1. Family: Fabaceae Lindl.
    1. Astragalus L.

      1. This genus is accepted, and its native range is Temp. & Subtropical Old World to Tropical African Mountains, N. America to Honduras, Ecuador to S. South America.

    [FTEA]

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

    Habit
    Herbs or subshmbs
    Leaves
    Leaves imparipinnate or, outside the Flora area, paripinnate with the rhachis ending in a spine; stipules entire, rather persistent; leaflets entire, not stipellate
    Flowers
    Flowers in axillary racemes; bracts, and sometimes bracteoles, present
    Calyx
    Calyx tubular, campanulate or inflated with 5 subequal teeth or, as in the Flora area, the upper 2 shorter
    Corolla
    Standard glabrous or rarely, outside the Flora area, silky or tomentose, much longer than wide, without appendages, tapering gradually to the base; blades of wings and keel auriculate, ± oblong, of about equal length, the keel slightly pouched at the sides and here adhering to the wings
    Stamens
    Vexillary filament free (except in one Himalayan species), straight at the base, the other 9 filaments united for most of their length, the free parts curved upwards, not dilated at the tip; anthers all alike
    Pistil
    Ovary sessile or stipitate, 2–∞-ovulate; style filiform, glabrous; stigma small, terminal, not pilose
    Fruits
    Pod sessile or stipitate, usually inflated, the lower suture usually, and the upper sometimes, impressed, usually wholly or partly divided lengthwise by a vertical septum growing up from the lower suture
    Seeds
    Seeds kidney-shaped, with a thread-like funicle.
    [LOWO]

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

    Vernacular
    locoweeds, milk vetches
    Habit
    Shrubs and herbs
    Ecology
    Mainly mediterranean, warm to cool temperate, and semi-arid to arid continental temperate grassland and shrubland, a few montane to dry tropical and subtropical
    Distribution
    mainly N temperate, from S and C Europe (chiefly Mediterranean region, including N Africa), Middle East, SW Asia and Sino-Himalayan region to W China (c. 1200 spp., most diverse in Turkey, Iran and Afghanistan); also E Europe to C Asia, Mongolia, Siberia, NE China and Japan (c. 620 spp., mostly in former USSR); western N America (c. 380 spp.); S America (c. 100 spp.); Africa (1 sp.)
    Note
    Astracantha has been returned to Astragalus on both molecular and morphological grounds (Liston & Wheeler, 1994; Sanderson & Liston, 1995; Zarre & Podlech, 1995); single taxa have been split off as Erophaca, Biserrula, Barnebyella and Ophiocarpus (q.v.). Kazempour Osaloo et al. (2003) find the latter two genera to be well supported within Astragalus; molecular data support the separation of the first two genera and several other isolated species, but wider sampling is still needed

    Polhill (1981a, h & i) recognised Galegeae and Carmichaelieae as distinct tribes. Within the Galegeae he distinguished four subtribes: Coluteinae, Astragalinae, Galeginae and Glycyrrhizinae. Polhill (1994) added a fifth subtribe Alhagiinae. More recent studies suggest that the Carmichaelieae are best treated as another subtribe within Galegeae, and this is followed here. However, these studies also point strongly to the polyphyly of Galegeae (Wagstaff et al., 1999; Wojciechowski et al., 2000; 2004). Polhill’s concept of Galegeae appears to be in the process of disintegration into a number of smaller tribes, but decisions on this would be premature until constituent taxa are sampled more comprehensively, and putative monophyletic groups can be substantiated by morphological and other systematic data.

    Glycyrrhizinae Rydb. was recognised by Polhill (1981h) as very distinct, but he left it as ‘an outlying subtribe of Galegeae’. Molecular analyses have confirmed the outlying position of Glycyrrhiza (Fig. 53). Although it is a member of the Inverted Repeat Lacking Clade (IRLC), it forms a basal grade or sister group, along with such woody millettioid genera as Callerya and Wisteria, to the whole of the rest of the IRLC (Sanderson & Liston, 1995; Wojciechowski et al., 2000; 2004).

    Galeginae, containing the single genus Galega (Fig. 53), is also isolated from the rest of ‘Galegeae’. In Wojciechowski et al. (2000), it falls near the base of their Vicioid clade (Fig. 53) and is sister to the Cicereae. In Wojciechowski et al. (2004), Galega is sister to a combined Cicereae-Trifolieae-Fabeae clade, although with relatively poor support. Such placements have not been suggested before and careful morphological studies are needed to see if they support the relationships suggested by molecular analyses. One of the prominent features of Cicereae is the craspedodromous venation of the leaflets and while the leaflet venation in Galega is not truly craspedodromous, Polhill (1981h) does give ‘leaflet-nerves to margin or nearly so’ as one of the characteristics of Galeginae.

