1. Family: Fabaceae Lindl.
    1. Pterocarpus Jacq.

      1. This genus is accepted, and is native to Asia-Tropical, Africa, Pacific, Southern America, Northern America and Asia-Temperate..

    [LOWO]

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

    Note

    The present circumscription of Dalbergieae sens. lat. contains radical changes to Dalbergieae sensu Polhill (1981d: 233–242). In the first instance it takes a traditional view of the tribe by including genera such as Vatairea and Vataireopsis (now in the Vataireoid clade, see Figs. 1& 40) and Andira and Hymenolobium, which in Wojciechowski et al. (2004) are sister to the combined Dalbergioid clade of Lavin et al. (2001a), plus Amorpheae. The bulk of the treatment, however, recognises the cryptic Dalbergioid clade (Lavin et al., 2001a) as comprising tribe Dalbergieae sens. lat., diagnosed by the synapomorphy of aeschynomenoid root nodules. This clade includes all the genera placed in the Dalbergieae sensu Polhill (1981d: 233– 242), the Aeschynomeneae sensu Rudd (1981) and the Adesmieae sensu Polhill (1981g: 355–356), plus subtribe Bryinae of the Desmodieae sensu Ohashi et al. (1981) as well as the genus Diphysa (tribe Robinieae sensu Polhill & Sousa (1981)).The placements of all members of the Dalbergioid clade within the classification presented here and in those of Polhill (1981d: 233–242; 1981g: 355–356), Polhill & Sousa (1981), Ohashi et al. (1981) and Rudd (1981) are listed in Fig. 40.

    Dalbergieae sensu Polhill (1981d) contained 19 genera defined by woody habit, supposedly plesiomorphic flowers, pods with a specialised seed chamber and seeds that accumulated alkaloids. Polhill (1981d) noted that there seemed to be two centres within the Dalbergieae: one around Andira with Hymenolobium, Vatairea, Vataireopsis, Dalbergia, and Machaerium, and one around Pterocarpus. He also highlighted evidence from wood anatomy (Baretta-Kuipers, 1981) which showed that Andira, Hymenolobium, Vatairea, and Vataireopsis have coarser wood structures more typical of members of the Sophoreae than the remaining members of the Dalbergieae. The study of fruit and seedling morphology by Lima (1990) further supported these two centres within the Dalbergieae: one including Andira, Hymenolobium, Vatairea, and Vataireopsis, and the second the remaining genera. Most recently several molecular and morphological studies (e.g., Lavin et al., 2001a; Pennington et al., 2001; Wojciechowski et al., 2004) confirm that these four genera do not belong in the Dalbergioid clade. Since there is, however, still much work to be done to resolve the phylogenetic relationships of these four genera, they have been kept in tribe Dalbergieae sens. lat. in this treatment to avoid tentative placements, which might be treated by users as formal.

    The tribe Aeschynomeneae sensu Rudd (1981) contained 25 genera characterised by lomentaceous pods, although some members lack loments (e.g., Arachis, Ormocarpopsis, Diphysa spp., Ormocarpum spp., and Pictetia spp.). None of the Aeschynomeneae had previously been considered closely related to the Dalbergieae, but the work of Lavin et al. (2001a) has resolved all the ‘aeschynomenoid genera’ within the Dalbergioid clade.

    The taxonomic history of the monogeneric tribe Adesmieae sensu Polhill (1981g) is very different from that of the Dalbergieae. The Adesmieae combines the presumed plesiomorphic trait of free stamen filaments, with presence of lomentaceous pods that are supposedly derived. This combination of features suggested a taxonomically isolated position far removed from the Dalbergieae. The analyses of Lavin et al. (2001a) based on molecular sequence and morphological data, however, support Adesmia (nested together with five genera of Rudd’s Aeschynomeneae) being sister to the Pterocarpus and Dalbergia clades. The neotropical genera Brya and Cranocarpus were placed together in a new subtribe Bryinae of Desmodieae in the classification of Ohashi et al. (1981). The features common to these genera are periporate pollen and glochidiate hairs or glandular trichomes. In the molecular studies of Doyle et al. (1995) and Bailey et al. (1997), Brya and Cranocarpus did not lack the intron for the chloroplast gene rpl2 nor for the open reading frame ORF184, which are characteristic of the other desmodioid genera studied. Bailey et al. (1997), therefore, suggested that Brya and Cranocarpus should be removed from the Desmodieae. Their findings were strongly corroborated by the three gene analyses of Lavin et al. (2001a) which place the two genera in the Pterocarpus clade (Fig. 40).

