1. Family: Fabaceae Lindl.
    1. Hymenaea L.

      1. This genus is accepted, and its native range is Mexico to Tropical America, Kenya to Mozambique, W. Indian Ocean.

    [LOWO]

    Legumes of the World. Edited by G. Lewis, B. Schrire, B. MacKinder & M. Lock. Royal Botanic Gardens, Kew. (2005)

    Note

    The Detarieae sens. lat. are pantropical in distribution, with c. 58% of the genera confined to Africa (incl. Madagascar), c. 20% to the Neotropics, and c. 12% to tropical Asia. Only Copaifera, Crudia and Cynometra are pantropical (and all possibly non monophyletic) and Afzelia, Guibourtia, Hymenaea, Intsia and Sindora are native to at least two of these regions. The apparent high level of diversity in the African tropics may in part be an artefact of the (relatively) greater taxonomic effort that has been invested in the study of the African taxa. Characteristic of African Detarieae taxonomy has been the splitting off of disparate elements as segregate genera, while this has often not been the case in the Neotropics. Both regions, however, currently contain large paraphyletic assemblages requiring detailed species-level analysis. Eighty-two genera and from (729) – 747 – (765) species are treated here in Detarieae sens. lat. (Fig. 21). Of the 132 (extant) species so far assessed for IUCN red data status, 97 have categories of threat. Of these 73 are assessed as vulnerable, 13 are endangered and 11 are critically endangered.

    The remarkable range and complex patterns of floral modifications found in the Detarieae sens. lat. have proved a considerable challenge to the establishment of widely accepted and clearly circumscribed generic groupings. Based on the work of Léonard (1957) and Cowan & Polhill (1981a & b), ten informal groups of genera were proposed; the Cynometra, Hymenostegia, Hymenaea, Crudia, Detarium and Brownea groups in tribe Detarieae, and the Berlinia, Macrolobium, Amherstia and Brachystegia groups in tribe Amherstieae. Genera with imbricate bracteole aestivation were assigned to Detarieae whilst those with valvate bracteole aestivation were placed in Amherstieae. Polhill (1994) retained these generic groupings with a few additions to accommodate recently described genera, and merged the two tribes into a single broadly defined tribe, Detarieae sens. lat. Breteler (1995) proposed the recognition of two tribes within Detarieae sens. lat., separated according to the relative size and position of the paired bracteoles before anthesis. Essentially, this resulted in the reassignment of Cowan & Polhill’s Amherstia group genera (Amherstia, Tamarindus and Humboldtia) from Amherstieae to Detarieae with the remaining genera forming tribe Macrolobieae. In 1999, Breteler (pers. comm.) proposed a modified Breteler (1995) tribal system in which Macrolobieae was maintained, the circumscription of Detarieae was greatly narrowed and the genera newly excluded from Detarieae were together recognised as Cynometreae sens. strict.

    The first comprehensive studies of phylogenetic relationships in tribe Detarieae sens. lat., were the analyses of Bruneau et al. (2000; 2001), based on nucleotide sequence data from the chloroplast trnL intron. They found that tribes Detarieae and Macrolobieae formed a well supported monophyletic group, which included all genera placed previously in Detarieae sens. lat., except Umtiza. Bruneau et al. (2000) examined 71 genera, with the African taxa most widely sampled. The key results of the analysis were that none of the generic groupings proposed by Cowan and Polhill (1981a & b) and Polhill (1994) were supported as strictly monophyletic, and the majority of the members of tribe Macrolobieae (although not Macrolobium) were placed as a monophyletic group derived within Detarieae sens. lat. These analyses also repeatedly recognised a second group of related genera made up entirely of resin-producing taxa, with the exception of some species of Guibourtia. The resins can be seen as translucent gland dots in the leaflets and (sometimes) other organs. Within the resin producing taxa, the genus Prioria and several members of Cowan & Polhill’s Crudia group were consistently placed together (see taxonomic notes under individual genera in main text). Another subclade within the resin-producing Detarieae comprising six members of Polhill’s Detarium group was repeatedly recognised (Bruneau et al., 2000; 2001; Fougère-Danezan et al., 2003). In addition, in the trnL intron analysis, the five sampled members of Cowan and Polhill’s Brownea group were consistently placed together with Macrolobium, although this grouping was not upheld in a more recent molecular and combined molecular-morphological analysis (Herendeen et al., 2003a). Within the exclusively African Macrolobieae of Bruneau et al. (2001), a well supported subclade of six genera was recognised by Gervais & Bruneau (2002) and as the ‘babjit’ clade sensu Wieringa & Gervais (2003). Bruneau et al. (2000; 2001) confirmed the view of Polhill (1994) that the two tribes Detarieae and Macrolobieae (sensu Breteler, 1995) are best considered a single entity. Evidence from ontogenetic studies by Tucker (2000, 2001, 2002a) challenged the validity of bracteole aestivation as a criterion for subdividing Detarieae sens. lat. and identified a set of character states associated with two modes of floral development (Circular and Omega) whose distribution amongst detarioid genera does not support Polhill’s groups.