    Astragalinae contains, inter alia, what is probably the largest genus of flowering plants — Astragalus — with 2300–2500 species. Wojciechowski et al. (1999; 2000) have shown that their Astragalean clade (Fig. 53) is itself made up of several distinct clades. The first of these includes the vast majority of the species of Astragalus. Wojciechowski et al. (1999) and Kazempour Osaloo et al. (2003) find good support for Astragalus sens. strict. as a monophyletic group, i.e., excluding only a very few outlying species (mentioned below). Wojciechowski et al. (1999) also showed that genera such as Oxytropis, Sutherlandia and Swainsona, which are morphologically very similar to Astragalus (although they have never been formally combined within it), are in fact distinct and not nested within Astragalus. Sanderson & Liston (1995) and Wojciechowski et al. (1999, and references therein) have clearly shown that the vast majority of New World Astragalus form a single clade in which most of the species have chromosome numbers in an aneuploid series: n = 11, 12, 13, 14 or 15 (Spellenberg, 1976). The Old World taxa, on the other hand, have chromosome numbers that are euploid: n = 8, 16, 32 etc., and polyploids are common (Spellenberg, 1976). All the studies of Liston and his group (e.g., Liston & Wheeler, 1994) demonstrate that Astragalus, far from being a taxonomic ragbag, does in fact form a monophyletic group in which speciation has been particularly active; possible mechanisms have been discussed by Barneby (1964) and Polhill (1981h).

    A second small clade, sister to Astragalus, includes Biserrula pelecinus L. and Astragalus epiglottis L. These are both annuals of the Mediterranean Basin and N Africa. Both have only five fertile anthers and dorsiventrally compressed pods. The position of Biserrula has long been disputed but it has often been regarded as no more distinct from Astragalus than some of the other monotypic genera that have from time to time been split off, usually on account of their distinctive pods (Barneby 1964: 26). The position of Biserrula at the base of the Astragalus clade (e.g., in Wojciechowski et al., 1999, 2001; Kazempour Osaloo et al., 2003), however, supports its current treatment as a separate genus. Astragalus epiglottis, however, does not seem to have been treated anywhere above sectional rank.

    The third clade (Fig. 53) within the Astragalean clade of Wojciechowski et al. (1999; 2001), i.e., the Coluteoid clade, contains much of Polhill’s (1981h) Coluteinae, plus Astragalus sinicus L. and A. complanatus Bunge and the Southern Hemisphere carmichaelioid group. The group has a very scattered distribution, with the two Astragalus species occurring in E Asia, Lessertia and Sutherlandia in southern Africa, the carmichaelioids (including Swainsona) in Australia and New Zealand, Colutea widespread in continental Eurasia and NE Africa, and various smaller genera such as Smirnowia and Eremosparton restricted to C Asia. Wojciechowski et al. (1999) point out that Astragalus sinicus and A. complanatus resemble other Coluteinae in possessing non-interlocking wing and keel petals and a ciliate style (Lavin & Delgado, 1990). However, they also note that Wenniger (1991) has found stylar hairs to be quite widely scattered among sections of Old World Astragalus, and suggest that the character may have arisen several times. Kazempour Osaloo et al. (2003) found that Astragalus vogelii (Webb) Bornm., from N Africa, Arabia, W Asia, Pakistan and India, grouped with Colutea in their analyses, and they erected a new genus Podlechiella Maassoumi & Kazempour Osaloo (p. 22) to account for this species. This decision appears premature before detailed molecular analyses of the Coluteinae are available, and it is not accepted here. The genera separated as Carmichaelieae by Polhill (1981i) also appear best placed in this clade (Heenan, 1998a; Wagstaff et al., 1999). They have been shown to form a monophyletic group, confined to Australia and New Zealand.

    The fourth clade (Fig. 53), is made up of a monophyletic and distinct Oxytropis. The isolation of Oxytropis is reassuring but somewhat surprising as members of the genus are often morphologically extremely similar to species of Astragalus and can only be distinguished by the pointed keel petals and by the pod septum that arises from the adaxial suture, not the abaxial as in Astragalus. Species of Oxytropis are widespread in the north temperate regions, often on mountains.

