1. Family: Fabaceae Lindl.
    1. Piliostigma Hochst.

      1. This genus is accepted, and is native to Asia-Tropical, Africa, Brazil West-Central, Australasia and Asia-Temperate..

    [FTEA]

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

    Habit
    Small deciduous trees, occasionally shrubby, not climbing
    Tendrils
    Tendrils absent
    Leaves
    Leaves simple, conspicuously bilobed
    Flowers
    Flowers medium to small, normally dioecious and unisexual, rarely monoecious, very rarely hermaphrodite (see F.C.B. 3: 278 (1952)), in terminal axillary or leaf-opposed racemes or panicles
    Calyx
    Calyx with a turbinate tube and 4–5 short acute lobes
    Corolla
    Petals 5
    Stamens
    Fertile stamens 10 (in ♂ flowers), reduced to staminodes in ♀ flowers; filaments villous below
    Pistil
    Stigma thick, capitate, flattened-globose, sessile on the ovary; funicle several times as long as ovule
    Fruits
    Pods linear to oblong, many-seeded, leathery or woody, indehiscent
    Seeds
    Seeds irregularly arranged, embedded in pulp, with a U-shaped line on one side; endosperm present.
    [FZ]

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

    Habit
    Deciduous trees or shrubs.
    Leaves
    Leaves composed of 2 united leaflets, bilobed, petiolate, with deciduous stipules.
    Flowers
    Flowers in terminal, leaf-opposed or axillary racemes or panicles, medium to small, unisexual with the plants dioecious; bracts small; bracteoles similar at top of the pedicel at an articulation.
    Hypanthium
    Hypanthium infilled.
    Calyx
    Calyx deeply cupular, with 4–5 short lobes.
    Corolla
    Petals 5, usually white to pink.
    Androecium
    Stamens 10(11), all fertile in male flowers, reduced to staminodes in female flowers; filaments villous below; anthers dehiscing by longitudinal slits.
    Stigma
    Stigma thick, capitate, flattened-globose, sessile on the ovary.
    Fruits
    Pods leathery to woody, oblong, indehiscent, many-seeded.
    Seeds
    Seeds embedded in pulp, irregularly arranged, with a U-shaped line below the small subapical hilum.
    [LOWO]

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

    Note

    Tribe Cercideae is basally branching in the Leguminosae (Bruneau et al., 2001; Herendeen et al., 2003a), as predicted by Wunderlin et al. (1981), and Cercis is the most basally branching genus in the tribe. While much taxonomic work has been carried out on the tribe in the past thirty years (e.g., Larsen et al., 1980, 1984; Wunderlin, 1976, 1979; Wunderlin et al., 1981, 1987; Zhang, 1995; Vaz, 2003; Vaz & Tozzi, 2003), few species have been included in phylogenetic analyses and inter- and intra-generic relationships are still largely unresolved with the exception of Cercis (Hao et al., 2001; Davis et al., 2002b).

    Wunderlin (1979) and Wunderlin et al. (1981) divided the tribe into two subtribes, Cercidinae and Bauhiniinae, based on seed, floral and fruit characters. Walpers (1842) had already down-ranked Bauhinieae Benth. (1840) to subtribal status, thus the combination Bauhiniinae (Benth.) Wunderlin (1979) is superfluous. Polhill (1994) kept the Cercideae unchanged with two subtribes and five genera. While the Cercidinae contains three small distinct genera, Cercis, Griffonia and Adenolobus, the Bauhiniinae houses the monospecific Madagascan genus Brenierea and the large, diverse pantropical genus Bauhinia sens. lat. which has been segregated into as many as twenty-six genera by various authors (Wunderlin, 1976).