    The analyses of Herendeen et al. (2003a) united a morphological dataset with the chloroplast trnL intron dataset of Bruneau et al. (2001). The combined analysis provided mixed results within Detarieae sens. lat. compared with the molecular dataset alone. Near the base of the clade, and within some subclades, greater resolution was provided but as several groups were not well supported, it would be premature to emphasise the details of this greater resolution. Elsewhere the addition of morphological characters produced weaker resolution and a less robust phylogeny due either to conflicting phylogenetic signal or increased homoplasy in the morphological data or both. The order of taxa followed here (Fig. 21) represents a synthesis of the present understanding of putative relationships within this (perhaps most morphologically diverse) tribe in the Leguminosae. Unsampled genera in the combined analysis are inserted into this order where morphological evidence appears to suggest close relationships.

    Whilst significant progress has been made since Polhill (1994), further studies (particularly including the non-African members of the larger and less well understood genera) are needed before a new comprehensive classification of the Detarieae sens. lat., based on a synthesis of all available data, can be established.

    Moderately well supported as sister to Guibourtia by Herendeen et al. (2003a) although sampling incomplete; Poinar (1991) described a fossil species (H. protera Poinar), from the Dominican Republic which he considered closely resembled the African H. verrucosa Gaertn.
    Vernacular
    jatoba, courbaril, guapinol, algarrobo, Zanzibar copal, S American copal
    Habit
    Trees or shrubs
    Ecology
    Tropical riverine and inundated forest to seasonally dry forest, woodland, thorn forest, bushland and thicket, often on slopes
    Distribution
    tropical S America with centres in Amazonia and coastal Atlantic forest (c. 6 spp.) and dryland NE to SE Brazil (5 spp.; one of which extends to Paraguay and Argentina); 1 sp. widespread in the Neotropics (extending to C America, the Caribbean and Mexico); 1 sp. endemic to Cuba, allied to a single species in coastal E Africa, Madagascar and the Mascarene Islands
    [FZ]

    Leguminosae, R.K. Brummitt, A.C. Chikuni, J.M. Lock & R.M. Polhill. Flora Zambesiaca 3:2. 2007

    Habit
    Trees, evergreen, unarmed.
    Leaves
    Leaves with a single pair of leaflets, with pellucid gland dots; petiolules short, twisted; stipules small, early caducous.
    Flowers
    Flowers in panicles, spirally arranged on the ultimate axes; bracts concave, caducous; bracteoles 2, enclosing the young bud, imbricate, caducous.
    Hypanthium
    Hypanthium short.
    Calyx
    Sepals 4, unequal (2 + 2), imbricate.
    Corolla
    Petals 3 large and 2 minute; sometimes all 5 large.
    Stamens
    Stamens 10; anthers dorsifixed, dehiscing by longitudinal slits.
    Pistil
    Ovary stipitate; ovules usually 4; style elongated, filiform; stigma terminal, small.
    Fruits
    Pods ovoid or ellipsoidal, the epicarp hard with numerous prominent resin-filled vesicles, indehiscent; endocarp pithy-powdery.
    Seeds
    Seeds 1–3, without areole or aril.
    [FTEA]

    Leguminosae, J. B. Gillett, R. M. Polhill & B. Verdcourt. Flora of Tropical East Africa. 1971