    Three further groups of genera, exemplified by a) Erophaca baetica Boiss. (=Astragalus lusitanicus Lam.); b) Chesneya plus Gueldenstaedtia and c) Caragana, Calophaca and Halimodendron, are sister either to the Astragalean clade, or to the Astragalean plus Vicioid clade (Wojciechowski et al., 2000). Relatively poor sampling within these groups has resulted in different analyses suggesting different divisions and placements. Some of the genera are poorly known and of restricted distribution (e.g., Oreophysa and Tibetia), but others are more widespread, including Chesneya and Gueldenstaedtia.

    Erophaca baetica, together with Chesneya and Gueldenstaedtia (note that G. himalaica Baker, the exemplar of the genus in the supertree, is now placed in Tibetia) are part of the sister group to the Astragalean clade (Wojciechowski et al., 2001). Caragana, Calophaca and Halimodendron, on the other hand, form a monophyletic group sister to the Hedysaroid clade, and are here placed in Hedysareae. Sanderson & Wojciechowski (1996) and Wojciechowski et al. (2000; 2004) show that Alhagi appears to be best placed in Hedysareae, in agreement with Hutchinson (1964).

    In a tribe that is so clearly polyphyletic it is difficult to know how to arrange the constituent genera. For the purposes of this book, genera recognised in the tribe largely follow Polhill (1981h; 1994), although with modifications resulting from more recent research. The two major realignments since Polhill (1994) are the transfer of the genera mentioned above to the Hedysareae and the inclusion of the Carmichaelieae (Polhill, 1981i) within Galegeae. The arrangement of genera in this treatment is that suggested by the supertree of Wojciechowski et al. (2001). Not all genera have been sampled; genera not represented in the supertree have been intercalated in positions that appear to be appropriate from other data in, e.g., Polhill (1981h). Likewise, with the exception of Astragalus lusitanicus, here treated as Erophaca, the isolated species of Astragalus revealed by the supertree have not been treated separately. Podlech’s (1994) placement of three isolated Astragalus species as genera (Biserrula, Ophiocarpus and Barnebyella) has been followed here.

    Although we have suggested that it is premature to re-circumscribe tribal limits in this group, the bulk of taxa (i.e., those comprising Polhill’s subtribes Astragalinae [for the most part] and Coluteinae, and tribe Carmichaelieae), are likely to become recognised as a more narrowly defined tribe Astragaleae. As treated here the Galegeae sens. lat. comprises 24 genera and (2880)–3030–(3180) species.

    [FZ]

    Leguminosae, various authors. Flora Zambesiaca 3:7. 2003

    Habit
    Herbs or subshrubs.
    Leaves
    Leaves imparipinnate, or paripinnate with a spine at the end of the rhachis (outside the Flora Zambesiaca area); leaflets entire; stipules entire, often foliaceous persistent; stipels absent.
    Inflorescences
    Inflorescence a many-flowered axillary raceme; bracts present; bracteoles present or absent.
    Calyx
    Calyx tubular, campanulate or inflated, divided into 5 subequal teeth or the upper two shorter.
    Corolla
    Standard much longer than wide, gradually narrowed to the base, glabrous, or rarely silky or tomentose (outside the Flora Zambesiaca area); wings oblong to oblong-ovate, clawed, the blades auriculate; keel petals ± as long as the wings, oblong, the lamina slightly pouched above the auricle and adhering to the wings.
    Stamens
    Vexillary stamen free (except in one Himalayan species), straight at the base; filament sheath long; free parts of the filaments curved upwards, filiform; anthers uniform.
    Pistil
    Ovary sessile or stipitate, 2–many-ovulate; style filiform, glabrous with a small terminal stigma.
    Fruits
    Pod sessile or stipitate, usually inflated, with the lower suture usually impressed, the upper one sometimes so, usually partly or wholly divided by a vertical false septum growing from the lower suture.
    Seeds
    Seeds reniform.
    [LOWO]

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

    Habit
    Herb
    Ecology
    Continental temperate and mediterranean grassland and desert
    Distribution
    Iraq to Afghanistan and NW India
    Note
    Kept distinct from Astragalus by Podlech pending molecular evidence, mainly because of the long narrow subcylindrical unilocular pods, often somewhat constricted between the seeds; Kazempour Osaloo et al. (2003), however, find O. aitchisonii (Baker) Podlech to be well supported within Astragalus

    Polhill (1981a, h & i) recognised Galegeae and Carmichaelieae as distinct tribes. Within the Galegeae he distinguished four subtribes: Coluteinae, Astragalinae, Galeginae and Glycyrrhizinae. Polhill (1994) added a fifth subtribe Alhagiinae. More recent studies suggest that the Carmichaelieae are best treated as another subtribe within Galegeae, and this is followed here. However, these studies also point strongly to the polyphyly of Galegeae (Wagstaff et al., 1999; Wojciechowski et al., 2000; 2004). Polhill’s concept of Galegeae appears to be in the process of disintegration into a number of smaller tribes, but decisions on this would be premature until constituent taxa are sampled more comprehensively, and putative monophyletic groups can be substantiated by morphological and other systematic data.

