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Pp.42-43 in the BOOK: Lupins

Geography, classification, genetic resources and breeding

 

 

 

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

 

The ovary contains two, four and more 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 100 up to 1000) requiring further analysis of their authenticity.

 

II. Subgen. Lupinus - A. Eulupinus Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221, p.p. - Old Worlds lupins.

 

The ovary has at least four or more 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.).

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. Basic characteristics of the genus Lupinus L. and its taxa are presented in the Table.

This genus is the principal object of our research. Geographic differences between the New and Old Worlds 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 Vavilovs 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). Vavilovs (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.

 

 

Table. The basic characteristics of taxa in the genus Lupinus L.

 

Taxa

The main characteristics and indices

 

Genus

(gen.)

Area. Main morphological characteristics of the genus:

The leaf is palmate compound, set on a long petiole. The inflorescence is polyanthous apical truss. The corolla of flowers is zygomorphic and papilionaceous. The pod is round or flat, coriaceous, straight or curved. The ovary has two, four or more seedbuds. .

Subgenus (subgen.)

Area; quantity of ovules and seedbuds in the ovary; degree of development of the embryo and endosperm in seed; shape of cotyledons and pods; definite range of variation in the chromosome number; way of pollination; cycle of development; type of branching; definite range of variation in protein fractions.

Species

(sp.)

Area; crossability; ability to produce fertile progeny with constant inheritance of the characters of both parents; identity of the karyotype. System of morphological, anatomic, cytological, caryological, paleobotanical, ontogenetic, biochemical, physiological, geographical, genetic and other characters (Vavilov, 1931, 1965)

Subspecies (subsp.)

Growing area or natural habitat; definite range variation in the shape and color of vegetative and generative organs; pubescence, shape and color of seed.

Variety

(var.)

Color of the seed coat in correlation with the color of the corolla.

 

Subvariety (subvar.)

Coloring of the cotyledons, vegetative organs, edge of the carina, and the presence/absence of anthocyan.

Form

(f.)

Determinate branching, fascicular stem, and other characters of breeding value.

 

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 and Lupinnelus. 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 to Lupinus. To the Platycarpos section were attributed some annual species with two ovules in the ovary and two seeds in the pod (L. densiflorus Benth., 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 (Tab.1), 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 (Wats.) Kurl. in our new combination integrates the numerous perennial and annual species from the Western hemisphere, both groups having two, four and more seedbuds in the ovary, while subgen. Lupinus L. includes 11 species from the Mediterranean region and Africa with as a minimum four and more seedbuds in the ovary.

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

 

The ovary contains two, four and more 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 100 up to 1000) requiring further analysis of their authenticity.

 

II. Subgen. Lupinus - A. Eulupinus Aschers. et Graebn. 1907, Mitteleurop. Fl. 6,2:221, p.p. - Old Worlds lupins.

 

The ovary has at least four or more 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.

 

In our opinion, it is debatable to reckon L. somaliensis Baker (Baker, 1895, Bull. Roy. Gard. Kew, 105:213) among the number of factual species, since no one of the contemporaries had even seen it. At present, its existence now seems rather doubtful. ?

 

You can read the results of our researches in more detail in the offered book:

 

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Geography, classification, genetic resources and breeding

 

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