Monday, April 25, 2016

CLASSIFICATION OF LUPINS





CLASSIFICATION OF LUPINS


B.S. Kurlovich and A. K. Stankevich



Basic characteristics of taxa in the genus Lupinus L. ..,,,... ,,,

Successes in genetic studies and breeding practice depend on the availability of a well-developed phylogenic system of a genus. Regretfully enough, by now there is no comprehensive monographic and systematic review of Lupinus L. The presence of such a breach may be explained by inaccessibility of the habitats of the American lupin species, large variability and weak differentiation of characters in lupins, and intricacy of their classifications. As a result, it is still unknown how many species of lupin exist in the nature. This circumstance hampers the solution of numerous theoretical and practical problems. Each form of lupin can be assessed as an object fitting in with a number of consecutive taxa. A taxon is a part of plant community consisting of a certain set of individual plants distinguished according to genetic principle by the uniformity of their genesis, and regarded as a formal unit at any level of hierarchic classification. This genus is the principal object of our research. Geographic differences between the New and Old World’s lupins are shown in Table 1 where the grouping of both subgenera is introduced. Different species, subspecies, varieties, subvarieties and forms were classified on the basis of Vavilov’s concepts (the law of homologous series in hereditary variation, studies on the problem of the species as a system, differential systematic and geographical method of crop studies, and others). Vavilov’s (1931, 1965) concept about the species as a complex multilateral and mobile phenomenon implies application of diverse methods for identification of differences between intraspecific categories. We recognize subspecies (subsp.) as an isolated group of individual plants within a population of a species. They occupy certain part of the area of a species, constitute together a mobile system, are able to cross among themselves and with plants growing in other parts of the area of this species, produce prolific progeny, possess distinctive morphological and inheritable characters in vegetative and generative organs with the uniform genetic base, and incorporate transient forms. (subsp. graecus,termis and albus within the limits of Lupinus albus L.). Allelism and character complementarity tests have shown that in lupins the color of seed is correlated with the color of the corolla. This linkage reflects the stability of genetic system, which corresponds to the rank of varieties (var.). A good diagnostic character is the color of vegetative parts, and the absence or presence of anthocyan, in particular. Being less stable, it could be used in identifying subvarieties (subvar.). Considerable practical interest for breeders may be generated by the plants with determinate branching, fascicular stem and other characters of breeding value. Such forms are theoretically possible in all the varieties and subvarieties systematized by us. Therefore, it seems justified to regard them in the rank of forma (f.). The detailed characteristics and indices of the genus Lupinus L. are presented also in the section «Description».

,,, Interspecific diversity of lupins ,,,

The genus Lupinus L. and, in particular, its North-American species, were divided by Watson (1873) into three parts: Lupinus, Platycarpos andLupinnelus. Differences in habit and in the number of ovules was accepted as the basis for this classification. The majority of perennial and annual species from the American continent described by Watson was referred toLupinus. To the Platycarpos section were attributed some annual species with two ovules in the ovary and two seeds in the pod (L. densiflorusBenth., L. micricarpus Sims. and others). Section Lupinnelus consisted of one species (L. uncialis), with axillary and solitary flowers, scarcely reflexed banner, and also with two ovules in the ovary. Presently, the existence of such species seems doubtful. This principle of classification was extended by Ascherson and Graebner (1907) to all lupins from the eastern and western hemispheres. Genus Lupinus L. was for the first time subdivided into two subgenera: A. Eulupinus and B. Platycarpos(Ascherson and Graebner, 1907). Quantity of ovules (seedbuds) in the ovary and seeds in the pod was also accepted as the criterion for this division. Majority of the described species from the eastern and western hemispheres were referred to subgenus A. Eulupinus. Subgenus 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 Watson). These works were a starting point for our researches. In connection with the definition of two secondary centers of formation of different species of lupin in the eastern and western hemispheres, and also with the essential morphological differences between lupins of the two hemispheres, we managed to revise the volumes of two subgenera in the genus Lupinus L. according to the geographic principle, however in view of the findings of the previous writers. Subgen. Platycarpos (S.Wats.) Kurl. in our new combination integrates the numerous perennial and annual species from the western hemisphere, both groups having two, four and more ovules in the ovary, while subgen. Lupinus L. includes 11 species from the Mediterranean region and Africa with as a minimum four and more ovules in the ovary.
,,,
I. Subgen. Platycarpos (S.Wats.) Kurl. 1989, Bull. N.I. Vavilov Inst. Plant Industry 193:24.   -   §2. PLATYCARPOS  S.Wats. 1873, Proc. Amer. Acad. Arts Sci. 8:522;    B. Platycarpos Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:232;   §1. LUPINUS  S.Wats. 1873, l.c.:518, p.max.p.;    
AEulupinus Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221, p.p.     –  New World’s or flat-fruited lupins.

,,, 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. 2n = 36, 48, 96.

,,, The type of subgenus: L. densiflorus Benth.

Geographic distribution: North, Central and South America, predominantly in the mining systems of the Andes and Cordillera. Some species are cultivated (L. mutabilis Sweet., L. polyphyllus Lindl.). This subgenus includes several hundreds of species (from 200 up to 1000) requiring further analysis of their authenticity. In the following subsection we have presented the descriptions of 34 most studied species from this subgenus.

,,, II. Subgen. Lupinus. 1989Bull. N.I. Vavilov Inst. Plant Industry 193:23.  -   A. Eulupinus  Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221, p.min.p.  -  Old World’s lupins. 

,,, The ovary has at least four and more ovules or seedbuds. The seed are predominantly large, with the well-developed embryo, without endosperm. Cotyledons are large, with a short caulicles. The first pair of true leaves is opposite. The stem is always pubescent; dominating type of branching is sympodial. Leaflets are pubescent to different extent; they are predominantly broad, and only one species (L. angustifolius L.) has narrow leaflets. Pods are orbicular, with four and more seeds. Represented by annual herbaceous forms. Plants are self-pollinated; some of them are predisposed to cross-pollination. 2n = 32, 36, 38, 40, 42, 50, 52.

The type of subgenus: L. albus L.

Geographic distribution: Mediterranean region and Africa. Some species are cultivated (L. albus L., L. angustifolius L., L. luteus L., etc.).

This subgenus includes 11 species:
1. L. albus L. 1753, Sp. Pl.:721.
2. L. angustifolius L. 1753, Sp. Pl.:721.
3. L. micranthus Guss. 1828, Fl. Sic. Prodr. 2:440.
4. L. luteus L. 1753, Sp. Pl.:722.
5. L. hispanicus Boiss. et Reut. 1842, Diagn. Pl. Nov. Hisp. 10.
6. L. cosentinii Guss 1828, Fl. Sic. Prodr. 2:440.
7. L. digitatus Forsk. 1775, Fl. Aegypt.:131.
8. L. princei Harms, 1901, Bot. Jahrb. 28:401.
9. L. pilosus Murr. 1774, Syst. Veg. ed 13:545.
10. L. palaestinus Boiss. 1849, Diagn. Pl. Or. Nov. 9:9.
11. L. atlanticus Gladstones, 1974, Techn. Bull. Dept. Agr. West. Austr. 26:30.
12. L. somaliensis Baker, 1895, Byll. Roy. Card. Kew 105:213.



,,, Description of the New World ’s lupins 

(Subgen. Platycarpos (Wats.) Kurl.)


Perennial fruticose and fruticulose plants

1. L. arboreus Sims, 1803, Bot. Mag., tab.682. – L. sericeus Eschsch. 1826, Mem. Acad. Sci. Petersb. 10:289. – L. macrocarpus Hook. et Arn. 1832, Bot. Beach. 138:121. – L. rivularis Aghardh, 1835, Syn. Gen. Lupin. : 24, non Dougl. 1833. – L. fruticosus Hort. ex Stend. 1841, Nom. ed. 2, 2:78. – L. propinguus Greene, 1893, Erythea, 1:126. – L. treetype.

Commonly called the Tree Lupin this is a bushy plant. Leaflets are narrowed down to the basis, and are concentrated at the top, being almost leveled with the calyx, almost naked from above and hirsute from below. Stipules are styliform, pubescent, 5-10 mm in length. Inflorescences are abnormally verticilate. Floral shoots are almost leveled with calyces. The corolla is yellow with delicate odor. The vexillum is spherical, twice wider than the wings. The wings are oval, yellow. The carina is white with a dark-violet edge, strongly pubescent. Labia of the calyx are almost equal, the upper ones are bicrenate, with an arched base, the lower ones are entire, more or less blunted. Floral bracts are lanceolate, pubescent. Pods are brown, pubescent, 4-5 cm in length. Seed are small-sized, spherically flattened, dark-brown, with dark hilum. Grows on the coast of the Pacific Ocean. Widespread in Western and Central California. Was naturalized in Ireland, Great Britain, Canada (British Colombia), USA (Washington), and also in Chile.

2. L.rivularis Dougl. ex Lindl. 1833, Bot. Reg., tab. 1595, non Torr. 1875. –L. labiatus Nutt. ex Torr. et Gray, 1840, Fl. N, Amer. 1:376. – L. lignipesHeller, 1912, Muhlenbergia 8: 66.

Leaflets blunted, acuminate, very weakly pubescent from above and slightly from below. Stipules are almost crescent, a little bit shorter than inL. arboreus. Inflorescences are verticilate-alternate with drooping flowers. Floral shoots are hardly longer than the lower labium of the calyx. The carina of the flower is pale-violet. The vexillum is round, cream-colored, with maculation specks at the base. Wings are widely oval, inflated with striae on the edge. The carina is wedge-shaped, white with a dark-violet end, pubescent only at the top. The calyx is deeply bilabiate, with almost integral fingers (weakly bipartite). Floral bracts are lanceolate, easily falling. Pods are brown, pubescent with short soft down, 6-8 seeded. Seed are spherically oval, dark-colored, 5,5 x 4 mm in diameter. Widespread in Canada and the U.S.(California, western Oregon, and western Washington).

3. L.multiflorus Desr. in Lam. 1789, Encycl. 3:624. – L. albescens Hook. et Arn. in Hook. 1833, Bot. Misc. 2:201.

The plant is densely covered by silky hair. The stalk is erect, branching. Leaflets (7) are lanceolate, silky from both sides, 20 x 34 mm in size. Petioles are 5-7.5 cm in length, with bristling stipules. The inflorescence is a polyanthous apical truss, with flowers almost sedentary. The calyx has a bicrenate upper labium and a tricrenate lower one. The corolla is blue, twice longer than the calyx. Wings are leveled with the vexillum, the base is yellowish. Floral bracts are short or oblong. Widespread in Argentina, Brazil (eastern part of Santa Catalina), and Uruguay.

4. L. littoralis Dougl. ex Lindl. 1828, Bot. Reg., tab. 1198. – L. versicolorLindl. 1837, Bot. Reg., tab. 1979. – L. variicolor Stend. 1841, Nomencl. Bot. ed. II, 2:78. – L. nutkatensis Cooper, 1860, Pacif. Rail. Rep. 12:58. –L. franciscanus Greene, 1887, Pittonia, 1:64.

Plants are silvery-silky. Stems are thin, prostrate. Leaflets are reduced at the base, slightly blunted and acuminate at the top, pubescent from both sides. Their number is 5-9. Petioles are twice longer than leaflets. Stipules are styliform. The inflorescence is a friable truss. The calyx is without bractlets, its labium being almost entire. The corolla is purple, with maculation specks in the basis of the vexillum. The wings are longer than the vexillum. Pods are linear, squeezed, pillar-shaped, 40 x 8 mm in size. Seed are speckled, 4-5 x 3-3.6 mm in size. Weight of 1000 seeds is 27.6 g. Widespread in Canada (southwest part of British Columbia) and in the northwest of the United States (western Oregon and western Washington).

5. L. paniculatus Desr. in Lam. 1789, Encycl. 3: 624.

The bush is covered with silvery-silky, almost felt-like pubescence. Leaflets are lanceolate, acuminate, green and naked from above, hirsute from below, 40 x 3 mm in size, 7-9 in number. The inflorescence is an abnormally verticillate truss up to 15 cm in length. Stipules are oblong, longer that floral shoots, non-falling. The calyx is with bractlets, its upper labium has a depression, and the lower one is entire and a little bit longer than the upper. The corolla is violet, twice longer than the calyx. Lobes are almost equal lengthwise. The vexillum is twice broader than the wings. Pods are 4-5-seeded, 65 x 3 mm in size. Widespread in Argentina (northwest part), Colombia, and Peru. May occur in Ecuador and Bolivia.

6. L. chamissonis Eschsch. 1826, Mem. Acad. Sci. Peters. 10:288. – L. sericeus Hook. et Arn. 1832, Bot. Beech. Voy.:138. – L. albifrons Bent. 1835, Trans. Hort. Soc. London, Ser. 2, 1:410, - L. macrocarpus Torr. 1857, in Pacif. Rail. Rep. 4:81. – L. ornatus Torr. 1857, l. c.:81, non Dougl. 1833. – L. douglasii Torr. 1858, Bot. Mex. Bound.:57. – L. holosericeus A. Gray, 1873, Proc. Amer. Acad. Arts Sci. 8:379. – L. sericatus Kellogg, 1877, Proc. Calif. Acad. Sci. 7:92.

The bush has almost felt-like pubescence. Stalks are prostrate, densely foliated. Leaflets are densely ciliated from both sides, obovate, wedge-shaped, 7-9 in number. Petioles are longer than leaflets, with stipules. The inflorescence is a verticillate truss, 4-5 flowers in each verticil. Floral bracts are ovate. The upper labium of the calyx is bilabiate, the lower is longer than the upper, entire. Bractlets are small. The pod is black, weakly pubescent, 4-seeded, 30 x 9.5 mm. Seed are 5-6 x 3.5-3.8 mm. Weight of 1000 seeds is 30-35 g. Widespread in the northern part of Mexico, U.S.A. (California and Oregon). Inhabits sandy areas up to 60 m above sea level.

