Monday, July 17, 2006

PREFACE


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« God said: Let the Earth produce vegetation, seed-bearing plants, trees bearing fruit with seed, each according to its kind, upon the Earth.
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And so it was.

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The Earth produced vegetation: plants bearing seed according to their kind and trees producing fruit which has seed, according to their kind.
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God saw that it was good. »
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.(Bible, Genesis 1:11,12)
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PREFACE
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The present work is a sequel and the first English publication among the unique series of volumes “Flora of cultivated plants” started by N.I. Vavilov. Justifying the issuing of volumes of this serial on separate crops, he specified this publication as an encyclopedia on cultivated plants, major guidebook and directory for every biologist, plant breeder and plant grower. By now, the N.I.Vavilov Institute of Plant Industry (St. Petersburg, Russia) has issued several dozens of volumes of this serial dedicated to main agricultural crops with a bias towards national specificity of Russia and in Russian.

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The volume devoted to long-term study of the collection of some annual grain legumes was for the first time published in 1937. However, due to a number of circumstances lupin was not included in this volume. The present book embraces the results of more than sixty years of monographic research on lupin genetic resources. Research materials are represented by the lupin collection of the N.I. Vavilov Institute of Plant Industry (VIR) storing now more than 2500 accessions of different lupin species collected by numerous expeditions over the globe. Theoretical basis for our research activities were N.I. Vavilov’s works on plant genetic resources.

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We have used the special methodology of the Vavilov Institute developed with regard to lupin (Kurlovich et al., 1990b).

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Many years of observing plants sown in various regions have enabled us to find out new regularities in the variability of characters depending on genetic features of the species and accessions, and also on ecological and agronomic conditions of their growth. We have also disclosed new characters and their combinations, both taxonomically significant and genetically determined. This factor as well as generalization of the data obtained by other researchers (Zhukovsky, 1929; Kazimierski and Novacki, 1961; Gladstones, 1974; Maissurjan and Atabiekova, 1974; Kazimierski and Kazimierska, 1975) made it possible to introduce essential corrections in the intra-specific taxonomic and eco-geographic classifications of lupins. These classifications have been developed on the basis of Vavilov’s concepts (the doctrine about the species as a system, the law of homological series in hereditary variation, differential systematic and geographical method of crop studies, etc.). The sites for studying lupin accessions were chosen with regard to the duration of vegetation periods, and soil and climate conditions required obtaining vigorous seed of each sample. The collection of white lupin was reproduced in the non-black-soil zone of the Ukraine (Kiev Province); yellow lupin in the forest-steppe zone of the Ukraine (Zhitomir Province) and on non-black soils in the Ukraine (near Kiev); narrow-leafed lupin in the non-black-soil region of Russia (Moscow Province) and in the black-soil zone of Russia (Tambov Province); multifoliate or Washington lupin near Pushkin (Leningrad Province, 20 km away from St.Petersburg); Pearl lupin (L. mutabilis Sweet.) and other species from America in the black-soil zone of Russia (Tambov Province). Besides, with the purpose of revealing the range of variability for specific characters and substances, the same accessions were also reproduced in other places. In particular, for a significant part of accessions replication were performed at the former Sukhumi Experimental Station of the Vavilov Institute (in Abkhazia) in the conditions of humid subtropics, at the former Central Asian Branch of VIR (near Tashkent, Uzbekistan) in the conditions of dry climate, in the black-soil zone of the Ukraine (Poltava Province), in the conditions of non-black-soil zone of Russia near the town of Pushkin (Leningrad Province, 20 km from St.Petersburg), and in Bryansk Province (at the All-Russian Institute of Lupin and Novozybkov Branch of the All-Russian Institute of Fertilizers and Agricultural Soil Science). Field investigations have been supplemented by herbarium studies and laboratory analyses. The purpose of this research is to accomplish comprehensive study of the global botanical and varietal diversity of different species in order to gain deeper insight into the problems of lupin geography, evolution and classification, and also to find out regularities in the variability of major valuable morphological and biological properties in various eco-geographic conditions of its cultivation.

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The existing worldwide shortage of protein is generating special attention to lupin. In Russia, where climatic resources for cultivation of soybean are limited, lupin in the long run can play the role that soybean has gained in the U.S., i.e. it may become a highly efficient source of vegetative protein. Special interest in lupin is stirred up by high content of protein in its seed (up to 50%), as well as of oil (from 5 up to 20%) whose quality is close to that of olive, and by the absence of digestion inhibitors and other anti-nutritional substances. Lupin seed have been used since ancient times as human food and animal feed (Gross, 1988; Lopez-Bellido and Fuentes, 1986; Beirão da Costa, 1994). Green material of low-alkaloid (sweet) varieties is also excellent forage. Due to their symbiosis with nodule bacteria lupins are capable to accumulate in soil up to 200 kg of nitrogen per hectare, thus being perfect symbiotic plants. Utilizing lupin as green manure helps to protect environments from pollution, go without expensive fertilizers, and obtain ecologically clean products (Lopez-Bellido and Fuentes, 1986; Lopez-Bellido, 1994). It should be marked that the progress achieved by man in domestication of lupins is rather modest. Of several hundred lupin species existing in a nature, only three annual species and one perennial (Lupinus polyphyllus Lindl.) are used in Russian agriculture for forage production. However, many wild species of lupin are promising for agricultural use. Besides, further efforts are needed to solve the problems of plant resistance to diseases and pests, higher adaptability to unfavorable environmental factors, and breeding of forms with high technological properties for agricultural production. With these goals in view, special significance is acquired by scientifically justified selection of breeding sources, diversity of such materials, and the degree of comprehensiveness in studying them. In the outcome of long-term studies of the lupin collection stored at the N.I. Vavilov Institute of Plant Industry, it became possible to accumulate a great deal of original data on this crop’s specific and varietal structure, its geography, ecology, systematization, genetics, biochemistry, immunology, and selection methods. All these issues are dealt with in the present book.
It is based on our own researches, as well as on the research data received by prominent scientists, such as P.M. Zhukovsky, E.I. Sinskaya, B.M. Libkind, V.S. Fedotov, N.I. Sharapov, N.N. Ivanov, M.I. Smirnova, V.N. Dyubin and other experts who worked in the experimental network of the Vavilov Institute in different times, N.A. Maissurjan and A.I. Atabiekova who conducted fundamental research on lupin in Moscow Agricultural Academy. The national (Russian) and world literature on lupin has also been generalized. Russian references are made here in the original (Cyrillic) alphabet. In our opinion, this may facilitate their search and additional study. Our work essentially supplements the book published in 1998 by the scientists of Australia «Lupins as crop plants» (edited by J.S. Gladstones, C. Atkins and J. Hambling, 1998). In their book, the materials dedicated to studying and cultivation of lupin in Australia and many other countries were summarized. However, information about lupin growing in the ex-USSR countries was provided there on a limited scale, probably due to certain difficulties in translating Russian literature. It should also be mentioned here that the world community on the whole has very limited access to scientific information on lupin cultivation in the former Soviet Union.
The present work is supposed to fill this gap, so that the data of the researches conducted by Russian scientists could be available to the world’s scientific community. We attempted to throw light on the global achievements in studying lupin by exposing the main developments of Russian, Polish, Byelorussian and Ukrainian scientists in this sphere. The contributors of this book are the liding scientists from Russia, Poland and Finland. Originally, the present work had been written in Russian ...(Kurlovich B.S., Rep'ev S.I., et al. 1995), but later was translated into English.

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The authors’ goal was to make scientific achievements of Russian scientists known to wider scientific circles of the world. Unfortunately, the English version of the text may contain discrepancies of translation and stylistic faults. The editor is making an apology for this; as it is actually his first attempt to present in English the existing voluminous materials about lupin in Russia.

Special thanks are addressed to Mr. A.G. Krylov for his help and counsel while translating this book. I would like to express my gratitude and compliments to all scientists of the Vavilov Institute and other institutes, who carry on the work with plant genetic resources started by N.I. Vavilov. We were able to generalize this research only owing to the close cooperation and dialogue with many of them.

Special gratitude is extended to Professors N.I. Korsakov, V.F. Dorofeev, M. G. Agaev, N.M. Chekalin, G.G. Davidjan, K.Z. Budin, B.N. Malinovsky, A.F. Merezko, B.V. Rygin, A.I. Shishkin, V.I.Bureniv and V.D. Kobyliansky, as well as to Drs. A.V. Khotyanovich, N.N. Nazarova, S.I. Pilipenko, V.A. Rybnikova, O.N. Korovina, M.P. Tarasov, T.V. Buravtseva, and F.T. Tarba.

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We do express deep appreciation to the Government and to the People of Finland for warm reception and for making favorable environment while this book was written. This work has been edited and published thanks to the financial support of Finish Company International North Express Co (Raimo Heinänen and Päivi Heinänen)

Boguslav S. Kurlovich

You can read our book on the Internet here!

You can buy the book here, and here!

The Russian version of the book on - http://lupin-rus.blogspot.com/

Intraspecific diversity of Lupins on - http://lupindiversity.blogspot.com/

Lupins as a potential crop in Finland - http://lupin-fin.blogspot.com/

Links to more Lupin info:

PLANTS Profile - Lupinus L. - lupine - http://plants.usda.gov/java/profile?symbol=LUPIN

Lupin - http://en.wikipedia.org/wiki/Lupinus

Search for: Lupinus L. - http://www.pfaf.org/database/search_name.php?ALLNAMES=Lupinus+L

Lupinus L. - http://www.itis.usda.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=25916

Tree Lupins - http://www.geocities.com/~gregmck/lupins/lupintax.htm#arboreus

Lupinus albus L. -
http://www.hort.purdue.edu/newcrop/nexus/Lupinus_albus_nex.html

L. angustifolius - http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?22805

Lupinus mutabilis - http://en.wikipedia.org/wiki/Lupinus_mutabilis

Gasid is one of the first italian companies to care about the distribution of LUPIN, promoting the consumption of this product - http://www.lupinor.com/300/e301.htm

We are waiting to more links on a theme: Lupins !!!


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THEORETICAL BASIS OF OUR RESEARSHES



THEORETICAL BASIS OF OUR RESEARSHES

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B.S. Kurlovich, S.I. Repyev and L.T. Kartuzova

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Theoretically, our research is founded on the works of Prof. N.I.Vavilov. Nikolai I. Vavilov is recognized as the foremost plant geographer, botanist and geneticist of this age. In the early 20th century, the world was yet unaware of the urgent necessity to protect the environments, and scientists thought little about gradual extinction of plant species. In fact, N.I. Vavilov was the first who understood the imperative of intensive plant collecting, studying and preservation, in spite of the hardest times he experienced in his life. In order to explore the major agricultural regions over the world, Vavilov organized and took part in over 100 collecting missions. During these expeditions, he always focused special attention on leguminous crops in general, and to lupin in particular, regarding it as a source of protein and means of increasing of soils fertility (Синская,1991). He considered this problem as the highest priority in biological and agricultural science and in the attempts to develop sustainable agricultural production. Collecting activities of N.I. Vavilov were started in 1916 with his foreign trip to Iran (Hamadan and Khorasan) and Pamir (Shungan, Rushan, Khorog), and never ceased until the end of his life. All expeditions undertaken by Vavilov and his followers were inspired with the idea of learning about the best plant resources of the world and placing them at the disposal of agricultural practice, thus enriching it with new valuable crops or varieties. As a result of these efforts, by 1940 about 200 thousand accessions had been stored in Vavilov’s collection. Such a rich diversity was thoroughly analyzed in various eco-geographic conditions, systematized and preserved. Vavilov’s foresight concerning possibility of quick disappearance of many valuable plant forms under the influence of human activity proved prophetic. Only for the last 20 years, dozens of plant species and forms have vanished from earth, but many species and forms were saved thanks to the collecting efforts of Vavilov and his followers. During our expeditions abroad, we were repeatedly addressed by the scientists of many countries (Brazil, Israel, Portugal, etc.) with requests to supply germplasm samples collected earlier in their countries by N.I. Vavilov. Such need was evoked by the situation when the requested plant forms with many valuable properties did not exist any more in natural or agricultural environments in these countries. Thanks to Vavilov, they were saved only in the collections of VIR, and are now freely available for everybody. In order to know the great diversity of plants, they ought to be systematized and put in order. Therefore, since the first years of its establishment and study, the plant collection has been used as basic material for working out vital laws and regularities of general biological nature, as well as theoretical fundamentals of plant breeding. The concept of global genetic diversity of cultivated plants remained the keystone of Vavilov’s work throughout his whole life. It incorporated a number of major theoretical conclusions, which played an important role in the development of botany, genetics and plant breeding. The law of homologous series in hereditary variation, idea of the species as a system, differential systematic and geographical method of crop research, botanical and geographical aspects of breeding, and the theory of the centers of origin of cultivated plants formed the core of these studies (Вавилов, 1920, 1926, 1935, 1940; Vavilov, 1997). Vavilov’s theory of plant introduction was developed on the basis of studying the earth’s vegetation. He identified a number of areas distinguished by exceptional diversity and richness of plant species and forms (Вавилов, 1965). Over 70 years ago (Vavilov, 1926), he substantiated the selection of five ancient foci of origin of cultural plants from local flora. Later (1935), he introduced additions and amendments to the decoding of these foci and outlined 6 foci and 2 centers of origin of cultivated plants. It is necessary to note that Vavilov always used to analyze his works in a critical way, developing and deepening their main conclusions in his subsequent publication. In 1940 (Вавилов, 1940), as well as in his Five Continents (published for the first time in Russian in 1962, and translated in English in 1997), Vavilov already named seven main primary centers of origin, placing foci within some of them. The Indian and Indo-Malaysian focus earlier described by him was, therefore, attributed to the South Asiatic tropical center (1940), which in 1962 was renamed Tropical. The Asia Minor and the Central Asian foci were attributed to the southwestern Asiatic center, and the South American center (1935) was renamed Andean (Вавилов, 1940, 1962, 1987). Finally, Vavilov offered the following seven centers: Tropical, East Asiatic, Southwestern Asiatic, Mediterranean, Abyssinian, Central American and Andean. Within the limits of some of these centers, specific geographical foci were also distinguished. According to Vavilov, a center of origin of cultural flora is determined by two prerequisite conditions: abundance of plants suitable for domestication in the local flora, and availability of an ancient agricultural civilization. The work on localization of the centers and foci of origin of cultivated plants was continued by Sinskaya (Синская, 1969), Zhukovsky (Жуковский, 1971), and other scientists from many countries. Sinskaja (1969), in her efforts to expand geographical connections and reciprocal interaction of cultural florae, suggested a broader notion of a historical-geographic area. She identified five areas and considerably updated the list of cultivated crops in each area. Zhukovsky (1971) accepted the theory of Vavilov’s centers, but increased their number to 12 and named them gene centers.
In our opinion (Kurlovich, 1998), there are centers of formation of wild plant species in the places of their origin after the last ice age, and also centers of origin (diversity) of cultivated plants in the places of their introduction into cultivation practice (domestication). Among the supporters of the existence of N.I. Vavilov’s centers there are many scientists from different countries (Kurth, 1957; Harris, 1967; Harlan, 1971; Брежнев and Коровина, 1981; Mathon, 1981). Some critics argue that it is very difficult to determine the very first geographical origin of a species. With this in view, they consider as more proper the term “centers of diversity” in lieu of Vavilov’s “centers of origin”. The term “centers of diversity” is regarded as a safer one (Pistorius, 1997). Thus, Vavilov’s theory of the centers of origin or diversity of cultivated plants has constantly been updated, improved, and even, in some respects, criticized, but it is nevertheless still used as a theoretical basis for collecting, study and utilization of crop genetic resources. So far as lupin is concerned, Vavilov considered the Mediterranean region and mountainous areas of Mexico, Peru and others American countries as native lands for this genus (Вавилов, 1926). Reviews performed on the basis of the approach offered by Vavilov helped to attain more precision in identifying centers of formation and origin for separate lupin species (L. albus L., L. luteus L. and L. angustifolius L.). Their centers were determined by applying the criterion of the greatest diversity of forms in a definite locality with primitive dominant characters. Distribution of lupins as crop plants also resulted in formation of their secondary centers in the areas of intensive cultivation. These findings made it possible to specify places of primary importance for collecting missions and obtain lupin samples valuable for future breeding for resistance to diseases, unfavorable conditions of cultivation, etc. Any plant species when studied in a wide range of geographical conditions, involving inbreeding, segregates into a wide range of hereditary forms, which is difficult to comprehend at first sight. Yet, in studying intraspecific diversity one can observe certain regularities and reveal similarities in accordance with to N.I. Vavilov’s law of homologous series in hereditary variation. The essence of this law may be described as follows. Species and genera more or less closely related to each other manifest similar series of variability lined up with such regularity that, knowing the succession of varieties in one species, one could forecast the existence of similar forms in other species or genera. The closer is the alliance between the species or genera within the general system, the more complete is their similarity in the series of variation. Whole plant families are in general characterized by a definite cycle of variability, which goes similarly through all genera and species of a given family. The laws discovered Vavilov help to put in order extensive materials of cultivated and wild vegetation. N.I. Vavilov marked out the congruity of variability in his description of the whole family of Fabaceae Lindl. The tables with the data and figures on this section are here. Proceeding from the study of separate genera within this family, he established the laws of their differentiation into varieties observable in the sets of characters displayed in seed, fruit, flowers, and vegetative organs. He provided detailed analysis of the variability of characters in the representatives of sections Vicieae, Trifolieae, Loteae, Galegae and Phaseoleae (Вавилов, 1920; 1987). It is clearly visible from his materials that, in spite of the differences existing between the above-mentioned sections, they demonstrate similar variability of characters, mandatory for all genera of the family. Although he presented no data as for the genus Lupinus L., subsequent investigations have shown, that Lupinus illustrates Vavilov’s law of homologous series in variation even better than any other genus of the Leguminosae family. Thus, conducting regular studies of the major lupin species one can observe not only immense diversity in many thousands of plant forms, but also parallel series even in pathological, mutagene and hybrid variability of a given plant. Describing the materials supporting his law of homologous series in hereditary variation, Vavilov paid considerable attention to the feature of plant color. Since various coloring of plants is an approved hereditary character transferred from generation to generation, it was introduced into systematic at a distinctive characteristic of separate species, varieties, subvarieties and smaller taxonomic units. Presently available data show that lupin has a correlation between the colors of seed, inflorescence, carina’s edge, and vegetative parts. These factors facilitate the task of making up intra - and interspecific systematics of plants (Вавилов, 1935). Combination of the coloring of seed, vegetative parts, inflorescence and carina’s edge in various species of lupin from both hemispheres, and complete parallelism in their age variability and in morphological and biological characters doubtlessly witness to their genetic closeness. Application of the above-mentioned laws allowed us to locate and describe several new forms so far unknown in different lupin species, and develop their intraspecific classification. The law of homologous series in hereditary variation gives an answer to the question what material should be looked for, while the theory of the centers of origin of cultivated plants provides an answer to the question where it could be found. To study intraspecific diversity and to determine the centers of origin of cultivated plants, Vavilov and his followers used a differential systematic and geographical method of crop studies (Вавилов, 1931), which meant as follows: · differentiation of a genus into species and intraspecific diversity with the help of morphological, hybridlogical, cytological and others methods; · determination of the genotypic composition of a species; · geographical localization of hereditary forms of a species as well as the centers of their diversity. Of great theoretical and practical importance and subject to further development is Vavilov’s concept of the species as a system (Вавилов, 1931,1965a). Previously, in the science prevailed the notion of Acad. Komarov about monotypic species. According to it, the species cannot include any systematic units of a lower rank (Комаров, 1931,1944). Also widely known was the concept of biological species which postulated impossibility of crossing between species (Grant, 1981,1984). Vavilov made a presentation of one of his fundamental concepts "Linnaean Species as a System". Practical study of several hundreds of species showed the absence of monotypic species, i.e. the species represented by one certain race alone or a certain form alone.
Each species appeared to incorporate a larger or smaller number of forms (genotypes). Vavilov regarded a species as a flexible isolated complex morpho-physiological system linked in its genesis to a certain environment and area of distribution (Вавилов, 1931, 1965a). The research on several hundreds of cultivated species performed by a great number of scientists according to a strictly regulated program lead him to the understanding of the Linnaean species as a definite complex system, i.e. an integrity consisting of closely interlinked components, where the whole and the parts are merged with each other (Вавилов, 1965a, Агаев, 1987, Коровина, 1987). Therefore, in the study of species on the basis of Vavilov’s theories (concept of the species as a system, and differential systematic and geographical method of crop studies) attention was focused not only on morphological characters, but also on geographical and ecological differentiation and other properties of plants. Such approach supported by the development of different intraspecific classifications helped to accomplish profound and comprehensive analysis of intraspecific and varietal diversity of cultivated plants, and find ways of efficient utilization. In this respect, the International Code of botanical nomenclature fixed such categories as subspecies (subspecies), varieties (varietas), subvarieties (subvarietas) and form (forma).
Besides, Vavilov gave special heed to eco-geographical differentiation of the species into ecotypes, geotypes, concultivars, etc. (Вавилов, 1931, 1965a). The postulate of the species as a complex multilateral phenomenon would urge a researcher to use diverse methods for identifying differences between intraspecific categories. In addition to the main morphological method, the scientists of the Vavilov Institute have widely used anatomic, cytological, caryological, paleobotanical, ontogenetic, biochemical, physiological, geographical, genetic and other methods. Such comprehensive approach is especially efficient when the intraspecific diversity of cultivated lupin forms is concerned. This Vavilov’s differential systematic and geographical method of crop studies is interconnected with other findings of N.I. Vavilov and is based on his law of homologous series in hereditary variation, his theory of the centers of origin (diversity) of cultivated plants, and his concept of the species as a system (Вавилов, 1920, 1926, 1935, 1987b). All Vavilov’s fundamental ideas interconnected among themselves and represent a complex doctrine about global genetic diversity of cultivated plants. On the basis of this doctrine, the experts of VIR have developed intraspecific classifications practically for all leguminous crops, including peas (Говоров, 1937; Макашева, 1979), mung bean (Попова, 1937), soybean (Корсаков, 1971, Теплякова, 1997), chickpea (Сеферова, 1997), and vetch (Станкевич and Репьев, 1999). These classifications make it possible to disclose completely the potential of leguminous crops, and they are widely used in breeding practice and plant science. We have also reviewed the system of Lupinus L. using Vavilov’s concept of the species as a system, and applied his differential systematic and geographical method in our studies. On this basis, we offer intraspecific and eco-geographical classifications for three cultivated annual species of lupin (L. albus L., L. luteus L. and L. angustifolius L.). This approach provided a possibility to perform a targeted search of genetic resources of lupins to solve problems of their evolution and selection. It enabled us not only to cast light on the diversity of lupin forms, but also to reveal a series of regularities in their variation depending on the degree of cultivation, geographic environments and soil conditions. Such outcome, in its turn, helped in finding and recommending valuable initial material for breeding. N.I. Vavilov laid a foundation of the lupin collection in VIR. Valuable accessions of white lupin (Lupinus albus L.) were collected by Vavilov during his trip to the Mediterranean in 1926. In Palestine, he managed to find very early, thermally neutral and small-seeded forms relating to the Jordanian ecotype. In particular, the duration of growing period of the sample Tel Karam (k-290) in the conditions of the Ukraine was only 105 days. From Sudan Vavilov brought, on the contrary, very late, but highly productive and large-seeded forms belonging to the Sudanese ecotype (k-486, k-495). Of special value as sources for lupin breeding in Russia, Poland and the Ukraine were samples of the Jordanian ecotype from Palestine. Their hybridization with samples of the Georgian ecotype, and also their involvement in combinations with application of mutagenesis, allowed V.I. Golovchenko to create early and high productive cultivars of white lupin, Kievsky mutant, Horizont and Druzba, suitable for the condition of Russia and the Ukraine (Головченко et al, 1984).
Not only did Vavilov collect and organize studying of the assembled plant materials, but also he constantly strove to improve methods of these studies. He took the initiative, in particular, to develop a simpler and more widely accessible method of determining alkaloids in lupin. This effort of his was inspired by the fact that the technique of detecting and determination of lupin forms with low alkaloid content elaborated in 1928-1929 by v. Sengbusch in Germany was kept secret, since sweet strains of lupin were sold to a private firm. As a result, under the leadership of Prof. N.N. Ivanov, M.I. Smirnova and other experts of VIR worked out an efficient rapid method for determination of alkaloids by means of different chemical reagents. It was immediately published for the first time in the world with the Introduction by N.I Vavilov (Иванов et al., 1932). This publication, together with the know-how of German scientists, launched the breeding work with fodder (sweet) low-alkaloid lupin on a worldwide scale. Numerous scientists in many countries continue the traditions and methods of Vavilov. On the field of cultivated crops, especially legumes, ideas of Vavilov have been further developed in Australia (Gladstones, 1974, 1998; Sweetingham, 1986, 1989; Cowling, 1994; Clements and Cowling, 1990, 1991), Finland (Hovinen, 1994; Tigerstedt, 1994), Germany (Diederichsen and Hammer, 1996), Italy (Laghetti et al., 1996; Saccardo, 1996), Poland (Swęcicki W, 1988; Swęcicki W.K., 1996; Kazimierski and Kazimierska, 1992; 1994), Portugal (Mota, 1984; Gusmão, 1988 ; Gusmão et al., 1996; Tavares de Sousa et al., 1992; Neves Martins; 1994; Dordio, 1996), Sweden (Blixt, 1970, 1996), UK (Polhill, 1976, Bisby, 1981; Ambrose, 1996) and many others.

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N.I. Vavilov symbolizes the glory of Russian and world science and, at the same time, personifies its tragedy. In the stifling atmosphere of Stalin’s totalitarian rule, the Institute of Plant Industry headed by Vavilov turned into a bulwark of resistance against the pseudoscientific ideas of Lysenko. Because of this controversy, Vavilov was arrested and died in prison in 1943. Nevertheless, the memory of Nikolai Vavilov was honored by his followers, and preserved in his collections of plant genetic resources, ideas and books. This priceless heritage of Vavilov not long ago was chronicled in his posthumous book Five Continents (Вавилов, 1962, 1987a, Vavilov, 1997). Scientific accomplishments presented further in this publication represent physical realization of Vavilov’s ideas with reference to lupin. More than 2500 accessions of lupin stored in the collection founded by Vavilov were used as materials for our research!
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THE REVIEW OF THE GENUS LUPINUS


B.S. Kurlovich, A.K. Stankevich and S.I. Stepanova


Lupinus L. 1753, Sp.Pl.: 721-722; idem 1754, Gen. Pl. ed 5:322; Wats. 1873, Proc. Amer. Acad. Arts Sci. 8:522; Aschers at Graebn. 1907, Mitteleurop. Fl. 6, 2:220; Zhukovsky, 1929, Bull.Appl. Bot. Gen. Pl-Breed., 21, 1:253; Palibin, 1945, Fl. URSS, 11:48; Atabiekova, 1959, Bjull. Glav. Bot. Sad., 33:62; Vassilczenko, 1987, Fl. Europ. Parts URSS, 6:213.

Lectotypus of genus: Lupinus albus L.

Description

The genus Lupinus L. concerns to divisio Magnoliophyta (Angiospermae), subclassis Magnoliatae (Dicotyledoneae), superordo Rosanae Takht., ordo Fabales Nakai, familia Fabaceae Lindl. s.str. ( = Papilionaceae Giseke), tribus Luppineae Hutch. (Vassilczenko, 1987). The name of the genus descends from the Latin word lupus - wolf, which is probably connected with palmate leaflets resembling a footprint of the wolf (Palibin, 1945). This genus consists of annual and perennial herbaceous, fruticulose and fruticose species (all biotic plant forms except trees). Lupin has a tap root system. The main root reaches the depth of 1-2 m. The roots, especially the main axis, bear nodule bacteria of Bradyrhizobium sp. (Lupinus) fixing the nitrogen from air and transforming it in the arrested state. Lupin stems are grassy or arborescent, with different degrees of leafiness and terete in cross-section. Some forms have a stem distinctly fasciated in the apex. Branches are erect or erectopatent. The surface of the main stem and lateral branches is either pubescent with various degrees of density or naked with waxen tinge. Leafs are palmate compound, on long petioles connected to the stem by a cushion with elongate stipules. The shape of leaflets is variable: oval oblong, ovate oblong, obovate, narrow linear, lanceolate, lanceolate, etc. The surface of leaflets is in most cases pubescent. The inflorescence of lupin is polyanthous apical truss with alternate, semi-verticillate or verticillate arrangement of flowers. Flower corolla is zygomorphous and papilionaceous. It consists of a vexillum (banner), wings, carina (keel) and ovary with a pistil and 10 stamens. The development of flowers proceeds in cycles. In early phases the calyx forms a compressed ring consisting of five sepals. The vexillum covers the wings and keel. Initially the stamens are joined with each other, but later one of them becomes slightly separated from the other nine. Flowers of lupin do not contain nectar; however insects are attracted by their bright color, presence of pollen and a smelling liquid dispensed to the vexillum and other parts of the flower. Wings of lupin flowers have concavo-convex shape, being narrow oval, oval or broad oval. They are densely clustered on the top and sparse on each side. The carina is falcate or falcate curved, less frequently it is more or less straight, and appreciably tapering. The topmost part of the carina forms a firm beak with a small dot-like aperture for pollen and the stigma of the pistil. The basis of the carina is always open to a small extent. The carina is transparent, colorless or weakly hued. The carina’s edge can be colorless, weakly hued or intensively colored, almost black with anthocyanin. Coloring of the carina’s edge is a simple and convenient distinctive character of different lupin forms and varieties. It is taken into account in the course of breeding process and taxonomic studies. The color of the carina depends from the presence of chloroplasts, anthocyanin and others pigments. Lupin bean is orbicular or flattened, straight or curved, and slightly squeezed. Pod surface is rough, frequently with outstanding nerves; pod color is cream, brown or black. Sometimes there are interceptions formed by crumbly-celled tissue that separates seeds from each other in the pod. The pod terminates with an arched columella. Lupin seed are very diverse in size, shape and color. Seed surface is smooth or fine-meshed. The seedstalk hangs over the micropyle. The platen, where the seedstalk is located, is much thicker than the rest of the hull. The bent embryo is at the top of the cotyledons where nutrient substances are reserved. Primary true leaves are opposite, while other leaves unseen before germination are alternate.

Distribution and Ecology

There is no clear notion about the quantity of true species within the genus Lupinus L. Their number is considered to be from 100-200 up to 800-1000 and even more. This may be explained by the extensive area of lupin’s distribution and wide ecological amplitude of its habitats. Wild lupins are concentrated in two large regions: Mediterranean-African (eastern hemisphere) and American (western hemisphere). According to Gladstones (1974, 1998), there are 12 species of lupin in the Mediterranean region and Africa. They are described as 11 annual species and 1 perennial, which probably has already disappeared. Lupin grows in this region on light-textured soils, mainly at small heights or on seacoasts. Most of the species are characterized by one-year-long or frequently winter mode of life and have large seeds. This is also typical for others plants of this center of distribution. Wild lupins are located on the outcrops of parent mountain rocks (granite, crystal slates), on acid volcanic clinkers and on the soils arisen as a result of eolation (Майсурян and Атабекова, 1974). Similar soil types occur in more or less large areas in the countries of the Mediterranean pool. They have acidic reaction and are suitable for growth and development of wilds lupins. Some of the rather small quantity of species in this center are domesticated and widely cultivated in Australia and in many countries of Europe, Asia and Africa. Most widely introduced in agricultural practice of many countries are: Lupinus albus L. (white lupin), Lupinus angustifolius L. (narrow-leafed lupin) and Lupinus luteus L. (yellow lupin). L. cosentinii Guss, L. atlaticus Gladstones and L. pilosus Murr. were domesticated in Australia quite recently and are now cultivated on sandy soils with alkaline reaction. In the western hemisphere, lupins are distributed from the ocean level up to 4800 m above see level and higher, spreading from the island of Tierra del Fuego to Alaska and from Pacific to the Atlantic Ocean. The greatest diversity of species is concentrated in the sub-Alpine and Alpine zones of the Andes/Cordilleras mountain system. Lupin plays in these zones a dominant role. It is usually represented by tall perennial herbaceous, fruticulose and fruticose species, plants quite often reaching a height of 4 m and more. On more open and dryer sites with less fertile soils one can meet dwarfish forms of lupin. At foothills and on plains of America more frequent are annual and biennial lupins, many of which inhabit rather poor soils and quite droughty areas. Among them there are ephemeredes. Until now it is not clear how many indigenous species of lupin are in America. This may be explained partly by inaccessibility of the places of lupin distribution, extraordinary wide variability and weak differentiation of characters, extensiveness of the area and inconsistency of literary data. According to Index Kewensis in New World there are in total about 1000 species of lupin, according to the data of C.P. Smith (1917,1924) - 847, and according to the earlier published data of Stepanova (Степанова, 1973, 1985) – 972. However, majority of the researchers agree that the real number of the existing lupin species, excluding synonyms, does not exceed 200. Of the American species, pearl lupin «Tarwi» (Lupinus mutabilis Sweet.) was domesticated by ancient American Indians as long ago as in the 7th - 6th centuries up to BC. Nowadays it is widely cultivated on different continents. Multifoliate or Washington lupin (Lupinus polyphyllus Lindl.) is also either naturalized or cultivated in many countries. Lupin plants are xeromesopfytes by nature, being distinguished by high drought resistance. Some species grow even in the deserts of Arizona, Oregon, Texas, California and New Mexico, on the plateaus of Mexico, in the deserts of Peru and Chile, and in the oases of Sakhara. In the Mediterranean countries lupin grows on seacoast sands, on meadows and deposits, in cliff fissures with cultivated plants. The majority of lupin forms are adapted to moderate temperatures, but several North-American species tolerate even very low temperatures (L. arcticus Wats.). In the damp tropics of Brazil lupin does not occur. The plants previously described as lupin have appeared to be a species of another close genus, Crotalaria (Майсурян, Атабекова, 1974).
Domesticated lupin species have been grown as a cultivated crop in many countries on five continents of the Earth. Subsequent sections will contain more detailed information on lupin cultivation.

Origin and Evolution

Fischer and Sengbusch (1935) supposed that there were three centers of origin of different lupin species, one in the Mediterranean region and two in the American continent – in North and South Americas. Their existence on different continents has allowed the authors to surmise that these centers were secondary ones. Having analyzed the specific structure of lupin, Zhukovsky (1929) presumed that lupin is a plant of the New World and only its separate branches spread to the Mediterranean area of the Old World. The center of development of this genus, according to Zhukovsky (1929), is located in North America.