    Glycyrrhizinae Rydb. was recognised by Polhill (1981h) as very distinct, but he left it as ‘an outlying subtribe of Galegeae’. Molecular analyses have confirmed the outlying position of Glycyrrhiza (Fig. 53). Although it is a member of the Inverted Repeat Lacking Clade (IRLC), it forms a basal grade or sister group, along with such woody millettioid genera as Callerya and Wisteria, to the whole of the rest of the IRLC (Sanderson & Liston, 1995; Wojciechowski et al., 2000; 2004).

    Galeginae, containing the single genus Galega (Fig. 53), is also isolated from the rest of ‘Galegeae’. In Wojciechowski et al. (2000), it falls near the base of their Vicioid clade (Fig. 53) and is sister to the Cicereae. In Wojciechowski et al. (2004), Galega is sister to a combined Cicereae-Trifolieae-Fabeae clade, although with relatively poor support. Such placements have not been suggested before and careful morphological studies are needed to see if they support the relationships suggested by molecular analyses. One of the prominent features of Cicereae is the craspedodromous venation of the leaflets and while the leaflet venation in Galega is not truly craspedodromous, Polhill (1981h) does give ‘leaflet-nerves to margin or nearly so’ as one of the characteristics of Galeginae.

    Astragalinae contains, inter alia, what is probably the largest genus of flowering plants — Astragalus — with 2300–2500 species. Wojciechowski et al. (1999; 2000) have shown that their Astragalean clade (Fig. 53) is itself made up of several distinct clades. The first of these includes the vast majority of the species of Astragalus. Wojciechowski et al. (1999) and Kazempour Osaloo et al. (2003) find good support for Astragalus sens. strict. as a monophyletic group, i.e., excluding only a very few outlying species (mentioned below). Wojciechowski et al. (1999) also showed that genera such as Oxytropis, Sutherlandia and Swainsona, which are morphologically very similar to Astragalus (although they have never been formally combined within it), are in fact distinct and not nested within Astragalus. Sanderson & Liston (1995) and Wojciechowski et al. (1999, and references therein) have clearly shown that the vast majority of New World Astragalus form a single clade in which most of the species have chromosome numbers in an aneuploid series: n = 11, 12, 13, 14 or 15 (Spellenberg, 1976). The Old World taxa, on the other hand, have chromosome numbers that are euploid: n = 8, 16, 32 etc., and polyploids are common (Spellenberg, 1976). All the studies of Liston and his group (e.g., Liston & Wheeler, 1994) demonstrate that Astragalus, far from being a taxonomic ragbag, does in fact form a monophyletic group in which speciation has been particularly active; possible mechanisms have been discussed by Barneby (1964) and Polhill (1981h).

    A second small clade, sister to Astragalus, includes Biserrula pelecinus L. and Astragalus epiglottis L. These are both annuals of the Mediterranean Basin and N Africa. Both have only five fertile anthers and dorsiventrally compressed pods. The position of Biserrula has long been disputed but it has often been regarded as no more distinct from Astragalus than some of the other monotypic genera that have from time to time been split off, usually on account of their distinctive pods (Barneby 1964: 26). The position of Biserrula at the base of the Astragalus clade (e.g., in Wojciechowski et al., 1999, 2001; Kazempour Osaloo et al., 2003), however, supports its current treatment as a separate genus. Astragalus epiglottis, however, does not seem to have been treated anywhere above sectional rank.

    The third clade (Fig. 53) within the Astragalean clade of Wojciechowski et al. (1999; 2001), i.e., the Coluteoid clade, contains much of Polhill’s (1981h) Coluteinae, plus Astragalus sinicus L. and A. complanatus Bunge and the Southern Hemisphere carmichaelioid group. The group has a very scattered distribution, with the two Astragalus species occurring in E Asia, Lessertia and Sutherlandia in southern Africa, the carmichaelioids (including Swainsona) in Australia and New Zealand, Colutea widespread in continental Eurasia and NE Africa, and various smaller genera such as Smirnowia and Eremosparton restricted to C Asia. Wojciechowski et al. (1999) point out that Astragalus sinicus and A. complanatus resemble other Coluteinae in possessing non-interlocking wing and keel petals and a ciliate style (Lavin & Delgado, 1990). However, they also note that Wenniger (1991) has found stylar hairs to be quite widely scattered among sections of Old World Astragalus, and suggest that the character may have arisen several times. Kazempour Osaloo et al. (2003) found that Astragalus vogelii (Webb) Bornm., from N Africa, Arabia, W Asia, Pakistan and India, grouped with Colutea in their analyses, and they erected a new genus Podlechiella Maassoumi & Kazempour Osaloo (p. 22) to account for this species. This decision appears premature before detailed molecular analyses of the Coluteinae are available, and it is not accepted here. The genera separated as Carmichaelieae by Polhill (1981i) also appear best placed in this clade (Heenan, 1998a; Wagstaff et al., 1999). They have been shown to form a monophyletic group, confined to Australia and New Zealand.

