Lupinus


Lupinus, commonly known as lupin, lupine, or regionally bluebonnet, is a genus of plants in the legume family Fabaceae. The genus includes over 199 species, with centres of diversity in North and South America. Smaller centres occur in North Africa and the Mediterranean. They are widely cultivated, both as a food source and as ornamental plants, but are invasive to some areas. For instance, lupine has become an ecological problem in Iceland since the beginning of the 21st century.

Description

The species are mostly herbaceous perennial plants tall, but some are annual plants and a few are shrubs up to tall. An exception is the chamis de monte of Oaxaca in Mexico, which is a tree up to tall.
Lupins have soft green to grey-green leaves which may be coated in silvery hairs, often densely so. The leaf blades are usually palmately divided into five to 28 leaflets, or reduced to a single leaflet in a few species of the southeastern United States and eastern South America.
The flowers are produced in dense or open whorls on an erect spike, each flower long. The pea-like flowers have an upper standard, or banner, two lateral wings, and two lower petals fused into a keel. The flower shape has inspired common names such as bluebonnets and quaker bonnets.
The fruit is a pod containing several seeds. The seeds contain alkaloids which lend them a bitter taste.
Extrafloral nectaries have been found on a species in Arizona.

Taxonomy

The genus Lupinus L. and, in particular, its North American species were divided by Sereno Watson into three sections: Lupinus, Platycarpos, and Lupinnelus. Differences in habitat and in the number of ovules were the basis for this classification. A majority of the perennial and annual species from the American continent described by Watson were referred to Lupinus. Some annual species with two ovules in the ovary and two seeds in the pod were attributed to the Platycarpos section. Section Lupinnelus consisted of one species, with axillary and solitary flowers, scarcely reflexed banner, and also with two ovules in the ovary.
While Watson's work was predominantly based on study of North American species, the later research of Ascherson and Graebner extended his principle of classification to cover all lupins from the Eastern and Western Hemispheres, also using number of ovules in the ovary as the criterion for this division. They described two subgenera, Eulupinus and Platycarpos. Most of the described species were referred to subgen. A. Eulupinus. Subgen. B. Platycarpos included several annual species from the Eastern Hemisphere with two seedbuds and seeds in the bean.
A current schema retains this distinction, but uses the nomenclature for the subgenera of Platycarpos and Lupinus. In this schema, subgenus Platycarpos Kurl. contains perennial and annual species from the Western Hemisphere, with a minimum two or more ovules or seedbuds. Subgenus Lupinus consists of 12 species from Africa and the Mediterranean, with a minimum of four ovules or seedbuds.
The taxonomy of Lupinus has always been confusing. How many distinct species exist or how they might be organized within the genus is not clear. The plants are variable and the taxa are not always distinct from one another. Some American taxa have been described as complexes rather than separate species. Estimates of the number of lupine species generally fall between 200 and 500. One authority places the estimate at approximately 267 species worldwide. Currently, two subgenera are recognized.

Subgenus ''Platycarpos''

The ovary contains two and more ovules or seedbuds. The seed are predominantly small-sized, with an underdeveloped embryo and small amount of endosperm. Cotyledons are small-sized, with long caulicles. The first pair of true leaves is alternate. The stem is predominantly naked with waxen coating. Dominating is the monopodial type of branching. Leaflets are smooth, with waxen coating or slight pubescence, predominantly narrow. Pods are flat or orbicular, with two or more seeds. Represented by frutcuilose, fruticose and herbaceous perennial forms, or less often annual ones. Plants are cross-pollinated. Chromosome number 2n is either 36, 48, or 96. This subgenus is distributed throughout North, Central and South America, predominantly in the mining systems of the Andes and Cordillera. Some species are cultivated. This subgenus includes several hundred species, requiring further analysis of their authenticity.
It comprises the following species:Lupinus aberrans C.P. Sm.Lupinus abramsii C.P. Sm. – Abrams' lupineLupinus acopalcus C.P. Sm.Lupinus adinoanthus C.P. Sm.Lupinus adsurgens Drew – Drew's silky lupineLupinus affinis J. Agardh – fleshy lupineLupinus agardhianus A. HellerLupinus alaristatus C.P. Sm.Lupinus albert-smithianus C.P. Sm.Lupinus albescens Hook. & Arn. – hoary lupineLupinus albicaulis Douglas – sickle-keel lupineLupinus albifrons Benth. – silver bush lupine