Commonly accepted supposition is that Lupinus evolved from the tropical and subtropical Sophoreae, a primitive generalized tribe of subfamily Papilionoideae (Gladstones, 1998). Sophoreae, in their turn, are believed to have arisen from a still more primitive legume subfamily, Caesalpinioideae, probably in the Palaeocene or Eocene epochs of the Tertiary, some 40-70 million years BC (Polhill et al., 1981). Concerning subsequent genesis of lupin, which is presumed to happen in the period from the middle to late Tertiary, different schools of thought diverge in their opinions.
The Northern Hemisphere Genisteae pathway. This school (Polhill, 1976, 1981a, 1981b; Bisby, 1981) places Lupinus in the tribe of Genisteae (sensu stricto). Bisby (1981) goes further stating that Lupinus should be elevated to a higher level to become monogeneric subtribe Lupininae. The remaining genera of the tribe, comprising the Cytisus-Genista complex, would make up subtribe Genistinae. Polhill (1976) and Turner (1981) suggested that Genisteae tribe evolved from the African Sophoreae via Thermopsideae, a tribe of temperate papilionoid shrubs and perennial herbs. Hence, the annual habit of the Old World lupins, together with the limited number of surviving species and wide genetic gaps between them, could be attributed to the increasing aridity from the Miocene (10 - 25 million years BC) onwards (Gladtones, 1998). The presence of lupin on both sides of the Atlantic points to early disjunction between the respective populations, but how early it happened remains unclear. Raven and Polhill (1981) supposed that Eurasia and North America remained in effective contact right up to the Miocene, via both the North Atlantic and Bering Strait. Legume seeds can be transported over quite long distances across the ocean by migrating birds, etc., due to their ability to resist water and digestive processes.
Plitmann (1981) postulated that lupin evolved in North America from a thermopsoid ancestor that had migrated earlier from the Old World, and that early species, in their turn, spread to the Mediterranean and South America by the above-mentioned long-distance transport. The latter route would have been supplemented by direct land migration once the Americas joined up about 5.7 million years BC (Raven and Polhill, 1981). The traditional diploid genetic behavior of the Old World lupin species shows that polyploidy among them, at least, must be ancient. Polyploidy was thus probably fundamental to the evolution of the genus, or of the broader ancestral group (Gladstones, 1998). The DNA studies of Käss and Wink (1998) suggest, on the other hand, that differentiation within the genus itself took place fairly recently, within the last 12-14 million years. If this is in any way close to the truth, it rules out the origin of this genus before the separation of Americas from Eurasia and Africa by continental drift. Dispersion of species or protospecies must therefore have taken place partially by long-distance seed transport across the ocean. The commonest explicit or implicit assumption (e.g. Nowacki and Jaworski, 1978; Raven and Polhill, 1981; Wolko and Weeden, 1990) is that it was directed westwards to America from Europe or Africa (Gladstones, 1998).
Opposing this idea, Cristofolini (1989) upheld the original evolution from Asian Sophoreae, along with other Genisteae, followed by independent migration of the ancestral American lupins eastwards across Bering Strait or a preceding land bridge, and of the ancestors of the Old World lupins westwards to the Mediterranean and Africa. Both of the presently existing ranges of distribution, according to this concept, may be regarded as refuges from climate deterioration in Asia itself, and have centers of independent evolution from the early stages of existence of the genus (Gladstones, 1998).
The Southern Hemisphere pathway. Dunn (1984) and Gross (1986) presented a radically different hypothesis: Lupinus originated in South America, from Crotalaria. Once included as a subtribe under the Genisteae sensu lato (Bentham, 1865), Crotalarieae are now regarded as a separate tribe, having a tropical or subtropical distribution centered largely on southern Africa, but with extension to neighboring continents (Polhill, 1976, 1981b). Lupinus, then, would have evolved from such a background, probably in South America following the Atlantic disjunction, and radiated secondarily from there. This hypothesis regards perennial simple-leaved lupins from subtropical Brazil as the ancestral form. It sees compound leaves and the am habit as evolving later, perhaps as opportunistic adaptation to shortness of seasons or aridity in the neighboring regions, such as the Andean Highlands. Migration to North America could have taken place both earlier, by long-distance transport, and later, directly by land. Evidence of the former is observed in the presence of two simple-leaved species in southeastern USA (Gladstones, 1998). Dunn’s (1984) consideration that Mediterranean lupins were introduced from South America by humans must surely be discounted, despite certain similarities between species. It is possible that the opposite thing occurred, i.e. that modern Mediterranean forms were introduced by early settlers and some became naturalized in South America. Either way, Zhukovsky (1929) considered L. linearis to be so much like L. angustifolius as to be probably identical with it. More detailed evidence is needed to settle this question. Presumable descent of Lupinus from Crotalaria would require large changes in the chromosome number. Goldblatt (1981) listed Crotalaria as having 2n = 14-16 and Crotalarieae in general as having 2n = 14 - 18. Little is known of the cytology of Atlantic forms of South American lupin species, but the chromosome numbers of the North American and Andean species are known to range from 2n = 36 to 2n = 96, with 2n = 48 being the most widespread. Dunn (1984) and Gross (1986) postulated the necessary changes by aneuploidy in the base number from 7 to 6, followed by repeated chromosome doubling associated with the evolution of annual and compound-leaved plants and adaptation to cooler or more arid climates. We believe that in both suppositions there is a grain of truth and something to speculate about and that they should not be disregarded when a synthesized hypothesis is developed.
Common ancient ancestor of lupin, which existed up to the divergence of continents (our hypothesis). We suggest a hypothesis of genesis and distribution of lupin arising from the geological past of our planet. Many researchers associate the evolution of life on the Earth with the theory of a drift of continents (Вегенер, 1925, Wegener, 1966). The modern theory of tectonic plates differs in several important aspects from that of Wegener's continental drift, however the former was developed on the basis of the latter (Dietz and Holden, 1972; Ушаков, Ясманов, 1984). According to the data of the above-mentioned authors, approximately 180 million years ago the ancient supercontinent of Pangea divided into two large parts – Lavrasia and Gondvana (Fig. 4A). About 135 million years ago there came splitting of Gondvana (Fig. 4B). As a result, South America was carved out of Africa. Nearly 50 million years ago partition of Lavrasia took place (Fig. 4C) which led to disconnection of North America from Europe. During the last 40 million years further extension of the basin of the Atlantic went on. In this period, South America was joined with North America and the continents acquired their modern configuration. There were isolated conditions on the continents, but because of those shifts the climate changed, which was mirrored in the evolution of florae. These epicyclical factors in combination with sporadic changes of solar radiation induced mutations of plants. They caused not only their extinction, but also uneven development. On different continents changes occurred differently and resulted in broad variability of florae. According to this hypothesis the unified primary center of formation for most ancient ancestors of lupin probably existed in the Cretaceous and subsequent periods until the splitting of continents initially on Pangea and afterwards on Lavrasia and Gondvana. Further on, as a result of subsequent division of the continents, the development of remote ancestors of lupin continued independently on the divided parts. It was homologous but not identical, pursuant to the Vavilov’s law (1920). As reported in publications (Ушаков, Ясманов, 1984), fast development and seizure of new biotic spaces by angiospermous vegetation started in the Cretaceous period. At that time there were many families of angiospermous plants, including the family Fabaceae Lindl. Proceeding from this thesis it is possible to surmise that a considerable proportion of ancient progenitors of lupins appeared also in the Cretaceous period in Lavrasia, and predominantly in its western part which subsequently turned into North America. It is confirmed by today’s presence on the American continent of a large number of species with more primitive features from the evolutionary viewpoint (long-term cycle of development, predominantly small-sized seed, monopodial branching). Because of the disjunction of continents the main part of the territory covered by the progenitors of lupin went to North America, while the second smaller part was left with Europe. This supposition is attested to by the presence of homologous series in heritable variation in the presently existing lupin forms in both hemispheres. Phylogenic proximity between them is confirmed by the combination of such attributes as coloring of seed, flowers and vegetative organs, and also by the parallelism in their age, pathological, mutational and hybridological variability. Further distribution of lupins or their progenitors from the north to the south, in particular from North America into Central and even South America and from Northern Europe into the countries of the Mediterranean and Africa, could have been provoked by periodically recurring changes of climate, chill and thaw, seismic activity or oscillations of the magnetic field. On the other hand, it is possible to explain the distribution of lupin in the indicated directions and their dissimilarities on different continents (in particular in Europe and Africa) by the previous drifts of the continents. Distribution and divergence of lupin progenitors could take place when Pangea divided into two large parts – Lavrasia and Gondvana – and later after the splitting of Gondvana. This problem provides free play to one’s fancy. When the continent drifted apart, however, lupin ancestors found themselves in isolated environments and started their independent development. The species of lupin currently existing in both hemispheres already emerged after the splitting of the continents. It explains the existence of two absolutely isolated lupin groups differing in morphological characters, development cycles, sets of chromosomes and having a genetic barrier against crossing.
Thus two secondary centers of formation of different lupin species were derived, one of which was in the Mediterranean area and the other one on the American continent. In the process of further development their areas underwent certain changes. In particular, the essential influence on the southward migration of vegetation was produced by the fall of temperature that preceded glaciating. With subsequent thawing plants step by step regained the areas freed from ice, although they remained in the former habitat as well. This phenomenon calls for a reference to the Vavilov’s work (Вавилов, 1987б) where he explains why such high concentration of florae is observed southwards of North America. According to Vavilov, it was determined by favorable environments and conditions that neutralized the effect of the glaciers which were moving from the north and exterminated all plant diversity in the most part of North America’s territory in the last geological epoch. Besides, a considerable influence on the distribution of species and their concentration in separate locales was rendered by the processes of orogenesis, which brought about contrasting environmental factors. Part of the species settled between mountains, while other ones came closer to mountain tops. For this reason, a large number of New World lupin species massed up in the mining belt of the Andes along their whole length. Practically the same climate-forming and geological processes went on in the European part. Migration of lupins or their progenitors to the south of Europe as well as into Asia and Africa could be influenced by the dehumidification of the Mediterranean basin area that took place about 5 million years ago and by the glaciation of the northern part of Europe. Initial progenitors of Lupinus genus developed in the western hemisphere in the conditions of a moderate climate (Gladstones, 1974). Predisposition of present-day annual lupins of the Old World to sedimentary and crystalline volcanic soils indicates that the center of their initial development was the pre-Cambrian massif of Central Sahara. Subsequent processes of mountain formation and increasing aridity of the climate resulted in desiccation and dehumidification of the ancient tertiary sea Tetis and the whole area of the ancient Mediterranean. This led to the loss of the subtropical flora, including the majority of lupin forms. Their ancient habitat, already with new arid conditions, was retained by solely two species, L. digitatus Forsk. and L. palaestinus Boiss., but the others had to retreat to narrow niches in outlying areas. Morphological and cytological links with the present-day lupin forms of the Old World could hardly be discerned. They can be considered as the products of the earlier differentiation of genera. Their small quantity testifies to the extinction of many species. Purely Mediterranean lupins developed more recently than African ones (apparently after the tertiary age). Old World lupins have clear genetic barriers and different chromosome numbers. Typical Mediterranean species (with smooth seeds) have more or less broad areas and wider morphological variability. The species of the African group (with rough seeds) are relicts and occupy very limited areas. All Mediterranean and African species are adapted to rather scanty non-carbonaceous soils, which is peculiar to most of tropical and subtropical legumes characterized by nitrogen-fixing ability with the help of Bradyrhizobium sp. (Lupinus) bacteria. Owing to their symbiosis with microorganisms, these lupins are tolerant to heat, acid soils and sand. Cultivated lupins from the Mediterranean area (white, yellow and narrow-leafed forms) negatively react to an increased content of carbonates in them. In the Mediterranean countries where dominating are the soils that demonstrate alkaline reaction of soil solution, wild forms of these species occur on the outcrops of acidic rocks (granites, gneisses, crystalline shales, volcanic long walls) and on the soils composed of the products of their weathering (including sand). In view of this, it seems possible to regard the mining site where a wild form occurs as a primary area of its origin. More ancient in this region are such infrequent connate lupins as L. digitatus Forsk. occurring in the oases of the Sahara and L. princei Harms growing in the mountains of Kenya, Ethiopia and Tanzania. Cultivated L. albus, L. luteus and L. angustifolius may be regarded as younger secondary species. Almost all Mediterranean lupins are annual, but among them some winter forms may be found. The American group of lupins is undergoing the process of segregation. In North and South Americas there are many annual and perennial species that quite often grow together. There is a set of ecotypes adapted to different soil and ecological conditions. Lupin species here are often poorly discernible from one another and easily cross between themselves. Thus, optimal conditions (moisture, heat, substratum, etc.) for their development are observed in mining areas of the tropics between the latitudes of 50°-55° north and south. Within these limits the greatest diversity of wild lupin forms is concentrated. Plants of American lupin are powerful and tall mesophytes and diploids. Most of them have obviously reached us in their initial shape with almost unchanged appearance. Processes of mountain formation here were not so disastrous as in Eurasia and Northern Africa. Relief of the mines, presence of natural isolators and variability of environments influenced the partition and differentiation of lupins. Different species of lupin gradually migrated from mountains to lowlands toward more moderate, cold and arid areas, thus spreading far northwards and southwards as far as the zone of tundra. The Andes served as the arena for the current intensive formation of lupin. This mining massif was an ancient center of genesis of many species and one of the centers of origin of cultivated plants (Вавилов, 1965b; Синская, 1969). Peru, Bolivia and Ecuador represent the most ancient center of primary agriculture in the New World. Until now the ancient culture of Andean pearl lupin (L. mutabilis Sweet) was preserved at the altitude of 3200-3400 m above sea level. This species occurs there also as wild plants in natural communities. Also widespread in North America is multifoliate or Washington lupin (L. polyphyllus Lindl.). Its area of distribution stretches from California northwards up to Alaska. It is distinguished for frost resistance and large yield of green matter. This lupin has been domesticated in European countries for green manure and as a fodder plant (sweet forms). This species was also naturalized in Belarus and other countries. American lupins have preserved all traits of their ancestors and relationship with the most ancient part of the genus. Lupins from the Old and New Worlds have different caryological, embryological and biochemical characters.
According to the number of chromosomes, they make two geographically oriented groups located in conformity with their polyploidy series. American lupin forms are somewhat less specialized than Mediterranean ones. They are characterized by a more primitive monopodial type of branching and by the cross-pollination habit (with the help of insects). Their plants are heterozygous and easily generate all possible mutations. Seeds are small, the embryo is gently differentiated, it is insufficient in endosperm and located close to the seed cover, with a lengthened hypocotyl. Rather specialized Mediterranean lupin forms are characterized by a more advanced type of branching (sympodial); for them, self-pollination is dominating. Seeds are larger, the embryo is well-formed, with two sedentary leaves or a very short hypocotyl. Lupins from the Mediterranean region and those from America are also contrasting in storage proteins, each of the groups having determinants specific for their native hemisphere. These distinctions between lupins of the two continents have induced us to search for a more accurate definition of their systematic position in the generic system. Considering the two secondary centers of formation of different lupin species identified in the Mediterranean areas of Europe and Africa and on the American continent and essential morphological differences between lupins of the two hemispheres, we made revision of two subgenera in Lupinus L. genus according to geographic principle. In our reclassification, subgen. Platycarpos (Wats). Kurl. incorporates numerous perennial and annual species of the western hemisphere, both types having two, four or more seedbuds in ovules, and subgen. Lupinus L. – 11 species from the Mediterranean region and Africa with four or more seedbuds.
Cultivated in Mediterranean lupins (L. albus, L. luteus and L. angustifolius) belong to the type of plants which may be characterized by only partial transition to domesticated conditions in the centers of their formation (Е.Н. Синская, 1969). Under cultivation all of them retained the customary structure and conventional large regional ecotypes typical for subspecies. White lupin was the first to be cultivated. It was the only cultivated lupin in the Ancient Greece. Its large seeds, well soaked, strongly salted, seasoned with vinegar and vegetable oil, served as food for humans. They also were used as forage for cattle. In Georgia L. albus was grown under the name of hancholy. This species was described in Egypt for the second time after Linnaeus as L. termis Forsk. In ancient Egypt and Palestine it was called thurmus, which means “hot”. Now Egyptian representatives of this lupin species differ from Greek forms of L. albus only by thinner stalks, small-sized seeds, blue flowers and early flowering. This circumstance makes it possible to rank L. termis as a subspecies within L. albus L. For a long time L. albus was regarded as a crop plant, which had lost connections with its wild relatives. However, we have got accessions with colored flowers and small-sized dark-brown mottled seed, earlier widely distributed in Greece (including Grecian islands) and described as L. graecus Boiss. et Reut. There are certain grounds to estimate them also as an initial subspecies for cultivated L. albus L. This subspecies occurs in the wild in the form of one-, two- and four-year plants with a leaf rosette, intensively colored flowers and smaller-sized dark-brown mottled seed. It is freely crossed with the cultivated forms of L. albus L. The Mediterranean lupin species are difficultly or cannot cross with each other and cannot cross with the American species. That testifies to the absence of close relationship between them. Successful crossing in such pairs of annual American species as L. mutabilis Sweet with L. ornatus Dougl. or L. pubescens Benth. confirms that they belong to two closely related, though geographically isolated groups. The annual species L. elegans H.B.K. is easily crossed with perennial L. mexicanus Cerv. (= L. hartwegii Lindl.). Apparently, they are also closely related, being, however, isolated in their altitudinal distribution. Difficulties in crossing perennial L. arboreus Sims. with L. mexicanus Cerv. and L. polyphyllus Lindl., experienced by Kazimierski and Novacki (1961a), prove that geographic isolation of these forms has resulted in certain genetic changes. It is possible to regard lupin as a primitive crop plant for the reason that it poorly grows on freshly plowed fields and its plantings do not require special maintenance. It usually prefers acid soil and sandy underground.

Only two species (L. albus L.) and (L. mutabilis Sweet) can be considered as completely domesticated. They are large-seeded species developed through thousands of years of selection. Ancient people had collected seed from plants of these species, sowed them in the disturbed soils around their dwellings and bred them. Other lupin species presently under cultivation are directly linked with wild forms. They have preserved the tendency of fast forfeiting their cultural characters when their selection terminates or becomes less intensive. Analysis of geographic and ecological distribution of lupin species with the help of N.I. Vavilov’s differential systematic geographical method of crop studies (Вавилов, 1926, 1931, 1935) provides for more accurate identification of the centers of development and origin of different lupin forms (Kurlovich, 1998).
The center of formation of wild white lupin (L. albus) and the primary center of origin (diversity) of its initial cultivated forms is the Balkan Peninsular, where an exceptionally rich diversity of wild forms, including those that turned wild, and local ones are concentrated. Three subspecies of white lupin (subsp. graecus Franko et Silva, subsp. termis Ponert., subsp. albus) grow in the Balkans. This region also harbors wild forms with dark-brown mottled seeds and dark-blue flowers (subsp. graecus). The Apennines and Egypt are also centers of origin of cultivated white lupin forms that have been cultivated there since the earliest recorded history. Moreover, widespread in the ancient Egypt were the forms with pink-and-blue or light-pink flowers (subsp. termis). The Apennines was abundant of the forms with grayish and light blue or white flowers (subsp. albus).
In the Pyrenees exist two closely cognate species (L. luteus and L. hispanicus Boiss. et Reut) having the same number of chromosomes (2n=52). There is wide diversity of wild and cultivated forms of yellow lupin (L. luteus) with a long historical period of growing there. All this supports the assumption that the Pyrenees was the center of formation of the wild forms of yellow lupin and the center of origin of its cultivated plants.
The Pyrenees are probably also the center of formation and origin of the blue lupin (L. angustifolius), because this peninsula shows the highest concen­tration and the widest diversity of its forms. As a result of wider utilization of lupin in agriculture, its area of cultivation gradually increased leading to the forma­tion of secondary macro- and microcentres of origin of culti­vated lupin forms. These correspond to the many geotypes we have described. Our researches on these problems are considered in more detail in following sections.

The history of study of the genus Lupinus L. and its species

Lupin was known long before C. Linnaeus; the botanists of the 16th -17th centuries wrote about it. Lobel (1591) mentioned three species: L.sativus (= L.albus L.), L. flore purpureo (= L.angustifolius L.) and L. flore luteo (= L. luteus L.). Clusius (1601) nameed L. albus, L. angustifolius and L. luteus. Bauhin (1651) listed four species: L.vulgaris semine et flore albo sativus (L. albus), L. exolicus hirsutissimus (L.micranthus), L. luteo florri semine compresso, vario (L. luteus) and L. sylvestris, purpureo flore, semine rotundo vario (L. angustifolius). Tournefort (1719) presented a description of the genus Lupinus and mentioned a number of species known at that time. Linnaeus (1753) described six species: L. albus L., L. angustifolius L., L. hirsutus L., L. perennis L., L. varius L. and L. luteus L. All of them, except L. perennis, represent Mediterranean lupins. Afterwards, Mediterranean species of lupin were described by Lamarck (1778, 1789) and Candolle (1825). Study of lupin forms from the New World began later. Lindley (1827, 1837) provided description of the following species of American origin – L. bicolor, L. polyphyllus, L.hartwegii and L.barkeri, – which became quite popular. In 1833-1859 the works of Bentham (1833, 1835, 1839, 1859) were published. He described a number of species, among which the most known were L. hilarianus, L. hirsutissimus, L. densiflorus and L. pubescens, cultivated as ornamental plants. Such species as L. albifrons, L. arvensis, L. bilineatus. L. bogotensis, L. caespitosus, L. coriaceus, L. foliolosus, L. humifusus, L. interruptus, L. laevigatus, L. lanatus, L. leptocarpus, L. leptophylius, L. lucidus, L. macrophyllus, L. ramosiseimus, L. sparsiflorus, L. subsessilis, L. tauris, L. vaginanus and L. velutinus, described by the same author, were less known, being in most cases synonyms of the species already described. For example, L. sparsiflorus Benth. is synonymous to L. micranthus Dougl., L. bilineatus Benth. to L. Hartwegii Lindl., L. macrophyllus Benth. to L. polyhypllus Lindl., L. albifrons Benth. to L. chamissonis Eschsch. The research works of Douglas (1828, 1829, 1835 and 1861) published from 1828 to 1861 in various editions are referred to the same period. Among the species described by him widely known in horticulture became L. ornatus Dougl., L. succulentus Dougl. and L. nanus Dougl., now represented in agriculture by a considerable number of varieties and forms. L. succulentus Dougl. is a synonym of L. Barkeri Lindl. However, the basic monograph dedicated to the analysis of genus Lupinus was the work of Agardh (1835), «Sinopsis Generis Lupini», based mainly on the materials from Lindley’s herbarium. In this general study the author provided detailed description of 83 lupin species, of which 7 were placed in the list «Species Inquireude».
Agardh (1835) analyzed the following species: L. affinis, L. aureus, L. concinnus, L. cytisoides, L. Douglassii, L glabratus, L. gracilis, L. Gussoneanus, L. Hornemanni, L. Kunthii, L. latifolius, L. leucopsis, L. Lindleyanus, L. Menziesii, L. Milleri, L. prostratus. L. rivularis and L. stripulatus. Within the genus Lupinus L. he identified ten tribes: L. lutei, L. pilosi, L. albi, L. angustifoli, L. polyphylli, L. nootkatensi, L. arborei, L. serigei, L. paniculati and L. microphylli. Some of the indicated species differed by integral leaves and were later referred to another tribe, Genisteae. As to the characteristics of the above-listed species, Agardh regarded many of them as synonymous or closely related to the already known ones, marking the identity or likeness of their traits. However, in his search for an objective overview of lupin systemization he was hindered by the convictions of other botanists with authority. Nevertheless, Agardh’s researches were based on a diversity of materials and presented very careful descriptions of plants, so they have exclusive value. His major mistake was complete neglect of the geographic principle. For example, in the same tribe of L. lutei he included, on the one hand, L. luteus which originated in the eastern hemisphere and, on the other hand, the species from the western hemisphere, L. Menziesii Agardh and L. densiflorus Behth. The tribe of L. angustifolii harbored spp. L. angustifolius L. and L. linifolius Roth. together with spp. L. nanus Dougl. and L. leptophylius Benth., notwithstanding different genesis of both pairs. The tribe of L. albi also incorporated lupin forms of both New and Old World’s origin. These circumstances, however, do not deprive Agardh’s monograph of its value; this work of his set up a solid foundation for botanical analysis of Lupinus L. genus. By the end of 1835, that is by the time when Agardh’s monograph was published, about a hundred of different lupin species were reported to have already been described. During the whole 19th century, as well as in the beginning of the 20th century, new and new descriptions of different lupin species appeared.
Among the most significant achievements worth mentioning are the works of Watson (1871, 1873 and 1890) who managed to present descriptions of 56 lupin species from various areas of the extra-tropical North America. Besides, the genus Lupinus L., and in particular its North-American species, were rearranged by Watson (1873) into three parts: Lupinus, Platycarpos and Lupinnelus. Differences in habit and in the number of ovules were accepted as the criteria for such classification. Most of the perennial and annual species from the American continent described by Watson were referred to Lupinus. The section of Platycarpos included some annual species with two ovules and seeds in the pod (L. densiflorus Benth., L.micricarpus Sims. and others). Lupinnelus section consisted of one species (L. uncialis) having axillary and solitary flowers, scarcely reflexed banner and also two ovules. The existence of such species now seems doubtful.
The same classification principle was later in the studies of Ascherson and Graebner (1906-1910) extended to all lupins from the eastern and western hemispheres. In the beginning of the 20th century this generalized arrangement of lupin diversity was the fullest and most detailed. It is often referred to by modern experts in plant taxonomy. This work recognized the achievements of the previous explorers (Lindley, Agardh, Watson, Boissier) and provided descriptions for most of the species known by that time in both eastern and western hemispheres, as well as the keys for the species’ definition. Our research is based on it as well. The genus Lupinus L. was then for the first time divided into two subgenera: A. Eulupinus and B. Platycarpos (Ascherson and Graebner, 1906-1910). This work was a starting point for our investigations. Taking into consideration the definition of two secondary centers of formation of different lupin species in the eastern and western hemispheres as well as essential morphological differences between lupin forms in both hemispheres, we managed to revise the volumes of two subgenera in the genus Lupinus L. according to the geographic principle. In our combination, subgen. Platycarpos (Wats). Kurl. incorporates numerous perennial and annual species of the western hemisphere, both groups having two, four and more seedbuds in the ovary, while subgen. Lupinus L. covers 11 annual species from the Mediterranean region and Africa with four and more seedbuds in the ovary.

Heller published his works «The North American Lupines» and «The Nevada Lupines» after 1900. He described 59 species, of which 17 were virtually new (Heller, 1905a, 1905b, 1906, 1912).
Beginning from 1917 Smith’s publications were coming forth. During a number of years his studies were published in the bulletins of Botanical Torr. club under the title «Studies in the genus Lupini» (1918-1924). This author described 56 new species and 33 new varieties. In 1925, Smith started to publish definitions of the species discovered by him in other periodicals. All these materials were united in his general monograph «Species Lupinorum» published in separate booklets from 1938 through 1953. They contained descriptions of different species from Mexico, North and South Americas, and also Mediterranean species, based of his study of herbarium materials and references. This research subsequently led to a real mess in the systematics of lupin from the western hemisphere. In the first issue of his monographic work, Smith made a catalog of the published names of lupin forms presented as species. There were more than 500 species (Smith, 1938). Since then this number has grown up and is still growing. A closer view on the materials of some species shows that there were dozens of synonyms. The latter circumstance has very strongly hindered the determination of their exact names. It can be also explained by the fact that till now there are explorers called ”splitters” who generously dispense the special title of “the species” to such forms that are rather poorly distinguished (Атабекова, 1962).

D. Dunn was for a long time engaged in studying lupins of North America. He made revision of the North American species, including Mexican ones (Dunn, 1955). As a result of his research, he made serious amendments and changes to the nomenclature of the genus Lupinus. In one of his works (Dunn, 1957) he included in a complex species L. arbustus Dougl. ex Lindl. described by other authors as independent. For example, L. laxiflorus sensu Agardh, which had been described by Agardh as a species, he attributed to L. arbustus subsp. neolaxiflorus, considering it to be a subspecies or a variant. All in all, he included in the named L. arbustus 6 previously described species. In his following work (Dunn, 1965) he revealed the links of relationship between L. arcticus, L. nootkatensis and L. polyphyllus. Research showed that all of them had hybrid origin, were very close to each other and constituted the most northern species of the genus. Their description and distribution were also presented in this work. In the next publication (Dunn, 1966) new names and synonyms of species included in the Lupinus concinnus - L.sparsiflorus complex were introduced: L.concinnus Agardh (subsp. concinnus), L. concinnus subsp. optatus (C.P.Smith) Dunn comb. nov., L. concinnus subsp. orcutti (Wats). Dunn comb. nov., Lupinus agardhianus Heller, L.pallidus Brandegee, L.brevior (Jeps). Chistian et Dunn comb. nov., L.sparsiflorus Benth. (subsp. sparsiflorus), L.sparsiflorus subsp. inopinatus (C.P.Smith) Dziekan. et Dunn comb. nov., L.sparsiflorus subsp. pondii (Greene) Dziekanowski et Dunn comb. nov., L.sparsiflorus subsp. pondii (Greene) Dziekan. et Dunn. Dunn (1984) described tentative group­ings of the 500-odd New World taxa, which he recognized as warranting descrip­tion at the specific or subspecific levels. The best studied are those of North and Central America.

Further efforts in analyzing and revising the described numerous lupin species in the American continent were undertaken by Planchuelo (1984, 1994, 1998). Planchuelo and Dunn (1984) made public the beginnings of work with the hitherto poorly documented lupins of South America. In Planchuelo’s next publication (1994) the areas of Lupinus (both in the eastern and western hemispheres) were determined by taxonomic studies, extensive bibliographical review and field trips. The centers of species formation were classified into three regions, each of them having two subregions, as follows: North and Central American region with South-Eastern (SEN) and Western Mountain Range (WMR) subregions; South American region: divided into Atlantic (ATL) and Andean (AND) subregions; and Mediterranean and African region with European Mediterranean (EUM) and African Mountainous (AFM) subregions. Geographic delimitation of these regions and subregions is presented in the world map along with approximate estimation of the number of species present. Possible relationships between wild and cultivated species are also pointed out. However, this complex problem is far from being completed. Nobody can yet make a precise summarization of the number of Lupinus species existing now in reality in the western hemisphere. Large variability of lupin-growing places and conditions and entomophilous nature of pollination account for the presence of exceptional diversity in the forms of this plant on the American continent. The situation seems much easier with botanical research on lupin from the eastern hemisphere. Studies by Merkenschlager (1928-1929), Zhukovsky (1929), Fisher and Sengbusch (1935), Gladstones (1974, 1998) made it possible to outline the distribution of wild lupin species in the zone of the Mediterranean Sea and Africa.
According to Gladstones (1974), there are 12 species of lupin in the eastern hemisphere, among which 11 are annual and one (most likely already extinct) perennial. However, the way toward this conclusion was difficult enough and full of contradictions. Study of lupin diversity of Mediterranean origin was also undertaken by Boissier (1842, 1843, 1849, 1872). In different times he made descriptions of L. hispanicus (Boisser and Reuter, 1842), L. graecus (Boisser, 843), L. palaestinus, L. Forskahlei and L. philistaeus (Boisser, 1849). Subsequent investigations clarified that L. Forskahlei is only a subspecies of L. pilosus L. and that L. philistaeus is one of the forms of L. angustifolius. Boissier’s (1872) research efforts were finalized in his «Flora Orientalis», in which he offered the description of ten species: L. albus L., L. angustifolius L., L. digitatus Forsk. L. graecus Boiss. and Sprunn., L. hirstitus L., L. hispanicus Boiss. Reut., L. palaestinus Boiss., L. pilosus L., L. reticulatus Desv. L. termis Forsk., and provided definition of three new species (L.graecus Boiss, L.hispanicus Boiss. and Reuter and L. palaestinus Boiss.). There was no L. luteus L. in this list, though further in the text the author identified the black-seeded form of yellow lupin as «Siberian» in spite of the fact that, in the author’s opinion, it had nothing to do with this geographic part of the globe. Research on the Mediterranean species was continued in the twentieth century by Hegi (1924).
Zhukovsky (1929) gave description to only most widespread species from the eastern and western hemispheres. Thus, he raked L. termis Forsk. as a subspecies of L. albus L. and L. digitatus Forsk. as a variant of L. pilosus Murr.
Libkind’s book „Lupin“ (Либкинд, 1932) analyzed the results of the research conducted on lupin collection accessions grown for a number of years at the Byelorussian and Ukrainian branches of the Institute of Plant Industry (VIR). On the basis of a cycle of observations she identified many new varieties and ecogeographic groups of ecotypes in the diversity of lupin forms. These investigations of hers made up a starting point also for our further research. Some information on the botanical analysis of lupin is contained in the book by Sharapov (Шарапов, 1935) where there is a special section with the description of the genus Lupinus and biological and botanical characteristics of its major species.
R. Mansfeld (1959) acknowledged the existence of the following species: Lupinus luteus L., L. albus L., L. termis Forsk., L. angustifolius L., L.hirsutus L., L. digitatus Forsk. and L. varius L. Kazimirski and Novacki (1961b) conducted detailed study of seeds and morphological attributes of 12 Mediterranean lupin species and finally came to the conclusion that only four species seemed indisputable: L. albus, L. angustifulius, L. luteus and L. rothmaleri. Other species (for example L. digitatus, L. pilosus, L. hirsutus and L. palaestinus) were considered by these authors debatable. On the basis of their studies and published data they united the species of Mediterranean origin in four sections: 1) species with yellow flowers - L. luteus and L. rothmaleri, 2) with blue flowers - L. angustifolius with subsp. L. linifolius Roth., 3) with white flowers - L. albus with subsp. L. termis Forsk., L. graecus Boiss. and L. jugoslavicus Kazim. et Novacki. 4) rough pubescent species - L. pilosus Murr., L. hirsutus L., L. digitatus Forsk. Others species, namely L. palaestinus, L. hispanicus Boiss. et Reut. and L. varius - they left outside of the sections for lack of data. Then, having continued their investigations on lupin together with Aniol, they found out that L. albus could be easily crossed with L. graecus Boiss. and L. termis Forsk. With this in view, they concluded that these taxa are only the subspecies of L. albus sensu lato (Kazimierski et al.,1968). They divided L. albus into five ecological subspecies: L. albus jugaslavicus, L. albus graecus, L. albus termis, L. albus albus, L. albus megalospermus.

