Fabaceae subfamilies

Subfamily Faboideae

Polhill (1981a, 1994a, 1997) has loosely divided the Faboideae into four main groups of tribes: 1. The basal tribes, Swartzieae (1) and Sophoreae (2), which are transitional to subfamily Caesalpinioideae; 2. the genistoid alliance, mainly temperate tribes occurring in both northern and southern hemispheres; 3. the tropical tribes with some woody tribes, such as Dalbergieae (4), Millettieae (7), and Robinieae (8), whose limits and phylogenetic history are unclear and with a number of predominantly herbaceous tribes such as Indigofereae (9), Desmodieae (11), Phaseoleae (10), and Aeschynomeneae (14), whose limits are better defined and which are considered to be relatively more advanced; and, 4. the temperate herbaceous tribes or epulvinate series which includes, for example, Galegeae (16), Hedysareae (18), Trifolieae (21), and Fabeae (Vicieae) (19).

The basal tribes are predominantly tropical and subtropical. Swartzieae has been placed in subfamily Caesalpinioideae or even considered to be a fourth subfamily, but the current, general consensus of opinion, among legume taxonomists, is that it should be in Faboideae (Cowan, 1981). Recent cladistic studies (Herendeen, 1995Herendeen, 1995:
Herendeen PS. 1995. Phylogenetic relationships of the tribe Swartzieae. In: M. Crisp and J.J. Doyle, eds., Advances in Legume Systematics 7: Phylogeny, pp. 123&-132. Royal Botanic Gardens, Kew, England.
), supported by rbcL data (Doyle et al., 1997), have indicated that Swartzieae and Sophoreae should be merged into a single tribe in the Faboideae.

The genistoid alliance was characterized (Polhill, 1994aPolhill, 1994a:
Polhill RM. 1994a. Classification of the Leguminosae. pp. xxx&-xlviii. In F.A. Bisby, J. Buckingham, and J.B. Harborne, eds., Phytochemical Dictionary of the Leguminosae. Chapman & Hall, London, England.
) by: progressive contraction of vegetative structures and inflorescences, progressive joining of stamens and dimorphic anthers, development on the seed of a hilar lobe from extension of the radicle, and an abundance of alkaloids as compared to other advanced tribal groupings. This alliance has three distinct regional groupings. Genisteae (30) and Thermopsideae (29) are in the northern hemisphere, Crotalarieae (27), Hypocalypteae (26), and Podalyrieae (25) are centered in southern Africa, and Bossiaeeae (23) and Mirbelieae (24) in Australia.

The tropical tribes are marked off by a series of character transitions (Polhill, 1994aPolhill, 1994a:
Polhill RM. 1994a. Classification of the Leguminosae. pp. xxx&-xlviii. In F.A. Bisby, J. Buckingham, and J.B. Harborne, eds., Phytochemical Dictionary of the Leguminosae. Chapman & Hall, London, England.
) to: strongly papilinoid flowers, staminal fusion late in development, more distinct stigmas, and the appearance of canavanine, a non-protein amino acid. The delimitation and separation of Millettieae from Sophoreae has remained difficult with the generic groupings within Millettieae unresolved. The more advanced tropical tribes centered in the Old World, Desmodieae (11), Indigofereae (9 ), Phaseoleae (10), and Psoraleae (12), are differentiated from those centered in the New World, Adesmieae (15), Amorpheae (6), and Aeschynomeneae (14), by their pollen wall structure (Ferguson, 1984Ferguson, 1984:
Ferguson I.K. 1984. Pollen morphology and biosystematics of the subfamily Papilionoideae (Leguminosae). In: Grant WF, ed. Plant Biosystematics. pp. 377&-394. Academic Press, Canada.
, and Ferguson and Skvarla, 1981Ferguson and Skvarla, 1981:
Ferguson IK and Skvarla JJ. 1981. The pollen morphology of the subfamily Papilionoideae (Leguminosae). In: R.M. Polhill and P.H. Raven, ed., Advances in Legume Systematics. Vol. 2. pt. 2. pp. 859&-896. Int. Legume Conf., Proc. 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England.
). The Old World tribes tend to have an increase in the thickness of the endexine and a reduction of the foot layer, and New World ones tend to have a reduction of the endexine and a thicker foot layer, usually associated with longer columellae.

The temperate herbaceous tribes are characterized by the lack of a foliar pulvinus, which correlates with a closed vascular system, and loss of secondary thickening (Polhill, 1981aPolhill, 1981a:
Polhill RM. 1981a. Papilionoideae. In R.M. Polhill and P.H. Raven, eds., Advances in Legume Systematics, v. 2, pt. 1, pp. 191&-208. Int. Legume Conf., Proc. 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England.
). This also correlates with a lack of the inverted repeat from the chloroplast DNA, stipels, ridge bundles in petioles and rhachides, anomalous secondary thickening, secretory reservoirs, and leucoanthocyanidins.