    The fourth clade (Fig. 53), is made up of a monophyletic and distinct Oxytropis. The isolation of Oxytropis is reassuring but somewhat surprising as members of the genus are often morphologically extremely similar to species of Astragalus and can only be distinguished by the pointed keel petals and by the pod septum that arises from the adaxial suture, not the abaxial as in Astragalus. Species of Oxytropis are widespread in the north temperate regions, often on mountains.

    Three further groups of genera, exemplified by a) Erophaca baetica Boiss. (=Astragalus lusitanicus Lam.); b) Chesneya plus Gueldenstaedtia and c) Caragana, Calophaca and Halimodendron, are sister either to the Astragalean clade, or to the Astragalean plus Vicioid clade (Wojciechowski et al., 2000). Relatively poor sampling within these groups has resulted in different analyses suggesting different divisions and placements. Some of the genera are poorly known and of restricted distribution (e.g., Oreophysa and Tibetia), but others are more widespread, including Chesneya and Gueldenstaedtia.

    Erophaca baetica, together with Chesneya and Gueldenstaedtia (note that G. himalaica Baker, the exemplar of the genus in the supertree, is now placed in Tibetia) are part of the sister group to the Astragalean clade (Wojciechowski et al., 2001). Caragana, Calophaca and Halimodendron, on the other hand, form a monophyletic group sister to the Hedysaroid clade, and are here placed in Hedysareae. Sanderson & Wojciechowski (1996) and Wojciechowski et al. (2000; 2004) show that Alhagi appears to be best placed in Hedysareae, in agreement with Hutchinson (1964).

    In a tribe that is so clearly polyphyletic it is difficult to know how to arrange the constituent genera. For the purposes of this book, genera recognised in the tribe largely follow Polhill (1981h; 1994), although with modifications resulting from more recent research. The two major realignments since Polhill (1994) are the transfer of the genera mentioned above to the Hedysareae and the inclusion of the Carmichaelieae (Polhill, 1981i) within Galegeae. The arrangement of genera in this treatment is that suggested by the supertree of Wojciechowski et al. (2001). Not all genera have been sampled; genera not represented in the supertree have been intercalated in positions that appear to be appropriate from other data in, e.g., Polhill (1981h). Likewise, with the exception of Astragalus lusitanicus, here treated as Erophaca, the isolated species of Astragalus revealed by the supertree have not been treated separately. Podlech’s (1994) placement of three isolated Astragalus species as genera (Biserrula, Ophiocarpus and Barnebyella) has been followed here.

    Although we have suggested that it is premature to re-circumscribe tribal limits in this group, the bulk of taxa (i.e., those comprising Polhill’s subtribes Astragalinae [for the most part] and Coluteinae, and tribe Carmichaelieae), are likely to become recognised as a more narrowly defined tribe Astragaleae. As treated here the Galegeae sens. lat. comprises 24 genera and (2880)–3030–(3180) species.

    [LOWO]

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

    Habit
    Herb
    Ecology
    Arid and semi-arid mediterranean and subtropical grassland and shrubland
    Distribution
    Iran, Afghanistan, Pakistan
    Note
    Originally described as a Dorycnium, but according to Podlech (1994) it is neither this nor Astragalus; Kazempour Osaloo et al. (2003), however, find B. calycina (Stocks) Podlech to be well supported within Astragalus; the pod is unilocular and 1-seeded, the petals are hairy and the calyx elongates during anthesis

    Polhill (1981a, h & i) recognised Galegeae and Carmichaelieae as distinct tribes. Within the Galegeae he distinguished four subtribes: Coluteinae, Astragalinae, Galeginae and Glycyrrhizinae. Polhill (1994) added a fifth subtribe Alhagiinae. More recent studies suggest that the Carmichaelieae are best treated as another subtribe within Galegeae, and this is followed here. However, these studies also point strongly to the polyphyly of Galegeae (Wagstaff et al., 1999; Wojciechowski et al., 2000; 2004). Polhill’s concept of Galegeae appears to be in the process of disintegration into a number of smaller tribes, but decisions on this would be premature until constituent taxa are sampled more comprehensively, and putative monophyletic groups can be substantiated by morphological and other systematic data.