Atabiekova (Атабекова, 1962a) made detailed characteristics of four cultivated species (L. albus L., L. angustifolius L., L. luteus L. and L. pilosus Murr.) with their intraspecific classification. She described the wide intraspecific diversity of the mentioned species and identified many varieties within them. However, this work did not follow the rules of the International Code of Botanical Nomenclature, so most of the names she gave to such varieties appeared invalid. In her other publications (Атабекова, 1962b) she provided a detailed review of the literature dedicated to systematization of lupin. Plitman (1966) published a description of the following lupin species: L. pilosus Murr., L. palaestinus Boiss., L. hirsutus L., L. angustifolius L. and L. termis Forsk. Within L. angustifolius he mentioned var. basalticus Zoh. et Plitm. with rather wide leaflets. Plitman considered L. linifolius Roth. to be the synonym of L. reticulatus Desv., the former representing by itself only a form within sp. L. angustifolius L. identified by narrow leaflets and pods. Plitman placed L. termis Forsk. among independent species. Having compared herbaria of L. termis and L.albus, he found out that they were not only similar in many respects, but also presented clearly expressed differences traced in the shape of inflorescence, flowers, petals and calyxes. Their areas of distribution were also set apart. However, in our opinion, these data are not enough to regard L. termis as an independent species. It is only one of the subspecies within L. albus L. Franco and Silva (1968) described Mediterranean species in their «Flora Europaea» and made a number of taxonomic changes. This work contained descriptions of L. luteus L., L. hispanicus Boiss. et Reuter, L. angustifolius with subsp. angustifolius and subsp. reticulatus (Desv). Coutinho, L. micranthus Guss., L. albus L. with subsp. albus and subsp. graecus (Boiss. et Sprun.) Franco et P. Silva and L. varius with subsp. varius and subsp. orientalis Franco et P. Silva (synonyms of L. digitatus Forsk. and L. pilosus Murr.). Species inhabiting Africa were not mentioned in this publication at all. Chamberlain (1970) presented description and classification of the following Mediterranean species: L. albus L., L. angustifolius L., L. micranthus Guss., L. varius L. and L. hispanicus Boiss. et Reuter. In L. albus L. he selected two varieties with synonyms: subsp. albus (L. termis Forsk.) and subsp. graecus (Boiss. et Sprun.) Franco et P.Silva. He considered Plitman to be wrong in regarding L. albus subsp. albus and L. termis as different species. He reckoned L. termis to be synonymous to L. albus subsp. albus, seeing considerable morphological likeness between them and relying on the data of «Flora Europaea». Maissurjan and Atabiekova (1974) guided by Agardh’s (1835) basic classification principles integrated all known species of lupin from the Mediterranean region and America into tribes. They classified Mediterranean lupin in four tribes: L. lutei, L. pilosi, L. albi and L. angustifolii. Each tribe included corresponding species: tribe L. lutei – spp. L. luteus L., L. hispanicus Boiss. et Reut., L. rothmaleri Klink. and L. palaestinus Boiss.; tribe L. pilosi – L. pilosus Murr., L. digitatus Forsk. and L. hirsutus L.; tribe L. albi – L. albus L., L. termis Forsk., L. vavilovi Atab. et Maiss. and L. graecus Boiss.; tribe angustifolii – L. angustifolius L., L. linifolius Roth. and L. opsianthus Atab. et Maiss. Maissurjan and Atabekova (1974) considered L. graecus Boiss., L. linifolius Roth. and L. rothmaleri Klink as genuine botanical species of the Mediterranean area in view of the presence of clearly distinctive traits in them. Many of the botanists, as was already indicated here, considered L. graecus Boiss. a subspecies, variety or form of L. albus L., L. linifolius Roth. a subspecies of L. angustifolius L., and L. rothmaleri as having very much in common with L. hispanicus Boiss. et Reut. Gladstones (1974) revised the diversity of lupin species from the

Mediterranean region and Africa and included one new species, L. atlanticus Gladstones. He states that though there are few lupin species in the Mediterranean countries and Africa, their taxonomy for a long time has been too complicated. Gladstones identified the following species: L. albus L., L.angustifolius L., L. micranthus Guss., L. luteus L., L. hispanicus Boiss. et Reut., L. cosentini Guss., L. digitatus Forsk., L. princei Harms, L. pilosus Murr., L. palaestinus Boiss., L. atlanticus Gladstones and L. somaliensis Baker. He pointed out many errors in the previous descriptions of the listed species and described the same species anew mentioning all their synonyms and introducing corrections and changes in the taxonomy of the Mediterranean lupin species. Gladstones (1974) indicated two varieties in L. albus L.: var. albus and var. graecus (Boiss. et Sprun.) J. S. Gladstones. The majority of botanists had previously regarded L. graecus as a separate species. In «Flora Europeae» it held the rank of a subspecies within L. albus L. White lupin (L. albus L.) is a crop plant cultivated in many regions. However, according to Gladstones (1974), it is necessary to regard graecus as the wild form of L. albus L. We completely agree with this judgement, as albus differs from graecus in its morphological and physiological traits that have appeared as a result of selection. For instance, albus is characterized by indehiscent pods, water-permeable seed coat, large seed of white colour, more straight growth habit and early flowering (similar changes were obtained by breeders in other lupin species). However, we suppose that attributing to graecus the rank of a subspecies would be more correct, as it has a definite area of distribution. In Gladstones’ description of var. graecus (Boiss. et Sprun.) J. S. Gladstones the author enumerates its synonyms: L. graecus Boiss. et Sprun. (1843), L. jugoslavicus Kazim. et Now. (1961), L. vavilovi Atab. et Maiss. (1962) and L. albus subsp. graecus (Boiss. et Sprun.) Franco et P.Silva (1968). As we see, Gladstones (1974) jettisoned one more species from the Mediterranean group, namely L. vavilovi, the description of which was published by Atabekova and Maissurjan (Атабекова and Майсурян, 1962). Furthermore, Gladstones also considered L. jugoslavicus (= L. vavilovi) to look very much like L. graecus. J.S. Gladstones expands the number of synonyms of L. angustifolius L. up to L. linifolilus Roth. and L.opsianthlis Atab. et Maiss. He founds in each of them many features in common with L. angustifolius; however, both of them have finer leaflets and seeds. Among the synonyms of L. angustifolius he also named L. reticulatus Desv. and L. philistaelus Boiss., thus disagreeing with Franco and Silva (1968). J.S. Gladstones updated the systematic position of L. micranthus Guss. Having analyzed the herbarium of Linnaeus, he came to the conclusion that L. hirsutus L. was named so by mistake and this name should be rejected in favor of a later, but more definite one – L. micranthus Cuss. It would agree with «Flora Europaea» where in the description of L. micranthus L. hirsutus L. appeared as its synonym. J.S. Gladstones did not introduce any changes in the systematization of L. luteus L. In Mediterranean countries alongside with L. luteus L also occurs L. hispanicus Boiss. et Reut. Gladstones partitioned this species in two subspecies: subsp. hispanicus and subsp. bicolor. In his description of new subsp. bicolor (Merino) J.S.Gladstones the author mentioned as its synonyms the following species: L. bicolor (Merino) Rothmaler, L. rothmaleri Klink., L. hispanicus sensu auct. non Boiss. et Reut.: Franco et P.Silva, L. versicolor Caballero with varieties L. luteus var. bicolor and L. hispanicus var. bicolor Merino. In Gladstones’ (1974) opinion, there are many inaccuracies in the taxonomy of L. cosentinii Guss. Some researchers erroneously determined it as L. varius L. This error is repeated in modern European and Australian literature. Other botanists attribute it as a subspecies to L. pilosus Murr., but it is not correct either. As it has become known, L. pilosus (n = 21) differs from L. cosentinii (n = 16), and crosses between them yield only non-viable seeds. Some authors considered that L. cosentinii is a synonym of L. digitatus Forsk., but it has appeared that the latter was described earlier. Gladstones gave a new status to these two taxa by placing them to the level of independent species: L. cosentinii Cuss. and L. digitatus Forsk. L. pilosus Murr. is also a chaotic species and it is not easy to disentangle its taxonomy. Gladstones named the following synonyms thereof: L. hirsutus L., L. varius L., L. varius subsp. orientalis Franco et P.Silva. Maissurjan and Atabekova (1974) reckoned among the synonyms of this species also L. varius L., L. silvestris Lam. and L. semiverticillatus Desr. They leave the authorship of L. pilosus to Linnaeus, though Lupinus pilosus Murr. has received almost everybody’s recognition. Until now some of the botanists go on applying the name of L. hirsutus instead of L. pilosus. It seems incorrect, as L. hirsutus (now L. micranthus Guss.) and L. pilosus are well differentiated. The reason of this confusion was that in the standard herbarium of Linnaeus the accession of L. pilosus (No 898.3) was lost and probably by accident was replaced by a sample of L. varius. The following consecutive number (No 898.4) was assigned to L. micranthus. Franco and Silva (1968) used the name of L. varius for L. pilosus Murr. (widespread in Greece, Syria, Israel, Eastern and Southern Turkey). They combined it with L. digitatus Forsk. under the common specific title of L. varius subsp. orientalis Franko et P.Silva Gladstones (1974) also described three less widespread species: L. atlanticus Gladstones, L. princei Harms and L. somaliensis Baker. L. atlanticus Gladst. is an annual species from the rough-seeded group. It has traits tightly cognate with L. pilosus, L. digitatus and L. cosentinii, but also manifests differences from them. It is widespread in Morocco, at the altitude of 900-1500 m. Thus, having conducted revision of lupin from the Mediterranean region and Africa, Gladstones (1974, 1998) numbered 12 species with this genus. He explained numerous taxonomic differences by discrepancies in previous classifications. In most cases he managed to clarify the taxonomy of the Mediterranean and African species. However, some problems remain disputable. We do not regard necessary to include in the list of Mediterranean and African species sp. L. somaliensis Baker which no one of the contemporaries noticed. There is vagueness in the taxonomy of L. princei Harms. Immunological genetic and electrophoretic research allow to specify many questions of systematic (Wolko and Weden, 1990, 1994; Swęcicki W.K. and Wolko, 1992; Salmanowich and Przybylska, 1994).
In particular, valuable research is presented in the work of Salmanowich and Przybylska (1994) about electrophoretic analysis. PAGE and SDS-PAGE electrophoretic analysis of 31 accessions of 10 Old-World Lupinus species, 5 smooth-seeded and 5 rough-seeded, covered total seed albumins and 2S albumins isolated by a solid-phase extraction. PAGE albumin patterns showed a distinct difference between the smooth- and rough-seeded lupins. SDS-PAGF analysis of seed albumins revealed interspecific differences, mainly due to the 2S albumnins. The differences were especially marked in the smooth-seeded species. In the rough-seeded lupins the fol­lowing subgroups were distinguished: (1) L. atlanticus, (2) L. cosentinii and L. digitatus, (3) L. palaestinus and L. pilosus. Evidence was provided that the 2S albumin class contains conglutin g, so far classified as a globulin. The results of the authors are discussed with reference to taxonomic relationships of the Old-World lupins and characterization of the lupin seed albumin fraction. Further development and perfecting of intraspecific classifications is required, especially for domesticated lupin species. This work presents intraspecific classifications designed by us for three species widely utilized in agriculture (L. albus L., L. angustifolius L. and L. luteus L.). These researches are based on Vavilov’s concept of the species as a certain complex system. Therefore, while studying species on the basis of Vavilov’s concepts we have taken into account not only morphological characters, but also geographical and ecological differentiation and other properties of plants (differential systematic geographical method of crop studies). Such approach based on the development of different intraspecific classifications has made it possible to conduct thorough and comprehensive investigation of intraspecific and varietal diversity of cultivated plants and to find ways of their efficient utilization. Many years of observing the plants of lupin sown in various regions have enabled us to find out new regularities in the variability of characters, depending on genetic features of species and accessions, and also on ecological and agronomic conditions of their growing. The authors of the present work array the identified rich diversity of forms in a definite system, founded on the steady combinations of several characters of generative and vegetative plant parts. As a result, subspecies, varieties, subvarieties, forms, geotypes and ecotypes have been established for the three annual domesticated lupin species. The collection of the N.I. Vavilov Institute served as a material for this purpose.

In summary, it should be mentioned here that the genus Lupinus L. is still waiting for researchers who could find out the true number of species and their intraspecific taxa within this polymorphous genus, especially on the American continent.
That will help to clear up many theoretical and practical problems.

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 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, 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 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 - 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.

,,, 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. lignipes Heller, 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 in L. 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. versicolor Lindl. 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. grandifolius Lindl. 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. cytisoides Agardh, 1835, l.: c.:81. – L. adsurgens Drev. 1889, Bull. Forrey Clab 16:150. – L. longipes Greene, 1891, Fl. Francisc.: 41. – L. viridifollius Heller, 1905, Muhlenbergia 2:65. – L. columbianus Heller, 1912, Muhlenbergia 8:84. – L. confusus Heller, 1912, ibid.:63. – L. barbatus Heller, 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. nutcanus Spreng. 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. laxiflorus Dougl. 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. argenteus S.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. foliosus Nutt. 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. ochroleucus Eastw. 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. brachypodus Piper, 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. argenteus Agardh, 1835, Syn. Gen. Lupin.:27, non Pursch, 1814. – L. leucapsis Agardh. 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. polyphyllus Lindl.

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. bimaculatus Hook. ex D. Don, in Sweet, 1835, Fl. Gard. Ser.2, tab. 314. – L. texensis Hook. 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 lower labium 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. rostrarus Eastw. 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. densiflorus Agardh, 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, 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.


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.albus var. 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.angustifolius var. 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 labeled L.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.hispanicus ssp. 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).

ECO-GEOGRAPHIC CLASSIFICATION OF LUPINS


ECO-GEOGRAPHIC CLASSIFICATION OF LUPINS (L. ALBUS L., L. ANGUSTIFOLIUS L. end L. LUTEUS L.)



B.S. Kurlovich



Effective utilization of plant genetic resources in plant breeding requires detailed information about different biological, physiological, biochemical and other properties of the collected accessions. Eco-geographic classification reflects the regularities of variability of these properties, based on the specific nature of geographic, historical, ecological and agronomic conditions. It cannot replace the botanical classification of the species, which requires clear distinction of morphological characters observed in the herbarium materials, but supplements it. Both classifications taken together provide for purposeful and conscious utilization of plant genetic diversity in breeding practice, to which N.I. Vavilov (1965, 1997) repeatedly paid attention.
Each species has the radical, i.e. genetically determined and correlated group of structural characters that may be observed in all its forms. The species is "... a flexible isolated complex morpho-physiological system linked in its genesis to a certain environment and area of distribution" (Вавилов, 1965a, p.248). Therefore, the species should be studied by morphological, physiological, biochemical, genetic and other methods.
The matter of N.I. Vavilov’s special attention and concern was ecological differentiation of a species. Many Linnaean species represent a complex system of ecotypes or climatypes. "An ecotype is a group of biotypes within one Linnaean species united by a series of constant hereditary characters and adapted to certain conditions of their habitat. It is quite natural that differing in space and submitting to the effect of selection, the basic potential of the Linnaean species is to develop and isolate the groups of hereditary forms most closely corresponding to the given environments..."(Vavilov 1965, p.245). The researches on the ecological pattern of many dozens of species were widely conducted on his initiative, as a result of which it became possible to reveal the regular nature of their variability within the limits of the areas of their distribution .
Much attention was devoted to the analysis of the species’ ecological pattern by Sinskaya (Синская, 1948, 1961, 1969) and Agaev (Агаев, 1958, 1987). Sinskaya assessed the species as a system of ecotypes.
We were guided in our research by N.I. Vavilov’s (1935, 1965) differential systematic and geographic method of analyzing cultivated plants, which provided a possibility to undertake targeted searching of valuable breeding materials in various regions and solve the problems of phylogenesis, taxonomy and evolution. This enabled us not only to disclose the diversity of forms, but also to reveal a series of regularities in their variation depending on the degree of cultivation, geographic environments and soil conditions. We arranged the accessions of different origin with ecologically and geographi­cally influenced differences in biological, physiological, bio­chemical and other properties into geotypes, eco-geographic groups of ecotypes, separate ecotypes and concultivars (varietal types). The accessions of each lupin species adapted to a definite and sufficiently vast habitat or geographic area of cultivation were grouped into geotypes, typical of the species with wide areas of distribution. Prof. Agaev (Агаев, 1987) defined the term “geotype” as "morphologically weakly differentiated but genetically determinate race that appeared in the composition of species as a result of their settlement and evolu­tion". This definition allowed us to develop a more detailed eco-geographical classification and revealed efficient ways of lupin genetic resources utilization in breeding.
Geotypes were divided into separate ecotypes or eco-geographic groups of ecotypes (when it was difficult to select sepa­rate ecotypes). Ecotypes are detected among plants adapted to specific niches in the wild conditions, e.g. in mountains, on littoral, close to vil­lages or roads) or in agriculture (e.g. different local forms). Breeding varieties were classified into concultivars (varietal types) on the basis of similar biological and economic properties.

It is necessary to mark that the proposed eco-geographic classification is conditional. It has been developed on the basis of the materials available in the collection of lupin from N.I. Vavilov Institute (about 2500 accessions). It is quite possible that it does not completely encompass all existing diversity of lupin’s forms in nature and in culture. Besides, the ecotypes existing in natural conditions are in the process of constant dynamic development. One of them would crop up and develop, but others tend to fade away because of the activity of man and other unfavorable factors. On the other hand, the humankind have already created many new cultivars of lupin for agricultural production. The breeding practice will be continued in future as well. With regard to these circumstances, such units of eco-geographic classification as ecotypes, concultivars, geotypes and others are less stable than taxonomic units. They should permanently be updated and perfected. However, they provide a better notion about the range of variability of the forms even at the given stage of analysis of genetic recourses and help to obtain a deeper insight in the potential of a concrete species. Using them makes it possible to reveal a number of regulations in the variability of characters within definite species and devise particular ways of their use in breeding practice. These regulations have found detailed consideration in this section.

White lupin (L. albus L.)

White lupin is distinct within the vast and polymorphous genus Lupinus L. for small variation of morphological characters. However, it has wide intraspecific variability in physiological plant properties: duration of vernalization time and growth rate, photoperiodic sensitivity, shape tolerance, drought resistance, cold- and winter-hardiness. There are winter and spring forms of white lupin. Besides, the lupin collection contains wild, undomesticated and local forms, modern breeding varieties, and genetic resources, having wide diversity. Libkind (Либкинд, 1931) classified gene pools of white lupin into two geographic groups: Mediterranean and Abyssinian. According to her, the Mediterranean group is characterized by lighter coloring of the corolla, large pods and seeds, and plants shaped like a sprawling scrub. It is widespread over all the Mediterranean region. The Abyssinian group has darker coloring of the corolla, and small-sized pods and seeds. The shape of plants is oblate and umbrella-like. The area of this geographic group is Abyssinia, with rare occurrence in Egypt. Further analysis of the lupin collection in the VIR network enabled Fedotov (Федотов, 1948) to arrange the accessions into six main types characterized by different physiological and morphological characters: Georgian (winter), Southwest (Algerian), Abyssinian, West-European (Swiss), Middle-European (German) and Palestinian types. This classification encompasses and characterizes the majority of the forms of white lupin having been available in the collection of VIR for 50 years. Up to the present, however, the collection has been abundantly replenished with numerous new accessions from various countries. As a result of their long-term analysis in different zones, many new data have been accumulated. In the outcome of this research, new ecogeographic classification of L. albus has been developed. We have grouped the gene pool of white lupin available now in the VIR collection into 11 geotypes and agrogeotypes, 4 ecogeographic groups of ecotypes, 17 separate ecotypes and 10 concultivares. The list and characteristics of different geotypes, ecotypes and concultivares are presented in the book Ecogeographic research has shown the ranges of changeability in different descriptors of white lupin forms. Duration of growing period under spring sowing varies from 106 to 180 days, seed mass per plant changes from 2.2 to 40 g, green mass yield per from 9 to 250 g, protein content in seed from 35.0 to 53.7%, and oil content from 6.2 to 12.0%.

The Balkan-Asian geotype

It comprises the ecotypes from Greece, Albania, Yugoslavia, Bulgaria, western coast of Turkey, Transcaucasia, and also from Crete and the Aegean Islands. This geotype differs from the others by an extremely broad diversity of forms (Cowling, 1986). All three subspecies of white lupin are contained here. This geotype harbors wild forms, local varieties cultivated in different historical periods, plant populations that turned wild, and also modern breeding cultivars. A rather short stalk, and large number of branches and pods characterize majority of forms. Wild forms differ from cultigens by a shorter stalk, dark blue flowers, dehiscent pods and colored seeds. There are true wild forms of white lupin with dotted dark brown seeds and dark blue flowers (subsp. graecus) in this group. This circumstance, as well as the presence of a large diversity of its other forms, has given us the grounds to surmise that the Balkan Peninsula is the center of formation of Lupinus albus L. and the primary center of origin (diversity) of its initial cultivated forms. The Balkan-Asian geotype consists of 7 ecotypes.
Winter mottled-seed ecotype. The plants grow in natural conditions at the foot of the mountains on carbonaceous soils in the north of Greece, in Yugoslavia, Albania and in the south of Bulgaria. For these plants, typical features are cyan-violet coloring of the corolla and speckled dark brown and dotted seeds with impermeable testa. They belong to subsp. graecus. Such forms were earlier considered by some Soviet and Polish scientists (Kazimierski and Nowacki., 1961b; Атабекова, Майсурян, 1962) as separate species (L. vavilovii Atab. et Maiss., L. jugoslavicus Kasim. et Nowacki.). Winter forms represent the plants. They stay during long time in condition of rosette at spring sowing and start to grow only after preliminary vernalization within 25-30 days at a temperature of 0... +1°C. In the absence of vernalization, these plants develop a short stalk and start to blossom only in autumn, when cold weather is approaching. In the fall planting near Sukhumi, Abkhazia, the growth of the stalk renews in spring and seed are maturing in late May or early June. Seed ripening goes on non-uniformly. These forms tolerate frost at about –15 ... –20°C, as was observed in the fall planting at the mountainous site in Zakatalsk, Azerbaijan (Курлович and Гаджиев, 1989). But the plants of this ecotype are characterized by intense seed shattering.
The plants are notable for a large number of pods on the main stem (up to 60), and rather small-sized depressed seeds (6... 8 mm in diameter). Their 1000 seed weight is 250... 270 g, and seed weight per plant is 30... 40 g. Nowadays valuable hybrids are obtained by crossing the forms of this ecotype with other forms of white lupin. Many of them are early ripening (101-125 days) in spring planting, and have higher seed yield on the main truss (k-2209). Samples of this ecotype are valuable as initial sources for breeding for higher seed productivity and frost resistance. The most representative accession of this ecotype is available under the name of Lupinus vavilovii (k-1788).
Semi-winter mottled-seed ecotype. It includes wild forms of lupin growing in the southern part of the Balkan Peninsula, Crete and the Aegean Islands, pertaining also to subsp. graecus. The plants of this ecotype are similar by morphological characters to the previous ecotype. However, in this ecotype taller and stronger branched plants with light blue corolla color and freckled brown seeds sometimes occur. They exhibit weaker reaction to vernalization, for which sufficing are 15 days with the temperature of +1... +2°C. However, these forms may also stay for 1.0-1.5 months in the rosette phase at spring planting without vernalization in the conditions of Russia. The rosette phase is essentially reduced when the spring is cold. The plants have a short period from flowering to maturing, which makes 60-70 days in Kiev Province, and 40-45 days in Abkhazia. It is quite possible that the plants of this ecotype have hybrid genesis, as in their natural habitats wild plants pertaining to other subspecies (subsp. termis and subsp. albus) frequently occur. Samples of this ecotype are valuable as initial breeding material for early maturity, winter hardiness, small seed size and high seed yield (k.k.-3018, 3118).
Early-ripening Balkan ecotype. Like the previous two ecotypes, it comprises wild and run-wild forms belonging, however, to subsp. termis. The plants have blue or pink flowers and white seeds. Such forms grow over the whole territory of the Balkan Peninsula. They are usually found near villages, on the sides of fields, vineyards, and roads. It is quite likely that these plants have turned wild. They develop in vivo as winter forms and have demonstrated in Kiev Province different reactions to vernalization depending on the weather at spring planting. Their vegetation period lasts 140-150 days. As observed on the fall plantings near Sukhumi, blossoming begins earlier than with the narrow-leafed lupin, and seed maturing starts by early June. The plants usually have thin and short stems (up to 50 cm), on which there are many pods (40-50 per plant) having shatter-protecting valves. Seed mass per plant is 25...27 g; mass of 1000 seed is 250-280 g. The plants are noteworthy for higher drought resistance.
The plants of this ecotype are interesting as sources of small seed size, high seed yield, and resistance to drought. (k.k.-3117, 3339).
Peloponnesus ecotype. Representatives of this ecotype are characterized by more powerful plants (up to 1 m high) and large white seeds. Their seed mass per plant is 28... 30 g, and mass of 1000 seeds is 300... 380 g. The plants are attributed to subsp. albus. Var. vulgaris Libk is the dominating variety of this ecotype. Such forms grow usually on the edges of vineyards, fields, and roads. It allows us to suppose that they are the forms of wild white lupin that has run wild. The plants exhibit predilection to vernalization at spring sowing. The period of vegetation is 140... 150 days in Kiev Province. The plants are notable for by irregular maturing of seed on the main stem and lateral branches. Many accessions are effective sources of resistance to Fusarium wilt (k-2864, 2865, 2881). The most representative accessions of this ecotype are Peloponnes - 127 (k-1441), Longa Peloponnes (k- 1807), k-1440.
Macedonian ecotype. It harbors domesticated local varieties related to subsp. albus. In the Antique period, they were widely cultivated in the territories of modern Greece, Yugoslavia and Albania. Nowadays, they would on a somewhat smaller scale used as green manure, for food after soaking and removal of alkaloids, in medicine and cosmetics. The plants of this ecotype are thermoneutral and more high-yielding in seed and green matter. The mass of the green part of a plant is 150... 200 g. Vegetation period lasts 120... 140 days when cultivated in the conditions of Kiev Province. The plants are characterized by rather simultaneous maturing of seeds on the main steam and lateral branches, and by non-shattering pods. Their seed mass per plant is 32... 38 g and 1000 seed mass is 380-500 g. The seed contain up to 40% of protein (k-2374 from Yugoslavia). Accession k-682 is resistant to Fusarium. Accessions of this ecotype are interesting as sources of early maturity and high productivity.
Turkish ecotype. It incorporates local forms of the western coast of Turkey cultivated there since ancient times. Their distinctive features include early maturity, great quantity of lateral branches, elevated attachment of pods to the main stem and lateral branches. Number of pods reaches 50-60 per plant. Position of pods on the main stem is 0.40... 0.50 m above ground. Their seed mass per plant makes 36... 39 g, and 1000 seed mass is 400... 450 g. Vegetational period is 115-140 days in the conditions of Kiev Province. These plants are enough thermoneutral. Accessions of this ecotype have breeding value in achieving early maturity or higher number of pods on the main stem and lateral branches. However, they are susceptible to Fusarium (k.k.- 529, 530, 532).
Georgian ecotype (Georgian type according the classification of Fedotov). The forms of this ecotype were cultivated as fall crop in western Georgia, mainly in Gurian and Mingrelian regions under local name "hanchkoly", which means "bitter bean". At present, however, as have been shown by our investigations, occurrence of hanchkoly in western Georgia is very limited. This ecotype is a little apart in the arrangement of the Balkan-Asian geotype system. However, Ancient Greece had close relations with Transcaucasia (Синская, 1969), and lupin was apparently distributed to western Georgia also from Greece. Our investigations have shown that hanchcoly in western Georgia is a typical winter form of white lupin planted in autumn. It tolerates low temperatures in winter at the temperature range of –12 … –15°C, reaches in spring 1 m or more in height, and ripens in late May or early June.
White lupin was used in Georgia mainly for food after removing bitter taste by means of washing and boiling. Boiled lupin seed were sold in the markets and were used for food as sweetmeats. The brew was splashed onto animals (cows, sheep) to repel insects. The seed were also used for preparing ersatz coffee. Now, however, white lupin is used in western Georgia mainly for green manure.
Our analysis has shown that accession k-2910 from Georgia is a typical winter form. It was in the rosette stage through­out the vegetation period under spring sowing and matured under autumn planting. Accession k-4423 also manifested itself as a winter or semi-winter form. The plants of this form were in the rosette stage for more than 3 months under spring planting, blossomed only by the end of August, and ripened in November. West Georgian acces­sions turned out to be earlier, taller and more productive in comparison with the winter form from France (k-3340), but their seed weight was less. Commercial cultivar Kievski mutant bred on the basis of Georgian accessions was available both in spring and winter forms. Its seed yield was higher in comparison with other acces­sions (Kurlovich, 1996).
Golovchenko (Головченко et al, 1984) has found out that the accessions of the Georgian ecotype are good material for mutagenesis. The most noteworthy are mutants adapted to the conditions of Ukraine and Russia, characterized by spring growth habit, early ripening, high productivity and low alka­loid content. Those valuable characters were transferred by crosses into a wide range of local lines and in accessions belonging to the Palestinian geotype, and on this basis new commercial cultivars of white lupin were bred (Kievski mutant, Gorizont, Solnechny etc.).

Palestinian geotype (Palestinian type according classification of Fedotov)

This geotype consists from wild and domesticated forms, which one grow in wild condition and are cultivated except for Palestine in Syria, Israel, Lebanon and Jordan. These forms take the first place among the wild forms of white lupin on rates of growth, especially in the first half of vegetation. They are characterized by enough high productivity of vegetative mass and its early forming. The period of vegetation becomes reduced at lengthy days. This circumstance opens a perspective of their usage as basic material for creation of early and acclimatized cultivars in conditions of Russia and Ukraine (Головченко et al., 1984). The samples of Palestinian geotype will widely be used by the breeders of Germany, Poland, Belarus, Russia and Ukraine for obtaining the material with long photoperiod and thermo neutrality. The Palestinian geotype is submitted in the gene pool of VIR by three ecotypes.
Palestinian wild ecotype. The plants of this ecotype concern to subsp. termis and meet in Palestine by separate populations at the foot of mountains and along fields. These plants probably were cultivated earlier in culture and at any stage turned wild. They diffuse by self-sowing and in this connection develop as winter plants. However, they are also weakly respond on vernalization at spring term of crop. Their period of vegetation is 105... 115 days. The plants are dwarfish (up to 20 sm high), with a thin stalk, small-sized leaflets (.,0 sm in length and 1.0 sm in width), short trusses, bluish or pink flowers and 1... 3-seeded beans. Mass of 1000 seeds is 280... 300g. The period of vegetation is reduced at cultivation in northern locales. These plants are more early and xeromorphous in comparison with plants of Balkan early ecotype, they have also a little bit large mass of seeds. The accessions of this ecotype have interest for an acclimatization in Russia in connection with early maturity and thermo neutrality (k.k.-3293, 3294).
Jordan ecotype. These plants also concern to subsp. termis. However, they are cultivated in culture for sideratio and for food purposes after debittering. These plants differ from wild forms by a habitus of plants. They are larger, higher (30 sm high), with leaflets of 2.5... 5.0 sm length and 1.0 sm width, with lengthy polyanthous trusses and 2-4-seeded beans. Their mass of 1000 seeds makes 300... 380 g, seed mass per plant 31... 35g. Plants are thermoneutral and early. The duration of period from plantling to maturing for a sample Tel Karam (k-290), collected by N.I. Vavilov, has averaged for 5 years of testing in conditions of the Kiev area 113 days (V=14.9 %). Many samples differ by high quality of seeds. So, the content of protein is more than 40 %, and content of oil 12 % for the sample k-298 (Каталог мировой коллекции ВИР, Вып. 496, 1989). Heightened (more than 40 %) contents of protein have the samples k-290 and k-302. The samples k-294 and k-295 contain up to 12 % of oil. This ecotype has value for acclimatization in Russia. Usage of the accessions of this ecotype in combination with applying of a mutagenesis has allowed creating highly productive cultivars of white lupin in the Ukrainian research institute of agriculture, and also in Poland (Головченко et al., 1984).
Israeli ecotype. These plants are similar on a habitus with the representatives of previous ecotype. They differ by thicker stalks and light flowers, and also concern to subsp. albus. The varieties vulgaris Libk. and albus L. are dominate. The plants have larger seeds. The mass of 1000 seeds makes 420... 480 g, seed mass per plant 38... 40g. The plants react weakly on vernalization. However, they begin to blossom on 3... 6 days earlier in conditions of vernalization. These plants differ by fast growth by the beginning of vegetation and as a result overtake growth of weeds. The period from plantling to maturing makes 110 … 125 days (V = 15.6 %) at spring crop in the Kiev area, and period from plantling to flowering makes only 32 … 37 days (V = 14.2 %). These plants are cultivated in Israel. They are probably delivered there from others countries. This ecotype has large perspectives in breeding as a source of early maturity and thermoneutrality (k.k.-3225, 3226, 3227).

Abyssinian geotype (Abyssinian type according classification of Fedotov)

It integrates ecotypes of Egypt, Sudan and Ethiopia. The majority of the forms concern to subsp. termis. The samples of this geotype have heightened resistance to drought, early maturity, and relative thermoneutrality. Many from them have resistance to fusarium. It is necessary to mean, however, that this geotype was formed in conditions of short day. The samples react on lengthening of photoperiod by a decrease of productivity in this connection. Abyssinian geotype is parted on 3 ecotypes.
Nile wild ecotype. It meets on all area of the Abyssinian geotype. The plants grow by separate populations in a wild condition, being diffused by self-sowing. It is quite possible that this ecotype has turned wild. N.I. Vavilov has collected a number of wild specimens of this ecotype near for initial sources of Blue Nile during his expeditions in Abyssinia in 1927. The collected plants differ by small-sized seeds, which one ripens more nonuniformly in comparison with cultivated forms of lupin from Egypt and Sudan. Their seed mass per plant makes 27... 31g, mass of 1000 seeds 260... 280g. The process of vernalization reduces growing period and green mass of plants. The period of vegetation changes from 105 to 140 days at spring crop. These plants differ from plants of Palestinian wild ecotype by smaller seeds and by lengthier stalk with smaller quantity of small-sized leafs and leaflets. The forms of this ecotype are most resistant to drought (k- 507, 509). The samples k - 484 and k - 494 are resistant also to fusarium.
Egyptian ecotype. It introduces domesticated part of white lupin in Egypt, where this crop is cultivated since ancient times as foodstuff. The majority of accessions concern to subsp. termis. There are also white-flowered shapes (subsp. albus), delivered apparently from other countries. Several of the intermediate shapes are obtained as a result of hybridization. These plants have small-sized leaflets and short phase of rosette, because they differ by fast growth after plantling. The period of vegetation makes 108-130 days at spring crop. Domesticated forms from Egypt have shorter period from flowering to maturing in comparison with the plants of Nile wild ecotype. It has compounded 62 days (V = 20.6 %) at the sample k-510 in conditions of the Kiev area, whereas this parameter changed from 75 to 95 days for wild forms. The mass of seeds from plant makes 35... 38 g, mass of 1000 seeds 300... 350 g. The samples of this ecotype are possible for using as sources of early maturity (k.k.- 510, 3105, 3114). Accession k-507 has high resistance to fusarium.
Sudanese ecotype. These forms are cultivated in Sudan and Ethiopia at autumnal crop on sandy soils. They were formed up in conditions of rainy autumn and arid summer. The plants fast pass a phase of rosette at spring crop as well as the plants of previous ecotype. Besides, they have enough short period from flowering to maturing. So, it has compounded 74 days (V = 19.3 %) for the sample k-1930 on the average for three years of investigation; for the sample k-1931 it has compounded 77 days (V = 14.3 %). The duration of all period of vegetation changes from 110 to 140 days at accessions of this ecotype in conditions of Kiev area. However, some samples do not ripen at cold and rainy autumn. These plants are more productive in comparison with the plants of previous ecotype. They have also large seeds. Their mass of 1000 seeds 450... 500 g, seed mass per plant 38... 40 g, mass of green part of plant in a phase of flowering 80... 100 g. Many samples have heightened resistance to drought (k-1642, 1646). The accessions k- 486 and k- 495, brought by N.I. Vavilov from Abyssinia, differ by the high contents of protein (Каталог мировой коллекции ВИР, Вып. 496, 1989).

Algerian geotype (Southwest or Algerian type according classification of Fedotov)

This geotype integrates the lupin samples of Algeria, Libya and Morocco. The plants of this geotype concern to subsp. albus. The majority of the forms is weakly differ among themselves. We do not allocate ecotypes in limits of this geotype in this connection. The forms of this geotype are cultivated as crop plants. They are characterized by sluggish growth on the first phases of development and by long period of vegetation (130 - 180 days at spring sowing in conditions of Kiev area). A number of accessions do not ripen here. The plants can stay in condition of a rosette for a long time, but to the beginning of flowering become very powerful, with a big quantity of branches and leafs. The high of plants is 1.2... 2.0 m. They strongly react on vernalization by reduction of vegetation period and by essential change of a habits of plants (Федотов, 1948). These plants have less thick stalk and faster ripen after vernalization or at autumnal crop in Sukhumi. They grow slowly on the shortened days. These plants provide a high yield of green mass and seeds. Their seed mass per plant makes 30... 35 g, mass of green part of plant in a phase of flowering 190... 240 g, mass of 1000 seeds 350... 400 g. The accessions of this geotype have interest at cultivation for sideratio, and also for breeding of winter cultivars for southern conditions - El Harrach 1 (k-3110), El Harrach 2 (k-3111), El Harrach 3 (k-3112), k.k.-2002, 2005, 2011, 2012.

Iberian geotype

It actuates the forms of lupin from Spain and Portugal relating to subsp. albus. White lupin was widely cultivated in Spain in past under a local title "chocho", and in Portugal under a title "tremoco" for human nutrition and on the fodder purposes. Areas of crops are reduced now. However, in those places where lupin was at one time cultivated in culture, meet its turned wild forms, which one usually grow on edges of fields, on shoulders of roads, for foot of mountains and are characterized by broad diversity. This geotype consists from macrospermous and microspermous, summer and winter ecotypes.
The size of beans and seeds tends to reduction at moving from a north to the south of these countries, and the duration of period of vegetation and flowering, number and size of leafs is augmented (Mota, et al., 1982, Mota, 1984; Martins, 1993). In the south of Spain and Portugal are grown in the main winter forms of white lupin, in a north and northwest are grown summer forms. The greatest diversity of the forms is detected in the south. (Mota et al., 1982; Simpson and Gibbon, 1982; Simpson and Martins, 1984). The forms of Iberian geotype are distinguished by heightened resistance to many diseases (fusarium, anthracnose etc.). They have the special value for breeding in this connection. However, all of them have a long period of vegetation in conditions of Russia. The separate small ecotypes are jointed in three eco-geographic groups in connection with extremely broad diversity of the forms of white lupin in the limits of Iberian geotype.
Iberian wild-grown eco-geographic group of ecotypes. It integrates wild forms of white lupin growing on all area of the Iberian geotype, which one probably turned wild in miscellaneous time. Their greatest concentration is detected in a southeast of Portugal on edges of fields. They develop in vivo as the winter shapes and differ by large diversity on predilection to vernalization, by habitus of plants and mass of seeds. This eco-geographic group can be subdivided on a number more small-sized soil and climatic ecotypes, adapted to definite places of growing in this connection. The majority of the forms of this ecotype differ from domesticated forms by big quantity of lateral branches, by non-uniformity of maturing of seeds and more continuous period of vegetation. The majority of them do not ripen at spring sowing in the Kiev area. Besides, the large oscillations in their productivity take place depending from changing weather conditions. Their seed mass per plant makes 260... 350 g, mass of green part of plant in a phase of flowering 150... 250 g, mass of 1000 seeds 250... 320g. The majority of accessions are resistant to fusarium and anthracnose (k.k.-201, 210, 516, 2356, 2364, 2623, 2624, 3260, 3262, 3264).
Algarvian eco-geographic group of ecotypes. It is submitted by domesticasted winter and intermediate forms, which one is cultivated at autumnal crop in the south of Spain and Portugal. A dominating variety is var. vulgaris Libk. The plants are dwarfish, with small-sized leafs. However, they have large seeds. The mass of 1000 seeds makes 400... 450g. The majority of the forms strongly react on vernalization by reduction of period of vegetation at spring crop. They can stay in a phase of rosette for a long time at a warm weather in the spring. These plants are exacting to soil and moisture conditions. The lack of moisture, especially on sandy soils, sharply reduce the yield of plants. The seed mass per plant on the average for 3 years of investigations in conditions of the Kiev area has compounded 20... 25 g, mass of green part of plant in a phase of flowering 200... 250g. The period of vegetation at spring crop is 150... 180 days. Many samples in conditions of the Kiev area do not ripen. This group of ecotypes has interest as sources of resistance to fusarium and high productivity (k.k-2625, 2626, 3250, 3261).
Santarem eco-geographic group of ecotypes. It integrates summer microspermous shapes, which are cultivated at spring sowing in the central and the north part of Spain and Portugal. This group of ecotypes is small-seeded. Their mass of 1000 seeds makes 250... 280 g, seed mass per plant is 5... 20 g. The plants are enough thermoneutral and are weackly react on change of duration of photoperiod. Their period of vegetation is 140... 180 days. A part of accessions does not ripen in conditions of the Kiev area. The plants are highly productive on green mass, differ by a thick high stalk (up to 1,5 m high) and by large leafs. Mass of green part of plant in a phase of flowering reaches 250 g. The accessions of Santarem eco-geographic group of ecotypes can be used as sources of small-seediness in breeding process. A number of accessions has heightened resistance to fusarium wilt (k.k.-2627, 2628, 3251, 3252, 3256, 3259, 3263, 3267).