Subgenus ''Lupinus''

In its current circumscription, subgenus Lupinus includes 12 species from the Mediterranean region and Africa with at least four ovules or seedbuds in the ovary:Lupinus albus L. 1753 – white lupine
  • * subsp. albus L.
  • * subsp. graecus Franco & P.Silva
  • * subsp. termis Ponert.Lupinus angustifolius L. 1753 – blue lupin, narrow-leafed lupin
  • * var. angustifolius L.
  • * var. albopunctatus Kurl. et Stankev.
  • * var. griseomaculatus Kurl. et Stankev.
  • * var. chalybens Kurl. et Stankev.
  • * var. corylinus Kurl. et Stankev.
  • * var. purpureus Kurl. et Stankev.
  • * var. rubidus Kurl. et Stankev.
  • * var. atabekovae Kurl. et Stankev.
  • * var. sparsiusculus Kurl. et Stankev.
  • * var. brunneus Kurl. et Stankev.
  • * var. albosyringeus Taran.
  • * var. albidus Kurl. et Stankev.
  • * var. candidus Kuptzov. et Kurl.Lupinus atlanticus Gladstones 1974Lupinus cosentinii Guss. 1828 – sandplain lupinLupinus digitatus Forsk. 1775Lupinus hispanicus Boiss. & Reut. 1842
  • * subsp. bicolor Gladst.
  • * subsp. hispanicus Boiss. & Reut.Lupinus luteus L. 1753 – yellow lupin
  • * var. luteus L.
  • * var. maculosus Kurl. et Stankev.
  • * var. kazimierskii Kurl. et Stankev.
  • * var. arcellus Kurl. et Stankev.
  • * var. sempolovskii Kurl. et Stankev.
  • * var. melanospermus Kurl. et Stankev.
  • * var. niger Kurl. et Stankev.
  • * var. cremeus Kurl. et Stankev.
  • * var. leucospermus Kurl. et Stankev.
  • * var. sulphureus Kurl. et Stankev.
  • * var. stepanovae Kurl. et Stankev.
  • * var. ochroleucus Kurl. et Stankev.
  • * var. aurantiacus Kurl. et Stankev.
  • * var. croceus Kurl. et Stankev.
  • * var. aureus Kurl. et Stankev.
  • * var. albicans Kurl. et Stankev.
  • * var. sinskayae Kurl. et Stankev.Lupinus micranthus Guss. 1828Lupinus palaestinus Boiss. 1849 – white-grey lupineLupinus pilosus Murr. 1774 – blue lupineLupinus princei Harms 1901Lupinus somaliensis Baker f. 1895