    One further transfer has been made in the Dalbergioid clade since Rudd (1981) and Polhill & Sousa (1981). Lavin (1987) transferred Diphysa from the Robinieae to tribe Aeschynomeneae based on the absence of canavanine in the seeds, a feature consistently present in the Robinieae (Lavin, 1986). Lavin (1987) also listed 14 morphological characters that placed Diphysa with the Aeschynomeneae rather than the Robinieae. In the phylogenetic analyses of Lavin et al. (2001a), Diphysa is resolved in the Dalbergioid clade nested within the ‘transatlantic clade’ (Fig. 40), first identified by Lavin et al. (2000).

    Finally, since 1981 four new genera have been published: the Brazilian monotypic Grazielodendron (Lima, 1983b), the possibly extinct Madagascan endemic Peltiera (Labat & Du Puy, 1997), Zygocarpum, the Horn of Africa – Arabian segregate of Ormocarpum (Thulin & Lavin, 2001) and Maraniona from northern Peru (Hughes et al., 2004). Three genera have also been placed in synonymy since 1981: the Caribbean genus Belairia which is a synonym of Pictetia (Beyra-Matos & Lavin, 1999), and the genera Pachecoa and Arthrocarpum which Thulin (1999) synonymised under Chapmannia. In this treatment 49 genera and (1319) –1325–(1331) species are recognised in Dalbergieae sens. lat. (including 4 basally branching dalbergioid genera comprising c. 58 species, and (1261)–1267– (1273) species in the 45 genera of the Dalbergioid clade [Lavin et al., 2001a]).The following informal groupings of genera are based on the work of Lavin et al. (2001a): Adesmia clade: 6 genera; c. 360 species; neotropical except Zornia, which is pantropical. Pterocarpus clade: 22 genera; c. 200 species centred in the Neotropics, Pterocarpus and Stylosanthes are pantropical, Inocarpus Asian, and Chapmannia transatlantic.Dalbergia clade: 17 genera; c. 706 species which are pantropical, but centred in Africa; Weberbauerella, Soemmeringia, Pictetia and Diphysa are neotropical, Machaerium transatlantic, Dalbergia and Aeschynomene are pantropical, and Geissaspis Asian. Isolated genera: 4 genera; 58 species, neotropical except Andira, which has one amphiatlantic species.

    The genus may be subdivided into two groups based on winged versus unwinged fruits. Polhill (1981d: 233-242) placed Pterocarpus in tribe Dalbergieae most closely related to Tipuana and Platypodium. This relationship is confirmed by the combined analyses of Lavin et al. (2001a), which resolved Pterocarpus in the Pterocarpus clade (Fig. 40), together with the majority of the genera in Dalbergieae sensu Polhill (1981d)
    Vernacular
    dragon's blood, kino gum, sangre de drago
    Habit
    Tall trees, some species are buttressed rain forest emergents
    Ecology
    Tropical lowland evergreen rain forest (less than 10 spp.) to seasonally dry forest, woodland, thicket and wooded grassland
    Distribution
    pantropical, with the greatest diversity in Africa; c. 20 spp. endemic to Africa, 1 sp. (P. santalinoides L'Her. ex DC.) also native to the Neotropics, and 1 sp. (P. indicus Willd.) native to Asia; 5 spp. restricted to S Asia, Indo-China and Madagascar; c. 11 spp. endemic to the Neotropics (Mexico and C America with 5 spp. and S America with 6 spp.)
    [FTEA]

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

    Habit
    Evergreen or deciduous trees; slash usually producing a red resinous exudate
    Leaves
    Leaves alternate, imparipinnate, with leaflets alternate to sub-opposite, very rarely (and not in East Africa) 1-foliolate; stipules variously developed, frequently caducous; stipels absent; leaflets often covered with minute glands beneath
    Flowers
    Flowers in terminal or axillary racemes or panicles; bracts and bracteoles generally small and caducous
    Calyx
    Calyx turbinate to campanulate, shortly 5-lobed, with the 2 upper lobes practically connate
    Corolla
    Corolla yellow to orange in Africa (elsewhere occasionally whitish and violet); standard generally with a well-developed claw and broadly expanded blade, glabrous or practically so (at least in Africa); wings free, oblong or often with the blade expanded, thus ± obovate or spathulate; keel-petals usually narrow and shorter, lightly coherent on lower side towards the tips
    Stamens
    Stamens connate into a sheath open above or above and below, sometimes the vexillary stamen free; anthers dorsifixed with longitudinal dehiscence
    Pistil
    Ovary sessile (not in Africa) or stipitate, few-ovulate; style little curved, tapered to small terminal stigma, glabrous at least apically
    Fruits
    Fruit compressed, indehiscent, with a variously thickened or hardened central seed-bearing part and a narrow to very broad wing, bearing the style subterminally, laterally or curved right round to the base, 1 (–3)-seeded
    Seeds
    Seeds reniform or oblong-reniform, smooth; hilum and rim-aril small.
    [FZ]

