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 (Lupinus jaimehintonianus) 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

thumb|right|[[Lupinus aschenbornii, Nevado de Toluca, Mexico]]

The genus Lupinus L. and, in particular, its North American species were divided by Sereno Watson (1873) 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 (L. densiflorus, L. microcarpus, etc.) were attributed to the Platycarpos section. Section Lupinnelus consisted of one species (L. uncialis), 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 (1907) extended his principle of classification to cover all lupins from the Eastern and Western Hemispheres, also using number of ovules (seedbuds) in the ovary (and thus of seeds in the pod) 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 (the same species, as the one specified by S. Watson).

A current schema retains this distinction, but uses the nomenclature for the subgenera of Platycarpos and Lupinus. In this schema, subgenus Platycarpos (S.Wats.) 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.

Lupinus aberrans C.P. Sm.

Lupinus abramsii C.P. Sm. – Abrams' lupine
Lupinus acopalcus C.P. Sm.
Lupinus adinoanthus C.P. Sm.

Lupinus adsurgens Drew – Drew's silky lupine

Lupinus affinis J. Agardh – fleshy lupine
Lupinus agardhianus A. Heller
Lupinus alaristatus C.P. Sm.
Lupinus albert-smithianus C.P. Sm.

Lupinus albescens Hook. & Arn. – hoary lupine
Lupinus albicaulis Douglas – sickle-keel lupine
Lupinus albifrons Benth. – silver bush lupine
var. albifrons Benth.
var. douglasii (J. Agardh) C. P. Sm.
var. hallii (Abrams) Isely
Lupinus albopilosus A. Heller
Lupinus albosericeus C.P. Sm.
Lupinus alcis-montis C.P. Sm.

Lupinus aliamandus C.P. Sm.
Lupinus aliattenuatus C.P. Sm.
Lupinus alibicolor C.P. Sm.
Lupinus aliceae C.P. Sm.
Lupinus alilatissimus C.P. Sm.

Lupinus alinanus C.P. Sm.
Lupinus alipatulus C.P. Sm.
Lupinus alirevolutus C.P. Sm.
Lupinus alivillosus C.P. Sm.
Lupinus allargyreius C.P. Sm.

Lupinus alopecuroides Desr.
Lupinus alpestris A. Nelson

Lupinus altimontanus C.P. Sm.
Lupinus altiplani C.P. Sm.
Lupinus amabayensis C.P. Sm.
Lupinus amandus C.P. Sm.

Lupinus amboensis C.P. Sm.
Lupinus ammophilus Greene
var. ammophilus Greene
var. crassus (Payson) Isely

Lupinus amnis-otuni C.P. Sm.
Lupinus ampaiensis C.P. Sm.

Lupinus amphibius Suksd.
Lupinus ananeanus Ulbr.
Lupinus anatolicus W. Święcicki & W. K. Święcicki
Lupinus andersonii S. Watson – Anderson's lupine
Lupinus andicola Gillies
Lupinus andinus Rose ex J. F. Macbr.
Lupinus angustiflorus Eastw. – narrowflower lupine
Lupinus antensis C.P. Sm.
Lupinus antiplani C. P. Sm.
Lupinus antoninus Eastw. – Anthony Peak lupine
Lupinus apertus A. Heller
Lupinus appositus C.P. Sm.

Lupinus arboreus Sims – yellow bush lupin, tree lupine
Lupinus arbustus Lindl. – longspur lupine
subsp. arbustus Lindl.
subsp. neolaxiflorus D.B.Dunn
subsp. pseudoparviflorus (Rydb.) D.B.Dunn
Lupinus arbutosocius C.P. Sm.
Lupinus archeranus C.P. Sm.
Lupinus arcticus S. Watson – Arctic lupine
subsp. arcticus S. Watson
subsp. subalpinus (Piper & Robinson)D.B.Dunn
Lupinus arenarius Gardner

Lupinus arequipensis C.P. Sm.
Lupinus argenteus Pursh – silvery lupine
var. argentatus (Rydb.) Barneby
var. argenteus Pursh
var. argophyllus (A. Gray) S. Watson
var. depressus (Rydb.) C. L. Hitchc.
var. fulvomaculatus (Payson) Barneby
var. heteranthus (S. Watson) Barneby – Kellogg's spurred lupine
var. hillii (Greene) Barneby
var. holosericeus (Torr. & A.Gray) Barneby
var. montigenus (A. Heller) Barneby
var. palmeri (S.Watson) Barneby
var. rubricaulis (Greene) S. L. Welsh
var. utahensis (S.Watson) Barneby

Lupinus argurocalyx C.P. Sm.
Lupinus aridorum McFarlin ex Beckner – scrub lupine
Lupinus aridulus C.P. Sm.
Lupinus aridus Lindl.
Lupinus ariste-josephii C.P. Sm.
Lupinus arizelus C.P. Sm.
Lupinus arizonicus (S. Watson) S. Watson
subsp. arizonicus (S. Watson) S. Watson – Arizona lupine
subsp. sonorensis Christian & D. Dunn – Sonora lupine
Lupinus arvensi-plasketti C.P. Sm.
Lupinus arvensis Benth.
Lupinus asa-grayanus C.P. Sm.
Lupinus aschenbornii S. Schauer
Lupinus asplundianus C.P. Sm.
Lupinus asymbepus C.P. Sm.
Lupinus atropurpureus C.P. Sm.
Lupinus attenuatus Gardner

Lupinus aureonitens Hook. & Arn.
Lupinus austrobicolor C.P. Sm.
Lupinus austrohumifusus C.P. Sm.