Species names with uncertain taxonomic status

The status of the following binomials is unresolved:Lupinus acaulis LarrañagaLupinus achilleaphilus C.P.Sm.Lupinus acutilobus A.HellerLupinus aegr-Aovium C.P.Sm.Lupinus africanus Lour.Lupinus agninus Gand.Lupinus agropyrophilus C.P.Sm.Lupinus alaimandus C.P.Sm.Lupinus albicaulis Douglas ex Hook.Lupinus alicanescens C.P.Sm.Lupinus aliclementinus C.P.Sm.Lupinus aliumbellatus C.P.Sm.Lupinus altissimus Sessé & Moc.Lupinus alturasensis C.P.Sm.Lupinus alveorum C.P.Sm.Lupinus amabilis A.HellerLupinus amniculi-cervi C.P.Sm.Lupinus amniculi-salicis C.P.Sm.Lupinus amniculi-vulpum C.P.Sm.Lupinus andersonianus C.P.Sm.Lupinus anemophilus GreeneLupinus angustifolius BlancoLupinus aphronorus Blank.Lupinus apodotropis A.HellerLupinus aralloius C.P.Sm.Lupinus arborescens Amabekova & MaisuranLupinus arceuthinus GreeneLupinus argyraeus DC.Lupinus atacamicus C.P.Sm.Lupinus aureus J.AgardhLupinus axillaris Blank.Lupinus barkeriae Knowles & Westc.Lupinus bartolomei M.E.JonesLupinus bassett-maguirei C.P.Sm.Lupinus beaneanus C.P.Sm.Lupinus biddleii L.F.Hend.Lupinus bimaculatus Hook. ex D.DonLupinus bimaculatus Desr.Lupinus bivonii C.PreslLupinus blankinshipii A.HellerLupinus blaschkeanus Fisch. & C.A.Mey.Lupinus brevior J.A. Christian & D.B. DunnLupinus brittonii AbramsLupinus caespitosus Nutt.Lupinus californicus K.KochLupinus campbelliae Eastw.Lupinus campestris Cham. & Schltdl.Lupinus campestris-florum C.P.Sm.Lupinus candicans Rydb.Lupinus canus Hemsl.Lupinus capitatus GreeneLupinus capitis-amniculi C.P.Sm.Lupinus carolus-bucarii C.P.Sm.Lupinus chachas Ochoa ex C. P. SmithLupinus chamissonis EschscholtzLupinus chiapensis RoseLupinus chihuahuensis S.WatsonLupinus christianus C.P.Sm.Lupinus chrysomelas Casar.Lupinus clementinus GreeneLupinus comatus Rydb.Lupinus consentinii Walp.Lupinus cymb-Aegressus C.P.Sm.Lupinus dasyphyllus GreeneLupinus davisianus C.P.Sm.Lupinus debilis Eastw.Lupinus decaschistus C.P.Sm.Lupinus diaboli-septem C.P.Sm.Lupinus dichrous GreeneLupinus dispersus A.HellerLupinus dissimulans C.P.Sm.Lupinus durangensis C.P.Sm.Lupinus eatonanus C.P.Sm.Lupinus equi-coeli C.P.Sm.Lupinus equi-collis C.P.Sm.Lupinus erectus L.F.Hend.Lupinus erminens S.WatsonLupinus ermineus S.WatsonLupinus falcifer Nutt.Lupinus falsoerectus C.P.Sm.Lupinus falsoformosus C.P.Sm.Lupinus falsograyi C.P.Sm.Lupinus fieldii Rose ex J. F. Macbr.Lupinus filicaulis C.P.Sm.Lupinus finitus C.P.Sm.Lupinus flavescens Rydb.Lupinus foliosus Hook.Lupinus foliosus Nutt.Lupinus forskahlei Boiss.Lupinus franciscanus GreeneLupinus fraxinetorum GreeneLupinus fruticosus Steud.Lupinus fruticosus Dum.Cours.Lupinus garcianus Bennett & DunnLupinus geophilus RoseLupinus geraniophilus C.P.Sm.Lupinus glabellus M.Martens & GaleottiLupinus graciliflorus C.P.Sm.Lupinus gratus GreeneLupinus gredensis Gand.Lupinus guadalupensis GreeneLupinus guadiloupensis Steud.Lupinus guatimalensis auct.Lupinus gussoneanus J.AgardhLupinus habrocomus GreeneLupinus haudcytisoides C.P.Sm.Lupinus helleri GreeneLupinus hexaedrus E. Fourn.Lupinus hintonii C.P.Sm.Lupinus huigrensis Rose ex C.P.Sm.Lupinus humicolus A.NelsonLupinus humifusus Benth.Lupinus humilis Rose ex PittierLupinus hyacinthinus GreeneLupinus idoneus C.P.Sm.Lupinus inamoenus Greene ex C.F.BakerLupinus indutus Greene ex C.F.BakerLupinus insignis Glaz. ex C. P. SmithLupinus integrifolius L.Lupinus intergrifolius Desr.Lupinus ione-grisetae C.P.Sm.Lupinus ione-walkerae C.P.Sm.Lupinus jamesonianus C.P.Sm.Lupinus javanicus Burm.f.Lupinus jorgensenanus C.P.Sm.Lupinus jucundus GreeneLupinus kellerrnanianus C.P.Sm.Lupinus kyleanus C.P.Sm.Lupinus labiatus Nutt.Lupinus lacticolor TamayoLupinus lacus-huntingtonii