ROLAND MCcKEE, Senior Agronomist,
A. J. PIETERS, Principal Agronomist, Division of Forage Crops and Diseases, Bureau of Plant Industry

[ABSTRACT.]  WHILE the legumes considered in this article occupy a place secondary to such crops as alfalfa and red clover, some have proved of great value in sections in which other legumes do not thrive.  The annual lespedezas, for example, grow on acid soils of low fertility, and without them a profitable agriculture in large areas from southern Iowa to the Gulf of Mexico would be difficult or impossible. The vetches and field peas likewise fulfill a valuable function in soil-improvement programs. Among the miscellaneous legumes about which comparatively little is now known, many forms especially suited to thrive in difficult situations could undoubtedly be found by intensive study and developed by systematic breeding.

The large family known as Leguminosae, or legumes, contains some of the most interesting and important crop plants. The legumes are distinguished from other plant groups particularly by their flowers and seed pods. The flowers are always irregular in shape, resembling a butterfly, while the pods have two valves or parts into which they readily divide, as in the case of garden beans or peas.  Most of the plants with pods that split into halves are legumes.  Another distinguishimg characteristic of legumes is their ability to take nitrogen directly from the air through association with bacteria that live on their roots, and to manufacture it into food for plant growth. By this means soils are enriched, soil fertility is maintained, and increased crop production is made possible. Some of the minor legumes used solely for cover crops and soil improvement are not so well known as crops grown for forage or grain, but they deserve much wider recognition and could well be used much more extensively.

Improvement work with legumes other than soybeans, alfalfa, and clovers has been very limited, but the importance of this group of plants justifies giving them serious consideration in any improvement program. They have been the subject of some genetic investigation, which will be discussed later.


While the legumes considered in this article occupy a place secondary to such crops as alfalfa and red clover, some have proved of great value in sections in which the other legumes do not thrive. The annual lespedezas, for example, because of their ability to grow on soils of low fertility and too acid for alfalfa and clover, have become the chief forage crops from southern Iowa to the Gulf of Mexico.  Without them a profitable agriculture in much of that region would be difficult or impossible.

Likewise the vetches and field peas fill an irreplaceable function in the soil-improvement programs of the Southern States, while the cowpea is a standard crop and the bur-clovers furnish winter grazing and soil improvement for millions of acres. Of the more recently introduced crops the crotalarias fill a place on sandy land not otherwise filled, and there are good reasons for believing that some of these species may become leading forage crops in the South.

The present soil-conservation program will require legumes for a variety of situations, but all must have in common the property of adaptation to reduced fertility and probably to soil acidity. Among these miscellaneous legumes now little known there may be some that with proper attention to selection will prove precisely suited for one or more such situations.

(Astragalus spp.)

The genus Astragalus contains a very large number of species, commonly called milk vetches. Most of them are especially adapted to dry and arid conditions. None of the species is of much commercial importance, although several are utilized locally and have forage value. No selection or other improvement work has been attempted in this genus, but the drought-resistant quality of the many species would seem to justify the conclusion that for dry-land and arid regions it is perhaps the most likely group from which to expect a legume of agricultural value.

(Meibomia spp.)

The beggarweed or tick trefoil group is composed of a fairly large number of species, few of which have been brought under cultivation.  The Florida beggarweed (Meibomia purpurea (Mill.) Vail) is the best known and the only one that is grown commercially in the United States. No attempt at improvement has been undertaken, although some natural selection no doubt has taken place, as the crop is harvested from cultivated stands. To what extent improvement is possible no one can say. The species are variable, however, and, no doubt, would respond readily to selection. One botanical variety, M. paniculata var. pubens (T. and G.) Vail, that has been grown in experimental plantings has growth habits that give promise of usefulness.

(Dolichos lablab L.)

The bonavist, which in habits, cultural requirements, and uses is much like the cowpea, has been used in Africa and southern Asia from ancient times, and many varieties are known to exist. Nowhere has improvement of the crop been attempted, although improvement with reference to nematode and wilt resistance and other characteristics no doubt could be effected (fig. 1).

Figure 1.—Bonavist (Dolichos lablab) showing general habit of growth.

(Medicago spp.)

Many species of Medicago that never are grown as cultivated crops enter into use for pasturage and are recognized as having very great value for this purpose. However, these have received no attention from the plant breeder. Other species that are grown for soil improvement and occasionally used for hay or seed have received some attention by experimenters. In a few instances selections of distinct forms have been made by practical growers who have increased their supply of seed and distributed it locally. Natural selection resulting from regional climatic differences, however, is responsible for much of the improvement in the medicagos.

In the case of spotted bur-clover (Medicago arabica (L.) All.) at least two new forms have appeared in the South in recent years.  One of these, Manganese bur-clover, was selected and named by A. Lee Andrews on his place at Lafayette in eastern Alabama. The other, Early Southern bur-clover, was selected and named by A. F. Ruff at Rock Hill, S. C. Both these varieties are earlier maturing than the commercial spotted bur-clover from which they were selected, and make as good a growth, if not better. A variety of bur-clover grown by H. H. Hopson in Coahoma County, Miss., under the name Giant bur-clover seems to be identical with the Early Southern and probably has the same origin. The Manganese bur-clover is somewhat earlier than the Early Southern and matures about 2 weeks ahead of the commercial spotted bur-clover. A spineless form of spotted bur-clover has been recognized as a subspecies, but this has never been grown commercially (fig. 2).

Figure 2.—Coiled seed pods of spiny and spineless forms of spotted bur-clover (Medicago arabica). In other species variation in length of spines is not uncommon.

California bur-clover (Medicago hispida Gaertn.) is very variable and responds to improvement by selection. Where bur-clovers occur spontaneously it is difficult to maintain pure stands of selected forms, and in pastures all that can be hoped for is to increase the relative amount of the improved varieties. Improved selections of bur-clovers were made by the United States Department of Agriculture some years ago and distributed from the United States Plant Introduction Garden at Chico, Calif., but supplies of seed have not been maintained and none are available commercially. The Texas Agricultural Experiment Station has made selections of the California bur-clover and is increasing seed of spineless forms for use in sheep pastures. The many other species are quite variable, like the species just mentioned, and could be readily improved by selection, but so far as is known most of these are in no way superior to the species now used commercially.  Thus there seems to be no reason to give them special attention in preference to the commercial forms. Black medic (Medicago lupulina is quite variable and could be improved readily by selection. No work has been reported on the improvement of this plant.

(Cicer arietinum L.)

Throughout the western United States attempts have been made grow the chickpea, which is native to western Asia, but nowhere has commercial production been successfully established, So far as known, no attempt has been made to develop superior varieties adapted to the United States, although it is known that the chickpea is variable and thus presumably could be improved by selection.

(Vigna sinensis (Torner) Savi)

A very large number of cowpea varieties are recognized. For the most part these have been developed through several hundred years by natural hybridization and incidental selection rather than by any planned improvement program. Introductions into the United States are recorded in the seventeenth century, but it is within the last century that specific reference is made tonamed varieties. In the United States in recent years considerable work has been done by experiment stations in the way of bringing existing varieties together for comparative testing.  This has resulted in a more extensive use of superior varieties and an elimination of inferior ones. While selection of superior plants has thus been the principal means of improving cowpeas, natural hybridization has played a very important part. Artificial hybridization in recent years has been attempted by a few plant breeders, and at least one outstanding variety has been developed in this way.

A cross between the Groit and Brabham varieties made by workers in the Department of Agriculture resulted in the Victor, characterized by its resistance to wilt and nematodes, which seriously damage most varieties. In the process of continuous growing and natural selection through long periods of time, varieties have changed to suit local conditions with reference to diseases, insects, and climate.  Apparently for this reason varieties that have been grown for years in the United States are much more resistant to bean rust than recently introduced varieties, and resistance to wilt and nematode in Iron, Brabham, Groit, and New Era varieties can be attributed to similar selection. In the case of the Victor, resistance to wilt and nematode has been inherited from its parents, the Brabham and Groit.

The origin of most of the commercial varieties of cowpeas is unknown. The Whippoorwill has been grown under that name since 1840, and the Iron has been known in South Carolina since 1888, but its origin is uncertain, and no information is available as to where the New Era may have come from. From a study of the characters of the Brabham it has been concluded that it is a hybrid between Iron and Whippoorwill, and the Groit is regarded as a hybrid of New Era and Whippoorwill. Improved selections from Brabham and Iron have been made in California and are being grown commercially.  Recently a variety of unknown origin was introduced into Florida and distributed to growers by the State Agricultural Experiment Station, under the name Suwanee. It has given high forage yields and seems to be especially well adapted to Florida conditions.