    Flora Zambesiaca Leguminosae subfamily Papillionoideae byJ.M. Lock*

    Habit
    Trees. Trees.
    Leaves
    Leaves imparipinnate, stipulate; leaflets alternate, opposite or irregularly subopposite. Leaves imparipinnate, stipulate; leaflets alternate, opposite or irregularly subopposite.
    Inflorescences
    Inflorescence racemose or paniculate. Inflorescence racemose or paniculate.
    Flowers
    Flowers yellow. Flowers yellow.
    Calyx
    Calyx turbinate at the base, incurved; upper teeth partially connate. Calyx turbinate at the base, incurved; upper teeth partially connate.
    Corolla
    Standard circular or broadly obovate; wings obliquely obovate; keel petals shorter than the wings, dorsally slightly connate or free. Standard circular or broadly obovate; wings obliquely obovate; keel petals shorter than the wings, dorsally slightly connate or free.
    Stamens
    Stamens either all 10 connate in a dorsally-slit sheath, or 9 fused with the upper one free, or in 2 groups (phalanges) of 5; anthers dorsifixed, longitudinally dehiscent. Stamens either all 10 connate in a dorsally-slit sheath, or 9 fused with the upper one free, or in 2 groups (phalanges) of 5; anthers dorsifixed, longitudinally dehiscent.
    Ovary
    Ovary few-ovulate; style somewhat incurved; stigma small, terminal.
    Fruits
    Pod circular, broadly winged or obovate with obsolete wings. Pod circular, broadly winged or obovate with obsolete wings.
    Pistil
    Ovary few-ovulate; style somewhat incurved; stigma small, terminal.
    Seeds
    Seed reniform or oblong-reniform; hilum small. Seed reniform or oblong-reniform; hilum small.
    [LOWO]

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

    Vernacular
    etabally
    Habit
    Trees
    Ecology
    Tropical lowland rain forest, often along riverbanks
    Distribution
    northern S America (Venezuela, Guyana and Brazil)
    Note
    Placed by Polhill (1981d: 233-242) in tribe Dalbergieae, but moved to the Sophoreae by Polhill (1994). Lavin et al. (2001a), based on DNA sequence data, resolved Etaballia in the Pterocarpus clade related to Inocarpus with which it shares radially symmetric flowers (Fig. 40)

    The present circumscription of Dalbergieae sens. lat. contains radical changes to Dalbergieae sensu Polhill (1981d: 233–242). In the first instance it takes a traditional view of the tribe by including genera such as Vatairea and Vataireopsis (now in the Vataireoid clade, see Figs. 1& 40) and Andira and Hymenolobium, which in Wojciechowski et al. (2004) are sister to the combined Dalbergioid clade of Lavin et al. (2001a), plus Amorpheae. The bulk of the treatment, however, recognises the cryptic Dalbergioid clade (Lavin et al., 2001a) as comprising tribe Dalbergieae sens. lat., diagnosed by the synapomorphy of aeschynomenoid root nodules. This clade includes all the genera placed in the Dalbergieae sensu Polhill (1981d: 233– 242), the Aeschynomeneae sensu Rudd (1981) and the Adesmieae sensu Polhill (1981g: 355–356), plus subtribe Bryinae of the Desmodieae sensu Ohashi et al. (1981) as well as the genus Diphysa (tribe Robinieae sensu Polhill & Sousa (1981)).The placements of all members of the Dalbergioid clade within the classification presented here and in those of Polhill (1981d: 233–242; 1981g: 355–356), Polhill & Sousa (1981), Ohashi et al. (1981) and Rudd (1981) are listed in Fig. 40.