Lupinus austrorientalis C.P. Sm.
Lupinus austrosericeus C.P. Sm.

Lupinus ballianus C.P. Sm.
Lupinus bandelierae C.P. Sm.
Lupinus bangii Rusby
Lupinus barbatilabius C.P. Sm.

Lupinus barkeri Lindl.
Lupinus bartlettianus C.P. Sm.
Lupinus benthamii A. Heller
Lupinus bi-inclinatus C.P. Sm.
Lupinus bicolor Lindl. – miniature lupine, bicolor lupine, Lindley's annual lupine
subsp. bicolor Lindl.
subsp. microphyllus (S. Watson) D. B. Dunn
subsp. pipersmithii (A. Heller) D. B. Dunn
subsp. umbellatus (Greene) D. B. Dunn

Lupinus bingenensis Suksd. – Bingen lupine
Lupinus blaisdellii Eastw.

Lupinus bogotensis Benth.
Lupinus bolivianus C.P. Sm.
Lupinus bombycinocarpus C.P. Sm.
Lupinus bonplandius C.P. Sm.

Lupinus boyacensis C.P. Sm.

Lupinus brachypremnon C.P. Sm.
Lupinus bracteolaris Desr.
Lupinus brandegeei Eastw.
Lupinus brevecuneus C.P. Sm.
Lupinus brevicaulis S. Watson – shortstem lupine
Lupinus brevior (Jeps.) Christian & D.B. Dunn
Lupinus breviscapus Ulbr.
Lupinus breweri A. Gray – Brewer's lupine

Lupinus bryoides C.P. Sm.
Lupinus buchtienii Rusby
Lupinus burkartianus C.P. Sm.
Lupinus burkei S. Watson – Burke's lupine
Lupinus burkeri Lindl.
Lupinus caballoanus B.L. Turner
Lupinus cachupatensis C.P. Sm.
Lupinus cacuminis Standl.
Lupinus caeruleus A. Heller
Lupinus caesius Eastw.
Lupinus caespitosus Torr. & A. Gray – stemless dwarf lupine
Lupinus calcensis C.P. Sm.
Lupinus caldasensis C.P. Sm.

Lupinus camiloanus C.P. Sm.
Lupinus campestris Schltdl. & Cham.

Lupinus carazensis Ulbr.
Lupinus carchiensis C.P. Sm.
Lupinus cardenasianus C.P. Sm.
Lupinus carhuamayus C.P. Sm.
Lupinus carlos-ochoae C.P. Sm.
Lupinus carpapaticus C.P. Sm.
Lupinus carrikeri C.P. Sm.
Lupinus caucensis C.P. Sm.

Lupinus cavicaulis C.P. Sm.
Lupinus ccorilazensis Vargas ex C. P. Smith
Lupinus celsimontanus C.P. Sm.
Lupinus cervinus Kellogg – Santa Lucia lupine
Lupinus cesar-vargasii C.P. Sm.
Lupinus cesaranus C.P. Sm.
Lupinus chachas C.P. Sm.
Lupinus chamissonis Eschsch. – Chamisso bush lupine
Lupinus chavanillensis (J.F. Macbr.) C.P. Sm.

Lupinus chipaquensis C.P. Sm.
Lupinus chlorolepis C.P. Sm.
Lupinus chocontensis C.P. Sm.
Lupinus chongos-bajous C.P. Sm.
Lupinus christinae A. Heller
Lupinus chrysanthus Ulbr.
Lupinus chrysocalyx C.P. Sm.
Lupinus chumbivilcensis C.P. Sm.
Lupinus citrinus Kellogg – orange lupine
Lupinus clarkei Oerst.

Lupinus cochapatensis C.P. Sm.

Lupinus colcabambensis C.P. Sm.
Lupinus collinus (Greene) A. Heller
Lupinus colombiensis C.P. Sm.

Lupinus compactiflorus Rose
Lupinus comptus Benth.
Lupinus concinnus J. Agardh
subsp. concinnus J. Agardh
subsp. orcuttii (S.Watson) D.B.Dunn
Lupinus condensiflorus C.P. Sm.
Lupinus confertus Kellogg

Lupinus congdonii (C.P. Sm.) D.B. Dunn
Lupinus conicus C.P. Sm.
Lupinus constancei T.W. Nelson & J.P. Nelson – Lassics lupine
Lupinus convencionensis C.P. Sm.
Lupinus cookianus C.P. Sm.
Lupinus coriaceus Benth.