(Crotalaria spp.)

Crotalaria, commonly called rattlebox, is one of the newest agricultural crops that has become of commercial importance in the United States within the last 10 years. In India and a few other tropical regions several species have been in use for a much longer period. Three species are now grown commercially in the United States. These are Crotalaria spectabilis Roth, C. striata DC., and C. intermedia Kotschy. Being variable in plant characters, they lend themselves readily to improvement by selection. Late maturity, which makes the saving of a good seed crop difficult, has been one of the principal objections to crotalaria, and this character was the first in which improvement was attempted. The Department of Agriculture and the North Carolina, South Carolina, Georgia, and Florida Agricultural Experiment Stations working in cooperation have made selections for earliness in all of these species. At Columbia, S. C., an early-maturing variety of C. spectabilis has been developed that ripens its seed quite uniformly and 2 weeks or more ahead of the original lot from which it was selected. This is known as F. C. 18096 and in South Carolina has been called locally Carolina crotalaria (fig. 3). Some progress has been made in selecting early varieties C. intermedia and C. striata, but additional improvement is needed before these can be called superior.

Figure 3.—Crotalaria spectabilis, F. C. 18096 (a), and the commercial strain from which it was selected (b).

Since most of the Crotalaria species have been observed only in the wild, it is not possible to say how much improvement may be expected. Up to this time, however, little work has been attempted.

(Trigonella foenumgraecum L.)

Fenugreek (the name means Greek hay) occurs in the Mediterranean region and east as far as India. As grown under cultivation it shows varietal differences that probably are the result of natural regional development rather than artificial selection. So far as known no special improvement work has been done in any country. In the United States fenugreek has succeeded only in California, and there it grown only occasionally. At the California Agricultural Experiment Station at Davis the continued propagation for years of one of the best introduced strains of fenugreek appears to have resulted in the development of a superior variety well suited to at least that part of the State.

(Pisum arvense L.)

The history of the field pea is closely associated with that of the garden pea, since the distinction between the two groups is more one of usage than of botanical characteristics. The more extended use of varieties of field peas, however, has been in comparatively recent times, while the use of the garden pea extends back to earliest history.  The development of pea varieties through all these years has been largely the work of the gardener and the commercial seed grower, while the field husbandman has merely grown the varieties apparently best suited to his conditions.

In more recent times a number of experiment stations have made selections from the more promising varieties and introduced them.  The experiment stations in Canada and in the United States, particularly in Wisconsin, Colorado, Idaho, and Washington, have been most active in this work. The varieties O. A. C. 121, Wisconsin Perfection, and others represent the results of such work.

The new interest in field pea varieties has been largely in connection with their use as a cover and green-manure crop for the South. The Austrian Winter variety serves well for this purpose, but its susceptibility to disease and its inability to mature a good seed crop under southern conditions has lessened its popularity and called attention to the need for breeding and selection to overcome these difficulties.  The Department in cooperation with the Georgia Experiment Station has recently inaugurated a program with such results as the objective.  The State experiment stations at Auburn, Ala., and Knoxville, Tenn., have begun similar programs.

By bringing varieties together from every possible source and growing them under southern conditions it is probable that varieties with superior disease resistance and heavy seeding qualities will be found.  These can then be used in a breeding program to combine the other desirable qualities needed in a cover and green-manure crop.

(Lathyrus sativus L.)
Lathyrus sativus, commonly known as grass pea, has never become of commercial importance in any part of the United States, although in many places it makes good growth and produces fair seed crops.  In India the crop is of some importance, the natives using the seed for food and the plant for forage. A large number of varieties exist, differing in flower and seed color, growth of the plant, and size and shape of the seed. Varieties with large white seeds are superior for human food, while those with strong vegetative growth are preferred for fodder. Varieties of L. sativus occurring in India have been studied by Howard and Khan (11). While inheritance studies have been carried on with the sweet pea (L. odoratus L.), no one has studied L. sativus or related species in this way.
(Cyamopsis psoraloides DC.)

Guar, a summer annual, is used in India quite commonly for food for man and beast. Varieties exist, and the plants are more or less variable, but so far as known no special work on improvement has been attempted. In the United States guar has been used in cultural experiments but has not been recognized as having commercial value.

(Anthyllis vulneraria L.)

No commercial plantings of kidneyvetch are made in the United States, and so far as known no selection or other improvement work has been attempted in this country. In Wales, selection work with this plant is in progress, but results have not yet been published. In Denmark the improvement of kidneyvetch has been attempted, and strains Tystofte No. 8 and Tystofte No. 28 when compared with commercial kidneyvetch were found to be more productive.

(Pueraria thunbergiana (Sieb. and Zuce.) Benth.)

No work has been done in the improvement of kudzu-bean, but there is no reason to suspect that it would not respond readily to improvement by selection.

(Lespedeza spp.)

The lespedezas or bush clovers as a group are still wild plants. Only one species, Lespedeza striata (Thunb.) H. and A., has been long known to agriculture. The others are of such recent introduction that the possibilities of improvement have not been adequately explored.

Agronomically there are two groups of lespedezas. The annual consists of two species, Lespedeza striata, or common lespedeza, and L. stipulacea Maxim., or Korean lespedeza. The second and much larger group consists of perennial plants, of which one species only, L. sericea (Thunb.) Benth., has recently been introduced to agriculture.

As is the case with most wild plants, the lespedezas are more or less variable, plants differing in size of leaflets and height of growth, in habit, and especially in date of maturity.

Lespedeza striata, common lespedeza, was first found in Georgia in 1846, spread rapidly over the lower South, and gradually worked north. The character that made this spread possible was undoubtedly difference in date of maturity between plants. As the species worked farther north the earliest forms seeded and reproduced themselves. This process went on generation after generation, until now the common lespedeza is established as far north as central Indiana.

The variations in habit of growth were made the basis of selection by the late S. H. Essary, of the Tennessee Agricultural Experiment Station. He began his study of-individual plants in 1912 and found a great variation, especially in the habit of growth. Among the variants was one having an erect habit and great productive capacity, which was segregated and put into experimental plantings in 1921.  This was introduced by the Tennessee station as Tennessee 76 and is one of the leading late-maturing varieties.

Another variety with larger leaflets and ranker growth than usual was found by J. B.S. Norton, an explorer of the Department of Agriculture, growing wild near the city of Kobe, Japan, and was introduced as Kobe.  It is somewhat earlier in maturity than Tennessee 76 but does not grow so erect except in thick stands. Another variety of common lespedeza, inventoried as no. 81742, was collected by Dorsett and Morse, of the Department, in Japan in 1929 and is the earliest maturing form of Lespedeza striata. In habit it is nearly as erect as Tennessee 76.

The possibilities for improvement in this species have not been exhausted, but its natural dependence on high temperatures will probably prohibit pushing the species much farther north than its present limits.

The Korean lespedeza also shows variations, especially in date of maturity. Two extra-early forms have been found growing wild in Manchuria. One of these, no. 65280, has been introduced as Harbin; the other, no. 59379, is a week later and makes a little larger growth than Harbin. Neither variety promises much usefulness, because the plant is low in growth habit and the yield is consequently small.  They do, however, show the range of possibilities in the development of earliness. Harbin has matured seed at Winnipeg.

Two varieties, an early Korean, no. 19604, maturing about 2 weeks earlier than standard Korean, and a late form, no. 19601, maturing 2 weeks later than the standard Korean, have been selected at the Department nursery at the Arlington Experiment Farm, Arlington, Va. (fig. 4). The early form, no. 19604, has been released for use in northern Iowa, northern Illinois, and adjacent areas. Its habit of growth and yielding ability are like that of standard Korean, and it differs only in earliness. The late form, which is matting in growth habit, is still under observation.

Figure 4.—Korean lespedeza (Lespedeza stipulacea). Selected plants showing different growth habits. The plant on the left (a), with horizontal lower branches, makes a low matting growth, while b is quite upright.

Here again the possibilities of selection have not been fully explored, and in the future better varieties, or varieties better suited to certain conditions, may be selected.

The group to which Lespedeza sericea, the perennial lespedeza, belongs consists of several species differing from one another in botanical characters and in habit of growth. Certain variations in L. sericea itself have been noted. The botanical group consists of L. sericea, an erect, rather strict plant with narrow leaflets; L. inschanica (Maxim.) Schindler, with larger leaflets and lax habit; L. latissima Nakai, a prostrate plant; and L. juncea, intermediate in habit between L. sericea and L. latissima. Variations in L. sericea are found in width of leaflets, height of growth, coarseness and number of stems, and earliness (fig. 5). Of the varieties studied, no. 04730 is early, tall, and coarse, no. 12087 is later, with finer stems, and no. 19284 is from a single plant selection out of no. 04730 and is somewhat more uniform than the parent. These variations are neither important nor significant, but they show that a more intensive study may uncover variations of agricultural significance.