7. L. sericeus Pursh, 1814, Fl. Amer. Sept, 2:468. – L. ornatus Nutt. 1834, Journ. Acad. Sci. Philadelphia, 7:20. – L. flexuosus S.Wats. 1871, Bot. King’s Exp.55,56.

Bush-type plant with different degrees of pubescence, from dense down to weak. Stems are erect. Leaflets are lanceolate, reduced toward the base, pubescent from both sides, numbering 7-9. Petioles are a little bit longer than leaflets or have the same length. The size of stipules is very small. The inflorescence is a verticillate truss. The upper labium of the calyx is bidentate, the lower one is integral. Bractlets are very small-sized. The corolla is blue, pink or purple, and twice longer than the calyx. The vexillum is smooth. Pods are expressly rough, 4-5-seeded, 30 x 6 mm in size. Seed are spherical, yellowish, 5-7 mm in size. Widespread in Canada, USA (Arizona, northern California, Colorado, Idaho, Montana, Nevada, New Mexico, Oregon, South Dakota, Washington and Wyoming).

Perennial herbaceous plants

8. L. polyphyllus Lindl. 1829, Bot. Reg., tab.1096. – L. grandifoliusLindl. ex Agardh, 1935, Syn. Gen.Lup.:18. – L. macrophyllus Benth. in Sweet, 1836, Brit.Flow. Garg. 2, ser.2:4, tab.356.- L. magnus Greene, 1897, Pittonia, 3:160. – L. superbus Heller, 1906, Myhlenbergia, 2;209. – L. elongatus Greene ex Heller, 1910, Myhlenbergia, 6:17.- L. pallidipes Heller, 1911, Myhlenbergia, 7:91. – L. piperita Davidson, 1927, Byll. S. Callif. Acad. Sci.26:70. – L. perglaber Eastw. 1940, Leafl.West. Bot. 2:268. – L. pallidipes Heller, subsp. superbus (Heller) Munz, 1959, A. California Flora:828. - multifoliate or Washington lupin.

During the first year of vegetation, the plants usually develop predominantly radical, rosette-like leaves. In the second year, many stalks are derived. Young leaflets, flower pedicles and calyces have silvery pubescence. Stalks are powerful and pubescent. The leaf consists of 9-10 almost lanceolate leaflets, which are large but twice shorter than petioles. Leaflets are naked from above or weakly pubescent; in the lower part pubescence is stronger. Stipules are styliform, pubescent; on ¾ they grow together with petioles. Inflorescences are very long and usually friable. Flowers are semi-verticillate, speckled. Labia of the calyx are integral or is weakly dentate. Floral bracts have early falling habit; they are not higher than the calyx. The corolla is three times longer than the calyx, and may have diverse colors, but more often violet. The carina is naked. Floral shoots are long, of almost the same length as flowers. Pods are flat, elongated, with 9 (6-7) seeds. Seed are oval, weakly squeezed, having brown, black or a different color. (2n=48). 1000 seed weight is 20-21 g. Inhabits fluvial detrital deposits and inundated soils of the rivers. Widespread in Canada (British Columbia), USA (Alaska, northern California, western Oregon and Washington). Introduced in Eastern Canada and New Zealand (Hill, 1988, 1994; Hill and Tesfaye, 1994). Cultivated in European countries as a horticultural and fodder crop and for green manure. Selected were the forms with miscellaneous beautiful coloring of flowers: f. albus – with white flowers; f. tricolor – with blue-white-yellow flowers; f. atripurpureus - with dark red flowers; f.fleribunur – with pale blue flowers; f. roseus – with pink flowers (Ascherson and Graebner, 1906-1910). Very frequently occurs in the turned-wild condition (Belarus, Poland and Germany). In many countries, including Russia, Belarus and Ukraine, it is cultivated for green manure. In the last years, low alkaloid (sweet) forms of this species were obtained. On their basis, commercial cv. Pervenec (first sweet cultivar) was released. It is now registered in the Russian State Catalogue of breeding achievements. Breeding activities with fodder multifoliate lupin have also started in the Ukraine, Poland, Finland and other countries. Lupinus polyphyllus represents special interest for northern areas where other species of lupin do not ripen.

9. L. latifolius J.Agardh, 1835, Syn. Gen. Lupin.:18. – L. cytisoidesAgardh, 1835, l.: c.:81. – L. adsurgens Drev. 1889, Bull. Forrey Clab 16:150. – L. longipes Greene, 1891, Fl. Francisc.: 41. – L. viridifolliusHeller, 1905, Muhlenbergia 2:65. – L. columbianus Heller, 1912, Muhlenbergia 8:84. – L. confusus Heller, 1912, ibid.:63. – L. barbatusHeller, ibid.: 6. - L. agninus Gand. 1913, Bull. Soc. Bot. Fr. 60: 461. – L. parishii Eastw. 1939, Leafl. West. Bot. 2:181. – L. pennelianus Heller, 1941, Leafl. West. Bot. 3 ; 42. - platyphyllous lupin.

Stalks are numerous, erect, almost naked, and hollow inside. Leaves are set 10 cm above ground at the lowest, or higher. Leaflets, numbering 5-9 (10), are obovate, acuminate, naked from above and covered by soft white down from below. Petioles are longer than leaves, and pubescent. Stipules are styliform; up to ½ they grow together with petioles. Inflorescence is friable; together with flower pedicles it reached 50 cm or more. Flowers are clustered in trusses 6 to 30 cm long, positioned on long floral shoots. Floral bracts are easily falling. The calyx is pubescent; the labia are almost equal to each other. The upper labium is chaffy; the lower is coriaceous, green, equal to the flower shoot. The vexillum is spherical, lilaclilac with specks. Wings are oval, of the same color. The carina is beaked, lilac, and strongly pubescent on the inner side. Pods are large, densely pubescent, 4 x 1 cm in size, 4-6-seeded. Seed are oval, brown with a mottled ornament, 5 x 4 mm in size. 1000 seed weight is 50 g. Widespread in the USA (northwestern Arizona, western and northern California, northwestern Nevada, western Oregon, and southwestern Washington).

10. L. minimus Dougl. ex Don f.G. 1832, Gen. Syst. 2:367.

The plant is very small, up to 20 cm tall, with silky pubescence. The main stem is almost aphyllous, the generatrix of the apical inflorescence is surrounded by a rosette of leaves. The leaf consists of 7-9 obovate and lanceolate leaflets, covered by silky pubescence on both sides, three times shorter that petioles. Stipules are bristly; on 1/3 they grow together with petioles. Inflorescences are dense, abnormally verticillate, about 10 cm in length. Flowers are purple, set on short floral shoots. Floral bracts are styliform, almost non-falling, equal to ½ of the calyx. The upper labium of the calyx is weakly bidentate, the lower one is tridentate, without pubescence, with the medium denticle longer than the other two. Widespread in Canada.

11. L. leucophyllus Dougl. ex Lindl. 1827, Bot. Red., tab.1124. – L. plumosus Dougl. ex Lindl. 1829, Bot. Reg. tab. 1217. – L. densiflorus Nutt. ex Torr. et Gray, 1840, Fl. N. Amer. 1:379. – L. albicaulis Hook. 1847, Journ.Bot. (London) 6:216.

Stalks are erect, with silky pubescence. Leaflets are lanceolate, pilose from both sides; their number is 7-9. Petioles are twice longer than leaflets, pubescent. The inflorescence is verticillate-alternate. The calyx is twice shorter than the corolla, with bractlets, its upper labium being bilabiate and the lower one one-pieced and strongly blunted. Floral bracts are lanceolate, green and red, pubescent. Wings are blue-lilac. The carina has a beak and a violet edge. Pods are yellow-brown, weakly pubescent, 3.2 x 102.5 cm in size., 4-seeded. Seed are rhombic, chestnut-colored, 4.2-5.6 x 2.7-4.4 mm in size. 1000 seed weight amounts to 26.5 g. Widespread in the USA (northwestern Arizona, northern and western California, Nevada, Oregon, and Washington).

12. L. nootkatensis Donn , 1840, Cat. Hort. Contab. ed. 111:135.- L. variegatus Poir. 1814, Encycl. Suppl. 3:520. – L. blaschkeamus Fisch. et Mey. 1824, Jdn. Sem. Hort. Bot. Petrop. 11, Supl.:57. – L. nutcanusSpreng. 1826, Syst. Veg. 3:227. – L. regius Rudolph ex Torr. et Gray, 1840, Fl. Amer. 1:376.

Stalks are densely white-haired. Leaflets are obovate-oblong, blunted, naked from the upper part, and pubescent from the lower one. Number of leaflets is 7 (5-8). Petioles are lanceolate, almost equal to leaflets. Stipules are large and long; up to ½ they grow together with petioles. The inflorescence is sparse, verticillate or abnormally verticillate. Flowers are large. The labia of the calyx almost equal, bractlets are very small. The upper labium is semi-bilabiate. Floral bracts are longer than flowers. The corolla is twice longer than the calyx, blue, with dark nerves and maculation of specks flecks in the middle of the vexillum. The carina is naked. Pods are black, weakly pubescent, with a long spear, 5.0 x 4.0 cm in size, with 10-11 seeds. Seed size is 3.5-4.2 x 2.6-3.0 mm. 1000 seed weight is 16 g. Widespread in Alaska and Canada (British Columbia). Naturalized in Iceland, Norway and Scotland. Introduced in New Scotland and Newfoundland.

13. L. affinis Agardh , 1835, Syn. Gen. Lupin.:20. – L. carnosulus Greene, 1886, Bill. Calif. Acad. 2,6:144.

This species has a relationship to L. nootkatensis Donn. Plants are pubescent. Leaflets are obovate, pubescent from both sides. Petioles are twice longer than leaflets. Stipules are bristly; the grow together with the basis of petioles. The inflorescence is friable, verticilate, with flowers set on floral shoots. The calyx is pubescent; its upper labium is divided, with bractlets. The corolla is twice longer than the calyx, blue in color. There is a light yellow spot in the middle of the vexillum. The carina is ciliated. Pods are yellow, weakly pubescent, with a long spear, 40 x 8.5 mm in size, 8-seeded. Seed are 3.7-4.6 x 3-3.5 mm. 1000 seed weight is 25 g. Widespread in the USA (inshore part of California, and Oregon).

14. L. mexicanus Cerv. In Lag. 1816, Gen. Sp. Nov.:22, non Torr. 1853. – Mexican lupin.

Hirsute plant, with short rough hair. Stalks are powerful. Leaflets are elongated, cuneiform-expanded in the middle part, naked from above and downy from below; their number is 5-7. Petioles are shorter then leaflets. Stipules are styliform, bristly, long. Inflorescences are abnormally verticilate, odorless. The calyx has small-sized bractlets. The upper labium of the calyx is with two notches; the lower one is almost integral. Floral bracts are bristly, easily falling. The corolla is blue or lilac-violet. Lengthwise the wings are almost equal to the vexillum, but they are wider and lighter, with dark nerves. The carina is white with a dark edge. Pods are small and flat. (2n=48). Seed are small-sized, glossy, diverse in color. 1000 seed weight is 13-16 g. Cultivated as ornamental plants. This species is very polymorphous. Widespread in Western Mexico.

15. L. perennis L. 1753, Sp.Pl. – perennial or sunny lupin.

In the first year of vegetation, derives only rosette-like radical leaves. Stalks are numerous, erect, striated, slightly pubescent. Leaflets are obovate, with a blunted apex or pointed spear, naked from above, sparsely pubescent from below; their number is 7-11. Petioles are longer then leaflets; stipules are very small, almost missing. The inflorescence is long, sparsely flowered, sometimes almost verticilate. The calyx is silky, without bractlets; its upper labium with a protuberant basis, is integral or weakly emarginate, the lower one is integral, almost twice longer then upper. Floral bracts are styliform, shorter then the calyx, early falling. The corolla is purple or white, three times longer than the calyx. The vexillum is shorter than the wings. The carina is weakly ciliate. Pods are yellow-grayish-brown, with straight lines, necklace-shaped, short and closely hirsute, easy shattered, with 5-6 seeds. Seed is oval with a light hilum. Widespread in the eastern part of the USA (from Florida to Canada), Canada (south of Ontario), and on the coasts of the Arctic Ocean. Lives on sand hills.

16. L. argenteus Pursch, 1814, Fl. Amer. Sept. 2:468. – L. laxiflorusDougl. in Lindl., 1828, Bot. Reg. tab. 1140. – L. decumbens Torr. 1828, Ann.Lyc. N. Jork, 2:191. – L. tenellus Dougl. ex G. Don, 1832, Hist. Dichl. Pl. 2:267. – L. laxiflorus Hook. 1847, Journ. Bot. (London) 6: 215. – L.foliosus Hook. 1847, ibidem. – L. decumbens Torr. 1857, Pacif. Rail. Rep. 4:81. – L. ornatus Newb. 1858, Pacif. Rail. Rep. 6:70. – L. argenteusS.Wats. 1871, ibidem. – L. laxiflorus A.Gray, Proc. Amerr. Acad. Arts Sci. 8:379. – silvery lupin.

Plants are pubescent. Stalks are slightly pubescent. Leaflets are obovate and linear lanceolate, silky from both sides, almost twice shorter then petioles; their number being 9-11. Stipules are awl-shaped, bristled, very small-sized. The inflorescence is friable, almost verticillate, slightly pubescent. The calyx has a rather long tube; its upper labium is bursiform at the basis, while the lower is integral. The corolla is purple or white, twice higher than the calyx. Widespread in Canada and in the western part of the USA.