Apennine geotype

There is an ancient culture of white lupin on Apennines up to a new aeon. Already then were selected and were acclimatized highly productive ecotypes. Some of them probably turned wild during a time, as a climate and part of soils of Apennine peninsula are favorable for existence of lupin in the nature. Many forms of Apennine geotype are characterized by high productivity of green mass and seeds, heightened contents of protein and oils. This geotype consists from two eco-geographic groups of ecotypes.
Neapolitan eco-geographic group of ecotypes. The wild growing forms submit it. Five main centers of their high concentration are detected on Apennine peninsula and off-shore islands (Zhukovsky, 1929; Fischer, 1938; Swęcicki, 1988): 1) near Naples, in region of the volcano Vesuvius and on some off-shore islands; 2) in the south of Italy, in area of Calabria; 3) on island Sicily, near cities Messina, Palermo, and also at foot of volcano Etna; 4) on island Sardinia; 5) on island Corsica.
Each of these centers is submitted probably by several ecotypes. Thus it is necessary to mark, that here grow also L. angustifolius, L. luteus and L.digitatus. However, white lupin usually places for foots of mountains and volcanos, frequently reaching beaches of the sea. The problem of belonging of this group of ecotypes to truly wild or turned wild forms is not solved, because the plants belong to subsp. albus. The truly wild forms of the Balkan Peninsula concern basically to subsp. graecus.
The plants of this group of ecotypes differ by small-sized seeds. Their mass of 1000 seeds makes 280... 350 g. They react on vernalization at spring sowing. The period of vegetation after vernalization in conditions of the Kiev area makes 130... 160 days. It is elongated for 2-3 weeks at sowing without vernalization, and part of accessions do not ripen per separate years in conditions of the Kiev area. The irregular maturing of seeds on a main steam and lateral branches takes place. The productivity of plants is increased on lengthy days. Their seed mass per plant makes 10... 50 g, mass of green part of plant in the phase of flowering 100 … 140 g. The most representative accessions are k.k.-218, 314, 2861, 2862, 2892.
Calabrian eco-geographic group of ecotypes. This group is submitted by local alkaloid forms domesticated by the inhabitants of ancient Roman empire, which one till now are cultivated in the south of Italy at autumnal sowing as green fertilizer and on a forage after deleting of alkaloids. The expressions of Varro Marcus Terentius, dwelling in 116-27 BC testify to their ancient genesis and usage. In the treatise "Agriculture" (book I.XXIII.3) he wrote: " Is necessary to sow lupin not so much for a present crop, but for the next year... Therefore lupin is ploughed to soil on the lean field instead of a dung" (It is quoted from: Скорняков, 1983).
The plants of this group of ecotypes differ by macrospermous. Their mass of 1000 seeds is 400... 440 g. The plants provide a high crop of green mass and seeds. Seed mass per plant is 23... 30 g, mass of green part of plant in a phase of flowering 190... 240 g. The high of plants reaches 1.5... 1.8 m. The plants have a thick stalk, large leafs, with powerfully branching. Duration of vegetational period makes 140 - 160 days at spring sowing in Russia. This period is reduced after realization of vernalization. These plants provide a large increase of a yield at cultivation on fertile soils. Accessions k-1600 and k-1601 differ by the heightened contents of protein. Ecotypes of this group are perspective as basic material for creation of highly productive varieties with resistance to low temperatures (k.k.- 313, 1600, 1601, 1988, 2013).

West-European agrogeotype

This agrogeotype integrates 3 separate ecotypes and 2 concultivars.
Swiss ecotype (West-European or Swiss type according classification of Fedotov). The forms of this ecotype are cultivated basically in Switzerland and in a north of Italy at spring sowing. These plants are more thermoneutral, dwarfish, weakly leafed and microspermous in comparison with the previous groups. Their mass of 1000 seeds is 300... 360 g. The plants provide more intensive growth and high productivity on short days (Федотов, 1948). The seed mass per plant makes 30... 36 g, mass of green part of plant in a phase of flowering 60... 80 g. The period of vegetation is 130... 150 days. These plants differ by low stability to a drought. Their usage is possible as microspermous initial material and sources of thermoneutrality (k.k.-94, 124, 240).
Toulon ecotype. It actuates landraces from the south of France, where there is an ancient culture white lupin. These plants are alkaloidal and have winter growth habit. The majority of the forms usually are in a condition of a rosette within all summer in conditions of the Kiev area and at spring sowing without vernalization. At autumnal sowing in conditions of Azerbaijan and Abkhazia these plants begin to blossom at the end of April, and the maturity of seed comes in June, however on 10-15 days later, than for the forms of Georgian ecotype. Their mass of 1000 seeds makes 280... 360 g. The winter forms from the south of France differ by a high potential of productivity. Their seed mass per plant makes 25... 35 g, mass of green part of plant in a phase of flowering 200... 250 g. However, they are more exacting to a climate and soils in comparison with the summer forms. The young plants maintain frosts up to -10 ...-12 С. Samples of Toulon ecotype can be utilized for creation of highly productive winter cultivars for conditions of Transcaucasia. The most representative accession is k-3340.
Northen France ecotype. The winter forms of white lupin do not always pass the winter in the north of France. The summer forms are cultivated in this part of country already for long time. The local ecotype was formed here distinguished by more small-sized seeds, relative thermoneutrality, and heightened contents of protein and oils. The duration of its vegetational period at tests in Russia and Ukraine was 130... 140 days. The mass of 1000 seeds makes 280... 350 g. The contents of protein in seeds is up to 40 %, oil up to 12 %. The seed mass per plant makes 10... 15g, mass of green part in a phase of flowering 91... 105 g. These samples differ by low resistance to a drought and redundant damp of soils, but have value as sources of high content of protein and oil (k-223 and k-1547).
Concultivar Lucku. It actuates intermediate and winter commercial cultivars from France, created in scientific entities and corporations and cultivated in the south of the country.
Cv. Lucku (k-2617). It is winter or intermediate low alkaloid (fodder) cultivar of white lupin. Duration of period of its vegetation at spring sowing in conditions of Russia and Ukraine after preliminary vernalization is 130... 140 days; including peroid from flowering to maturing 80... 90 days. Its seeds do not ripen without vernalization at spring sowing in Russia at separate years. The plantlets maintained frost about -100 at testing in mountain region of Zakatalsk (Azerbaijan) at autumnal sowing. The seeds ripen in these conditions, and in conditions of autumnal crop near Sukhumi (Abkhazia), in middle of July. The mass of 1000 seeds makes 380... 400 g, seed mass per plant 20... 30 g, mass of above the ground green part in a phase of flowering 210... 240 g. The productivity of seeds in condition of France is up to 4.0 t/ha (Lemaire and Poulain, 1984).
Concultivar Lublanc. It integrates commercial cultivars of France cultivated in a north of country at spring sowing.
Cv. Lublanc (k-2589). It is fodder low alkaloid cultivar, belonging to var. vulgaris Libk. Duration of period of its vegetation at spring crop in Russia is 130... 150 days, including from flowering to maturing 70... 95 days. These plants react weakly on vernalization. Their mass of 1000 seeds makes 340... 350 g. The contents of protein in seeds is more than 40 %, oil 11.2... 12.0 % (Каталог мировой коллекции ВИР, Вып.496, 1989). The seed mass per plant makes 15... 18 g, mass of green above the ground part in a phase of flowering 50... 60 g. Productivity of seeds in condition of France 3.5... 3.8 t/ha (Lemaire and Poulain, 1984).
The accessions of this concultivar have interest as a source of the high contents of protein and oils. Cvs. Lutop (k-2959) and Lukrop (k-2960) concern also to this concultivar.

German agrogeotype (Middle-European or German type according classification of Fedotov)

Lupin has penetrated to Germany from eastern Mediterranean in ХVШ. (Zhukovsky, 1929; Дюбин, 1974). Here for the first time in a world in 1927-1930 by v. Sengbusch were discovered low alkaloid (sweet) forms of lupin, which have been used as an initial material for creation of the commercial fodder cultivars. Lupin received the status of valuable fodder crop after this discovery. The long-lasting breeding and cultivation of lupin in Germany promoted formation of adapted to local condition agrogeotype, consisting from alkaloid and sweet cultivars. German cultivars are characterized by early maturity, macrospermous and manufacturability to field mechanization.
Concultivar Snezhinka. It actuates alkaloidal (bitter) cultivars created until 1930 and used as green fertilizer. First impetus came from King Frederick 11 of Prussia, who sent for seed of L. albus from Italy with a view to its use for improving the poor soils of northen Germany (Hondelmann, 1984, 1986). The large worth in their intrusion in production belongs to Karl von Wulfen, which widely advocated lupin for sideratio (Майсурян, Атабекова, 1974).
Cv. Snezhinka (k-1596). It is characterized by high productivity of green mass, which one makes up to 80... 90 t/ha (Юхимчук, 1963). The mass of 1000 seeds makes 300... 320 g, mass of seeds from 1 plant 10... 16 g. The contents of protein in seeds is 38.5-45.1 %, oil – 9.3 %, methionine 0.51 g on 100 g of dray mass. Resistance to fusarium wilt is medium (24 % of defeat on an infected background). The duration of vegetational period in Russia changes from 130 to 170 days, plants do not ripen per separate years. The productivity sharply changes depending on weather conditions during flowering and maturing of seeds. The accessions Kheine 703 (k-1426), Herkvell (k-1496) etc. concern to this concultivar.
Concultivar Kraftquell. This concultivar consists from fodder (low alkaloid) cultivars created in Germany after discovering of low alkaloidal (sweet) forms of lupin by v. Sengbusch.
Cv. Hadmerslebener Kraftquell (k-1661). It is one of first cultivars made by the widely known Hadmerslebener Corporation. It belongs to var. vulgaris Libk. and is characterized by comparatively low productivity of seeds (10-15 g/plant) and green mass (60-90 g/plant). However, their seeds contain 47.3-52.8 % of protein, 6.2 % oil, 0.212 g methionine and 1,94 g lysine for 100g of dray matter. The mass of 1000 seed makes 320... 380 g. Vegetational period in conditions of Russia is 140... 175 days. It is necessary to attribute to lacks of this old cultivar very late and non-uniform maturity, limitlessness of branching and low resistance to fusarial wilt. The cultivars Hansa (k-2172), Ultra (k-1660), Blanca (k-2295) etc. are concerned also to this concultivar.

Polish agrogeotype

It integrates alkaloidal and fodder cultivars of white lupin in Poland. Lupin has come to this country from Germany and is cultivated from XVШ century with the purposes of fertilizing of pour acid soils. In 1930ies, a turn in cultivation and breeding of lupin took place upon finding by v. Sengbush the first sweet plants. Several years later, upon publishing by Ivanov (Иванов et al., 1932) the method of detection of non-bitter plants and finding of low alkaloid forms lupin began to be used widely on the fodder purposes
(Kubok, 1988). The lupin breeding in Poland is intimately connected with the names of Drs. Barbacki, Frencel, Jach, Jaranowski, Łastowski, Novacki, Ewa i Tadeusz Kazimierski, Mikołajczyk, Sempołowski, Stawiński, Swęcicki etc., which one have made the large contribution in lupin breeding not only in Poland, but also in other countries. Polish agrogeotype is characterized by early maturity, comparative thermoneutrality and stability to fusarial wilt.
Concultivar Gorzki. It integrates alkaloidal (bitter) cultivars. Accessions from Eastern Mediterranean and cultivars from Germany were by the basis for their creation, among which several biotypes were separated, differing by most suitable for local conditions habitus of plants and growth rates.
Cv. Palidus gorzki (k-1984) differs by enough high productivity of green mass and seeds. Their seed mass per plant makes 25... 32 g, mass of green overgrown part in a phase of flowering 180... 210 g. Their seeds contain 36-39 % of protein and 7.0-8.2 % of oil. The mass of 1000 seeds makes 280 … 320 g. The period of vegetation changes in Russian and Polish conditions depending on weather conditions from 120 to 180 days. Late and non-uniform maturity, and low stability to fusarial wilt concerns to lacks of this cultivar. The cultivars and accessions Węlkopolski (k-104), Local from Poland (k-414), Lubin bialy późny (k-1539) etc. are included in this concultivar.
Concultivar Kalina
. It consists from fodder (sweet) forms created after the Second World War. The breeding activity with fodder lupin in Poland was begun after publication method of the mass analysis of plants on low alkaloidness in USSR (Kubok, 1988).
Cv. Kalina (k-2056) is created by Dr. Mikiłajczyk on experimental station Przebędowo by hzbridization accessions Biały 1 x Biały V. It differs by early maturity. Period of vegetation in Russia is 130... 160 days. Their seed mass per plant makes 20... 26 g, mass of green part in a phase of flowering 150... 180 g. This cultivar showed enough high resistance to fusarium on infected background (only 19 % of infected plants). The mass of 1000 seeds makes 350... 400 g.
This concultivar actuates following cultivars and accessions: Przebędowski Wczesny (k-1313), Kali (k-2057), serial of accessions under a title Bialy: Bialy 1 (k-602), Bialy 4 (k-1603), Bialy 6 (k-1604), Bialy 7 (k-1605). Ultra early cultivar Start is created in Timiryazev Agricultural Academy in Moscow with usage of accession from serial Bialy (Гатаулина, 1984).
Concultivar Wat. This concultivar integrates ultra modern fodder commercial cultivars distinguished by early maturity, manufacturability at growing and harwesting, thermoneutrality, immunity to fusarium.
Cv. Wat (k-2585) is created by the breeders T. Kazimierski and W. Swęcicki with usage of wild forms (ssp. graecus). It differs by heightened early maturity and manufacturability at the expense of biological capacity to drop leafs before harvesting. This property eliminates necessity of defoliation. The period of vegetation of this cultivar makes 120... 140 days. It provides high productivity of seed. Their seed mass per plant makes 30... 35 g, mass of green above the ground part in a phase of flowering 40... 50 g. The seeds contain 39.6-41.1 % of protein, 9.3-9.8 % of oil, and 0.200-0.204 g of methionine for 100 g. The mass of 1000 seeds makes 260... 320 g. Cv. Hetman concern also to this concultivar. It is created on the basis of Wat and is distinguished by higher productivity and early maturity. These cultivars have value for breeders of Russia, Belarus and Ukraine as basic material for creation of highly productive forms of lupin of a grain direction.

East-European agrogeotype

It actuates cultivars of white lupin from former USSR. The first item of information about lupin cultivation in Russia have appeared in works of the Free Economical Society in 1811 as about decorative plant. It began to be used also for sideratio in southern regions of Russian Empires subsequently. Cv. Snezhinka from Germany was culrivated at first in these purposes. The local varieties Chernigovsky, Nosovsky 3 etc. were then created also. The new stage in lupin breeding has set in after discovery of the method of finding of low alkaloidal plants. However, method designed in 1928 -29 by v. Sengbusch in Germany were kept secret. In this connection was elaborated new expresses-method of determination of alkaloids in plants in the Institute of Plant Industry (VIR). It was immediately and for the first time in the world published with the foreword of N.I Vavilov (Иванов et al., 1932). This publication, and also the invention of German scientists, was a central to the beginning of breeding work with fodder (sweet) lupin in all world.
East-European agrogeotype is characterized by early and rather even maturity of seeds on a main truss and lateral branches and by restricted branching. The samples of Palestinian geotype, Georgian ecotype, and also cultivars from Germany and Poland were by the initial material for its formation. The mutagenesis was widely used also at creation of Ukrainian and Russian cultivars.
Concultivar Chernigovsky. It integrates alkaloidal cultivars, which is widely cultivated for sideratio.
Cv. Chernigovsky (k-1642) is created on the Chernigov agricultural experimental station (Ukraine). It belongs to var. vulgaris Libk. and differs by high productivity green mass, which one makes 70... 100 t/ha. Duration of vegetation is 135... 165 days, including period from plantling to flowering - 75-80 days, and period from flowering to maturing - 60... 80 days. Their seed mass per plant is 18... 25g, mass of 1000 seeds 350... 420g. This cultivar now is strongly struck by fusarium. Cvs. Nosovsky 3, Zakarpatye local etc. are entered also to this concultivars.
Concultivar Kievski mutant. This concultivar includes fodder (sweet) cultivars bred under the supervision of Dr. V.I. Golovchenko at the Ukrainian Research Institute of Agriculture, and also the cultivars created in Volynsk and Chernigov Agricultural Experimental Stations, which are now cultivated basically on the Ukraine.
Cv. Kiev mutant (k-1904) was developed by the techniques of hybridization and induced mutagenesis with the use of accessions belonging to the Georgian ecotype and Palestinian geotype. It produces seed yield on the level of 2.5... 3.0 t/ha in the conditions of the Ukraine. The protein content in seed amounts to 42.6 %, oil content to 10.7 %, and methionine content to 0.232 g per 100 g. Duration of vegetation is 115... 130 days, including 35... 50 days from planting to flowering, and 65... 80 days from flowering to maturity. Their leaves have a number of new properties connected with mutagenesis: turgid tropism, "dormancy" of leaves, and physiological defoliation of leaf plates (Головченко, 1984). These properties provide fast growth of plants at the expense of effective usage of solar energy, early maturity, and harvest ability. This cultivar is widely cultivated not only on the Ukraine but also abroad, and is used as standard when testing new cultivars in many countries of the world. Cvs. Primorsky (k-2240), Horizont (k-2026), Ukrainian (k-2604), Dnieper (k-2239), Lotos (k-2222), Pischevoy (k-2602) etc. are included in this concultivar.
Concultivar Start. This concultivar includes ultra-early fodder cultivars created in the Timiryazev Agricultural Academy in Moscow under the supervision of Prof. G.G.
Gataulina. They are cultivated in Bryansk, Moscow, Oryol, Tambov and other regions of Russia.
Cv. Start (k-2498) was developed from Polish accession Bialy 7 with the using of mutagenesis. It is one of the earliest cultivars of white lupin. Theшк period from planting to harvesting extends 107... 115 days, including 27... 35 days from planting to flowering, and about 80 days from flowering to maturity. It provides seed yield on the level of 3.0 up to 5.0 t/ha. The protein output is 1.2 t/ha. The mass of 1000 seeds is 200... 250 g. High productivity, early seed maturity, high protein content (40 %) and oil content (12-14 %) make it possible to grow this cultivar quite effectively in most northern areas (Гатаулина, 1984). Besides, this cultivar would find wide application in breeding programs as an efficient source of early maturity. Cvs. Tambov early (k-2717), Tambov 86 (k-2806), Ortam (k-2848), Manovitsky (k-3491) etc. are included in this concultivar.

Chilean agrogeotype

It encompasses white lupin cultivars bred by the well-known breeder, Erik von Baer: Amiga, Prima, Victoria etc. They are grown in Chile and other countries of Latin America, have winter growth habit, long period of vegetation and are poorly studied in the conditions of Russia in this connection.

Narrow-leafed lupin (Lupinus angustifolius L.)

Narrow-leafed lupin is characterized by broad polymorphism in both morphological and physiological characters. It is widespread over all Mediterranean regions as wild plant, and is cultivated in Australia, Europe, and Northern America. Variability of characters depends on eco-geographic conditions of their growing and on the degree of domestication. Wild forms are, as a rule, smaller, with narrow leaflets and pods, and small-sized seeds. Such forms basically occur on inshore sands and in the mountains, on unbroken soils. There are also macro-seeded and wide-leafed plants that grow predominantly close to arable lands. This circumstance evokes the supposition that such forms had once been cultivated in the field and afterwards returned to wild-growing state. Micolajczyk (1963, 1966) marked that the plants of wild types with sluggish initial growth, narrow leaflets and small-sized seeds are more adapted to severe conditions of growing, including frost and drought. Macro-seeded and wide-leafed forms with fast initial growth have more advantages on fertile soils and are good for cultivation in agricultural conditions. Probably such forms were the first to be selected and introduced into culture. Linnaeus described this species using numerous domesticated examples.
Macro- and microspermic plants and the forms with broad and narrow leaflets occur over all Mediterranean area. Besides, no wide gap was observed between cultivated and wild forms. In view of this, there is no reason to insist on the existence of two species: L. linifolius Roth. and L .opsianthus Atab. et Maiss.; and also two subspecies: subsp. angustifolius and subsp. reticulatus (Desv.) Franco et Silva, which have been set apart only because they have different width of leaflets and size of seeds (quantitative characters). We suppose that it would be more correct to subdivide this species into varieties, subvarieties and forms on the basis of morphological variation, and into geotypes, ecotypes, ecogeographic groups of ecotypes and concultivars on the basis of physiological, biochemical, economic and other parameters, similar to the white lupin. This approach provided us with a possibility to identify a series of similar ecotypes within the limits of different geotypes, where variations have definite geographic regularities. The diversity of narrow-leafed lupin is grouped into 12 geotypes and agrogeotypes, 16 separate ecotypes, 3 ecogeographic groups of ecotypes, and 13 concultivars.

Iberian geotype

It is represented by lupin forms from Portugal and western part of Spain. Narrow-leafed lupin occurs in this region basically in wild state, its local name being "tremoco bravo" (Mota at al., 1982; Swęcicki W., 1985, 1988). It exhibits an extremely broad diversity of forms, but dominating are microspermic ecotypes, which have been growing for a long time on one place. They are characterized by primitive dominant characters (mostly small-sized leaflets and seeds, grain hardness, monopodial branching). Macrospermic and broad-leafed forms occur basically on field margins and near roads. Here in Iberia probably was a center of formation of wild narrow-leafed lupin. Wild Iberian lupin forms are resistant to low temperatures and fungal diseases (Forbes and Well, 1966). These positive properties are already utilized in the breeding programs of Australia and USA. The Iberian geotype includes 4 ecotypes.
Iberian waterside ecotype. Plants of this ecotype grow in wild conditions on sandy soils of coastal Portugal at the altitude of 80-120 m above sea level, and are characterized by sluggish growth at initial phases of development, sympodial branching, narrow leaflets and mostly small-sized seeds. Their 1000 seed mass makes 30... 70g. Seed mass per plant, when cultivated in Sukhumi, was 2.0... 4.5 g, the mass of green matter of the above-surface part of the plant in the flowering phase was 16... 20 g. Length of leaflets is 40.0... 42.0 mm, width is 3.0... 4.0 mm. Plant height is 0.8... 1.0 m. It is a completely wild ecotype that has been growing in one place for a long time, as seashores gentle changed their delineation in the course of time (Mota, 1982). Reproduction of the accessions belonging to this ecotype in Russian environments was possible only in the fall plantings in Krasnodar Region (on the Black Sea coast) as well as in hothouses and artificial environment rooms with prior scarification and vernalization. Accessions P.I.168530 and P.I. have been resistant to gray mould and anthracnose (Gladstones, 1977, 1987). They were utilized in the breeding programs of Australia and USA.
Iberian rock ecotype
. Plants belonging to this ecotype grow in separate wild populations in the mountains of Portugal and western Spain at the altitudes up to 1500 m. These plants are similar to those of the previous ecotype in habit, having narrow leaflets, shattering pods and small-sized seeds with testa hardly permeable for water. However, they are distinguished by their dwarfism (up to 0.5 m high), a shorter and denser truss, clearly expressed reaction to vernalization, and increased frost and winter hardiness. The plants of this ecotype survived frosts of about –10 ... –15°C when cultivated in the conditions of Zakatalsk area of Azerbaijan. Their 1000 seed mass ranges between 30... 80 g, seed mass per plant being 18... 22 g. Length of leaflets is 38.0... 43.0 mm, width is 3.0... 4.0 mm. It is possible to grow out the samples of this ecotype, in the same way as with the previous one, only in the fall plantings in southern regions of Russia or in artificial climate conditions. They have value as basic material in breeding for resistance to frost and diseases (k.k.-2970, 3074).
Iberian roadside ecotype. These plants mainly grow as separate populations on the edges of fields together with yellow lupin. This circumstance supports the assumption that in the past they were cultivated for as a green manure crop. Their distinctive features are broad leaflets and rather large seeds. The masss of 1000 seeds is 80... 120 g, seed mass per plant is 8... 12 g, green matter mass of the above-surface part of the plant in the phase of flowering is 80... 110 g. Plant height is 0.80... 1.0 m. Length of leaflets is 48.0... 52.0 mm, their width is 6.0... 6.3 mm. There are samples with miscellaneous coloring of flowers and seeds, and they pertain to different varieties (vars. angustifolius, albopunctatus, chalybens, purpurens). The plants are marked for having faster growth habit, combination of monopodial and sympodial branching, and non-uniform seed maturing. They react to vernalization by reducing their vegetation period. Many forms are resistant to frost, like the samples of the previous ecotype. Accession Elvas (k-2673) has exhibited high resistance to Fusarium wilt when tested against an infested background (only 8% was affected). The samples of this ecotype have higher yield of green matter, and are valuable for breeding cultivars for use as green forage or green manure, and as initial material in frost resistance breeding. (k.k.-3080, 3081, 3083-3087).
Iberian green manure ecotype. It is represented by alkaloidal forms, not long ago cultivated as sideral fertilizer. The plants are notable for higher productivity of green mass (80... 120 g/plant), and for simultaneous seed maturity. Vegetational period lasts 140-160 days when tested in the conditions of Kiev Province. 1000 seed mass is 85... 120 g, and seed mass per plant is 9.2... 14.5 g. Length of leaflets is 50.0... 53.0 mm, their width is 6,0... 6.4 mm. The plants exhibit predisposition to vernalization, surviving frosts of about –5 ... –10ºC. The most representative accessions of this ecotype, k- 511 and Sierra de Aracena (k-3275), are valuable as initial material for breeding cultivars with high green matter yield for use in forage and green manure production.

Moroccan geotype

It integrates the forms of lupin, growing in the northwest part of Africa, basically in Morocco. They differ by a relative thermoneutrality, fast initial growth and heightened contents of protein (Gladstones and Crosbi, 1987). The stability to early frosts is much lower at them than at the forms of Iberian geotype. They grow basically in a wild condition. Klinkowski (1938) however marked, that peasants had collected the seeds of wild narrow-leafed lupin and had grown them on plantations of citron for sideratio in near past. Many forms of narrow-leafed lupin from Morocco have resistance to gray mould, as well as Iberian lupins. The Moroccan geotype consists from 3 ecotypes.
Moroccan waterside ecotype. It actuates wild, small-leafed and small-seeded forms, growing on coast of the Atlantic Ocean and Mediterranean Sea. These plants have more fast rates of growth, early flowering, heightened (38-43 %) content of protein in comparison with Iberian waterside ecotype. Their mass of 1000 seeds makes 45... 85 g, seed mass per plant 3... 5 g, mass of green part of plant 18... 21g. High of plants is 60... 80 sm. Length of leaflets 41.0... 44.0 mm, width 3.5... 4.5 mm. Vegetational period makes 140... 160 days at vernal crop in conditions of the Kiev area. Seeds do not ripen per separate years in this area. The flowering of the plants comes in beginning of May in Sukhumi (Abkhazia) at autumnal sowing, and maturing of seeds start by the beginning of July. Many samples differ by resistance to gray mould.
Moroccan rock ecotype. The forms of this ecotype grow in mountains, at the level of 800... 1200m. The plants are dwarfish (up to 50 sm) and have short and narrow leaflets. Their length is 40.0... 43.0 mm, width 3.5... 4.5 mm. The mass of 1000 seeds makes 50... 90 g, seed mass per plant is 2.5... 6.0 g, mass of green part of plant in a phase of flowering is 15... 23g. The plants are inclined to vernalization in the greater degree in comparison with the forms of previous ecotype. Their seeds do not ripen at vernal crop without vernalization in most cases. The accessions of this ecotype have interest as sources of stability to gray mould (k-3092).
Moroccan roadside ecotype. The plants are tall (up to 1 m) with sympodial branching, broad leaflets and large seeds. The mass of 1000 seeds makes 100... 180 g, seed mass per plant is 5,6... 11,3 g, mass of green part of plant in a phase of flowering is 75... 115 g. Length of leaflets is 50.0... 54.0 mm, width 6.2... 6.7 mm. Vegetational period makes 140... 160 days at vernal crop in the Kiev area. The seeds of this ecotype ripen in June at autumnal crop in Sukhumi. It is possible to suspect that forms relating to this ecotype were cultivated earlier in quality of green manure and then become wild. They grow basically on edges of fields on fertile soils and can be utilized in breeding as sources of high productivity of green mass and stability to diseases (k.k.-2003, 3091, 3093, 3094).

Algerian geotype

This geotype actuates wild forms of the Mediterranean coast of Algeria and Tunis. Available in the collection of VIR accessions from this region are weakly differ among themselves. The geotype is not differentiated on ecotypes in this connection. The plants of this geotype have sluggish growth in initial phases of development, broad and rather short leaflets. Their width is 7.3... 7.5 mm, length 45.0... 48.0 mm. A relative thermoneutrality and high productivity of green mass and seeds characterize these plants. Their seed mass per plant makes 20... 25 g, mass of above the ground part of plants in a phase of flowering 80... 120 g, mass of 1000 seeds 130... 210 g. The samples of this geotype can be utilized in breeding as sources of thermoneutrality and high productivity (k.k.-91, 371, 373, 1939).

Apennine geotype

This geotypes actuates the forms of Western and Southern Italy, and also Sicily, Sardinia and Corsica islands. The extremely broad diversity of the forms of blue lupin is detected within the limits of this geotype. They differ on a habitus of plants, width of leaflets, mass of seeds and other parameters. Parallel to wild plants, which are massed in five main regions, narrow-leafed lupin was cultivated in Italy as green fertilizer in olive plantings and as a substitute of coffee (Maisurjan and Atabiekova, 1974). Alongside with speckled seeds with a vague maculation (var. angustifolius and purpureus), at a part of plants meets beige seeds (var. corylinus and brunneus). Under the data of Swęcicki (1988), the plants are characterized by generically heightened number of seeds in a bean (up to 7). Stability to fusarium wilt is weakly for the majority of the forms.
All diversity of ecotypes in the limits of Apennine geotype are jointed into 3 eco-geographic groups in connection with presence of large assortment of forms and with insufficiency of information about their genetic feature in certain conditions of growing on Apennines.
Neapolitan eco-geographic group of ecotypes. It actuates the wild forms, growing in the region of Naples, in Calabria, and also on Sicily, Sardinia and Corsica islands. Narrow leafed lupin grows here usually on depositions of lava on declines of mountains. It is situated a little bit higher then white lupin and lower then yellow lupin. It is quite possible that separate local ecotypes were formed up in each of the indicated places.
The majority of the forms have predilection to vernalization. Their seed is small-sized, however is a little bit larger, than at Iberian and Moroccan waterside and rock ecotypes. Their mass of 1000 seeds makes 90... 120 g, seed mass per plant 8... 11g, mass of green above the ground part of plant in a phase of flowering 14... 18 g. Length of leaflets is 45.0... 50.0 mm, width 4.5... 5.0 mm. The plants have monopodial type of branching. Vegetational period makes 130... 145 days at vernal crop in the Kiev area. Their seeds however do not ripen per separate years near Kiev.
Apennine roadside eco-geographic group of ecotypes. The plants belonging to this group differ by more powerful development, combination of monopodial and sympodial branching and by large leaflets and seeds. They grow near of roads and villages by separate populations. They are presumptively turned wild shapes, which were at one time cultivated on green fertilizer. Their mass of 1000 seeds makes 110... 170 g, seed mass per plant 10... 14 g, mass of green above the ground part of plant in a phase of flowering 60... 90 g. Length of leaflets is 52.0... 55.0 mm, width – 6.4... 6.8 mm. These plants react on vernalization by reduction of vegetational period, which one changes from 125 to 160 days at vernal crop in the Kiev area.
Apennine green manure eco-geographic group of ecotypes. It is shown by local races, which are cultivated in Italy from XVП of century as green manure for fertilizer of gardens and vineyards. They have appeared as highly productive both on green mass (for green fertilize) and on seeds at conditions of the Kiev area. Their seed mass per plant makes 16... 25 g, mass of green part in a phase of flowering 80... 110 g, mass of 1000 seeds 130... 200 g. Sympodial branching dominates at these plants. Length of leaflets is 53.0... 55.0 mm, width 6.5... 7.0 mm. The period of vegetation is much shorter at these plants than at the representatives of the previous groups and makes 120... 130 days at vernal crop. The accessions of this group have value as sources of high productivity (k.k.-169, 2868, 2921).

Balkan-Asian geotype

This geotype integrates wild and cultivated forms of the Balkan Peninsula, western coast of Turkey, Crete and other Aegean islands. Early maturity, macrospermous and higher seed productivity characterize the majority of ecotypes belonging to this geotype. Many forms have good capacity to reallocating of nutrient substances between miscellaneous organs and have thin valves of beans. Majority of the plants have beige colouring of seed (var. corylinus and brunneus). The sources of resistance to fusarium and drought are detected among them. The accessions of this geotype have perspective as basic material for creation of lupin cultivars for grain. They will widely be used in the breeding programs of Russia, Belarus, Ukraine, Germany and Poland. Balkan-Asian geotype actuates 4 ecotypes.
Balkan wild ecotype. The representatives of this ecotype grow in a wild type in Greece and on Aegean islands. The plants are dwarfish (0.3-0.5 m high), differ by monopodial branching, very shattering beans and small-sized seeds. Their mass of 1000 seeds makes 100... 130 g. However they provide enough high productivity. The seed mass per plant makes10... 13 g, mass of green part of plant in a phase of flowering 12... 20 g. Length of leaflets is 46.2... 50.5 mm, width 4.8... 5.2 mm. These plants exhibit predilection to vernalization. Vegetational period is 120... 140 days at vernal crop. These accessions can be utilized in breeding as sources of small-seediness, early maturity and high productivity of seeds. The most representative accession is k-3346.
Izmir ecotype. It actuates wild forms of western and southern coast of Turkey. They differ from the accessions of previous ecotype by even greater early maturity, dwarfish and by greater mass of seeds. Duration of vegetational period makes 115... 135 days at vernal crop. Mass of 1000 seeds is 110... 140 g, seed mass per plant 11... 14 g, mass of green part of plant in a phase of flowering 12... 19 g. High of plants is about 0,25... 0,40 m. Length of leaflets is 46.0... 51.2 mm, width 5.0... 5.3 mm. The plants have thin valves of beans and differ by heightened drought resistance. Their seeds contain up to 5.6 % of oil. The forms of this ecotype have interest for breeding on early maturity, resistance to drought and high seed productivity. The most typical accession is Izmir 1.
Balkan roadside ecotype. The forms of this ecotype grow by separate populations on edges of fields and on shoulders of roads on all area of geotype. They presumptively were at one time cultivated and after that turned in wild condition. These forms differ from the representatives of previous ecotype by more powerful habits of plants and by large beige seeds. Duration of vegetational period makes 110... 130 days at vernal crop near Kiev. Their mass of 1000 seeds makes 120... 145 g, seed mass per 1 plant 15... 26 g, mass of green part of plant in a phase of flowering 50... 80 g. Length of leaflets is 51.0... 54.0 mm, width 5.0... 5.4 mm. These plants have thin valve of beans and heightened number of seeds in a bean (up to 7). The forms of this ecotype are characterized by xeromorphism. Many from them are resistant to fusarium. The accession Apendrilon (k-2666) is most widely used in breeding programs as source of high seed productivity, resistance to fusarium and drought.
Balkan green manure ecotype. It actuates the local forms, which are cultivated in Greece for sideratio on olive plantations with XVП century (Gladstones, 1974). They are characterized by early maturity and high seed and green mass productivity. Duration of vegetational period is 100-120 days at vernal crop. These plants have broad leaflets, sympodial branching and large seeds. Length of leaflets is 53.0... 55.0 mm, width 6.5... 7.0 mm. Their mass of 1000 seeds makes 140... 200 g. Seed mass per plant is 18... 27 g, mass of green part of plant in a phase of flowering 60... 90 g. The accession k-3345 has interest as source of high productivity and drought resistance.