    Glycyrrhizinae Rydb. was recognised by Polhill (1981h) as very distinct, but he left it as ‘an outlying subtribe of Galegeae’. Molecular analyses have confirmed the outlying position of Glycyrrhiza (Fig. 53). Although it is a member of the Inverted Repeat Lacking Clade (IRLC), it forms a basal grade or sister group, along with such woody millettioid genera as Callerya and Wisteria, to the whole of the rest of the IRLC (Sanderson & Liston, 1995; Wojciechowski et al., 2000; 2004).

    Galeginae, containing the single genus Galega (Fig. 53), is also isolated from the rest of ‘Galegeae’. In Wojciechowski et al. (2000), it falls near the base of their Vicioid clade (Fig. 53) and is sister to the Cicereae. In Wojciechowski et al. (2004), Galega is sister to a combined Cicereae-Trifolieae-Fabeae clade, although with relatively poor support. Such placements have not been suggested before and careful morphological studies are needed to see if they support the relationships suggested by molecular analyses. One of the prominent features of Cicereae is the craspedodromous venation of the leaflets and while the leaflet venation in Galega is not truly craspedodromous, Polhill (1981h) does give ‘leaflet-nerves to margin or nearly so’ as one of the characteristics of Galeginae.

    Astragalinae contains, inter alia, what is probably the largest genus of flowering plants — Astragalus — with 2300–2500 species. Wojciechowski et al. (1999; 2000) have shown that their Astragalean clade (Fig. 53) is itself made up of several distinct clades. The first of these includes the vast majority of the species of Astragalus. Wojciechowski et al. (1999) and Kazempour Osaloo et al. (2003) find good support for Astragalus sens. strict. as a monophyletic group, i.e., excluding only a very few outlying species (mentioned below). Wojciechowski et al. (1999) also showed that genera such as Oxytropis, Sutherlandia and Swainsona, which are morphologically very similar to Astragalus (although they have never been formally combined within it), are in fact distinct and not nested within Astragalus. Sanderson & Liston (1995) and Wojciechowski et al. (1999, and references therein) have clearly shown that the vast majority of New World Astragalus form a single clade in which most of the species have chromosome numbers in an aneuploid series: n = 11, 12, 13, 14 or 15 (Spellenberg, 1976). The Old World taxa, on the other hand, have chromosome numbers that are euploid: n = 8, 16, 32 etc., and polyploids are common (Spellenberg, 1976). All the studies of Liston and his group (e.g., Liston & Wheeler, 1994) demonstrate that Astragalus, far from being a taxonomic ragbag, does in fact form a monophyletic group in which speciation has been particularly active; possible mechanisms have been discussed by Barneby (1964) and Polhill (1981h).

    A second small clade, sister to Astragalus, includes Biserrula pelecinus L. and Astragalus epiglottis L. These are both annuals of the Mediterranean Basin and N Africa. Both have only five fertile anthers and dorsiventrally compressed pods. The position of Biserrula has long been disputed but it has often been regarded as no more distinct from Astragalus than some of the other monotypic genera that have from time to time been split off, usually on account of their distinctive pods (Barneby 1964: 26). The position of Biserrula at the base of the Astragalus clade (e.g., in Wojciechowski et al., 1999, 2001; Kazempour Osaloo et al., 2003), however, supports its current treatment as a separate genus. Astragalus epiglottis, however, does not seem to have been treated anywhere above sectional rank.

    The third clade (Fig. 53) within the Astragalean clade of Wojciechowski et al. (1999; 2001), i.e., the Coluteoid clade, contains much of Polhill’s (1981h) Coluteinae, plus Astragalus sinicus L. and A. complanatus Bunge and the Southern Hemisphere carmichaelioid group. The group has a very scattered distribution, with the two Astragalus species occurring in E Asia, Lessertia and Sutherlandia in southern Africa, the carmichaelioids (including Swainsona) in Australia and New Zealand, Colutea widespread in continental Eurasia and NE Africa, and various smaller genera such as Smirnowia and Eremosparton restricted to C Asia. Wojciechowski et al. (1999) point out that Astragalus sinicus and A. complanatus resemble other Coluteinae in possessing non-interlocking wing and keel petals and a ciliate style (Lavin & Delgado, 1990). However, they also note that Wenniger (1991) has found stylar hairs to be quite widely scattered among sections of Old World Astragalus, and suggest that the character may have arisen several times. Kazempour Osaloo et al. (2003) found that Astragalus vogelii (Webb) Bornm., from N Africa, Arabia, W Asia, Pakistan and India, grouped with Colutea in their analyses, and they erected a new genus Podlechiella Maassoumi & Kazempour Osaloo (p. 22) to account for this species. This decision appears premature before detailed molecular analyses of the Coluteinae are available, and it is not accepted here. The genera separated as Carmichaelieae by Polhill (1981i) also appear best placed in this clade (Heenan, 1998a; Wagstaff et al., 1999). They have been shown to form a monophyletic group, confined to Australia and New Zealand.

