded that the name Acacia is derived from the Greek, ake (= a point), in reference to the spiny stipules that characterised the first (African) species described as Acacia; these species now belong to the genus Vachellia.
Acacias have had extensive utilisation for economic, social and environmental purposes. A discussion of these uses is presented by
Commercially, the most important use of Acacia is for wood products where the species A. auriculiformis A. Cunn. ex Benth., A. crassicarpa A. Cunn. ex Benth., A. mangium Willd. and A. × mangiiformis Maslin & L.A.J. Thomson are especially important in tropical forestry plantation industries, particularly in south-east Asia and India (
As noted by
Several species of Acacia have displayed significant invasiveness in places where they have been introduced. Examples include A. cyclops A. Cunn. ex G. Don, A. baileyana F. Muell., A. dealbata Link, A. mangium, A. melanoxylon, A. pycnantha Benth. and A. saligna. Factors promoting the spread of such species as weeds include the production of large quantities of tough, long-lived seeds, prolific seedling recruitment following fire or other environmental disturbances, and the absence of natural invertebrate predators and fungal pathogens.
Indigenous Australians have long used species of Acacia as a source of food and medicine, tools and weapons, and various other purposes (e.g.,
As discussed at
In recent years there have been substantial changes to both the classification and nomenclature of Acacia. The reasons for this are twofold. Firstly, molecular and other evidence have shown that the former broadly circumscribed, pantropical genus Acacia is polyphyletic and should be treated as comprising at least seven genera (see below). Secondly, the name Acacia is now conserved with a new type, namely, the Australian species Acacia penninervis Sieber ex DC. which replaces the Afro-Asian species, Acacia nilotica (L.) Willd. ex Delile. This retypification has meant that about half of the non-Australian species formerly called Acacia are now Vachellia and the name Racosperma Mart. [preferred by
For several years prior to the fragmentation of Acacia s.l. the genus was commonly viewed as comprising three subgenera as defined by
Although all genetic studies have shown Acacia s.s. to be monophyletic, the morphological and other characters that separate this genus from the six genera excised from Acacia s.l. have been poorly studied. Nevertheless, 95% of Acacia s.s. species possess phyllodes which readily distinguishes them from the bipinnate-leaved species of the other six genera (but there are also 73 bipinnate-leaved species in Acacia s.s.;
The infrageneric classification adopted for Acacia today is that of
Felipe da Silva Santos2, Carolina Lima Ribeiro2, Luciano Paganucci de Queiroz2
Citation: Santos FS, Ribeiro CL, Queiroz LP (2024) 32. Cedrelinga. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 430–433. https://doi.org/10.3897/phytokeys.240.101716
Cedrelinga cateniformis (Ducke) Ducke [≡ Piptadenia cateniformis Ducke]
Phylogenomic studies support Cedrelinga Ducke as related to other genera of the Ingoid clade but its phylogenetic position is not fully resolved as it appears in a mostly isolated position in a polytomy including the genus Pseudosamanea Harms and the Samanea, Jupunba, Inga and Albizia clades (
Unarmed emergent trees, 25–60 (65) m, to 2.5 m diameter, the trunk reddish with a rugous and striated bark, buttressed at base. Stipules absent. Leaves bipinnate, foliar glands at or near the insertion of each pair of pinnae and on the pinnae between the insertion of leaflets; pinnae 2–4 pairs, opposite; leaflets 3–4 pairs per pinna, opposite, elliptical with acute or acuminate apex, pinnately veined, glabrous. Inflorescences 8–20 flowered hemispherical capitula, grouped in terminal panicles or pseudoracemes; bracts ovate or obovate-spatulate, puberulent, persistent. Flowers 5-merous, greenish-white, glabrous except for ciliate calyx-teeth, and sometimes papillate on corolla lobe tips; calyx gamosepalous, short-campanulate; corolla gamopetalous with ovate lobes; stamens 24–30, greenish-white, united to the middle; intrastaminal disc absent; pollen in 16-celled acalymmate polyads, each pollen grain 6-porate with a finely reticulate surface; ovary shortly stipitate, ellipsoid, abruptly conic at apex, style a little longer than stamens. Fruits pendulous, indehiscent, lomentiform, linear but with deeply constricted margins forming 2–5 (6) oblong-elliptic, plano-compressed and one-seeded articles, twisted through ±90° at each isthmus, but plane and straight between them; valves brownish, glabrous, sinuously venulose, separating through the transverse fission of the isthmus and dispersed individually. Seeds disciform.
Unknown.
Monospecific (C. cateniformis), across equatorial latitudes of the Amazon delta region, reaching the main tributary rivers in Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Suriname, Venezuela and near the Andes in Peru (
Cedrelinga cateniformis (Ducke) Ducke, the only species in the genus A tree in a natural park environment in Guyana B trunk showing the rough and striate bark C flowering branch with foliage and inflorescences D extrafloral nectaries at the point of insertion on the leaflets (arrowheads) E flowers grouped into hemispherical capitula F loment, showing the articles rotated between them G separate mature articles. Photo credits A, B, F © S Sébastien Sant / Parc amazonien de Guyane (Cedrelinga cateniformis (Ducke) Ducke, 1922-Description, detailed sheet (mnhn.fr) C, D, E unknown-BIOWEB (Galería Bioweb Ecuador) G RB Foster © Field Museum of Natural History - CC BY-NC 4.0 (Cedrelinga cateniformis | Fotos de Campo | The Field Museum).
Wet or seasonally dry primary forests, especially along streams.
The name Cedrelinga refers to the similarity of its trunk with that of Cedrela fissilis Vell. and C. odorata L. (‘cedro’ in Brazilian Portuguese, family Meliaceae) and the Tupi ‘ingá’, which is an indigenous name for several South American species of rainforest mimosoid legume genera.
Cedrelinga cateniformis can be used as timber, in construction, carpentry, as paper and cellulose, and in agroforestry, due to its rapid seedling development and association with nitrogen-fixing bacteria (Loureiro et al. 1979; Baluarte and Alvarez 2015). It is reported as a medicine but without information on which part of the plant is used (Lewis and Rico Arce 2005). It is known by the vernacular names “chuncho” (Ecuador), “cedrorana” (Brazil, literally fake cedar in Tupi language), “iacaiacá” (Brazil), “cachicana”, “mure”, “guaura” (Venezuela), “don-ceder” (Suriname), “huayracaspi” and “tornillo” (Peru) (
When describing Cedrelinga,
Andrés Fonseca-Cortés2
Citation: Fonseca-Cortés A (2024) 33. Pseudosamanea. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 434–436. https://doi.org/10.3897/phytokeys.240.101716
Pseudosamanea guachapele (Kunth) Harms [≡ Acacia guachapele Kunth]
The phylogenetic position of Pseudosamanea is not well resolved in the Mimoseae phylogeny (
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.
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-Cortés 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).
2n = 26 [P. carbonaria (Britton) E.J.M. Koenen and P. guachapele] (Rico Arce 1992).
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.
Pseudosamanea carbonaria grows in dry and humid forests (
From the Greek pseudo (= false) and Samanea, due to the similarity to that genus (Lewis and Rico Arce 2005).
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 (
As defined by
João Iganci20,25, Marcos Vinícius Batista Soares20, Marli Pires Morim8
Citation: Iganci J, Soares MVB, Morim MP (2024) 34. Jupunba clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 437–444. https://doi.org/10.3897/phytokeys.240.101716
Jupunba clade
Figs
Included genera (4). Hydrochorea Barneby & J.W. Grimes (10 species), Jupunba Britton & Rose (37), Pseudalbizzia Britton & Rose (17), Punjuba Britton & Rose (5).
Description. Trees, treelets and shrubs, unarmed. Stipules puberulent to glabrous, deltate, elliptic, lanceolate, linear, ligulate, ovate, subulate, triangular or triangular-ovate, veinless or faintly 3-veined, falling early to tardily, or absent. Leaves bipinnate, pinnae 1–16 (19) pairs, leaflets 1–52 (63) pairs, alternate, variable in size and shape, the first pair of leaflets often reduced to paraphyllidia in Pseudalbizzia; extrafloral nectaries sessile or rarely stipitate, between the pairs of pinnae, or close below the proximal pair of pinna-pulvines, campanulate to cupular, orbicular, patelliform, elliptic, vertically elongate, verruciform or obsolete. Inflorescences congested or lax racemes, capitate, corymbose-umbellate or spiciform. Flowers homomorphic or heteromorphic, pedicellate or sessile; sepals 5–8, united; petals 5–8, joined into a gamopetalous corolla; stamens numerous, filaments fused into a tube, stemonozone present, anthers rimose; pollen 16, 18, 28-celled polyads; ovary glabrous or pubescent. Fruit a dehiscent legume, lomentiform or with the endocarp breaking into 1-seeded envelopes. Seeds disciform, oblong, elliptic, ovate, obovate, orbicular or flattened, translucent, pleurogram present or absent.
