Lupinus
Wild Perennial Lupin (Sundial lupine, Lupinus perennis).
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Eudicots
(unranked):	Rosids
Order:	Fabales
Family:	Fabaceae
Subfamily:	Faboideae
Tribe:	Genisteae
Subtribe:	Lupininae
Genus:	Lupinus L.
Species
See text

Lupinus, commonly known as Lupins or lupines (North America), is a genus in the legume family (Fabaceae). The genus comprises about 280 species (Hughes), with major centers of diversity in South and western North America (Subgen. Platycarpos (Wats.) Kurl.), parts of the Southern Hemisphere (New Zealand and parts of Australia) and the Andes and secondary centers in the Mediterranean region and Africa (Subgen. Lupinus).^[1][2]

Description [link]

The species are mostly herbaceous perennial plants 0.3–1.5 m (0.98–4.9 ft) tall, but some are annual plants and a few are shrubs up to 3 m (10 ft) tall—see also bush lupin^{[disambiguation needed ]}—with one species (Lupinus jaimehintoniana from the Mexican state of Oaxaca) a tree up to 8 m (26 ft) high with a trunk 20 cm (7.9 in) in diameter. They have a characteristic and easily recognised leaf shape, with soft green to grey-green leaves which in many species bear silvery hairs, often densely so. The leaf blades are usually palmately divided into 5–28 leaflets or reduced to a single leaflet in a few species of the southeastern United States. The flowers are produced in dense or open whorls on an erect spike, each flower 1–2 cm long, with a typical peaflower shape with an upper 'standard' or 'banner', two lateral 'wings' and two lower petals fused as a 'keel'. Due to the flower shape, several species are known as bluebonnets or quaker bonnets^{[disambiguation needed ]}. The fruit is a pod containing several seeds.

Usage [link]

Culinary [link]

The yellow legume seeds of lupins, commonly called lupin beans, were popular with the Romans, who spread the plant's cultivation throughout the Roman Empire; hence common names like lupini in Romance languages. The name 'Lupin' derives from the Latin word lupinus (meaning wolf), and was given with regard to the fact that many found that the plant has a tendency to ravage the land on which it grows. The peas, which appear after the flowering period, were also said to be fit only for the consumption of wolves. Lupin beans are commonly sold in a salty solution in jars (like olives and pickles) and can be eaten with or without the skin.

Lupini dishes are most commonly found in Mediterranean countries, especially in Portugal, Egypt, and Italy, and also in Brazil. In Portugal, Spain and the Spanish Harlem they are popularly consumed with beer. In Lebanon, salty and chilled Lupini Beans are called "Termos" and are served pre-meal as part of an aperitif. The Andean variety of this bean is from the Andean Lupin (tarwi, L. mutabilis) and was a widespread food in the Incan Empire. The Andean Lupin and the Mediterranean L. albus (White Lupin), L. angustifolius (Blue Lupin),^[3] and Lupinus hirsutus^[4] are also edible after soaking the seeds for some days in salted water.^[5] Lupins were also used by Native Americans in North America, e.g., the Yavapai people. Lupins are known as altramuz in Spain and Argentina. In Portuguese the lupin beans are known as tremoços, and in Antalya (Turkey) as tirmis.^{[verification needed]} Edible lupins are referred to as sweet lupins because they contain smaller amounts of toxic alkaloids than the bitter lupin varieties. Newly bred variants of sweet lupins are grown extensively in Germany; they lack any bitter taste and require no soaking in salt solution. The seeds are used for different foods from vegan sausages to lupin-tofu or baking-enhancing lupin flour.

Agriculture [link]

Given that lupin seeds have the full range of essential amino acids and that they, contrary to soy, can be grown in more temperate to cool climates, lupins are becoming increasingly recognized as a cash crop alternative to soy.

Three Mediterranean species of lupin, Blue Lupin, White Lupin, and Yellow Lupin (L. luteus) are widely cultivated for livestock and poultry feed.