    While many of the Bauhinia segregates are based on minor morphological differences, others are distinguished morphologically by a suite of characters. Britton and Rose (1930), in their account of the Caesalpiniaceae for the North American Flora, divided Bauhinia into several segregate genera, including Schnella Raddi which here is treated as a synonym of Phanera, but might prove to be distinct as indicated in recent molecular analyses by Forest (unpublished data). Britton and Killip (1936) recognised Schnella as distinct from Bauhinia in Colombia. De Wit (1956), treating ‘Malaysian Bauhinieae’, recognised Bracteolanthus, Lysiphyllum, Gigasiphon, Piliostigma, Lasiobema and Phanera as separate genera and this was largely followed by subsequent flora writers in Africa and New Guinea (e.g., Brenan, 1967; Coetzer & Ross in Ross, 1977; Verdcourt, 1979). Others have retained a more inclusive Bauhinia proposed by Wunderlin et al. (1981, 1987), e.g., Macbride (1943: 207–220) for Peru; Larsen et al. (1980) for the Flora of Cambodia, Laos and Vietnam; Larsen et al. (1984) for the Flora of Thailand; Chen (1988) for China, and Larsen & Larsen in Hou et al. (1996) in Flora Malesiana. Zhang (1995) published a morphological cladistic analysis of the series of Bauhinia sens. lat., but few species of Bauhinia have been included in molecular studies. It remains equivocal as to whether Bauhinia sens. lat. is monophyletic, but preliminary molecular results indicate that some elements should be reinstated as distinct genera (Bruneau et al., in prep.; Forest, unpubl.). This runs contrary to the findings of Larsen & Larsen in Hou et al. (1996) who concluded “that Bauhinia in the sense of Linnaeus, Bentham, De Candolle, Taubert and Hutchinson is an evolutionary unit and a very natural genus”. Larsen and Larsen also noted that Bauhinia sens. lat. presents a reticulate pattern of variation across its pantropical range (this apparently conflicting somewhat with its status as a “natural genus”). While this is undoubtedly true if the genus is considered as all-inclusive, recent studies of legume distributions in general (Schrire et al., this volume and 2005) have revealed repeated patterns of generic distribution which appear to be duplicated by at least some of the segregates of Bauhinia. If these segregates are recognised as distinct genera (as several are in this treatment) then the reticulate pattern of variation of Bauhinia is far less pronounced. More sampling at the species level in molecular analyses and more morphological studies are needed across the full pantropical range of Bauhinia sens. lat. before inter- and intra-generic relationships are clearly resolved. In the current account genera that have been recognised as distinct from Bauhinia in at least one flora treatment that post-dates De Wit (1956) have been treated as separate genera, especially where these are supported by the preliminary results from a chloroplast trnL (intron and spacer) sequence analysis (Forest, unpubl.). The reader’s attention is also alerted to the detailed infra-generic division of Bauhinia by Wunderlin et al. (1987) in their reorganisation of the Cercideae which also forms a sound basis for sampling in future studies.

    Palynological studies of Bauhinia (Larsen, 1975; Schmitz, 1977; Ferguson & Pearce, 1986) have all stressed the considerable variation in pollen morphology within the genus sens. lat. and there are clear correlations between pollen exine ornamentation, floral morphology and pollination. It remains to be seen just how closely these correspond to evolutionary relationships of species. Nevertheless, Schmitz (1977) made several new combinations in segregate genera of Bauhinia based on palynological type. These included new names in Lasiobema, Lysiphyllum, Pauletia, Perlebia and Phanera (Pauletia and Perlebia here considered as synonyms of Bauhinia). Zhang (1995), who analysed morphologically the series of Bauhinia proposed by Wunderlin et al. (1987), concluded that while some supraspecific segregates of the genus were supported, none of the subgenera appeared to be monophyletic. Several realignments were proposed.

    The Cercideae as presented here includes 12 genera and (322)–335–(348) species. This treatment differs from Wunderlin et al. (1981, 1987) and Polhill (1994) in that Barklya, Gigasiphon, Lasiobema, Lysiphyllum, Phanera, Piliostigma and Tylosema are considered distinct from Bauhinia. While some of these may well be reincluded in Bauhinia after further study, yet other genera may be reinstated from within Bauhinia. Bracteolanthus, treated as distinct by De Wit (1956), is here included in Lysiphyllum following Wunderlin et al. (1987), while Barklya, considered congeneric with Bauhinia by Wunderlin (1979) and Wunderlin et al. (1981, 1987) is considered distinct following George (1998b) and Forest (unpublished data). The reinstatement of Lasiobema appears least well supported (Forest, unpubl.).