Distribution. In the Neotropics from Argentina to Colombia in South America, Central America, and West Indies, with two species of Hydrochorea from Democratic Republic of the Congo and Senegal, in Africa.
Clade based definition. The most inclusive crown clade containing Punjuba racemiflora (Donn. Sm.) Britton & Rose and Pseudalbizzia berteroana (Balb. ex DC.) Britton & Rose, but not Pseudosamanea guachapele (Kunth) Harms, Samanea saman (Jacq.) Merr. or Albizia retusa Benth. (Fig.
Notes. Although fruit morphology has been used in the past to group taxa in the former tribe Ingeae sensu
Hydrochorea and Balizia Barneby & J.W. Grimes were recognised by
Arthrosamanea Britton & Rose, Ann. New York Acad. Sci. 35: 128. 1936. Type: Arthrosamanea pistaciifolia (Willd.) Britton & Rose ex Britton & Killip [≡ Mimosa pistaciifolia Willd. (≡ Pseudalbizzia pistaciifolia (Willd.) E.J.M. Koenen & Duno)]
Albizia sect. Arthrosamanea (Britton & Rose) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 206. 1996. Type: Albizia pistaciifolia (Willd.) Barneby & J.W. Grimes [≡ Mimosa pistaciifolia Willd. (≡ Pseudalbizzia pistaciifolia (Willd.) E.J.M. Koenen & Duno)]
Pseudalbizzia berteroana (Balb. ex DC.) Britton & Rose [≡ Acacia berteroana Balb. ex DC.]
Trees or rarely treelets, unarmed. Stipules puberulent to glabrous, deltate, narrowly triangular, triangular-ovate, narrowly ovate, or narrowly lanceolate, veinless or faintly 3-veined, falling early to tardily. Leaves bipinnate, extrafloral nectaries sessile between the first pair of pinnae, near or well below mid-petiole, round, elliptic or vertically elongate, cupuliform to obsolete; pinnae (1) 2–15 (19) pairs; leaflets (2) 16–52 (63) pairs, alternate, variable in size and shape, the first pair of leaflets often reduced to paraphyllidia. Inflorescences capitate or corymbose-umbellate. Flowers heteromorphic, short-pedicellate or sessile; calyx gamosepalous; corolla gamopetalous, petals 5 (6); stamens numerous, filaments fused into a tube, stemonozone present, anthers rimose; pollen in 16-celled polyads; ovary glabrous or pubescent. Fruit lomentiform or the endocarp fragmented into 1-seeded envelopes, the valves papery, coriaceous, or glossy ligneous, straight or nearly so, decurved or plano-compressed, tardily and inertly dehiscent through both sutures or indehiscent. Seeds disciform, oblong-ellipsoid, elliptic, flattened, translucent, pleurogram present.
2n = 26 (Rico Arce 1992;
Seventeen species, in the Neotropics, from Argentina to Colombia in South America, and from north-west Mexico to Panama in North America, extending weakly into the West Indies (Fig.
Jupunba clade flowers (A–E) and fruits (F–K) A Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes B Hydrochorea uaupensis M.P. Morim, Iganci & E.J.M. Koenen C Jupunba langsdorffii (Benth.) M.V.B. Soares, M.P. Morim & Iganci D Pseudalbizzia niopoides (Spruce ex Benth.) E.J.M. Koenen & Duno E Pseudalbizzia polycephala (Benth.) E.J.M. Koenen & Duno F Pseudalbizzia inundata (Mart.) E.J.M. Koenen & Duno G Hydrochorea corymbosa H Hydrochorea uaupensis I, J Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim K Jupunba campestris (Benth.) M.V.B. Soares, M.P. Morim & Iganci. Photo credits A, G, I–K MVB Soares B, C, H J Iganci D–F RT Queiroz https://rubens-plantasdobrasil.blogspot.com/.
Humid, semi-deciduous, seasonally inundated, and seasonally dry tropical and extratropical forests and woodlands.
In reference to Albizia Durazz.
The wood of P. polycephala (Benth.) E.J.M. Koenen & Duno (‘farinha-seca’ in Brazil), is used for laminate wood floors and for internal use in civil construction (
The name Pseudalbizzia was recently resurrected mostly to accommodate species of Albizia sect. Arthrosamanea (
A new phylogenetically-based infrageneric classification, comprising five sections, was presented by
Punjuba racemiflora (Donn. Sm.) Britton & Rose [≡ Pithecellobium racemiflorum Donn. Sm.]
Shrubs and trees, unarmed. Stipules absent. Leaves bipinnate, extrafloral nectaries sessile, between the first pair of pinnae or along the leaf rachis, orbicular, patelliform, cupuliform; pinnae 1–3 pairs; leaflets 2–7 pairs, alternate, variable in size and shape. Inflorescences racemes or spiciform racemes. Flowers homomorphic, pedicellate or sessile; sepals 5, united; corolla gamopetalous, petals 5; stamens numerous, filaments fused into a tube, stemonozone present, anthers rimose; pollen in 18-grained polyads; ovary glabrous. Fruit a legume dehiscing through both margins, the valves chartaceous, curved to spiral, generally with a red endocarp, epicarp glabrous. Seeds obovate, orbicular or oblong, translucent, pleurogram absent.
2n = 26 (P. racemiflora) (
Mainly in mountain and Andean forests and secondary rainforests.
An anagram of Jupunba Britton & Rose.
Firewood.
Punjuba was described by
Balizia Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 34–35. 1996. Type: Balizia pedicellaris (DC.) Barneby & J.W. Grimes [≡ Inga pedicellaris DC. (≡ Hydrochorea pedicellaris (DC.) M.V.B. Soares, Iganci & M.P. Morim)]
Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes [≡ Mimosa corymbosa Rich.]
Shrubs and trees, unarmed. Stipules linear, linear-lanceolate, ligulate or subulate, persistent or caducous. Leaves bipinnate, extrafloral nectaries sessile between the first pair of pinnae or along the leaf rachis, orbicular, patelliform or cupuliform; pinnae 1–15 pairs; leaflets 2–33 pairs, alternate, variable in size and shape. Inflorescences corymbose-umbellate. Flowers heteromorphic, the peripheral flowers pedicellate, the terminal flower sessile; calyx gamosepalous, 5–8 toothed; corolla gamopetalous, 5–8 toothed; stamens (10) 12–30 (36), filaments fused into a tube, stemonozone present, anthers rimose; pollen in 16 or 28-celled polyads, plano-compressed disc-shaped; ovary pubescent or glabrous. Fruits straight or recurved, either lomentiform, the seeds released in one-seeded articles, or woody and indehiscent, or follicular, or crypto-lomentiform with follicular dehiscence. Seeds ovate to oblong, with pleurogram.
Unknown.
Riparian habitats, inundated and non-inundated wet tropical forests, swamp forests, seasonally inundated forests, riverbanks, mangrove swamps, and gallery forests.
From Greek, hydro (= water) and chorein (= to travel), in allusion to water dispersed propagules.
Firewood.
The genus Hydrochorea was recently revised based on phylogenomic analyses (Soares MVB et al. 2022) that confirmed Balizia as paraphyletic in relation to Hydrochorea, along with two African species formerly placed in Albizia. Soares MVB et al. (2022) suggested that rapid initial divergence between Hydrochorea and the closely related Jupunba led to extensive incomplete lineage sorting and poor phylogenetic resolution between the two genera.
Klugiodendron Britton & Killip, Ann. New York Acad. Sci. 35(3): 125. 1936. Type: Klugiodendron laetum (Benth.) Britton & Killip [≡ Pithecellobium laetum Benth. (≡ Jupunba laeta (Benth.) M.V.B. Soares, M.P. Morim & Iganci)]
Jupunba jupunba (Willd.) Britton & Rose [≡ Acacia jupunba Willd. (= Jupunba trapezifolia (Vahl) Moldenke)]
Trees, treelets and shrubs, unarmed. Stipules commonly small, narrow, lanceolate, linear, elliptic, caducous. Leaves bipinnate, petiolar nectaries sessile or stipitate (rarely) between the pairs of pinnae, or close below the proximal pair of pinna-pulvinules, campanulate to cupular, patelliform, or verruciform; pinnae 12–16 (19) pairs; leaflets 1–40 pairs, alternate, variable in size and shape. Inflorescences congested or lax racemes, rarely capitate or spiciform racemes. Flowers homomorphic or heteromorphic, pedicellate or sessile; calyx gamosepalous, 5-merous; corolla gamopetalous, 5-merous; stamens numerous, filaments fused into a tube, stemonozone present, anthers rimose; pollen in acalymmate large polyads containing 16 pollen grains, circular or elliptic, suboblate, sexine rugulate, as thick as nexine; ovary glabrous or pubescent. Fruit a legume, the valves chartaceous, curved to spiral, generally with a red endocarp, epicarp glabrous. Seeds obovate, orbicular, or oblong, translucent, pleurogram generally present, rarely absent.