Lupins are also cultivated as forage and grain legumes. Like most members of their family, lupins can fix nitrogen from the atmosphere into ammonia via a rhizobium-root nodule symbiosis, fertilizing the soil for other plants, this adaption allows lupins to be tolerant of infertile soils and capable of pioneering change in barren and poor quality soils. The genus Lupinus is nodulated by Bradyrhizobium soil bacteria.^[6] Some species have a long central tap roots, or have proteoid roots.

Companion plant [link]

Ornamental lupins, Ushuaia

Lupins make good companion plants for crops that need significant amounts of nitrogen in their soil and can be intercropped properly, like cucumbers, squash, broccoli, and spinach.

Horticulture [link]

Lupin population, Lake Tekapo, New Zealand

Lupins are popular ornamental plants in gardens. There are numerous hybrids and cultivars. Some species, such as Garden Lupin (Lupinus polyphyllus) and hybrids like the Rainbow Lupin (L. × regalis) are common garden flowers. Others, such as the Yellow Bush Lupin (L. arboreus) are considered invasive weeds when they appear outside their native range.

In New Zealand Lupinus polyphyllus have escaped into the wild and grow in large numbers along main roads and streams on the South Island. Although considered attractive by some it is also seen as an invasive species. A similar spread of the species has occurred in Finland after the non-native species was first deliberately planted alongside main roads as part of the road landscaping.

Ecology [link]

Canadian Tiger Swallowtail on wild lupine, Gatineau, Quebec

For several Lepidoptera (butterflies and moths), lupins are an important larval food. These include:

• Callophrys irus (Frosted Elfin)^[7][8]
• Chesias legatella (The Streak)^[9]
• Chionodes braunella
• Glaucopsyche xerces (Xerces Blue) – extinct
• Icaricia icarioides missionensis (Mission Blue)^[8][10]
• Lycaeides melissa samuelis (Karner Blue)^[7][8]
• Melanchra persicariae (Dot Moth)
• Phymatopus behrensii
• Schinia suetus^[11]

The endangered Apodemia mormo langei (Lange's Metalmark) mates on Silver Bush Lupin (L. albifrons).

The most significant diseases of lupins are anthracnose as well as wilting and root rot diseases caused by Fusarium and other pathogens, and some bacterial and viral diseases.^[12]

Potential harms [link]

Lupins contain significant amounts of certain secondary compounds like isoflavones and toxic alkaloids, e.g. lupinine and sparteine. On 22 December 2006, the European Commission submitted directive 2006/142/EC, which amends the EU foodstuff allergen list to include "lupin and products thereof".

Both sweet and bitter lupins in feed can cause livestock poisoning. Lupin poisoning is a nervous syndrome caused by alkaloids in bitter lupins, similar to neurolathyrism. Mycotoxic lupinosis is a disease caused by lupin material that is infected with the fungus Diaporthe toxica;^[13] the fungus produces mycotoxins called phomopsins, which cause liver damage. Poisonous lupin seeds cause annually the loss of many cattle and sheep on western American Ranges.^[14]

People with peanut allergy should generally avoid lupins. In one study^[15] 44% of people with peanut allergy had a positive allergy test for lupin allergy and 7 of 8 who had a positive test and were fed lupin as part of a study reacted to this food.

Taxonomy [link]

Lupin and other wildflowers cover the mountaintop of Raspberry Island (Alaska)

Lupinus flavoculatus

Lupinus texensis, state flower of Texas

Overall the taxonomy of this genus has been traditionally confusing. The last major monograph, by Agardh in 1835 recognised 83 species, yet about 1800 names are in use.(Hughes) Some of the most recent phylogenetics indicates the presence of 13 Old World species, where the genus is thought to have originated, and two groups of New World species. The smaller of these two clades comprises c. 35 species distributed largely in the Eastern lowlands, and the larger of c. 222 in Western highland regions.(Hughes)^[16]

History [link]

Watson (1873) originally divided the genus Lupinus into three sections, Platycarpos, Lupinus and Lupinellus based on habitat and the number of ovules. Most of the species found in the Americas were assigned to Lupinus. Platycarpos consisted of some annuals with two ovules and two seeds (e.g., L. densiflorus, L. micricarpus), while Lupinellus had only one species (L. uncialis).

