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.

The part of the plant which is eaten is the leguminous seed. It is nutritious, sustainable, and affordable. Because they are bitter and contain strong alkaloids, they must be pickled prior to consumption, to avoid toxicity. They can also be made into a flour and used as fertilizer. They were historically eaten in the Mediterranean, North Africa, and Latin America, but have now gained popularity in other parts of the world, such as Denmark and Belarus. In spite of lupins not being native to Australia, in the past decade it has produced 85 percent of commercially grown lupins.

There are ancient texts which refer to the utilization of lupine seeds, such as the Mishnah, and the Roman writer Pliny the Elder. Evidence of their consumption have been found in Egyptian Pharonic tombs and in the Western Galilee, Israel, with the latter dating back to hunter and gatherer Natufian cave residents.

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

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.Kurlovich, B. S. and A. K. Stankevich. (eds.) Classification of Lupins. In: Lupins: Geography, Classification, Genetic Resources and Breeding. St. Petersburg: Intan. 2002. pp. 42–43. Accessed 2 August 2013.

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.Naganowska, B., et al. (2005). 2C DNA variation and relationships among New World species of the genus Lupinus (Fabaceae). Plant Systematics and Evolution 256(1–4), 147–57. 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 (L. mutabilis, L. polyphyllus). 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' 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. NelsonSome sources treat Lupinus alpestris as a synonym of Lupinus argenteus.

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 lupineSome sources treat Lupinus aridorum as a synonym of Lupinus westianus. Lupinus aridulus C.P. Sm. Lupinus aridus Lindl.Some sources treat Lupinus aridus as a synonym of Lupinus lepidus. 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 GardnerSome sources treat Lupinus attenuatus as a synonym of Lupinus coriaceus.

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 lupineSome sources treat Lupinus brevicaulis as a synonym of Lupinus grisebachianus. 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 lupineSome sources treat Lupinus burkei as a synonym of Lupinus polyphyllus. 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 lupineSome sources treat Lupinus caespitosus as a synonym of Lupinus lepidus. 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.