2n = 26, 30 (
Riparian habitats, inundated and non-inundated wet tropical forests especially on the Amazon basin.
Sieber collected the type specimen of Acacia jupunba and recorded the name ‘jupunba’ on the sheet, probably in reference to a vernacular name.
Animal feed, firewood, folk medicine, soap making (
Jupunba is the largest genus in the Jupunba clade, and is mostly composed of species previously described under Pithecellobium, and later transferred to Abarema. The genus is closely related to Hydrochorea but is distinguished by its fruits with reddish endocarp and seeds bicoloured with translucent testa, even though those are apparently plesiomorphic characters (
Gwilym P. Lewis10
Citation: Lewis GP (2024) 35. Samanea clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 445–450. https://doi.org/10.3897/phytokeys.240.101716
Samanea clade
Figs
Included genera (2): Chloroleucon (Benth.) Britton & Rose (10 species), Samanea (Benth.) Merr. (3).
Description. Unarmed (Samanea) or usually armed with axillary spines (Chloroleucon) trees or shrubs. Stipules often lanceolate, caducous or lacking. Leaves bipinnate, extrafloral nectaries present. Inflorescences umbelliform capitula or racemose spikes, in Chloroleucon emerging from a cone of imbricate perules. Flowers in each inflorescence (occasionally) homomorphic or more commonly dimorphic with the terminal (central) flower modified and stouter, androecium with 10–36 (52) whitish, greenish, pink, or reddish stamens; pollen in polyads of 32 (Samanea) or 16–32 grains (Chloroleucon); ovary sessile or subsessile. Fruits variable, indehiscent, or tardily dehiscent, straight, falcate, coiled or randomly twisted, with or without a pulpy mesocarp. Seeds with a hard testa and a pleurogram.
Distribution. Mostly South America, more diverse in seasonally dry forests, extending northward to Central America, Mexico and the Caribbean.
Clade-based definition. The most inclusive crown clade containing Samanea saman (Jacq.) Merr. and Chloroleucon mangense (Jacq.) Britton & Rose, but not Jupunba leucophylla (Spruce ex Benth.) M.V.B. Soares, M.P. Morim & Iganci, Albizia retusa Benth. or Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes (Fig.
Notes.
Pithecellobium sect. Samanea Benth., London J. Bot. 3: 197. 1844. Type: Pithecellobium saman (Jacq.) Benth. [≡ Mimosa saman Jacq. (≡ Samanea saman (Jacq.) Merr.)]
Samanea saman (Jacq.) Merr. [≡ Mimosa saman Jacq.]
Unarmed trees (Fig.
Flower, fruit and vegetative characters of the Samanea clade A Samanea inopinata (Harms) Barneby & J.W. Grimes, tree, Brazil B, C Samanea saman (Jacq.) Merr. B inflorescences in bud and mature flower, El Salvador (Hughes 1241) C bipinnate leaves, Sri Lanka D Samanea inopinata, rough bark of trunk and main branches, Brazil E, F Samanea saman E flowers, cultivated in Hawaii F fruits G Chloroleucon mangense (Jacq.) Barneby & J.W. Grimes, trunk bark, Comayagua valley, Honduras H, I Chloroleucon acacioides (Ducke) Barneby & J.W. Grimes H fruits, Mato Grosso, Brazil I foliage and inflorescences, Brazil J Chloroleucon foliolosum (Benth.) G.P. Lewis, foliage and fruit, Bahia, Brazil (Lewis 1972). Photo credits A, D, F E Ner B, G CE Hughes C P Rajatewa E GD Carr H D Sasaki I D Cardoso J GP Lewis.
2n = 26 (Samanea saman) (
Three species, mostly circum-Amazonian in tropical continental Central and South America, native from El Salvador in Central America south eastwards through Venezuela, Colombia, Peru and Ecuador, to north-eastern Bolivia, southern, eastern and north-eastern Brazil, and Paraguay. Samanea saman is widely cultivated and partly naturalised as far north as Mexico, and long established in the West Indies in parks and gardens (Fig.
Seasonally dry deciduous to moist evergreen forest, woodland, and wooded grassland.
‘Saman’ is derived from the French Caribbean vernacular ‘zamang’ or ‘rain tree’, the leaves fold up at dusk or at the approach of storms.
Widely planted pantropically for shade (especially coffee), preserved in pastures as cattle-shade, planted as an ornamental and for nutritious fruits (for animal fodder, human food and for beverages), also for medicine and timber (for a wide range of uses from construction and panelling to furniture and veneers), and for bee forage (Lewis and Rico Arce 2005).
The species now included in Samanea were included in several different genera of the old sense tribe Ingeae.
Pithecellobium sect. Chloroleucon Benth., Lond. J. Bot. 3: 197, 221. 1844. Type: Pithecellobium vincentis Benth. [≡ Chloroleucon mangense var. vincentis (Benth.) Barneby & J.W. Grimes]
Chloroleucon vincentis (Benth.) Britton & Rose [≡ Pithecellobium vincentis Benth. (≡ Chloroleucon mangense var. vincentis (Benth.) Barneby & J.W. Grimes)]
Trees and shrubs, with dimorphic vegetative branches, the bark usually smooth and a patchwork of grey, white, and green (Fig.
2n = 26 [Ch. tenuiflorum (Benth.) Barneby & J.W. Grimes;
Ten species, north-west Mexico (one species) to the Antilles (two species), with the remaining species in South America as far south as Argentina and south-east Brazil (Fig.
Warm temperate and tropical lowland, less often in submontane seasonally dry forest, xeromorphic brush-woodland, coastal thicket, wooded grassland, shrubland and desert.
From Greek, chloro- (= greenish yellow) and leuco- (= white), probably referring to the patchwork bark of most species.
Used for construction timber, as a medicinal tea, and the fruits as human food [Ch. dumosum (Benth.) G.P. Lewis in Bahia, Brazil;
The species of Chloroleucon separate into two well supported groups based on whether the flowers in the inflorescence are homomorphic or heteromorphic (Almeida et al., in prep.).
Erik J. M. Koenen26
Citation: Koenen EJM (2024) 36. Albizia clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 451–465. https://doi.org/10.3897/phytokeys.240.101716
Albizia clade
Figs
Included genera (3). Albizia Durazz. (ca. 90 species), Boliviadendron E.R. Souza & C.E. Hughes (1), Enterolobium Mart. (8).
Description. Small to very large trees to 45 m with large spreading crown or sometimes shrubs, rarely scandent or lianescent shrubs or a liana (one species of Albizia), usually unarmed except for a few species of Albizia with spinescent shoots, prickles/hooks or recurved thorn-like stipules. Stipules usually small, linear to linear-triangular and caducous, sometimes bigger and leaf-like and persistent, or in one species recurved and thorn-like and caducous. Leaves bipinnate, micro- or macrophyllidinous, with an extrafloral nectary and usually with further glands between the pinnae and leaflets; leaflets usually asymmetrical with palmate-pinnate venation and the midrib displaced to either side of the leaflet, or, particularly in several macrophyllidinous species, symmetrical with pinnate venation. Inflorescences globose, sub- or hemiglobose, or corymbiform capitula, usually dimorphic with one or a few enlarged central flowers with broad nectariferous base and exserted staminal tube, or homomorphic, either fasciculate from leaf axils or mostly aggregated into larger paniculate compound inflorescences with leaf development suppressed. Flowers 5-merous, sessile or often especially the peripheral flowers of capitula pedicellate, with (slenderly) campanulate or funnel-shaped calyx and corolla, stamens fused into a tube for about the same length as the corolla or slightly exserted, sometimes the tube much longer and exserted well beyond the corolla lobes; pollen shed in 16- or 32-celled flat disk-shaped polyads. Fruits usually flat and straight or slightly curved non-septate, with papery valves, dehiscent or tardily dehiscent along both sutures or indehiscent, or septate reniform-auriculiform, annular or contorted indehiscent, with thick coriaceous to woody valves and then often with soft mesocarp, or lomentiform, and then sometimes breaking up into segments or tardily so, or straight to slightly curved, thick, woody and indehiscent. Seeds elliptical and laterally flattened, with a hard smooth testa and usually with open or closed pleurogram, sometimes lacking, rarely (Boliviadendron) the testa thin and the seed with a minute wing, pleurogram lacking.
Distribution. Pantropical, with the two smaller genera Boliviadendron and Enterolobium restricted to the Americas and the large genus Albizia distributed across Africa, Madagascar, tropical Asia, Malesia, Australia and some Pacific islands, and a few species extending into the temperate zone in continental Asia.