While Watson's work was predominantly based on study of N American species, the later research of Ascherson and Graebner (1907) was more global. They described two subgenera, Eulupinus and Platycarpos using similar criteria. Most species fell into the subgenus Eulupinus, while Platycarpos included the annual species from the Eastern Hemisphere in Watson's classification.

Current schema (Kurlovich and Stankevich 2002) retain this distinction but use the nomenclature for the subgenera of Platycarpos and Lupinus. In this schema Subgen. Platycarpos (Wats.) Kurl. contains perennial and annual species from the Western hemisphere, two or more ovules. Subgen. Lupinus consists of eleven species from Africa and the mediterranean, with a minimum of ovules or seedbuds.

[edit] Subgenus Platycarpos (circa 270 species)

(Wats.) Kurl., comb.nova. - §2. Platycarpos Wats. 1873, Proc. Amer. Acad. Arts Sci. 8:522; B. Platycarpos Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:232. - §1. Lupinus Wats. 1873, Proc. Amer. Acad. Arts Sci. 8:522, p.p.; A. Eulupinus Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221 p.p. (New World’s or flat-fruited lupins)^[1]

Selected species

• Lupinus adsurgens – Drew's silky lupine
• Lupinus affinis – fleshy lupine
• Lupinus albicaulis – sickle-keel lupine
• Lupinus albifrons – silver bush lupine
• Lupinus × alpestris
• Lupinus andersonii – Anderson's lupine
• Lupinus angustiflorus – narrowflower lupine
• Lupinus antoninus – Anthony Peak lupine
• Lupinus arboreus – yellow bush lupin, tree lupine
• Lupinus arbustus – longspur lupine
• Lupinus arcticus – Arctic lupine
• Lupinus argenteus – silvery lupine
  • Lupinus argenteus var. palmeri
• Lupinus aridorum – scrub lupine
• Lupinus arizonicus – Arizona lupine
• Lupinus benthamii
• Lupinus bicolor – miniature lupine, bicolor lupine, Lindley's annual lupine
• Lupinus bingenensis – Bingen lupine
• Lupinus brevicaulis – shortstem lupine
• Lupinus breweri – Brewer's lupine
• Lupinus burkei – Burke's lupine
• Lupinus caespitosus – stemless dwarf lupine
• Lupinus caudatus – Kellogg's spurred lupine
• Lupinus cervinus Kellogg – Santa Lucia lupine (= L. latissimus)
• Lupinus chamissonis – Chamisso bush lupine
• Lupinus citrinus – orange lupine
• Lupinus concinnus
• Lupinus constancei – The Lassics lupine
• Lupinus covillei – shaggy lupine
• Lupinus croceus – saffron-flowered lupine
• Lupinus dalesiae – Quincy lupine
• Lupinus densiflorus – (Subgenus type species)
• Lupinus duranii – Mono Lake lupine
• Lupinus diffusus – spreading lupine, Oak Ridge lupine, sky-blue lupine
• Lupinus elatus – tall silky lupine
• Lupinus elegans – elegant lupine
• Lupinus elmeri – Elmer's lupine
• Lupinus excubitus – grape soda lupine
• Lupinus flavoculatus
• Lupinus foliolosus
• Lupinus formosus – summer lupine
• Lupinus grayi – Sierra lupine
• Lupinus guadalupensis – Guadalupe Island lupine
• Lupinus havardii