17. L. sulphureus Dougl. ex G. Donf. 1832, Gen. Syst. 2:367. – L. sericeus Torr. et Gray, 1840, Fl. N. Amer. 1:379 p.p.

Stalks are thin and pubescent. Leaflets are linear-lanceolate, with dense silky pubescence on both sides, shorter then petioles, 13-15 in number. Stipules are small, styliform. Inflorescences are broad, abnormally verticilate. Floral bracts are early falling. The calyx is without bracklets; the upper labium is protuberant at the basis and deeply emarginate at the apex, the lower one is integral, and a little bit longer than the upper. The corolla is orange-yellow, surpassing the calyx by 1/3 lengthwise. The carina is naked. Widespread in Canada (British Columbia) and the USA (eastern Oregon and eastern Washington).

18. L. albicaulis Dougl. in Hook. 1833, Fl. Bor. Amer. 1:165. – L. foliosusNutt. ex Torr. et Gray, 1840, Fl. N. Amer, 1:377. – L. falcifor Nutt. ex Torr. et Gray, 1840, l. c.: 378. – L. mexicanus Torr. 1853, Sigr. Rep. :158, non Cerv. 1816. – L. laxiflorus Torr. 1857, Pacif. Rail. Rep. 4:81, non Dougl. 1828, nec Hook. 1847. – L. sylvestris E. Drev. 1889, Byll. Torr. Club.:150. – L. elmeri Greene, 1897, Pittonia, 3:159. – L. purpurascens Heller, 1905, Myhlenbergia, 2:66. – L. shastensis Heller, 1905, ibid.:67. – L. ochroleucusEastw. 1942, Leafl. West. Bot. 3:171.

Stalks are almost naked, but leafed. Leaflets are obovate-oblong, having silky pubescence on both sides silky-hirsute; their number is 9. Petioles are equal to leaflets. Stipules are bristly, very small-sized. The inflorescence is rich, more or less verticillate, elongated, and broad. The calyx is reddish, silky, without bractlets. The labia of the calyx are almost equal lengthwise; the upper one is bilabiate, the lower is integral. Floral bracts are lanceolate, equal in length to floral shoots, with late falling habit. The corolla is yellow, twice longer than the calyx. The carina is strongly bent, naked. Widespread in the USA.

19. L. aridus Dougl. in Lindl. 1829, Bot. Reg, tab 1242, - L. brachypodusPiper, 1902, Bull. Torr. Clab, 29:642. – L. abortivus Greene, 1912, Myhenbergia, 8:177. – L. volutans Greene, 1912, l. c. :178.

Plants are dwarfish with hard pubescence. The stem is strongly branching at the basis. Leaflets are obovate-oblong, covered by long rigid hair, thrice shorter then petioles; their number is 7. Stipules are styliform, growing together with leafstalks in the basis. The inflorescence is conical, abnormally verticillate, and dense. Flowers are set on short floral shoots. Floral bracts are styliform, shorter then bractlets of the calyx, not falling even during pod formation. The upper labium of the calyx is bilabiate, the lower one is integral. The corolla is blue (?), surpassing the calyx by 1/3. The carina is closely ciliated. Pods are black, strongly pubescent, 4-seeded. Seed are spherical, yellowish, 5.3-6.3 x 3.2-3.8 mm. 1000 seed weight is 29.5 g. Widespread in the USA. Occurs on fluvial soils and lake sandstones.

Annual herbaceous plants

20. L. mutabilis Sweet. 1825, Brit. Flower Gard. Ser.1, 2. tab.130. -L.cruckshanksii Hook. 1831, Bot. Mag. tab 3056. – L. mutabilis Lindl. 1832, Bot. Reg. tab. 1532. – mutable, pearl or Andean lupin.

Annual or biennial plant, up to 2 m high, naked, with waxen raid. Branching and leaf bearing are inconsiderable. There are 11-18 leaves on the main stem. Most strongly developed are 3-4 uppermost shoots. Leaflets are obovate-oblong, acuminate, blunted; their number is 7-9 (11). Stipules are small-sized, growing together at the basis with a pulpy cushion of petioles. Inflorescences are friable, semi-verticillate or alternate. Flower pedicles are long (7-10 cm); the inflorescence is 8-20 cm in length. Flowers are clustered in 9-10 verticils on 5 flowers. The calyx has almost equal labia; the lower one is integral, while the upper one is bilabiate. Bractlets are bristly. Floral bracts are styliform, small. The corolla is odorous, with diversified coloring (blue, white, pink, violet, etc.), 1.8-2.0 cm in size. The vexillum and the wings are broad, longer than the carina. Pods are pubescent, non-shattering, 5-6-seeded. Seed have high oil content (more than 20%), they are white or may have other various colors, large, smooth, sometimes with a brown spot. Most widespread in Peru, Colombia, and Chile. Domesticated by ancient inhabitants of Peru as early as B.C. Its seed were used for food after soaking in water and cooking in order to remove toxic alkaloids. In recent years, low alkaloid (sweet) forms were identified. This species undergoes intensive breeding in many countries. The first stable low alkaloid variety Inti was bred in Chile (Römer and Jachn-Deesbach, 1988; Baer E. and Baer D., 1988). The form with determined branching has been obtained in Germany (Römer, 1994). Cultivated in many countries of America and Europe (Baer, D. von et al., 1994; Baer, E. von et. al., 1994; Römer, 1994). In the conditions of Russia, is distinguished by lateness and susceptibility to diseases (especially to anthracnose).

21. L. ornatus Dougl. ex Lindl. 1828, Bot. Reg. tab. 1216. – L. argenteusAgardh, 1835, Syn. Gen. Lupin.:27, non Pursch, 1814. – L. leucapsisAgardh. 1835, l. c. :29. – L. sericeus Cooper, 1860, Pacif. Rail. Rep.12:52, non Pursch, 1814.

Annual plant, generically close to L. mutabilis, pubescent with weakly pressed hair. The stem is powerful, 40-80 cm in height. Branching initially goes in an ascending direction. After formation of the apical inflorescence, lateral branches develop from the top downward. Leaflets are obovate-oblong, from both sides covered by silky pubescence; their number is 7-9. Petioles are 2-3 times longer than leaves. Stipules are styliform, growing up to middle together with petioles. The inflorescence is long, verticillate or semi-verticillate. The upper labium of the calyx is divided, the lower one is integral. Floral bracts are lanceolate, falling. The corolla is blue, white, or violet, with nerves. The vexillum is yellowish in the middle, with silky pubescence outside. The wings and the carina are white. Pods are pubescent, 3-5-seeded. Seed are small-sized, with various coloring. This species is rather polymorphous, with outstanding ornamental qualities. Widespread in the USA.

22. L. elegans H.B.K. 1823, Nov.Gen.Sp.6:447. – L. campestris Schlecht. et Cham. 1830, Linnaea, 5, 5:89. – L. pulchellus Sweet, 1831, Brit. Fl. Gard. 2:67. – L. exaltatus Zucc. 1832, Abh. Acad. Wiss. (Munchen), 1:345. – L. moritzianus Kunth, 1852, Gartenflora :10. – L. californicus Hort. in C. Koch, 1858, Wochenschr. 1:80. – refined lupin.

The plant with silky pubescence and powerful stems is up to 100 cm in height. Leaflets are oblong-oval, almost naked from above and hirsute from below. Petioles are equal to leaves. Stipules on one-third grow together with petioles. The inflorescence is elongated, dense, having 6-10 verticils. The upper labium of the calyx is weakly bidentate, the lower is integral. Floral bracts are falling. The corolla varies from blue to violet, and is preserved at the time of pod formation. On the vexillum there is a yellow speck. Pods are 16-18 x 3 mm in size, 3-6-seeded. Seed are small-sized, grayish-brown with a maculation. 1000 seed weight is 20 g. Widespread in Mexico and Guatemala at an altitude of 1700-3000 m above sea level. Bears some resemblance to North American sp. L. polyphyllusLindl.

23. L. hartwegii Lindl. 1839, Bot. Reg. tab, 31. – L. bilineatus Benth. 1839, Pl. Hartw.:11.

The plant is roughly hirsute, erect. Branching is monopodial in the first phases of development. After formation of the apical inflorescence, the development of sympodial branches begins in the direction from the top downward. There are forms, in which the ascending and descending branches are interlocked. Stalks are powerful, more or less round in cross-section. The number of leaves on the main stem varies from 9 to 18. The leaf consists of 4-9 or, more often, 8 wide lanceolate leaflets pubescent only underneath. Their length is 4.5-5.5 cm, width 0.8-1.2 cm. Petioles are round-shaped, 9-13 cm long. Stipules are styliform, strongly consolidated and very long. Plants are closely hirsute, pubescence is long, hairy. Pubescence is especially strongly expressed on stipules, densely boarded by long upright hair. The inflorescence is long, polyanthous, with alternate arrangement of flowers, with feeble scent. Coloring of the corolla is quite diverse: blue, lilac, violet, etc. At the base of the vexillum there is a white spot that would turn pinker step by step with the aging of the flower. The wings are a little lighter than the sail, with darkly outlined nerves. The carina is white or very weakly colored, with a light or dark edge. The calyx is large, pubescent, emarginate down to the base, its upper labium is bidentate, the lower one is integral and longer than the upper. The floral shoot is long, drooping, and pubescent. Floral bracts are styliform, very large, twice longer than flowers, with hirsute pubescence, easily falling. Pods are small and flat. Seed are small-sized, with glossy surface, variable in color, with different ornamentation or single-colored. Widespread in Mexico and other countries of America. Cultivated in gardens as an ornamental plant. L. hartwegii is a very polymorphous species showing variability in the color of generative organs (flowers and seed), vegetative bodies (stems and leaves), and other morphological characters (branching habit, width of leaves, pubescence, etc.). Maisurian and Atabekova (Майсурян, Атабекова, 1974) identified seven varieties in this species on the basis of differences in the color of flowers and seeds.

24. L. subcarnosus Hook. 1836, Bot. Mag. tab. 3467. – L. bimaculatusHook. ex D. Don, in Sweet, 1835, Fl. Gard. Ser.2, tab. 314. – L. texensisHook. 1836, Bot. Mag. tab. 3492.

Rather dwarfish annual or biennial plant, up to 40 cm in height, with silky pubescence. Stems are prostrate and upstanding, thin, branching, with shortened sub-crown internodes. Leaflets are pubescent, oblong-oval, 3-9 in number. Petioles are short, 5-7 cm. Stipules are styliform, elongated, pubescent. The inflorescence is short and semi-verticillate. Flowers are set on long pubescent floral shoots. The calyx is small, pubescent, the lowerlabium is tridentate, the upper one is bidentate, with a deep notch. The corolla is dark-blue with a cream-white spot in the middle part of the sail. The corolla is colorless, with a dark edge. Pods are slightly bent, pubescent. Seed are depressed. (2n=36). Cultivated as an ornamental plant. Widespread in the USA (Texas).

25. L. pubescens Benth. 1845, Pl. Hartw.:169. – L. speciosus Hort. ex Vilm. 1844, Blimend, ed.3.

Plants are erect, strongly leafed, pubescent. Stems are powerful, branching. Leaflets are elongated, ovate-oblong, pubescent from both sides, 5-11 in number. Stipules are wedge-shaped, densely pubescent. Petioles are 7-9 cm long. The inflorescence is alternate or verticillate, polyanthous. The calyx is small, notched over more than one half. Floral bracts are easily falling. Flowers have feeble scent. The corolla is blue-violet with a white spot in the middle of the vexillum. The carina is colorless with a dark edge. Pods are hazel; easy shattered, strongly pubescent, 3-4 cm x 7-9 mm. Seed are small-sized, oval, weakly depressed, light-brown, frequently with a crescent gap. Widespread in Colombia. Naturalized in Uganda and Zaire. Cultivated as a horticultural plant in Europe and America.

26. L. nanus Dougl. ex Benth. 1835, Trans. Hort. Soc. 1:409. - L. sparsiflorus Torr. 1857, Pacif. Rail. Rep. 5:360. – L. bicolor A. Gray, 1857-61, Proc. Bost. Soc. Nat. Tlist. 7:146.

Plants are prostrate in the beginning and raised afterwards, strongly branching. Active development of lateral branches starts after the formation of the apical inflorescence. Stems are thin, up to 40 cm in height. Leaflets are linear-lanceolate, grooved, 6-8 (7) in number, weakly pubescent from above and stronger from below. Petioles are up to 7 cm long, twice longer than leaflets. Stipules are rather large, pubescent. The inflorescence is elongated, verticillate, polyanthous. The calyx has no bractlets, its upper labium is deeply bidentate, the lower being almost integral. Floral bracts are equal in size to flowers, with a falling habit. The corolla is dark-blue, blue, or white. The carina is white with a light or dark edge. Pods are linearly bent, light-yellow, easily opened, 6-10-seeded. (2n=48). Widespread in the USA (western California). Cultivated for ornamental purposes.

27. L. bicolor Lindl. 1827, Bot.Reg.:tab. 1109, non A.Gray, 1859-61. – L. micranthus var. bicolor Benth. 1843, Proc. Amer. Acad. Arts Sci.8:536. –L. trifidus Torr. ex Wats. 1877, Proc. Amer. Acad. Arts Sci.12:250. – L. umbellatus Greene, 1866, Bull. Calif. Acad. Sci. 2:145. – L. rostrarusEastw. 1896, Proc. Calif. Acad. Sci. 2,6:424. – L. plebeins Greene ex Baker, 1902, West. Amer. Pl. 1;11. – L. inanoemus Greene ex Baker, 1903, West. Amer. Pl. 2:14. – L. hirsutus Greene, 1911, Leafl. Bot. Obs. 2:152. –L. sabulosus Heller, 1911, Myhlenbergia, 7:9. – L. pipermsmithii Heller, 1911, Myhlenbergia, 7:93, fig. 16-17.