Clade based definition. The most inclusive crown clade containing Albizia julibrissin Durazz. and Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes, but not Samanea saman (Jacq.) Merr., Blanchetiodendron blanchetii (Benth.) Barneby & J.W. Grimes, Inga vera Willd. or Jupunba trapezifolia Moldenke (Fig.
Notes. The Albizia clade includes a more narrowly defined Albizia that is restricted to the Old World and its two closest relatives Boliviadendron and Enterolobium that are only found in the Americas. The clade is well supported in recent phylogenomic studies (
The clade is relatively uniform in vegetative (Figs
Habit diversity of the Albizia clade A Albizia atakataka Capuron, a densely branched small tree in dry thorn-scrub in south-western Madagascar B A. anthelmintica Brongn., a small tree flowering before the leaves appear, in semi-arid regions of southern Africa C A. corniculata (Lour.) Druce, a scandent shrub or liana in humid forests in continental South East Asia, the Philippines and northern Borneo D Albizia sp. (Koenen 434), a large tree in semi-deciduous forest in north-western Madagascar E A. tanganyicensis Baker f., a small tree in dry savanna woodland and (sub)montane grassland in southern and eastern Africa F Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes (Hughes 2608), a tree in seasonally dry tropical forest in Bolivia G Enterolobium cyclocarpum (Jacq.) Griseb. (Hughes 1254) H trunk of E. contortisiliquum (Vell.) Morong, both trees of seasonally dry and semi-deciduous forests of South and Central America. Photo credits A, D, E, H E Koenen B A Dreyer, www.africanplants.senckenberg.de C Biobank Lantauhk (Hong Kong), iNaturalist (https://www.inaturalist.org/photos/66503444) F, G CE Hughes.
General and vegetative morphology of the Albizia clade A Albizia chinensis (Osbeck) Merr., flowering branch (Koenen 190) B cultivated A. julibrissin Durazz., flowering branch C A. glaberrima Benth., foliage and inflorescences D A. zygia J.F. Macbr., flowering branch E A. polyphylla Fourn., finely bipinnate foliage (Koenen 256) F A. atakataka Capuron, dense branching with the leaves arising from brachyblasts (Koenen 229) G A. laurentii De Wild., flower buds and quadrifoliolate bipinnate leaves H A. tanganyicensis Baker f., leaf I A. gummifera C.A. Sm., detail of pinna with rhombic leaflets J Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes, foliage and flowers (Hughes 2423) K Enterolobium contortisiliquum (Vell.) Morong, foliage L E. timbouva Mart., detail of pinna. Photo credits A, E, F, H E Koenen B M Fitzgerald C G Baumann D W Dijkstra G D Harris I I Dinter J CE Hughes K A Martinez Ponte, iNaturalist (https://www.inaturalist.org/photos/195358543) L F Acaz Sonntag, iNaturalist (https://www.inaturalist.org/photos/216600814) B–D, G, I www.africanplants.senckenberg.de.
Inflorescences of the Albizia clade A Albizia grandibracteata Taub., strongly dimorphic capitula (Koenen 159) B A. mainaea Villiers of Albizia section Zygia, strongly dimorphic capitula (Koenen 426) C A. chinensis (Osbeck) Merr., more cryptic dimorphic capitula (Koenen 190) D A. lebbeck (L.) Benth., more cryptic dimorphic capitula E A. procera (Roxb.) Benth., homomorphic capitula F A. anthelmintica Brongn., dimorphic capitula G A. atakataka Capuron, dimorphic capitula (Koenen 229) H A. forbesii Benth., dimorphic capitula I A. zimmermannii Harms, homomorphic capitula J Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes, homomorphic capitula (Hughes 2423) K Enterolobium cyclocarpum (Jacq.) Griseb., homomorphic capitula (Hughes 1254). Photo credits A–C, G, H E Koenen D M Schmidt E R Cumming, iNaturalist (https://www.inaturalist.org/photos/168379143) F I Dinter I C Boucher Chisale J, K CE Hughes D, F, I www.africanplants.senckenberg.de.
Fruits of the Albizia clade A Albizia lebbeck (L.) Benth., flat papery fruits and B A. procera (Roxb.) Benth., flat papery fruits C A. moniliformis (DC.) F. Muell., lomentaceous fruits D A. ferruginea (Guill. & Perr.) Benth., flat papery fruits (Harris 9726) E A. boivinii Fourn., flat papery fruits (Koenen 376) F A. commiphoroides Capuron, moniliform fruits G of A. masikororum R. Vig., thick woody fruits H Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes, flat papery fruits (Hughes 2423) I Enterolobium contortisiliquum (Vell.) Morong, thick coriaceous ear-shaped fruits (Queiroz 15579) J E. monjollo (Vell.) Mart., thick coriaceous ear-shaped fruits (Lima 7911). Photo credits A G Baumann www.africanplants.senckenberg.de B I Cowan, iNaturalist (https://www.inaturalist.org/photos/54408314) C R Cumming, iNaturalist (https://www.inaturalist.org/photos/200251197) D D Harris www.africanplants.senckenberg.de E, I, J E Koenen F Désiré Ravelonarivo, Tropicos (https://www.tropicos.org/ImageFullView.aspx?imageid=100291301) G SE Rakotoarisoa, iNaturalist (https://www.inaturalist.org/photos/3113769) H CE Hughes.
Sassa Bruce ex J.F. Gmel., Syst. Nat., ed. 13[bis]. 2(2): 1038. 1792. Type: Sassa gummifera J.F. Gmel. [≡ Albizia gummifera (J.F. Gmel.) C.A. Sm.]
Besenna A. Rich., Tent. Fl. Abyss. 1: 253. 1847. Type: Besenna anthelmintica A. Rich. [≡ Albizia anthelmintica (A. Rich.) Brongn.]
Arthrosprion Hassk., Retzia i: 212. 1855. Type: Arthrosprion stipulatum (DC.) Hassk. [≡ Acacia stipulata DC. (= Albizia chinensis (Osbeck) Merr.)]
Cathormion (Benth.) Hassk., Retzia 1: 231. 1855. Type: Cathormion umbellatum (Vahl) Kosterm. [≡ Albizia umbellata (Vahl) E.J.M. Koenen]
Parasamanea Kosterm., Organ. Natuurw. Onderz. Indonesië 20: 11. 1954. Type: Parasamanea landakensis (Kosterm.) Kosterm. [≡ Albizia rosulata subsp. landakensis (Kosterm.) I.C. Nielsen]
Serialbizzia Kosterm., Organ. Natuurw. Onderz. Indonesië 20: 15. 1954. Type: Serialbizzia acle (Blanco) Kosterm. [≡ Albizia acle (Blanco) Merr.]
Parenterolobium Kosterm., Organ. Natuurw. Onderz. Indonesië 20: 19. 1954. Type: Parenterolobium rosulatum (Kosterm.) Kosterm. [≡ Albizia rosulata Kosterm.]
Albizia julibrissin Durazz.
Small to very large trees to 45 m with large rounded or flat-topped spreading crown or rarely shrubs, and then sometimes scandent or lianescent, or a liana to 45 m [Albizia corniculata (Lour.) Druce], usually unarmed except for a few species with spinescent shoots [A. anthelmintica Brongn., A. moniliforme (DC.) F. Muell. and A. umbellata (Vahl) E.J.M. Koenen] or prickles/hooks (A. corniculata, A. myriophylla Benth. and A. rufa Benth., these species with a scandent shrubby or lianescent habit) or recurved thorn-like stipules (in A. pedicellata Baker ex Benth.), short shoots or brachyblasts sometimes present. Stipules small linear to triangular, usually caducous, sometimes larger, rounded, leaf-like and persistent or caducous, or recurved, thorn-like and caducous in one species (A. pedicellata). Leaves bipinnate, macro- or microphyllidinous, with 1–25 pairs of pinnae, the petiole with an extrafloral nectary and usually with further nectaries between the apical pairs of pinnae and leaflets; leaflets 1–47 pairs, small and linear to large and elliptical, rounded or rhombic, frequently (especially in microphyllidinous species) the midrib displaced to either side of the leaflet blade and often several additional strong veins starting from the leaflet base (i.e., palmate-pinnate venation), in the macrophyllidinous species often with pinnate venation, base oblique to equal, apex rounded, acute or mucronate. Inflorescence units sub- or hemiglobose, often corymbiform, or sometimes globose, capitula, usually aggregated into terminal or axillary panicles with the leaves suppressed, otherwise fascicular in leaf-axils. Flowers 5-merous,mostly white or cream, or often bicoloured with pink to red upper half of stamens, sessile or often especially the peripheral flowers pedicellate, dimorphic or sometimes monomorphic, often the central flower enlarged and more robust with a broad cupular nectariferous base and longer staminal tube; calyx slenderly campanulate or funnel-shaped; corolla funnel-shaped; stamens numerous, fused into a tube not or slightly exserted beyond the corolla lobes, in terminal flowers usually well-exserted and in some species the tube very long in all flowers and exserted well beyond the corolla lobes, anthers dorsifixed, eglandular; pollen shed in 16-celled flat disk-shaped polyads (but only a few species have been studied); ovary sessile or shortly stipitate, the style approximately the same length as the stamens or extending slightly beyond the anthers, stigma funnel-shaped. Fruit usually straight, flat, papery, dehiscent or inertly dehiscent along both sutures or indehiscent, or straight or slightly curved, indehiscent and thick woody (in several Madagascan species), or straight to curved or twisted and moniliform (in A. atakataka Capuron and A. commiphoroides Capuron) or straight to twisted, septate, lomentiform, breaking up into one-seeded segments (in A. moniliforme and A. umbellata) or straight or curved, septate and coriaceous [in A. acle (Blanco) Merr.] or contorted, septate and woody [in A. rosulata (Kosterm.) I.C. Nielsen and A. dolichadena (Kosterm.) I.C. Nielsen] tardily breaking up into articles. Seeds elliptical to rounded, laterally compressed, usually with a large subcircular or narrowly elliptic closed pleurogram, or a small, rounded deltoid to subcircular pleurogram in the upper half of the seed, or sometimes a U-shaped pleurogram open towards the hilum.