• Lupinus hirsutus
• Lupinus hirsutissimus – stinging lupine
• Lupinus holmgrenianus – Holmgren's lupine
• Lupinus hyacinthinus – San Jacinto lupine
• Lupinus incanus – Hoary lupine
• Lupinus jaimehintoniana
• Lupinus kuntii
• Lupinus kuschei – Yukon lupin
• Lupinus lapidicola ; Mt. Eddy lupine
• Lupinus latifolius – broadleaf lupine
  • Lupinus latifolius var. barbatus – Klamath lupine, bearded lupine
• Lupinus lepidus – prairie lupine
• Lupinus leucophyllus – woolly-leaf lupine
• Lupinus littoralis – seashore lupine
• Lupinus longifolius – longleaf bush lupine
• Lupinus luteolus – butter lupine, pale yellow lupine
• Lupinus lyallii – Lyall's lupine
• Lupinus macbrideanus
• Lupinus michelianus
• Lupinus microcarpus – wide-bannered lupin, chick lupin
  • Lupinus microcarpus var. densiflorus – dense-flowered lupin
• Lupinus minimus – Kettle Falls lupin
• Lupinus mutabilis – Andean lupin, pearl lupin, South American lupin, tarwi, tarhui, chocho
• Lupinus nanus – dwarf lupin, field lupin, sky lupin, Douglas' annual lupin
• Lupinus nevadensis – Nevada lupine
• Lupinus nipomensis – Nipomo Mesa lupine
• Lupinus niveus
• Lupinus nootkatensis – Nootka lupin
• Lupinus nubigenus
• Lupinus obtusilobus – bluntlobe lupine
• Lupinus odoratus – royal Mojave lupin
• Lupinus onustus – Plumas lupine
• Lupinus oreganus – Oregon lupin
• Lupinus padre-crowleyi – DeDecker's lupine, Father Crowley's lupine
• Lupinus parviflorus – lodgepole lupin
• Lupinus peirsonii – Peirson's lupine, long lupine
• Lupinus perennis – wild perennial lupin, sundial lupin, Indian beet, old maid's bonnets
• Lupinus plattensis
• Lupinus polycarpus – smallflower lupin
• Lupinus polyphyllus – largeleaf lupin, bigleaf lupin, garden lupin
• Lupinus pratensis – Inyo Meadow lupine
• Lupinus prunophilus – hairy bigleaf lupin
• Lupinus pusillus – small lupin
• Lupinus × regalis – rainbow lupin
• Lupinus rivularis – riverbank lupin
• Lupinus rupestris
• Lupinus saxosus – rock lupine
• Lupinus sericatus – Cobb Mountain lupine
• Lupinus sericeus – Pursh's silky lupin
• Lupinus shockleyi – purple desert lupine
• Lupinus smithianus
• Lupinus sparsiflorus – desert lupin, Coulter's lupin, Mojave lupin
• Lupinus spectabilis – shaggyhair lupine
• Lupinus stiversii – harlequin annual lupine
• Lupinus subcarnosus – buffalo clover
• Lupinus succulentus – succulent lupin, arroyo lupin, hollowleaf annual lupin
• Lupinus sulphureus – sulphur lupin, sulphur-flowered lupin
  • Lupinus sulphureus ssp. kincaidii – Kincaid's lupin (formerly in L. oreganus)
• Lupinus texensis – Texas bluebonnet
• Lupinus tidestromii – Tidestrøm's Lupin
• Lupinus toratensis – Warwanzo, Lito
• Lupinus tracyi – Tracy's lupine
• Lupinus truncatus – collared annual lupine
• Lupinus vallicola – open lupin
• Lupinus variicolor – varied lupin
• Lupinus villosus
• Lupinus wyethii – Wyeth's lupin

Subgenus Lupinus (11 species) [link]

Blue Lupin (L. angustifolius)

