1. Family: Fabaceae Lindl.
    1. Sophora L.

      1. This genus is accepted, and is native to Asia-Tropical, Europe, Africa, Tristan da Cunha, Pacific, Southern America, Australasia, 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

    In Polhill’s (1994) treatment the following informal groups were recognised: the Myroxylon group (11 genera; 10 Neotropics, one Africa); Ormosia group (3 genera; Neotropics, Africa, Asia); Angylocalyx group (4 genera; Neotropics, Africa, Australia); Baphia group (6 genera; Africa to Asia); Dussia group (9 genera; Neotropics) and Sophora group (14 genera; Africa, Asia, Neotropics).

    The only formal change made to the classification of Sophoreae since Polhill (1994) is the transfer of Bowringia and Baphiastrum to Leucomphalos (Breteler, 1994b). In this account we maintain Bowringia and Baphiastrum, not because we disagree with Breteler (1994b), but in the spirit of this volume, to encourage future workers to verify the monophyly of Leucomphalos sens. lat. with new data. We also do not follow Polhill’s (1994) suggestion that Riedeliella, Etaballia and Inocarpus belong in Sophoreae. It has been generally accepted (e.g., Polhill, 1981b) that these belong in Dalbergieae, which is confirmed by the recent study of Lavin et al. (2001a) that places them in the Dalbergioid clade. They are therefore treated as Dalbergieae in this volume (see page 307).

    Cladistic analyses of overall morphology (Chappill, 1995; Herendeen, 1995) and pollen data (Ferguson et al., 1994) showed Sophoreae to be non-monophyletic because Swartzieae genera were mixed in the same monophyletic groups as Sophoreae. These results have been corroborated by molecular studies. Doyle et al. (1996) showed Sophoreae to be heterogeneous for a large inversion in the chloroplast genome. This suggests that Sophoreae is non-monophyletic if it is assumed that the inversion arose only once. Doyle et al.’s (1997) DNA sequencing study of the chloroplast gene rbcL included 18 genera of Sophoreae. Cladistic analysis showed these to be scattered widely across the papilionoid tree. More recently, these results have been corroborated by another chloroplast locus, the trnL intron (Pennington et al., 2001). This study sampled more putatively basal genera of Papilionoideae (26 of 41 Sophoreae; 14 of 15 Swartzieae and all Dalbergieae and Dipterygeae). The trnL tree (summarised in Fig. 29) is also largely congruent with other molecular studies that include some taxa of basal Papilionoideae (e.g., Hu et al., 2000; Ireland et al, 2000; Lavin et al., 2001a; Kajita et al., 2001; Wojciechowski et al., 2004). It clearly shows genera of Sophoreae to be members of disparate papilionoid clades.

    Diverse datasets now indicate Sophoreae to be non-monophyletic as Polhill (1981b; 1994) predicted. If the trnL results are corroborated, it seems likely that Sophoreae will be dismembered with its genera scattered across several tribes. This would entail extensive taxonomic changes. Yakovlev (1972b; 1991) split Sophoreae into five and nine tribes respectively. These classifications have not been widely accepted, and although they are not congruent with the most recent molecular topologies, they will need to be considered in any formalisation of new tribal names. In any new scheme, Sophoreae sens. strict. will comprise a group of genistoid clade genera from among Polhill’s (1994) Sophora group (Fig. 29), but published molecular phylogenetic studies have not yet sampled sufficient genera to suggest its delimitation.

    A new classification for Sophoreae requires sampling of the genera not included by Pennington et al. (2001; see Fig. 29) and other authors, in future molecular systematic studies. Some of the clades discovered by DNA sequence data (Fig. 29) are cryptic in that they are not marked by obvious macro-morphological features, and it is therefore perilous to attempt to determine the affinities of genera based upon macro-morphology alone. It may be that these clades are defined by anatomical or chemical characters. For example, quinolizidine alkaloid accumulation may be a synapomorphy for the Genistoid clade (Pennington et al., 2001; Kite & Pennington, 2003), and lack of these chemicals in Styphnolobium species supports the segregation of this genus from Sophora sens. strict. The presence of quinolizidine alkaloids in Calia, which is not placed amongst the genistoids, suggests that this genus is a strong candidate as sister group to the Genistoid clade, a relationship that might be resolved by more robust molecular phylogenies. Such phylogenies should incorporate information from nuclear genes (Lavin et al., 1998; Doyle & Doyle, 2000) which would be particularly useful to test hypotheses that are currently based solely upon evidence from chloroplast DNA. Careful integration of morphology, preferably as part of a simultaneous cladistic analysis, is also critical. Such morphological study may be best achieved by focusing on separate monophyletic groups because assessment of homology of morphological features across all Papilionoideae is difficult. The monophyletic groups discovered in the trnL analysis provide a framework for starting these future studies. In all 45 genera and (393) – 396 – (398) species are treated here (including c. 76 basally branching, c. 262 genistoid and c. 58 baphioid species of Sophoreae; Fig. 29).