2n = 26, one species (A. polyphylla Fourn.) is reported to be an octoploid with 2n = 104 (
ca. 90 species, in Africa, Madagascar, (sub)tropical Asia, Malesia, Australia and several Pacific Islands (Fig.
Albizia has a wide ecological range with species found in coastal, riverine and swamp forests, terra firme rainforest, seasonally dry deciduous forest, savannas and (sub)montane forest. Although most species of Albizia are tropical, a few Asian species, including the type species A. julibrissin Durazz., are mildly frost-tolerant and extend into the subtropical and warm temperate zone, as well as cooler (sub)montane regions. Most species are medium sized trees, in either primary vegetation or as coloniser species in secondary vegetation, with a few species occurring as large canopy trees while some of the semi-arid species remain small shrubby trees and a few Asian species are scandent shrubs to lianas at forest edges and disturbed sites or in primary and secondary forests (Fig.
Named for Filippo degli Albizzi, an 18th century Italian naturalist from the wealthy Albizzi family of Tuscany, who reportedly brought seeds of the species from Constantinople to Florence in 1749. The spelling “Albizzia” has historically been used and would be the correct spelling given the Albizzi family name, however, in the original publication the spelling Albizia was used and hence this has to be considered the correct name (
The wood of several species is used for various purposes, including house construction, furniture, boat building, musical instruments, as well as for firewood and charcoal. Several species are planted as shade trees in coffee and tea plantations and for soil improvement, and as street trees or ornamentals in parks and gardens (Fig.
Albizia was described by
Following these recent taxonomic updates, the species that remain accommodated in Albizia under the circumscription that is presented here, have been shown to form a monophyletic group in preliminary phylogenomic analyses that sampled 75 of the ca. 90 species (Koenen et al., unpubl. data;
Nearly all species of Albizia are fundamentally trees (Fig.
As in many Mimoseae genera, the bipinnate leaves vary widely in the numbers of pinna and leaflet pairs per leaf, as well as the size and shape of leaflets (Fig.
Most species have straight flat papery fruits inertly dehiscent along both sutures or indehiscent, that are sometimes slightly swollen around the seeds, but markedly different fruits are found in some Asian and Madagascan species, these placed in clades 1 and 4, respectively (Suppl. material
First, the species A. moniliformis (Indonesia to Australia) and A. umbellata (continental South East Asia) that are placed in clade 1, have septate lomentiform fruits (Fig.
Secondly, the five Asian species that
Thirdly, two unusually densely branched (Figs
Finally, seven of the ca. 24 Albizia species endemic to Madagascar, all also placed in clade 4, have thick woody indehiscent fruits (Fig.
Bentham (1985);
Boliviadendron bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes [≡ Leucochloron bolivianum C.E. Hughes & Atahuachi]
Small, unarmed multi-stemmed tree to 5–6 (10) m and 20–35 cm diameter with an irregular spreading crown; resting buds perulate, ovoid to 1.5 mm long and densely pilosulous (as for shoots), but these generally lacking. Stipules linear-triangular, caducous. Leaves bipinnate with 4–5 (6) pairs of pinnae, petiole with a circular or weakly ellipsoid cupular nectary at or slightly above mid-petiole position; similar smaller nectaries variably present between distal 1–2 pairs of pinnae or sometimes between all pinnae pairs; leaflets 15–20 pairs per pinna, slightly decrescent proximally and distally, linear-oblong, obliquely truncate at base, obtuse at apex, the tip sometimes acute, venation indistinct, palmate-pinnate brochidodromous, with 1–2 (weakly 3) primary veins at base, the main primary vein distinctly asymmetric, dividing the blade 1:2–2.5, giving rise on each side to 3–4 sinuous secondary veins, the first anterior or first pair of leaflets reduced to small paraphyllidia. Flowers white, sessile, homomorphic and arranged in lax globose capitula on slender peduncles, each capitulum with 20–25 flowers, the capitula immersed in new foliage, solitary or in fascicles of 2–3 in axils of coeval leaves; calyx narrowly campanulate, corolla tubular, slightly funnel-shaped; androecium of ca. 40 stamens, fused basally into a short tube that is not or hardly exserted beyond the corolla; pollen shed in 16-celled flat disk-shaped polyads; ovary sessile, the style held below anthers of the stamens. Fruit 1 per capitulum, sessile, (3) 4–6 (8)-seeded, flat plano-compressed, linear-oblong or oblong, obtuse at each end, sometimes with a short apiculum at apex, the valves straight or with slightly undulate margins, stiff papery, tardily dehiscent along both sutures. Seeds flat disciform, broadly suborbicular, the testa thin, lustrous, translucent castaneous, pleurogram lacking, surrounded by a narrow dark marginal nerve or minute wing.
Unknown.
Monospecific (B. bolivianum), endemic to Bolivia, where it is only known from the eastern flanks of the Andes at mid-elevations in interior Andean valleys in the Departments of La Paz, Cochabamba and Santa Cruz (
Boliviadendron is known only from the slopes of interior valleys of Bolivia between 2150 and 2770 m elevation, around the transition from seasonally dry tropical inter-Andean valley forests to more moist mid-elevation montane Ceja de Monte Yungeña vegetation, where it is locally common in fence-lines, remnant patches of subhumid or seasonally dry Yungas forest and secondary thickets (
From ‘Bolivia’ and dendron (Greek = tree), named as such for the endemicity of the genus to the country Bolivia.
Unknown.
The type species of Boliviadendron was first described as Leucochloron bolivianum C.E. Hughes & Atahuachi, but phylogenetic results of
Enterolobium timbouva Mart.
Medium to large trees to 40 m with large rounded spreading crown, unarmed, bark dark grey or brown, smooth or with lenticels. Stipules absent or small and mostly caducous. Leaves bipinnate, petiole canaliculate or angular, rachis with a pallid indumentum, petiolar nectaries present, additional nectaries sometimes on the leaf rachis and rachillae, usually sessile, globose, transversely elliptic, elliptical-oval to patelliform, paraphyllidia present or absent; pinnae 4–14 pairs; leaflets 4–20 (30) pairs per pinna, symmetrical or asymmetrical, oblong to linear-falcate, both sides glabrous or pilose, venation conspicuous on both sides, palmate, palmate-pinnate, rarely dimidiate-palmate. Inflorescences pedunculate globose capitula, in fascicles or aggregated into pseudoracemes, homomorphic or heteromorphic. Flowers 5-merous, white, sessile to pedicellate, calyx and corolla tubular or campanulate, petals and sepals outside with a pallid, puberulent, rarely sericeous, strigose-pilose indumentum or glabrous; stamens 20–68 per flower, monadelphous, staminal tube included or exserted, intrastaminal disc usually absent, anthers dorsifixed, longitudinal; pollen shed in 16- or 32-celled flat disk-shaped polyads; ovary at anthesis glabrous, pubescent after fertilization. Fruits reniform-auriculiform or annular, rarely contorted, thick coriaceous with a septate papery endocarp, an either dry-mealy or resinous-pulpy mesocarp, and a glabrous or pilose thin exocarp with smooth surface. Seeds uniseriate or biseriate, ovoid, ellipsoid or obovoid, pleurogram present or absent, fissure line present or absent.
2n = 26 (
Eight species, from southern Mexico and the Greater Antilles to northern Argentina (
Lowland and submontane tropical rainforests in Amazonia (especially Venezuela and Brazil) and the Atlantic forest of Brazil, in semi-deciduous and seasonally dry tropical forests from southern Mexico and the Greater Antilles to northern Colombia and in southern Brazil, northern Argentina, Bolivia, Paraguay and Uruguay, as well as in the Brazilian Caatinga and Cerrado biomes.