Plants are prostrate in the beginning and raised afterwards, pubescent with silvery hair. Leaflets are linear-lanceoliate, grooved, and pubescent on both sides; their number is 7-8. The calyx has a bilabiate upper labium and tridentate lower one, without bractlets. Floral shoots are almost equal to the calyx. The inflorescence is small, up to 6 cm long. The corolla is blue, longer than the calyx by one-third; the vexillum is yellowish in the middle. Pods are weakly pubescent, yellow, 6-seeded, 1.5 cm. x 4.4 mm in size. Seed are small-sized, 2.3-2.4 x 1.8-2.1 mm. 1000 seed weight is 3.4 g. Widespread in Canada (British Columbia), USA (northwestern California, western Oregon, western Washington). Cultivated for ornamental purposes.

28. L. truncatus Nutt. ex Hook. et. Arn. 1839, Bor. Beech. Vog.:336.

Plants are erect, more then 0.5 m high. Stems are reddish, pubescent rather sparsely with scanty short hair. Cotyledon leaves are non-falling. Usually, up to 8 leaves would be developed on the main stem. The leaf consists of 3-6 (7) very narrow linear grooved leaflets, 2.5-4.3 cm long and 2.0-4.0 mm wide. Length of petioles greatly varies depending on the location a leaf on the plant. The apical leaves are brachypetiolate, petioles of the lower leaves being more than twice longer than leaflets. Stipules are hardly noticeable, short, styliform, growing together with petioles. The upper part of leaflets is almost naked, the lower is sparsely pubescent. The inflorescence is short (7 cm), sparsely flowered, with 7 up to 17 flowers. The corolla is violet, blushing. The vexillum is broad, almost cordate, with a light spot and specks in the middle. The wings are oval, and also violet. The carina is light, with a crimson edge, broad, strongly pubescent on the internal edge, ciliated. Floral shoots are short. The calyx is bilabiate, its lower labium is integral, while the upper one is bipartite. Floral bracts are small, styliform, pubescent, green, preserved during flowering. Pods and seeds are small. 1000 seed weight is 3-5 g. Widespread in the countries of Central America.

29. L. polycarpus Greene, 1888, Pittonia, 1:171. – L. micranthus Dougl. ex Lindl. 1829, Bor. Reg. tab. 1251. – L. douglasii Guss. 1844, Fl. Sis. Syn. 2:852. – L. gracilis Durand et Hilg, 1858, Pacif. Rail. Rep. 5:7.

Plant prostrate in the beginning and raised afterwards, with silky pubescence. Leaflets are linear-lanceolate, grooved, almost naked from above and strongly hirsute from below. Petioles are twice longer than leaves, pubescent. Stipules are large, styliform; up to one half they grow together with petioles. The inflorescence is abnormally verticillate, sparsely flowered. Flowers are small. The corolla is blue and white; it is longer that the calyx only by one-third. Flowers are set on short floral shoots. The calyx is pubescent, its upper labium is emarginate, the lower one is integral. Bractlets are bristly. Pods are narrow-linear, beige, strongly pubescent, 7-9 seeded, 3 x 0.5 cm in size. Seed are small, gray-beige, with a brown strip and speckles, 2.2-3.0 x 1.7-2.3 mm. 1000 seed weight is about 6 g. Widespread in Canada (British Columbia), USA (California, Oregon, and Washington).

30. L. succulentus Dougl. ex C. Koch, 1867, Index Sem. Hort. Bot. Berol. App. 1:11.

Scarcely foliated, pubescent plant. Stems reach 40 cm in hight. Branching on the main stem proceeds sequentially from the bottom upward. Inflorescences are developed after the formation of 8-11 leaves. Leaflets are obovate, 6-7 in number. Petioles are 5-7 cm long. Stipules are lanceolate, small, pubescent or nearly naked. The inflorescence is ordinary looking, sparsely flowered, with an alternate arrangement of flowers. The upper labium of the calyx is equal in length to 2/3 of the lower, with anthocyan. Floral bracts are styliform, and easily falling. The corolla is blue-violet, turning orange-red in the process of aging. The pubescent carina has a dark-violet beak. Pods are yellow, weakly pubescent, 9-seeded. Seed are 4-5 x 3-4 mm in size, brown-gray with a darker pattern, or dark-violet. 1000 seed weight is 26-30 g. It is one of the earliest species of lupin. Widespread in Mexico and the USA (California, Arizona).

31. L. stiversii Kellogg, 1863, Proc. Calif. Acad. Sci. 2:192, fig. 58.

The plant is dwarfish. Stems are red-brown, leaves are green, petioles and stipules are reddish. Leaflets are obovate, without pubescence; their number is 6-7. Petioles are equal in size to leaves. The vexillum is egg-shaped, with speckles. The wings are oval, crimson-pink with a darker spot in the base. The carina has a pink-orange edge. Floral bracts are lanceolate, pubescent. Pods are yellow-brown, without pubescence, polyspermous; usually the corolla is retained. Seed are spherical- rhombic, with a spear, gray-brown in color, with marbling pattern and a light spear. Mostly, it is an ornamental plant. Widespread in the USA (California).

32. L. microcarpus Sims, 1823, Bot. Mag. tab. 2413. – L. densiflorusAgardh, 1835, Syn. Gen. Lupin.:3. – L. menziessi Torr. 1858, Mex. Bound:57. – L. lacteus Kellog, 1875, Proc. Calif. Acad. Sci. 5:37. – L. palustris Kellog, 1875, l. c. :36.

Plants are low-level, branched in the basis, pubescent. Leaflets are obovate-oblong, slightly curled, naked from above and pubescent from below. Petioles are 2-3 times longer than leaves. Floral bracts are styliform, up to 10 mm long, up to 1/2-1/3 they grow together with petioles. The inflorescence is set on a very long flower pedicle, with 7-11 verticils. Flowers are up to 17 mm long, set on short pubescent floral shoots. The upper labium of the calyx is chaffy, bidentate, and thrice shorter than the lower, which is coriaceous. The corolla is violet. The vexilum is wider than the wings. The carina is lilac, narrow, with equal wings having a violet edge, pubescent. Floral bracts are longer than the upper labium of the calyx, styliform, pubescent. Pods are small, down to 20 x 12 mm., 1-2 seeded, bright-yellow, silver-silky. Seeds are spherically domed, up to 5 mm in diameter, brown with speckles, and striped. Widespread in Argentina and Chile.

33. L. pusillus Pursh, 1814, Am. Sept. 2:468. – L. rubens Rydb. 1907, Bull. Torr. Club, 34:45. – L. intermontanus Heller, 1912, Mylhenbergia, 8: 87.

The plant is small, tiny, scleroid, and hirsute. Stems are strongly branched from below; the style of branching is horizontal. Leaflets are elongated, blunted at the top. and reduced from top to bottom, naked from above and pubescent from below; their number is 5. Petioles are longer than leaves. Stipules almost entirely grow together with petioles. The inflorescence is friable, with alternate arrangement of flowers. Floral bracts are egg-shaped, equal to the tube of the calyx. The upper labium is almost bipartite, the lower one is vaguely tridentate (the middle finger being longer). Bractlets are missing. The corolla is blue, and twice longer than the calyx. Pods are almost rhombic, 2-seeded. Seed are squeezed, light-colored. Widespread in Canada and the USA (Arizona, California, Colorado, Idaho, western Kansas, Montana, western Nebraska, Nevada, New Mexico, western Dakota, Washington, and Wyoming).

34. L. densiflorus Benth. 1835, Trans. Hort. Soc. London, Ser. 2,1:410. –L. menziesii Agardh, 1835, Syn. Gen. Lupin. :2. – L. tracyi Eastw. 1940, Leafl. West. Bot. 2: 268.

Plants are slightly fluffy, covered with hair. The stem is 40-60 cm high. The leaf consists of 9-11 oblong-lanceolate leaflets. The inflorescence is 15-20 cm lengthwise, compactly verticillate, and odorless. Floral bracts are non-falling. Flowers are sulphur-yellow, but later they would turn a little bit orange and ultimately up to dirty-brown. The vexillum has expressly bent back edges. The ovary contains two seedbuds. Pods are short, egg-shaped, flat-fruited, with two seeds. Seed are light- waxy-yellow with brown marbling. Widespread in North America. Grown as a flower-bearing plant in the USA (California).
L. densiflorus Benth. is selected as the type of subgenus Platocarpos(Wats) Kurl.

We have presented the description of only those 34 species of lupin from the New World that are available in the collection of the Vavilov Institute and have been studied thoroughly enough. Lupins from American continent have identical chromosome number (48) and they are easily crossed among themselves. This phenomenon opens ample opportunities (prospects) for them morphogenesis. There are a huge number of the various hybrid forms in the nature described by different authors as independent species (up to 1000) in this connection. Real number of existing lupin species in America practically it is not possible to establish in this connection. ,,,,,





Description of the Old World ‘s lupins and their intraspecific diversity
(Subgen. Lupinus)

Key to the Mediterranean and African Species (Gladstones, 1974,1998)

1. Seeds ± smooth. 2. Lower flowers of inflorescence alternate.
3. Upper lip of calyx entire; seeds ± square, compressed; flowers white to violet- blue. ..........1. L. albus L.

3. Upper lip of calyx deeply 2-partite; flowers normally blue.
4. Leaflets linear; lower lip of calyx entire to slightly 3-toothed; seeds round; seeds sparsely sericeous. ........................................2. L. angustifolins L.

4. Leaflets short and broad; lower lip of calyx deeply 3-toothed; seeds lenticular or compressed, brownish; plant coarsely hirsute. ............................3. L. micranthus Guss.

2. Flowers fully verticillate.
5. Flowers golden yellow, scented; upper surface of leaflets villous. ..........4. L. luteus L.

5. Flowers cream to mauve, not scented; upper surface of leaflets ± glabrous. ............................................................................5. L. hispanicus Boiss. and Reut.

1. Seeds rough; flowers subverticillate to verticillate, usually blue.
6. Hairs on stems < or =" 1" or =" 1/3"> 1 mm; lower lip of calyx entire. 7. Lower lip of calyx shallowly 3-toothed; tip of keep blunt, pigmented blue; seeds about 8х6х3 mm, brown or grayish with blackish markings                                                                  …………6. L. cosentinii Guss.

7. Lower lip of calyx ± entire; tip of keel pointed, not pigmented blue. 8. Pods 9-12 mm broad; seeds about 7х6х3 mm, mottled reddish brown with a smooth hump over the hilum………7. L. digitatus Forsk.

8. Pods 16-18 mm broad; seeds about 10 x 8 x 4 mm, mottled reddish brown on pale yellow. ………..
                                                                              ...................8. L. princei Harms

6. Hairs on stem > 1 mm; lower lip of calyx entire. 9. Hairs on stem 3-4 mm; pedicel > or = 1/2 length of calyx, lax.
10. Flowers very large, deep blue (rarely pink) with a white band up the centre of the standard; leaflets softly villous; seeds 10-14 x 9-12 x 6- 8 mm, mottled brownish red; plants erect. ......9. L. pilosus Murr.

10. Flowers pale yellow to pale pink, tinged with blue; leaflets sericeous; seeds slightly smaller than 9, mottled brown; plants rosetted.                                                 ..........10. L. palaestinus Boiss.

9. Hairs on stem 1-2 mm; pedicel 1/2length of calyx, rigid.

11. Leaflets sericeous; pedicel; standard with a central white band, broadening towards the upper margin; seeds about 8-11 x 6-8 x 5 mm, mottled brown to pinkish brown with a smooth hump over the hilum.
                                                                                          .....11. L. atlanticus Gladstones.
12. Upper surface of leaftlets glabrous; pedicel about 1/2 length of calyx; rounded white spot on standard, not approaching the upper margin. Highlands of Somaliland, possibly Ethiopia.
                                                                                              .....12. L. somaliensis Baker


Description of individual species.
1. L.albus L. 1753, Sp. Pl.:721; Willd. 1803, Sp.Pl. ed. 4, 3:1022; DC. 1825, Prodr. 2:407; Boiss. 1872, Fl. Or. 2:29; Willk. at Lange,1880, Prodr. Fl. Hisp.3:466; Halacsy, 1901, Consp. Fl. Graec. 1:341; Fiori, 1925, Nuov. Fl. Anal. Ital. 1:804; Zhukovsky, 1929, Bull.Appl. Bot. Gen. Pl-Breed., 21, 1:270, figs.13, 13a; Franco et Silva, 1968, in Fl. Europ. 2:105; Chamberlain in Davis, 1970, Fl. Turkey 3:38; Zohary, 1972, Fl. Palaest. 2:42; Gladstones, 1974, W. Austral. Dep. Agr. Techn. Bull. 26:5. – L. varius Gaertn. 1791, Fl. Fr. 2 : 324, tab.150, fig. 4, non L. 1753. – white lupin.

Typus: Herb. Linn. № 898 - 2 (LINN).