Palestinian geotype

Broad-leafed and macro-seeded forms submit it on the one hand and narrow leafed and micro-seeded forms on the other. Plitmann (1966) considers that broad-leafed forms (var. basalticus) are endemic in Palestine. The accessions of this geotype differ by early maturity, fast rate of growth in an initial stage, resistance to drought and heightened contents of oil in seeds. As well as accessions of previous geotype, they are valuable as initial material for creation of cultivars for grain. This geotype consists from 2 ecotypes.
Palestinian narrow-leafed ecotype. These forms grow in a wild condition on sandy soils, frequently in mountains. Duration of their vegetational period makes 95... 120 days. These plants have small-sized narrow leaflets and enough small-sized seeds. The mass of 1000 seeds makes 100... 130 g, seed mass per plant 15... 20 g, mass of green part of plant in a phase of flowering 10... 15 g. These forms differ by heightened drought resistance. The contents of oil in seed makes 5.5... 6.4 %. The accessions of this ecotype have interest in breeding on early maturity and drought resistance.
Broad leafed ecotype. It actuates broad-leafed and macrospermous forms growing on fertile soils in apron plains. They differ by early maturity, fast initial growth and broad dark leaflets. Their length is 54.0... 56.0 mm, width 6.5... 7.5 mm. The duration of vegetational period makes 90... 110 days at cultivation in the Kiev area. Their mass of 1000 seeds makes 140... 210 g, seed mass per plant 25... 30 g, mass of green part of plant in a phase of flowering 45... 53 g. The seed contain up to 6.5 % of oil. These samples are earliest in Mediterranean and can be use as valuable initial material in breeding on high seed productivity, early maturity, drought resistance. The most representative accession was collected by N.I. Vavilov in 1924 (k-288).

South-African geotype

This geotype is submitted by the forms growing in the Republic of South Africa (mainly in Cape Province). It was formed up as a result of naturalization here the forms of narrow-leafed lupin from Mediterranean, as the Republic of South Africa and Mediterranean are located on an identical geographic latitude and have similar climatic conditions. The available in our disposal forms from the Republic of South Africa are looking alike as large-seeded ecotypes from Iberian and Moroccan geotypes. Their mass of 1000 seeds makes 150... 180 g, seed mass per plant 20... 28 g, mass of green part of plant in a phase of flowering 71... 80 g. Duration of vegetational period makes130-150 days at cultivation in the Kiev area. . Length of leaflets is 50.0... 54.0 mm, width 6.0... 6.6 mm. These forms react on vernalization by reduction of vegetational period. This geotype is not differentiated on separate ecotypes in connection with unessential distinctions between various accessions and their small quantity in the collection of VIR (k.k.-1992, 1993, 1994).

Australian agrogeotype

It was formed up partially by the same way as previous geotype as a result of naturalization of narrow-leafed lupin from Western Mediterranean. There were favorable conditions for growth and development for lupin in a nature in Australia, located on same geographic latitude with Mediterranean. Besides, the effective breeding activity is conducted in Western Australia since 1954. Highly productive and resistant to frost and diseases commercial cultivars of narrow-leafed lupin are created as a result (Gladstones, 1977). These cultivars are widely grown in agriculture and used in the breeding programs of many countries as sources of non-shattering beans and high productivity. Australian agrogeotype consists from one ecotype and two concultivars according our classification.
West-Australian ecotype. It is submitted by naturalized forms from Mediterranean which are growing now mainly in wild and turned wild condition. Available in the collection of VIR accessions are similar to the forms of Iberian, Moroccan and Apennine geotypes. They are characterized by late maturity and susceptibility to fusarium at cultivation in Russian conditions. Duration of vegetational period makes 130... 160 days at vernal crop. They react on vernalization by acceleration of growth and development. Their mass of 1000 seeds is 80... 150 g, seed mass per plant 10... 15 g, mass of green part of plant in a phase of flowering 30... 55 g. Length of leaflets changes from 45.0 up to 55.0 mm, width from 4,5 up to 6,0 mm.
Concultivar New Zealand Blue. It was formed from wild Mediterranean ecotypes by selection of alkaloidal plants with high productivity of green mass and seeds. Such forms were cultivated in Australia and New Zealand in quality of green manure at autumnal crop.
Cv. New Zealand Blue (k-1732) differs by cyan colouring of a corolla and by motley seed with unclear spotting (var. angustifolius). Their seed coat is easily permeable for water (gene moll). It is characterized by high productivity of green mass and by large seeds. The mass of 1000 seeds makes 160 g, seed mass per plant 14... 18 g, mass of green part of plant in a phase of flowering 70... 120 g. The duration of vegetational period from plantling to maturing makes 106-125 days at vernal crop in the Kiev area, including 40... 45 days from plantling to flowering and 66... 80 days from flowering to maturing. High of plants makes 0.60... 1.10 m. They have interest as basic material for breeding of highly productive on green mass cultivars for green manure. Local variety without name (k-1873) from Australia concerns also to t his concultivar.
Concultivar Unicrop. It integrates fodder (sweet), white flowered and white seeded cultivars of narrow-leafed lupin created under the leadership of Dr. J.S. Gladstones (1977, 1989, 1998). They are characterized by a number of valuable characters and properties: non-shattering pods (genes ta and le), early flowering (Ku), resistance to anthracnose (An) and grey mould (gl). All cultivars concern to var. albidus. They are cultivated in Australia at autumnal sowing.
Cv. Unicrop (k-2096) is created with usage of wild forms from Mediterranean, accessions from Sweden, cv. Muncheberg Sweet Blue from Germany and cv. New Zealand Blue. The duration of its period of vegetation makes 78... 110 days at vernal crop in Russia, high of plants is 0.60... 0.95 m and mass of 1000 seeds is160 g. Valuable character of this cultivar is its non-shattering pods. However it is not adapted to conditions of cultivation in Russia at vernal crop and is strongly struck by fusarium. It is possible to use this cultivar in hybridization with domestic initial material with the purpose of obtaining adapted in Russian conditions forms with non-shattering pods. The conducted researches have shown (Курлович, 1988), that practically non-shattering material possible to receive only by overlapping in a genotype all two genes controlling this characters (le and ta). This procedure can be executed by looking up of non-shattering material in F2 among the obtained hybrids with pigmented beans, as one gene (le) has marker characters (reddish pigmentation of a bean and its swelling). It is possible to find among them the plants possessing also the gene ta, by means of long delay with harvesting, by drying the beans, or with the help of anatomic researches.
Cvs: Marri (k-2270), Uniharvest (k-2090), Illyarie (k-2586), Chittick (k-2631), Yandee (k-2632), Danja (k-3056), Yorrel (k-3057), Warrah (k-3058), Gungurri (k-3059) etc. concern to this concultivar. They can be used in future breeding as sources of non-shattering pods and resistance to anthracnose, gray leaf spot, phomopsis and frost.

North-American agtogeotype

This concultivar is submitted by fodder (sweet) cultivars of narrow-leafed lupin cultivated in USA. The majority of them are created as a result of implementation of joint breeding program with Australia. Wild forms from Mediterranean, and commercial cultivars from many other countries served as basic material for their creation. Narrow-leafed lupin is cultivated in USA as well as in Australia at autumnal term of sowing. The cultivars of North-American agrogeotype are adapted to transferring of frost and have resistance to many diseases.
Concultivar Rancher. It integrates cultivars with resistance to anthracnose (Glomerella cingulata) and gray maculation of leaflets (Stemphylium vesicarium). The sources of these tags were taken from Portuguese accessions P.I.168530 and P.I.168535 (Forbes, Well, 1966).
Cv. Rancher (k-3039) is created by hybridization of cv. Blanco with theforms resistant to anthracnose and gray maculation of leaflets. It has white flowers and seeds (var. albidus). The mass of 1000 seeds makes140... 160g. This cultivar differs by late maturity and low productivity at vernal crop in conditions of Russia and the Ukraine. Its seeds do not ripen per separate years. Besides, the plants are strongly struck by fusarium. It can be utilized in breeding programs as a source of resistance to anthracnose and gray leaflets maculation. The accessions k-1716 and G-93 (k-1878) concern also to this concultivar.
Concultivar Frost. It actuates cultivars resistant to early frosts, fusarium and other deseases.
Cv. Frost (k-2258) is created in USA with usage of wild forms from Iberian geotype. It has blue flowers and small-sized and white seeds with sparse dark-brown and gray spots (var. chalybeus). The mass of 1000 seeds makes 108... 110 g. This cultivar differs by high productivity of green mass, big quantity of seeds and stability to fusarium in some locales of Russia and Ukraine at vernal crop. However, stability to diseases is exhibited not in all places. So, disease incidence of plants has compounded only 3 % on strongly contaminated by fusarium experimental field of Pavlovsk experimental station (the Leningrad area). However, 38 % of the ill plants were detected on the infected plots near Kiev, and the plants were completely struck (100 % of a defeat) on the infected plots in Bryans region. Сv. Frost can be involved in hybridization with others accessions steady to fusarium. It opens perspective to identify the transgressive forms in F2 and subsequent generation, which have higher resistance to fusarial wilt in comparison with the initial forms. This problem is considered in more detail in еру section «Genetics of quantitative characters». Сv. Frost can be utilized in the breeding programs as a source of high productivity and stability to low temperatures and fusarium. The accessions Tifblue (k-2630), Frost blue (k-1750) and others concern to this concultivars.

German agrogeotype

Narrow-leafed lupin as well as white was spread in Germany for sideratio from Eastern Mediterranean. Highly productive on green mass and earliest forms were selected in these purposes. As a result of their acclimatization were formed up adapted to local conditions cultivars of lupin for green manure. The first in the World fodder (sweet) cultivars were created also in Germany after v. Sengbusch’s discovery. German agrogeotype is submitted by two concultivars, integrating the sorts for sideratio and the fodder cultivars, which are adapted to cultivation at vernal crop. The majority of them are not resistant to fusarial wilt.
Concultivar Pflug. It integrates local alkaloidal sorts cultivated in quality of green fertilizer. They differ by high productivity of green mass and seeds.
Cv. Pflug (k-167) is created by selection of highly productive alkaloidal plants from a material of an unknown genesis. Its mass of 1000 seeds makes 150... 160 g, seed mass per plant is 18... 21g, mass of green part of plant in a phase of flowering is 80... 115 g. Vegetational period makes 140... 160 days at cultivation in conditions of Ukraine. The seeds of this cultivar do not ripen in conditions of the Moscow area. The plants react on vernalization by reduction of vegetational period. Their resistance to fusarium is very weakly. Its usage is possible in breeding programs as a source of high productivity of green mass. The cultivars Bismark, Edelweiss, Pflug blue and others concern to this concultivars.
Concultivar Muncheberg. This concultivar consists from fodder (sweet) cultivars.
Cv. Muncheberg Sweet Blue (k-1618) is created by one of the first on the basis of discovered by v. Sengbusch low alkaloidal (sweet) forms. The mass of 1000 seeds makes 160 …180 g, seed mass per plant is 15... 18 g, mass of green part of plant in a phase of flowering is 45... 58g. Vegetational period makes 130... 155 days at cultivation in the Kiev and Moscow areas. This cultivar is strongly struck by fusarial wilt, has predilection to lodging and very shattering beans. However, it served as initial material for creation of many more highly productive fodder cultivars of narrow-leafed lupin not only in Germany but also in other countries (Australia, Poland, USA, Sweden). Cvs. Sengbusch stamm 411, Stamm 415, Guelzower gelbe suesslupine etc. concern also to this concultivar.

Polish agrogeotype

It integrates alkaloidal and fodder cultivars of narrow-leafed lupin from Poland. This species as well as white lupin was delivered to Poland from Germany and was cultivated since 1850 as green fertilizer, as decorative plant, and also on forage for sheep. Creation of the fodder cultivars of narrow-leafed lupin was began after finding the first fodder (sweet) forms. The cultivars of Polish agrogeotype are characterized by heightened early maturity in comparison with German geotype. Many from them are resistant to fusarium. Polish cultivars are jointed into 3 concultivars.
Concultivar Pulawski. This concultivar actuates local alkaloidal varieties cultivated on green fertilizer.
Cv. Pulawski (k-335) is created on experimental station Puławy by the breeder J. Sypniewski in 1920 from a local material. It differs by high productivity of green mass and seeds. The seed mass per plant makes 16... 20 g, mass of green part of plant in a phase of flowering is 75... 110 g, mass of 1000 seeds makes 180 g. It differs by early maturity, but it is not resistant to fusarium. Vegetational period makes 120... 140 days in conditions of the Kiev area. It can be utilized as a source of high productivity. The varieties Pulawski Wczesny, Pulawski Wysoki , Wielkopolski Gorzki etc. are included to this concultivar.
Concultivar Mirela. It integrates modern alkaloidal cultivars used as green fertilizer, and on the fodder purposes by mixing of their green mass and seeds with other forages (Kubok, 1988). Early maturity, high productivity and stability to fusarium characterize these cultivars.
Cv. Mirela (k- 2570) is created in 1981 by Dr. J. Mikołajczyk on experimental station Przebędowo by a hybridization of accessions (Rozowy Past. x Krasnopolistny) x Ignis. It differs by pink flowers and beige seeds (var. brunneus). Early maturity and high productivity of seeds characterize it. Their seed mass per plant makes 26... 30 g, mass of green part of plant in a phase of flowering is 60... 71 g, mass of 1000 seeds is 140... 150 g. This cultivar has demonstrated high stability to fusarium in many locales of the former USSR. Duration of vegetational period makes 85... 110 days. It will widely be used in the breeding programs as a source of high productivity, early maturity and stability to fusarium. The cultivar Turkus (k-2148) concerns also to this concultivar.
Concultivar Emir. It consists from fodder (sweet) breeding cultivars.
Cv. Emir (k-2662) is created in 1981 by Dr. J. Mikołajczyk on experimental station Przebędowo by a hybridization of accessions Procerus Róż. Wezesny x LA-111. Thermoneutrality and high resistance to fusarium characterizes it. Their seed mass per plant makes 20... 25 g, mass of a green part of plant in a phase of flowering is 50... 60 g, mass of 1000 seeds is 120... 130 g. Duration of vegetational period is 90... 115 days. This cultivar has value as a source of early maturity, thermoneutrality and resistance to fusarium. The cultivars Kazan (k-2569), Ignis (k-2183), Remik (k-3122) and others are included in this concultivar.

East-European agrogeotype

It integrates bitter and fodder (sweet) cultivars from Belarus, Russia, Lithuania and Ukraine. These cultivars are characterized by even greater early maturity in comparison with the Polish forms, by large-seediness and by fitness to local conditions of cultivation. The accessions from Poland and Germany, and also the samples of Balkan-Asian and Palestinian geotypes were as initial material for their creation. East European agrogeotype consists from 4 concultivars.
Concultivar Bienyakonsky. It is submitted by alkaloidal cultivars cultivated on green fertilizer.
Cv. Bienyakonsky 484 (k-1457) differs by blue flowers and motley and gray seeds with unclear spotting (var. angustifolius). It is characterized by high productivity of green mass (95... 120 g/plant) and seeds (25... 29 g/plant). The mass of 1000 seeds makes 150... 155 g, the contents of protein in seed is 32... 34 %, duration of vegetational period is 100... 130 days. It is not resistant to fusarium. Its usage is possible in breeding as a source of high productivity of green mass. Cvs. Bienyakonsky 335 (k-1477), Nemchinovsky cyan (k-1632), Pink 339 (k-1342), Cyan 173 (k-1507) and others are included in this concultivar.
Concultivar Nemchinovsky. This concultivar integrates fodder (sweet) cultivars created before 1985 year in the former USSR. They differ by early maturity and high productivity of seeds and green mass. However, they are not resistant to fusarial wilt.
Cv. Nemchinovsky 846 (k-1981) is created in Enterprise ’’Podmoskovie’’ by hybridization of Russian variety Беляк with the German sample Ку-2. The plants have pale-violet flowers and white seeds without a triangular spot and strip (var. albosyringeus). The seed mass per plant makes 23... 26 g, mass of green part in a phase of flowering is 80... 100 g. This cultivar provides productivity of seeds up to 2.3 t/ha. That is much higher in comparison with widely distributed in Russia cultivar of yellow lupin Быстрорастущий 4 (Дебелый, Калинина, 1974). It is adopted also as the standard at testing of new cultivars and widely has used in the breeding programs with the purpose of its improvement, has value for genetic researches. The cultivars Byelorussian 155 (k-1486), Немчиновский кормовой (k- 1980), Северный 3 (k-712) and others are included in this concultivar.
Concultivar Danko. It integrates new breeding cultivars created after 1985 year in the countries of the former USSR. Higher productivity and stability to deseases (fusarium) characterize them. Some of them have non-shattering beans.
Cv. Danko (k-2949) is created by Dr. N.S. Kuptsov in Byelorussian Institute of Agriculture by individual selection from a hybrid combination Ban x Лаф 4. It differs by high productivity, resistance to root rot and has non-shattering beans. The average productivity of seeds for 3 years of tests has compounded 3.4 t/ha, the yield of green mass has made 47.9 t/ha. The content of protein in seeds makes 33.7 %. Duration of vegetational period is 96... 119 days (Купцов, 1987). Cvs. Reserve 884 (k-2747), White consul (k-3029), Grey consul (k-3026), Натальевский (k-3027) concern to this concultivar.
Concultivar Ladny. This concultivar integrates cultivars of narrow-leafed lupin with determinate branching. This character causes uniform of seed maturity, non-lodging and manufacturability at harvesting.
Cv. Ladny (k-2648) is created in Timiryazev Agricultural Academy in Moscow in cooperation with Enterprise ’’Podmoskovie’’ by processing of cv. Nemchinovsky 846 by gamma rays. The plants have pale-violet flowers and white seeds without a triangular spot and strip. Their lateral branches are absent or shortened, flowers are axillary (var. albosyringeus, f. kloczkovii). This cultivar is characterized by early maturity and large stability and productivity of seeds on the main truss. The coefficient of variation of mass of 1000 seeds makes only 9 % at it, whereas this index has 22 % at initial cultivar Nemchinovsky 846 (Коновалов et al., 1985). Their seed productivity is at the level of Nemchinovsky 846 or even exceeds it. However implementations of high seed productivity need the heightened seeding rate (1.6... 2.0 million seeds on 1 ha). Duration of vegetational period makes for this cultivar is 75... 90 days. The cultivars and samples: Дикаф 14, Гелена, Силена, Lanedeks 1 etc. concern to this concultivars.

Yellow lupin (Lupinus luteus L.)

This species is characterized by a rather smaller diversity of physiological and commercial characters if compared with the white and narrow-leafed lupins. However, it may also be divided into different geotypes, ecotypes, eco-geographic groups of ecotypes, and concultivars on the basis of variation in productivity of green matter and seeds, duration of vegetation period, predisposition to vernalization, seed weight, etc. Yellow lupin probably originated, according to Gladstones (1974), in Portugal and western Spain, whence it was distributed as cultivated and horticultural plant. Merino (1905) observed that plants of yellow lupin in the Spanish province of Galicia are often clustered around villages, which underlines the role of cultivation in the distribution of this species even within its present natural range. In the same way (by semi-naturalization) it appeared on the Island of Madeira (Lowe, 1968) and in the west of France, in Southern Africa and Western Australia (Gladstones, 1974). Yellow lupin is cultivated as fodder crop on acid and, basically, sandy soils in Belarus, Russia, Ukraine, Germany, Poland, Hungary, Netherlands, Brazil and other countries. The diversity of yellow lupin is classified into 8 geotypes, 10 ecotypes, 2 eco-geographic groups of ecotypes, and 10 concultivars.

Iberian geotype

It actuates wild, turned wild and cultivated forms from Portugal and Western Spain. Alkaloidal yellow lupin is widely used in these countries in the miscellaneous purposes:
as standing hay (dry pasture) for seep, as green forage, as hay or silage, as a grain crop, on green fertilizer and other uses (Abreu, 1994). It is used in these purposes most widely in area of „Montados” in Portugal, where the climate is typical for Mediterranean and the soils have lower productive capacity. The „Montado” is domesticated in Portugal by a highly valued oak – the cork oak, which has been „gardened” by man through centuries. This process of domestication, where is not only a plant species that is being domesticated, but the entire agroforestry system, found a special role for the Lupinus luteus species (Pinto, 1994). The plants of Iberian yellow lupin differ by high productivity of green mass, small seediness, and resistance to low temperatures and diseases. Iberian lupin already has ensured advance of breeding on resistance to fusarial wilt, and probably will ensure advance on resistance to anthracnose (Colletotrichum gloesporioides Penz.). Besides, accessions of Iberian geotype are perspective as initial material for breeding of fodder cultivars for green mass and silage, which should provide a high yield of vegetative mass. Wild forms of lupin from Iberian geotype have most small-sized seeds. However many forms can be grown on seeds only at autumnal crop in Russian conditions in the Southern region (on Caucasus) or in artificial conditions after vernalization and scarification.
Lupinus hispanicus Boiss. et Reuter is other related to yellow lupin species growing also on Pyrenees. Both species have identical number of chromosomes but differ serologically and on colouring of flowers. They had probably one initial shape, however divergented for a long time. Hybrids obtained between them distinguished by a number of valuable characters. Existence on Pyrenees two close species (L.luteus L. and L. hispanicus Boiss.et Reut.) with same number of chromosome (2n=52), wide diversity of wild and cultivated forms of yellow lupin and long historical period of their growing give the reasons to suppose that on Pyrenees was the center of formation the wild forms of yellow lupin and the center of origin (diversity) of its cultivated forms. Iberian geotype integrates 4 ecotypes.
Lisbon ecotype. The wild small-seeded forms growing on sandy acid soils of coast of Portugal and Spain submit it. One of their large concentrations is situated in neighborhoods of Lisbon. Analysis of collected accessions has shown, that the plants concerning to this ecotype long time stay in a phase of rosette in conditions of Russia. However, this phase of development is essentially reduced after vernalization. High of plants is 0.6... 0.7 m. They have basically monopodial branching. The seed mass per plant makes 2... 5 g, mass of green part of plant in a phase of flowering is 100... 110 g, mass of 1000 seeds makes 75... 90 g. The contents of protein in seeds is 41.9 %, oil 5.4 % (Gladstones and Crosole, 1987). Their seeds have the property of grain hardness. The majority of accessions do not ripen at vernal crop in conditions of Russia. The flowering comes in May and maturing of seeds in July at autumnal crop in Transcaucasia. These plants are resistant to low temperatures. The part of plants maintained frosts up to -10 ...-150 within all winter in mountains regions of Azerbaijan. Tomaszewski in Poland utilized the wild Portugal form № 679 of this ecotype for creation of resistant to fusarial wilt varieries Cyt and Afus (Kubok, 1988; Swęcicki, 1988). The forms of Lisbon ecotype have value as basic material for creation of highly productive fodder green mass cultivars (for silage) with high immunity to fusarial wilt. The most representative accession of this ecotype is k-3099.
Madrid ecotype. This ecotype actuates wild forms growing by separate populations on declines of mountains, at the altitude from 500 up to 1000 m. One of the centers of their high concentration is reviled in neighborhoods of Madrid. These plants differ by unpretentiousness to conditions of growing and by resistance to cold. They have smaller high (up to 0,5 м) and stronger monopodial branching in comparison with the forms of previous ecotype. The reacting on vernalization for plants is strongly exhibited by reduction of vegetational period and by change of habits of plants. Their seeds do not ripen without preliminary vernalization at vernal crop in the Kiev area. The plants maintain frosts up to -12 ...-150С in conditions of autumnal crop in mountains regions of Azerbaijan. Their seed mass per plant makes 3.5... 6.0 g, mass of 1000 seeds is 70... 100 g. The seeds have coat difficultly permeable for water. The contents of protein in seeds makes 38... 42 %, oil 4.5... 5.7 %. Such forms succumb on productivity of green mass the plans of Lisbon ecotype (80... 90 g/plant). They are valuable basic material at breeding on small-seediness, frost, cold and diseases resistance (k.k.-3072, 2289, 2291).
Callician ecotype. It is submitted by growing in wild condition more large-seeded and tall forms, which grow near villages and along fields. That testifies (Merino, 1967), that they had been cultivated at one time and then they turned wild. These plants are with monopodial and sympodial branching; their high is up to 0.8 m. They differ by high productivity of seeds and green mass. Their seed mass per plant makes10... 12 g, mass of green part of plant in a phase of flowering is 200... 210 g. Their seeds ripen not uniformly on a central truss and on lateral branches. The mass of 1000 seeds makes 90... 120 g. These plants to a lesser degree exhibit predilection to vernalization, differ by high nitrogen fixing capacity, resistance to fusarium and anthracnose. Duration of vegetational period makes 150... 175 days in conditions of the Kiev area at vernal crop. These accessions have perspective as initial material for creation of highly productive on green mass (for silage) and resistant to diseases cultivars (k.k.-2290, 2292, 2869, 2870, 3279, 3280).
Seville ecotype. It consists from local high productive alkaloidal local races of Spain and Portugal, which are cultivated on fields as green fertilizer and on the forage for sheep. They are subjected to long-lived selection, are adapted to cultivation on good reclamationed soils, have large stalks and leaflets with mesomorphic pattern. They provide high productivity of green mass (up to 245 g/plant). Their seed mass per plant makes 11... 12 g, mass of 1000 seeds is 100... 120 g. The seeds ripen more uniformly. Duration of vegetational period is 140... 180 days at vernal crop in the Kiev area. Some samples do not ripen per separate years. The accessions of this ecotype are resistant to fusarium, have high nitrogen fixing capacity and positive reaction to applying of the commercial strains of Bradyrhizobium lupini. They are perspective for creation of high productive and resistant to fusarium and anthracnose cultivars (k.k.-2865, 2992, 3274, Tromusillo, MKW-2).

Moroccan geotype

It integrates the forms growing in wild or turned wild condition on Northwest coast of Africa, basically on the territory of Morocco. They differ by weak reacting on vernalization (thermoneutrality). Besides, vegetational period of these plants is elongated at cultivation in conditions of lengthy day. Many forms are characterized by the heightened contents of protein in seed (40... 45 %), by short period of rosette phase and fast initial growth. This geotype consists from 2 ecotypes.
Tangier ecotype. It is submitted by wild small-seeded forms, which grow on sandy and petrous soils on coast and declines of mountains. The plants are characterized by enough high productivity of green mass (160... 170 g/plant). Their seed mass per plant makes 4.0... 6.5 g, mass of 1000 seeds is 80... 120 g. The part of seeds has property of grain hardness. The plants differ by gentle reacting on vernalization. Duration of vegetational period makes 150... 160 days at vernal crop in conditions of the Kiev area, and about 180 days in conditions of the Moscow area. The accessions of this geotype have interest as sources of a thermoneutrality and small-seediness. The most representative accession is Anamoro (k-3126).
Rabat ecotype. It consists from more large-seeded forms growing on fertile soils, frequently on edges of roads and fields. These plants are gentle react on vernalization in Russian condition at vernal sowing, differ by fast initial growth, strong monopodial and sympodial branching and high productivity of green mass. The maturity of seeds goes no uniformly on lateral branches of the different orders. Their seed mass per plant makes 12... 13 g, mass of green part of plant in a phase of flowering is 190... 200 g, mass of 1000 seeds is 100... 130 g. The content of protein in seeds makes up to 44... 45 %. Duration of period of vegetation is 130... 155 days at vernal crop in Kiev area and 140... 160 days in conditions of Moscow area. The forms of this ecotype have value at selection on thermoneutrality, high contents of protein and productivity (k-2004).

Apennine geotype

It actuates the forms growing on Apennine peninsula and on islands Sicily, Sardinia and Corsica. This geotype is characterized by broad scope of variability. It is submitted by cultivated for a long time local alkaloidal forms, which are used as green fertilizer, and by numerous wild populations of plants, which are located in miscellaneous ecological conditions, frequently near to plants of white and narrow-leafed lupins. Many wild forms of yellow lupin have similarity with local varieties (large seeds, sympodial branching). Heightened number of seeds in a bean characteristically for many forms of this geotype (Swęcicki, 1988). It is possible to suspect that separate ecotype were formed in different foci of them growing, and especially on islands. But we have not found the sufficient basis for partition of this geotype on separate ecotypes in connection with restricted differences between available in the collection of VIR accessions of this geotype. We have aggregated all available diversity of accessions into two eco-geographic groups of ecotypes.
Messina eco-geographic group of ecotypes. It is submitted by wild forms growing on the island Sicily (near city Messina), on islands Sardinia, Corsica, and also in the south of Apennine peninsula, in Calabria and near Naples. These plants are characterized by broad polymorphism of variability. Their seed mss per plant changes from 6 up to 18 g, mass of green part of plant in a phase of flowering from 50 up to 85 g and duration of vegetational period at vernal crop in the Kiev area from 120 to 150 days. These plants react on vernalization by reduction of vegetational period on 10... 15 days. The majority of the forms are not resistant to fusarium (k-1447, Sycylia Kl.1, Sycylia Kl.2, Sycylia Kl.3).
Italian green manure eco-geographic group of ecotypes. It integrates local alkaloidal varieties. They differ by high productivity of green mass and by large seeds. Their mass of 1000 seeds makes 140... 150 g, seed mass per plant is 10... 12 g and mass of green part of plant in a phase of flowering is 100... 130 g. Duration of vegetational period makes 135... 155 days at vernal crop in the Kiev area. The forms with resistance to fusarium are not detected among accessions of this group. The typical accessions of this group are k- 3341 and k- 2074.

Balkan-Asian geotype

It integrates forms of yellow lupin collected on the Balkan peninsula and in the western part of Turkey. They differ by more shorts period of vegetation, heightened drought resistance and small-seediness. The genetic resources of lupin belonging to Balkan-Asian geotype are widely used in the breeding programs of Germany, Poland, Russia and Ukraine. Balkan-Asian geotype consists from 2 ecotypes.
Greek ecotype. This ecotype is submitted in the collection of VIR now only by one wild form (k-3342) collected on Balkan Peninsula. Some forms of this ecotype having been available in the VIR collection before Second World War have appeared lost during blockade of Leningrad. But, Sibthort and Smith (1830) mentioned and illustrated L. luteus in the Flora of Greece. Merkenschlager has found the plants of L. luteus near Sparta on acid soils derivated from crystalline shales (Майсурян, Атабекова, 1974). Klinkowski (1938) also has informed about apparently native Greek L. luteus L. However, Gladstones (1998) has exposed to doubt presence of wild yellow lupin on Balkan. We consider on this basis that yellow lupin meets now in Greece very seldom. It is on the verge of disappearance, probably as a result of human activity. The available in our disposal sample is characterized by relative early maturity, high seed productivity and by enough large seeds. Its mass of 1000 seeds makes 110... 150 g, seed mass per plant is 18... 22 g and mass of green part of plant in a phase of flowering is 45... 50 g. Duration of vegetational period makes 120... 130 days.
Anatolian ecotype. The wild forms collected in Turkey represent it. They are submitted in the collection of VIR mainly by accessions collected by Klinkowski (Klinkowski Anatolian, Klinkow.Aschersleb.31555, 31657, 32155, and 32457). The plants of this ecotype differ also by early maturity, even bigger seed productivity and mass of seeds in comparison with the previous ecotypes and heightened resistance to drought. Their mass of 1000 seeds makes 130... 160 g, seed mass per plant is 19... 23 g and mass of green part of plant in a phase of flowering 30... 40 g. Duration of vegetational period is 115... 130 days in conditions of the Kiev area. The accessions of this geotype have interest as sources of high seed productivity and resistance to drought (k.k.-2076-2081 and k-3343).

Palestinian geotype

This geotype actuates the forms of yellow lupin from Israel, Palestine and Lebanon. The isolation was rendered essential influencing on formation of yellow lupin in this region. That has resulted in differences of growing here plants of lupin (Kazimierski and Kazimierska, 1975). The seeds of same plants have brown tint (gene fuscus). For some of the forms are typical fast initial growth, early maturity, small-seediness and resistance to drought. Kazimierski and Kazimierska (1975) mark following special characters of the Palestinian forms of yellow lupin: small-sized leaflets, short inflorescence, first flower is arranged below the vercicil, low plants, smaller number of seeds in a bean (3-4) and brown seed. The presence of these characters has given the basis to the writers to dedicate the Palestinian forms in separate subspecies (subsp. orientalis Kazim. et Kazim.).
However, conducted analysis of available in the collection of VIR forms has allowed to establish a number of essential distinctions between accessions of this geotype. Parallel to plants described by Kazimierski and Kazimierska (1975) as subsp. orientalis, the forms with broad leaflets, black- doted large seeds, fast initial growth and early flowering are detected. We suppose, that described by Kazimierski and Kazimierska (1975) forms more correctly to esteem as one of the local ecotype on an edge of area of the species in connection with available big diversity of the forms within the limits of same area. The Palestinian geotype consists from 2 ecotypes in our opinion.
Oriental ecotype. It actuates the forms with brown tints of seed (var. kazimierskii), with small-sized narrow leaflets, short dense inflorescence, and with small quantity of seeds in a bean (3-4). These forms were described as subsp. orientalis Kazim. et Kazim. Late maturity and low productivity of green mass and seeds characterize them. Their seed mass per plant makes 10... 15g, mass of green part of plant in a phase of flowering is 20... 30g and mass of 1000 seeds is 90... 105g. Duration of vegetational period makes 120... 150 days in conditions of the Kiev area. These forms are very susceptible to fusarium. We consider that they are endemic in the territory of Israel and Palestine. They have interest for further genetic researches (Israel 8a, Israel 8b).
Jerusalem ecotype. This ecotype consists from the forms with black-doted large seeds (var. luteus and var. maculosus), with broad leaflets, fast initial growth and early flowering. They differ by early maturity, resistance to drought and by high seed productivity. Their seed mass per plant makes 20... 24 g, mass of green part of plant in a phase of flowering is 40... 45g and mass of 1000 seeds is 120... 160g. Duration of period of vegetation makes 108... 120 days at cultivation in the Kiev area. We concider that accessions belonging to this ecotype were delivered from other countries. They have interest as initial material in the breeding programs on early maturity, resistance to drought, tolerance to virus diseases and high seed productivity (Palestyna 1, Palestyna 3).

German agrogeotype

Yellow lupin was spread in Germany from Mediterranean as well as white and narrow-leafed lupins. It began to cultivate as flower plant under a title "the Turkish violet" with XVI century (Zhukovsky, 1929). It has received broad distribution as crop plant for sideratio in subsequent. The first fodder (sweet) cultivars were created in Germany also after finding of low alkaloidal forms by v. Sengbusch. Thay have received widespread occurrence in many countries of a world (concultivar Weiko). The German geotype is characterized by early maturity, small-seediness and relative thermoneutrality in comparison with the gene pool from Mediterranean. It consists from 3 concultivars.
Concultivar Lueneburger. It is shown by alkaloidal cultivars used on green fertilizer and in horticulture. It is characterized by high productivity of green mass and by large brightly yellow flowers.
Cv. Lueneburger gelbe (k-391). This cultivar was created by the method of selection of highly productive plants from a local material. It concerns to var. luteus. Their seed mass per1 plant makes 10... 15 g, mass of a green part of plant in a phase of flowering is 200... 250 g and mass of 1000 seeds is 160 g. Duration of vegetational period makes 125... 135 days in conditions of Kiev area. This cultivar is unstable to fusarium wilt and virus diseases. Its usage is possible in breeding as the source of high productivity of green mass (for silage). Cvs. Von Kolbens Vienauer and Paulsens Gelbe concern also to this concultivar.
Concultivar Weiko. It integrates first fodder (sweet) cultivars of the Weiko serial, created by v. Sengbusch, Troll and Zimmerman. The method of stepwise hybridization has widely used in selection of this serial, which has permitted by step-by-step to improve valuable characters and to receive a number of new valuable characters and properties of plants (fast pace of initial growth, white colouring of seed, non-shattering beans without pubescence on them.
Cv. Weiko IV (k-1773) is characterized by the fast rate of growth, not-shattering and weakly pubescent beans, yellow flowers and white seeds (var. leucospermus). Its seed mass per plant makes 20... 24 g, mass of green part of plant in a phase of flowering is 80... 90 g and mass of 1000 seeds is 140... 170 g. The contents of protein in seeds is 42.2 %. Duration of period of vegetation makes 120... 135 days in conditions of the Kiev area. This cultivar has ensured the productivity of green mass on the level of 42.4 t/ha, seed productivity has made 2.14 t/ha on testing plots in Zhitomir area. However, it is not resistant to fusarium. This cultivar was used as initial material for creation of more highly productive cultivars in many countries of the world. The cultivars Weiko I (k-1402), Weiko II (k-1594), Weiko III (k-1656), Muencheberger Gelbe suesslupine (k-337) etc. concern to this concultivar.
Concultivar Borluta. It integrates modern German breeding cultivars created in main on base of concultivar Weiko, but distinguished by resistance to fusarium and by higher productivity.
Cv. Borluta (k-2200) is the highly productive cultivar in agriculture of many countries. It is also the high-performance source of resistance to fusarium it breeding programs. The character of resistance is transferred to it from wild Portuguese and Italian forms collected by expeditions of Wuttke (1943) and Lamberts (1955). The character of resistance is controlled by one recessive gene (Лукашевич, 1989). The seed mass per plant at this cultivar makes 20... 24 g, mass of green part of plant in a phase of flowering is 130... 140g. The content of protein in dry matter of green mass is 19 %. The maximal productivity of green mass has compounded 105.0 t/ha; seeds yield has made 2.9 t/ha at production-line testings in Russian conditions. Duration of vegetational period makes 120... 140 days. Cvs. Bornova, Trebatch, Topaz, Refusa etc. concern to this concultivar.