Figure 5.—Lespedeza sericea, showing variation in size and habit of growth. Plants in each row are from seed of a single mother plant.

One of the most important fields for selection with Lespedeza sericea is that of finding forms with a low tannin content. Studies made show that the tannin content varies among individuals, ranging from about 5 or 6 to 10 or 11 percent. From the standpoint of palatability as well as forage value, it is important that a form with low tannin content be found and introduced. Work with this object is being carried on.

The shrubby species, as Lespedeza bicolor Turcez. and its allies, have not been studied intensively enough to determine the range of variation, and it does not seem probable at this time that they will offer great possibilities for agricultural usefulness.

(Lotus spp.)

Several species of Lotus (not to be confused with the water-lily of that name) are recognized as being of some commercial importance in several European countries and in Australia, but in the United States none is recognized as having special value. Seed of Lotus corniculatus L. and L. uliginosus Schkuhr, two perennial species that are used for hay and pasturage, is available through the seed trade, but no varieties are listed. So far as known no special improvement of these plants has ever been attempted, although varieties with low cyanophoric glucoside content have been reported. The elimination of this glucoside is desirable, since in the course of digestion it is changed to hydrocyanic acid, which is injurious to animals.

(Lupinus spp.)

Several species of Lupinus are grown commercially as field crops in European countries, and at least one species is being used in Australia. None, however, has ever been commercialized in the United States, since experimental plantings have indicated that in most places they are not well adapted and cannot be used as economically as legumes now commonly grown.

Varietal improvement in this group of plants has been confined largely to work in European countries, particularly Germany and the Union of Soviet Socialist Republics, where in recent years attempts have been made to produce lupines devoid of alkaloid poisons. Both German and Russian workers report having developed strains with little or no alkaloid, which they call “sweet lupines.” The object has been to produce plants and seed that could be used for both livestock and human consumption without injurious results.

PEANUT (Arachis hypogaea L.)
The improvement of the peanut for forage has, received little attention. In the United States the experiment stations of Florida and Texas have undertaken hybridization studies, but the results of this work have not yet been published. Arachis nambyquare Hoehne and A. rastiero A. Cheval. have been used in crossing with the common peanut (A. hypogaea), and attempts have been made to introduce other wild species. These offer possibilities but as yet are too little known to justify a statement regarding results.
(Cajanus indicus Spreng.)

In many tropical countries the pigeonpea is recognized as one of the most valuable legumes. In India a large number of varieties exist and the superior value of some of these is recognized. In the Hawaiian Islands, where the pigeonpea was introduced, improvement of the crop has been undertaken and selection and breeding work have resulted in the production of superior varieties. New Era strain X is recognized in Hawaii as one of the most desirable and much superior to the strains from which it was produced. In the United States varieties have been grown for selection work at several southern stations. The plantings at the Florida Agricultural Experiment Station at Gainesville have been the most extensive. Wide variation in the plants has been observed and early-maturing varieties have been selected, but no variety has been found sufficiently well adapted to justify commercial use. One variety has matured as far north as Washington, D. C., but seed production has been light.

(Onobrychis vulgaris Hill.)

Sainfoin has been grown in France and other European countries for over 300 years. In the United States it has never become of commercial importance, although in experimental trials at a number of experiment stations it has made good growth. A number of botanical varieties are known to exist, but improvement of most of these for agricultural use so far as known has never been attempted. Commercial sainfoin is quite variable and includes both biennial and perennial forms. Improvement by selection in both of these no doubt could be easily effected. The Washington Agricultural Experiment Station at Pullman is making selections of sainfoin with the idea of obtaining better adapted varieties.

(Ornithopus sativus Brot.)

In central Europe serradella is used on acid sandy soils, but in the United States production or use on a commercial scale has never been successful. Improvement of varieties is reported from Germany, although the amount of work seems to have been very limited.

(Sesbania macrocarpa Muhl.)

The seed of sesbania as found in the commercial trade is gathered wild and volunteer plants and represents the species as it is ind growing naturally. The plant grows in wet or moist soils in the Southern States. No attempts at improvement have been made.

(Hedysarum coronarium L.)

Attempts to grow sulla in the United States have indicated that it succeeds fairly well in a number of places so far as growth is concerned, it does not appear to have a place in competition with legumes now commonly grown. In several European countries it is grown as a commercial crop and is considered as having superior value for special localities. While varieties are known to exist, few attempts at varietal improvement seem ever to have been undertaken. In South Wales a special strain was selected for many years and tested on the experimental farms, but for other countries no work is reported.

(Stizolobium spp.)

The Florida velvetbean (Stizolobium deeringianum Bort) is one of leading legume crops in the southeastern United States. It was introduced in Florida previous to 1875 and for many years was confined largely to that State, since it would seldom mature much farther north.  Attempts to grow the crop in Georgia and Alabama, later resulted in the selection of mutants that required a much shorter season to mature and were well adapted otherwise. In 1906 Clyde Chapman, of Sumner, Ga., observed an early-maturing plant that he selected and grew under the name Hundred Day Speckled. This variety later became known as Georgia and is the earliest maturing commercial variety now grown.

In 1908 R. W. Miller, of Broxton, Ga., selected an early-maturing variety that was grown as Clark’s velvetbean, but this proved to be solike the selection made by Chapman that it later was considered identical with the Georgia.

In 1911 a Mr. Blount, of Flomaton, Ala., selected an early-maturing variety, which he called Alabama. This was not quite so early maturing as the Georgia but was sufficiently good to become commercialized and and is still being grown.

A variety of velvetbean known as the bush velvetbean (fig. 6) was selected on the farm of Roan Beaseley at Kite, Ga., about 1914.  As its name implies, it is a bush or bunch variety, lacking the twining habit of other velvetbean varieties. In work with this variety at the Mississippi branch experiment station at McNeill, H. R. Reed noted a white-seeded variant and made selections of it with the idea of using the white seed character to identify the bush variety. This selection did not prove to be stable as to seed color, and subsequent selection was continued. Now after 10 years a variety seems to have been obtained that reproduces true to color.

Figure 6.—Bush velvetbean.

At the Florida station a hybrid variety known as Osceola was produced by John Belling, who made a cross between the Florida velvetbean (Stizolobium deeringianum) and the Lyon velvetbean (S. niveum (Roxb.) Kuntze). This is a vigorous, heavy-yielding variety with a medium season, so that it is adapted as far north as central Georgia.

The velvetbean offers opportunity for much further improvement, and the importance of the crop suggests that such work could be done with profit.

(Vicia spp.)

A large number of species of Vicia are in general use, all of which go under the general name vetch. The species that are of importance commercially are Vicia villosa Roth, V. sativa L., and V. pannonica Crantz. Others that are occasionally used are V. atropurpurea Desf., V. calearata Dest., V. monantha Desf., V. dasycarpa Ten., and V. angustifolia Grufberg. So far as known, vetches are close-pollinated, and seldom, if ever, does crossing take place. All species seem to be variable, but differ somewhat in this respect.  Hairy vetch (V. villosa) is much more uniform than common vetch (V. sativa), and while several species seem to be less variable than hairy vetch, none are so uniform but that improvement can be made by selection.

Common vetch has been grown in the Mediterranean region for centuries, and through regional selection and otherwise a large number of varieties have been developed, differing in seed color and growth characteristics. Much of the improvement in this crop, no doubt, traces to selections made by local growers, but natural selection due to continued regional production probably also played a part. In later years experiment stations have developed improved varieties for local use, but published statements regarding such work are very meager.

Most of the work in vetch improvement in the United States has been carried on at Corvallis, Oreg., by cooperation between the Department and the Oregon station.

   In this region common vetch has been grown for many years, and the commercial strain now grown is the result of natural selection through this long period. The winter temperatures of western Oregon and western Washington represent about the extreme of cold that the most hardy strain of common vetch will endure. Thus the variety that has survived and been developed and increased here in commercial production represents one of the most winter-hardy of the common vetches.

Since other vetches are of very recent introduction commercially, natural selection has played little if any part in the development varieties.

In the case of both hairy vetch and common vetch the work of the Oregon station, cooperating with the Department, has resulted in improved varieties that already have been or are becoming commeralized.  Here a vigorous growing variety of smooth vetch (V. villosa var.), lacking the heavy pubescence of hairy vetch, was selected in 1926 by H. A. Schoth and is now grown quite extensively. A good deal of the seed of V. villosa imported from central Europe under the name hairy vetch is smooth vetch and resembles the smooth vetch variety grown in Oregon. Of the common vetch (V. sativa) selections that have been developed, a white-flowered variety, F. C. 02830, that is somewhat superior in vigor and winter hardiness to commercial Oregon common vetch and was selected by Schoth in 1915, is perhaps outstanding. The white-flower character offers a ready means of identification in the field and will enable the grower to keep his seed pure.