Annual, more or less pubescent plants, 30 - 120 cm high. Stems and petioles are sparsely sericeous. Stipules are subulate, concrescent with the petioles over 1/3 of their length. Leaflets (5 – 11) are oblong or obovate, glabrous above and villous below, margins ciliate. Their size is 20 - 60 x 10 - 20 mm. Colour of the corolla is white, grayish and light blue, less frequently pink blue, dark blue or violet blue. Lower flowers of the inflorescence are alternate. The lower lip of the calyx is lobed with 3 teeth, the upper one is entire. Pods are 70-160 x 10-20 mm, 3 - 6 seeded, seeds are square, compressed, white with a variable tinge of salmon pink, or dotted, dark brown. 2n = 50. Widely spread as wild plants throughout Greece including islands, in Albania, on the islands of Sicily, Corsica and Sardinia, as well as in Israel, Palestine and Western Turkey (European and Western part of Asia Minor). Occurs in meadows, pastures, and grassy slopes, predominantly on sandy and acid soils. Cultivated over all the Mediterranean region and also in Egypt, Sudan, Ethiopia, Syria, Central and Western Europe, USA and South America, Tropical and Southern Africa, Russia, and Ukraine. The ancient culture of white lupin under the local name «hanchcoly» until recently existed in Western Georgia (Kurlovich, 1996). Easily turns wild, especially in the Mediterranean region.Lupinus albus L. has been selected as a type of genus Lupinus L. The closest described neighbors of this species are L.graecus Boiss. et Sprun. and L. termis Forsk. The first is the wild plant on Balkan Peninsula; the second is grown in Egypt, Libya, Sudan, Ethiopia, Israel, Palestine and Syria. Information accumulated by scientists made it possible to revise and integrate the volumes of these species. All three species differ only in the colour and size of flowers and seeds (quantitative characters). Cultivated forms of white lupin were introduced in agricultural practice long time ago, before the ancient times, on the place of present-time Greece (Gladstones, 1974; Майсурян, Атабекова, 1974). From Greece they spread throughout the Mediterranean region and elsewhere. To the south (Egypt, Libya, Palestine) mainly the forms with white seeds and pink-and-blue or light-pink flowers (L. termis) were spread, while westwards (Apennine Peninsula and farther) mostly the forms with white seeds and grayish-blue or white seeds (L.albus) dispersed. White lupin (L.albus) is still grown in Greece, where wild L.graecus is also spread. The absence of distinct specific characters makes it possible to consider all three species harbored within the limits of one species of Linnaeus (L.albus L). But the existence of geographical and physiological differentiation is sufficient to recognize them as having the rank of subspecies in L.albus L: subsp.albus, subsp. graecus (Boiss. et Sprun.) Franco et Silva and subsp. termis(Forsk.) Ponert. Small species such as L. vavilovii Atab. et Maiss. and L.jugoslavicus Kazim. et Nowacki, recognized by some plant growers of the former USSR and Poland, are only nomenclature synonyms of Balkan subspecies graecus.

The scheme of classification of Lupinus albus L.

I. Subsp. graecus (Boiss. et Spun.) Franko et Silva
2. Subsp. termis (Forsk.) Ponert.
I. Var. abissinicus Libk.
2. Var. subroseus Libk.
3. Subsp. albus L.
3. Var. albus 
4. Var.vavilovii (Atab.) Kurl. et Stanke
v. 5. Var vulgaris Libk.
I. f. libkindae Kurl. et Stankev.
Key to the subspecies of Lupinus albus L.

1. Rosettes are formed initially from the leaves, stalks appear at the time of flowering. Corolla is dark-blue with a white centrobasal spot at the base of the vexillum. Pods are 60-80 x 11-14 mm, shattering at maturity; seed are 7-10 x 6-8 x 2-3 mm, dark brown, dotted, with impermeable testa. Native to Balkan Peninsula …………1. Subsp. graecus

+ Rosette is lacking. The colour of the corolla is pink, blue, grayish, light blue or white. Pods are 80-150 x 16-20 mm in size, indehiscent at maturity; seed are pinkish white or white with permeable testa …………………. 2.

2. Plants are shaggy, sericeous. The inflorescence is stretched, crumbly.
Flowers are pink or blue. The calyx is with a bract. Found in Egypt, Libya, Sudan, Ethiopia, Israel …………..................................… 2. Subsp. termis

+ Plants are softly villous. The inflorescence is short, more or less compact. Flowers are grayish, light blue or white. The calyx is without bract (it falls quickly). Cultivated in the Mediterranean area, in Europe, North and South America, Asia .. ……………………...………………… 3. Subsp.albus

..1. Subsp. graecus (Boiss. et Sprun.) Franco et Silva, 1968, Feddes Repert. 79 :52; idem, 1968, in Fl. Europ. 2 : 105; Chamberlain in Davis, 1970, Fl. Turk. 3 :39. - L.graecus Boiss. et Sprun. 1843, Diagn. pl. or. nov. 2:15; Halacsy, 1901, l. c.:341. - L.jugoslavicus Kasimierski et Nowacki, 1961, Gen. Polon. 2:115. - L.vavilovii Atab. et Maiss. 1962, Изв. ТСХА, 44 (1):239. - L.albus var. graecus (Boiss. et Sprun.) Gladstones, 1974, l. c.: 7.

Typpus: In cultis Graeciae, inter Spartam et Mistra, E. Boisser et W. Spruner, April 1842 (G).

Two- to four-year plant in the nature and annual in culture. It is a direct wild ancestor of white lupin. This subspecies is spread in Greece, on Crete and other Aegean islands, in Albania and western Turkey (Gladstones, 1974) and can also be found in Italy (Hammer et al., 1992). It differs from other subspecies by having small diversity of forms.

..2. Subsp. termis (Forsk.) Ponert, 1973, Feddes Rep. 83 (9 - 10) :619. - L. termis Forsk, 1775, Fl. Aegypt:131; Willd. 1803, l.c.:1023; Guss. 1828, Fl. Sic. Prodr. 2:397; Bertol. 1847, Fl. Ital. 7:411; Boiss. 1872, Fl. Or. 2:29; Post. 1896, Fl. Syr. Pal. Sin.:213; Halacsy, 1901, Fl. Graec. 1:341; Plitmann, 1966, Isr. Journ. Bot. 15:26. - L.albusvar. termis (Forsk). Fiori, 1925; Nouv. Fl. Anal. Ital. 1:804; Hanelt, 1960, Lupinen 10, fig. 4, 12.Typus : In Delta ad pagum Nedjel. Copiose in agris, an spontaneus, nesсio ( C? ).

Subsp. termis differs from the standard subspecies (subsp. albus) by earlier flowering, finer stems and smaller flowers and seed. The flowers have a different colour. The name of this subspecies takes place from the Greek term «thermos». In the Mediterranean countries it is usually called «turmus» (or alternative versions). Spread and cultivated in Egypt, Libya, Sudan, Ethiopia and Israel. Easily turns wild. We have at our disposal accessions from Libya. Egypt and Ethiopia. Libkind (Либкинд, 1931) described it as varietas but she did not indicate any types.

1. Var. abissinicus Libk. 1931, Lupin: 59, sine typo. - Flowers pink-and-blue. Vegetative parts with anthocyan. Neotypus: `B -5`, received from Belarus, k-2237, reproduction of Pushkin`s laboratories of VIR, 28.07.1989, L.T. Kartuzova (WIR). Distribution - the upper reaches of the Nile.
2. Var. subroseus Libk. 1931, l. c.: 59, sine typo. - Flowers light -pink. Vegetative parts green. Neotypus: Libya, k-2094, reproduction of Pushkin`s laboratories of VIR, 28.08.1989, L.T. Kartuzova (WIR). Distribution - Libya, Egypt, Israel, Syria.
3. Subsp. albus. Distinctive features are thick meaty stalks, large leafs and seeds. The genepool of subsp. albus has wide variations in physiological properties of plants. Cultivated in many countries of Europe, Asia and America.
3. Var. albus Kurl. et Stankev. 1990, Bull. Appl. Bot. Gen. Pl.-Breed., Leningrad, 135:33. - Flowers white, carina`s edge without anthocyan. We put the accession `Mutant 47` down to this varietas.
4. Var. vavilovii Kurl.et Stankev. 1990, l. c.:33. - Flowers white, carina`s edge with anthocyan. Typus: `Bialy - 1`, Poland, k-1602, reproduction of Pushkin`s laboratories of VIR, 9.08.1986, A.K. Stankevich (WIR).
5. Var. vulgaris Libk. 1931, l. c.: 59, sine typo. - Flowers grayish-blue, carina`s edge with anthocyan. Neotypes: `Kievsky mutant`, Ukraine, k-1904, reproduction of Pushkin`s laboratories of VIR, 9.08.1989, B.S. Kurlovich (WIR). This varietas is the most spread.
1. f. libkindae Kurl. et Stankev. 1990, l. c.:33. - Side shoots absent or shortened. Flowers axillary. Typus: `EP-1`, Poland, k-2890, reproduction of Pushkin`s laboratories of VIR, 24.08.1989, B.S. Kurlovich (WIR).

...2. L.angustifolius L. 1753, Sp. Pl:721; Willd. 1803, l. c.:1024; DC. 1825, l. c.:407; Boiss. 1872, l. c.:28; Willk. et Lange, 1880, l. c.:466; Halacsy, 1901, l. c.:340; Aschers. et Graebn. 1907, Syn. Mitteleur. Fl. 6 (2):231; Fiori, 1925, l. c.:804; Plitmann, 1966, Israel J. Bot. 15:26; Chamberlain in Davis, 1970, l. c.:39; Zohary, 1972, l. c.:43, t. 57; Gladstones, 1974, l. c.:9; Vass. 1987, in Fl. Part. Eur. URSS, 6:214. –L. linifolius Roth, 1787, Bot. Abh. 14, t. 5. - L.reticulatus Desv. 1835, Ann. Sc. Nat. ser. 2,3:100. - L. leucospermus Boiss. 1849, Diagn. Pl. Or. Nov. 9:8. - L. philistaeus Boiss. 1849, l.c.: 9. – L. angustifolius var. sativus et var. spontaneus Libk. 1931, Lupinus :53. - L.opsianthus Atab. et Maiss., 1968, Bjull. Glav. Bot. Sad. Acad. Nauk SSSR. : 75. - L.angustifolius subsp. angustifolius et subsp.reticulatus (Desv). Franco et Silva, 1968, in Fl. Europ. 2:105. – narrow-leafed or blue lupin.
Typus: Herb. Linn. No 898-7 (LINN).

Plants are 20-150 cm high. Stalks are sparsely sericeous, with numerous lateral branches. Leaves are developed from 5-9 linear-lanceolate or narrow-linear leaflets, 20-40 х 2,5 mm in size, the upper surface is glabrous, the lower surface is sericeous. Stipules are linear and linear-lanceolate. The inflorescence is dense, almost sedentary, 5-20 cm long, placed on a short floral pedicle. Flowers are 11-15 mm long, almost sedentary, lower flowers are alternate, upper ones more or less subverticilate inside. Floral bracts are small-sized, easily falling. The calyx is bilabiate, profoundly bipartite, the lower lip is longer, entire or irregularly two- or three-toothed. The corolla is blue, violet, less frequently pink and white. Pods are 35-50 х 7-10 mm in size, oblong, slightly inflated, 4- or 7-seeded, with oblique partitions between seeds. The seed is 4-8 х 3-7 х 3-6 mm, globular, with smooth testa, variously coloured, with a triangular spot and a stria close to the hilum. 2n = 40.
Occurs on meadows, among rocks, in bushes, on seaside sands and near reservoirs, along the roads and, as a weed, in the field. Tends to grow on well-ventilated soils. Widespread in all Mediterranean countries and also in Asia Minor, Transcaucasia and Iran. Naturalized in Southern Africa and Southwest Australia. Widely cultivated in Northern Europe (Germany, Holland, Poland, CIS), in the southeast part of USA and in New Zealand. It is a rather polymorphic species. The variability of characters depends on eco-geographic conditions. Wild-growing races have, as a rule, narrow pods, smaller-sized leaflets and seeds as well as lower plant height. The size of seeds and their colouring vary on all area of growing. Large-seeded plants usually occur closer to arable soils, while small-seeded ones on inshore sands and slopes. The species is described using a large cultivated form. The colouring of seeds varies from dark grey (almost black) with light spots and specks of miscellaneous size, grayish brown and brown up to light grey and white. Usually seeds with coloured testa correlate with cyan and pink flowers. In these two characters (traits) homologous variability is observed. Unpigmented (white) seeds are typical for white-flowered and lilac plants. These combinations of characters do not demonstrate any geographic arrangement. There is no abrupt gap between cultivated and wild forms. On the contrary, transition between them appears very smooth. Therefore there is nothing to justify the existence of two species (L. linifolius Roth. and L. opsianthus Atab. et Mais.) and also two subspecies (subsp. angustifolius and subsp. reticulatus(Desv.) Franko et Silva.) differing only on in the width of leaflets and the size of seeds (quantitative characters). In our opinion, it would be more correct to subdivide this species in a different way. Using the developed criteria of intraspecific taxa, the spreading of VIR’s blue lupin collection disclosed 13 varieties of L. angustifolius L. distinctly differing from each other by the colour of seeds and the corolla. Variations in the colour of cotyledons, vegetative parts and carina have been used for identification of 12 subvarieties. Besides, the plants with determined branching and fascicular stems are described as 8 separate forms.
The scheme of classification of Lupinus angustifolius L.

I. Var. angustifolius 
I. Subvar. angustifolius
1. f. angustifolius
2. Subvar. viridulus Kurl. et Stankev.
2. Var. albopunctatus Kurl. et Stankev.
3. Var. griseomaculatus Kurl. et Stankev.
2. f. belorussicus Kurl. et Stankev.
4. Var. chalybens Kurl. et Stankev.
5. Var. corylinus Kurl. et Stankev.
3. f. zhukovskii Kurl. et Stankev.
6. Var. purpureus Kurl. et Stankev.
3. Subvar. purpureus 
4. f. deramosus Taran. et Busch.
4. Subvar. rhodanthus Kurl. et Stankev.
7. var. rubidus Kurl. et Stankev.
5. Subvar. rubidus 
6. Subvar. maissuriani (Atab.) Kurl. et Stankev.
5. f. epigonalus Taran. et Busch.
8. Var. atabekovae Kurl. et Stankev.
9. Var. sparsiusculus Kurl. et Stanke
v. 10. Var. brunneus Kurl. et Stankev.
11. Var. albosyringeus Taran.
7. Subvar. albisyringeus
6. f. kloczkovii Kurl. et Stankev.
7. f. confertus Kloczko et Kurl.
8. Subvar. polonicus Kurl. et Stankev.
9. Subvar. lilacinus Kurl. et Stankev.
12. Var. albidus Kurl. et Stankev.
8. f. kuptzovii Kurl. et Stankev.
13. Var. candidus Kuptzov. et Kurl.
10. Subvar. candidus
11. Subvar. virescens Kuptzov. et Kurl.
12. Subvar. violaceus Kuptzov. et Kurl.