Polish agrogeotype

It integrates fodder (sweet) cultivars of Poland. They were created after publication in 1932 in USSR method of rapid analysis of plants on low alkaloidness. Yellow lupin is the species, which is most widely cultivated in Poland in view of its low requirements towards soil conditions. It is cultivated in Poland on acidic sandy soils with the purpose of them reclamation and obtaining of highly-quality forage. The cultivars of this agrogeotype are characterized by shorter period of vegetation in comparison with German agrogeotype. This agrogeotype actuates 3 concultivars.
Concultivar Ekspress. It actuates fodder cultivars with not falling beans and fast paces of initial growth, but not resistant to fusarium. They are created on the basis of the German cultivars from the serial Weiko and from local materials also.
Cv. Ekspress (k-1684) is created by Winiarski on experimental station Przebędowo by hybridization of accessions Poznański x Oleski. It is characterized by enough high productivity of green mass (200... 210 g/plant) and fast growth rate (gene promtus). The seed mass per plant at this cultivar makes 15... 19 g, mass of 1000 seeds is 120 g. The contents of protein in seeds is 40.0 %. The plants of this cultivar are not resistant to fusarium. 86 % of plants have appeared staggered by fusarium at test on an infected background. Duration of vegetational period makes 120... 130 days in the Kiev area. Cvs. Sam (k-1736), Mazowiecki (k-1738), Lima (k-1969), Bas (k-2174) concern to this concultivar.
Concultivar Afus. It integrates cultivars with high resistance to fusarium and created with usage as initial material of the Portuguese forms from Iberian geotype.
Cv. Afus (k-2084) is created by Z.Tomaszewski and I.Kubok by hybridization of the Portuguese form M - 385 with Polish low alkaloidal and fast growing sample N 679 (Таранухо, 1977, 1980; Kubok, 1988). This cultivar has yellow flowers and white seeds with black spots and two light arcs (var. maculosus). Their resistance to fusarium is conditioned by dominant alleles of two genes (Лукашевич, 1989). It is rather late cultivar but with fast growth rate on the initial stages (gene Rapidus). The plants have small seeds and high green mass yield. Beside resistance to fusarium also gramine alkaloid was brought to Polish breeding by this cultivar, which, despite low toxicity causes a certain worsening of the fodder palatability (Kubok, 1988). The seed mass per plant makes at this cultivar 22... 24 g, mass of green part of plant in a phase of flowering is 230... 250 g and mass of 1000 seeds is 125 g. The contents of protein in seeds is 41.9 % (Каталог мировой коллекции ВИР, вып.511, 1990). Duration of vegetational period makes 130... 155 days. The cultivar is the good donor of high productivity and resistance to fusarium, which one will widely be used in the breeding programs of many countries. Cvs. Cyt (k-2398), Tomik (k-2146) etc. concern also to this concultivars.
Concultivar Iryd. It consists from modern breeding cultivars distinguished by early maturity, high productivity, thermoneutrality, immunity to fusarium and tolerance to virus deseases.
Cv. Iryd (k-3020) is created on experimental station Wiatrowo by a hybridization of spontaneous early mutant with resistant to fusarium sample Refusa Nowa (Swięcicki, 1985). It differs by fast rate of initial growth, immunity to fusarium and resistance to lodging and virus diseases. The virus infection is not transmitted through its seeds. Its seed mass per plant makes 20... 22 g, mass of green part of plant in a phase of flowering is 180... 220 g. The contents of protein in seeds is up to 44 % and in dry matter of green mass about 20 %. The mass of 1000 seeds is 130 g. This cultivar has the broad usage in the breeding programs of Belarus and Russia as the source of complex mentioned above useful characters. Cvs. Aga, Baltyk, Orbit, Reda, Topaz, Ventus, Juno etc. are included in this concultivar.




East-European agrogeotype

It actuates fodder cultivars of Belarus, Lithuania, Russia and Ukraine. The basic material for their creation was served accessions and cultivars from Poland and Germany, and also wild forms and local varieties from eastern Mediterranean. The cultivars relating to this agrogeotype are created for different soils and climatic zones and have difference in the assignation (for grain, silage or combined usage), in the level of productivity and in duration of vegetational period (Дюбин, 1974; Таранухо, 1980).
Conculivar Akademichesky. It integrates breeding cultivars of silage-grain usage created in the period from 1945 to 1990 years. They differ by enough high productivity of seeds and green mass. However, thay are not resistant to fusarium.
Cv. Akademichesky 1 (k-1947) is created in the Byelorussian Agricultural Academy by the method of individual selection of early plants with their subsequent hybridization among themselves (Таранухо, 1980). It belongs to var. luteus. This cultivar takes until now the big areas of sowing in Belarus and Russia, and also is the standard at tests of new cultivars. It is productive both on seeds and on green mass. The average yield of seeds has compounded from 1.25 up to 2.66 t/ha and yield of green mass from 32.7 up to 61.9 t/ha during the tests. The content of protein in seeds makes 42.3... 47.2 %, and it is on the level of 18.5... 20.4 % in dry matter of green mass (Таранухо, 1977). The mass of 1000 seeds makes 130 g and duration of vegetational period is 95... 120 days. This cultivar has rather resistance to virus diseases, however is not resistant to fusarium. It is the effective course of early maturity in breeding programs of many countries. Cvs. Быстрорастущий 4 (k-1509), Быстрорастущий 81 (k-1943), Балтеи (k-1631), Рокинсkий 58 (k-2053), Носовский белосемянный (k-1531) are included in this concultivar.
Concultivar Kastrychnik. It integrates modern resistant to fusarium and others diseases breeding cultivars of Belarus and Russia with duration of vegetational period 90... 120 days.
Cv. Kastrychnik (k-2651) is created in Byelorussian Institute of Agriculture by hybridization of cvs. Afus and Akademichesky 1 with subsequent individual selection on infected by fusarium plots (Лукашевич, 1988). It belongs to var. luteus and differs by fast rate of growth and heightened resistance to fusarium. Their seed mass per plant makes 23... 25 g, mass of green part of plant in a phase of flowering is 200... 250 g, mass of 1000 seeds is 115 g. The contents of protein in the seeds is on the level of 42.3... 46.5 %, the content of protein in dry matter of green mass is 16.5... 18.3 %. The maximal yield of seeds (3.3 t/ha) and green mass (86.2 t/ha) is obtained in the Gomel area of Belarus. Duration of vegetational period makes 90... 115 days. This cultivar is widely cultivated in Belarus, Ukraine and Russia. It is widely used also in the breeding programs as a source of early maturity and high productivity. Cvs. БСХА-382, Bryansk 6, Нарочанский and others concern to this concultivar.
Concultivar Kopylovsky. It actuates modern resistant to fusarium cultivars created on the Ukraine with duration of vegetational period 105... 130 days.
Cv. Kopylovsky (k-2601) is created in Ukrainian Institute of Agriculture by stepwise hybridization of cvs. Niko and Shwako with resistant to fusarium mutants. It has black seeds (var. niger). This cultivar has exhibited high resistance to fusarium at test on different artificial infected backgrounds, created by miscellaneous methods in many locales of Russia, Belarus and Ukraine. It testifies about complex resistance of this cultivar to the different strains and races of this pathogen. The cultivar differs by high productivity of seeds and green mass. The seed mass per plant makes 24... 26 g, mass of green part of plant in a phase of flowering is 240... 250 g, mass of 1000 seeds is 105... 120 g. The contents of protein in seeds makes 47 %, its content in dry matter of green mass is 18 %. The maximal seed productivity has compounded 2.5 t/ha and yield of green mass has reached 66.0 t/ha at production-line testing (Синицын et al., 1983). This cultivar is highly effective source of resistance to fusarium. Cvs. Volynsk 1, Martin 2 , Факел etc. are included in this concultivar.
Concultivar Zhitomirsky
. It integrates sorts of yellow lupin with determinate branching, all lateral branches at which are finished by flowers or flower truss. This character causes early maturity and manufacturability at harvesting. Such cultivars introduce the special value at cultivation and usage on a grain. The forms of yellow lupin with the determinate branching are obtained for the first time by breeder Valovnenko (Валовненко, 1974).
Cv. Zhitomirsky anniversary (k-2149) is created on Zhitomir agricultural experimental station with usage of mutagenesis. The plants have yellow axillary flowers and white seeds. The lateral shoots absent or shortened (var. leucospermus, f. ucrainicus). The seed mass per plant makes 12... 15 g, mass of green part of plants in a phase of flowering is 40... 45 g and mass of 1000 seed is 135 g. The contents of protein in seeds makes 41.35 % (Каталог мировой коллекции ВИР, вып. 511, 1990). This cultivar differs by short vegetational period (90... 105 days). The plants ripen on 12... 20 days earlier, than at others cultivars with conventional branching. The productivity of this lupin form with determinate branching usually is lower than productivity of conventional forms with traditional branching in calculation on one plant. However, this cultivar provides the yield of seeds from a unit of area at the level of others cultivars with usual branching at the thickened sowing rate, and the quality of seeds is much higher at it. Concerning green mass, this cultivar succumbs to cultivars with traditional branching on productivity (30... 35 t/ha). Cvs. Искорость, Детер 5774/83, Лимонный, Юбилейный etc. concern also to this concultivar.

THE HISTORY OF LUPIN DOMESTICATION


THE HISTORY OF LUPIN DOMESTICATION



B.S. Kurlovich



Two conditions are essential for the formation of Vavilov’s centers of origin (diversity) of cultivated plants (Вавилов, 1987B): the existence of plants suitable for introducing into cultivation in the local plant resources, and the presence of an age-old agricultural civilization. For lupin, both of these conditions are found in the Mediterranean region and on the American continent (territory of the modern Peru). These two centers of development of wild lupins were simultaneously the first places of them domestication. The history of lupin domestication has embraced more than four thousand years. Although lupin development processes in the Mediterranean and American centers went separately, they were similar enough.

The earliest archaeological reports on lupins are referred to the XII dynasty of Egyptian Pharaohs (over 2 thousand years BC). In their tombs, seeds of Lupinus digitatus Forsk., already domesticated in those times, were discovered. Several empty pod valves seven seeds of this species were also retrieved in the tombs of this dynasty dated back to the 22nd century BC. They are the most ancient evidence of lupin in the Mediterranean (Zhukovsky, 1929).

Andean pearl lupin (L. mutabilis Sweet.) was domesticated on the American continent by ancient inhabitants of the territory of the present-day Peru. The imprints of its leaves and seeds on the rock stored in the national museum of Lima testify to the ancient origin of this species. This rock dates back to the pre-Inca culture which existed in the 6th – 7th centuries BC (Майсурян, Атабекова, 1974). The forms of lupin cultivated in this period were a little different from the present-day ones. Their seeds were larger than those of wild forms. This may serve as proof that lupin was introduced into cultivation on both hemispheres even earlier.

Modern domesticated lupin plants have partially changed their former exterior and biological properties. Breeding practice enhanced the development of plant qualities useful for man, and induced the loss of a number of traits characteristic of wild forms. There were repeated upsurges of common interest in lupin cultivation and its abatements in connection with arising difficulties.
It is possible to split the history of lupin domestication into the following conventional stages:

· Primary domestication of white lupin in ancient Greece and Egypt (earlier than 2000 BC) with the purpose of producing grain which was used after soaking as food for man and feed for animals, and also utilized in cosmetics and medicine.

· Beginning of utilization of white lupin as green manure in ancient Rome and, subsequently, in others Mediterranean countries (about 800-1000 BC).

· Primary domestication of Andean pearl lupin ”Tarwi” on the American continent (600-700 BC).

· Beginning of domestication of yellow and narrow-leafed lupins for green manure production in Mediterranean countries and afterwards in Germany.

. Obtaining of the first low-alkaloid (sweet) forms of lupins in Germany (1927-1930) and in Russia (1929-1932). The method of their production was for the first time published in Russia (Иванов et al., 1932). After this discovery, lupin received the status of a fodder crop.

. Domestication of new lupin species in Australia, Russia, Finland and other countries (L. cosentini Guss, L. atlanticus Gladstones, L. polyphyllus Lindl. etc.).

· The modern stage of lupin cultivation, when lupin received the status of an alternative to soybean in world agriculture as a source of protein and another useful substances. Lupin resources will be utilized in many branches of the world economy.

The beginning of the history of lupin cultivation in the Old World is often associated with the times of the ancient Egyptian civilization (Zhukovsky, 1929; Майсурян, Атабекова, 1974; Таранухо, 1980). It is, however, more likely (Kurlovich, 1998) that originally white lupin was introduced into cultivation in ancient Greece where its greatest biodiversity was concentrated and wild-growing forms have been preserved until nowadays (ssp. graecus). Here, on the Balkan Peninsula, representatives of another subspecies of white lupin (ssp. termis and ssp. albus) turned wild and grows now in natural environments. Besides, the Grecian genesis of cultivated lupin is testified by lupin’s Greek name “termis”, that may be translated as “ardent”. Until now, in may countries of the world water-soaked and boiled lupin seeds are sold on markets and in bars as delicacies (like sunflower seeds). Utilization of white lupin was probably the same in ancient Greece.
From the rich diversity of lupins on the Balkan Peninsula, ancient Greeks obviously selected a wild white lupin form with white large seeds and light flowers. Other cultivated plants were also selected and domesticated basically according to the same traits. In view of this, it is possible to regard the Balkan Peninsula as the primary center of origin (diversity) of white lupin. Domesticated lupin is mentioned by Hippocrates of Cos, a Greek scientist (400-356 BC), who noticed that the pods of lupin were less harmful than those of Cicer arietinum. He considered lupin as most nutritious food. The flour made of lupin, in his opinion (Bd. VIII, p. 369), makes the face beautiful. The works of Theophrastus (372-288 BC) already contained fragments of information concerning cultivation of lupin on sandy soils (Zhukovsky, 1929). Theophrastus also marked the wild-growing habit of lupin, owing to which it did not require maintenance. It is possible to conclude from the descriptions of Theophrastus that cultivated lupin in Ancient Greece belonged to sp. L. albus. The detailed description of plant morphology and growing methods testifies that even in those times the culture of white lupin was widely spread in Greece. White lupin dispersed step-by-step from Greece to adjacent countries, in particular, to Egypt and Ancient Rome. The forms with white seeds and pink-and-blue or light-pink flowers (L. termis) spread mainly towards the south (Egypt, Libya and Palestine), while the forms with white seeds and grayish-blue or white flowers (L. albus) moved to the west (Apennine Peninsula and farther). White lupine (L. albus ) is still grown in Greece, where the wild ssp. graecus is also spread. The main purpose of lupin cultivation in Egypt was production of seeds widely used for food. However, there is no information about utilization of lupin in Ancient Egypt for feeding cattle or producing green manure (Майсурян and Атабекова, 1974). It is not revealed by the imprints of lupin on historical monuments anywhere in Egypt In view of this, there are grounds to assume that lupin started to be cultivated in Egypt not earlier than in 330 BC. Egyptians received this crop from Greeks in already domesticated condition (Gladstones, 1974). The broad and full overview of the developed culture of lupin in Ancient Rome is presented in the work of Zhukovsky (1929) where the materials of many ancient writers were generalized. For example, Plinius (23-79 BC) wrote about lupin (Book. XVIII, 133):
“… It is used both by man and by ungulates. It is necessary to clean it after rain. In this case, its grains do not fall out and are not lost at harvesting. It is a plant so wonderfully agreeable with the soil. At first, during the day it rotates together with the sun and shows time to the farmer even under a cloudy sky. It is the only plant sown without plowing (!). Lupin loves stony, dry and even sandy places. It does not require any maintenance at all. Fields and vineyards are improved by this crop. It does not require manure at all, being in itself the best fertilizer. It is the only plant which does not require any expenses or labor. It is the first to be sown and the last to be harvested, approximately in September. Modey (unit of weight or volume) of lupin per day makes an ox stout and strong. ”. Famous Marcus Terentius Varro (116 – ca. 27 BC) reported: “lupin is plowed instead of manure in the lean soil while it has not yet produced pods and sometimes stalks with beans”. Galen (AD 129-199) wrote about numerous applications of lupin, mainly about leaching its seeds in sweet water in order to use them for food. Pedanius Dioscorides (AD ca.20 – 70) in his 20th book recommends lupin as a remedy against abscesses, birthmarks and scab of sheep. He advised to sweeten the leached seeds, mill them into flour, and to add several drops of vinegar for whetting appetite. Columella (fl. 1st century AD) put lupin above other legumes. He recommended lupin as the best fertilizer “for depleted vineyards and fields, for exhausted soils”. Thus, it is possible to surmise that the value of lupin as green manure and in general its wide usability in agriculture was first acknowledged in Classic Rome, when the new second stage of lupin cultivation began. Romans, as well as Greeks, were fully aware of the medical and pharmaceutical value of lupin. Seeds, plants and decoction left after seed soaking were used as drugs. Plinius mentioned sixteen ways of applying lupin in medicine. It was also used as means of protection fruit trees from diseases. From Greece, and subsequently from Egypt and Ancient Rome, lupin was spread over all Mediterranean regions. In literature there are crumbs of information about the well-developed culture of lupin in the ancient and medieval Italy, France, Spain, Portugal, Algeria, Tunisia, Sudan, Israel and Turkey (Gladstones, 1974; Maissurjan and Atabiekova, 1974; Swęcicki, 1988).
Concurrently with white lupin, yellow, narrow-leafed and other species of this crop started to be cultivated as sources of green manure and as ornamental plants, acquiring most high-yielding and beautiful forms among wild plants. Domesticated lupin spread from the Mediterranean area to other countries included in the so-called spheres of influence (Sinskaja, 1969). To these spheres of influencing she attributed, first of all, Central and Northern Europe, as well also Asia and Africa. In 1927, N.I. Vavilov collected white lupin at the river-head of Blue Nile in Abyssinia, where lupin had most likely come from Egypt. From Greece and Turkey white lupin penetrated northwards and eastwards, and reached Western Georgia, where the culture of white lupin has survived until recent times after many ages of cultivation under the local name of “hanchcoly”, which in Georgian means a bitter bean (Либкинд, 1931; Майсурян and Атабекова, 1974; Kurlovich, 1996).

The parallel history of Lupinus mutabilis in the Andean highlands of South America is documented to a lesser degree. Zhukovsky (1929), Hondelmann (1984), Gladstones (1998) cited archaeological evidence of its cultivation in the 6th or 7th century BC, while in the Nazca (AD 100-800) and subsequent civilizations L. mutabilis formed a regular part of the crop rotation. Brücher (1968) notified that the Indians had removed bitter taste of seeds by washing them. It was impossible to remove alkaloids completely, and poisoning sometimes still occurred.
Lopez-Bellido and Fuentes (1986) commented fertility-enhancing properties of lupins and reported their use in Andean religious rites and festivals. Well-known was also the curative role of alkaloids in the treatment of cardiac diseases, rheumatism, malaria and internal parasite infections, similar to the functions attributed to white lupin in Rome.
The appearance of the Spanish and Portuguese conquistadors in the 16th century started a gradual decline in native Andean agriculture. The new society ignored lupins, and their cultivation became restricted to marginal subsistence agriculture in a few sites on very high altitudes. The interest toward lupin has been revived only in the last decade or two (Gladstones, 1998).
The scraps of information about lupin, at first as a medicinal plant, date back to the Middle Ages, like in Central European countries. However, special agronomic literature on lupin in Central Europe came forth later. The agricultural history of bitter lupins in northern Europe began in 1781. The first impetus came from King Frederick II (Frederick the Great) of Prussia, who personally sent for seed of white lupin from Italy with a view to use it for improving the poor soils of northern Germany (Maissurjan and Atabiekova, 1974; Hondelmann 1984, 1996). But this attempt, as well as later northern European efforts with L. albus, for the most part failed, because the soils were too poor for it and the available genotypes was too late maturing in the region's short summer growing season. In this respect, the idea has appeared to use more early-ripening lupins: yellow lupin (L.luteus) and narrow-leafed lupin (L.angustifolius). Earlier, these species had already been effectively used as green manure in Morocco, Portugal and other Mediterranean countries (Klinkowski, 1938).
Then, in 1841 a farmer named Borchard tried growing the garden yellow lupin, L. luteus. This was much more successful, and despite the continuing official support of L. albus, the cultivation of L. luteus and L. angustifolius by 1860 had become an essential part of agriculture all over the acid sandy soils of the Baltic coastal plain (Hondelmann 1984; Gladstones, 1998). These species showed extremely high effectiveness as green manure crops, as witnessed by the experience of Dr. Albert Schltz of Lupitz (known as Schutz-Lupitz). On poor sandy soil in his estate, with the help of lupin he managed in 10-12 years to double the yield of potatoes and rye, and to pass overt to wheat on the best sites. The results are widely known. Owing to him, yellow and narrow-leafed lupins began to be cultivated on a wide scale in many countries with the purpose of increasing fertility of poor sandy soils (Maissurjan and Atabekova, 1974). Besides, the experience of Schutz-Lupitz served as a catalyst to the discovery of the phenomenon of lupin nitrogen-fixing ability, as a result of which his practice has also obtained theoretical explanation.

After these experiments (the middle of the 19th century), the next stage in cultivating several lupin species for green manure and other purposes was initiated. A little later, however, with the beginning of mineral fertilizer production, the interest in lupin cultivation for green manure in Germany declined once more, though, on the other hand, the need for seeking low-alkaloid fodder forms emerged.
In the middle of the 19th century, lupin spread from Germany to Poland where there was abundance of mild sandy soils requiring reclamation. Cultivated at that time were bitter-tasted populations of yellow and narrow-leafed lupin, which were plowed down as green manure, or used, despite their bitter taste, as a feed for sheep. After the initial expansion of lupin cultivation, its temporary decrease, especially in seed production, occurred at the end of the 19th century, probably due to frequent poisoning of animals with alkaloids (Kubok, 1988). During World War I, lupin cultivation began to grow continuously owing to the necessity of widening food production areas to light soils, with simultaneous decrease of mineral fertilizer production. At that time, lupin breeding activities were started in Poland.
The first breeding works involving populations of bitter narrow-leafed lupin were carried out as early as at the end of the 19th century by Sempołowski at Sobieszyn. Sypniewski, who bred a number of bitter narrow-leafed cultivars such as Puławski Wezes­ny and Puławski Wysoki, and continued his works in l920s at the Institute of Puławy. He was one of the first breeders, who began systematic genetic studies on this species (Kubok, 1988).
In Russia, the first mention about lupin as an ornamental plant appeared in 1811 in the papers of the Free Economical Society. Utilization of lupin for green manure came to Russia from Poland when it had been divided into three parts (1772, 1793, 1795) and partially incorporated in the Russian empire. The first harvests of lupin for green manure were reported beginning from 1903 in Chernigov province. In the following years the centers of lupin studies and breeding were organized: Novozybkov Experimental Station in Bryansk province, Bieniakonsk Experimental Station on sandy soils of the forest-steppe zone in the western part of the Russian Empire (now it is a territory of Poland), and also a number of facilities in Byelorussia and the Ukraine. Thanks to their research activities and released cultivars, the area under lupin in the European part of the Soviet Union by 1935 reached 100 thousand hectares (Майсурян and Атабекова, 1974).
Lupin has always attracted attention by its unpretentiousness and ability to grow on sandy acid soils where cultivation of other crops would be economically inexpedient, as well as by its high content and quality of protein and oils. Prof. D.N. Pryanishnikov (Прянишников, 1931, 1962) called lupin a surprising plant and compared it with “the barrel of honey”, in which, however, there is a hateful “spoon of tar”. He meant the presence of alkaloids in lupin, causing not only bitter taste but also toxicity for man and animals. In view of this, the problem of finding alkaloid-free forms of lupin was repeatedly roused in the history of this crop (Roemer, 1919; Прянишников, 1920,1923). Prof. E. Baur who in 1920ies was Director of the Kaiser Wilhelm Institute in Berlin later supported this idea. He cited Vavilov’s Law of homological series in hereditary variation as the reason to connect low alkaloid content with mutations, which sometimes happened in the nature or were induced synthetically. One of the students who in 1927 listened to Baur’s lectures was R. von Sengbusch. After studying the earlier works of Prof. Pryanishnikov and hearing the lectures of Prof. Baur, he took up the challenge of devising a method to be used in screening the diversity of lupins. R. von Sengbusch worked out the method of fast definition of alkaloids, analyzed great diversity of plants and found among them low-alkaloid (sweet) forms. These forms were promptly propagated and used as initial material for creating the first fodder cultivars of yellow and narrow-leafed lupin. The seeds quickly multiplied. Finally, before World War II, the area under fodder (sweet) lupins in Germany exceeded 100 thousand hectares.
But the new methods and results of German breeding had been secret and were published only in 1942 (Sengbusch, 1942). The exclusive right for the sale of seeds was delegated to a private German corporation. In the meantime, the export of seed was impeded by strict clearance check-up. Hence, a fast method of detecting alkaloids by means of solution of Burhard was developed in the Institute of Plant Industry (VIR) under the leadership of Prof. N.N. Ivanov. It was immediately and for the first time in the world published with the foreword by N.I Vavilov. Vavilov expressed that "... We do not conceal the results obtained by us but make them generally known, in order to interest in this discovery the scientific workers of our country, as well as those abroad. In the present work the Institute of Plant Industry publishes the results of its researches as well as an instruction for the determination of alkaloidless lupins" (Иванов et al., 1932). With the help of the new method, the first sweet cultivar of yellow lupin, Yubileiny, and also many low-alkaloid forms of L polyphyllus were bred in 1932 in VIR.
The above-mentioned publication by VIR as well as the discovery of German scientists was a capstone of the modern breeding work with low-alkaloid fodder (sweet) lupin in the whole world. Lupin obtained the status of a valuable fodder crop plant.
In the former USSR, the first breeding program of releasing fodder lupin cultivars was launched by Fedotov, Sharapov and other scientists of VIR, by Bozenova of Novozybkov Experimental Station in Bryansk province, and by Swirski of Minsk Experimental Station. Successful work with different species of lupin was organized in the following years at Chernigov and Polessk Experimental Stations in the Ukraine, at Moscow Agricultural Academy, Byelorussian Agricultural Academy, Byelorussian Institute of Agriculture, Byelorussian State University, Grodno Experimental Station, Ukrainian Institute of Agriculture, Institute of Agriculture of the Central Non-Black-Soil Zone, etc. As a result of these works, the area under lupin in the former USSR by 1959 had reached 1132 thousand hectares (Майсурян, Атабекова, 1974). Particular progress in yellow lupin breeding was attained in postwar years by K. Savichev (Novozybkov Experimental Station) and G. Taranukho (Byelorussian Agricultural Academy). Savichev’s cv. Bystrorastushchy 4 in due time was cultivated in 30 provinces of the former USSR.
Later, Prof. Taranukho released an ultra-early cultivar, Academichesky 1, that is still recognized as the reference point of new breeding achievements. Breeder V. Valovnenko was the first in the world to discover the form of yellow lupin with determinate branching. He was the author of a series of cultivars with this valuable character (Zhitomir anniversary, Iskorost, etc.). It is for a long time already that the breeding of narrow-leafed and white lupins is conducted in Moscow Agricultural Academy. Super-early cultivars of narrow-leafed lupin were created there: Northern-3 (by Majsurian), Timir-1 (by N. Pukhalskaja), Ladny (by N. Klochko). Prof. G. Gataulina in cooperation with other research institutions bred ultra-early fodder cultivars of white lupin (Start, Manovitsky, etc). Great progress was made by the Ukrainian Institute of Agriculture (V.Golovchenko, O. Golovchenko, N. Solodyk and other breeders) in white lupin breeding. Cvs. Kievsky mutant, Primorsky, Gorizont, Ukrainsky, Dnieper, Pishchevoy etc. were produced by the method of hybridization and induced mutagenesis with the use of accessions of the Georgian ecotype and Palestinian geotype. The Institute of Agriculture of the Central Non-Black-Soil Zone released a series of fodder cultivars of narrow-leafed lupin, among which cv. Hemchinovcky 846 (authors: G.Debely, V. Fedotov and L. Kalinina) became the most widely known.
Fusarium resistant cultivars of yellow lupin (Kastrychnik, Narochansky, Zhodinsky) were created by Dr. M. Lukashevich, breeder of the Byelorussian Institute of agriculture, while cultivars of narrow-leafed lupin (Gelena, Selena, etc.) by the breeder N. Kuptsov. With the purpose of complex research on lupin-related problems, the specialized All-Russian Institute of Lupin was founded in 1987 in Russia (near Bryansk). It has undertaken large-scale work on breeding, seed production, cultivation technology and utilization of lupin. In 1960’s-70’s, lupin was cultivated for grain in ex-USSR on the area of about 600 thousand hectares, and the areas under this crop for green forage and green manure approached 2 million hectares (Такунов и др.1993).
Regretfully enough, now the areas under lupin in Russia have reduced (for more details see the last section). The method of fast alkaloid determination published in Russia was effectively used in Poland (Kubok, 1988). In a short period of time, plenty of fodder (sweet) cultivars of different lupin species were released in this country. More detailed information on the results of breeding work with lupin in Poland is presented in the section „Eco-geographic classification of lupins”. As a result of these works, the area under fodder lupins in Poland was expanded to 350 thousand ha (Kubok, 1988).
Situation with cultivation of lupins in other countries now is following. Australia Prominent success in acclimatization of narrow-leafed lupin and its other species was achieved by scientists in Australia. As a result of implementing the scientific project under the supervision of Dr. J.S. Gladstones (launched in the fifties), the area under fodder lupin in this country grew to exceed 15 million ha (Nelson, 1994). According to the data of Cox (1998), sweet lupin seed has been used as a feed since it was first produced within Australia. Initially the most favored application was its use as a sheep feed. Ongoing research investigating its nutritional value across a range of ruminant and monogastric livestock types resulted in significant expansion of its use as a feed to the point wherein sweet lupin seed is utilized in many intensive and non-intensive Australian livestock industries. Australian domestic usage of sweet lupin seed is estimated to be in excess of 350,000 t per annum.
Competitive protein products include imported soybean meal, canola meal, field peas and other grain legumes. Sweet lupin has been successfully bred into a new crop plant around the world. International markets for lupin seed have been established. Australia, as the world's largest exporter of lupin (L. angusifolius), is the dominant force in the international marketing and trade of lupin.
We have used the information of Baer, E. von, (1986, 1989, 1994) and Cox (1998) to illustrate volumes of utilization of sweet lupin seed (primarily of Australian and Latin American origin) in different countries and on major markets:

Europe
The Netherlands, Belgium and Germany. 150,000-250,000 t per annum – mostly compound feed formulae and dairy rations. Sweet lupin seed is commonly used in dairy rations. It competes as a protein and energy source with competitive products often including corn gluten feed, palm and corn products, rapeseed meal, citrus and beetpulps. The highly sophisticated industry, with access to a broad range of ingredients, creates a very competitive market environment. The industry is generally not committed to any commodity and the major companies constantly trade in and out of materials, depending on price relativity. The market has the depth to absorb large quantities of sweet lupin seed at varying price levels, depending on ration formulation (Cox, 1998).
Spain and Portugal. 120,000 t per annum – all livestock types. Consumption is highest in the ruminant sector, with limited utilization in monogastric diets at low inclusion rates. Chief protein competitors include soybean meal, corn gluten feed, rapeseed and sunflower meals, meat and bonemeal. Sweet lupin seed was initially targeted into limited geographical areas specializing in cattle production in order to extract the highest market price (Cox, 1998).
Italy. 20,000-50,000 finishing beef cattle. Ready access to inexpensive domestic soybean and sunflower meals pre­clude wide-scale usage on a cost comparative basis. High freight costs for small parcel shipments is also a significant limiting factor (Cox, 1998).

Asia (Cox, 1998).
Republic of Korea. 105,000 t per annum – all livestock types, aquaculture. A major trading company in Korea established a sweet lupin seed dehulling plant in the late 1980s to enable the seed fractions to be used most effectively over all animal types. The major protein competition in this market includes domestic and imported soybean meal.
Japan. 115,000 t per annum – cattle. The whole seed is flaked and incorporated into a balanced diet of partially processed ingredients typically comprising flakes of corn, sorghum and vitamin/ mineral supplements. Domestic and imported soybean meals have been partially displaced by sweet lupin seed.
Indonesia. 5000-8000 t per annum – human consumption, cattle. The hull material is utilized in cattle diets. Indonesia is the first commercial export market for Western Australian lupins for human consumption.
Israel. Sales were conducted for several years in the mid 1980s for use in cattle diets. Marketing opportunities tend to be limited because of price competition from cheap alternative protein sources, particularly cottonseed meal. High freight costs from Western Australia often present difficulties (Cox, 1998).
Taiwan. Significant quantities were sold to feed compound in the 1980s and now this market is regularly taking small parcels of sweet lupin seed (Ford, 1990, Cox,1998).
In addition to the key and regular market outlets of the European Union, Japan and Korea a number of other markets have also been investigated by Australian farmers and businessmen (Cox, 1998). The Middle East, United Arab Emirates, South America, Thailand and the Philippines have feed industries capable of taking a significant portion, if not the bulk, of the Australian sweet lupin seed crop. Although many of these countries have imported sweet lupin seed in the past, they have not been able to compete consistently against the price offered for sweet lupin seed by key traditional markets. Often this is due to the availability of cheaper protein meals from countries in close proximity, restrictions on the allowable import volume of imported protein meals or high tariff levels on sweet lupin seed. These markets continue to evolve and may at some later stage provide valuable opportunities for Australian producers (Cox, 1998).

America

South America. South American farmers produce a number of lupin species (L. albus, L. luteus, L. angusti­fohus). However, greatest attention is given to local Andean lupin (L. mutabilis Sweet.), which is cultivated from ancient times (Gross, 1982). Many of these species have alkaloid levels in the seed that are much higher than those in Australian sweet lupin cultivars. But low-alkaloid (sweet) forms were found out also at this ancient species, which also has received the status of fodder crop and is widely cultivated in Latin America and in Europe. The work on breeding, development of cultivation technology, use and sale of lupins is most intensively developed in Chile, under the management of Erik von Baer. The large scale production and industrial utilization of lupin in South America will only be possible if a dynamic organization can be achieved that combines, in a profitable way, research, production, processing and merchandising of this product. This is feasible only with an integrated private and public effort, like in Australia. Chile achieved the stepwise increase of the area cultivated with lupins, from 329 ha in the season 1973/74 to 16,000 ha in 1992/93 (Baer E, 1986, 1989, 1994). The specific achievements and limitations are in the countries of Latin America. To make industrial utilization of lupin possible in Peru, it is necessary to grow sweet cultivars or to utilize the alkaloids from bitter varieties. The priority to obtain disease-resistant forms is established in Brazil (Baier, A.C. and Linhares, 1990). Lupins have to go directly into competition with other protein sources like soya-bean, fish meal and energetic sources like corn and others. Considering the lower prizes of these, it has to be calculated how much Lupin production can cost, and then it is necessary to guarantee to the user a steady provision of a standardized product all over the year (Baer E, 1994). Production made from lupin in Latin America (L. albus and L. mutabilis) is also exported as a food to countries such as Egypt, Israel and the Middle East in competition with Australian resources. Products from sweet lupins have been cleared for use as food supplements in milk substitutes for intervention programs (Aguilera, 1988). An industrial plant has been producing up to 50% of protein requirements for milk substitutes with typical formulations being 6~73% wheat flour, 12% lupin meal, 12% skim milk and 5% soybean oil. Other uses are as extrusion - pre-cooked lupin flours and additives in sausages and jams (Cox, 1998). Thus, South America keeps presenting good possibilities for the production, consumption and perhaps exportation of lupins (Ford, 1994). In the future, it can become the worthy competitor of Australia.
North America. Narrow-leafed lupin is growing in the USA in spite of the huge potential of this country in producing soybean. Trials on others lupin species will be made there (Clapham and Willcott, 1995). Characteristics of the cultivars released in the USA are given in the section “Eco-geographic classification of lupins”.