    Habit
    Unarmed evergreen tree
    Leaves
    Leaves with a single pair of leaflets; stipules lateral, free, very small, very quickly falling off; leaflets opposite, with numerous pellucid gland-dots
    Inflorescences
    Inflorescence paniculate; flowers spirally arranged along the racemose ultimate branches; pedicels jointed at base; bracteoles large, well-developed, concavo-convex, ± completely concealing the very young flower-buds, one bracteole overlapping the other by its margins, both caducous before the flower opens
    Receptacle
    Receptacle short
    Calyx
    Sepals 4, imbricate (2 outer, 2 inner)
    Corolla
    Petals 5: upper 3 large, clawed, lower 2 very small, or occasionally all 5 well-developed and subequal
    Stamens
    Stamens 10, free
    Pistil
    Ovary stipitate; ovules 4; style long; stigma small, capitate
    Fruits
    Pod indehiscent, thick, woody, ± resinous-warted, with a pithy endocarp, 1–3-seeded
    Seeds
    Seeds hard, ± ellipsoid.
    [LOWO]
    Use
    Resin (referred to as S American or Zanzibar copal , depending on the species) is used for incense, glue, varnish, shellac and as traditional medicines; other uses include edible fruits, timber ( algarrobo, jatoba, courbaril, guapinol ) for construction, high quality furniture, cabinetry, veneers, joinery, panelling, turnery, musical instruments and boat building, the bark is used for canoes, and some species are cultivated as ornamentals

    Images

    Distribution

    Native to:

    Belize, Bolivia, Brazil North, Brazil Northeast, Brazil South, Brazil Southeast, Brazil West-Central, Colombia, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, Haiti, Honduras, Jamaica, Kenya, Leeward Is., Madagascar, Mauritius, Mexico Central, Mexico Gulf, Mexico Northwest, Mexico Southeast, Mexico Southwest, Mozambique, Nicaragua, Panamá, Paraguay, Peru, Puerto Rico, Réunion, Suriname, Tanzania, Trinidad-Tobago, Venezuela, Venezuelan Antilles, Windward Is.

    Introduced into:

    Benin, Borneo, China Southeast, Congo, Ghana, Guinea, Guinea-Bissau, India, Ivory Coast, Jawa, Madagascar, Malaya, Philippines, Senegal, Seychelles, Sri Lanka, Taiwan, Uganda, Zimbabwe

    Hymenaea L. appears in other Kew resources:

    Date Reference Identified As Barcode Type Status
    Feb 5, 2008 Freire, E. [54], Pernambuco K000834250
    Feb 5, 2008 Hora, M.J. [49], Pernambuco K000834251
    Feb 5, 2008 Tschá, M. [642], Pernambuco K000834252
    Jan 1, 1998 Clarke, D. [4426], Guyana K000404329
    Jan 1, 1988 Thomas, W. [5313], Brazil K000404320
    Jan 1, 1985 Rabelo, B.V. [3052], Brazil K000404296
    Dec 12, 1978 Philcox, D. [4116], Brazil K000404307
    Jan 1, 1976 Ramos, J.F. [P22546], Brazil K000404308
    Hunt, D.R. [5713], Brazil K000404313
    Hunt, D.R. [5713], Brazil K000404314
    Stannard, B. [5355], Brazil K000404326
    Harley, R. [3702], Bahia K000834253
    Hatschbach, G. [38970], Brazil K000404299
    Hatschbach, G. [45959], Brazil K000404302
    Hatschbach, G. [38369], Brazil K000404303
    Freire, E. [60], Pernambuco K000834248
    Milliken, W. [822], Brazil K000404124
    Rico, L. [1460], Bolivia K000295377
    Rico, L. [1480], Bolivia K000295368
    Thomas, W. [9571], Brazil K000404316
    Silva, J.M. [5956], Minas Gerais K000834247
    Giulietti, A.M. [6074], Brazil K000404330
    Souza, V.C. [5378], Bahia K000834243
    Souza, V.C. [5378], Bahia K000834244
    Souza, V.C. [5378], Bahia K000834245
    Queiroz, L.P. [4128], Brazil K000404294
    Queiroz, L.P. [3447], Brazil K000404325
    Schatz, G.E. [801], Brazil K000404310
    Schatz, G.E. [846], Brazil K000404311
    Carvalho, A.M. [2087], Brazil K000404309
    Carvalho, A.M. [3162], Brazil K000404315
    Guedes, M.L. [PCB 5425], Brazil K000404331
    Arbo, M.M. [7482], Brazil K000404327
    Heringer, E.P. [988], Brazil K000404298
    Hora, M.J. [52], Pernambuco K000834241
    Hora, M.J. [52], Pernambuco K000834242
    Passos, L. [PCD 4951], Brazil K000404332
    Saar, E. [PCD5318], Brazil K000404333
    Ferreira, L.V. [67], Brazil K000404300
    Bernardi, L. [18272], Paraguay K000404297
    Figueirêdo, L. [253], Pernambuco K000834249
    Balée, W.L. [880], Brazil K000404312
    Silva, M.A. [3838], Brazil K000404328
    Cordeiro, J. [2730], Santa Catarina K000834246
    Ferreira, C.A.C. [9492], Brazil K000404301
    Ferreira, C.A.C. [7090], Brazil K000404321
    Ferreira, C.A.C. [7946], Brazil K000404323
    Vilhena, R. [1012], Brazil K000404319
    White, C.E. [s.n.], Amazonas K000834991
    Jansen-Jacobs, M.J. [1365], Guyana K000404324
    Ezechias, P. [14064], Brazil K000404295
    Vargas, A.A. [131], Peru K000404317
    Arostegui V., A. [111], Peru K000404318
    Ferreira, C.A.C.. [7712], Brazil K000404322