In the Netherlands, Denmark, and other European countries where common vetch is grown, improved varieties adapted to local conditions have been developed, but none of these, so far as they have been tested in the United States, has proved superior to varieties developed in this country.


INHERITANCE studies have been made in few of the so-called miscellaneous forage legumes. From general observation and in some cases from definite experimental demonstrations it is known that a number legumes are self-pollinated and rarely if ever are cross-fertilized.  Whether or not these can be crossed or hybridized has in many cases not been determined.

Studies of several species of Phaseolus in India indicate that the urd bean (P. mungo L.) and the mung bean (P. aureus Roxb.) are usually self-pollinated, although the mode of anthesis in many cases would permit of [sic] cross-pollination.

Unpublished observation of legume plantings in the United States indicates that Crotalaria, Vicia, Lathyrus, and the annual species of Medicago are largely self-pollinated and seldom if ever cross-fertilize.  A close study of anthesis in such cases, however, might indicate a means of effecting crossing.

No work on crossing lespedezas has been done. The technical difficulties are of the same order as those presented by the clovers and are due to the fact that the flowers are small and difficult to manipulate. There is another difficulty in addition. The three commonly known species of Lespedeza—L. striata (Thunb.) H. and A., L. stipulacea Maxim., and L. sericea (Thunb.) Benth.—have flowers of two kinds, and both kinds occur in the same cluster. One set of flowers bears a corolla and is therefore conspicuous; the other and more numerous kind has closed flowers with reproductive parts complete but with no corolla. These flowers are consequently self-fertile.  Although definite data are wanting, the conclusion drawn from observation is that these species are self-fertile. No sign of hybridizing has been observed, though the species have been grown side by side for years. The progeny from the seed of individual plants is always true to the mother plant.

In the case of serradella, which has been reported as self-fertile, plants inbred for four generations did not lose vigor.

While crossing in many legumes seems to be uncommon, there are others that cross readily and are naturally cross-fertilized. Species that have been used in inheritance studies are the cowpea (Vigna sinensis (Torner) Savi), chickpea (Cicer arietinum L.), adzuki bean (Phaseolus angularis (Willd.) Wight), bonavist (Dolichos lablab L.), horsebean (Vicia faba L.), pigeonpea (Cajanus indicus Spreng.), lupine (Lupinus spp.), peanut (Arachis hypogaea L.), velvetbean (Stizolobium deeringianum Bort), and field pea (Pisum arvense L.).

(Cicer arietinum L.)

While self-pollination is the general rule in Cicer arietinum, varietal crosses have been recorded. In 1915 Howard, Howard, and Khan (10) reported the growing of selections that split in the F2 generation, indicating natural crossing. Their observations were not made in sufficient detail, however, for the deduction of the various color factors present.

More recently inheritance studies were made by Khan and Akhtar (13) relating to color and number of flowers. In making artificial hybrids it was found that in order to prevent accidental crossing emasculation should be effected the evening of the second day preceding fertilization. The flowers open naturally on bright days between 9 and 10 a. m., and the pollen should be applied at this time.  Five crosses were studied and the following results reported (13, p. 155):

  1. The flower color depends upon the interaction of several factors.
  2. Blue color depends on a single factor B.
  3. Pink color is produced by a factor P in the presence of B.
  4. In the absence of B the flower is white whether P is present or absent.
  5. Greenness in the standard is developed in the absence of the factor W.  Greeness is therefore recessive to nongreen.
  6. Singleness depends upon a factor S and is dominant to doubleness.
(Vigna sinensis (Torner) Savi)

The cowpea has been recognized as one of the legumes that offer excellent facilities for inheritance studies, and investigations by Spillman and Sando, and in particular by S. C. Harland, have indicated what may be expected by more extended research. In studies made by Spillman and Sando (16) flower color was found to be correlated with coloration in the seed coat, joints, peduncles, stipules, and petioles, and complete linkage was observed in certain seed-coat-color factors.

It was determined that the presence of anthocyanin coloration in the stem and leafstalk is due to a single unit factor, dominant to its absence.

In the case of certain seed-coat-color patterns, two factors that are inherited independently were found to influence the color pattern, resulting in the expected 9:3::3:1 ratio.

Seventeen Mendelizing factors of cowpeas were definitely identified.  These factors with the characteristic effect they produce are as follows:

   A.  Seed pod curved after the manner of the alfalfa seed pod.
   B.  Brown seed coat.
   D.  Dense speckling, characteristic of the New Era variety.
   E.  Narrow eye.
   F.  Very fine and dense speckling, giving rise to blue seed coat.
   G.  Dotting; converts Holstein spots into numerous small ones.
   H.  Holstein type of seed-coat spotting.
   I.  Eye with indefinite margin.
   L.  Longitudinal furrowing of the surface of the seed.
   N.  Presence of anthocyanin pigment factor.
   P.  Purple seed coat.
   R.  Red seed coat. (This is the general factor for color, the absence of which determines white seed coat, white flowers, and absence of pigment in vegetative parts.)
   S.  Black spotting on certain types of seed coat.
   T.  Less dense speckling, characteristic of the Taylor variety.
   U.  Buff, or clay-colored, seed coat.
   W.  Whippoorwill type of seed-coat spotting.
   X.  Taylor inhibitor cancels (crosses out) the effect of T.
The eight factors B, F, N, P, R, TF, U, and X, either singly or in combination, give rise to the ten distinguishable seed-coat colors, purple, black, dull black, blue, coffee, maroon, buff, red, pink, and white. The factors P, B, D, T, and F and probably S belong to a linked group. The factors D, T, and F restrict the distribution of color pigments in the seed coat. Three independent eye factors I, H and E, either singly or in combination, give rise to the five distinct types of eyes, Watson, Holstein, small eye, narrow eye, and very small eye.
  Indications were found that when all of the pigment factors are absent, including factor R, there is a tendency for the seeds to be small, weak, or abortive.
  The flower color has been found to be correlated with the coloration in the seed coat, joints, peduncles, stipules, and petioles (16, pp. 282-283).

Experiments reported by Haigh and Lochrie (6) indicate a progressive variation with age of a simple Mendelian ratio in the cowpea.  The results in the F, cultures from successive days of flowering showed an orderly variation in the simple Mendelian 3:1 ratio, an excess of recessives in the first 9 days being compensated for by an excess of dominants as the plants grew older. No cause for this phenomenon was discovered.

Hofmann (9), in experiments at the University of Illinois, found that crosses made in the greenhouse between California Blackeye and Blue Goose show definite evidence of hybrid vigor.

(Dolichos lablab L.)

In studying Dolichos lablab, Harland (8) found that dehiscence of the anthers takes place at least 1 and sometimes 2 days before the flowers open. Studies in inheritance showed that in the case of determinate and indeterminate growth the segregating ratio in the F2 generation was 3:1, with complete dominance of the indeterminate factor.

Two factors were found to influence color, each being transmitted independently, resulting in the expected 9:7 ratio. One of these factors has no effect except in the presence of the other, when it converts white flower into purple, and brown seed into black, and causes pigmentation of the nodal region.

(Lupinus spp.)

Burlingame (2) reports studies of Lupinus species with reference to variation and inheritance. His findings show that races with dark-blue and pink flowers breed true and that races with striped white flowers are heterozygous for a single factor, which in the homozygous condition produces white flowers. Light-blue flowers are due to a single dominant factor, indistinguishable in the homozygous and heterozygous condition.

   Dark seed coats are linked with dark-blue flower color, but probably due to separate factors.
   The factors for light-blue and striped-white flowers are both allelomorphic to that for dark-blue and not improbably constitute a system of multiple allelomorphs.
   Mutations are frequent, some are already known to be dominant, and others appear to be in the nature of additions of new characters and factors and so progressive in the sense of de Vries (2, p. 447).

Hallqvist (7) studied seven different types of flower color and five types of seed color in Lupinus angustifolius L. His conclusions were as follows (7, p. 344):

   One fundamental colour factor has been demonstrated (pure red). A synthesis of blue colour has been obtained from crosses between bluish red and violet flower colours. One "dilution" factor has been found to be present.
   Pleiotropic correlation has been demonstrated between certain flower and seed colours.
   Three flower colour factors have been found to form a linkage group. The linkage between two of the factors is very close, if not complete. The other linkage value represents a crossover percentage of about 22%.