    The fourth clade (Fig. 53), is made up of a monophyletic and distinct Oxytropis. The isolation of Oxytropis is reassuring but somewhat surprising as members of the genus are often morphologically extremely similar to species of Astragalus and can only be distinguished by the pointed keel petals and by the pod septum that arises from the adaxial suture, not the abaxial as in Astragalus. Species of Oxytropis are widespread in the north temperate regions, often on mountains.

    Three further groups of genera, exemplified by a) Erophaca baetica Boiss. (=Astragalus lusitanicus Lam.); b) Chesneya plus Gueldenstaedtia and c) Caragana, Calophaca and Halimodendron, are sister either to the Astragalean clade, or to the Astragalean plus Vicioid clade (Wojciechowski et al., 2000). Relatively poor sampling within these groups has resulted in different analyses suggesting different divisions and placements. Some of the genera are poorly known and of restricted distribution (e.g., Oreophysa and Tibetia), but others are more widespread, including Chesneya and Gueldenstaedtia.

    Erophaca baetica, together with Chesneya and Gueldenstaedtia (note that G. himalaica Baker, the exemplar of the genus in the supertree, is now placed in Tibetia) are part of the sister group to the Astragalean clade (Wojciechowski et al., 2001). Caragana, Calophaca and Halimodendron, on the other hand, form a monophyletic group sister to the Hedysaroid clade, and are here placed in Hedysareae. Sanderson & Wojciechowski (1996) and Wojciechowski et al. (2000; 2004) show that Alhagi appears to be best placed in Hedysareae, in agreement with Hutchinson (1964).

    In a tribe that is so clearly polyphyletic it is difficult to know how to arrange the constituent genera. For the purposes of this book, genera recognised in the tribe largely follow Polhill (1981h; 1994), although with modifications resulting from more recent research. The two major realignments since Polhill (1994) are the transfer of the genera mentioned above to the Hedysareae and the inclusion of the Carmichaelieae (Polhill, 1981i) within Galegeae. The arrangement of genera in this treatment is that suggested by the supertree of Wojciechowski et al. (2001). Not all genera have been sampled; genera not represented in the supertree have been intercalated in positions that appear to be appropriate from other data in, e.g., Polhill (1981h). Likewise, with the exception of Astragalus lusitanicus, here treated as Erophaca, the isolated species of Astragalus revealed by the supertree have not been treated separately. Podlech’s (1994) placement of three isolated Astragalus species as genera (Biserrula, Ophiocarpus and Barnebyella) has been followed here.

    Although we have suggested that it is premature to re-circumscribe tribal limits in this group, the bulk of taxa (i.e., those comprising Polhill’s subtribes Astragalinae [for the most part] and Coluteinae, and tribe Carmichaelieae), are likely to become recognised as a more narrowly defined tribe Astragaleae. As treated here the Galegeae sens. lat. comprises 24 genera and (2880)–3030–(3180) species.

    [LOWO]
    Use
    A few species are significant economically; A. gummifer Labill. (gum tragacanth) , is used for its gum in a wide range of industries (at least 20 other species are noted to produce gums [Duke, 1981]); A. mongholicus Bunge var. dahuricus (DC.) Podlech (=A. membranaceus Bunge) is a widely used traditional medicine; other species ( milk vetches ) are used for human food (edible pods, but some species toxic), forage, erosion control, fuelwood and as selenium and uranium indicators (Allen & Allen, 1981); some species are poisonous to livestock ( locoweeds )

    Images

    Distribution

    Native to:

    Afghanistan, Alabama, Alaska, Albania, Alberta, Aleutian Is., Algeria, Altay, Amur, Argentina Northeast, Argentina Northwest, Argentina South, Argentina South, Arizona, Arizona, Arkansas, Assam, Austria, Baleares, Baltic States, Belarus, Belgium, Bolivia, British Columbia, British Columbia, Bulgaria, Burundi, Buryatiya, California, Canary Is., Cape Verde, Central European Rus, Chad, Chile Central, Chile North, Chile South, China North-Central, China South-Central, China Southeast, Chita, Colorado, Connecticut, Corse, Cyprus, Czechoslovakia, Denmark, Djibouti, East Aegean Is., East European Russia, East Himalaya, Ecuador, Egypt, Eritrea, Ethiopia, Finland, Florida, France, Free State, Galápagos, Georgia, Germany, Great Britain, Greece, Greenland, Guatemala, Gulf States, Honduras, Hungary, Idaho, Illinois, India, Indiana, Inner Mongolia, Iowa, Iowa, Iran, Iraq, Ireland, Irkutsk, Italy, Japan, Kamchatka, Kansas, Kazakhstan, Kentucky, Kenya, Khabarovsk, Kirgizstan, Korea, Krasnoyarsk, Kriti, Kriti, Krym, Krym, Kuril Is., Kuwait, KwaZulu-Natal, Labrador, Lebanon-Syria, Libya, Louisiana, Madeira, Magadan, Maine, Malawi, Mali, Manchuria, Manitoba, Maryland, Mauritania, Mexico Central, Mexico Gulf, Mexico Northeast, Mexico Northwest, Mexico Southeast, Mexico Southwest, Michigan, Minnesota, Mississippi, Mississippi, Missouri, Mongolia, Mongolia, Montana, Montana, Morocco, Mozambique, Myanmar, Nebraska, Nebraska, Nepal, Netherlands, Nevada, Nevada, New Brunswick, New Hampshire, New Jersey, New Mexico, New Mexico, New York, Newfoundland, Niger, North Carolina, North Caucasus, North Dakota, North European Russi, Northern Provinces, Northwest European R, Northwest Territorie, Norway, Nunavut, Ohio, Ohio, Oklahoma, Oman, Ontario, Oregon, Pakistan, Pakistan, Palestine, Pennsylvania, Peru, Poland, Portugal, Primorye, Qinghai, Québec, Romania, Rwanda, Sakhalin, Sardegna, Saskatchewan, Saudi Arabia, Sicilia, Sinai, Socotra, Somalia, South Carolina, South Carolina, South Dakota, South Dakota, South European Russi, Spain, Sudan, Sweden, Switzerland, Tadzhikistan, Taiwan, Tanzania, Tennessee, Texas, Tibet, Transcaucasus, Tunisia, Turkey, Turkey-in-Europe, Turkmenistan, Tuva, Uganda, Ukraine, Uruguay, Utah, Utah, Uzbekistan, Vermont, Vietnam, Virginia, Washington, West Himalaya, West Siberia, West Virginia, Western Sahara, Wisconsin, Wyoming, Xinjiang, Yakutskiya, Yemen, Yugoslavia, Yukon, Zambia, Zaïre, Zimbabwe

    Introduced into:

    Masachusettes, New South Wales, New York, Queensland, South Australia

    Astragalus L. appears in other Kew resources:

    Date Reference Identified As Barcode Type Status
    unknown [s.n.], USA K000264015
    Rico, L. [1277], Bolivia K000295096
    Rico, L. [1511], Bolivia K000295174
    Rico, L. [1645], Bolivia K000295097
    Rico, L. [1879], Tunisia K000296052
    Rico, L. [1880], Tunisia K000296056
    Rico, L. [1915], Morocco K000564900
    Rico, L. [1925], Morocco K000564891
    Rico, L. [1927], Morocco K000564889
    Rico, L. [1928], Morocco K000564888
    Rico, L. [1940], Morocco K000564878
    Rico, L. [1709b], Armenia K000661661
    Rico, L. [1712], Armenia K000661665
    Rico, L. [1713], Armenia K000297241
    Rico, L. [1787], Armenia K000297288
    Rico, L. [1800], Armenia K000661343
    Rico, L. [2233], Syria K000764180
    Rico, L. [2261], Syria K000764205

    First published in Sp. Pl.: 755 (1753)

    Accepted by

    • Arevschatian, I.G. (2011). The subgenus Astragalus of the genus Astragalus in Southern Transcaucasia Takhtajania 1: 69-84.
    • Govaerts, R. (1995). World Checklist of Seed Plants 1(1, 2): 1-483, 529. MIM, Deurne.
    • Tutin, T.G. & al. (eds.) (1968). Flora Europaea 2: 1-469. Cambridge University Press.

    Literature

    Flora Zambesiaca
    • Gen. Pl., ed. 5: 335 (1754).
    • Sp. Pl.: 755 (1753)
    Flora of Tropical East Africa
    • L., Gen. Pl., ed. 5: 335 (1754)
    • Sp. Pl.: 755 (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 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