    First published in Sp. Pl.: 1192 (1753)

    Literature

    Kew Bulletin
    • BFG (2015). Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguesia 66(4): 1085 -1113.
    • Bruneau, A., Breteler, F.J, Klitgaard, B. B., Prenner, G., Fougere-Danezan, M. & Tucker, S. C. (2014). Floral evolution in the Detarieae (Leguminosae): Phylogenetic evidence for labile floral development in an early-diverging legume lineage. Int. J. Pl. Sci. 175(4): 392 - 417.
    • Carneiro-Torres, D. S., Cordeiro, I., Giulietti, A. M., Berry, P. E. & Riina, R. (2011). Three new species of Croton (Euphorbiaceae s.s.) from the Brazilian Caatinga. Brittonia 63: 122 - 132.
    • Bruneau, A., Breteler, F.J, Mercure, M., Lewis, G. P. & Herendeen, P. S. (2008). Phylogenetic patterns and diversification in the caesalpinioid legumes. Botany 86: 697 - 718.
    • Cardoso, D. B. O. S., Queiroz, L. P. & Lima, H. C. (2008). Three new species of Luetzelburgia (Leguminosae, Papilionoideae) from the caatinga of Bahia, Brazil and an identification key to all species of the genus. Kew Bull. 63: 289 - 300.
    • Bruneau, A., Breteler, F.J., Wieringa, J.J., Gervais, G.Y.F. & Forest, F. (2000). Phylogenetic relationships in tribes Macrolobieae and Detarieae inferred from chloroplast trnL intron sequences. In: P. S. Herendeen & A. Bruneau (eds), Advances in legume systematics, part 9, pp. 121 - 149. Royal Botanic Gardens, Kew.
    Flora Zambesiaca
    • Lee & Langenheim in Univ. Calif. Publ. Bot. 69: 1–109 (1975).
    • Gen. Pl., ed. 5: 499 (1754).
    • Sp. Pl.: 1192 (1753)

    Sources

    Flora Zambesiaca
    Flora Zambesiaca
    http://creativecommons.org/licenses/by-nc-sa/3.0

    Flora of Tropical East Africa
    Flora of Tropical East Africa
    http://creativecommons.org/licenses/by-nc-sa/3.0

    Herbarium Catalogue Specimens
    'The Herbarium Catalogue, Royal Botanic Gardens, Kew. Published on the Internet http://www.kew.org/herbcat [accessed on Day Month Year]'. Please enter the date on which you consulted the system.
    Digital Image © Board of Trustees, RBG Kew http://creativecommons.org/licenses/by/3.0/

    Kew Backbone Distributions
    The International Plant Names Index and World Checklist of Selected Plant Families 2018. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
    © Copyright 2017 World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

    Kew Names and Taxonomic Backbone
    The International Plant Names Index and World Checklist of Selected Plant Families 2018. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
    © Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

    Legumes of the World Online
    http://creativecommons.org/licenses/by-nc-sa/3.0