    Some authors maintain a broad circumscription of Sophora; Yakovlev (1967), however, split it into a number of genera and there is a tendency to recognise some of these segregates, e.g., Calia and Styphnolobium have been shown by molecular and morphological data to be distinct genera; sect. Edwardsia (c. 16-18 spp. with various hybrids in New Zealand) is most distinctive biogeographically, comprising the Australasian, Pacific, Mascarenes and S American element of the genus; the centre of the remaining species is China, India and Indo-China. On the basis of DNA sequence data, Sophora sens. strict. belongs with the Genistoid clade (Doyle et al., 1997; Pennington et al., 2001; Kajita et al., 2001; Wojciechowski et al., 2004)
    Habit
    Trees, shrubs and herbs
    Ecology
    Seasonally dry tropical to warm temperate lowland and upland forest or dry vegetation types and sand dunes
    Distribution
    SE Europe to W, C & E Asia and south through tropical regions to Australasia and the Pacific; c. 3-4 spp. in sect. Edwardsia in western S America (Chile, Argentina and Juan Fernandez Is.); largely introduced in Africa; 1 sp. endemic from coastal Kenya south to S Africa and Madagascar; S. tomentosa L. widespread in the coastal Palaeotropics and also in coastal E Brazil
    [FZ]

    Leguminosae, R.K. Brummitt, D.K. Harder, G.P. Lewis, J.M. Lock, R.M. Polhill & B. Verdcourt. Flora Zambesiaca 3:3. 2007

    Habit
    Trees or shrubs or rarely perennial herbs.
    Leaves
    Leaves imparipinnate, with 4–18(32) leaflets per side, glabrous to densely tomentose; stipels absent.
    Inflorescences
    Inflorescences terminal or axillary, consisting of few- to many-flowered racemes; bracts often fairly large; bracteoles small when present but usually apparently absent.
    Flowers
    Flowers markedly perigynous, with a distinct hypanthium.
    Calyx
    Calyx campanulate to tubular, with very shallow to prominent and acute lobes, the upper 2 often fused.
    Corolla
    Petals yellow, white, blue or purple, small to rather large; standard usually gradually narrowed below into a short claw, the limb ± reflexed; keel petals overlapping or joined on the lower side.
    Stamens
    Stamens free to shortly joined at the base; anthers dorsifixed.
    Ovary
    Ovary shortly stalked.
    Fruits
    Pod moniliform, often winged, with 1–14 seeds, dehiscent or tardily breaking up irregularly.
    Seeds
    Seeds ovoid, ellipsoid or globose, usually without a distinct radicular lobe and with a small hilum; radicle short, ± straight or incurved.
    [FTEA]

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

    Habit
    Trees, shrubs or rarely perennial herbs
    Leaves
    Leaves imparipinnate, 8–36(–64)-foliolate; stipels setaceous or often absent; leaflets 0.5–5(–9) cm. long
    Flowers
    Flowers in terminal or axillary few- to many-flowered racemes; bracts often fairly large; bracteoles small when present, but usually apparently absent; pedicels often swollen or jointed near the top
    Hypanthium
    Hypanthium often well developed
    Calyx
    Calyx campanulate to tubular, with very shallow to prominent and acute lobes, the upper 2 often fused
    Corolla
    Petals yellow, white, blue or purple, up to 5 cm. long; standard usually narrowed into a short claw, the limb ± reflexed; wings ± obliquely oblong; keel-petals usually overlapping or joined on the lower side
    Stamens
    Stamens free or shortly joined at the base; anthers dorsifixed
    Pistil
    Ovary shortly stipitate, with several to numerous ovules; style incurved, with a small terminal stigma
    Fruits
    Fruit moniliform, often winged, 1–14-seeded, dehiscent or coriaceous to fleshy and indehiscent
    Seeds
    Seeds obovoid or globose, usually with a small hilum; radicle straight or incurved.
    [LOWO]

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

    Habit
    Shrubs (dwarf)
    Ecology
    Dry continental temperate grassland and desert
    Distribution
    C Asia (Kazakhstan, Tadzhikistan, Uzbekistan and Turkmenistan)
    Note
    Some authors consider this genus to be a section within Sophora (e.g., S. gibbosa (DC.) Yakovlev and S. pachycarpa C.A.Mey. from W Asia and the Middle East), but within its area of distribution it is regarded as a distinct genus

    In Polhill’s (1994) treatment the following informal groups were recognised: the Myroxylon group (11 genera; 10 Neotropics, one Africa); Ormosia group (3 genera; Neotropics, Africa, Asia); Angylocalyx group (4 genera; Neotropics, Africa, Australia); Baphia group (6 genera; Africa to Asia); Dussia group (9 genera; Neotropics) and Sophora group (14 genera; Africa, Asia, Neotropics).