C.P.Sm.Lupinus lacuum-trinitatum C.P.Sm.Lupinus larsonanus C.P.Sm.Lupinus lassenensis Eastw.Lupinus latissimus GreeneLupinus laxifolius A.GrayLupinus leptostachyus GreeneLupinus lesueurii Standl.Lupinus linearifolius LarrañagaLupinus lingulae C.P.Sm.Lupinus longilabrum C.P.Sm.Lupinus lorentzianus C.P.Sm.Lupinus louise-bucariae C.P.Sm.Lupinus louise-grisetae C.P.Sm.Lupinus lucidus Benth. ex LoudonLupinus lyman-bensonii C.P.Sm.Lupinus lysichitophilus C.P.Sm.Lupinus macrocarpus Hook. & Arn.Lupinus macrocarpus Torr.Lupinus macrophyllus Benth.Lupinus macrorhizos GeorgiLupinus magnistipulatus Planchuelo & DunnLupinus maissurianii Atabek. & PolukhinaLupinus marcusianus C.P.Sm.Lupinus mariae-josephae H.PascualLupinus markleanus C.P.Sm.Lupinus marschallianus SweetLupinus mearnsii C.P.Sm.Lupinus meli-campestris C.P.Sm.Lupinus meridanus Moritz ex C. P. SmithLupinus mexiae C.P.Sm.Lupinus micensis M.E.JonesLupinus micheneri GreeneLupinus milleri J.AgardhLupinus minearanus C.P.Sm.Lupinus minutissimus TamayoLupinus molle A.HellerLupinus mollissifolius DavidsonLupinus monettianus C.P.Sm.Lupinus muellerianus C.P.Sm.Lupinus multicincinnis C.P.Sm.Lupinus neglectus RoseLupinus nemoralis GreeneLupinus niger WehmerLupinus noldekae Eastw.Lupinus nutcanus Spreng.Lupinus nutkatensis J.G.CooperLupinus obtunsus C.P.Sm.Lupinus octablomus C.P.Sm.Lupinus opsianthus Amabekova & MaisuranLupinus pavonum C.P.Sm.Lupinus pendeltonii A.HellerLupinus pendletonii A.HellerLupinus perconfertus C.P.Sm.Lupinus perplexus C.P.Sm.Lupinus philistaeus Boiss.Lupinus pinus-contortae C.P.Sm.Lupinus piperi B.L.Rob. ex PiperLupinus piperitus DavidsonLupinus platanophilus M.E.JonesLupinus plebeius Greene ex C.F.BakerLupinus prato-lacuum C.P.Sm.Lupinus prolifer Desr.Lupinus propinquus GreeneLupinus proteanus Eastw.Lupinus psoraleoides PollardLupinus pumviridis C.P.Sm.Lupinus puroviridis C.P.Sm.Lupinus purpurascens A.HellerLupinus pygmaeus TamayoLupinus quercus-jugi C.P.Sm.Lupinus quercuum C.P.Sm.Lupinus rainierensis Eastw.Lupinus regius Rudolph ex Torr. & A.GrayLupinus rhodanthus C.P.Sm.Lupinus rickeri C.P.Sm.Lupinus rivetianus C.P.Sm.Lupinus rydbergii Blank.Lupinus sabuli C.P.Sm.Lupinus salicisocius C.P.Sm.Lupinus salinensis C.P.Sm.Lupinus sativus GaterauLupinus scaposus Rydb.Lupinus scheuberae Rydb.Lupinus schickendantzii C.P.Sm.Lupinus schiedeanus Steud.Lupinus schumannii C.P.Sm.Lupinus seclusus C.P.Sm.Lupinus semiaequus C.P.Sm.Lupinus semiverticillatus Desr.Lupinus sergenti Tamayo ex PittierLupinus sergentii TamayoLupinus serradentum C.P.Sm.Lupinus shrevei C.P.Sm.Lupinus sierrae-zentae C.P.Sm.Lupinus sileri S.WatsonLupinus sinus-meyersii C.P. Sm.Lupinus sparhawkianus C.P.Sm.Lupinus spatulata LarrañagaLupinus speciosus VossLupinus spruceanus C.P.Sm.Lupinus standleyensis C.P.Sm.Lupinus stationis C.P.Sm.Lupinus stiveri KelloggLupinus stoloniferus L.Lupinus strigulosus Gand.Lupinus subhirsutus DavidsonLupinus subvolutus C.P.Sm.Lupinus suksdorfii B.L. Rob. ex PiperLupinus summersianus C.P.Sm.Lupinus sylvaticus Hemsl.Lupinus thermis Gasp.Lupinus thermus St.-Lag.Lupinus tilcaricus C.P.Sm.Lupinus timotensis TamayoLupinus tricolor GreeneLupinus tricolor G.NicholsonLupinus trifidus Torr. ex S.WatsonLupinus tristis SweetLupinus trochophyllus Hoffmanns.Lupinus tuckeranus C.P. Sm.Lupinus vaginans Benth.Lupinus valdepallidus C.P.Sm.Lupinus vandykeae Eastw.Lupinus variegatus A.HellerLupinus variegatus Poir.Lupinus varneranus C.P.Sm.Lupinus vavilovii Atabekova & MaissurjanLupinus venustus BaillyLupinus violaceus A.HellerLupinus viridicalyx C.P.Sm.Lupinus volcanicus GreeneLupinus watsonii A.HellerLupinus westiana SmallLupinus wolfianus C.P.Sm.Lupinus yanlyensis C.P.Sm.Lupinus yaruahensis C.P.Sm.