From Greek, entero (= intestine) and lobion (= pod), referring to either the shape or contents of the fruit.
The timber of several species is used for high quality furniture, cabinet work, joinery, panelling, veneers and water resistant construction. Also often planted as shade trees, ornamentals or for pasture improvement and livestock fodder (although the fruits of some species are toxic to cattle). Further uses include for fibre (paper), gum extraction and the bark is used as a soap substitute and medicinally (Lewis and Rico Arce 2005).
Enterolobium was described by
João Iganci20,25, Ethiéne Guerra20, Marli Pires Morim8, Ana Carla da Silva Oliveira2, Andrés Fonseca-Cortés2, Carolina Lima Ribeiro2, Felipe da Silva Santos2, Filipe Gomes Oliveira2, Julia Ferm15, Luciano Paganucci de Queiroz2
Citation: Iganci J, Guerra E, Morim MP, Oliveira ACS, Fonseca-Cortés A, Ribeiro CL, Santos FS, Oliveira FG, Ferm J, Queiroz LP (2024) 37. Inga clade. In: Bruneau A, Queiroz LP, Ringelberg JJ (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae. Part 2: Higher-level classification. PhytoKeys 240: 466–486. https://doi.org/10.3897/phytokeys.240.101716
Inga clade
Figs
Included genera (8). Abarema Pittier (2 species), Blanchetiodendron Barneby & J.W. Grimes (1), Inga Mill. (ca. 300), Leucochloron Barneby & J.W. Grimes (4), Macrosamanea Britton & Rose ex Britton & Killip (12), Osodendron E.J.M. Koenen (3), Robrichia (Barneby & J.W. Grimes) A.R.M. Luz & E.R. Souza (3), Zygia P. Browne (ca. 60).
Description. Unarmed trees or shrubs, rarely with spine-like projections (some Osodendron); axillary perulate resting buds absent or present and then with imbricate and striate perules. Stipules mostly caducous. Leaves bipinnate or paripinnate, with glands on the petiole and/or leaf rachis and then between the pinnae or leaflets insertion. Inflorescences capitate, umbelliform, spicate or racemose, isolated or clustered in axillary fascicles or terminal panicles. Flowers mostly 5-merous; calyx gamosepalous; corolla gamopetalous; stamens basally fused in a staminal tube, exserted or included in the corolla; pollen in 16, 18, 20, 24, 32 (35)-celled polyads; gynoecium 1 or pluricarpellate. Fruits indehiscent, and then straight or encircled, or passively dehiscent through one and both margins and then with mostly papery or chartaceous valves, rarely lomentiform and breaking up in 1-seeded articles (Osodendron). Seeds flattened and discoid, ovoid, obovoid, oblong, elliptic to orbicular, transverse, narrowly winged in Blanchetiodendron and Leucochloron, slightly compressed and bicoloured with testa partially translucent, showing the dark bluish cotyledons in Abarema, slightly compressed or terete in cross section, fleshy, thin-walled, the testa developing a thick white sugary sarcotesta in Inga; pleurogram U-shaped, O-shaped or absent.
Distribution. New World and Africa. The clade is especially diverse in American wet forests, mostly from northern South America in the Amazon basin, extending northward to the Guiana shield and to Mexico and the Antilles, and eastern South America extending to northern Argentina, Paraguay, Uruguay and Bolivia. Also frequent in the riparian forests across the Neotropics. Blanchetiodendron and Leucochloron are more characteristic of seasonally dry forests and woodlands of eastern and central Brazil. Osodendron occurs disjunctly in West and Central Africa, mostly in wet forests.
Clade-based definition. The most inclusive crown clade containing Blanchetiodendron blanchetii (Benth.) Barneby & J.W. Grimes and Inga vera Willd., but not Albizia retusa Benth., Samanea saman (Jacq.) Merr. or Punjuba racemiflora (Donn. Sm.) Britton & Rose (Fig.
Notes. The Inga clade brings together a heterogeneous set of ingoid legume genera that were placed in disparate groups in the taxonomic system of
The clade is morphologically diverse in several aspects and presents some traits rarely found in tribe Mimoseae, such as paripinnate leaves [the entire genus Inga and Zygia inundata (Ducke) H.C. Lima ex Barneby & J.W. Grimes] and a polycarpellate gynoecium (some species of Inga and Macrosamanea). Inflorescence units are compact capitate or umbelliform in most genera but spicate or racemose inflorescences are also common. Fruits are very diverse and sometimes diagnostic for particular genera. Indehiscent legumes with a fleshy tissue (sarcotesta) around the seed are characteristic of Inga. Robrichia also presents indehiscent fruits but they have an encircled body like an ear. Lomentiform fruits breaking up into one-seeded articles are found in Osodendron. The remaining genera have dehiscent fruits opening through one or both margins.
Generic taxonomy of the Inga clade has been shaken by recent phylogenetic studies of the mimosoid legumes that demonstrated the rampant non-monophyly of numerous genera (
Blanchetiodendron blanchetii (Benth.) Barneby & J.W. Grimes [≡ Enterolobium blanchetii Benth.]
Unarmed trees; perulate buds at some leaf axils, the perules imbricate and striate. Stipules absent. Leaves bipinnate; petiolar nectary near or above mid-petiole, sessile, cupular or shallow-cupular; pinnae 2–4 pairs; leaflets 8–15 pairs, (sub)opposite, variable in size and shape. Inflorescence units capitate or umbelliform, grouped in axillary pseudoracemes or terminal panicles (Fig.
Foliage, inflorescence and fruits in Inga clade A, B Blanchetiodendron blanchetii (Benth.) Barneby & J.W. Grimes A flowering branch showing the capitate inflorescences clustered in efoliate pseudoracemes B detail of the heteromorphic capitate inflorescence C Leucochloron limae Barneby & J.W. Grimes leafless branch showing homomorphic capitate inflorescences and follicle fruits D Abarema diamantina E. Guerra, Iganci & M.P. Morim with axillary homomorphic capitate inflorescences E, F Robrichia schomburgkii (Benth.) A.R.M. Luz & E.R. Souza E young branch with the characteristic rusty pubescent indumentum F detail of a homomorphic capitate inflorescences G Osodendron altissimum (Hook. f.) E.J.M. Koenen detail of a homomorphic capitate inflorescence H, I Macrosamanea amplissima (Ducke) Barneby & J.W.Grimes H flowering branch with a homomorphic capitate inflorescence I bracts with showy extrafloral nectaries and visiting ants. Photo credit: A, B, D D Cardoso C GP Lewis E G Perez Huertas F F Oviedo-Brenes G E Bidaut H, I J Iganci.
Fruit diversity in Inga clade A young pod of Blanchetiodendron blanchetii (Benth.) Barneby & J.W. Grimes B, C Leucochloron incuriale (Vell.) Barneby & J.W. Grimes B dehisced follicular fruits exposing the seeds (C) D lomentiform fruit of Osodendron altissimum (Hook. f.) E.J.M. Koenen E indehiscent and internally septate pod of Osodendron leptophyllum (Harms) E.J.M. Koenen F auriculiform indehiscent pod of Robrichia schomburgkii (Benth.) A.R.M. Luz & E.R. Souza G young fruit of Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes H dehisced twisted pod of Abarema diamantina E. Guerra, Iganci & M.P. Morim exposing the reddish endocarp and bicoloured seeds I dehisced follicle of Macrosamanea amplissima (Ducke) Barneby & J.W.Grimes J Young fruits of Zygia brenesii (Standl.) L. Rico K dehisced pods of Zygia cognata (Schltdl.) Britton & Rose L–O Indehiscent pods of Inga, with fleshy and sweety sarcotesta around the seeds L Inga grazielae (Vinha) T.D. Penn. M Inga capitata Desv. N Inga ingoides (Rich.) Willd. O Inga edulis Mart. Photo credits A, H LP Queiroz B NA Escobar C J Vieira D D Harris E P Latham, www.africanplants.senckenberg.de F LCampbell G, M, N RT Queiroz http://rubens-plantasdobrasil.blogspot.com/ I J Iganci J F Chinchila Romero K S de Jesus Calva L, O D Cardoso.
Unknown.
Monospecific (B. blanchetii), endemic to north-eastern Brazil mostly from central Bahia state, extending to northern Minas Gerais and southern Piauí (Fig.
Seasonally deciduous and semi-deciduous forests and woodlands from the Caatinga and Atlantic Forest phytogeographical domains, from 450 to 1000 m elevation (
The genus is named after the Swiss naturalist Jacques Samuel Blanchet, and dendron (Greek = tree).
Unknown.
The genus Blanchetiodendron was described as a monotypic genus by
Leucochloron incuriale (Vell.) Barneby & J.W. Grimes [≡ Mimosa incurialis Vell.]