Lupinus costaricensis D.B. Dunn
Lupinus cotopaxiensis C.P. Sm.
Lupinus couthouyanus C.P. Sm.
Lupinus covillei Greene – shaggy lupine
Lupinus crassulus Greene
Lupinus crassus Payson
Lupinus croceus Eastw. – saffron-flowered lupine
Lupinus crotalarioides Benth.
Lupinus crucis-viridis C.P. Sm.

Lupinus cuatrecasasii C.P. Sm.
Lupinus culbertsonii Greene
subsp. culbertsonii Greene
subsp. hypolasius (Greene) B.J.Cox

Lupinus cumulicola Small
Lupinus cusickii S. Watson
subsp. abortivus (Greene) B.J.Cox
subsp. brachypodus (Piper) B.J.Cox
subsp. cusickii S. Watson
Lupinus cuspidatus Rusby

Lupinus cuzcensis C.P. Sm.

Lupinus cymboides C.P. Sm.

Lupinus czermakii Briq. & Hochr.

Lupinus dalesiae Eastw. – Quincy lupine
Lupinus decemplex C.P. Sm.
Lupinus decurrens Gardner

Lupinus deflexus Congdon
Lupinus delicatulus Sprague & Riley
Lupinus densiflorus Benth. – dense-flowered lupin
subsp. densiflorus Benth.
subsp. lacteus (Kellogg) R.M.Beauch.
Lupinus depressus Rydb.
Lupinus diasemus C.P. Sm.
Lupinus diehlii M.E. Jones
Lupinus diffusus Nutt. – spreading lupine, Oak Ridge lupine, sky-blue lupine
Lupinus disjunctus C.P. Sm.

Lupinus diversalpicola C.P. Sm.

Lupinus dorae C.P. Sm.
Lupinus dotatus C.P. Sm.

Lupinus duranii Eastw. – Mono Lake lupine
Lupinus dusenianus C.P. Sm.
Lupinus eanophyllus C.P. Sm.

Lupinus edysomatus C.P. Sm.
Lupinus egens C.P. Sm.

Lupinus elaphoglossum Barneby
Lupinus elatus I.M. Johnst. – tall silky lupine
Lupinus elegans Kunth – elegant lupine
Lupinus elegantulus Eastw.
Lupinus ellsworthianus C.P. Sm.
Lupinus elmeri Greene – Elmer's lupine

Lupinus eramosus C.P. Sm.
Lupinus erectifolius C.P. Sm.

Lupinus eremonomus C.P. Sm.
Lupinus eriocalyx (C.P. Sm.) C.P. Sm.
Lupinus eriocladus Ulbr.
Lupinus evermannii Rydb.
Lupinus espinarensis C.P. Sm.
Lupinus exaltatus Zucc.
Lupinus excubitus M.E. Jones – grape soda lupine
subsp. austromontanus (A.Heller) R.M.Beauch.
subsp. excubitus M.E. Jones
Lupinus exochus C.P. Sm.
Lupinus expetendus C.P. Sm.
Lupinus extrarius C.P. Sm.
Lupinus falsomutabilis C.P. Sm.
Lupinus falsoprostratus C.P. Sm.
Lupinus falsorevolutus C.P. Sm.
Lupinus famelicus C.P. Sm.

Lupinus fiebrigianus Ulbr.
Lupinus fieldii J.F. Macbr.

Lupinus fissicalyx A. Heller

Lupinus flavoculatus A. Heller

Lupinus foliolosus Benth.

Lupinus formosus Greene – summer lupine
var. bridgesii (S.Watson) Greene
var. formosus Greene

Lupinus fragrans A. Heller
Lupinus francis-whittieri C.P. Sm.
Lupinus fratrum C.P. Sm.
Lupinus fulcratus Greene

Lupinus gachetensis C.P. Sm.

Lupinus garfieldensis C.P. Sm.
Lupinus gaudichaudianus C.P. Sm.
Lupinus gayanus C.P. Sm.

Lupinus gentryanus C.P. Sm.
Lupinus geophilus Rose
Lupinus gibertianus C.P. Sm.
Lupinus giganteus Rose
Lupinus glabratus J. Agardh

Lupinus goodspeedii J.F. Macbr.
Lupinus gormanii Piper
Lupinus gracilentus Greene

Lupinus grayi S. Watson – Sierra lupine
Lupinus grauensis C.P. Sm.