Lupinus pilosus, Judean Foothills, Israel

A. Eulupinus Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221, p.p. (Old World Lupins)^[1]

• Lupinus albus L. 1753 – white lupine (Subgenus type species)
• Lupinus angustifolius L. 1753 – blue lupin, narrowleaf lupine
• Lupinus atlanticus Gladstones, 1974
• Lupinus cosentinii Guss. 1828
• Lupinus digitatus Forsk. 1775
• Lupinus hispanicus Boiss. et Reut. 1842
• Lupinus luteus L. 1753 – yellow lupine
• Lupinus micranthus Guss. 1828
• Lupinus palaestinus Boiss. 1849 –White-grey lupine, indigenous to Israel
• Lupinus pilosus Murr. 1774 – Blue Mountain lupine, indigenous to Israel
• Lupinus princei Harms, 1901

Symbolic uses [link]

Bluebonnet lupins, notably the Texas Bluebonnet (Lupinus texensis) are the state flower of Texas, USA.

See also [link]

• Alice Eastwood
• Lupin bean
• Lupin milk
• Lupin poisoning

Footnotes [link]

References [link]

• Ainouche, Abdel-Kader & Bayer, Randall J. (1999): Phylogenetic relationships in Lupinus (Fabaceae: Papilionoideae) based on internal transcribed spacer sequences (ITS) of nuclear ribosomal DNA. Am. J. Bot. 86(4): 590-607. PDF fulltext
• Eastwood, R.J., Drummond, C.S., Schifino‐Wittmann, M.T. & Hughes, C.E. 2008. Diversity and evolutionary history of lupins – insights from new phylogenies. Pp. 346‐354, in: Palta, J.A. & Burger, J.B. (Eds.) Lupins for Health & Wealth, Proceedings 12th International Lupin Conference, Fremantle, Australia, International Lupin Association, Canterbury, New Zealand.
• Golubev, A.A. & Kurlovich, Boguslav S.: Diseases and Pests. In: Kurlovich 2002: 287-312.
• Hedrick, U.P. (ed.) (1919): Sturtevant's Edible Plants of the World
• Hughes, Colin, Dept Systematic Botany, University of Zurich
  • Lupinus
• Kurlovich, Boguslav S. (ed.) Lupins: Geography, classification, genetic resources and breeding. Department of Leguminous Crops of N.I. Vavilov Institute of Plant Industry, 2002, ISBN 5-86741-034-X ; 468 pages.
• Kurlovich BS and Stankevich AK. Description of the New World's lupins (Subgen. Platycarpos (Wats). Kurl.), in Kurlovich BS and Stankevich AK: Classification of Lupins, in Kulovich BS (ed.) 2002 pp 44-61
• Kurlovich BS and Stankevich AK. Description of the Old World's lupins and their intraspecific diversity (Subgen. Lupinus), in Kurlovich BS and Stankevich AK: Classification of Lupins, in Kulovich BS (ed.) 2002 pp 62-87
• Kurlovich, Boguslav S.; Tikhonovich, I.A.; Kartuzova, L.T.; Heinänen, J.; Kozhemykov, A.P.; Tchetkova, S.A.; Cheremisov B.M. & Emeljanenko, T.A.: Nitrogen fixation. In: Kurlovich 2002: 269-286.
• Murcia, José & Hoyos, Isabel (1998): Características y aplicaciones de las plantas: ALTRAMUZ AZUL (Lupinus angustifolius) [in Spanish]. Retrieved 2007-10-09.
• Williamson, P.M.; Highet, A.S.; Gams, W.; Sivasithamparam, K. & Cowling, W.A. (1994): Diaporthe toxica sp. nov., the cause of lupinosis in sheep. Mycological Research 98(12): 1364-1365. HTML abstract ADRIS record
• University of Melbourne: Sorting Lupinus Names

External links [link]

Wikimedia Commons has media related to: Lupinus

Wikispecies has information related to: Lupinus

• Classification of lupins
• Lupins resource website
• Alternative field crops manual: Lupine
• Seed proteome reference maps at Lupin Protein Team