Fabaceae () or Leguminosae, International Code of Nomenclature for algae, fungi, and plants. Article 18.5 states: "The following names, of long usage, are treated as validly published: ....Leguminosae (nom. alt.: Fabaceae; type: Faba Mill. =); ... When the Papilionaceae are regarded as a family distinct from the remainder of the Leguminosae, the name Papilionaceae is conserved against Leguminosae." English pronunciations are as follows: , and . commonly known as the legume, pea, or bean family, is a large and agriculturally important family of . It includes , , and perennial or annual plant , which are easily recognized by their fruit (legume) and their compound, stipule leaves. The family is widely distributed, and is the third-largest land plant family in number of species, behind only the Orchidaceae and Asteraceae, with about 765 genera and nearly 20,000 known species.Judd, W. S., Campbell, C. S. Kellogg, E. A. Stevens, P.F. Donoghue, M. J. (2002), Plant systematics: a phylogenetic approach, Sinauer Axxoc, 287–292. .
The five largest genera of the family are Astragalus (over 3,000 species), Acacia (over 1,000 species), Indigofera (around 700 species), Crotalaria (around 700 species), and Mimosa (around 400 species), which constitute about a quarter of all legume species. The c. 19,000 known legume species amount to about 7% of flowering plant species. Fabaceae is the most common family found in tropical rainforests and dry forests of the Americas and Africa.
Recent molecular and morphological evidence supports the fact that the Fabaceae is a single monophyletic family.Lewis G., Schrire B., Mackinder B. and Lock M. 2005. (eds.) Legumes of the world. The Royal Botanic Gardens, Kew, Reino Unido. 577 pages. 2005. . This conclusion has been supported not only by the degree of interrelation shown by different groups within the family compared with that found among the Leguminosae and their closest relations, but also by all the recent phylogenetic studies based on DNA sequences.Doyle, J. J., J. A. Chappill, C.D. Bailey, & T. Kajita. 2000. Towards a comprehensive phylogeny of legumes: evidence from rbcL sequences and non-molecular data. pp. 1–20 in Advances in legume systematics, part 9, (P. S. Herendeen and A. Bruneau, eds.). Royal Botanic Gardens, Kew, UK. These studies confirm that the Fabaceae are a monophyletic group that is closely related to the families Polygalaceae, Surianaceae and Quillajaceae and that they belong to the order Fabales.
Along with the , some fruits and tropical roots, a number of Leguminosae have been a staple human food for millennia and their use is closely related to human evolution.Burkart, A. Leguminosas. In: Dimitri, M. 1987. Enciclopedia Argentina de Agricultura y Jardinería. Tomo I. Descripción de plantas cultivadas. Editorial ACME S.A.C.I., Buenos Aires. pages: 467–538.
The family Fabaceae includes a number of plants that are common in agriculture including Glycine max (soybean), Phaseolus (beans), Pisum sativum (pea), Cicer arietinum (), Vicia faba (Vicia faba), Medicago sativa (alfalfa), Arachis hypogaea (peanut), Ceratonia siliqua (carob), Trigonella foenum-graecum (fenugreek), and Glycyrrhiza glabra (liquorice). A number of species are also weedy pests in different parts of the world, including Cytisus scoparius (broom), Robinia pseudoacacia (black locust) , Ulex europaeus (gorse), Pueraria montana (kudzu), and a number of Lupinus species.
Many species have leaves with structures that attract which protect the plant from herbivore insects (a form of mutualism). Extrafloral nectaries are common among the Mimosoideae and the Caesalpinioideae, and are also found in some Faboideae (e.g. Vicia sativa). In some Acacia, the modified hollow stipules are inhabited by ants and are known as domatia.
In the Caesalpinioideae, the flowers are often zygomorphic, as in Cercis, or nearly symmetrical with five equal petals, as in Bauhinia. The upper petal is the innermost one, unlike in the Faboideae. Some species, like some in the genus Senna, have asymmetric flowers, with one of the lower petals larger than the opposing one, and the style bent to one side. The calyx, corolla, or stamens can be showy in this group.
In the Mimosoideae, the flowers are actinomorphic and arranged in globose inflorescences. The petals are small and the stamens, which can be more than just 10, have long, coloured filaments, which are the showiest part of the flower. All of the flowers in an inflorescence open at once.
In the Faboideae, the flowers are zygomorphic, and have a specialized structure. The upper petal, called the banner or standard, is large and envelops the rest of the petals in bud, often reflexing when the flower blooms. The two adjacent petals, the wings, surround the two bottom petals. The two bottom petals are fused together at the apex (remaining free at the base), forming a boat-like structure called the keel. The stamens are always ten in number, and their filaments can be fused in various configurations, often in a group of nine stamens plus one separate stamen. Various genes in the CYCLOIDEA (CYC)/DICHOTOMA (DICH) family are expressed in the upper (also called dorsal or adaxial) petal; in some species, such as Cadia, these genes are expressed throughout the flower, producing a radially symmetrical flower.