Lupinus grisebachianus C.P. Sm.
Lupinus guadalupensis C.P. Sm. – Guadalupe Island lupine
Lupinus guaraniticus (Hassl.) C.P. Sm.
Lupinus guascensis C.P. Sm.
Lupinus guggenheimianus Rusby

Lupinus hamaticalyx C.P. Sm.
Lupinus hartmannii C.P. Sm.
Lupinus hartwegii Lindl.
Lupinus haughtianus C.P. Sm.
Lupinus hautcarazensis C.P. Sm.
Lupinus havardii S. Watson
Lupinus hazenanus C.P. Sm.
Lupinus hendersonii Eastw.

Lupinus heptaphyllus (Vell.) Hassl.
Lupinus herreranus C.P. Sm.
Lupinus herzogii Ulbr.
Lupinus hieronymii C.P. Sm.
Lupinus hilarianus Benth.
Lupinus hillii Greene
Lupinus hinkleyorum C.P. Sm.
Lupinus hintoniorum B.L. Turner
Lupinus hirsutissimus Benth. – stinging lupine

Lupinus holmgrenianus C.P. Sm. – Holmgren's lupine

Lupinus honoratus C.P. Sm.
Lupinus horizontalis A. Heller
Lupinus hornemanni J. Agardh
Lupinus hortonianus C.P. Sm.
Lupinus hortorum C.P. Sm.
Lupinus howard-scottii C.P. Sm.
Lupinus howardii M.E. Jones
Lupinus huachucanus M.E. Jones
Lupinus huancayoensis C.P. Sm.
Lupinus huariacus C.P. Sm.
Lupinus huaronensis J.F. Macbr.
Lupinus huigrensis Rose ex C. P. Sm.

Lupinus humifusus Sessé & Moc. ex G. Don
Lupinus hyacinthinus C.F. Baker – San Jacinto lupine
Lupinus hybridus Lem.

Lupinus ignobilis C.P. Sm.
Lupinus imminutus C.P. Sm.

Lupinus indigoticus Eastw.
Lupinus inflatus C.P. Sm.

Lupinus insignis C.P. Sm.
Lupinus insulae C.P. Sm.

Lupinus interruptus Benth.
Lupinus intortus C.P. Sm.
Lupinus inusitatus C.P. Sm.
Lupinus involutus C.P. Sm.
Lupinus inyoensis A. Heller
Lupinus isabelianus Eastw.

Lupinus jahnii Rose ex Pittier
Lupinus jaimehintoniana B.L. Turner
Lupinus james-westii C.P. Sm.
Lupinus jean-julesii C.P. Sm.
Lupinus jelskianus C.P. Sm.
Lupinus johannis-howellii C.P. Sm.
Lupinus jonesii Rydb.
Lupinus jujuyensis C.P. Sm.
Lupinus juninensis C.P. Sm.

Lupinus kalenbornorum C.P. Sm.

Lupinus kellermanianus C.P. Sm.
Lupinus kerrii Eastw.
Lupinus killipianus C.P. Sm.
Lupinus kingii S. Watson

Lupinus klamathensis Eastw.
Lupinus kunthii J. Agardh

Lupinus kuschei Eastw. – Yukon lupine

Lupinus lacus C.P. Sm.

Lupinus laetus Wooton & Standl.
Lupinus laevigatus Benth.
Lupinus lagunae-negrae C.P. Sm.
Lupinus lanatocarpus C.P. Sm.
Lupinus lanatus Benth.
Lupinus lapidicola A. Heller – Mt. Eddy lupine

Lupinus latifolius J. Agardh
subsp. dudleyi (C.P.Sm.) P.Kenney & D.B.Dunn
subsp. latifolius J. Agardh
var. latifolius J. Agardh – broadleaf lupine
var. barbatus – Klamath lupine, bearded lupine
subsp. leucanthus (Rydb.)P.Kenney & D.B.Dunn
subsp. longipes (Greene) P.Kenney & D.B.Dunn
subsp. parishii (C.P.Sm.) P.Kenney & D.B.Dunn
subsp. viridifolius (A.Heller) P.Kenney & D.B.Dunn

Lupinus laudandrus C.P. Sm.

Lupinus lechlerianus C.P. Sm.
Lupinus ledigianus C.P. Sm.
Lupinus lelandsmithii Eastw.
Lupinus lemmonii C.P. Sm.

Lupinus lepidus Lindl. – prairie lupine
var. aridus (Douglas) Jeps.
var. confertus (Kellogg) C. P. Sm.
var. lepidus Lindl.
var. lobbii (A. Gray ex S. Watson) C. L. Hitchc.
var. sellulus (Kellogg) Barneby
var. utahensis (S. Watson) C. L. Hitchc.
Lupinus leptocarpus Benth.
Lupinus leptophyllus Cham. & Schltdl.

Lupinus lespedezoides C.P. Sm.

Lupinus leucophyllus Lindl. – woolly-leaf lupine

Lupinus lilacinus A. Heller
Lupinus lindenianus C.P. Sm.
Lupinus lindleyanus J. Agardh
Lupinus linearis Desr.
Lupinus littoralis Lindl. – seashore lupine
Lupinus lobbianus C.P. Sm.