Gradual domestication of other new species of lupin began after the methods of searching low-alkaloid forms had been discovered. One of these species is the multifoliate or Washington lupin (Lupinus polyphyllus Lindl.), which is frequently called perennial or long-term lupin. It was brought in Europe from America and migrated to many European countries as a wild plant or the one turned wild. It has often been grown for green manure and for ornamental purposes. Prof. Pryanishnikov was an eager initiator of introducing multifoliate lupin in agriculture as a fodder plant. However, repeated attempts to develop fodder (sweet) varieties of multifoliate lupin (L. polyphyllus Lindl.) and other perennial species (L. perennis L., L. nootkatensis Donn, L. arboreus Sims.) cannot be recognized as essentially successful because of the difficulties induced by their perennial lifestyle and cross pollination habit (Kurlovich et al., 1995). It proved impossible to stabilize low alkaloid content in these cross-pollinated species. Owing to complimentary interaction of different genes determining this character, the alkaloid content tended to restore (Maissurjan and Atabiekova, 1974; Turbin and Anohina,1974). That is why bitter plants are found in the crop, preventing wide use of grass and seed for fodder. However, in the last years it has at last become possible to develop a technique for obtaining sufficiently stable fodder forms of L. polyphyllus at the N. I. Vavilov Institute of Plant Industry (Чекалин and Курлович, 1989; Курлович and Чекалин, 1992; Kurlovich et al, 1995). This technique is discussed in detail in the section “Methods of breeding”.
It helped to breed a fodder variety of multifoliate lupin called Truvor, which now undergoes state trials in the Ukraine. Under the conditions of Northwestern Russia, positive results have been reported with the commercial cultivar Pervenec (first sweet variety), bred by us and included in the State Catalogue of Breeding Achievements in Russia. Since 1996, we have been conducting breeding work with multifoliate Washington lupin in Finland. The cultivar of fodder multifoliate Washington lupin SF/TA has been bred also in Finland by Aniszewski (1993).

Application of the developed methods of breeding fodder (sweet) multifoliate Washington lupin (L. polyphyllus Lindl.) for low alkaloid content provides a possibility to produce high-yielding fodder (sweet) cultivars of this perennial species for different eco-geographical conditions with low alkaloid content and a set of valuable biological and commercial characters. Multifoliate Washington lupin can be grown for fodder and seed in the countries with limited agricultural resources and with a deficit of vegetation period for other lupin species, particularly in the northwestern areas of Russia (Kurlovich et al., 1995), Finland, (Aniszewski,1993), Sweden (Maissurjan and Atabiekova, 1974), and other northern countries. Introduction of perennial fodder (sweet) multifoliate Washington lupin into agricultural production of many countries will stimulate a significant increase in soil fertility, production of protein-rich fodder and, consequently, productivity of animal husbandry. These methods may also be applied to such cross-pollinated lupin species as L. nootkatensis Doon.. L. arboreus Sims., L. perennis L., L. elegans H.B.K., L.hartwegii Lindl., etc., promising for utilization in agricultural production.

The project on domestication of the rough-seeded lupins is successfully realized in Australia (Buirchell, 1994). There are very few cultivated lupin species adapted to fine-textured and alkaline soils. Some rough-seeded lupin species grow naturally on these soils in the Mediterranean region. Breeding of rough-seeded lupins in Western Australia has resulted in the first filly domesticated species L. cosentinii and is close to domesticating L. atlanticus and L. pilosus. In 1970s J.S. Gladstones bred Errequella-S, the first cultivar of L. cosentinii, which is characterized by low alkaloids, early flowering and soft-seediness. Now this cultivar is widely cultivated in the coastal area of Western Australia. Besides, low-alkaloid and white-flowered forms were identified in L. atlanticus and L. pilosus, which are included in large-scale breeding programs. These two species have great potential for cutivation on fine-textured alkaline soils in southern Australia (Buirchell, 1994, Buirchell and Cowling, 1998). Interspecific crossing with the fully domesticated L. cosentinii was used to transfer soft-seediness genes into L. atlanticus. Crosses between L. atlanticus and L digitatus have produced early flowering lines. Shatter-resistant pods, while not being detected in primary crosses of L. atlanticus and L. cosentinii, have been selected in crosses where L. digitatus is part of the combination. L. atlanticus now has all the domestication genes in a number of lines which will be combined into the fully domesticated cultivars. Mutants of L. pilosus have been also selected with soft seeds and low alkaloid levels. Further development of these species may provide alternative grain legume species for fine-textured and alkaline soils (Buirchell, 1994, Buirchell and Cowling, 1998).
There are also similar references on the attempts to introduce such species as L. arboreus, L. nootcatensis, L. succulentus etc. into cultivation process (Шутов, 1982, 1984).
At the 7th International Lupin Conference in Portugal, the Polish scientists W. Swęcicki, B.Wolko and K. Jach have reported the discovery of a new Mediterranean species Lupinus atlanticus Swiec. (Swęcicki et al., 1994).
The present stage in the development of lupin science and production is characterized by ever increasing interest in lupin as an alternative to soybean in the world agriculture in connection with the deficit of protein. Many countries (Australia, USA, Germany, Poland, France, Spain, Portugal, Chile, Israel, South Africa, New Zealand, etc.) are developing national breeding programs on different lupin species. International conferences on lupin are regularly held, and the International Lupin Association (ILA) has been founded by UNO. In Russia, where climate resources for cultivation of soybean are limited, lupin in the long term can play the same role as soybean has in the USA, i.e. it may become a highly effective source of protein, the means to increase the fertility of soils, a tool of environmental protection, raw material for medicine, perfumery, etc. Studying the history of lupin domestication leads to a number of generalizations that may be useful for obtaining new forms by means of conscious selection.
Two species of lupin (white and Andean) entered the agricultural production in two completely separate parts of the globe. Their domestication processes went on independently for thousands of years, but they were nevertheless similar enough. These processes were both connected with primitive agriculture with little involvement of breeding techniques. The other two crop species (yellow and narrow-leafed lupins) were domesticated much later and faster, when the agriculturists learned to carry out breeding work more consciously. And finally, such species as Lupinus cosentinii, Lupinus atlanticus and others are now being domesticated by methods of modern scientific breeding and biotechnology. The domesticated species (except maybe L. mutabilis) have their wild relatives growing in natural environments, and their habitat in the Mediterranean region significantly overlaps the areas of cultivated forms. These circumstances make it possible to compare directions and purposes of lupin breeding in different times and with different species. The basic purpose of cultivation of the majority of lupin species was and is the production of seed used for food even in the ancient times after being cooked and macerated. The changes that occurred in the process of lupin domestication have been connected, first of all, with the size and color of seed. Domesticated forms have, as a rule, larger seed of white color. For example, on the Balkan Peninsula there still remains the wild ancestor (ssp. graecus) of white lupin. Its plants have dark-brown and dotted seeds, much smaller in size than those of the domesticated white lupin forms. In the western hemisphere, from Canada to Argentina, several hundreds of lupin species are grown, and all of them are characterized by small seed size. Ancient Peruvians obtained the cultivated large-seeded species (L. mutabilis) from such a diversity of forms (Zhukovsky, 1929; Либкинд, 1931). Now it is not an easy task to establish a direct wild ancestor of this species. The absence of the forms of similar size in the wild vegetation of America, and easy crossing of L. mutabilis with many other wild-growing species of lupin testify that it has a hybrid origin (Майсурян and Атабекова, 1974). On the other hand, according to Blanko (1982), wholly wild ancestors use to grow within the area of traditional cultivation of L. mutabilis. However, their leaves are smaller and have narrow leaflets, the pods are smaller and dehiscent at maturity, and the seed are also much smaller, black or marbled, and water-proof. Natural cross-pollination is very common between wild and cultivated species (Pakendorf, 1970; Gladstones, 1998). However, on the American continent breeding practice also pursued the size of seed and their white color. White coloring of seeds occurs in the wild species in America seldom enough, although it can be found in many other plant genera as well. In the course of selection, such traits have appeared as soft-seediness, facilitating fast and amicable seed germination, and non-dehiscence of pods. There were anatomic changes in the structure of pods that provided the farmers with protection against shattering in the process of domestication. There were also similar changes in the chemical composition of seed. The same domesticated forms, besides the high content of protein (up to 40-50%), also acquired rather high oil content (up to 15-20%). Such content of protein and oil cannot be found in wild forms. Domesticated forms, as a rule, display a shorter vegetation period, and their prevailing type of branching is sympodial. Finally, fodder low-alkaloid forms have been obtained in both Mediterranean and American species. These facts testify to the similarity of requirements set by the farmers in different parts of the globe. It is possible to find out the same regulations with the changes that domesticated yellow and narrow-leafed lupins have undergone, though the selection process in these species advanced more consciously and speedily. The breeders were also striving to select large-seeded forms and preferably with white and soft testa of seed, increased protein content, sympodial and limited branching, non-dehiscent pods, low alkaloid content, and other characters useful for man.

The present-day process of domestication of new species (L. cosentinii, L. atlanticus etc.) is also going in the above-specified directions. However, owing to the achievements of modern science and with the help of novel bioengineering methods (biotechnology), it runs faster.

In view of this, it is possible to take for granted that the history of lupin domestication is not finished, and that many new valuable species within the genus Lupinus will be selected for human use.

BIOLOGICAL FEATURES



BIOLOGICAL FEATURES



B.S. Kurlovich, V.N. Dyubin and J. Heinänen



The response of lupins to different environmental factors

Response to light. It was shown by the researches conducted by the scientists of VIR, that the majority of grain legumes including lupin belong to the group of long-day plants, though there are some forms with neutral photoperiodic reaction (Дорошенко, Разумов, 1929). Our research (Kurlovich and Ivanova, 2000) has revealed positive but not identical reaction of the majority of the investigated accessions of narrow-leafed lupin to the lengthening of the photoperiod.
For example, the line of narrow-leafed lupin Mut-1 (k-2803) from Poland developed flowers and formed pods only in long daylight, i.e. it manifested a long-day photoperiodic reaction. The rest of accessions were early maturing in both photoperiodic modes. However, under short-day conditions the plants of these accessions were less in height.
Variation of characteristic features is observed in the accessions of lupin with different ecogeographic origin. The forms of lupins from northern latitudes and mountain areas responded to a short 10- and 12-hour day much stronger, than southern forms. The greatest changes in short-day environments were marked in wild forms, especially the ones from Portugal, Spain and Algeria. The samples from Greece, Turkey and Palestine were more neutral (Курлович, 1991б). Photoperiodic response of lupins was usually combined with the effect of other factors, especially with the response to vernalization (Kurlovich and Ivanova, 2000). Table 7 presents the influence of photoperiod duration and the complex effect of both factors (vernalization and photoperiod duration) on the growth and development of Mut-1 accession.
Mut-1 accession, which had shown long-day photoperiodic response, under the influence of short-day conditions and after vernalization showed determinate branching not on the main stem, but on its lateral branches. Ladny accession appeared to be the most thermo- and photo-neutral.

Photoperiodic response is also closely connected with spectral structure and intensity of the light. Long-day plants pass their reproductive phase faster with 24-hour illumination when in addition they are exposed to red light. Prevalence of long-wave rays in the light spectrum also provided for acceleration of plant development (Синская, 1946; Кашманов, 1970).
Besides, lupin plants respond to the intensity of illumination. Byszewski (1959) studied the effect of different light intensity of 7200, 10800 and 14400 lx on plants. With increased intensity of light, the plants grew faster, developed a more advanced root system, and produced higher yield at the increased intensity of light. Alkaloid content in seed was thus reduced. High light intensity produced an even greater effect on plants than lengthening of the daylight. Practically all lupin species are characterized by strongly expressed heliotropism, and turn their leaflets to the sun. Leaf dormancy is observed in many forms in the night when leaves lose turgor and droop. Lupin most effectively uses active photosynthetic radiation in comparison with other crops. Coefficient of its utilization in lupin is 4.79%, whereas in rye it is 2.42%, in wheat 2.68%, in oats 2.74%, and in vetch 1.98% (Шарапов, 1935). It is connected with high photosynthetic activity of plants and high calorific capacity of protein and oil contained in plants.
Response to heat. Multifoliate or Washington lupin (L.polyphyllus Lindl.) is the most resistant to cold among cultivated species. In Russia, the northern border of its cultivation for seed production passes along the line from Petrozavodsk to Syktyvkar. Narrow-leafed lupin is the least demanding of temperature among annual species. Its early forms ripen even in the northwest of Russia. The most heat loving is white lupin. Yellow lupin occupies an intermediate position. The data of Dyubin (Дюбин, 1981) describing heat requirements of different lupin species are presented in Tab. 8. In Russia, the sum of average daily temperatures ensuring maturity of narrow-leafed lupin is 1650-1690°C; for yellow lupin this figure is 1810-1950°C; and for white lupin it is about 2140°C. The sum of average daily temperatures for multifoliate lupin (L. polyphyllus Lindl.) for the period from the beginning of spring growth to the ripeness of seed makes 1325°C. The northern borders of growing areas for the considered lupin species for different maturity ensuring levels are also shown in Tab. 8. The vegetation period of cultivated lupins is constantly reduced, and accordingly aggregate heat requirement is decreased in the process of breeding more and more early-ripening cultivars and enhancement of their cultivation techniques. Already available are early thermo- and photo-neutral cultivars with determinate branching, which would make it possible to expand the area of lupin cultivation ever farther northwards.
For a majority of lupin species, minimum temperature for seed germination is low (+1… +2°C). Optimum temperature is +4 … +6°C. According to Sawichev (Саввичев, 1961), low air temperature at the time of shooting brings about reduction of the period from young plantlets to flowering. In the phase of stem growth and branching, lupin prefers moderately warm temperature. Flowering goes most intensively and effectively at moderate air temperatures (+15 … +25°C). Pollen sterility increases at higher temperatures and with absence of moisture. Seed ripening time is considerably reduced at higher temperatures, but is essentially longer with low temperatures and high humidity, especially in white lupin. In the north, tolerance of annual lupins to low temperature plays a significant role in their progress. According to Sharapov (Шарапов, 1935), annual species can tolerate the following temperature: L. albus down to –4°C, L. luteus down to –6°C, and L. angustifolius to –7°C. Barbacki (1960) states that lupin species can endure even more severe frosts: down to –6°C for white lupin, down to –8°C for yellow lupin, and down to –9°C for narrow-leafed lupin. These data testify that the researcher dealt with newer cultivars that had not only shorter vegetation period, but also, as a rule, increased tolerance to low temperatures. Resistance to low temperatures also depends on ecogeographic features of accessions and on the phases of plant development.
Our investigations (Курлович and Гаджиев, 1989) showed that in fall planting a winter form of white lupin of the Georgian ecotype endured the temperatures of –10°C down to –15°C during the whole winter period in Sheki-Zakatalsk zone of Azerbaijan. Lupin plants are more resistant to frost on the initial stages of development both in spring and fall plantings. Early frosts in autumn exert stronger influence on yellow lupin at the moment of seed ripening in spring planting. As to narrow-leafed and white lupins, they are able to go on growing even after long frosts. Perennial lupin species, in general, tolerate frosts quite well owing to their biological features, though some variance between species and variety also exists.
In most of the lupin species there are spring, intermediate and winter forms. Majority of wild forms belong to the winter type. In natural environments, seed of wild forms use to fall down on the ground from dehiscent pods. In autumn, with the beginning of a rainy season, these seed germinate, and young shoots remain for the whole winter in the state of rosette. During the Mediterranean winter with average temperatures between –5°C and +5°C, the plants stop growing and pass a stage of vernalization. They renew the process of growth in spring when air temperature becomes higher. Such cycle of development had been evolving for a long time in wild forms, and helped them to exist without any intervention from outside. When the plants are sown in spring without preliminary vernalization, majority of them would remain in the phase of rosette during the whole summer, and only with coming of colder weather in the fall some of them may start growing. Their cultivation as a spring crop is possible only after artificial vernalization within 30-40 days at a temperature of 0 … +1°C (Курлович, 1991б). It is necessary, however, to take into account that among wild forms there are plants with different demand for duration and intensity of vernalization. There are also typically summer forms with thermo-neutral habit used by breeders in Russia, Poland and other northern countries in spring plantings. Domesticated forms also manifest different responses to vernalization, which is testified by our experience in the study of genetic and environmental effects on branching in narrow-leafed lupin (L. angustifolius L.). The greatest effect of vernalization (Tab.9) was observed in accession Lanedeks-1.
Accession Mut-1 was transformed into the form with usual non-determinate branching under the influence of vernalization. Besides, it displayed determinate branching not on the main stem, but on lateral branches in these conditions.
But in the majority of the countries with warmer climate (Australia, southern Portugal, southern France, USA, and Chile) lupin is sown in the fall. For these purposes, winter and intermediate forms are used (Cowling, 1994; Erik von Baer, 1994; Huyghe et al., 1994; Nelson, 1994). Their cultivation with autumn sowing, wherever it is possible, provides for more efficient utilization of agroclimatic resources, higher productivity of lupin, and lesser degree of disease and pest incidence.
Response to moisture. The amount of free and bound water required for normal life of lupin reaches 80-85%. Lupin is xeromesophyte by nature, with sufficiently high resistance to drought and moisture. Its different species grow in different ecological conditions. However, if domesticated species are concerned, they are moisture-loving plants, suitable for cultivation in the areas supplied with moisture. Their transpiration rate is 600-700. Rather high moisture requirement of lupin is explained, first of all, by the fact that these plants accumulate much green matter. Secondly, plenty of moisture (120% of seed weight) is needed for the swelling of seed at germination. It is twice more than in cereal crops. On the other hand, lupin often endures drought quite well due to the well-developed tap root system, which supplies a plant with moisture and nutrients from deep underground. The first critical period in relation to the absence of moisture happens in lupin at the time of seed germination when the root system of plants does not work yet. The second period comes at the time of budding before flowering and fructification. The lack of moisture in this period results in considerable underdevelopment of pods and reduction of productivity. In the seed-ripening phase, higher humidity noticeably lengthens the vegetation period. The native land of domesticated annual lupin species is the Mediterranean region where they grow in mountainous areas. Rather damp and cool weather prevails there during the period of plant growth, and drought comes in the seed-ripening period. In the process of evolution lupin has adapted to such conditions. In Russia, on the contrary, in the time of seed maturing it often rains, so the period of ripening grows considerably longer, and in northern areas in some years seed do not ripen at all. Therefore, the plants are treated by defoliants and desiccants before grain harvest.
Laboratory evaluation of lupin accessions with various geographic origin for drought resistance based on the principle of seed germination in sucrose solutions has revealed (Курлович and Чернышева, 1986) the existence of interspecific and intraspecific variability in this parameter. Among the lupin species cultivated in Russia, higher relative resistance was found in the accessions of yellow lupin. The accessions of white and narrow-leafed lupins were characterized, on the whole, by lower resistance to drought, though among them there were some forms with high resistance. Yellow lupin shows rather low requirements to humidity in the soil and even in the air, as its native Mediterranean region it grows in dry mountainous areas. On the other hand, domesticated yellow lupin is grown basically on sandy soils than warm up quickly and are incapable to preserve moisture for a long time. In view of this, frequent spring droughts in the areas of its cultivation produce harmful effect on its seed and plantlets. This fact has created the problem of drought resistance for lupin, especially at early stages of its development, as seed of this crop require significant amount of moisture for the swelling. By now, too little effort has been made to solve this problem. Optimum soil humidity at a level of 60-80 % from full moisture capacity plays an essential role in formation of nitrogen-fixing nodules, promotes accumulation of higher protein content, and decreases the content of alkaloids (Byczwski, 1959; Barbacki, 1960). Especially high requirements to soil and air humidity are found in the cultivars with fast initial growth rate. The forms of lupins from the eastern Mediterranean region are, as a rule, more resistant to drought than the forms from the western Mediterranean. Eastern Mediterranean forms have leaflets with more xeromorphic structure (less stomas, presence of pubescence, etc.).
Excessive humidity of soil results in oppression of the root system and nodule bacteria that also lead to a decrease in the productivity of plants.
Responses to soil types and fertilizers. Lupin has aroused special interest because it is able to grow on poor light-textured sandy soils where the conditions are economically unfavorable for cultivation of other crops. Owing to its well-developed root system and high nitrogen-fixing ability, it is also widely used for reclamation of poor sandy soils. It is specifically so with yellow lupin, which shows in Russian conditions the least demand for soils. Most of all light loamy soils and sandstone may meet its requirements. Narrow-leafed lupin is a little more demanding of soil types. It prefers grounds with more coherent texture. However, narrow-leafed lupin like yellow one badly endures heavy clay soils, and suffers from superfluous humidity and high level of ground waters. White lupin is different in having the greatest requirement to soils among the domesticated species. Most favorable for it appear coherent loamy soils or sandstone, clay and even black prairie. Major criterion in the assessment of soil suitability for lupin cultivation is the reaction of soil environment. Lupins, even wild ones, grow only on acid and neutral soils where parent breeds or lava are on the ground surface. Almost all domesticated species of lupin also prefer subacid soils. Yellow lupin is considered as the most attached to subacid soils. There are very few species (L. casentinii, L. atlanticus and L. digitatus) adapted to fine-textured and alkaline soils (Buirchell, 1994). As to other domesticated species, they show high stability to soil acidity, prefer subacid or neutral reaction of soil environment, and poorly endure alkaline reaction. It is considered that рН 5.0-6.0 is the optimum of soil acidity for lupin (Майсурян and Атабекова, 1974). Thus, the response of lupin to soil reaction varies with the age of plants. In the early periods of development, optimum soil reaction is рН 4.6-5.0, while after flowering it is neutral, рН 6.0-7.0. The problem of lupin’s optimal reaction to soil environment is linked with sensitivity of lupin to the content of calcium in soil. It was generally accepted that lupin negatively responded to increased calcium content. However, it has also been established that its deficiency produces negative effect on the productivity of plants. Nonetheless, the limit of favorable effect produced by calcium on this crop is lower than in other plants. Negative effect of large amounts of lime results in significant accumulation of ammoniac nitrogen in plants, greater combustion of carbohydrates, and intoxication of plants with ammoniac (Майсурян and Атабекова, 1974).
Especially negative is lupin’s reaction to lime when the soil lacks for magnesium and other trace elements. However, profound and all-round study of the role of calcium conducted in the past years disproved the common opinion concerning incompatibility between lupin cultivation and calcification. It was observed that the negative effect of calcium is removed by applying magnesium into the ground, and iron on the same soils (Шутов, 1982). Therefore, in the fields where lupin is included in crop rotation, it has been recommended to calcify soil with flour containing magnesium that would neutralize the negative effect of lime (Курлович, 1985). Besides, it is undesirable to plant lupin on the sites that have recently been calcified. During a number of years after calcification, it is better to grow other crops prior to lupin. Different sensibility of lupin to the content of calcium in soil is also connected with the activity of nodule bacteria whose development goes well only with neutral and weak alkaline reaction of soil environment. They develop poorly with acid reaction of environment, and the process of nitrogen fixation proceeds inadequately. It is necessary to take into account the specified inconsistent factors of lime effect when working out an optimum technology of lupin cultivation as well as in breeding practice. The study of the genetic diversity of lupins preserved in the collection of VIR has revealed interspecific and intraspecific variation in the response of different forms to the content of calcium in soil. It opens prospects for breeding cultivars with good reaction to an excessive content of this element in soil. One of the valuable features of lupin as a leguminous crop is its ability to make symbiosis with nodule bacteria responsible for fixing free nitrogen from the atmosphere. Therefore, retention of nutrient elements is closely connected with the ability to provide biological nitrogen fixation by Bradyrhizobium sp. (Lupinus). These bacteria are especially sensitive to a surplus of mineral nitrogen. But its large amounts lead to abrupt reduction of nitrogen-fixing ability. Biological nitrogen fixation processes reach their greatest efficiency at the phase of plant flowering. To enhance this process, special preparations containing the most effective race of bacteria are applied. On the other hand, lupin is also capable to grow and develop at the expense of mineral nitrogen, especially if lupin is planted for the first time on a given site, and bacteria are absent in the soil. In this case, application of nitric fertilizers provides an essential increase in the yield. More efficient, however, is treatment of seed by an effective strain of nodule bacteria, or its application into the soil (Kurlovich et al., 1995, 1996). The process of biological nitrogen fixation begins only after the emergence of leaves. In view of this, initial dozes of mineral nitrogen are also effective at the first stages of plant development. The intensity of biological nitrogen fixation depends on the specific and varietal features of lupin, conformity of the applied strain of bacteria to a definite lupin cultivar, and also the conditions of supplying plants with other nutrient elements. Phosphoric and potassium fertilizers also render strong influence on the development of lupin plants, content of nitrogen, and formation of nodules. The lack of these fertilizers would greatly reduce general plant productivity. Phosphoric fertilizers stimulate growth of the root system; in particular its absorbing root hairs through which bacteria penetrate from soil, and enhance nitrogen-fixing ability by reducing the unfavorable effect of increased amounts of mineral nitrogen (Макашева, 1979). Nodule bacteria capable of transforming nearly insoluble forms of phosphorus into more absorbable forms provide for enrichment of plants with not only nitrogen, but also phosphorus. Great effect is produced by potassium, which also helps to increase assimilation of phosphorus. Phosphorus, in its turn, does not exert essential influence on the exchange of potassium. Plants absorb almost all potassium from soil in the flowering phase. The lack of potassium also causes a decrease in nitrogen fixation and impedes the growing processes. Low content of this element severely hampers the movement of nitrogen substances and carbohydrates from leaves to seed. Application of phosphoric and potassium fertilizes provides for an increase in the chlorophyll content in leaves and intensified photosynthesis and transpiration. As a result, general plant productivity becomes higher.
Not only simple presence of nutrients is important, but also their ratio which varies depending on various conditions. Significant prevalence of phosphorus against potassium in soil disrupts the exchange of substances and the transition of plants into the stage of reproductive development (Гукова, 1962). Best conditions for high-grade development of lupin are created by an increased supply of potassium and rather low level of phosphoric nutrients. In this respect, application of phosphoric and potassium fertilizers in the required proportions is very important for obtaining a high yield of lupin. It is necessary to take into account that lupin consumes twice more potassium, than phosphorus (Курлович, 1985). Besides, it responds well to magnesium (20 kg/ha), and also to the treatment of seed by micro fertilizers containing iron, molybdenum, boron, cobalt, zinc and manganese. Magnesium contains in the structure of chlorophyll. Its deficiency may lead to premature falling of leaves, breach in nitrogen supplies, and reduction of the efficiency of phosphoric and potassium fertilizers and lime (Магницкий, 1967) Iron also plays an important role during photosynthesis and nitrogen fixation. However, when it is present in excessive amounts, decomposition of chloroplasts is observed (Рубин and Германова-Гавриленко, 1956; Трепачев, 1967).
Molybdenum raises efficiency of symbiosis with nodule bacteria. It is accumulated in roots and nodules. This element is especially effective in neutralizing the negative effect of excessive calcium content in a plant.
Boron also enhances growth of the root system in all legumes. Cobalt makes part of the structure of vitamin B12, stimulating the formation of chlorophyll in plants. Microelements have an essential effect, specifically on the development of generative organs in lupin. Responsiveness of plants to their application depends on the type of soil, dozes and ways of application of fertilizers, and also on specific and varietal features of plants.

Biology of development

Prof. Kuperman (Куперман, 1961, 1977) identified 12 stages of organogenesis in individual development of all angiospermous plants. These 12 stages of organogenesis have also been accepted for lupin (Наймарк, 1976). Description of these stages was given in the following works: Жуков, 1961; Наймарк, 1976; Наймарк and Таранухо, 1982; Пронин et al., 1961; Шалыганова, 1961. At stage I, when the plants are in the phase of cotyledons, there is no differentiation in the apical point of plants. Stage II is characterized by formation of leaves and the rudimentary stalk at the basis of the apical point. At stage III, sheathing leaves are formed, the apical point is extended and its size increases. At stages IV and V, floral tubercles and bracts are developed, and also differentiation in the floral tubercle is observed on the bodies of flowers. At stages VI-IX, the processes of formation of sexual cells and fertilization take place. At stages Х-ХII, the process of sees formation and ripening takes place; then the plants grow older and die. Analysis of visible morphological changes in separate organs and plant habit in lupin development has shown that the process of plant development has a number of phenological phases.
As a result of the study of ontogenesis, the following phenological phases have been described for lupin (Наймарк, 1976): 1) sowing / sprouting; 2) rosette (8-10 leaves in lupin); 3) stem formation / branching; 4) budding; 5) flowering and fruit formation (on the central truss); 6) grain formation / grain plumpness (on the central truss); 7) green ripeness of grain; 8) whitish or waxen ripeness of grain; and 9) yellow, complete, firm or fully ripened stage of grain. Duration of the phases of growth and development depends on genetic features of species and cultivars, soil type, climate conditions, and the level of agriculture.
Sowing / sprouting. The duration of this phase varies in yellow lupin, on the average, from 10 to 16 days (Наймарк, 1976). Such variation depends on the level of daily average temperatures in soil, presence of moisture, and depth of sowed of seeds. Differences between varieties have had no effect on the duration of this phase under identical conditions of cultivation. In this phase, the growing point is on stages I-II of organogenesis.
The phase of rosette (stage II of organogenesis) comes after the complete emergence of young shoots, and lasts depending on the temperature regime, e.g. from 22 to 26 days with cv. Academichesky 1 (L. luteus L.), and from 22 to 31 days with cv. Bystrorastushchy 4 (also L. luteus L.). By the end of this phase, 8-10 leaves are formed, roots penetrate down to 30-35 cm deep, and nodules are developed.
The phase of stem formation / branching begins when plants are coming out of the rosette. This period lasts in different years from 9 to 14 days with cv. Academichesky 1 and 13-23 days with cv. Bystrorastushchy 4 (stages III-V of organogenesis).
The phase of budding is characterized by appearance of flower buds on the central truss and lasts until the beginning of flowering. With cv. Academichesky 1, its duration is 8-9 days, while with cv. Bystrorastushchy 4 it is 10-15 days (stages VI-VII-VIII of organogenesis).
The phase of flowering and fruit formation starts with the opening of the first flower and comes to an end with the development of pods on the central truss. With cv. Academichesky 1 it lasts 9-12 days, and with cv. Bystrorastushchy 4 it is 10-13 days (stage IX). This whole period is delayed in lupin when flowering takes place on lateral branches.
The phase of grain formation / grain plumpness on the central truss (stages X-XI) lasts in yellow lupin for 17-23 days with cv. Academichesky 1, and 16-22 days with cv. Bystrorastushchy 4. After complete plumpness of grain, three phases of ripeness take place, i.e. green, whitish (waxen), and yellow or firm (stage XII of organogenesis).
Green ripeness of grain. This phase lasts 14-18 days with cv. Academichesky 1 and 13-19 days with cv. Bystrorastushchy 4. By the end of this phase, grain cotyledons become firm, and the radicle of the embryo turns white. This is an evidence of physiological maturity of seed. If necessary, it is possible to apply plant defoliants during this phase. In this period, it is also possible to harvest seed with normal germination.
Whitish (waxen) ripeness of grain. This phase lasts 7-12 days. In this period the valves of pods turn dirty-brown, leaves of the main stem and lateral branches of the first order die off, the embryonic radicle in seed becomes yellow, and the seed is completely shaped. The inflow of nutritive substances in seed has not yet stopped, but is considerably slowed down. Such plants are quite suitable for separate harvesting.
The phase of yellow or firm ripeness of grain lasts 5-9 days. In this period, the plants are completely dried up, the pods become brown, and the cotyledons turn yellow (Наймарк, 1976). Essential influence on the duration of certain phases of development and the period of vegetation on the whole is rendered by agro-meteorological conditions, mainly by daily average air temperatures in combination with soil humidity level. Besides, the duration of the vegetation period is determined by a plant genotype.
In Russia and adjacent countries lupin is cultivated basically as a spring crop. Early maturity is an extremely valuable trait that makes it possible to expand the area of cultivation of this crop. As shown by the results of the long-term study of VIR’s lupin collection, the duration of the growth period for summer forms of narrow-leafed lupin sown in spring varies depending on the cultivar, year of planting and place of cultivation with an amplitude from 72 to 170 days. With yellow lupin, this character varies from 90 to 175 days, and with white lupin, from 106 to 180 days. Besides, duration of separate intervals is also signifying. For instance, in yellow lupin the period from young growth to flowering is the longest; in white lupin, on the contrary, it is short, but the period from flowering to maturity is the most drawn out. Great variability of the vegetation period is also observed in different years of cultivation. Narrow-leafed lupin begins to blossom simultaneously with white lupin, and a little earlier than the yellow one. It ripens much earlier then white and yellow forms. In Kiev Province, in damp years yellow lupin has blossomed on the 63rd day after the emergence of young shoots, white lupin on the 45th day, and narrow-leafed lupin on the 62nd day, whereas the duration of the flowering/maturity period in these three forms is respectively 54, 97 and 52 days. Processing of mathematical data of these observations has shown that there is a close positive correlation between the period from young growth to flowering and the whole period of vegetation. Coefficient of correlation (r) is +0.50.
In the environments of Russia, white lupin is the most thermophilous among all cultivated annual species. Its period of vegetation is longer than those of two other species. The study of white lupin collection was conducted in Kiev Province (Ukraine) where accessions were sown in spring sowing and at the former Sukhumi Station of VIR (Abkhazia) where they were planted in autumn. In the conditions of Kiev Province, variability of the vegetation period of early forms was studied, because late accessions do not ripen there. The period of their vegetation varied from 106 to 180 days. Majority of the accessions available in the collection are late-ripening, so they were studied near Sukhumi in autumn plantings. Their vegetation period there showed variation within the range of 230-260 days. Sown in autumn in damp subtropics near Sukhumi, white lupin produced high yield of seeds and green matter. In these environments, the vegetation period also displayed strong variability under different weather conditions (Tab.11).

In the damp subtropics of Abkhazia, the most valuable winter and intermediate forms of white lupin showed an increase in the yield of seed and green matter for green manure. Among them, the most promising are the accessions from France (k-1547), Argentina (k-1582 and 1583), Australia (k-1779 and 1791), and the forms of Georgian ecotype (k.k-1423, 2910, 3292).

ANATOMIC STRUCTURE


ANATOMIC STRUCTURE

M.V. Petrova

Anatomic structure of different lupin species was studied by a number of researchers. Some scraps of information can be found in the monographs of Metcalfe and Chalk (1950), and Maissurjan and Atabiekova (1974). Other authors studied the structure of plantlets (Compton, 1912), roots (Тен, 1973; Михайловская, 1960, 1968), stems (Ball, 1949; O’Neil, 1961; Tomaszewska, 1966), and petioles (Petit, 1887; Борисова, 1975). Petrova (Петрова, 1983) analyzed anatomic structure of leaves in one perennial and eight annual species of lupin from various centers of origin. The results of the study of physiological and anatomical features of leaves of L. аlbus and L. luteus are presented in the work of Latykova and Borisova (Латыковa and Борисовa, 1989). Watari (1934) described the path of conducting bunches in petioles of two species. The structure of nodes was investigated in annual and perennial species (Гуленкова, 1982), and the effect of cotyledon and leaf traces on the development of conducting systems in plantlets was described (Денисова and Дмитриева, 1980). Some of the authors wrote about the structure of vegetative and generative organs of lupin when studying the variability of characters in this genus (Майсурян and Атабекова, 1974; Карпицкая et al., 1980). Great attention was focused on the structure of pods (Атабекова, 1958; Карпицкая and Таранухо, 1976; Белов and Лотова, 1991; Tomaszewska, 1954) and seed coat (Михайлова, 1958; Циклаури, 1962; Карпицкая and Таранухо, 1974; Редькина, 1976, 1979; Лотова, 1991; Zimmermann, 1936a, b; Heyn and Herrnstadt, 1977).