Unarmed trees (Fig.
Habitat and habit diversity in Inga clade A Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes, arborescent shrub in sandy coastal scrub B Robrichia schomburgkii (Benth.) A.R.M. Luz & E.R. Souza, buttressed emergent tree in rainforests C Leucochloron incuriale (Vell.) Barneby & J.W. Grimes, tree with thick corky bark in semi-deciduous forests D Inga bifoliolata D.B.O.S. Cardoso & Amorim, scandent treelet of rainforest understory E Zygia juruana (Harms) L. Rico treelet of the Amazonian inundated forests (igapó) F Osodendron dinklagei (Harms) E.J.M. Koenen, giant tree from the Guinean forests in West tropical Africa. Photo credits A RT Queiroz https://rubens-plantasdobrasil.blogspot.com/ B G Perez Huertas C F Ventura D, E D Cardoso F W Hawthorne.
Unknown.
Four species, probably endemic to Brazil, occurring along eastern Atlantic coastal states (Bahia to Paraná) and inland to planaltine Minas Gerais, and Distrito Federal (Fig.
Seasonally dry forests, campo, caatinga, open woodland, bush-islands in campo, and in cerrado-gallery ecotone and at elevations of 710–1500 m. Leucochloron minarum (Glaz. & Harms) Barneby & J.W. Grimes and L. limae Barneby & J.W. Grimes are endangered species (
An anagram of Chloroleucon.
Leucochloron incuriale wood used in luxury furniture, parquet flooring, interior decoration, sheets for plywood coverings, wainscoting; civil construction, such as rafters, frames, slats, floorboards; in rural construction and in external works, such as sleepers, stakes, fence posts, and beams (Campos Filho and Sartorelli 2015).
The genus Leucochloron was proposed by
Pithecellobium sect. Abaremotemon Benth., London J. Bot. 3: 203. 1844. Lectotype (designated by Britton & Killip, 1936): Pithecellobium auaremotemo Mart. [= Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes]
Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes [≡ Mimosa cochliocarpos Gomes]
Unarmed shrubs and trees (Fig.
Unknown.
Two species, A. cochliacarpos and A. diamantina E. Guerra, Iganci & M.P. Morim, endemic to Brazil, along the Atlantic Forest, in coastal south-east and north-east Brazil, from São Paulo to Ceará states. In north-east Brazil the genus also occurs in the Caatinga, in inland Bahia state (Fig.
Sandy soil in Caatinga (Bahia state) with caatinga and cerrado vegetation elements (carrasco), ombrophilous forest, semi-deciduous forest and coastal scrub (restinga) (
From Tupi-Guarani “abaré” – priest; “motimbora” – make smoke (
Abarema cochliacarpos, usually known as “barbatimão” in north-east Brazil, is rich in phenolic secondary compounds. The bark is used in infusions to treat ulcers, sores, gastritis, inflammation, leukorrhea, and vaginal discharge (
The first mention of Abarema is attributed to
Abarema cochliacarpos is widely distributed throughout the geographic range of the genus, and has a high degree of morphological variation across its distribution (
Enterolobium sect. Robrichia Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 249. 1996. Type: Enterolobium schomburgkii (Benth.) Benth. [≡ Pithecellobium schomburgkii Benth.]
Robrichia schomburgkii (Benth.) A.R.M. Luz & E.R. Souza [≡ Pithecellobium schomburgkii Benth.]
Trees 8–50 m high (Fig.
Unknown.
Three species known to date, R. glaziovii (Benth.) A.R.M. Luz & E.R. Souza, R. oldemanii (Barneby & J.W. Grimes) A.R.M. Luz & E.R. Souza and R. schomburgkii (
Robrichia species are large trees extending to the rainforest canopy.
The name of this genus is composed of the initial parts of two names “Rob” and “Rich”, which
In Brazilian Amazonia, Robrichia schomburgkii is used for civil and naval construction (Le Cointe 1947); in Oaxaca, Mexico this species is also used in teas to relieve stomach ache (
Osodendron altissimum (Hook. f.) E.J.M. Koenen [≡ Albizia altissima Hook. f.]
Trees, rarely shrubs, 5–35 m tall (Fig.
2n = 26 (
Three species, O. altissimum, O. dinklagei (Harms) E.J.M. Koenen and O. leptophyllum (Harms) E.J.M. Koenen, in tropical West and Central Africa, from Senegal and Guinea Bissau to Zambia and Angola (Fig.
Osodendron is found mainly in tropical wet forests extending to the savanna belt. Osodendron dinklagei and O. leptophyllum can also be found in swampy areas, forest edges or gallery forest (
The word “oso” refers to a recipe from African cuisine, mainly from the area of Ghana and Guinea, which uses O. altissimum seeds as a base for preparation of a local fermented food (Jolaoso 2014).
Osodendron is important for food in Africa, where O. altissimum seeds are used in cooking and human nutrition (Jolaoso 2014). The leaves of this species also have anti-inflammatory and analgesic properties, confirmed by phytochemistry and animal testing (
The species currently assigned to Osodendron were placed in Cathormion (Benth.) Hassk. and Albizia, but the phylogenomic study of
Osodendron species are restricted to Africa, whereas Robrichia species have a Neotropical distribution, and they can be differentiated from Robrichia mainly by the shape of their fruits, which are straight, slightly curved, coiled and/or twisted (vs. contorted or “ear-shaped” in Robrichia) (
Jolaoso (2014);
Pithecellobium ser. Coriacea Benth., Trans. Linn. Soc. 30: 589. 1875. Type: Pithecellobium adiantifolium (Kunth) Benth., nom. illeg. [≡ Macrosamanea discolor (Willd.) Britton & Killip]
Macrosamanea discolor (Humb. & Bonpl. ex Willd.) Britton & Rose ex Britton & Killip. [≡ Inga discolor Humb. & Bonpl. ex Willd.]
Unarmed shrubs (sometimes semi-scandent), trees or lianas. Stipules caducous. Leaves bipinnate, extrafloral nectaries absent or present in basal or median region of petiole, sometimes between pairs of pinnae, sessile, elliptic, patelliform, scutiform, round; pinnae 1–17 pairs; leaflets 2–31 pairs, opposite, variable in size and shape. Inflorescence units capitate or spicate, congested, lax, or umbelliform, axillary or terminal, rarely cauliflorous (Fig.
Unknown.
Riparian habitats, and in forests and open areas subject to seasonal flooding in the Amazon basin (savanna or “campinarana”).
From Greek, macro (= great), and Samanea (from the aboriginal samán = rain tree).
Unknown.
Macrosamanea was described by
Pithecellobium sect. Caulanthon Benth., London J. Bot. 3: 197. 1844. Type: Pithecellobium unifoliolatum Benth. [≡ Zygia unifoliolata (Benth.) Pittier]
Calliandra sect. Caulanthon (Benth.) Griseb., Fl. Brit. W. I.: 225. 1864. Type: Calliandra latifolia (L.) Griseb. [≡ Zygia latifolia (L.) Fawc. & Rendle]
Marmaroxylon Killip in Record, Trop. Woods 63: 3. 1940. Type: Marmaroxylon racemosum (Ducke) Killip [≡ Pithecellobium racemosum Ducke (≡ Zygia racemosa (Ducke) Barneby & J.W. Grimes)]
Pithecellobium sect. Callozygia Barbosa, Caldasia 14: 400. 1986. Type: Pithecellobium lehmannii Harms [≡ Zygia lehmannii (Harms) Britton & Rose]
Zygia sect. Callozygia (Barbosa) L. Rico, Kew Bull. 46: 495. 1991. Type: Zygia lehmannii (Harms) Britton & Rose [≡ Pithecellobium lehmanii Harms]
Zygia sect. Macrophylla L. Rico, Kew Bull. 46: 495. 1991. Type: Zygia macrophylla (Spruce ex Benth.) L. Rico [≡ Pithecellobium macrophyllum Spruce ex Benth.]
Zygia latifolia (L.) Fawc. & Rendle [≡ Mimosa latifolia L.]
Unarmed shrubs or small trees, rarely reaching 20 m height, rarely lianas (Fig.
Leaf and inflorescence diversity in Zygia and Inga (Inga clade) A Zygia juruana (Harms) L. Rico, dense cauliflorous capitate inflorescences B Zygia selloi (Benth.) L. Rico with elongate cauliflorous spikes C part of a leaf of Inga ingoiodes (Rich.) Willd. showing the winged leaf rachis and extrafloral nectaries between leaflet insertions D dense pyramidal spike of Inga grazielae (Vinha) T.D. Penn. E elongate cylindrical spikes of Inga marginata Willd. F pedunculate capitate inflorescence of Inga cordistipula Mart. G terminal panicle of Inga vera Kunth. Photo credits A, D, F D Cardoso B M Magenta C, E, G RT Queiroz https://rubens-plantasdobrasil.blogspot.com/.