Greb (5), in studying Lupinus albus L., found that there are at least two genetically different rootlet types in this species. These differ in rate of rootlet elongation in young seedlings and in time of development of the root hairs. The ratio in the F2 was 2:1, which suggested that the homozygous nonhairy might be lethal.

(Phaseolus angularis (Willd.) Wight)

Kakizaki (12), in a study of crosses between Miyako and Donsu varieties of adzuki bean, found that reddish purple in the stems, black spotting on red seed coats, and blackish brown in ripe pods were dominant over their recessive allelomorphs, green stems, unspotted seed, and brown pods. From the segregating ratios in the F2 it was concluded that color of stem was due to the interaction of two factors, while the black spotting of the seed coat and color of ripe pods were influenced by only one factor pair.

Colored stems are completely correlated with black-spotted seed coats, and colorless (green) stems with unspotted seed coats.

The factor P for reddish-purple color of stems and its recessive allelomorph p (green stems) are very closely linked with S, a factor for black-spotting of seed-coats, and its recessive allelomorph s, respectively, and hardly any crossing over occurs between them.
The I factor, which intensifies the purple color of the stem, and its recessive alleomorph # are also very closely linked with the B factor, which produces a blackish-brown color of the ripe pods, and b, its recessive allelomorph, and this linkage is also very close, so that crossing over hardly ever occurs between them.
The P-S linkage group is independent of the J-B linkage group.
Presence of S in a homozygous condition produces more intense spotting of the seed coats than when it is present in a heterozygous condition (12, p. 177).

HORSEBEAN (Vicia faba L.)

The horsebean is known to cross readily but has been used comparatively little in inheritance studies. Darlington (3) studied variegation and albinism and found that variegation is a heterozygous type of which albino and normal are the homozygous types. Sirks (15) in a study of quantitative inheritance in Vicia faba presented evidence to indicate that quantitative factors do exist.

(Stizolobium spp.)

Several species of Stizolobium have been hybridized. Inheritance studies made at the Florida Agricultural Experiment Station a number of years ago were reported in the annual reports of that station for 1910, 1912, 1913, and 1915. Species that have been hybridized are S. deeringianum Bort X S. pachylobium Piper and Tracy, S. deeringianum X S. niveum, and S. deeringianum X S. hasjoo.

The inheritance studies are concerned mostly with the cross S. deeringianum X S. niveum, which are the species of most economic significance. In these crosses Belling (1) found that color in wings and standard, length of seed, curve in pods, and open and closed pods were influenced by a single factor pair, and size of pod and length of pubescence on pod by two or more factors, while mottling was due to three independent factors. Correlations were established between lateness of flowering and number of flowers in a raceme and between length of pod and seed.

(Cajanus indicus Spreng.)

Inheritance studies with the pigeonpea carried on at the Hawaiian Agricultural Experiment Station have been reported by Krauss (14, p. 18). According to his findings—

   * * * red flower standards are dominant over yellow; blotched or speckled seed dominate over solid colored, and maroon-blotched pods are dominant over solid light-tinted pods. Pubescent pods are dominant over glabrous; large, flat pods are dominant over small round pods, and large seeds over small seeds.
   Four and five seeded pods are dominant over 3 and 4 seeded pods. Round seeds, slightly flattened, dominate over all others of widely different shapes, including spherical, oval, flattened, and irregular. The axillary flowers and pods dominate over terminal inflorescence. In stature blended inheritance is observable, very dwarf varieties when crossed with very tall varieties produce an intermediate type, and two varieties when crossed almost invariably produce a type that is taller and more vigorous than either parent. Crossing an annual type on a perennial type appears to produce perennial forms. This behavior has been found to remain constant, practically complete dominance for some well-defined differentiating unit characters being the rule.
   When red dorsal standard sorts were crossed with red types, it was noticed that the solid red changed to red lacing, and when extremely tall and dwarfed forms were crossed, the first generation was of intermediate height. Wherever dominance is apparent the second generation shows fairly definite Mendelian segregation as well as definite linkage between some characters. Dihybrid crosses appear to adhere rather closely to the 9:3::3:1 ratio.

Studies in Cajanus indicus relating the inheritance of color in the flower and seed coat have been reported by Dave (4). In most cases one factor pair controlled color inheritance, but in others two independently inherited factors were operative, in agreement with the general results obtained by Kraus. Complete linkage was noted between orange-yellow flowers and purplish-black seed, yellow flowers and back of standard with purple veins, and base diffused purple and green pods; and purple color at the back of the standard was closely linked with maroon color of the pod.


(1) Belling, J. 1911. SECOND GENERATION OF THE CROSS BETWEEN VELVET AND LYON BEANS. Fla. Agr. Expt. Sta. Rept. 1911: lxxii-civ, illus.
   (2) Burlingame, L. L. 1921. VARIATION AND HEREDITY IN LUPINUS. Amer. Nat. 55: 427-448, illus.
   (3) Darlington, D. C. 1929. VARIEGATION AND ALBINISM IN VICIA FABA. Jour. Genetics 21:  [161]-168, illus.
   (4) Daves, B. B. 1934. INHERITANCE OF CHARACTERS IN CAJANUS INDICUS. Indian Jour. Agr. Sci. 4: 674-691, illus.
   (5) Greb, R. J. 1935. TWO GENETICALLY DIFFERENT ROOTLET TYPES IN THE LUPINE. Jour. Heredity 26: 503-504, illus.
   (8) Harland, S. C. 1920. INHERITANCE IN DOLICHOS LABLAB., L., PARTI. Jour. Genetics 10:219-226.
   (9) Hormann, F. W. 1926. HYBRID VIGOR IN cow PEAS. Jour. Heredity, 17: 209-211, illus.
   (10) Howard, A., Howard, G. L. C., and Khan, A. R. 1915. SOME VARIETIES OF INDIAN GRAM (CICER ARIETINUM L.). India Dept. Agr. Mem., Bot. Ser. 7: 213-235, illus.
   (11) Howard, G. L. C., and Khan, A. R. 1928. THE INDIAN TYPES OF LATHYRUS SATIVUS L. (KHESARI, LAKH, LANG, TEORA). Indian Dept. Agr. Mem., Bot. Ser. 15: 51-77, illus.
   (12) Kakizaki, Y. 1923. LINKED INHERITANCE OF CERTAIN CHARACTERS IN THE ADZUKI BEAN. Genetics 8: [168]-177.
   (13) Khan, A. R., and Akhtar, A. R 1934. THE INHERITANCE OF PETAL COLOUR IN GRAM (CICER ARIETINUM L.). Agr. and Livestock in India 4: 127-155, illus.
   (14) Krauss, F.G. 1932. THE PIGEON PEA (CAJANUS INDICUS), ITS IMPROVEMENT, CULTURE, AND UTILIZATION IN HAWAII. Hawaii Agr. Expt. Sta. Bull. 64, 46 pp., illus.
   (15) Sirks, M. J. 1929. GROWTH AND INHERITANCE OF LEAF DIMENSIONS IN THE BROADBEAN (VICIA FABA L.). K. Akad. Wetensch. Amsterdam, Proc. Sec. Sci. 32: 1066-1084.
   (16) Spillman, W. J., and Sando, W. J. 1930. MENDELIAN FACTORS IN THE COWPEA (VIGNA SPECIES). Mich. Acad. Sci., Arts, Letters, and Papers 11: 249-283, illus.