Lupinus longifolius (S. Watson) Abrams – longleaf bush lupine

Lupinus lorenzensis C.P. Sm.
Lupinus ludovicianus Greene – San Luis Obispo County Lupine
Lupinus luetzelburgianus C.P. Sm.

Lupinus luteolus Kellogg – butter lupine, pale yellow lupine
Lupinus lutescens C.P. Sm.
Lupinus lutosus A. Heller
Lupinus lyallii A. Gray
subsp. alcis-temporis (C.P. Sm.) B.J.Cox
subsp. lyallii A. Gray – Lyall's lupine
subsp. minutifolius (Eastw.) B.J.Cox
subsp. washoensis (A.Heller) B.J.Cox

Lupinus macbrideanus C.P. Sm.

Lupinus macranthus Rose

Lupinus maculatus Rydb.
Lupinus madrensis Seem.
Lupinus magdalenensis C.P. Sm.
Lupinus magnificus M.E. Jones
Lupinus magniflorus C.P. Sm.
Lupinus magnistipulatus Planchuelo & D.B. Dunn
Lupinus malacophyllus Greene
Lupinus malacotrichus C.P. Sm.
Lupinus maleopinatus C.P. Sm.
Lupinus mandonanus C.P. Sm.
Lupinus mantaroensis C.P. Sm.

Lupinus marinensis Eastw.
Lupinus mariposanus Eastw.
Lupinus martensis C.P. Sm.
Lupinus martinetianus (C.P. Sm.) C.P. Sm.
Lupinus mathewsianus C.P. Sm.
Lupinus matucanicus Ulbr.
Lupinus meionanthus A. Gray
Lupinus melaphyllus C.P. Sm.
Lupinus menziesii J. Agardh
Lupinus meridanus C.P. Sm.

Lupinus metensis C.P. Sm.
Lupinus mexicanus Lag.
Lupinus michelianus C. P. Sm.

Lupinus microcarpus Sims
var. densiflorus
var. microcarpus – wide-bannered lupin, chick lupin
Lupinus microphyllus Desr.

Lupinus minimus Hook.

Lupinus mirabilis C.P. Sm.

Lupinus misticola Ulbr.
Lupinus mollendoensis Ulbr.
Lupinus mollis A. Heller
Lupinus monensis Eastw.
Lupinus monserratensis C.P. Sm.
Lupinus montanus Kunth
subsp. glabrior (S.Watson) D.B.Dunn & Harmon
subsp. montanus Kunth
subsp. montesii (C.P.Sm.) D.B.Dunn & Harmon

Lupinus monticola Rydb.
Lupinus montigenus A. Heller

Lupinus moritzianus Kunth
Lupinus mucronulatus Howell
Lupinus muelleri Standl.

Lupinus multiflorus Desr.

Lupinus munzianus C.P. Sm.
Lupinus munzii Eastw.
Lupinus mutabilis Sweet – Andean lupin, pearl lupin, South American lupin, tarwi, tarhui, chocho

Lupinus nanus Benth. – dwarf lupin, field lupin, sky lupin, Douglas' annual lupin
Lupinus navicularius A. Heller
Lupinus nehmadae C.P. Sm.

Lupinus neocotus C.P. Sm.
Lupinus neomexicanus Greene
Lupinus nepubescens C.P. Sm.
Lupinus nevadensis A. Heller – Nevada lupine
Lupinus niederleinianus C.P. Sm.
Lupinus nipomensis Eastw. – Nipomo Mesa lupine

Lupinus niveus S. Watson
Lupinus nonoensis C.P. Sm.
Lupinus nootkatensis Sims – Nootka lupin
Lupinus notabilis C.P. Sm.
Lupinus nubigenus Kunth
Lupinus nubilorum C.P. Sm.

Lupinus obscurus C.P. Sm.
Lupinus obtusilobus A. Heller – bluntlobe lupine
Lupinus ochoanus C.P. Sm.
Lupinus ochroleucus Eastw.
Lupinus odoratus A. Heller – royal Mojave lupin
Lupinus onustus S. Watson – Plumas lupine
Lupinus opertospicus C.P. Sm.
Lupinus oquendoanus C.P. Sm.

Lupinus oreganus A. Heller – Oregon lupin
Lupinus oreophilus Phil.
Lupinus ornatus Lindl.
Lupinus oscar-haughtii C.P. Sm.
Lupinus ostiofluminis C.P. Sm.

Lupinus otto-buchtienii C.P. Sm.
Lupinus otto-kuntzeanus C.P. Sm.
Lupinus otuzcoensis C.P. Sm.
Lupinus ovalifolius Benth.

Lupinus pachanoanus C.P. Sm.
Lupinus pachitensis C.P. Sm.
Lupinus pachylobus Greene
Lupinus padre-crowleyi C.P. Sm. – DeDecker's lupine, Father Crowley's lupine
Lupinus pallidus Brandegee

Lupinus paniculatus Desr.
Lupinus paraguariensis Chodat & Hassl.
Lupinus paranensis C.P. Sm.