1. Var. angustifolius, 1990. Bull. Appl. Bot. Gen. Pl.-Breed., Leningrad, 135:23. - L.angustifolius var. coeruleus Asch. et Graebn. ex Atab. 1962, Lupinus: 129, nom. illeg. - Flowers blue. Seed motley, gray with unclear spotting.
1. Subvar. angustifolius. - Cotyledons and the carina`s edge anthocyan - coloured. We do not know, what kind of seed had C. Linnaeus in the disposal, but we think, that most likely described by him accessions was represented by domesticated example. It is possible to consider cv. Benyakonsky 484 (k-1457) from collection of VIR as typical accessions of this taxon.
1. f. angustifolius. - Side shoots absent or shortened, flowers axillary. Typus: ‘Mut-1’, Poland, k-2803, reproduction of Pushkin`s laboratories of VIR, 12.7.1989, B.S. Kurlovich (WIR).
2. Subvar. viridulus Kurl. et Stankev. 1990, l.c.:24. - L.angustifolius var.subcoeruleus Sypn. ex Atab. 1962, l. c.:129, nom. illeg. - Cotyledons and the carina`s edge green. Typus: Poland, k-334, reproduction of Pushkin`s laboratories of VIR, 14.7.1989, B.S. Kurlovich (WIR).
2. Var. albopunctatus Kurl. et Stankev. 1990, l.c.:24. - L.angustifolius var.maculatus Atab. l. c.:129, nom. illeg. - Flowers blue. Seed almost black with tiny white dots and spots. Typus: ‘Benyakonsky 335 ’, Belarus, k-1477, reproduction of Pushkin`s laboratories of VIR, 10.7.1989, B.S. Kurlovich (WIR).
3. Var. griseomaculatus Kurl. et Stankev. 1990, l. c.:24. - L.angustifoliusvar. subalbidus Sypn. ex Atab. 1962, l. c.:129, nom. illeg. - Flowers blue. Seed gray with white spots. Typus: Belarus, k-3148, reproduction of Pushkin`s laboratories of VIR, 14.8.1989, B.S. Kurlovich (WIR).
2. f. belorussicus Kurl. et Stankev. 1990, l. c.:24. - Side shoots shortened, flowers axillary. Typus: ‘Першацвет’, Belarus, k-2955, reproduction of Pushkin`s laboratories of VIR, 7.7.1989, B.S. Kurlovich (WIR).
4. Var. chalybeus Kurl. et Stankev. 1990, l. c.:24. - Flowers blue. Seed white with sparse dark-brown and gray spots. Typus: ‘Frost’, USA, k-2258, reproduction of Pushkin`s laboratories of VIR, 21.7.1989, B.S. Kurlovich (WIR).
5. Var. corylinus Kurl. et Stankev. 1990, l. c.:24. - Flowers blue. Seed beige with browm spots. Typus: ‘Apendrilon’, Greece, k-2666, reproduction of Pushkin`s laboratories of VIR, 20.7.1989, B.S. Kurlovich (WIR).
3. f. zhukovskii Kurl. et Stankev. 1990, l. c.:24. - Side shoots absent or shortened, flowers axillary. Typus: ‘Line AT - Д-2-Т/5’, Belarus, k-2837, reproduction of Pushkin`s laboratories of VIR, 14.8.1986, A.K. Stankevich (WIR).
6. Var. purpureus Kurl. et Stankev. 1990, l. c.:24. - L.angustifolius var. roseus Semp. ex Atab. 1962. l.c.: 129, nom. illeg. - Flowers pink. Seed motley or gray with unclear spotting. Is close to var. angustifolius (1).
3. Subvar. purpureus - Cotyledons anthocyan - coloured, vegetative parts dark-violet. Typus: ‘Borre’, Sweden, k-1593, reproduction of Pushkin`s laboratories of VIR, 21.7.1989, B.S. Kurlovich (WIR).
4. f. deramosus Taran. et Busch. f.nova - Rami laterallii nullum esse vel valde abbreviati. Flores axillaria. Typus: Belarus, k- 3527, reproduction of Pushkin`s laboratories of VIR, 14.8.2001, L.T. Kartuzova (WIR). Side shoots absent or shortened, flowers axillary.
4. Subvar. rhodanthus Kurl. et Stankev. 1990, l. c.:25. - Cotyledons and vegetative parts green. Typus: Poland, k-1341, reproduction of Pushkin`s laboratories of VIR, 19.8.1989, B.S. Kurlovich (WIR).
7. Var. rubidus Kurl. et Stankev. 1990, l. c.:25. - L.angustifolius var. lastowski Atab. 1962, l. c.:129, nom. illeg. - Flowers pink. Seed almost black with tiny white dots and spots. It is close to var.albopunctatus (2).
5. Subvar. rubidus. - Cotyledons anthocyan-coloured, vegetative parts dark-green. Typus: ‘Вада-18’, Belarus, k –2679, reproduction of Pushkin`s laboratories of VIR, 18.7.1989, B.S. Kurlovich (WIR).
6. Subvar. maissurianii Kurl. et Stankev. 1990, l. c.:25. – L. angustifolius var. maissurianii Atab. 1962, l. c.:129, nom. illeg. - Cotyledons and vegetative parts anthocyan-coloured. Typus: ‘Northern - 3’, Moscow province, k- 1712, reproduction of Pushkin`s laboratories of VIR, 17.7.1989, B.S. Kurlovich (WIR).
5. f. epigonalus Taran. et Busch. f.nova - Rami laterallii nullum esse vel valde abbreviati. Flores axillaria. Typus: Belarus, k- 3528, reproduction of Pushkin`s laboratories of VIR, 14.8.2001, L.T. Kartuzova (WIR). Side shoots absent or shortened, flowers axillary.
8. Var. atabekovae Kurl. et Stankev. 1990, l. c.:25. - Flowers pink. Seed with white spots. It is close to var.griseomaculatus (3). Typus: ‘Гюльцовский’, Germany, k -1498, reproduction of Pushkin`s laboratories of VIR, 28.7.1989, B.S. Kurlovich (WIR).
9. Var. sparsiusculus Kurl. et Stankev. 1990, l. c.:25. - Flowers pink. Seed white with sparse dark-brown and gray spots. It is close to var. chalybeus (4). Typus: ‘Лаф-рбс/2’, Belarus, k-2831, reproduction of Pushkin`s laboratories of VIR, 20.7.1989, B.S. Kurlovich (WIR).
10. Var. brunneus Kurl. et Stankev. 1990, l. c.:25. - Flowers pink. Seed beige with brown spots. It is close to var. corylinus (5). Typus: ‘Mirela’, Poland, k-2570, reproduction of Pushkin`s laboratories of VIR, 5.7.1989, B.S. Kurlovich (WIR).
11. Var. albosyringeus Taran. ex Kurl. et Stankev. 1990, l. c.:26. - Flowers pale-violet. Seed white, dull at the scar, without a triangular spot or strip.
7. Subvar. albosyringeus. - Cotyledons without anthocyan, vegetative parts dark-green. Typus: ‘ Немчиновский 846 ’, Moscow province, k-1981, reproduction of Pushkin`s laboratories of VIR, 6.7.1989, B.S. Kurlovich (WIR).
6. f. kloczkovii Kurl. et Stankev. 1990, l. c.:26. - Side shoots absent or shortened, flowers axillary. Typus: ‘Ладный’, Moscow province, k-2648, reproduction of Pushkin`s laboratories of VIR, 2.8. 1986, A.K. Stankevich (WIR).
7. f. confertus Kloczko et Kurl. 1990, Bull. VIR, 206:82. - Stem fascicular. Flowers, pods and side shoots clustered at the top of the stem. Typus: ‘5M2-253’, ТСХА, k - 2979, reproduction of Timiryazev Agricultural Academy in Moscow, 10.7.1990, N.A. Klochko and N.F. Anikeeva (WIR).
8. Subvar. polonicus Kurl. et Stankev. 1990, Bull. Appl. Bot. Gen. Pl.-Breed., Leningrad, 135:26. - Cotyledons and vegetative parts anthocyan – coloured. Typus: ‘ Muzin bialy ’, Poland, k-1851, reproduction of Pushkin`s laboratories of VIR, 25.7. 1989, B.S. Kurlovich (WIR).
9. Subvar. lilacinus Kurl. et Stankev. 1990, l. c.:26. - Cotyledons and vegetative parts light-green. Typus: ‘Тимир-1’, Moscow province, k-2664, reproduction of Pushkin`s laboratories of VIR, 20.7. 1989, , B.S. Kurlovich (WIR).
12. Var. albidus Kurl. et Stankev. 1990, l.c.:26. - L.angustifolius var. leucanthus Harz. ex Atab. 1962, l. c.:129, nom. illeg. - Flowers white. Seed white with sparse brown spots. Typus: ‘Unicrop’, Australia, k-2096, reproduction of Pushkin`s laboratories of VIR, 5. 8. 1989, B.S. Kurlovich (WIR).
8. f. kuptzovii Kurl. et Stankev. 1990, l. c.:26. - Side shoots absent or shortened, flowers axillary. Typus: ‘Ланедекс-1’, Belarus, k-2687, reproduction of Pushkin`s laboratories of VIR, 3. 7. 1989, B.S. Kurlovich (WIR).
13. Var. candidus Kuptzov et Kurl. 1990, Bull. Appl. Bot. Gen. Pl.-Breed., Leningrad, 135:26. - L.angustifolius var. albescens Atab. 1962, l. c.:129, nom. illeg. - Flowers white. Seed sheen-white, glossy.
10. Subvar. candidus. - Cotyledons and and vegetative parts without anthocyan. Typus: ‘ЗЛНР-2’, Belarus, k-3184, reproduction of Pushkin`s laboratories of VIR, 19. 7. 1989, B.S. Kurlovich (WIR).
11. Subvar. virescens Kuptzov et Kurl. 1990, l. c.:26. - Cotyledons anthocyan –coloured. Vegetative parts without anthocyan. Typus: ‘DM - green’, Belarus, k-2972, reproduction of Pushkin`s laboratories of VIR, 19. 7. 1989, B.S. Kurlovich (WIR).
12. Subvar. violaceus Kuptzov et Kurl. 1990, l. c.:27. - Cotyledons and vegetative parts anthocyan - coloured. Typus: ‘DM - anthocyaned’, Belarus, k-2971, reproduction of Pushkin`s laboratories of VIR, 19. 7. 1989, B.S. Kurlovich (WIR).
The quantity of new intraspecific taxa of Lupinus angustigolius L., and specially its subvarietas and forms, can be greater in the process of further investigation of its intraspecific diversity.

3. L. micranthus Guss. 1828, Fl. Sic. Prodr. 2:440; Franco et Silva. 1968, l. c.:105; Chamberlain in Davis, 1970, 1. c.:39; Zohary, 1972, l. c.:43. - L.hirsutus L. Sp. Pl. ed. 2:1015 non ed.1 1753; Aschers. et Graebn. 1907, l. c. :226; Zhukovsky, 1929, l. c.:265: Gladsones, 1974, l. c.:14-15. Typus: Herb. Linn. No 898-5 and 898-6 labeledL.hitsutus, and No 898-4 labeled L. varius. (LINN).
Plants are 10-50 sm high, resettled in the beginning and then becoming erect, with basal branching. Stalks, petioles and leaflets are hirsute. Stipules are linear-subulate, on 2/3 grow together with petioles. Leaflets are obovate-wedge-shaped or oblong. Inflorescence is 3-12 sm long, on short flower peckers, lower flowers alternate, upper more or less vercicilate. Floral bracts are styliform, not falling. Bractlets are linear. The upper labium of a calyx short, is deep 2-divided, lower is twice longer, is deep 3—lobed. Corolla is strongly varies on size, from short and barely exceeding the calyx up to twice longer it, blue with a white spot on middle of vexillum. The vexillum is rather small and angled at about 450 to the wings. Carina is white below, and blue above. Beans 30-50 х 9-12 mm, brownish, coarsely hirsute, 2-5 seeded. Seeds 5-8 х 4-6 х 3-4 mm, lentiform, slightly square, squeezed, smooth, hazel with one more dark marbling, sometimes with dark brown arcs around hilum. 2n=50.? Grows on coast, frequently on limier soils than L.angustifolius. It is widespread on all Mediterranean. This species is described by Linnaeus and by many other writers as L.hirsutus and L. varius. The titles of Linnaeus were changed in order to prevent tangle with other species.


4. L.luteus L. 1753, Sp. Pl. :722; Willd. 1803, l. c.:1024; DC. 1825, l.c.:407; Willk. et Lange, 1880, l. c.:468; Franco et Silva, 1968, l. c.:105; Zohary, 1972, l. c.:44; Gladstones, 1974, l. c.:17; Vass. 1987, l. c.:214. – yellow lupin.Typus: Herb. Linn. No 898-8 (LINN). Protologus: « In Siciliae arenosis ».