TABLE 1.—Chromosome numbers in legumes
Name*Chromosome numbers**Reference no.
Acacia arabica Willd.----±52, ±104***(26), (28)
A. baileyana F. Muell.13(59, 60)
A. cyanophylla Lindl.26(26), (28)
A. dealbata Link
A. decurrens Will(27, 28)
A. dermatophylla F.13(15)
A. eburnea Willd±52, ±104(27, 28)
A. farnesiana Will(26, 28)
A. horrida
A. longifolia Willd26(27, 28)
A. lophantha24(74)
A. nilotica±52, ±104(26)
A. podalyiaefolia A. Cunn26(26, 28)
A. saligna Wendl(27)
A. scorpioides A. Chev., var. adstringens (Schum. and Thon.) A. Chev.52, 104, 208(27, 28)
A. scorpioides A. Chev. var. nilotica Benth.±52, ±104(28)
A. scorpioides A. Chev. var. pubescens Benth(27,28)
Aeschynomene indica L.20(44)
Amorpha californica Nutt10(48)
A. fruticosa L.20
A. microphylla10
Amphicarpa monoica (L.) Ell(11)
Anthyllis alpestris Kit.12(7)
A. barba-jovis L.14
A. gerrardi L.16
A. maritima Schweigg12(7)
A. tetraphylla L.16(7)
A. vulneraria612(1)
Arachis hypogaea20e40e(44)
A. hypogaea (Spanish and small Japan peanuts)10(55, pp. 338-348)
A. nambyquare40(37)
A. prostrata Benth. var. rasteir±40(28)
A. rasteiro Chevalier (?)40(38)
Astragalus alopecurioides8(48)
A. altaicus Bunge.16(4)
A. baeticus L8(47)
A. candidissimus Led16(4)
A. echinus DC64(4)
A. edulis Durψ14(48)
A. exscapus B. transsilvanicus A. and G.= A. transsilvanicus Barth.164
A. falcatus Lam847
A. galegiformis847
A. hamosus L2448
A. hypoglottis L.164
A. massiliensis Lam1648
A. membranaceus Fisch. {Bunge}164
A. mollis878
A. monspessulanus L.847
A. secundus DC4866
A. sesameus L847
A. sieversianus Pall.164
A. sinicus L844
A. transsilvanicus164
A. vulpinus Willd847
Baptisia australis R. Br9c71
B. australis R. Br. var exalta Sweet185
B. sulphurea Engelm971
B. tinctoria R. Br185
Biserrula pelecinus L.847
Cajanus indicus Spreng.1165
Calophaca wolgarica Fiscl847
Canavalia ensiformis DC1144
C. gladiata DC227
Caragana arborescens Lam16c7
C. frutescens DC324
Carmichaelia australis1548
Cassia didymobotrya142870
C. dimidiata1674
C. fistula1277, pp.541-623
C. leschenaultiana DC2444
C. mimosoides L8, 1644
C. occidentalis L1356
C. purpurea Roxb1031
C. sophera L1244
C. tomentosa L12e
C. tora L1336
Cercis siliquastrum716
Cicer arietinum L71413
C. kabulium81655
Clitoria ternatea L167
Colutea arborescens L164
C. halevica Lam848
C. media Willd. (C. arborescens L. X C. orientalis Lam.)
C. orientalis Lam
Crotalaria alata Ham.44
C. anagyroides H. B.44
C. arenaria Senth32
C. juncea162
C. obovata G. Don832
C. retusa L44
C. usaramoensis Back. {Baker}
C. valetonii Back
Cyamopsis vscraloides DC71462
Cystisus canariensis O. Kuntze465
C. nigricans L2481
C. purpureus4873
C. scovarius Link24c44
C. scovarius4874
C. sessilifolius L.525
Desmodium grandiflorum (Walt.) DC1111
D. perpesium DC§1144
Dolichos biflorus2463
D. lablab L11c78
D. lubia Forsk227
D. multiflorus2458
D. niloticus Del227
D. ornatus Wall227
Dorcynium herbacium Vill147
D. hirsutum Ser147
D. rectum Ser147
D. suffructicosum Vill147
Erythrina crista-galli L447
Galega officinalis L847
G. orientalis Lam847
Genista ferox Poir485
G. pilosa L24
G. sagittalis L44(42-45)
G. tinctoria v. angustifolia Ledeb48
G. trangularis Kit48(48-50)
Glycine gracilis Skvortozow204024
G. hispida3830
G. hispida Max204024
G. soja Sieb. and Zucc20c40d44
G. soja var. akasaya3886
G. ussuriensis407
Glycyrrhiza aspera Pall164
G. echinata L847
G. uralensis Fisch164
Hedysarum elongatum Fisch. var. albiflorum Ledeb1466
Hymenocarvus circinnatus Savi167
Indigofera asvera Perr832
I. decora Lindl484
I. diphylla832
I. gerardiana Wall2447
I. kirilowi Maxim844
I. parviflora Heyne732
I. pseudotinctoria Matsum844
I. pseudotinctoria1674
I. suffructicosa Mill1644
I. sessiliflora DC1632
I. viscosa Lam832
Laburnum adami4873
L. alpinum Griseb48(50)5
L. vulgare4873
Lathyrus angulatus L771
L. annuus L771
L. avhaca L7c12
L. articulatus L7c14c14
L. cicera L7c1413
L. cirrhosus Ser771
Lathyrus ciymenum L771
L. crassipes Gillies1469
L. dumetorum Phillippi1469
L. ensifolius Bad771
L. grandiflorus Sibth. and Sm7c1414
L. heterphyllus L771
L. hirsutus L771
L. latifolius L7c1484
L. macropus Gillies1469
L. magellanicus Lam1453
L. maritimus Bigel7c44
L. niger7c14c(15)
L. nigrivalis A. Burkart1469
L. nissolia L.771
L. numidicus Batt771
L. ochroleucus Hook1469
L. ochrus DC7c1413
L. odoratus L7h14d84
L. palustris769
L. pannonicus [Garcke]1453
L. paranensis A. Burkart1469
L. parodii A. Burkart1469
L. pratensis L71453
L. pubescens Hook. and Arnot771
L. quadrimarginatus Bory and Chaub1469
L. rotundifolius Willd771
L. sativus L7d1414
L. sessifolius Hook. and Arnot1469
L. setifolius L769
L. sylvestris L771
L. sphaericus Retz771
L. tingitanus L7c1453
L. tuberosus L721
L. venosus Muhl2869
L. vernus Bernh771
Lens esculenta Moench778
Lespedeza bicolor Turcz944
L. cyrtobotrya Miq9
L. daurica Schindl3611
L. homoloba Nakai944
L. sericea Benth1811
L. sieboldi Miq944
L. stipulacea Maxim2011
L. tomentosa Siebold2011
L. variegata Cambess1811
Lotus angustissimus L127
L. corniculatus L12c45
L. corniculatus L var. alpestris Lamotte247
L. corniculatus L. japonicus Regel6c44
L. creticus L287
L. cytisoides14
L. filicaulis Dur12
L. hispidus Desf24
L. ornithopodiodes L14
L. requieni Mauri
L. siliquosus L
L. tetragonolobus L
L. uliginosus Schkuhr
Lupinus albicoccineus4879
L. albus L5079
ψ 4072
L. angustifolius L2085
L. barkeri Lindl50c5
L. densiflorus Benth4879
L. douglasii Agar
L. elegans H. B. and K. T. H.
L. hartwegii Lindl48-5079
L. hirsutus L. var. micranthus Boiss5079
L. luteus Lψ2378
L. micranthus Dougl4879
L. mutabilis Sweet2485
L. nanus Dougl
L. ornatus Dougl
L. pilosus L. [Murr.]42
L. polyphyllus Lindl2411
L. pubescens Benth48(79)
L. subcarnosus Hook36(79)
L. succulentus48(79)
L. varius L48(5)
L. venustus Vilm48(79)
Medicago apiculata Willd16(29,28)
M. arabica All16c(28)
M. arborea L32(29, 28)
M. carstiensis Wulf16(23)
M. ciliaris Krock16d(29, 28)
M. coronata Desr16(23)
M. denticulata Willd(29,28)
M. disciformis DC(29,28)
M. dzawakhetica Bordz(11)
M. echinus CD16c(25,28)
M. falcata L16(78)
32d29, 28
M. gerardi Waldst. and Kit32
M. glutinosa M. Bieb32(11)
M. helix Willd16(25,28)
M. hemicycla Grossh32(11)
M. hispida14c(23)
M. intertexta Mill16c
M. laciniata Mill16d29, 28