    A dioecious genus; species from tropical America tentatively referred to the informal Piliostigma group of Bauhinia by Wunderlin et al. (1981) were excluded from Bauhinia section Piliostigma by Wunderlin et al. (1987). Most African floras retain Piliostigma as a distinct genus although Wunderlin et al. (1987) recognise section Piliostigma in Bauhinia subgenus Elayuna. Phylogenetic studies by Sinou et al. (2009) indicate that Piliostigma should be recognised at the generic level, sister to Brenierea and Bauhinia and that within the Piliostigma-Brenierea-Bauhinia clade there is a unique loss of the rpl2 plastid region. Studies by Banks et al. (2013) show that the pollen of Piliostigma is also very distinct from the other segregate genera of Bauhinia s. l. in having pores with spiny opercula and spinose ornamentation.
    Habit
    Trees and shrubs
    Ecology
    In Malesia P. malabaricum is typically found in drier, deciduous, monsoon forest, usually at altitudes up to 300m (Hou et al., 1996), whilst in Australia it occurs in scrubland, grassland, forest and rainforest (George, 1998b), and in India it occurs in deciduous and evergreen forests.The genus occupies various habitats in Africa: P. reticulatum occurs in relatively dry savanna, whereas P. thoningii prefers moister climates (Burkhill, 1995) and inhabits woodland, wooded grassland and open, often disturbed sandy areas. P. foveolatum occurs in evergreen forest.
    Distribution
    P. thonningii is widespread, ranging from Senegal to Yemen in the north to South Africa's Northern Provinces in the south. P. reticulatum is found across west and central Africa to Ethiopia and Sudan. P. malabaricum is widespread from Pakistan, throughout southern Asia, to northern Australia. P. foveolatum is endemic to India and P. tortuosum is endemic to Myanmar.
    [LOWO]
    Use
    The bark, roots, seeds and pods of most species yield a dye. The bark and roots of P. thonningii are rich in tannin and have numerous medicinal uses, including as a gonorrhoea treatment, and the leaf sap is used to treat eyes infected with cobra spit. Fruit pulp from the drying pod is edible and has a sweet-acid taste, and the young shoots are added to porridge to give a sour taste. Fibre yielded by the bark is widely used for cordage and is also used for binding arrowheads to shafts and making bows. Resin from green pods is used as glue for fixing spear heads to the shaft. The dried leaves are pulverised with those of Parinari curatellifolia, to constitute an ingredient in the making of gunpowder. The species is also used as livestock fodder and fuel wood. In the Central African Republic it is considered as 'women's firewood' because it is excellent for the hearth and cooking place. It is alleged to have various magical powers including being added to charms against witchcraft as it is bitter and considered to be disliked by witches. P. reticulatum yields a fibre from the bark that is widely used for cordage and is used in drop nets by the Serer fishermen of Senegal. There are various medicinal uses across the range of this species. Preparations of the bark or leaf are given as treatments for earache, lumbago and urethral discharge amongst many other things. It also has a symbolic role in the fudngo ceremony, of the Niger Fula people, which is said to increase the fertility of cattle. P. malabaicum is often used as cattle fodder.

    Images

    Distribution

    Native to:

    Algeria, Andaman Is., Angola, Assam, Bangladesh, Benin, Botswana, Brazil West-Central, Burkina, Burundi, Cambodia, Cameroon, Caprivi Strip, Central African Repu, Chad, China South-Central, China Southeast, Congo, East Himalaya, Eritrea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, India, Ivory Coast, Jawa, Kenya, Laos, Lesser Sunda Is., Malawi, Mali, Mauritania, Mozambique, Myanmar, Namibia, Nepal, Nicobar Is., Niger, Nigeria, Northern Provinces, Northern Territory, Pakistan, Philippines, Queensland, Rwanda, Senegal, Sierra Leone, Sri Lanka, Sudan, Tanzania, Thailand, Togo, Uganda, Vietnam, West Himalaya, Western Australia, Yemen, Zambia, Zaïre, Zimbabwe

    Introduced into:

    Iraq, Windward Is.

    Piliostigma Hochst. appears in other Kew resources:

    First published in Flora 29: 598 (1846)

    Literature

    Flora of West Tropical Africa
    • Milne-Redhead in Hook. Ic. Pl. t. 3460 (1947).
    • in Flora 29: 598 (1846)
    Flora Zambesiaca
    • De Wit in Reinwardtia 3: 530 (1956).
    • in Flora 29: 598 (1846).
    Flora of Tropical East Africa
    • De Wit in Reinwardtia 3: 530 (1956)
    • Milne-Redh. in Hook., Ic. Pl. 35, t. 3460 (1947)
    • in Flora 29: 598 (1846)

    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

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