    Dalbergieae sensu Polhill (1981d) contained 19 genera defined by woody habit, supposedly plesiomorphic flowers, pods with a specialised seed chamber and seeds that accumulated alkaloids. Polhill (1981d) noted that there seemed to be two centres within the Dalbergieae: one around Andira with Hymenolobium, Vatairea, Vataireopsis, Dalbergia, and Machaerium, and one around Pterocarpus. He also highlighted evidence from wood anatomy (Baretta-Kuipers, 1981) which showed that Andira, Hymenolobium, Vatairea, and Vataireopsis have coarser wood structures more typical of members of the Sophoreae than the remaining members of the Dalbergieae. The study of fruit and seedling morphology by Lima (1990) further supported these two centres within the Dalbergieae: one including Andira, Hymenolobium, Vatairea, and Vataireopsis, and the second the remaining genera. Most recently several molecular and morphological studies (e.g., Lavin et al., 2001a; Pennington et al., 2001; Wojciechowski et al., 2004) confirm that these four genera do not belong in the Dalbergioid clade. Since there is, however, still much work to be done to resolve the phylogenetic relationships of these four genera, they have been kept in tribe Dalbergieae sens. lat. in this treatment to avoid tentative placements, which might be treated by users as formal.

    The tribe Aeschynomeneae sensu Rudd (1981) contained 25 genera characterised by lomentaceous pods, although some members lack loments (e.g., Arachis, Ormocarpopsis, Diphysa spp., Ormocarpum spp., and Pictetia spp.). None of the Aeschynomeneae had previously been considered closely related to the Dalbergieae, but the work of Lavin et al. (2001a) has resolved all the ‘aeschynomenoid genera’ within the Dalbergioid clade.

    The taxonomic history of the monogeneric tribe Adesmieae sensu Polhill (1981g) is very different from that of the Dalbergieae. The Adesmieae combines the presumed plesiomorphic trait of free stamen filaments, with presence of lomentaceous pods that are supposedly derived. This combination of features suggested a taxonomically isolated position far removed from the Dalbergieae. The analyses of Lavin et al. (2001a) based on molecular sequence and morphological data, however, support Adesmia (nested together with five genera of Rudd’s Aeschynomeneae) being sister to the Pterocarpus and Dalbergia clades. The neotropical genera Brya and Cranocarpus were placed together in a new subtribe Bryinae of Desmodieae in the classification of Ohashi et al. (1981). The features common to these genera are periporate pollen and glochidiate hairs or glandular trichomes. In the molecular studies of Doyle et al. (1995) and Bailey et al. (1997), Brya and Cranocarpus did not lack the intron for the chloroplast gene rpl2 nor for the open reading frame ORF184, which are characteristic of the other desmodioid genera studied. Bailey et al. (1997), therefore, suggested that Brya and Cranocarpus should be removed from the Desmodieae. Their findings were strongly corroborated by the three gene analyses of Lavin et al. (2001a) which place the two genera in the Pterocarpus clade (Fig. 40).

    One further transfer has been made in the Dalbergioid clade since Rudd (1981) and Polhill & Sousa (1981). Lavin (1987) transferred Diphysa from the Robinieae to tribe Aeschynomeneae based on the absence of canavanine in the seeds, a feature consistently present in the Robinieae (Lavin, 1986). Lavin (1987) also listed 14 morphological characters that placed Diphysa with the Aeschynomeneae rather than the Robinieae. In the phylogenetic analyses of Lavin et al. (2001a), Diphysa is resolved in the Dalbergioid clade nested within the ‘transatlantic clade’ (Fig. 40), first identified by Lavin et al. (2000).

    Finally, since 1981 four new genera have been published: the Brazilian monotypic Grazielodendron (Lima, 1983b), the possibly extinct Madagascan endemic Peltiera (Labat & Du Puy, 1997), Zygocarpum, the Horn of Africa – Arabian segregate of Ormocarpum (Thulin & Lavin, 2001) and Maraniona from northern Peru (Hughes et al., 2004). Three genera have also been placed in synonymy since 1981: the Caribbean genus Belairia which is a synonym of Pictetia (Beyra-Matos & Lavin, 1999), and the genera Pachecoa and Arthrocarpum which Thulin (1999) synonymised under Chapmannia. In this treatment 49 genera and (1319) –1325–(1331) species are recognised in Dalbergieae sens. lat. (including 4 basally branching dalbergioid genera comprising c. 58 species, and (1261)–1267– (1273) species in the 45 genera of the Dalbergioid clade [Lavin et al., 2001a]).The following informal groupings of genera are based on the work of Lavin et al. (2001a): Adesmia clade: 6 genera; c. 360 species; neotropical except Zornia, which is pantropical. Pterocarpus clade: 22 genera; c. 200 species centred in the Neotropics, Pterocarpus and Stylosanthes are pantropical, Inocarpus Asian, and Chapmannia transatlantic.Dalbergia clade: 17 genera; c. 706 species which are pantropical, but centred in Africa; Weberbauerella, Soemmeringia, Pictetia and Diphysa are neotropical, Machaerium transatlantic, Dalbergia and Aeschynomene are pantropical, and Geissaspis Asian. Isolated genera: 4 genera; 58 species, neotropical except Andira, which has one amphiatlantic species.