M. lappacea Desr16
M. littoralis Rhode16c
M. lupulina L8c78
16d25, 28
M. lupulina typica Urban326
M. maculata Willd1625, 28
M. marina L1629, 28
M. media Pers32, 3523
M. minima L16c25, 28
M. murex Willd16d29, 28
M. muricata (L.) All1623
M. nigra Krock1629, 28
M. obscura Retz16, 17, or 1823
M. olivaeformis Guss1629, 28
M. obicularis All16d25, 28
M. ovalis Urban (syn. Trigonella ovalis Boiss.)326
M. pentacycla DC1629, 28
M. platycarpa (L.) Trautv878
M. radiata L. (syn. Trigonella radiata Boiss.)166
M. rigidula DC1625, 28
M. rigidula (L.) Desr1423
M. rotata Boiss.1623
M. rugosa Desr3223
M. ruthenica Trautv1623
M. sativa L16d78
M. scutellata Mill32c23, 28
M. soleirolii Duby1623
M. sphaerocarpa Bertol1625, 28
M. tenoreana Ser
M. tornata Mill
M. tribuloides Desr29, 28
M. trunculata Gaertn25, 28
M. tuberculata Willd16d29, 28
M. turbinata Willd16c
Melilotus alba Med. (Dest .(?)]8d16g19
M. dentata Pers.166
M. indica All16c23
M. italica (L.) Lam.168
M. melilotus indica A. and G. (syn. M. parviflora Desf.)6
M. messanensis All8
M. neapolitana Ten. (syn. M. gracilis DC)6
M. officinalis810
M. segetalis Ser168
M. speciosa Dur9
M. suaveolens Ldb6
M. sulcata Desf16d23
M. taurica Ser166
M. wolgica Poir16c6
Millettia japonica A. Gray844
Mimosa pudica L2444
Onobrychis crista-galli Lam71414
O. viciaefolia Scop1112
Ononis alopecuroides L32(6)
O. biflora Desf32
O. fruticosa L32
O. hircina Jacq32 (30)
O. natrix L32
O. ornithopodiodes L32
O. reclinata L64
O. rotundifolia L32
O. spinosa L32 (30)
O. viscosa32
Ornithopus sativus Brot816(44)
Oxpiropis halleri Bunge16(4)
O. rishiriensis Matsumψ5366
O. uralensis Pall. [DC.]16(4)
O. vaginata Fisch164
Prachyrhizus angulatus1165
Parochetus communis16(12)
Phaseolus aconitifolius Jacq227
P. acutifolius A. Gray2241
P. angularis Willd. [((Willd.) W. F. Wight]
P. aureus Roxb
P. capensis Thunb227
P. chrysanthos Sav.1156
P. lunatus1144
P. multiflorus Willd1246
P. mungo L2241
P. nigerrimus Juss227
P. radiatus L1143
P. trilobus2241
P. vulgaris L1144
Piptanthus nepalensis Sweet185
Pisum arvense L7c85
P. elatius Bieb7
P. fulvum Sibth. [Sibth. and Sm.]
P. humile Boiss. [Boiss. and Noé]
P. jomardi Schrank
P. sativum L7s14t3
Psoralea bituminosa L1048
ψ 20c47
P. glandulosa Lψ 2047
P. macrostachya2048
P. palaestina Lψ 2047
Rhynchosia phaseoloides DC227
Robinia boyntonii Ashe1583
R. fertilis Ashe1083
R. hartwigii Koehne1083
R. hispida L1583
R. kelseyi Hutchins1083
R. luxurians (Dieck) Schneid1083
R. pseudoacacia L1047
R. viscosa Vent1083
Securigera coronilla DC127
Sesbania aculeata Pers1644
Soja hispida Mönch4041
S. max, Illini variety204080
Sophora angustifolium Sieb. and Zucc944
S. chinensis G. [G. Don]285
S. davidii Kon165
Sophora flavescens Ait185
S. japonica L285
S. moorcroftiana Benth1649
Spartinum junceum L48 (48-52)5
Swansonia galegifolia R. Br. var. albiflora Lindl1611
Tephrosia hookeriana Wit. and A1644
Thermopsis alterniflora Regel [Regel and Schmalh.]185
T. montana Nutt.971
Trifolium albopurpureum T. and G1682
T. alexandrinum1682
T. alpestre L81
T. ambiguum M. B1642
T. augustifolium L1442
T. arvense L71
T. badium71
T. campestre71
T. ciliolatum Benth. (T. ciliatum Nutt.)1682
T. dichotomum H. and A3282
T. filiorme L1442
T. fragiferum L81
T. fulcatum1682
T. glomeratum71
T. hybridum L8c1
T. incarnatum L81
T. lappaceum L81
T. lupinaster L4842
T. maritimum Huds1642
T. medium Lψ 48-491
ψ 8042
T. microcephalum Pursh1682
T. minus141
T. montanum L9 (?)1
T. obtusiflorum Hook1682
T. ochroleucum81
T. pannonicum Jacqψ 48-491
ψ 13042
T. parviflorum Ehrh1642
T. pratense L7c1
ψ 1252
T. procumbens L1442
T. reflexum L1682
T. repens Lψ 1252
(24) 2861
T. resupinatum L81
T. rubens L16
T. scabrum L16
T. spadiceum L14
T. squarrosum L14
T. thalii81
T. tumens Stev1642
T. variegatum Nutt1682
T. wormskoeldii Lehm48(?)82
Trigonella balansae Boiss. {Boiss. and Reut.}1676
T. calliceras Fisch76
T. coerulea (L.) Ser1623
T. corniculata L1676
T. cretica (L.) Desr16c23
T. foenumgraecum L16c23
T. glomerata Hort. (syn. Medicago brachycarpa Fisch.)166
T. melilotus coerulea A and G (syn. Melilotus coerulea Desf.)
T. monspeliaca L.
T. polycerata L286
T. striata L. (syn. T. cancellata Desf.)165
Ulex europaeus L.965
U. nanus Forst465
U. parviflorus Pourr965
Vicia alpestris Steph. (Stev.)2834
V. amoena Fisch.1278
V. amphicarpa L51076
V. angustifolia L612d75
V. atropurpurea Desf775
V. aurantia Boiss1434
V. bithynica L14d75
V. calcarata Desf.1434
V. cracea L.6c12c67
V. dasycarpa Ten714c75
V. disperma DC714d75
V. dumetorum1434
V. erviformis Boiss1434
V. ervilia Willd714d75
V. faba L6j50
V. gracilis Lois778
V. grandiflora Scop778
V. hirsuta S. F. Gray7c14d75
V. hybrida L612c75
V. hyrcanica Fisch. and Mey1234
V. lathyroides L.1234
V. lutea L778
V. macrocarpa678
V. monantha Desf778
V. musquinez Bosc1434
V. narbonensis L778
V. orobus DC678
V. pannonica Crantz612c75
V. peregrina L778
V. picta Fisch. and Mey778
V. pisiformis1234
V. pseudocracea Bertol775
V. pseudoorobus1268
V. pyrenaica Pourr1434
Vicia sativa L6d68
V. sepium L714d75
V. serratifolia Jac778
V. sicula Guss1434
V. silvatica L778
V. tenuifolia Roth1278
V. tetrasperma Moench7c78
V. unijuga A. Br678
V. villosa Roth778
V. catjang (Burm.) Walp2241
V. glabra Savi227
V. owahuensis Vog227
V. sesquipedalis A.I. Pierters {F. Ageaoili}1244
V. sinensis Endl1244
V. unguiculata (L.) Walp2241
V. vexilata Benth227
Wistaria brachybotrys Sieb. and Zucc8c40
W. floribunda DC8d40
W. frutescens (L.) Poir864
W. macrostachya Nutt864
W. multijuga Van Houtte (W. chinensis var. multiflora Hook.)484
W. sinensis Sweet864
W. venusta Rehder and Wilson864
*Names are given as in the articles cited except for obvious misspelling. Where the wrong authority for a name is given it is followed by the correct authority in brackets.
**Letters following numbers denote number of times verified by other authors. c=1; d=2; e=3; f=4; g=5; h=6; j=8; k=9; s=17; t=18.
*** Italic numbers in parentheses refer to References for Chromosome Numbers. In the case of more than 1 determination, the earliest author giving the established number is listed.
ψ Approximately
§ Apparently name is wrong.
   (1)   Bleier, H. 1925. CHROMOSOMENSTUDIEN BEI DER GATTUNG TRIFOLIUM. Jahrb. Wiss. Bot., 64: [604]-636, illus.