Unknown.
ca. 60 species from the Neotropics, distributed from Mexico to north-east Argentina, including the Caribbean, with the greatest diversity in Central America, Colombia, the Guianas and north-west Amazon (Fig.
Typically plants from wet forest understories, occurring in riparian forests and coastal habitats. They can extend from sea level mangroves up to 750 m elevation, in a few cases reaching 2600 m in northern Andean forests (
From the Greek Zygo referring to the single pair of joined pinnae and stamen filaments that are partially fused (Lewis and Rico Arce 2005).
As nitrogen fixers, Zygia species are used for recovery of degraded areas, and they are used for carpentry in civil constructions and furniture making (
Zygia was described by
The taxonomic limits of Zygia and Marmaroxylon have been controversial, the two having been considered synonyms (
Phylogenetic studies also demonstrate the polyphyly of Zygia (including Marmaroxylon), but the taxonomy still needs to be updated (Ferm et al. in prep.). The non-cauliflorous and multipinnate-leaved Zygia (= Marmaroxylon) ocumarensis is resolved as sister to Macrosamanea (
Affonsea A. St.-Hil., Voyage Distr. Diamans Brés, 1: 385. 1833. Type: Affonsea juglandifolia A. St.-Hil. [≡ Inga globularis T.D. Penn.]
Feuilleea Kuntze, Revis. Gen. Pl. 1: 182. 1891. Type: Feuilleea inga (L.) Kuntze [≡ Mimosa inga L. (≡ Inga vera Willd.)]
Inga vera Willd. [≡ Mimosa inga L.]
Unarmed trees or treelets to ca. 40 m high, with a smooth cylindrical bole (Fig.
2n = 26, and in some tetraploid species 2n = 52 (
ca. 300 species, restricted to tropical America. The species are distributed in Mexico, southern Central America, western South America, Venezuela and the Guianas, the coastal states of Brazil and the West Indies. The highest species diversity is concentrated in the Andean foothills of Peru, Ecuador, Colombia and in southern Central America, occupying a wide variety of habitats from sea level to 3000 m (
The genus is largely confined to wet forests, having a strong relationship with the germination process of the seeds, as they need shade and high humidity for immediate germination (
Derived from “ingá”, the Brazilian name of several species of mimosoid legumes, especially of the genus Inga. Originated from the Tupi word in-gá, which probably means “soaked, stewed”, due to the consistency of the pulp that surrounds the seeds (
The genus has many uses. The fruits are edible, and the fleshy sarcotesta surrounding the seed can be eaten fresh or in recipes (Fig.
Inga has been supported as monophyletic in several phylogenetic studies of the mimosoid legumes although only about one third of the species have been sampled (
The infrageneric taxonomy has changed importantly over the years.
We thank Andrew Schnabel for discussion of the native distribution of Gleditsia triacanthos, Russell Barrett and Rafaël Govaerts for information on the nomenclature of Erythrophleum, Ellie Becklund for information about Neptunia, Ilia Leitch and Gustavo Souza for chromosome data for Caesalpinieae, David Mabberley for the suggested etymology of the genus Biancaea, Mauricio Diazgranados for help with the preparation of the photograph figures for the Ceratonieae, Campsiandreae and Samanea clade treatments, Ana Flávia T. Duarte, Luis Carlos Casas Restrepo and Francisco de Assis R. dos Santos for sharing pollen grain photos for the glossary, and Braam van Wyk for excellent photographs of South African legumes. We are deeply grateful to all those who generously permitted use of their photographs throughout the monograph.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This project was funded by multiple sources, including the Natural Sciences and Engineering Research Council of Canada, Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq/Brazil (Processes 305230/2021-2, 311847/2021-8), Swiss National Science Foundation grants (310003A_156140 and 31003A_182453/1) and Early.Postdoc.Mobility (P2ZHP3_199693) and Postdoc.Mobility (P500PB_211111) fellowships, Claraz Schenkung Foundation of Switzerland, and the University of Guam Center for Island Sustainability and Sea Grant.
Authors contributed different taxonomic treatments to this monograph. Contributions are noted in each treatment.
Anne Bruneau https://orcid.org/0000-0001-5547-0796
Luciano Paganucci de Queiroz https://orcid.org/0000-0001-7436-0939
Jens J. Ringelberg https://orcid.org/0000-0003-0567-5210
Leonardo M. Borges https://orcid.org/0000-0001-9269-7316
Roseli Lopes da Costa Bortoluzzi https://orcid.org/0000-0002-7445-7244
Gillian K. Brown https://orcid.org/0000-0002-7940-5435
Domingos B. O. S. Cardoso https://orcid.org/0000-0001-7072-2656
Ruth P. Clark https://orcid.org/0000-0001-9974-2933
Adilva de Souza Conceição https://orcid.org/0000-0002-8800-422X
Matheus Martins Teixeira Cota https://orcid.org/0000-0003-0654-7501
Else Demeulenaere https://orcid.org/0000-0002-1815-3051
Rodrigo Duno de Stefano https://orcid.org/0000-0003-1707-4121
Julia Ferm https://orcid.org/0000-0002-8762-3942
Andrés Fonseca-Cortés https://orcid.org/0000-0001-7207-9940
Edeline Gagnon https://orcid.org/0000-0003-3212-9688
Rosaura Grether https://orcid.org/0000-0003-2673-665X
Ethiéne Guerra https://orcid.org/0000-0002-9495-1717
Elspeth Haston https://orcid.org/0000-0001-9144-2848
Patrick S. Herendeen https://orcid.org/0000-0003-2657-8671
Héctor M. Hernández https://orcid.org/0000-0002-1741-5515
Helen C. F. Hopkins https://orcid.org/0000-0003-4984-8224
Isau Huamantupa-Chuquimaco https://orcid.org/0000-0002-4153-5875
Colin E. Hughes https://orcid.org/0000-0002-9701-0699
Stefanie M. Ickert-Bond https://orcid.org/0000-0001-8198-8898
João Iganci https://orcid.org/0000-0002-5740-3666
Erik J. M. Koenen https://orcid.org/0000-0002-4825-4339
Gwilym P. Lewis https://orcid.org/0000-0003-2599-4577
Haroldo Cavalcante de Lima https://orcid.org/0000-0003-2154-670X
Alexandre Gibau de Lima https://orcid.org/0000-0002-9168-2507
Melissa Luckow https://orcid.org/0009-0007-2543-0516
Brigitte Marazzi https://orcid.org/0000-0003-3252-5816
Bruce R. Maslin https://orcid.org/0000-0002-3039-0973
Matías Morales https://orcid.org/0000-0001-5540-9725
Marli Pires Morim https://orcid.org/0000-0003-0872-8429
Daniel J. Murphy https://orcid.org/0000-0002-8358-363X
Shawn A. O'Donnell https://orcid.org/0000-0003-0731-7425
Filipe Gomes Oliveira https://orcid.org/0000-0003-0244-32620
Ana Carla da Silva Oliveira https://orcid.org/0000-0001-7042-5360
Juliana Gastaldello Rando https://orcid.org/0000-0002-3714-8231
Pétala Gomes Ribeiro https://orcid.org/0000-0002-0070-9971
Carolina Lima Ribeiro https://orcid.org/0000-0001-9508-2894
Felipe da Silva Santos https://orcid.org/0000-0002-1068-0578
David S. Seigler https://orcid.org/0009-0003-5177-5893
Guilherme Sousa da Silva https://orcid.org/0000-0002-4250-0017
Marcelo F. Simon https://orcid.org/0000-0002-5732-1716
Marcos Vinícius Batista Soares https://orcid.org/0000-0003-2660-1771
Vanessa Terra https://orcid.org/0000-0001-5669-1304
All of the data that support the findings of this study are available in the main text or Supplementary Information. Occurrence datasets are available at https://zenodo.org/doi/10.5281/zenodo.8407862.
Methods for distribution maps
Data type: pdf
Explanation note: text with one table, and references.
Phylogeny of Caesalpinioideae including all accessions
Data type: pdf
Explanation note: figure S1. Phylogeny of Caesalpinioideae including all accessions. Branch lengths are expressed in coalescent units, terminal branches were assigned an arbitrary uniform length for visual clarity. Tribes (underlined) and Mimoseae clades are labeled. Full details about phylogeny and tree file are available in
Phylogeny of Caesalpinioideae including all accessions showing full gene tree support
Data type: pdf
Explanation note: figure S2. Phylogeny of Caesalpinioideae including all accessions showing full gene tree support. Pie charts show the fractions of supporting and conflicting gene trees per node (blue representing supporting gene trees, green gene trees supporting the most common alternative topology, red gene trees supporting further alternative topologies, grey gene trees uninformative for this node), and numbers above pie charts are Internode Certainty All support values [both calculated with PhyParts (