Lupinus paruroensis C.P. Sm.
Lupinus parviflorus Hook. & Arn. – lodgepole lupin
subsp. myrianthus (Greene) Harmon
subsp. parviflorus Hook. & Arn.
Lupinus parvifolius Gardner
Lupinus pasachoensis C.P. Sm.
Lupinus pasadenensis Eastw.
Lupinus patulus C.P. Sm.
Lupinus paucartambensis C.P. Sm.
Lupinus paucovillosus C.P. Sm.

Lupinus paynei Davidson
Lupinus pearceanus C.P. Sm.
Lupinus pendentiflorus C.P. Sm.
Lupinus peirsonii H. Mason – Peirson's lupine, long lupine
Lupinus penlandianus C.P. Sm.

Lupinus perblandus C.P. Sm.
Lupinus perbonus C.P. Sm.
Lupinus perennis L. – wild perennial lupine, sundial lupine, Indian beet, old maid's bonnets
subsp. gracilis (Nutt.) D.B.Dunn
subsp. occidentalis S. Watson
subsp. perennis L.
Lupinus perglaber Eastw.
Lupinus perissophytus C.P. Sm.
Lupinus persistens Rose
Lupinus peruvianus Ulbr.
Lupinus philippianus C.P. Sm.
Lupinus physodes Douglas
Lupinus pickeringii A. Gray

Lupinus pilosellus Eastw.
Lupinus pilosissimus M. Martens & Galeotti
Lupinus pinguis Ulbr.
Lupinus pipersmithianus J.F. Macbr.
Lupinus pisacensis C.P. Sm.
Lupinus piurensis C.P. Sm.
Lupinus platamodes C.P. Sm.
Lupinus plattensis S. Watson
Lupinus platyptenus C.P. Sm.

Lupinus polycarpus Greene – smallflower lupin
Lupinus polyphyllus Lindl. – largeleaf lupine, bigleaf lupine, garden lupin, many-leaved lupine
var. burkei (S. Watson) C. L. Hitchc.
var. humicola (A.Nelson) Barneby
var. pallidipes (A. Heller) C. P. Sm.
var. polyphyllus Lindl.
var. prunophilus (M. E. Jones) L. Ll. Phillips
Lupinus poopoensis C.P. Sm.
Lupinus popayanensis C.P. Sm.

Lupinus potosinus Rose
Lupinus praealtus C.P. Sm.
Lupinus praestabilis C.P. Sm.
Lupinus praetermissus C.P. Sm.
Lupinus pratensis A.Heller – Inyo Meadow lupine
Lupinus pringlei Rose
Lupinus proculaustrinus C.P. Sm.
Lupinus prostratus J. Agardh
Lupinus protrusus C.P. Sm.
Lupinus prouvensalanus C.P. Sm.
Lupinus prunophilus M.E. Jones – hairy bigleaf lupin

Lupinus pseudopolyphyllus C.P. Sm.
Lupinus pseudotsugoides C.P. Sm.
Lupinus pubescens Benth.
Lupinus pucapucensis C.P. Sm.

Lupinus pulloviridus C.P. Sm.
Lupinus pulvinaris Ulbr.

Lupinus punto-reyesensis C.P. Sm.
Lupinus puracensis C.P. Sm.
Lupinus purdieanus C.P. Sm.
Lupinus pureriae C.P. Sm.

Lupinus purosericeus C.P. Sm.
Lupinus pusillus Pursh – rusty lupine or dwarf lupine
subsp. intermontanus (A.Heller) D.B.Dunn
subsp. pusillus Pursh
Lupinus puyupatensis C.P. Sm.
Lupinus pycnostachys C.P. Sm.
Lupinus quellomayus C.P. Sm.
Lupinus quitensis C.P. Sm.
Lupinus radiatus C.P. Sm.
Lupinus ramosissimus Benth.
Lupinus reflexus Rose
Lupinus regalis Bergmans
Lupinus regnellianus C.P. Sm.
Lupinus reineckianus C.P. Sm.
Lupinus reitzii Burkart ex M. Pinheiro & Miotto

Lupinus retrorsus L.F. Hend.
Lupinus revolutus C.P. Sm.
Lupinus richardianus C.P. Sm.
Lupinus rimae Eastw.
Lupinus rivularis Lindl. – riverbank lupin
Lupinus romasanus Ulbr.

Lupinus roseolus Rydb.
Lupinus roseorum C.P. Sm.
Lupinus rotundiflorus M.E. Jones

Lupinus rowleeanus C.P. Sm.