    [LOWO]
    Use
    Pterocarpus indicus (narra, Solomon's padauk, Papua New Guinea rosewood, amboyna) may be the most economically important legume timber species. The wood of several species including P. angolensis DC. (African teak, bloodwood, kiaat, muninga) and P. soyauxii Taub. (padauk) is highly valued and used for fine furniture, cabinet making, panelling, joinery, musical instruments, implements and curios. The resin supplies dyes (e.g., P. tinctorius Welw.), and in the Amazon is known as ' sangre de drago' (= dragon ' s blood ); the resin is also a widely used folk medicine (the source of Kino gum ); other uses are in revegetation and soil improvement, as shade trees, ornamentals, cosmetics and baskets (from the inner bark)

    Images

    Distribution

    Native to:

    Andaman Is., Angola, Argentina Northeast, Bangladesh, Belize, Benin, Bismarck Archipelago, Bolivia, Borneo, Botswana, Brazil North, Brazil Northeast, Brazil South, Brazil Southeast, Brazil West-Central, Burkina, Cabinda, Cambodia, Cameroon, Caprivi Strip, Central African Repu, Chad, China South-Central, China Southeast, Colombia, Congo, Costa Rica, Cuba, Dominican Republic, Ecuador, El Salvador, Equatorial Guinea, Eritrea, Ethiopia, French Guiana, Gabon, Gambia, Ghana, Guatemala, Guinea, Guinea-Bissau, Gulf of Guinea Is., Guyana, Haiti, Honduras, India, Ivory Coast, Jamaica, Jawa, KwaZulu-Natal, Laos, Leeward Is., Lesser Sunda Is., Liberia, Malawi, Malaya, Maldives, Mali, Maluku, Mauritania, Mexico Central, Mexico Gulf, Mexico Southeast, Mexico Southwest, Mozambique, Myanmar, Namibia, Nansei-shoto, New Guinea, Nicaragua, Niger, Nigeria, Northern Provinces, Panamá, Paraguay, Peru, Philippines, Puerto Rico, Santa Cruz Is., Senegal, Sierra Leone, Solomon Is., Sri Lanka, Sudan, Sulawesi, Sumatera, Suriname, Swaziland, Taiwan, Tanzania, Thailand, Togo, Trinidad-Tobago, Uganda, Vanuatu, Venezuela, Vietnam, Windward Is., Zambia, Zaïre, Zimbabwe

    Introduced into:

    Assam, Kenya, Madagascar, Mauritius, New South Wales, Northern Territory, Queensland, Seychelles

    Pterocarpus Jacq. appears in other Kew resources:

    Date Reference Identified As Barcode Type Status
    Jun 1, 2012 Pollard, B.J. [417], Cameroon K000746682
    Apr 1, 2002 Cheek, M. [10210], Cameroon K000027490
    Oct 1, 2001 Cheek, M. [8928], Cameroon K000027491
    Pennington, T.D. [15016], Ecuador 62949.000
    Ducke, A. [s.n.], Brazil K001076593
    Ule, E. [9449], Peru K001076596
    Burchell, W.J. [3860], Brazil K001076595
    Blanchet, J.S. [2818], Brazil K001076594

    First published in Select. Stirp. Amer. Hist.: 283 (1763)

    Literature

    Flora of West Tropical Africa
    • —F.T.A. 2: 237.
    Flora Zambesiaca
    • Select. Stirp. Amer. Hist.: 283, t.183, fig. 93 (1763), nom. conserv.
    Flora of Tropical East Africa
    • Select. Stirp. Am. Hist.: 283, t. 183/92 (1763), nom. conserv. propos.

    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

    Kew Science Photographs
    Digital Image © Board of Trustees, RBG Kew http://creativecommons.org/licenses/by/3.0/

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