Root. The root of lupin in its primary structure consists of the epidermis, primary cortex and central cylinder formed by the pericycle, xylem, phloem and parenchyma. The root is covered from outside by the epidermis forming root hairs, which are short-lived and quickly die off. The greatest area is occupied by the parenchyma of the cortex, consisting of thin-walled live cells different in shape and size. The internal layer of the primary cortex is the epidermis which differs from all other cells of the cortex by its shape and by its small and tightly serried cells. Not only one-layered endoderm was found in the genus Lupinus L., but also two-layered (L. angustifolius), three-layered (L. albus) and tetra-layered endoderm (L. luteus) (Тен, 1973). The pericycle is the outside layer of the central cylinder. It consists of one-layered densely located cells with thin parenchymal cell walls. The number of the rows of cells in this tissue is increased in front of the rays of the xylem, and there are lateral roots.
The root system of annual lupin species has a diarchic structure. The arrangement of lateral radicles in lupin has two-rowed layout. The first root vessels are annulate and spiral. The larger vessels of a porous type are differentiated closer to the center and in the central part of a root. There are the groups of phloem between the rays of the xylem. Transition to the secondary structure begins when the cambium is formed between protoxylem and protophloem as a result of division of the procambium. It forms the outside elements of secondary phloem and elements of secondary xylem inside. Cambium forms a wide continuous ring in the root of secondary structure. The root of lupin has a xylem type of structure, i.e. it has more xylem in comparison with phloem. The secondary xylem of the root consists of vessels, libriform fiber, parenchyma and radial rays. The vessels are basically porous in the secondary xylem. They are surrounded with fibers of libriform shape and cells of ligneous parenchyma. The radial rays are primary and secondary. The number of primary rays corresponds to the number of rays of the primary xylem. Both primary radial rays proceed from the primary xylem located in the center of the root. They connect the central part of the root with the cortex. They are wide and consist of several rows of parenchymal cells of various shapes. The secondary rays are shorter and narrower. The continuous ring of phloem is situated outside of the cambium submitted by bast parenchyma, bast fibers, sieve tubes and cells - satellites. Intensive activity of the cambium causes growth of the central cylinder; therefore, there is a flattening of the cow part of the root and abscission of the primary cortex.
Ten (Тен, 1973) has noticed that abscission of the primary cortex is partial in lupin; it is located in a zone positioned against the rays of the primary xylem. The root is covered by fuse and by secondary cortex lies. It is formed by cellular parenchyma, with groups of rather thick-walled and slightly lignescent bast fibers between its cells. The root of long-term miltifoliate lupin (L.polyphyllus) differs from the root of annual species by the presence of lignifying primary xylem. There are five primary radial rays in the root of this species. Secondary xylem consists of numerous layers of vessels formed with regular intervals. These layers are separated from one another by profuse homogenous tissue, often thin-walled and poorly lignescent. Considerably advanced ligneous parenchyma surrounds the vessels. The main root contains plenty live tissues of parenchyma in the secondary structure. They consist of ligneous and bast parenchyma and also of the cells of numerous radial rays of the root. A large number of long radial rays are formed in the root of perennial lupin during the first year of its life, and also short radial rays start their development. They become significantly longer only in the next year. The primary radial rays have greater length and width. In the second year and next years of lupin’s life, new short radial rays are set again. A series of radial rays are well discernible on a cross-section of an aged root, especially of its ligneous part. They are formed in different years of the life of the root. The rays of each year are always set at one level. This regularity in the formation of radial rays gets broken with the aging of a plant. It is connected with reduction of the cambium activity. The bast part of the root is solidly developed, especially in the roots of a young plant. Lignescent mechanical tissues are always present in the bast part, in addition to lignifying tubes and abundant bast parenchima.
Mechanical tissues are set down in the periphery of the bast fiber in tangential layers interrupted by radial rays of the root. A certain periodicity is observed in the formation of fibrous mechanical tissues in the bast part of the root of an adult plant. It is considered that year-by-year growth of lupin roots proceeds with more difficulty as compared with other plants. It is necessary to take into account a number of anatomic characters while determining the age of a root. The study of the annual increment of bast tissue and the arrangement of narrow radial rays can help get an answer to this question (Михайловская, 1960, 1968).
There is an early developed cambium between phloem and xylem. Therefore, the primary structure of the root is very quickly replaced by the secondary in young plantlets. The cambium is active in the main root during its whole life. In view of this, the zone of cambium is large in adult plants. The activity of cambium decreases with aging of the plant and year-by-year gained layers become narrower. The primary cortex in perennial species of lupin dies off and falls in the first year of plant life. Only the secondary cortex remains surrounded by a layer of a cork.

Stem. The stem of lupin has a dissimilar structure throughout all its length. In the top part of the stem, the conducting system consists of separate bunches divided by parenchyma. In the bottom part of the stem, the conducting system represents a continuous ring. The annual species of lupin differ from each other by the shape of the cross-section of the stem and by its size. In the basis, the stem of lupin is round, but at some species, such as L. albus and L. luteus, the sides are appreciable. The smallest size of the cross-section of the stem is found in L. truncatus and L. nanus, and the greatest in L. luteus and L. hartwegii.
Considerable similarity is observed in the stem structure of the investigated annual lupins. They epidermis is single-row and is covered by thin wavy cuticle. Characteristic of the epidermis are stomas and simple acuminate hairs. There are 1-2 rows of collenchyma set under the epidermis. Behind them there is cortical parenchyma formed by 5-9 rows of roundish thin-walled cells. The first 1-3 rows of these cells have chloroplasts. On the sides of the stem there is a reinforced rim of collenchymal cells. Their presence makes the stems of lupin cultivars stronger and provides them with resistance to lodging. Fragility of the stem, characteristic of some species, for example L. subcarnosus and other species of the western hemisphere, is preconditioned by the lack of this mechanical tissue replaced in them by friable parenchyma (Майсурян and Атабекова, 1974).
The cordial parenchyma comes to an end by the axis. Rather powerful rays of sclerenchyma are located behind it. The conducting system represents a continuous ring. Some species are characterized by very powerful xylem (L. angustifolius, L. mutabilis). The cambium of 4-8 cell rows covers the xylem. A continuous ring of phloem is set at the external part of the cambium, above which there are rather powerful rays of sclerenchymal fibers. The medulla formed of rather large cells of parenchyma surrounds the internal cavity of the stem, which has different size in different species. The largest cavity is observed in L. cosentinii (up to 40% of the whole cross-section) and L. truncatus (53.3%), whereas the smallest is found in L. atlanticus and L. hartwegii (5.7 and 7.7 %). The cavity was not present in all studied accessions of L.angustifolius. The crystals are absent in all species of lupin. Thus, all examined annual lupin species differ from each other by the shape of the stem’s cross-section, outlines of the conducting cylinder, sizes of cells in different tissues, presence of collenchymal rims in the cortex, and percentage of various tissues. The stem of perennial L. polyphyllus has a round cross-section, sometimes with a small protuberance. One layer of collenchyma is located under the epidermis with cuticle and hairs. Farther behind it, a cortical parenchyma of 5-8 roundish thin-walled cells follows. The first one or two rows of these cells contain chloroplasts. The conducting system in this species represents a continuous ring, but the rays are well distinguishable. Sclerenchymal layer of 3-8 rows of cells are located above the phloem. Xylem and phloem occupy smaller area (about 10%) of the cross section in this perennial species as compared with the annual species, where xylem and phloem occupy from 16 up to 44%. The cambium consists from 4-5 layers in L. polyphyllus. The central part of the stem (up to 66-70%) is occupied by a large cavity.
Maissurjan and Atabekova (1974) have noticed that the species of lupin from the eastern hemisphere are more large-celled than the species from the western hemisphere, where all plant components are smaller-celled. The largest cells are observed in the stems of the Mediterranean species, such as L. pilosus, L. albus, L. angustifolius and L. luteus. Of all species from the western hemisphere, a large-celled stem is reported only in L. mutabilis. Besides, all species from the western hemisphere differ from those from the eastern one by the presence of suberification in the bottom part of the stem. The nodes of the stem are ternary-lacunar and ternary-fascicular in all annual species of lupin. They are single-fascicular only in the area of the inflorescence. The node of the cotyledon is also single-lacunar or single-fascicular. Three rays go into one lacuna, of which two extreme rays are large and the middle ray is smaller in size, and no one of them has bast fibers. The last rays are often advanced poorly.
The nodes of the cotyledons in perennial multifoliate lupin (L.рolyphyllus) are similar to those of annual species. However, nodes of vegetative burgeons are penta- lacunar and penta-fascicular. Besides, the numbers of rays and lacunae vary for different nodes (Гуленкова, 1981). Quite often it is possible to find nodes with six, seven or eight lacunae. One ray enters into each of the lacunas. The same author has reported that in the nodes of vegetative burgeons the minimal number of rays on the leaf trace is three. But usually the nodes have 5- or 7-fascicular rays. The number of rays, as a rule, corresponds to the number of lacunae.

Leaf. The pubescence of lupin leaves is formed by simple, acuminate, three-celled hairs of various length. Pubescence is present on the bottom part of a leaf in all studied species, but it was mainly on the top in three species: L. atlanticus, L. luteus and L. nanus. In other species, individual hairs occur, or they are in general absent on the top epidermis of the leaf. The greatest number of hairs on the bottom surface of a leaf is presented at the Mediterranean species and at two American species (L.ornatus and L.mutabilis). The hairs are longer on the top of epidermis than on bottom.
Long hairs are observed at the American species L.hartwegii and L.elegans and shortest at L.mutabilis, L.angustifolius and L.сosentinii (Tab. 12 in the book!). The least number of hairs is at L.polyphyllus. Sometimes they are absent and from time to time available individual hairs on top epidermis at this species. Some variation is marked in a degree of sinuosity of walls of epidermal cells (Fig. 11). The walls of cells are twisting on both surfaces of a leaf. Exception is L.albus, L.atlanticus, L.cosentinii and L.mutabilis, at which the majority of cells are rectilinear-roundish on the top party of a leaf. The cells of L.hartwegii are weakly twisting. By the most twisting outlines of cells differ L.truncatus and L.elegans. Distinctive feature of perennial lupins is the large number of fine cells and also largely wavy and zigzag walls of cells. At all species of lupin, especial at L.albus and L.polyphyllus, the coats of epidermal cells have thickenings. For epidermis of the leaf are characteristic usually anomocytic type of the stomas. They are settling down on both leaf surfaces. However, the stomas at perennial multifoliate lupin are available only on the bottom surface of a leaf. The smallest number of stomas is marked at L.angustifolius and greatest is at L.polyphyllus, L.atlanticus, L.cosentinii, L.albus and L.mutabilis on bottom epidermis. The number of stomas at L.luteus and L.angustifolius on the top surface of a leaf approximately is twice more, than on bottom; at L.elegans and L.hartwegii, on the contrary, the quantity of stomas on bottom epidermis is more than on the top.
The number of stomas at other species is approximately identical on both parties of a leaf. In greatest length of stomas differ L.angustifolius, L.elegans, L.hartwegii, L.ornatus and L.luteus. The thickness of the leaf plates varies from 179 up to 528 microns at all investigated species. The leafs are dorsoventral at L.luteus, L.albus, L.atlanticus, L.cosentinii, L.ornatus and L.elegans (Fig. 12). The number of lines of palisade tissue changed from 2 up to 3, and porous - from 3 up to 5. The cells of palisade tissue at L.luteus are rather wide, high and more serried in comparison with other species. The leaf is isolateral at others annual species. The most xeromorfous structure of leafs has only L.angustifolius among the investigated Mediterranean species. It has the highest palisade’s factor (81,8-83,7%). The palisade tissue is submitted by two - three lines of cells from the top party and by two - four lines with bottom. The spongy tissue is friable. The highest palisade’s coefficient between American species is marked at L.hartwegii (86,5%) and L.mutabilis (81,4%). The palisade tissue consists from both parties of a leaf at them from two-three lines of short and wide cells with large intercellular spaces. Three - four lines of short wide cells of palisade tissue from the top party of a leaf and two - three lines of narrower cells with large intercellular spaces with bottom are characteristic for L.nanus. The palisade tissue at L.truncatus is submitted from the top party of a leaf by two - three lines of high, basically wide and dense serried cells. This species has only one line of cells from the bottom party, which remind spongy tissue with very large intercellular spaces. The porous parenchyma is very friable and consists from four-five twisting cells. The palisade tissue at L.mutabilis consists from two-three lines of rather dense cells from the top party of a leaf, and from two shorter and more friable cells located with bottom. The palisade cells at L.micranthus are not dense from the bottom party of a leaf and have intercellular spaces. The leaf of perennial multifoliate lupin is very thin and has most mesomorphic structure in comparison with all investigated annual species. The palisade tissue consists from two lines of short cells. The second line has large intercellular spaces. The spongy tissue also consists from four-six lines. The lowest palisade’s factor is typical at this species. The middle vein is very similar at all species on anatomic structure. The orbed ledge is well excreted from the bottom party almost at all species . It is smoothed at L. angustifolius. There is one line of collenchimal cells in the ledge behind of epidermis. This line is swept poorly up at same species (L.angustifolius, L.truncatus). Some lines of orbed parenchymal cells are situated behind it. With the opposite parties from epidermal cells follow two-three lines of palisade cells with chloroplasts. The conducting system of the middle vein is formed by one vascular-fibrous beam located at the centre of vein. This beam consists from xylem, phloem and mechanical tissue. Last is submitted by collenchyma and sclerenchyma. The sclerenchyma is very weak and is present not at all species. Several cells at L.luteus, L.nanus and L.mutabilis submit it under phloem . It is frequently weakly lignescent at these species and it is absent absolutely at other species. Collenchyma is located from the party of xylem and limits phloem from two parties, sometimes incorporating under phloem by a thin line. The mechanical coat is absent completely at a conducting beam from the party of xylem at L.polyphyllus. The petiole of leafs differs on the size, form and anatomic structure at different species (Fig.13). They have the cylindrical form in cross section (at L.polyphyllus, L.hartwegii and L.mutabilis), ovate-triangular (at L.albus, L.truncatus, L.nanus and L.elegans) or weakly canaliculate (at L.angustifolius and L.luteus). Some variation is observed on the area of cross section, which depends from the sizes of leaf. The greatest area of cross section is observed at L.polyphyllus and least at L.angustifolius. Simple and acuminate hairs of various lengths cover the petiole at almost all species. All species of lupin, except for L.truncatus, have at leafs one line of collenchyma cells which follow behind epidermis (Fig. 14). Further medium-sized orbed cells with chloroplasts are located, which go on all circle of the petiole, obstructing only under a large beam. Number of their lines in this place is decreased. More deeply behind chlorophyllous tissue settles down some lines of large parenchymal thin-walled cells. Some differences are observed on number and form of chlorophyllous cells. So, at L.albus, L.elegans and L.polyphyllus this tissue consists from one, sometimes from two lines of orbed cells, and at L.luteus, L.nanus and L.mutabilis it consists from three-four lines. The part of chlorophyllous cells has the form of palisade cells (at L.angustifolius and L.truncatus). The chlorophyllous tissue of L.angustifolius consists from two or three lines of palisade cells, behind which follows one line of orbed cells. L.truncatus has one line of orbed cells at once for epidermis, and then two lines of palisade cells. Sometimes directly after epidermis settle down two-three lines of palisade cells. Large and fine beams form the conducting system of petiole. Some species have only three conducting beams (L.luteus and L.angustifolius), others have except for three large some more fine beams. There are many beams especially at perennial multifoliate lupin (8-10 large and 9-15 fine beams). The number of large beams reaches up to 5 at L.cosentinii and L.ornatus. It is many fine beams at L.truncatus (12-15). The beams are formed from xylem, phloem and sklerenchyma, which consist from two-four lines of cells. The xylem differs with size of vessels and degree of lignification of parenchymal cells. The xylem has almost not lignescent parenchyma at L.mutabilis and L.hartwegii . L.truncates and L.angustifolius on the contrary has strongly lignescent xylem. Large thin-walled parenchymal cells submit the central zone of the petiole. They partially break down forming a cavity. There is a large cavity at L.polyphyllus, L.truncatus, L.mutabilis, L.ornatus, L.micranthus and L.cosentinii at the centre of the petiole. But the cavity is absent at L.atlanticus and L.angustifolius. The anatomic study of a leafs at lupin has allowed to reveal the species with mesomorphfic and xeromorphic structure of this organs. The most mesomorphfic structure of leaf has perennial multifoliate lupin (L.polyphyllus). It has dorsovental type of structure, the large interrcellular cpace in mesophyll of leaf, very small palisade’s factor, large leaf plate and small thickness of leafs. Its stomas are only on the bottom party of a leaf. The presence of almost all listed attributes at L.albus, L.luteus, L.ornatus, L.cosentinii and L.elegans allows also relating these species to mesophits. The structure of their leafs shows, that all these species, in particular L.polyphyllus, differ by the increased insistence to a moisture and less resistant to a drought. Other species also have an anatomic structure of a leaf characteristic for mesophytes, though they bear also some attributes of xeromorphity, such as isolateral structure of a leaf, well advanced palisade tissue, fine leaf, high percent of lignescent parenchyma in the xylem of petiole, chlorophyllous parenchyma in the petiole as palisade tissue. The most xeromorphic characters in a structure of a leaf differs differ among annual species L.angustifolius. It has small leafs with thick plates, isopalisadity, high palisade’s factor, high percent of lignescent parenchyma in the xylem of petiole, numerous narrow vessels and chlorophyllous parenchyma in the petiole as palisade tissue. All species with xeromorphic characters have at the same time large intercellular spaces not only as spongy tissue but also as palisade tissue. It can specify that they are resistant to the air drought but require to the soil moisture. Formation of them occurred probably in conditions of sufficient safety of moisture and plant was exposed to strong action of isolation. That, in opinion of a number of the researchers, results in a thickening of leaf plate and greater development of palisade tissue. So, as a result of study of physiological-anatomic features of assimilation at two species of lupin is established, that the large thickness of plate of leaf, palisade mesophil and epidermis, and also greater number of stomas and chloroplasts characterizes L.albus as more photophilios species in comparison with L.luteus (Латыкова and Борисова, 1989). Perennial multifoliate lupin differs from annual by absence of stomas on top epidermis of a leaf, by thin plate of leaf, structure of mesophil, low factor of palisadity, type of conducting system and large number of conducting bunches in the petiole. The research of 13 species has allowed to reveal, that such characters as number and distribution of stomas and the sizes of epidermal cells on both surfaces of a leaf, character of outline of epidermal cells, presence, number and length of hears, structure of mesophil, form of cross section of petiole, number of conducting bunches and structure of chlorophyllous tissue in the petiole can play an essential role in the systematic of the genus Lupinus L. (Майсурян and Атабекова, 1974). But, the analysis of the anatomic characters of separate species not given an opportunity to carry out precise border between Mediterranean and American subgenera of Lupinus L.

Bean. The pubescence of beans can be rare or on the contrary rather strong depending from a species. The hairs at bean are usually simple unicellular; at one species they are shorter and at others are longer. The valves of beans consist from outside epidermis, the layer of parenchyma, pergameneous layer and internal epidermis. The cells of outside epidermis are dense serried, slightly extended, with club-shaped by outside walls covered by cuticle. 1-3 lines of thickly walled hypodermic cells are located under epidermal cells at some species (L.mutabilis, L. hilarianus, L.succujentus) (Майсурян and Атабекова, 1974; Белов and Лотова, 1991). The parenchyma is formed by several lines of orbed or ovate thin-walled cells, behind which is situated the pergameneous layer, consisting from prosenchymal dense serried cells with thick lignescent boundaries. They are advanced poorly at one species and their thickness is rather significant at others. The walls of cells are covered by lengthened-ovate pinholes. The cells of internal epidermis, which are behind pergameneous layer, early collapse or keep in very small quantity. The deep drills pass lengthways of abdominal seam and middle vein at many species. The middle vein of a bean has one or two conducting bunch usually divided by one - two layers of thin-walled parenchymal cells. There are also two conducting bunches on abdominal seam of a bean, between which also settle down the parenchymal cells. There are two bands of sclerenchymal cells outside of conducting bunches from the party of abdominal seam and middle vein. Between them are situated not only thin-walled parenchymal cells but also clean collenchymal bands of the various sizes. The sclerenchymal bands are available at some species not only outside of conducting bunches but also from the inside, as at beans of L.mutabilis (Майсурян and Атабекова, 1974).
The species of lupin differ from each other on anatomic structure of beans. There are species with easy shattering, non-shattering and weakly shattering beans both in the Mediterranean and in American groups. This ability is closely connected with an anatomic structure of a bean. The shattering of beans is provided by a number of the reasons: by presence of deep grooves lengthways of abdominal seam and middle nein, thick pergameneous layer, heterogeneity of tissues in area of abdominal seam and middle vein, formation of two layers of cells from the party of abdominal seam being as though continuation of cells of outside parenchyma, faltering sclerenchymal beams, absence of hypoderma under outside parenchyma etc. Such characters as absence or weak development of grooves lengthways of abdominal seam and middle nein, weaker development of pergameneous layer , presence of hypoderma from cells with club-shaped by cellulose bounady under outside epidermis, as at L.mutabilis, presence of continuous beam of sclerenchymal cells above conducting bunches, formation of cubepidermal collenchyma resist to shattering of pods. The direct bean is shattering usually more difficult than bent. The reasons ensuring durability of beans at various species of lupin are not identical (Белов and Лотова, 1991). These authors noticed that by the same set of characters can be characterized both easily shattering beans and difficultly shattering beans. These writers consider that it is impossible to judge about presence of the precisely expressed tendency to easy or complicated shattering pods on the basis of separate anatomic characters . More important is a degree of reduction of pergameneous layer, which can be connected with features of submicroscopic structure of its cells and membranes. The character of changes in other cells of pericarp tissue at its drying also influences on this character. Other authors consider, that the significant role in the mechanism of shattering is played an arrangement of pinholes in the cells of pergameneous layer (Майсурян and Атабекова, 1974).
The effective sources of non shattering pods at L. angustifolius were revealed by Gladstones (1967) at cv. New Zealand Вluе as natural mutants. This character is caused by two genes tardus () and lentus (). The gene limits shattering for the account of accretion of beans valves by formation of solid beam of sclerenchymal cells on all perimeter of a bean. The gene reduces shattering at the expense of structural changes in the valves. The layer of dense pigmented tissue as a grid form inside them, which interferes a curling of valves. The valves of such beans get brightly expressed reddish shade to beginning of seed maturing.

Seed. The outside layer of the seed coat is palisade epidermis covered by cuticle. The thickness of the cuticle may vary from one species to another. The palisade epidermis occupies the most part of the whole seed coat depth. It is formed by one row of cells having elongated shape.
The size of the palisade epidermis is not the same with different species and correlates with the size of seed (Карпицкая and Таранухо, 1974). Thus, it is the smallest in L. nanus and the largest in L. pilosus and L. albus. The cavities of cells in the palisade epidermis are very narrow; sometimes, in some species, they are slightly extended in the middle part. The shape of their cross-section is 5-6-edged. The cells of this tissue contains barked matter and various pigments. Seed color depends on the distribution of the species. There are species (L. aridus, L. elegans and others) where some cells of the palisade layer are missing, and funnel-shaped channels are formed (Карпицкая and Таранухо, 1974; Лотова, 1991). The cuticle sags above these channels and the seed surface becomes rough. The palisade cells are crossed by one or two light lines of another cells. Under the palisade layer there is hypodermis formed of one row of club-shaped cells with more or less regular intervals, cells which look like sand-glass or coils. They are joined with each other by their ends leaving large intercellular spaces. The hypodermis has unequal thickness in various parts of seed.
Maissurjan and Atabekova (1974) considered that the structure and size of hypodermic cells in the seed coat of different species are variable and may be regarded as a proof of their specific character. A thicker hypodermis and larger cells are observed in the seed of L. pilosus, L. digitatus, L. mutabilis, L. albococcineus and L. angustifolius (Карпицкая and Таранухо,1974). There are three layers of cells behind the hypodermic layer. The first is thick- walled parenchyma with cells filled by nutrient substance in unripe seed; the second is thin-walled parenchyma through which conductive bunches pass; the third layer is formed by strongly squeezed cells of thick-walled parenchyma. Maissurjan and Atabiekova (1974) reported that different species differ from one another in the structure of these layers, variable likeness, and unequal thickness. Another author (Лотова, 1991), on the contrary, stated that the variation of anatomic characters of the seed structure is very wide, and they have no diagnostic importance for the taxonomy of the genus Lupinus. Seed hardness characteristic of a majority of leguminous crops is found also in lupin species (Михайлова, 1958; Циклаури, 1964; Карпицкая and Таранухо, 1974; Редькина, 1976; Лотова, 1991). The thickness of palisade tissue in hard seed is greater than in normal swelling seed and makes about half of the whole thickness of seed coat. Such seed may be found in L. pilosus, L. digitatus and L. subcarnosus. In other species, palisade tissue occupies approximately one third of the thickness of seed coat (Карпицкая and Таранухо, 1974).
The cells of the palisade layer of hard seed have thicker membrane and narrower cavities. It was found out (Лотовa, 1991) that the epidermis and hypodermis are the tissues determining seed hardness. An increase in the total thickness of these two tissues, if compared with the depth of all spermoderm, plays an important role in increasing the hardness of seed. The cells of the hypodermis in hard seed are densely linked with each other. It is considered that the pigments contained in the cells of the palisade epidermis, hypodermis and parenchymal layer render an effect on the permeability of seed coat as well (Редькина, 1976). The hilum is represented on Figure 16. There are two layers of palisade cells outside of the hilum: the first internal palisade layer is the rest of seed stalk, and the second internal layer is the continuation of palisade tissue of seed coat. Both layers are broken up forming the hilum cleft which is pressed in the tracheal islet consisting of cells of various length and with numerous pinholes looking like tracheids. Mesh tracheids are typical for L. luteus and L. pollyphyllus, ladder-like tracheids are typical for L. albus and spiral ones for L. pollyphyllus and spiral ones for L. elegans (Редькина, 1979). Hygroscopic properties of seed are regulated with the help of the hilum cleft. Under the palisade epidermis, on both sides of the tracheal island there is a hypodermic layer consisted of cells of various shapes: fusiform, cylindrical, etc. Significant part of the hilum is occupied by asterinoid tissue which adjoins hypodermis and consists of thick-walled cells of various shapes with broad intercellular intervals. The asterinoid tissue in some species (L.angustifolius and L. pollyphyllus) directly adjoins the tracheal islet, while in other species it also has a two- or three-layered thin-walled parenchyma which surrounds the tracheal islet. In asterinoid tissue cells there can be pigments, sometimes specific to a certain species. Adjacent to both asterinoid tissue areas is porous tissue made of thin-walled cells of various shapes. This tissue has large intercellular spaces in L. angustifolius, and is strongly condensed in L. polyphyllus (Редькина, 1979). There is a layer of extended, slanting and strongly obliterated cells below the porous and asterinoid tissues. These are the remnants of a nutritive layer that separated cotyledons and seed coat.

BIOLOGY OF FLOWERING, EMBRYOLOGICAL AND CARYOLOGICAL PECULIARITIES


BIOLOGY OF FLOWERING, EMBRYOLOGICAL AND CARYOLOGICAL PECULIARITIES

E.M. Kazimierska, T. Kazimierski

Biology of flowering and pollinating

Lupin is an entomophilous plant attracting insects by multi-colored flowers, nutritious pollen and fragrance. The flower of lupin is monoecious, zygomorphous and hermaphroditic. It has concrescent stamens, superior ovary and apocarpous gynaeceum (Дорофеев et al., 1990). Among lupin species of the eastern hemisphere there are strictly self-pollinated species (L. angustifolius L.) and self-pollinated species with facultative cross-pollination. Cross-pollination prevails in lupins of the western hemisphere. The main stem and the lateral branches of lupin terminate in inflorescences of the apical truss type, whereas a majority of other representatives of fam. Leguminosae bear flowers on leaf axils. The arrangement of inflorescences and their sequence of development also have peculiarities with different species. For example, in many perennial lupins peduncles are direct continuation of the stem, which means that their inflorescences are originally apical. However, there are also perennial species where trusses or even individual flowers are formed on leaf axils. With the annual forms of lupin, inflorescences are also apical, but in addition to accessory buds there are branches which, after their growth and development, would end in lateral inflorescences, often considerably exceeding the apical inflorescences. Annual forms usually have much more inflorescences than perennial ones, but they normally comply with the latter in length and number of flowers. The length of the flower truss is a quite reliable specific character. In some species the length of axial inflorescences reached 40-50 cm (Майсурян and Атабекова, 1974).
The flowers set in a normally developed inflorescence are veticillate or semi-verticillate. However, there are trusses where the strictly verticillate arrangement of flowers in the lowest part of the inflorescence goes up in a spiral. Hence, the arrangement of flowers in an inflorescence can be verticillate and, at the same time, semi-verticillate in some lupin species. The flower corolla of lupin is zygomorphic and papillionaceous. It consists of a vexillum (flag), wings, a keel and an ovary with a pistil and 10 stamens (Fig.17 in the book). The development of the flower goes on in cycles. In early phases the calyx forms a compressed ring consisting of five sepals. The vexillum covers the wings and keel (keel). Initially the stamens are joined together, but later on one of the stamens becomes slightly isolated from the other nine. The flowers of lupin do not contain honey, but they attract insects with their bright color, presence of pollen and secretion of a smelling liquid from the vexillum and other parts. The vexillum densely covers other parts of the corolla before the flower opens. In the middle, between two halves of the vexillum, there is a small vallecula, making it almost keeled. The vexillum has a rounded or oval contour; it is straightened in the middle during flowering, and becomes prone to strong curving at the edges. Claw of the vexillum is dense and fleshy. The cells of the epidermis in the vexillum are elongated from the outside, with straight walls. Specific features are inherent in the epidermis structure of the inner side of the vexillum. In the middle of the vexillum there are small nipple-shaped outgrowths emanating odoriferous liquid, which attracts insects. Towards the top of the vexillum, its cells are appreciably shorter and outgrowths are narrower, so there are two types of nipple-shaped outgrowths on its inner surface. Various species of lupin differ on the structure of epidermis and the size and structure of nipple-shaped outgrowths. These outgrowths are smaller and less numerous in completely non-odoriferous or insufficiently odoriferous species (Майсурян and Атабекова, 1974).
The flowers of some species have a strong insect-attracting smell. The intensity of this odor depends on the size and number of nipple-shaped outgrowths on the epidermis of the vexillum and corolla wings. There are well-advanced and closely placed nipple-shaped outgrowths on the epidermis of the vexillum and on the wings of odoriferous flowers (L. luteus), while in odorless flowers (L. albus) small obtuse nipple-shaped outgrowths are only at the basis of the vexillum on its inner side. Towards the top of the vexillum in odorless flowers, these outgrowths gradually decrease in size and finally disappear. The wings of lupin flowers are concave-convex, having narrow, normal or wide oval shape. They are usually linked at the top edge and loose on both sides. The outer part of the wings is always convex and forms an arch. In the closed hollow of the wings the keel is located. The epidermis on the outer side of the wings consists of rectangular cells with wavy-surfaced walls. Two types of nipple-shaped outgrowths can be found here: the first type is smaller, wide and blunt, while the second is larger and pointed. The inner side of the wings is covered by smooth epidermis and by blunt and pointed outgrowths. The size of cells markedly decreases towards the top part of the wings. The flower keel is falcate or falcate-curved, less often it is more or less straight, narrowed toward the top. The uppermost part of the keel accretes into a firm beak. There is a tiny aperture for pollen and the pistil’s stigma in it. The base of the keel is always open. The edges of the keel in some lupin species are ciliated. The keel is transparent, colorless or just a little hued. The keel’s edge can be colorless, slighly hued, or intensively colored, almost black with anthocyanin. The color of the keel’s edge is a simple and convenient distinctive feature of different lupin forms and varieties. It is taken into account by breeders and taxonomists. The color of the keel depends on the presence of chloroplasts, anthocyanin and other pigments. Besides, in all parts of the flowers it is possible to find rather large crystals, insoluble in hot water, alcohol or nitric acid (Майсурян and Атабекова, 1974).
There are a pistil and 10 stamens in the keel. In lupin flowers, all the stamens are joined at the bottom in the staminate tube, while their upper ends are loose. They are arranged in two circles. The anthers of five stamens of the outer circle located against the sepal are more elongated and much larger then the inner ones. The anthers of five inner stamens situated closer to the petals of the corolla are reniform; they are developed later than the outer ones. Pollen grains in the anthers of both groups have the same size, being rather large, and more or less triangular-obovate. The ovary is apical, with two or more ovules; the pistil is rounded, slightly curved, naked. The stigma consisting of friable elongated cells is capitate in the middle, and covered by numerous nipple-shaped outgrowths. It can be smooth enough (without hairs), or surrounded with a ring of firm hairs, longer at the side of the vexillum and shorter at opposite sides (in cross-pollinated species). The stigma of lupin is open, since a rather long and wide channel goes from it inside the plant. The formation of the pistil happens simultaneously with the development of the stamens and corolla. The calyx of the flower is bilabiate, the cut of the lips is deep, either reaching the base of the calyx, or less frequently ending halfway. The upper lip is usually shorter than the lower one, and has a deep incision or bilabiate shape. The lower lip is integral and pointed. Less frequently, both lips of the calyx are integral. The tube of the calyx is quite short, and more or less cylindrical. The color of the calyx in lupin is more often not the same. The base part of the lips has the same color as the floral shoot, while the top part of both lips has another color, closer to the color of the corolla. Thus, the calyx can be either all green, or similar to the corolla. The shape of the calyx is an important common descriptor and is widely used in classification of the species. The floral bract of lupin usually falls down early. Specific differences of floral bracts (their size and shape) are also very characteristic and serve as an excellent customary descriptor. In various lupin species, the shape of floral bracts can be obovate, rhombic or other. The texture of floral bracts is also variable, from paleaceous and transparent to rough-skinned and dense. Pubescence on floral bracts, as well as on the calyx, corresponds to the type of pubescence on the whole plant. The color of floral bracts can be extensively variable: cream, salad, green, green with anthocyanin and dark-anthocyanin, almost black.

Flowering. Most of Lupinus spp. are cross-pollinated. However, there is a species which is almost exclusively self-pollinated (L. angustifolius). Within the genus Lupinus L. it is possible to observe in different species all transitional pollination types – from cross-pollination to self-pollination. Reports on frequent visiting of lupin flowers by bees and on cross-pollination opportunities are found in the works of Fruwirth (1919, 1927). One of the conditions enabling cross-pollination is the property of pollen to be separated from the flowers during the contact with insects. The mechanism of this process has been known for a long time and is called the technology of the swinging pump (Szafer, 1968). The essence of this process is that anthers push out in the keel’s edge constantly falling pollen during growth. A newly opened flower is usually abundantly covered with plenty of pollen and, as a result, is readily visited by insect. As a rule, the insects sit on the vexillum and wings and by their weight stir the wings and the adjoining keel. Consequently, the anthers hidden in it become straightened and start working as a pump pushing out pollen upon the abdomen and legs of an insect. Detailed information on the pollination of lupin by insects was published by Maissurjan and Atabiekova (1974). This work presents consistent description of the pollination process in L. ornatus Dougl. It goes on in the following sequence. Prior to flowering, under the dense closed petals of flower buds there are two circles of stamens: the stamens with longer anthers are colored more richly in yellow-orange color, and the stamens with reniform anthers are cream-green. One of the stamens of the lower circle is isolated from the other four. The stigma of the pistil develops a little earlier that the stamens. In the beginning of flowering process, the distance between the upper and lower circles of anthers becomes markedly reduced. The elongated anthers in the upper circle are the first to be opened lengthwise. The reniform anthers of the lower circle turn yellow; on their surface very sparse and hardly noticeable fractures appear. Afterwards, the stamens of the lower circle of anthers also extend to the top level. Then the anthers of the upper circle start to crack completely lengthwise. At this phase of development, a small quantity if pollen occurs on the keel’s edge. The extreme stamen of the lower circle, being slightly isolated from the others, develops markedly faster than the other four. This stamen is intermediate between the upper and lower stamens. Then, the lower circle outruns the upper one in its development. In this period the keel is filled with pollen. The flowering phase is terminated when all ten stamens completely wither. By that time, the ovary turns into a pod, and a light stain at the base of the flag acquires bright color. More accurate comparative analysis showed a number of essential differences in the manner of pollination between different species of lupin. For example, in L. mutabilis Sweet pollination happens in an opened flower. In L. nanus Dougl., some setback in flowering is from time to time observed, when the wings of the corolla do not fall from top to bottom, but rising above the keel straighten out to a part of the inflorescence axis. Thus, the transfer of androecium and gynaecium can go in opposite directions. Long-blossoming inflorescence at L. elegans is of an unusual kind. Drooping of the floral shoot occurs during flowering, and it carries away the wings with the keel, pistil and stamens. Fertilization comes to pass in the lowered flower; then the ovary turns into a pod and begins to grow and develop vigorously. With its new size, the pod does not find sufficient place in the keel and is gradually put forward beyond the borders of the keel. Then the floral shoot straightens and becomes a fruit spur. L. angustifolius is reported by many authors to be a self-pollinated species. This is justified, in particular, by the absence of pubescence on the stigma (Fig.19). As a result, pollen can freely penetrate the stigma of the pistil.
Fruwirth (1925) marked that artificial isolation had no negative effect on the seed yield of this species. This opinion is shared by other researchers: Hallquist, 1921; Sypniewski, 1930; Шарапов, 1937; Mackiewicz, 1958. Roemer (1924) also attributed L. angustifolius to self-pollinated crops, because cross-pollination happens in this species extremely seldom, approximately in one case out of one thousand. However, that situation can change depending on the growing conditions and especially on the availability of bees. Sengbusch (1931) also expressed that classification of narrow-leafed lupin as a self-pollinated species is more justified than with yellow lupin. Flowering and fertilization in self-pollinated species, for example in L. angustifolius, occur only in closed flowers and on the earliest phases of their development. L. luteus is peculiar for the presence of not only self-pollination but also cross-pollination habit. Pollination occurs in this species also on very early phases of flower development. Thus, the arrangement of flowers in this species is verticillate, while in L. angustifolius and L. albus it is alternate. However, most part of lupins, especially from the American continent, is cross-pollinated. There are grounds to believe that the polymorphic genus of Lupinus L. includes all possible biological forms: from a self-pollinated up to mandatory cross-pollinated species. The flowers of lupin are pollinated by bees gathering pollen with their abdomen (Megalichidae). They have a special brush on the abdomen for collecting pollen. This group of insects includes many wild bees living alone (Wojciechowska, 1976, 1993). Honeybee and bumblebees also take part in the pollination process of yellow lupin. There are very interesting observations concerning the variation of color in the corolla during the flowering phase of L. pilosus Murr. and many other species of lupin (Wo