Lupinus ruber A. Heller

Lupinus rubriflorus Planchuelo

Lupinus ruizensis C.P. Sm.
Lupinus rupestris Kunth
Lupinus rusbyanus C.P. Sm.
Lupinus russellianus C.P. Sm.
Lupinus sabinianus Lindl.
Lupinus sabinii Hook.
Lupinus sabulosus A. Heller
Lupinus salticola Eastw.
Lupinus sandiensis C.P. Sm.
Lupinus santanderensis C.P. Sm.
Lupinus sarmentosus Desr.
Lupinus saxatilis Ulbr.
Lupinus saxosus Howell – rock lupine
Lupinus schwackeanus C.P. Sm.
Lupinus seifrizianus (C.P. Sm.) C.P. Sm.
Lupinus sellowianus Harms
Lupinus sellulus Kellogg
var. lobbii (S.Watson) B.J.Cox
var. sellulus Kellogg
var. ursinus (Eastw.) B.J.Cox
Lupinus semiprostratus C.P. Sm.
Lupinus semperflorens Benth.
Lupinus sericatus Kellogg – Cobb Mountain lupine

Lupinus sericeus Pursh – Pursh's silky lupin
var. barbiger (S.Watson) S.L.Welsh
var. sericeus Pursh
Lupinus setifolius Planchuelo & D.B. Dunn
Lupinus shastensis Lupinus albicaulis
Lupinus shockleyi S. Watson – purple desert lupine
Lupinus sierrae-blancae Wooton & Standl.
subsp. aquilinus (Wooton & Standl.) L.S.Fleak & D.B.Dunn
subsp. sierrae-blancae Wooton & Standl.
Lupinus simonsianus C.P. Sm.
Lupinus simulans Rose
Lupinus sinaloensis C.P. Sm.
Lupinus sitgreavesii S. Watson

Lupinus smithianus Kunth
Lupinus solanagrorum C.P. Sm.
Lupinus sonomensis A. Heller
Lupinus soratensis Rusby
Lupinus soukupianus C. P. Smith ex J. F. Macbr.
Lupinus sparsiflorus Benth. – desert lupin, Coulter's lupin, Mojave lupin

Lupinus spectabilis Hoover – shaggyhair lupine
Lupinus splendens Rose
Lupinus spragueanus C.P. Sm.

Lupinus staffordiae C.P. Sm.

Lupinus stipulatus J. Agardh
Lupinus stiversii Kellogg – harlequin annual lupine

Lupinus storkianus C.P. Sm.
Lupinus subacaulis Griseb.

Lupinus subcarnosus Hook. – buffalo clover
Lupinus subcuneatus C.P. Sm.
Lupinus subhamatus C.P. Sm.
Lupinus subinflatus C.P. Sm.
Lupinus sublanatus Eastw.
Lupinus submontanus Rose
Lupinus subsessilis Benth.
Lupinus subtomentosus C.P. Sm.

Lupinus subvexus C.P. Sm.
Lupinus succulentus K. Koch – succulent lupin, arroyo lupin, hollowleaf annual lupin
Lupinus sufferrugineus Rusby
Lupinus suksdorfii Robinson
Lupinus sulfureus Douglas
subsp. kincaidii (Suksd.) L. Ll. Phillips – Kincaid's lupin
subsp. subsaccatus (Suksd.) L. Ll. Phillips
subsp. sulfureus Douglas – sulfur lupin, sulfur-flowered lupin
Lupinus surcoensis C.P. Sm.

Lupinus syriggedes C.P. Sm.
Lupinus tacitus C.P. Sm.
Lupinus tafiensis C.P. Sm.
Lupinus talahuensis C.P. Sm.
Lupinus tamayoanus C.P. Sm.

Lupinus tarapacensis C.P. Sm.
Lupinus tarijensis Ulbr.
Lupinus tarmaensis C.P. Sm.
Lupinus tatei Rusby
Lupinus taurimortuus C.P. Sm.
Lupinus tauris Benth.
Lupinus tayacajensis C.P. Sm.
Lupinus tegeticulatus Eastw.

Lupinus tetracercophorus C.P. Sm.
Lupinus texanus Hook.
Lupinus texensis Hook. – Texas bluebonnet
Lupinus thompsonianus C.P. Sm.
Lupinus tidestromii Greene – Tidestrøm's lupin
var. layneae (Eastw.) Munz
var. tidestromii Greene

Lupinus tolimensis C.P. Sm.
Lupinus tomentosus DC.
Lupinus tominensis Wedd.
Lupinus toratensis C.P. Sm. – warwanzo, lito
Lupinus tracyi Eastw. – Tracy's lupine
Lupinus triananus C.P. Sm.

Lupinus truncatus Hook. & Arn. – collared annual lupine
Lupinus tucumanensis C.P. Sm.
Lupinus ulbrichianus C.P. Sm.
Lupinus uleanus C.P. Sm.
Lupinus ultramontanus C.P. Sm.
Lupinus umidicola C.P. Sm.
Lupinus uncialis S. Watson
Lupinus uncinatus Schltdl.
Lupinus urcoensis C.P. Sm.