Subfamily Faboideae consists of 30 tribes (Polhill, 1981aPolhill, 1981a:
Polhill RM. 1981a. Papilionoideae. In R.M. Polhill and P.H. Raven, eds., Advances in Legume Systematics, v. 2, pt. 1, pp. 191&-208. Int. Legume Conf., Proc. 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England.
, 1994a, 1994b; Polhill and Raven, 1981Polhill and Raven, 1981:
Polhill RM and Raven PH, eds. 1981. Advances in Legume Systematics, v. 2, pt. 1, pp. 1–464. Int. Legume Conf., Proc. 1978, Kew, England Minis. Agric., Fisheries and Food, Richmond, England.
), 452 genera, and more than 12,725 species. Polhill (1981a) proposed merging tribe Coronilleae into Loteae, and did so in his latest classification scheme for the legumes (Polhill, 1994aPolhill, 1994a:
Polhill RM. 1994a. Classification of the Leguminosae. pp. xxx&-xlviii. In F.A. Bisby, J. Buckingham, and J.B. Harborne, eds., Phytochemical Dictionary of the Leguminosae. Chapman & Hall, London, England.
, 1994b), which reduced the number of tribes to 30. Faboideae are distributed throughout the world in all habitats, including aquatic ones. Three hundred and five genera have ten or fewer species, of which 102 are monotypic. One hundred and sixteen genera have more than ten and less than 100 species, 22 genera have 100 to 200 species, and nine genera have more than 200 species. Astragalus C. Linnaeus (3.16.15), with more than 2,000 species, is the largest genus of legumes and probably the largest genus of seed plants. The distributions, generic names, and parameters follow Polhill and Raven (1981)Polhill and Raven (1981):
Polhill RM and Raven PH, eds. 1981. Advances in Legume Systematics, v. 2, pt. 1, pp. 1–464. Int. Legume Conf., Proc. 1978, Kew, England Minis. Agric., Fisheries and Food, Richmond, England.
, except as noted.

At the Fourth International Legume Conference, 2–6 July 2001, Canberra, Australia, numerous cladistic analyses employing molecular and morphological data were presented for various Faboideae tribes and genera. In some cases, they supported the current classification, and in others rejected it. Extensive additionally studies are required to understand the Faboideae, and apparently the traditional classification will be significantly changed.

Neither mature fruit or seed material nor published data were available for the following 13 genera:

Burkilliodendron A.R.K.R. Sastry (3.7, subfamily Faboideae, tribe Millettieae)
Carrissoa E.G. Baker (3.10.79, subfamily Faboideae, tribe Phaseoleae)
Clitoriopsis R. Wilczek (3.10.17, subfamily Faboideae, tribe Phaseoleae)
Erichsenia W.B. Hemsley (3.24.05, subfamily Faboideae, tribe Mirbelieae)
Exostyles H.W. Schott (3.1.10, subfamily Faboideae, tribe Swartzieae)
Luzonia A.D.E. Elmer (3.10.23, subfamily Faboideae, tribe Phaseoleae)
Neocollettia W.B. Hemsley (3.11.26, subfamily Faboideae, tribe Desmodieae)
Petaladenium W.A. Ducke (3.2.25, subfamily Faboideae, tribe Sophoreae)
Sartoria P.E. Boissier & T.H.H. von Heldreich (3.18.05, subfamily Faboideae, tribe Hedysareae)
Sellocharis P.H.W. Taubert (3.30.07, subfamily Faboideae, tribe Genisteae)
Spongiocarpella G.P. Yakovlev & N. Ulziykhutag (3.16.13, subfamily Faboideae, tribe Galegeae)
Stirtonanthus B.-E. Van Wyk & A.L. Schutte (3.25.07, subfamily Faboideae, tribe Podalyrieae)
Weberbauerella O.E. Ulbrich (3.14.18, subfamily Faboideae, tribe Aeschynomeneae)

Only fruits or valves represented the following 7 genera: 

Dahlstedtia G.O.A. Malme (3.7, subfamily Faboideae, tribe Millettieae)
Margaritolobium H.A.T. Harms (3.7, subfamily Faboideae, tribe Millettieae)
Nephrodesmus A.K. Schindler (3.11.04, subfamily Faboideae, tribe Desmodieae)
Oryxis A. Delgado Salinas & G.P. Lewis (3.10.69A, subfamily Faboideae, tribe Phaseoleae)
Podolotus J.F. Royle (3.13.08, subfamily Faboideae, tribe Loteae)
Spirotropis E.L.R. Tulasne (3.2.32, subfamily Faboideae, tribe Sophoreae)
Sylvichadsia D.J. Du Puy & J.-N. Labat (3.7, subfamily Faboideae, tribe Millettieae)

For subfamily Faboideae unpublished data (pers. commun.) were supplied by reviewers of tribes: Aeschynomeneae, Vela E. Rudd; Amorpheae, Rupert C. Barneby; Cicereae, L.J.G. Van Der Maesen; Dalbergieae, R.T. Pennington, Vela E. Rudd, and K. Thothathri; Desmodieae, Shinobu Akiyama; Euchresteae, Hiroyoshi Ohashi; Galegeae, Rupert C. Barneby, Aaron Liston, Dieter Podlech, and Andrey Sytin; Genisteae, Chaia Clara Heyn; Hedysareae, K. Thothathri; Loteae, Ana M. Arambarri and Chaia Clara Heyn; Millettieae, Frits Adema and K. Thothathri; Phaseoleae, Paul R. Fantz, Alfonso Delgado Salinas, and L.J.G. Van Der Maesen; Robinieae, Matt Lavin; Sophoreae, Patrick S. Herendeen and Velva E. Rudd; Swartzieae, Patrick S. Herendeen; Thermopsideae, B.L. Turner; Trifolieae, Chaia Clara Heyn and Ernest Small; and Fabeae, Nigel Maxted.