The Fabaceae have an abundant and diverse fossil record, especially for the Tertiary period. Fossils of flowers, fruit, leaves, wood and pollen from this period have been found in numerous locations.Crepet, W. L., and P. S. Herendeen. 1992. Papilionoid flowers from the early Eocene of south eastern North America. Pages 43–55 in Advances in Legume Systematics, part 4, the fossil record (P. S. Herendeen and D. L. Dilcher, eds.). Royal Botanic Gardens, Kew, UK.Herendeen, P. S. 1992. The fossil history of Leguminosae from the Eocene of south eastern North America. Pages 85–160 in Advances in Legume Systematics, part 4, the fossil record (Herendeen, P. S., and D. L. Dilcher, eds.). Royal Botanic Gardens, Kew, UK.Herendeen, P. S. 2001. The fossil record of the Leguminosae: recent advances. In Legumes Down Under: the Fourth International Legume conference, Abstracts, 34–35. Australian National University, Canberra, Australia.Herendeen, P. S., and S. Wing. 2001. Papilionoid legume fruits and leaves from the Palaeocene of north western Wyoming. Botany 2001 Abstracts, published by Botanical Society of America (http://www.botany2001.org/).Wing, S. L., F. Herrera, and C. Jaramillo. 2004. A Palaeocene flora from the Cerrajón Formation, Guajíra Peninsula, north eastern Colombia. Pages 146–147 in VII International Organization of Paleobotany Conference Abstracts (21–26 March). Museo Egidio Feruglio, Trelew, Argentina. The earliest fossils that can be definitively assigned to the Fabaceae appeared in the early Palaeocene (approximately 65 million years ago). Representatives of the 3 sub-families traditionally recognised as being members of the Fabaceae – Cesalpinioideae, Papilionoideae and Mimosoideaeas well as members of the large clades within these sub-familiessuch as the genistoideshave been found in periods later, starting between 55 and 50 million years ago. In fact, a wide variety of taxa representing the main lineages in the Fabaceae have been found in the fossil record dating from the middle to the late Eocene, suggesting that the majority of the modern Fabaceae groups were already present and that a broad diversification occurred during this period. Therefore, the Fabaceae started their diversification approximately 60 million years ago and the most important clades separated 50 million years ago.Bruneau, A., Lewis, G. P., Herendeen, P. S., Schrire, B., & Mercure, M. 2008b. Biogeographic patterns in early-diverging clades of the Leguminosae. Pp. 98–99, in Botany 2008. Botany without Borders. Botanical The age of the main Cesalpinioideae clades have been estimated as between 56 and 34 million years and the basal group of the Mimosoideae as 44 ± 2.6 million years. The division between Mimosoideae and Faboideae is dated as occurring between 59 and 34 million years ago and the basal group of the Faboideae as 58.6 ± 0.2 million years ago. It has been possible to date the divergence of some of the groups within the Faboideae, even though diversification within each genus was relatively recent. For instance, Astragalus separated from the Oxytropis 16 to 12 million years ago. In addition, the separation of the aneuploidy species of Neoastragalus started 4 million years ago. Inga, another genus of the Papilionoideae with approximately 350 species, seems to have diverged in the last 2 million years.Wojciechowski, M. F. 2003. Reconstructing the phylogeny of legumes (Leguminosae): An early 21st century perspective. Pp. 5–35, in Klitgaard, B. B. & Bruneau, A. (eds), Advances in Legume Systematics, Part 10, Higher Level Systematics. Royal Botanic Gardens, Kew.Wojciechowski, Martin F., Johanna Mahn, and Bruce Jones. 2006. Fabaceae. legumes. Version 14 June 2006. The Tree of Life Web Project, http://tolweb.org/
It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along the Tethys Ocean during the Palaeogene Period.
The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event. Several different pathways have been implicated as donating duplicated genes to the pathways need for nodulation. The main donors to the pathway were the genes associated with the arbuscular mycorrhiza symbiosis genes, the pollen tube formation genes and the haemoglobin genes. One of the main genes shown to be shared between the arbuscular mycorrhiza pathway and the nodulation pathway is SYMRK and it is involved in the plant-bacterial recognition. The pollen tube growth is similar to the infection thread development in that infection threads grow in a polar manner that is similar to a pollen tubes polar growth towards the ovules. Both pathways include the same type of enzymes, pectin-degrading cell wall enzymes. The enzymes needed to reduce nitrogen, nitrogenases, require a substantial input of ATP but at the same time are sensitive to free oxygen. To meet the requirements of this paradoxical situation, the plants express a type of haemoglobin called leghaemoglobin that is believed to be recruited after a duplication event. These three genetic pathways are believed to be part of a gene duplication event then recruited to work in nodulation.