Hybrids

The following hybrids have been described:Lupinus ×alpestris D.B. Dunn & J.M. GillettLupinus ×hispanicoluteus W.Święcicki & W.K.ŚwięcickiLupinus ×hybridus Lem.Lupinus ×insignis Lem.Lupinus ×regalis Bergmans—rainbow lupin Lupinus ×versicolor Caball.

Etymology

While some sources believe the origin of the name to be in doubt, the Collins Dictionary definition asserts that the word is 14th century in origin, from the Latin lupīnus "wolfish" from lupus "wolf" as it was believed that the plant ravenously exhausted the soil.
But a more likely explanation is that lupinus meant that the plants were as dangerous to livestock as wolves, because the alkaloid poisons of lupines can sicken or kill grazing animals, especially sheep. Farmers have known since ancient Rome that lupines improve soil by adding nitrogen and loosening compacted earth with their strong root systems, so the Collins explanation is improbable.

Ecology

Certain species, such as the yellow bush lupin, are considered invasive weeds when they appear outside their native ranges. In New Zealand, lupines are viewed as invasive and a severe threat in some cases. L. polyphyllus has escaped into the wild and grows in large numbers along main roads and streams on the South Island. A similar spread of the species has occurred in Sweden, Finland and Norway after the non-native species was first deliberately planted in the landscaping along the main roads. Lupins have been planted in some parts of Australia with a considerably cooler climate, particularly in rural Victoria and New South Wales.
Lupins are important larval food plants for many lepidopterans. These include:Iraricia icarioides missionensis, larvae limited to LupinusCallophrys irus, recorded on L. perennisErynnis persius

Cultivation

Lupinus polyphyllus, the garden lupin, and Lupinus arboreus, the tree lupin, are popular ornamental plants in gardens, and are the source of numerous hybrids and cultivars in a wide range of colours, including bicolors. As legumes, lupins are good companion plants in gardens, increasing the soil nitrogen for vegetables and other plants. As well as growing in the ground, lupins can do well in pots on balconies or patios.