    The only formal change made to the classification of Sophoreae since Polhill (1994) is the transfer of Bowringia and Baphiastrum to Leucomphalos (Breteler, 1994b). In this account we maintain Bowringia and Baphiastrum, not because we disagree with Breteler (1994b), but in the spirit of this volume, to encourage future workers to verify the monophyly of Leucomphalos sens. lat. with new data. We also do not follow Polhill’s (1994) suggestion that Riedeliella, Etaballia and Inocarpus belong in Sophoreae. It has been generally accepted (e.g., Polhill, 1981b) that these belong in Dalbergieae, which is confirmed by the recent study of Lavin et al. (2001a) that places them in the Dalbergioid clade. They are therefore treated as Dalbergieae in this volume (see page 307).

    Cladistic analyses of overall morphology (Chappill, 1995; Herendeen, 1995) and pollen data (Ferguson et al., 1994) showed Sophoreae to be non-monophyletic because Swartzieae genera were mixed in the same monophyletic groups as Sophoreae. These results have been corroborated by molecular studies. Doyle et al. (1996) showed Sophoreae to be heterogeneous for a large inversion in the chloroplast genome. This suggests that Sophoreae is non-monophyletic if it is assumed that the inversion arose only once. Doyle et al.’s (1997) DNA sequencing study of the chloroplast gene rbcL included 18 genera of Sophoreae. Cladistic analysis showed these to be scattered widely across the papilionoid tree. More recently, these results have been corroborated by another chloroplast locus, the trnL intron (Pennington et al., 2001). This study sampled more putatively basal genera of Papilionoideae (26 of 41 Sophoreae; 14 of 15 Swartzieae and all Dalbergieae and Dipterygeae). The trnL tree (summarised in Fig. 29) is also largely congruent with other molecular studies that include some taxa of basal Papilionoideae (e.g., Hu et al., 2000; Ireland et al, 2000; Lavin et al., 2001a; Kajita et al., 2001; Wojciechowski et al., 2004). It clearly shows genera of Sophoreae to be members of disparate papilionoid clades.

    Diverse datasets now indicate Sophoreae to be non-monophyletic as Polhill (1981b; 1994) predicted. If the trnL results are corroborated, it seems likely that Sophoreae will be dismembered with its genera scattered across several tribes. This would entail extensive taxonomic changes. Yakovlev (1972b; 1991) split Sophoreae into five and nine tribes respectively. These classifications have not been widely accepted, and although they are not congruent with the most recent molecular topologies, they will need to be considered in any formalisation of new tribal names. In any new scheme, Sophoreae sens. strict. will comprise a group of genistoid clade genera from among Polhill’s (1994) Sophora group (Fig. 29), but published molecular phylogenetic studies have not yet sampled sufficient genera to suggest its delimitation.

    A new classification for Sophoreae requires sampling of the genera not included by Pennington et al. (2001; see Fig. 29) and other authors, in future molecular systematic studies. Some of the clades discovered by DNA sequence data (Fig. 29) are cryptic in that they are not marked by obvious macro-morphological features, and it is therefore perilous to attempt to determine the affinities of genera based upon macro-morphology alone. It may be that these clades are defined by anatomical or chemical characters. For example, quinolizidine alkaloid accumulation may be a synapomorphy for the Genistoid clade (Pennington et al., 2001; Kite & Pennington, 2003), and lack of these chemicals in Styphnolobium species supports the segregation of this genus from Sophora sens. strict. The presence of quinolizidine alkaloids in Calia, which is not placed amongst the genistoids, suggests that this genus is a strong candidate as sister group to the Genistoid clade, a relationship that might be resolved by more robust molecular phylogenies. Such phylogenies should incorporate information from nuclear genes (Lavin et al., 1998; Doyle & Doyle, 2000) which would be particularly useful to test hypotheses that are currently based solely upon evidence from chloroplast DNA. Careful integration of morphology, preferably as part of a simultaneous cladistic analysis, is also critical. Such morphological study may be best achieved by focusing on separate monophyletic groups because assessment of homology of morphological features across all Papilionoideae is difficult. The monophyletic groups discovered in the trnL analysis provide a framework for starting these future studies. In all 45 genera and (393) – 396 – (398) species are treated here (including c. 76 basally branching, c. 262 genistoid and c. 58 baphioid species of Sophoreae; Fig. 29).