The rhizobia and their hosts must be able to recognize each other for nodule formation to commence. Rhizobia are specific to particular host species although a rhizobia species may often infect more than one host species. This means that one plant species may be infected by more than one species of bacteria. For example, nodules in Acacia senegal can contain seven species of rhizobia belonging to three different genera. The most distinctive characteristics that allow rhizobia to be distinguished apart are the rapidity of their growth and the type of root nodule that they form with their host. Root nodules can be classified as being either indeterminate, cylindrical and often branched, and determinate, spherical with prominent lenticels. Indeterminate nodules are characteristic of legumes from temperate climates, while determinate nodules are commonly found in species from tropical or subtropical climates.
Nodule formation is common throughout the Fabaceae. It is found in the majority of its members that only form an association with rhizobia, which in turn form an exclusive symbiosis with the Fabaceae (with the exception of Parasponia, the only genus of the 18 Ulmaceae genera that is capable of forming nodules). Nodule formation is present in all the Fabaceae sub-families, although it is less common in the Caesalpinioideae. All types of nodule formation are present in the subfamily Papilionoideae: indeterminate (with the meristem retained), determinate (without meristem) and the type included in Aeschynomene. The latter two are thought to be the most modern and specialised type of nodule as they are only present in some lines of the subfamily Papilionoideae. Even though nodule formation is common in the two Monophyly subfamilies Papilionoideae and Mimosoideae they also contain species that do not form nodules. The presence or absence of nodule-forming species within the three sub-families indicates that nodule formation has arisen several times during the evolution of the Fabaceae and that this ability has been lost in some lineages. For example, within the genus Acacia, a member of the Mimosoideae, A. pentagona does not form nodules, while other species of the same genus readily form nodules, as is the case for Acacia senegal, which forms both rapidly and slow growing rhizobial nodules.
Their ability to fix atmospheric nitrogen reduces fertilizer costs for farmers and gardeners who grow legumes, and means that legumes can be used in a crop rotation to replenish soil that has been depleted of nitrogen. Legume seeds and foliage have a comparatively higher protein content than non-legume materials, due to the additional nitrogen that legumes receive through the process. Legumes are commonly used as natural fertilizers. Some legume species perform hydraulic lift, which makes them ideal for intercropping. Preview available at Google Books.
Farmed legumes can belong to numerous classes, including Fodder, cereal, blooms, pharmaceutical/industrial, fallow/green manure and timber species, with most commercially farmed species filling two or more roles simultaneously.
There are of two broad types of forage legumes. Some, like alfalfa, clover, vetch, and Arachis, are sown in pasture and grazed by livestock. Other forage legumes such as Leucaena or Albizia are woody shrub or tree species that are either broken down by livestock or regularly cut by humans to provide fodder.
Grain legumes are cultivated for their , and are also called pulses. The seeds are used for human and animal consumption or for the production of oils for industrial uses. Grain legumes include both herbaceous plants like , , , and ,The gene bank and breeding of grain legumes (lupine, vetch, soya and beah) / B.S. Kurlovich and S.I. Repyev (Eds.), - St. Petersburg, The N.I. Vavilov Institute of Plant Industry, 1995, 438p. - (Theoretical basis of plant breeding. V.111) and trees such as carob, mesquite and tamarind.
Lathyrus tuberosus, once extensively cultivated in Europe, forms tubers used for human consumption.
Bloom legume species include species such as lupin, which are farmed commercially for their blooms, and thus are popular in gardens worldwide. Laburnum, Robinia, Gleditsia (honey locust), Acacia, Mimosa, and Delonix are Ornamental plant and .
Industrial farmed legumes include Indigofera, cultivated for the production of Indigo dye, Acacia, for gum arabic, and Derris, for the insecticide action of rotenone, a compound it produces.
Fallow or green manure legume species are cultivated to be tilled back into the soil to exploit the high nitrogen levels found in most legumes. Numerous legumes are farmed for this purpose, including Leucaena, Cyamopsis and Sesbania.
Various legume species are farmed for timber production worldwide, including numerous Acacia species, Dalbergia species, and Castanospermum australe.
Melliferous plants offer nectar to and other insects to encourage them to carry pollen from the of one plant to others thereby ensuring pollination. Many Fabaceae species are important sources of pollen and nectar for bees, including for honey production in the beekeeping industry. Example Fabaceae such as alfalfa, and various clovers including Trifolium repens and Melilotus, are important sources of nectar and honey for the Apis mellifera.
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