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
World’s 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 World’s 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 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».
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. |
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 World’s 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 World’s 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:
All materials of the book are here!