    [LOWO]
    Use
    Cultivated as ornamentals (in sect. Edwardsia, with yellow flowers, e.g., S. tetraptera J.F.Mill. [kowhai ] and S. microphylla Aiton [small-leaved kowhai ]; and in sect. Pseudosophora, with blue to white flowers, e.g., S. davidii (Franch.) Skeels); also used for its durable timber (for bearings, turnery and cabinet work), as medicine (e.g., S. flavescens Aiton [ku shen ], S. tonkinensis Gagnep. [shan dou gen ] and S. microphylla); some species are toxic and used as insecticides
    [LOWO]
    Use
    Used for medicine, dyes, forage, ornamentals, preventing erosion and as bee crops for honey

    Images

    Distribution

    Native to:

    Afghanistan, Aldabra, Altay, Amur, Andaman Is., Argentina Northeast, Argentina Northwest, Arizona, Arkansas, Aruba, Assam, Bahamas, Bangladesh, Belize, Bermuda, Bismarck Archipelago, Brazil North, Brazil Northeast, Brazil South, Brazil Southeast, Cambodia, Cayman Is., Central African Repu, Chatham Is., Chile Central, Chile South, China North-Central, China South-Central, China Southeast, Chita, Colombia, Colorado, Cook Is., Costa Rica, Cuba, Dominican Republic, East European Russia, East Himalaya, Fiji, Florida, Ghana, Gilbert Is., Guinea, Gulf of Guinea Is., Hainan, Haiti, Hawaii, Honduras, India, Inner Mongolia, Iran, Iraq, Irkutsk, Ivory Coast, Jamaica, Japan, Jawa, Juan Fernández Is., Kansas, Kazakhstan, Kenya, Khabarovsk, Kirgizstan, Korea, Krym, KwaZulu-Natal, Laccadive Is., Lebanon-Syria, Leeward Is., Lesser Sunda Is., Liberia, Louisiana, Madagascar, Maldives, Manchuria, Mexico Gulf, Mexico Northeast, Mexico Northwest, Mexico Southeast, Mongolia, Mozambique, Mozambique Channel I, Myanmar, Nansei-shoto, Nepal, Netherlands Antilles, New Caledonia, New Guinea, New Mexico, New South Wales, New Zealand North, New Zealand South, Nicaragua, Nicobar Is., Nigeria, Norfolk Is., North Caucasus, Northern Territory, Ogasawara-shoto, Oklahoma, Oregon, Pakistan, Panamá, Philippines, Primorye, Puerto Rico, Qinghai, Queensland, Romania, Réunion, Samoa, Saudi Arabia, Senegal, Seychelles, Sierra Leone, South China Sea, South Dakota, South European Russi, Sri Lanka, Sumatera, Tadzhikistan, Taiwan, Tanzania, Texas, Thailand, Tibet, Togo, Tonga, Transcaucasus, Trinidad-Tobago, Tristan da Cunha, Tuamotu, Tubuai Is., Turkey, Turkey-in-Europe, Turkmenistan, Turks-Caicos Is., Utah, Uzbekistan, Vanuatu, Venezuela, Vietnam, Wallis-Futuna Is., West Himalaya, Windward Is., Wyoming, Xinjiang, Zimbabwe

    Extinct in:

    Easter Is.

    Introduced into:

    Great Britain, Mauritius, Peru, Rodrigues, Ukraine, West Siberia

    Sophora L. appears in other Kew resources:

    First published in Sp. Pl.: 373 (1753)

    Literature

    Flora of West Tropical Africa
    • —F.T.A. 2: 253.
    Flora Zambesiaca
    • Gen. Pl., ed.5: 175 (1754).
    • Sp. Pl.: 373 (1753)
    Flora of Tropical East Africa
    • L., Gen. Pl., ed. 5: 175 (1754)
    • Sp. Pl.: 373 (1753)

    Sources

    Art and Illustrations in Digifolia
    Digital Image © Board of Trustees, RBG Kew

    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

    Kew Backbone Distributions
    The International Plant Names Index and World Checklist of Selected Plant Families 2019. 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 2019. 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
    Digital Image © Board of Trustees, RBG Kew http://creativecommons.org/licenses/by-nc-sa/3.0/
    http://creativecommons.org/licenses/by-nc-sa/3.0