Lupinus urubambensis C.P. Sm.

Lupinus valerioi Standl.
Lupinus vallicola A. Heller – open lupin
subsp. apricus (Greene) D.B.Dunn
subsp. vallicola A. Heller
Lupinus vargasianus C.P. Sm.
Lupinus varicaulis C.P. Sm.

Lupinus variicolor Steud. – varied lupin
Lupinus velillensis C.P. Sm.
Lupinus velutinus Benth.
Lupinus venezuelensis C.P. Sm.
Lupinus ventosus C.P. Sm.
Lupinus verbasciformis Sandwith
Lupinus verjonensis C.P. Sm.
Lupinus vernicius Rose

Lupinus viduus C.P. Sm.
Lupinus vilcabambensis C.P. Sm.
Lupinus villosus Willd.

Lupinus visoensis J.F. Macbr.
Lupinus volubilis C.P. Sm.

Lupinus weberbaueri Ulbr.

Lupinus werdermannianus C.P. Sm.
Lupinus westianus Small
var. aridorum (McFarlin ex Beckner) Isely
var. westianus Small
Lupinus whiltoniae Eastw.
Lupinus wilkesianus C.P. Sm.
Lupinus williamlobbii C.P. Sm.
Lupinus williamsianus C.P. Sm.

Lupinus xanthophyllus C.P. Sm.
Lupinus xenophytus C.P. Sm.

Lupinus yanahuancensis C.P. Sm.
Lupinus yarushensis C.P. Sm.

Lupinus ynesiae C.P. Sm.

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 (Boiss. & Spruner) Franco & P.Silva
subsp. termis (Forsk.) 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 1974
Lupinus cosentinii Guss. 1828 – sandplain lupin

Lupinus digitatus Forsk. 1775

Lupinus hispanicus Boiss. & Reut. 1842
subsp. bicolor (Merino) 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 (Atab) Kurl. et Stankev.
var. melanospermus Kurl. et Stankev.
var. niger Kurl. et Stankev.
var. cremeus Kurl. et Stankev.
var. leucospermus Kurl. et Stankev.
var. sulfureus (Atab.) 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. 1828
Lupinus palaestinus Boiss. 1849 – white-grey lupine
Lupinus pilosus Murr. 1774 – blue lupine
Lupinus princei Harms 1901

Lupinus somaliensis Baker f. 1895

Species names with uncertain taxonomic status

The status of the following binomials is unresolved: 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

thumb|upright|[[Papilio canadensis|Canadian tiger swallowtail on wild perennial lupine, Gatineau, Quebec]]

Certain species, such as the yellow bush lupin (L. arboreus), 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 (butterflies and moths). These include:

Iraricia icarioides missionensis (Mission blue butterfly), larvae limited to Lupinus
Callophrys irus (frosted elfin), recorded on L. perennis
Erynnis persius (Persius duskywing)
†Glaucopsyche xerces (Xerces blue)
Glaucopsyche lygdamus (silvery blue)
Plebejus melissa samuelis (Karner blue)
Erynnis persius persius (eastern Persius duskywing)
Schinia sueta, larvae limited to Lupinus

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.

File:Lupinus Polyphyllus Hybrid 4.png|Russell hybrid lupin Lupinus polyphyllus, UK

File:Mountaintop Lupin overlooking Raspberry Strait, Alaska 2009 114.jpg|Lupinus sp., Raspberry Island, Alaska, United States

File:Lupinus in Hokkaido 20080630.jpg|Lupins in Hokkaido, Japan

File:Lupin Leaf.jpg|Lupin cultivar "My Castle"

File:Flores de lupino (Ushuaia).jpg|Ornamental lupins, Ushuaia, Argentina

File:Lupins at lake tekapo.jpg|Lupins at Lake Tekapo, New Zealand

File:Lupins, Lake Tekapo, NZ.jpg|Lupins at Lake Tekapo, New Zealand

File:Bluebonnet-8100.jpg|Texas Bluebonnet, Texas, United States

File:Lupins in Pacific Northwest Washington.png|alt=Lupins_PacificNorthwest_Washington_USA|Lupins in Pacific Northwest Washington, United States

</gallery>

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.

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 (low alkaloid) 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, 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, 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

thumb|right|[[Lupinus angustifolius]]

thumb|right|[[Lupinus pilosus in Tel Aviv University, Israel]]

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 .

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 (Lupinus mutabilis) 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 or , 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 , Spanish domination led to a change in the eating habits of the indigenous peoples, and only recently (late 20th century onward) has interest in using lupins as a food been renewed. and L. hirsutus (blue lupin) 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 (L. texensis), are the state flowers of Texas.

Notes

References

External links

Sorting Lupinus Names Multilingual Multiscript Plant Name Database. University of Melbourne.
Biodiversity of LUPINS
Hughes, C. Lupinus.
Lupins.org
Superhydrophobicity in Lupins – video and commentary.