Plants are 20-80 cm high, rosetted in the beginning and becoming erect in subsequent, with vigorous basal branching. Stems are short and hirsute, with intesive branching in the lower part. The leaf consists of 7-9 (11) ovate-oblong or lanceolate leaflets, prolated at the basis, densely villous on both sides, sized 30-60 х 8-15 mm. Stipules of the rosetted leaves are crescent and chuffy on stalks, linear-obovate in shape. The inflorescence is an elongated truss, 5-25 cm long, set on a peduncle of 5-12 cm. Flowers are verticillate, odorous. Floral bracts are small-sized, obovate, silky-pubescent, easily falling. The upper labium of a calyx is bipartite, the lower one has 3 small denticles. The corolla is 14-16 mm long, bright goldish-yellow in color. Pods are elongated, 40-60 х 10-14 mm, densely villous, 4-6-seeded, with oblique partitions between seeds. Seeds are 5,5-6,5 mm in diameter, spherical-reniform, oblate, variable colouring - from pinkish, brown and yellowish up to dark-violet and mottled. 2n=52. Occurs on mild sandy and volcanic soils in mining belts. As a wild plant, it is widespread over the coastal area in the western part of Iberian Peninsula, Morocco, Tunisia, and Algeria, on the islands of Corsica, Sardinia and Sicily and in southern Italy. In Israel and Lebanon it has most likely turned wild. It is cultivated in Northern Europe and CIS (Belarus and Ukraine) as well as, on a smaller scale, in Western Australia and South Africa. Having previously been cultivated in southern France and on Madeira, it has turned wild there.
Usually this species is considered as an annual one, but in wild environments it is sometimes possible to find two- and four-year plants. The variability of characters in this species is less expressed than in L.angustifolius; however a homologous series can be modeled on the color of seeds, which is more or less similar to L.angustifolius. Dominating coloring of flowers is yellow or, less frequently, lemon-yellow, orange and whitish. Using combinations of such characters as the color of the corolla, the carina`s edge, vegetative organs and seeds, 18 varieties, 4 subvarieties and 6 forms have been identified.


The scheme of classification of Lupinus luteus L..
1.Var. luteus 
I. f. volovnenkoae Kurl. et Stankev.
2. f. compactus Kazim. et Kaz.
2. Var. maculosus Kurl. et Stankev.
3. Var. kazimierskii Kurl. et Stankev.
4. Var. arcellus Kurl. et Stankev.
5. Var. sempolovskii (Atab.) Kurl. et Stanke
6. Var. melanospermus Kurl. et Stankev.
7. Var. niger Kurl. et Stankev.
8. Var. cremeus Kurl. et Stankev
9. Var. leucospermus Kurl. et Stankev.
I. Subvar. leucospermus
3. f. ucrainicus Kurl. et Stankev.
2. Subvar. taranuchoi Kurl. et Stankev.
10. Var. citrinus Kurl. et Stankev.
4. f. lukasheviczii Kurl. et Stankev.
11. Var. sulphureus (Atab.) Kurl. et Stankev.
12. Var. stepanovae Kurl. et Stankev.
13. Var. ochroleucus Kurl. et Stankev.
3. Subvar. ochroleucus
4. Subvar. chloroticus Kurl. et Stankev.
5. f. bernatzkayae Kurl. et Stankev.
14. Var. aurantiacus Kurl. et Stankev.
15. Var. croceus Kurl. et Stankev.
16. Var aureus Kurl. et Stankev.
6. f. golovczenkoi Kurl. et Stankev.
17. Var. albicans Kurl. et Stankev.
18. Var. sinskayae Kurl. et Stankev.


1. Var. luteus, 1990. Bull. Appl. Bot. Gen. Pl.-Breed., Leningrad, 135:28. - L.luteus var. maculatus Zhuk. ex Atab. 1962, Lupinus: 151, nom. illeg. - Flowers yellow. Seed white with black dots without arcs (doted). Typical representative accession of this varietas is cv. Academichesky 1 (k-1947).
1. f. volovnenkoae Kurl. et Stankev. 1990, l. c.:28. - Side shoots absent or shortened, flowers axillary. Typus: ´Искорость´, Ukraine, k-2437, reproduction of Pushkin`s laboratories of VIR, 6. 7. 1989, B.S. Kurlovich (WIR).
2. f. compactus Kazim. et Kaz. f. nova. – Flores et leguminis laterali apice conferta. Typus: ´Compactus´, Poland, reproduction of Institute of Plant Genetic, Polish Academy of Science, 10.08.1995, T. Kazimierski (WIR). Flowers and pods clustered at the top of the stem.
2. Var. maculosus Kurl. et Stankev. 1990, l. c.:28. – Flowers yellow. Seed white with black spots and two light arcs. Typus: ‘ Afus’, Poland, k-2106, reproduction of Pushkin`s laboratories of VIR, 28. 7. 1989, L.T. Kartuzova (WIR).
3. Var. kazimierskii Kurl. et Stankev. 1990, l. c.:28. – Flowers yellow. Seed white with brown spots without arcs. Typus: ‘И - 511562’, Israel, reproduction of Pushkin`s laboratories of VIR, 28. 7. 1989, L.T. Kartuzova (WIR).
4. Var. arcellus Kurl. et Stankev. 1990, l. c.:28. – Flowers yellow. Seed white with brown spots and two dark arcs. Typus: ‘И - 511504’, (Popularion B 469/79), Spain, reproduction of Pushkin`s laboratories of VIR, 7. 8. 1989, B.S. Kurlovich (WIR).
5. Var. sempolovskii Kurl. et Stankev. 1990, l. c.:28. – L.luteus var. sempolovski Atab. 1962, l. c.:151, nom. illeg. - Flovers yellow. Seed white with black spots and wide clear space round the scar. Typus: ‘Полуцкий’, Poland, k-2399, reproduction of Pushkin`s laboratories of VIR, 27. 7. 1989, B.S. Kurlovich (WIR).
6. Var. melanospermus Kurl. et Stankev. 1990, l. c.:29. - L.luteus var. melanospermus Körn. ex Atab. 1962, l. c.:151, nom.illeg. - Flower yellow. Seed brown-and-black with light arcs. Typus: ‘Schwako’, Hungary, k-1835, reproduction of Pushkin`s laboratories of VIR, 18. 7. 1989, B.S. Kurlovich (WIR).
7. Var. niger Kurl. et Stankev. 1990, l. c.:29. - Flowers yellow. Seed black, without arcs. Typus: ‘Копыловский’, Ukraine, k-2601, reproduction of Pushkin`s laboratories of VIR, 8. 9. 1989, B.S. Kurlovich (WIR).
8. Var. cremeus Kurl. et Stankev. 1990, l. c.:29. - Flowers yellow. Seed cream with white arcs. Typus: ‘ Müncheberger Süsslupine ’, Germany, k-1653, reproduction of Pushkin`s laboratories of VIR, 10. 7. 1989, B.S. Kurlovich (WIR).
9. Var. leucospermus Kurl. et Stankev. 1990, l. c.:29. - L.luteus var. leucospermus Körn. ex Atab. 1962, l. c.:151, nom.illeg. - Flowers yellow. Seed white.
1. Subvar. leucospermus. - Carina`s edge without anthocyan. Typus: ‘Weiko’, Germany, k-1310, reproduction of Pushkin`s laboratories of VIR, 20. 7. 1989, B.S. Kurlovich (WIR).
3. f.ucrainicus Kurl. et Stankev. 1990, l. c.:29. - L.luteus var. leucospermus Köern. subvar. deramosum Taran. 1980, Cел. и сем. люп.:34, nom. illeg. - Side shoots absent or shortened, flowers axillary. Typus: ‘Юбилейный’, Ukraine, k-1935, reproduction of Pushkin`s laboratories of VIR, 28. 9. 1989, L.T. Kartuzova (WIR).
2. Subvar. taranuchoi Kurl. et Stankev. 1990, l. c.:29. - L.luteus var. leucimelanus Taran. 1980, Cел. и сем. люп.:34, nom. illeg. - Carina`s edge with anthocyan. Typus: ‘Горецкий’, Belarus, k-2169, reproduction of Pushkin`s laboratories of VIR, 9. 8. 1986, A.K. Stankevich (WIR).
10. Var. citrinus Kurl. et Stankev. 1990, l. c.:29. - Flowers lemon-yellow. Seed white with black dots without arcs (doted). Typus: ‘ БСХА-287 ’, Belatus, k-2738, reproduction of Pushkin`s laboratories of VIR, 16. 7. 1989, B.S. Kurlovich (WIR).
4. f. lukasheviczii Kurl. et Stankev. 1990, l. c.:29. - Side shoots absent or shortened, flowers axillary. 11. Var. sulphureus Kurl. et Stankev. 1990, l. c.:30. - L.luteus var. sulphureus Atab. 1962, l. c.:151, nom. illeg. - Flowers lemon-yellow. Seed white with black spots and two light arcs. Typus: ‘ № 12099 ’, Germany, k-2556, reproduction of Pushkin`s laboratories of VIR, 9. 8. 1986, A.K. Stankevich (WIR).
12. Var. stepanivae Kurl. et Stankev. 1990, l. c.:30. - Flowers lemon-yellow. Seed creme-coloured with two white arcs. Typus: ‘WTD-6049’, Poland, to - 2585, reproduction of Pushkin`s laboratories of VIR, 21. 7. 1989, B.S. Kurlovich (WIR).
13. Var. ochroleucus Kurl. et Stankev. 1990, l. c.:30. - Flowers lemon-yellow. Seed white.
3. Subvar. ochroleucus. - Carina`s edge with anthocyan. Typus: ‘Припять’, Belarus, k-2652, reproduction of Pushkin`s laboratories of VIR, 9. 8. 1986, A.K. Stankevich (WIR).
4. Subvar. chloroticus Kurl. et Stankev. 1990, l. c.:30. - Carina`s edge without anthocyan. Typus: ‘Салатный’, Ukraine, k-2156, reproduction of Pushkin`s laboratories of VIR, 9. 8. 1986, A.K. Stankevich (WIR).
5. f. bernatzkayae Kurl. et Stankev. 1990, l. c.:30. - Side shoots absent or shortened, flowers axillary. Typus: ‘Лимонный’, Ukraine, k- 2153, reproduction of Pushkin`s laboratories of VIR, 9. 8. 1989, B.S. Kurlovich (WIR).
14. Var. auranticus Kurl. et Stankev. 1990, l. c.:30. - Flowers orange. Seed white with black dots without arcs. Typus: Belarus, Minsk province, k-2476, reproduction of Pushkin`s laboratories of VIR, 19. 7. 1989, L.T. Kartuzova (WIR). 1
5. Var. croceus Kurl. et Stankev. 1990, l .c.:30. - Flowers orange. Seed white with black dots and light arcs. Typus: Belarus, Minsk province, k-2823, reproduction of Pushkin`s laboratories of VIR, 24. 9. 1989, L.T. Kartuzova (WIR).
16. Var. aureus Kurl. et Stankev. 1990, l. c.:30. - Flowers orange. Seed white. Typus: Poland, k-3135, reproduction of Pushkin`s laboratories of VIR, 20. 8. 1990, B.S. Kurlovich (WIR).
6. f. golovczenkoi Kurl. et Stankev. 1990, l. c.:30-31. - Side shoots absent or shortened, flowers axillary. Typus: ‘Пламенный’, Ukraine, k-2155, reproduction of Pushkin`s laboratories of VIR, 20. 8. 1990, B.S. Kurlovich (WIR).
17. Var. albicans Kurl. et Stankev. 1990, l. c.:31. - Flowers whitish. Seed white with black spots and two light arcs. Typus: ‘ Белоцветковый Т ’, Belarus, k-2973, reproduction of Pushkin`s laboratories of VIR, 21. 7. 1989, L.T. Kartuzova (WIR).
18. Var. sinskayae Kurl. et Stankev. 1990, l .c.:31. - Flowers whitish. Seed white. Typus: ‘ Белоцветковый C ’, Belarus, k-2974, reproduction of Pushkin`s laboratories of VIR, 20. 9. 1989, L.T. Kartuzova (WIR).
The quantity of new intraspecific taxa of Lupinus luteus L., and specially its subvarietas and forms, can be greater in the process of additional investigation.



5. L. hispanicus Boiss. et Reut. 1842, Diagn. Pl. Nov. Hisp. 10; Boiss. 1872, l. c.:28; Willk. et Lange, 1880, l. c.:467; Merino, 1905, Fl. Galic. 1:411; Zhukovsky, 1929, l. c.:265; Gladstones, 1974, l. c.:19.Isotypus: K, « In pascuis arenosis, ad radicis Sa. de Guadarrama, prope Escorial et Colmenar ».

Plants have medium leaves and silvery pubescence. Leaflets are obovate, almost naked from above and pubescent from below. Petioles are twice longer than leaves. Stipules are growing together with petioles on some extent. The inflorescence is sparsely vercicilate. The upper labium of the calyx is bifurcated, the lower one is almost integral and slightly longer. Floral bracts are small, pubescent, with early falling habit. The corolla’s hue spectrum is from cream-colored to lilac and violet; it has no odor. The carina is cariniform with a brown edge. Pods are flattened, sclerotic, hirsute, 40-60 х 6-12 mm in size, with 4 (5-7) seeds. Seeds are equal to, slightly larger or substantially smaller than those of L. luteus. They may be white, olive brown or cream-colored to light reddish or greenish brown with darker brown specks and an arc around the hilum. 2n=52. Grows on moderately and highly acidized soils, on granite and shaly mountain slopes as high as up to 1500 m above sea level, and also on sandy soils. Widespread in southern and central Spain. Presence of L. hispanicus was recorded in Greece, Turkey and Algeria (Zhukovsky, 1929, Gladstones, 1998). Close to L. luteus, but differs from it by a more or less smooth upper part of leaflets, cream, lilac or violet flowers, a little narrower pods, and smaller-sized, slightly tubercular rough (ssp. hispanicus) or smooth (ssp. bicolor) seeds.

Key to the subspecies of Lupinus hispanicus Boiss. and Reuter (Gladstones, 1974).