Agriculture

Like other legumes, lupines can fix nitrogen from the atmosphere into ammonia via a rhizobium–root nodule symbiosis, fertilizing the soil for other plants. This adaptation 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.
In the early 20th century, German scientists attempted to cultivate a sweet variety of lupin lacking the bitter taste, making it more suitable for both human and animal consumption.
Many annual species of lupins are used in agriculture and most of them have Mediterranean origin.
While originally cultivated as a green manure or forage, lupins are increasingly grown for their seeds, which can be used as an alternative to soybeans. Sweet lupins are highly regarded as a stock feed, particularly for ruminants, but also for pigs and poultry and more recently as an ingredient in aqua-feeds. Three Mediterranean species of lupin, blue (narrow-leafed) lupin, white lupin, and yellow lupin, are widely cultivated for livestock and poultry feed.
The market for lupin seeds for human food is currently small, but researchers believe it has great potential. Lupin seeds are considered "superior" to soybeans in certain applications and evidence is increasing for their potential health benefits. They contain similar protein to soybean, but less fat. As a food source, they are gluten-free and high in dietary fibre, amino acids, and antioxidants in food|antioxidants], and they are considered to be prebiotic.
About 85% of the world's lupin seeds are grown in Western Australia.

Toxicity

Some lupins contain certain secondary compounds, including isoflavones and toxic alkaloids, such as lupinine, anagyrine and sparteine. With early detection, these can be removed through processing, although lupins containing these elements are not usually selected for food-grade products.
A risk of lupin allergy exists in patients allergic to peanuts. Most lupin reactions reported have been in people with peanut allergy. Because of the cross-allergenicity of peanut and lupin, the European Commission, as of 2006, has required that food labels indicate the presence of "lupin and products thereof" in food.
Lupin plants can be colonized by the fungus Diaporthe toxica which can cause a mycotoxicosis known as lupinosis when ingested by grazing animals.

Uses

The legume seeds of lupins, commonly called lupin beans, were popular with the Romans, who cultivated the plants throughout the Roman Empire where the lupin is still known in extant Romance languages by names such as lupini.
Seeds of various species of lupins have been used as a food for over 3,000 years around the Mediterranean and for as long as 6,000 years in the Andes. Lupins were also used by many Native American peoples of North America such as the Yavapai. The Andean lupin or tarwi was a widespread food in the Incan Empire; but they have never been accorded the same status as soybeans, dry peas and other pulse crops. The pearl lupin of the Andean highlands of South America, L. mutabilis, known locally as tarwi or chocho, was extensively cultivated, but no conscious genetic improvement other than to select for larger and water-permeable seeds seems to have been made. Users soaked the seed in running water to remove most of the bitter alkaloids and then cooked or toasted the seeds to make them edible, or else boiled and dried them to make kirku, reported as a pre-Columbian practice in Relaciones geográficas. Spanish domination led to a change in the eating habits of the indigenous peoples, and only recently has interest in using lupins as a food been renewed.
Lupins can be used to make a variety of foods both sweet and savoury, including everyday meals, traditional fermented foods, baked foods, and sauces. The European white lupin beans are commonly sold in a salty solution in jars and can be eaten with or without the skin. Lupini dishes are most commonly found in Europe, especially in Portugal, Spain, Greece, and Italy. They are also common in Brazil and Egypt. In Egypt, the lupin is known in Arabic as ترمس termes, and is a popular street snack after being treated with several soakings of water, and then brined. In Portugal, Spain, and the Spanish Harlem district of New York, they are consumed with beer and wine. In the Levant and Palestine regions, the salty and chilled beans are called turmus and are served as part of an apéritif or a snack. Other species, such as L. albus, L. angustifolius, and L. hirsutus also have edible seeds.

Culture

Consumed throughout the Mediterranean region and the Andean mountains, lupins were eaten by the early Egyptian and pre-Incan people and were known to Roman agriculturalists for their ability to improve the fertility of soils.
In the late 18th century, lupins were introduced into northern Europe as a means of improving soil quality, and by the 1860s, the garden yellow lupin was seen across the sandy soils of the Baltic coastal plain.
The successful development of lupin varieties with the necessary "sweet gene" paved the way for the greater adoption of lupins across Europe and later Australia.
Further work carried out by the Western Australian Department of Agriculture and Food during the 1950s and '60s has led to more sweet lupin crops being produced in Western Australia now than anywhere else in the world.
Bluebonnets, including the Texas bluebonnet, are the state flowers of Texas.