1. Corolla violet: upper surface of leaflets glabrous; pods 9-12 mm broad, 3-6 seeded; seeds moderately large, pure white or olive brown, slightly tuberculate …… 1. Subsp. hispanicus 

2. Corolla cream becoming lilac; upper surface of leaflets sparsely villous near margins; pods 6-8 mm broad, 5-7 seeded. Seeds smaller, cream to light reddish or greenish brown with variable darker brovn markings, or occasionally pure white, smooth ….............. 2. Subsp. bicolor



1. Subsp. hispanicus. Meets in Southern and Central Spain on granite rocks to about 1200 m above sea-level and on moderately and very acid soils. 2. Subsp. bicolor (Merino) J.S.Gladstones. 1974, W.Austral. Dep. Agr. Techn. Bull. 26:21. - L.luteus var. bicolor Merino, 1905, Contr. Fl. Gal. 72. - L.hispanicus var. bicolor Merino, 1905, Fl. Gal. 1:41. - L.bicolor (Merino) Rothmaler, 1935, Cavan. 7:114, nom. illeg. non Lindley (1827). - L.rothmaleri Klinkowski, 1938, Züchter 10:124, fig. 17. It is widespread in Central and Northwestern Spain and Northern Portugal basically on granite rocks and plateaus to 1500 m. Grows on sands and sandy looms, moderately strongly acid, sometimes on poorly drained soils. Many breeders of Belarus, Russia and Poland call this subspecies as L. rothmaleri Klink. (synonym). L. hispanicus is close, but isolated species in relation to L. luteus. It has not yellow flowers (colouring which one is conditioned by pigments from group of carotenoids), but violet, lilac and cream (from group of anthocyanins). These species have same number of chromosomes (2n=52), but they miscellaneous serologically. The hybrids between them frequently are sterile (Glagstones, 1974), however are obtained also fertile hybrids (Lamberts, 1955, 1958; Kazimierski and Kazimierska, 1970; Майсурян, Атабекова, 1974; Swiecicki, 1988). More viable descendants receives under hybridization of L.luteus x L.hispanicusssp. bicolor, and also L.luteus with hybrids ssp. hispanicus x ssp. bicolor (Cortero et al., 1988). L.hispanicus and L.luteus represent two interacted species (altitudinal and ecological vicariants. They though grow on same Pyrenean peninsula, but L.luteus takes a seaside part of Portugal, and L.hispanicus grows in a mining belt of Spain and Portugal. Probably there were same initial forms at these species, however they had divergent for a long time. Future detail cytological and molecular-biological researches should promote in creation of the new forms, donors of genes of resistance to cold, drought and different diseases on the basis of these species.

6. L. cosentinii Guss. 1828. Fl. Sic. Prodr. 2:440; Cesati, Passer and Gibelli, 1867, Comp. Fl. Ital. 729; Gladstones, 1974, l. c.: 21-24. - L.pilosus ssp. cosentini (Guss). Rouy and Fouc. 1887, Fl. Fr. 4:190. - L.digitatus auct. non Forskal: Lojac. 1891, Fl. Sic. 1 (2):33; Gladstones, 1958, Roy. Soc. W.Aust. 41:31; Kazimierski, Nowacki, 1961, Genet. Polon. 2:115. - L.pilosus ssp. digitatus auct. non Forskal: Fiori and Paol. 1899, Fl. Anal. Ital. 2:11; Zhukovsky, 1929, l. c.:263.
Typus: Catania. In herbosis inter vulcanicos lapides, F.Cosintini.

It is a powerful erect plant 20-120 cm high, with vigorous lateral branching. The stem and petioles are villous, with short hair up to 1 mm long. Stipules are styliform. Leaflets are 9-11, 25-60 х 7-12 mm, oblong–oblanceolate, pubescent on both sides. The inflorescence is 5-15 cm long, on short pedicles. Arrangement of flowers is verticillate, less often subalternate. Flowers are set on short pedicles measuring 1/3 of the calyx’s length. Floral bracts are lanceolate, caducous, with linear bractlets. The upper labium of the calyx is strongly bipartite, the lower labium is 3-toothed. The corolla is 12-17 mm long and 14-19 mm wide, light blue from the sides and yellowish in the center, with a white spot positioned at about 3 mm from the upper margin. The carina is short, blunt, with cyanic pigmentation; it may be mildly odored or odorless. Pods are 40-55 х 13-16 mm in size, ranging from densely villous to softly hirsute, 3-5 seeded. Seeds, 6-9 х 4-7 х 3-4 mm, are square, compressed, tubercular, light grey to more brown with black marks, having narrow arcs around the hilum (2n=32). Widespread in Tunis, Morocco, southwest Spain, southern Portugal, on isolated areas of Corsica, Sicily and Sardinia. Naturalized, domesticated and cultivated in Australia. The major wild populations of western Australia extend along 800 km of mainly coastal sandy soils (Gladstones, 1998). Commonly occurs on disturbed sites. Less frequently occurred in the south of Australia, it may, in certain cases, have been naturalized there in earlier times, but this is uncertain. Any plant of this species found in western Victoria and coastal New South Wales is almost certainly a descendant from later plantings for forage production or dune stabilization. Plants of the Western Australian population (cv. Chapman) are brown-seeded, and it was this type, which early in the present century was taken into cultivation for soil improvement and summer grazing (Gladstones, 1998). The history of this species’ taxonomy has been persistently accompanied by confusion. Botanists attributed this species to miscellaneous taxa: L. pilosus, L. digitatus, L. hirsutus, L .varius.



7. L.digitatus Forsk. 1775. Fl. Aegypt.:131; Gladstones, 1974, l. c.:24.Typus: « In Delta ad pagum Nedjel. Copiose in agris. an spontaneus, nescio ».

The plant is intensively pubescent with soft silky silvery hair. Stems are erect, up to 40 cm high, sparsely branched. Leaflets are oblong-obovate, acuminate on the apex, densely sericeous on both sides. Petioles are twice longer than leaves. Stipules are styliform, grow together with petioles on some extent. The inflorescence is 3-15 cm long, oliganthous, abnormally verticillate. Floral bracts are obovate-oblong, pubescent, falling not instantly. The upper labium of the calyx is bipartite, the lower one is longer and entire. The calyx has bractlets. The flowers are large. The corolla is about 16 mm long and 18 mm high, blue with a white or yellow spot in the middle of the vexillum. The wings are cyanic, matching with the vexillum, of oval shape. The carina is white with a violet edge. Pods are dehiscent, beige, velvety, 5,5-6 х 2,5 cm in size, with 3-4 seeds. Seeds are oval-spherical, compressed, with a prominent smooth hump over the hilum. The testa is scabrous, reddish brown with darker mottling and the arc, 7 х 6 х 3 mm. (2n=36). Widespread in Egypt (in the delta of Nile), Senegal, Central Saharan uplands, and also in the semi-desert near-shore area of the North-Western Sahara. Grows on sandy soils, on the bottom of brooks, among wilderness grasses and acacias, in deserts as a weed of winter crops.



8. L.princei Harms. Harms in Engler, 1901, Bot. Jahrb. 28:4-1; Gladstones, 1974, l. c.:25-26.Typus: « Iringa, Savanne, Hauptmann Prince, 1898 ».

Robust annual erect plant, 30-120 cm high, with sparse branching. Stalks and petioles are covered with short villi. Stipules are linear. Leaflets are linearly–oblanceolate, densely and shortly sericeous from both sides. The inflorescence is 5-20 cm long, on stout peduncles, verticillate. Floral bracts are lanceolate, with early falling habit. Bractlets are lanceolate. The upper labium of the calyx is 2-partiate; the lower one is almost integral and longer than the upper. The corolla is large, 17-18 х 18-20 mm, bright blue. The vexillum has yellow stria in the middle. The carina is bent at 80-900, with a blunt edge, unpigmented. Pods are 50-70 х 16-18 mm in size, softly hirsute, with 3-4 seeds. Seeds, 9-10 х 8-4 mm in size, are square, squeezed, with a slight hump near the hilum, rough, mottled dark brown on a yellow background, with a brown arc around the hilum. (2n=38). Widespread in the highlands of Kenya and Tanzania, at altitudes of 1700-3000 m above sea level, and also in Southern Ethiopia. This species is close to L. digitatus. Its seed is very large, which is typical for traditionally cultivated species, although no indications have been found concerning its cultivation.



9 . L.pilosus Murrey. 1774, Syst. Veg. Ed, 13:545; Boiss. 1872, l. c.: 27; Kazimierski and Nowacki, 1961, Gen. Polon. 2:115; Kazimierski and Kazimierska, 1970, Gen. Polon. 11:208; Gladstones, 1958, J. Roy. Soc. W. Austral. 41:31, pl. 1, 1; Gladstones, 1974, l. c.: 26-29. - L.hirsutus L. 1753, Sp. Pl.:721 (diagn. inaccurate), non L. 1763, Sp. Pl. ed. 2, 1015.

Typus: non published.

Large, up to 80 cm, but sparsely foliated plant, densely hirsute. Branching is both monopodial and sympodial. Stems are powerful. Leaflets of 4.0-5.5 х 1.2-1.7 cm are obovate, slightly bent along the main nerve. Stipules are large, up to the middle they grow together with petioles. The inflorescence is small, with 5-6 verticils, and 5-6 flowers in evach of them. Floral bracts fall off early. The calyx is with bractlets. The upper labium is deeply bipartite and arched back in the basis, the lower one is integral. The corolla is white, blue, cyanic, pink or red. The vexillum with a white spot in the middle is twice wider than the wings. The carina is white with a violet edge. Pods are very large, hirsute, 2-4 seeded. Seeds are very large, 10-14 х 9-12 х 6-8 mm, rough, brownish red in color, with a darker crescent around the hilum. (2n=42). Grows on inshore sands and in mountains up to 1200 m above sea level. Occurs as a weed in crops, on fallow lands and on disturbed soils. Probably it was once cultivated in Africa and from there was dispersed in the Mediterranean area. Widespread in Greece, on Crete and the Aegean Islands, Turkey, Syria, Israel and Lebanon. The taxonomy of L. pilosus Murr. for a long time remained confused. Gladstones found for it the following synonyms: L.hirsutus L., L. varius L., L.varius subsp. orientalis Franco et P.Silva. Maissurjan and Atabekova (1974) considered as synonyms of this species also L. varius L., L. silvestris Lam. and L. semiverticillatus Desr. They leave Linnaeus as the author of L. pilosus, though Lupinus pilosus Murr. has received almost general recognition. Some of the botanists until now go on using the name L. hirsutus instead L.pilosus. The reason of these misinterpretations lies in the fact that in the standard herbarium of Linnaeus the accession of L.pilosus (No 898.3) was lost and probably incidentally replaced by a sample of L.varius. The following sequence number (No 898.4) is assigned to L. micranthus (Gladstones, 1974).


10. L. palaestinus Boiss. 1849, Diagn. Pl. Or. Nov. Ser. 1, 9:9; Boiss. 1872, l. c :27; Zhukovsky, 1929, l. c.:265; Zohary, 1972, l. c :42; Gladstones, 1974, l.c.:29.Typus: « In cultis regionis Philistaeirum et in Arabia Palaestinae Contermina ».

The plant resembles L. pilosus, though it is much smaller, up to 40 cm in hight. Initially it develops as a rosette, from which short stalks and protracted inflorescences are issued. Stems and petioles are hirsute. Leaflets are linearly obovate, acuminate, and pubescent from both sides. The inflorescence is 5-20 cm long, verticillate or almost alternate, friable. Floral bracts and bractlets are linear. The upper labium of the calyx is profoundly bipartite, the lower one is integral, narrow, and longer. The corolla is pale yellow or pale pink, with blue tint. The carina has a violet edge. Pods, 50-70 х 18-10 mm in size, are coarsely hirsute, 3-4 seeded. Seeds are squeezed, rough, pale with a thick brown crescent and dark spots around the hilum. (2n=42). Occurs on semi-desert sands and in the deserts of Sinai Peninsula. It probably grows as a winter or ephemeral crop in moist environments on sandy soils. Widespread in Palestine and Israel.

11. L. atlanticus Gladstones, 1974, W. Austral. Dep. Agr. Techn. Bull. 26:30-32.Holotypus: Loc. Amizmiz. Morocco, alt. 4500 feet, June 16, 1936, E.K. Balls, № 2846 (K).

It is a powerful plant up to 60 cm high. The stalk and petioles are softly villous, hair is white, up to 2 mm in length. Stipules are long, 8-18 х 1 mm. Leaflets are 9-11 in number. Light green, they are oblong–oblanceolate or linear-oblanceolate, with pubescence on both surfaces. The inflorescence is set on a long peduncle. The arrangement of flowers is subverticillate or verticillate. Flowers are large, about 18 mm in length and 20-22 mm in width. The upper labium of the calyx is profoundly bipartite, the lower one is long and integral. Pedicles about 1/3 length of calyxes; the vexillum is blue with a white or yellowish medial band reaching the upper edge. The wings are blue, the keel is blunt with a blue edge. Pods are hairy, 40-70 х 15-22 mm in size, 3-5 seeded. Seeds are oblong, compressed, 8-11 х 6-8 х 4-5 mm in size, mottled brown, or pinkish with a pale arc around the hilum, slightly rough. (2n=38). L. atlanticus differs from other species by the shape and colouring of seeds. Its large flowers are sometimes similar to the flowers of L. pilosus, and the light green colouring of leaves makes it like L. digitatus. Large seeds and usual proximity to cultivated lands indicate that this species was very likely in the past and is perhaps at present used for human consumption (Gladstones, 1998). Distributed in Anti-Atlas, on the foothills of the High Atlas Mountains, and in Djebilet, Morocco, at the altitudes of 900-1500 m above sea level (Gladstones, 1974).


12. L. somaliensis Baker, 1895, J.G. Baker, Bull. Roy. Gard. Kew 105:213.
Holotypus: Somaliland, Mrs. lord Phillips (K).
Upper surface of leaflets glabrous, pedicel about 1/2 lenght of calux, rounded white spot on standard, not approaching the upper margin. Annual (or doubtfully perennial).  Distribution: Highlands of Somalyland, perhabs extending westward into Ethiopia.

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