   (2)  Breslavetz, L., Medwedewa, G., and Magitt, M. 1934. ZYTOLOGISCHE UNTERSUCHUNGEN DER BASTPFLANZEN (APOCYNUM, BOEHMERIA, HIBISCUS, ABUTILON UND CROTALARIA). Pflanzenzüchtung 19: [229]}-234, illus.
   (3)  Cannon, W. A. 1903. STUDIES IN PLANT HYBRIDS: THE SPERMATOGENESIS OF HYBRID peas. Bull. Torrey Bot. Club 30: 519-543, illus.
   (4)  Chekhov, V. P. (Tschechow, W.). 1930. KARYOLOGISCH-SYSTEMATISCHE UNTERSUCHUNG DES TRIBUS GALEGEAE, FAM. LEGUMINOSAB (VORLAUFIGE MITTEILUNG). Planta, Arch. Wiss. Bot. 9: [673]-680, illus.
   (5)  ―1931. KARYOLOGISCH SYSTEMATISCHE UNTERSUCHUNG DBR TRIBUS SOPHOREAE, PODALARIBAE UND GENISTEAE. Izv. Tomsk. Otdel. Gosud Ruesk. Bot. Obshdr. (Mitt. Tomsk. Abo. Russ.) Bot. Gesell. 3: 121-131, illus. [In Russian. German summary.]
   (6)  ―1932. KARYO-SYSTEMATICAL ANALYSIS OF THE TRIBE TRIFOLIEAE D. C. (FAM. LEGUMINOSAE). Trudy Prikl. Bot., Genetike i Selek. (Bull. Appl. Bot., Genetics, and Plant Breeding) (2) 1: [119}-146, illus. [In Russian. English summary, pp. 144-146.]
   (7)  and Kartashova, N. N. 1932. KARYO-SYSTEMATISCHE UNTERSUCHUNG DER TRIBUS LOTEAE BENTH. UND PHASEOLEAE BRONN. FAM. LEGUMINOSAE Juss. Trudy Tomsk. Gosud. Univ. (Bull. Tomsk. State Univ.) 85: 1-22, illus. {In Russian. German summary.]
   (8)  Clarke, A. E. 1932. SOME CYTOLOGICAL AND GENETICAL STUDIES IN THE GENUS MELILorus. 6th internatl. Cong. Genetics Proc. 2: pp 20-21.
   (9)  Clarke, A. E. 1934. THE NUMBER AND MORPHOLOGY OF THE CHROMOSOMES IN THE GENUS MELILOTUS. Calif. Univ. Pubs., Bot. 17: 435-443, illus.
  (10)  Cooper, D. C 1933. MACROSPOROGENESIS AND EMBRYOLOGY OF MELILOTUS. Bot,; Gaz. 95: 143-155, illus.
  (11)  ―1936. CHROMOSOME NUMBERS IN THE LEGUMINOSAE. Amer. Jour. Bot. 23: 231-233, illus.
  (13)  ―1930. NUOVI REPERTI SULLA CARIOLOGIA DI ALCUNE LEGUMINOSE. Nuovo Giorn. Bot. Ital. 37: 679-680.
  (14)  ―1931. OSSERVAZIONI CARIOLOGICHE SU ALCUNE LEGUMINOSE. Nuovo Giorn. Bot. Ital. 38: 230.
  (16)  Datta, R. M. 1933. THE POLLEN DEVELOPMENT IN CASSIA TORA LINN. (Abstract.) Indian Sci. Cong. Proc. 20: 313.
  (17)  Dixit, PD. 1932. STUDIES IN INDIAN PULSES. A note on the cytology of "kabuli" and “desi” gram. types. Indian Jour. Agr. Sci. 2: 385-390, illus.
  (18)  Dombrowsky-Sludsky, L. 1927. LA CYNÈSE SOMATIQUE DE CICER ARIETINUM L. Jour. Soc. Bot. Russie 12: [163]}-172, illus. [In Russian. Summary in French, p. 171-172.)
  (19)  Elders, A. T. 1926. SOME POLLINATION AND CYTOLOGICAL STUDIES OF SWEET CLOVER. Sci. Agr. 6: 360-365, illus.
  (20)  Erith, A. G. 1924. WHITE CLOVER (TRIFOLIUM REPENS L.); & monograph. 150 pp., illus. London.
  (21)  Fisk, E. L. 1931. THE CHROMOSOMES OF LATHYRUS TUBEROSUS. Natl. Acad. Sci. Proc. 17: 511-513, illus.
  (22)  Fraser, H. C. L., and Snell, J. 1911. THE VEGETATIVE DIVISIONS IN vicIA FABA. Ann. Bot. [London] 25:[845]-855, illus.
  (23)  Fryer, J. R. 1930. CYTOLOGICAL STUDIES IN MEDICAGO, MELILOTUS AND TRIGONELLA. Canad. Jour. Research 3: 3-50, illus.
  (24)  Fukuda, Y. 1933. CYTO-GENETICAL STUDIES ON THE WILD AND CULTIVATED MANCHURIAN SOY BEANS (GLYCINE L.). Japan. Jour. Bot. 6: [489] 506, illus.
  (25)  Ghimpu, V. 1928. CONTRIBUTION À L’ÉTUDE CARYOLOGIQUE DU GENRE MEDICAGO. Compo. Rend. Acad. Sci. [Paris], 187: 245-247, illus.
  (26)  ―1929. CONTRIBUTION À L’ÉTUDE CHROMOSOMIQUES DES ACACIA. Compt. Rend. Acad. Sci. [Paris] 188: 1429-1431.
  (29)  ―[1932?] RECHERCHES CHROMOSOMIQUES SUR LES LUZERNES, VIGNES, CHÈNES ET oRGES. 14th Cong. Internatl Agr. Bucarest, 1929, v. 4, pp. 557-564, illus.
  (30)  ―1933. LES NOMBRES CHROMOSOMIQUES DE QUELQUES ANGIOSPERMS CULTivées. Compt. Rend. Soe. Biol. [Paris] 112: 1115-1117.
  (31)  Ghose, S. L., and Alagh, R. 1933. MICRO- AND MEGA-SPOROGENESIS IN CASSIA PURPUREA ROXB. (Abstract) Indian Sci. Cong. Proc. 20: 315-316.
  (34)  ―1931. NUKLEOLEN UND CHROMOSOMEN IN DER GATTUNG Vicia. Planta, Arch. Wiss. Bot. 15: [495}-505, illus.
  (35)  Hunter, H., and Leake, H. M. 1933. RECENT ADVANCES IN AGRICULTURAL PLANT BREEDING. 361 pp., illus. London.
  (36)  Hus, H. T. A. 1904. SPINDLE FORMATION IN THE POLLEN-MOTHER-CELLS OF CASSIA TOMENTOSA L. Calif. Acad. Sci. Proc. Bot. (3) 2: 329-354, illus.
  (37)  Husted, L. 1931. CHROMOSOME NUMBER IN SPECIES OF PEANUT, ARACHIS. Amer. Nat. 65: 476-477, illus.
  (39)  Ishikawa, M. 1916. A LIST OF THE NUMBER OF cHROMOSOMES. Bot. Mag. [Tokyo] 30:404-448, illus.
  (40)  Jimbo, T. 1927. THE CHROMOSOMES OF WISTARIA. Bot. Mag. [Tokyo] 41: 487-489, illus. [In Japanese. Title also in English.]
  (41)  Karpechenko, G. D 1925. ON THE CHROMOSOMES OF THE PHASEOLINAE. Trudy Prikl. Bot. i Selek. (Bull. Appl. Bot. and Plant Breeding) 14: [143}-148, illus. [In Russian. English summary, pp. [147]-148.]
  (42)  ―1925. KARYLOGOISCHE STUDIEN UBER DIE GATTUNG TRIFOLIUM L. Trudy Prikl. Bot. i Selek. (Bull. Appl. Bot. and Plant Breeding) 14: [271]-279, illus. [In Russian. Résumé in German, p. 279.]
  (44)  Kawakami, J. 1930. CHROMOSOME NUMBERS IN LEGUMINOSAE. Bot. Mag. [Tokyo] 44: [319]-328, illus. [In Japanese. Title also in English.]
  (45)  Kihara, H., Yamamoto, Y., and Hosono. 1931. A LIST OF CHROMOSOME, NUMBERS OF PLANTS CULTIVATED IN JAPAN. Pp. 195-330. Tokyo.
  (46)  Kleinmann, A. 1923. UEBER KERN- UND ZELLTEILUNGEN IM caMBIUM. Bot. Arch. 4:[113}-147, illus.
  (47)  Kreuter, E. 1929. CHROMOSOMENSTUDIEN BEI DEN GALEGEEN (VORLÄUFIGE MITTEILUNG). Ber. Deut. Bot. Gesell., 47: 99-101.
  (48)  ―1930. BEITRAG ZU KARYLOGISCH-SYSTEMATISCHEN STUDIEN AN GALEGEEN. Planta, Arch. Wiss. Bot. 11: 1-44, illus.
  (51)  Lutkov, A. N. 1930. INTERSPECIFIC HYBRIDS OF PISUM HUMILE BOISS. X PISUM SATIVUM L. U.S.S. R. Cong. Genetics Plant and Anim. Breeding, Proc. v. 2, pp. 353-367, illus. [In Russian. English summary, pp. 366-367.]
  (52)  Martin, J. N. 1914. COMPARATIVE MORPHOLOGY OF SOME LEGUMINOSAE. Bot. Gaz. 58:154-167, illus.
  (53)  Melderis, A., and Viksne, A. 1931. NOTES ON THE GENUS LATHYRUS. Acta Horti Bot. Univ. Latviensis, Riga, 6, (2-3:) 90-94.
  (54)  Milovidov, P. F. 1926. ÜBER EINIGE NEUE BEOBACHTUNGEN AN DEN LUPINENKNOLLCHEN, Centbl. Bakt. [ete.] 68: 333-345, illus.
  (57)  Nemec, B. 1904. UEBER DIE EINWIRKUNG DES CHLORALHYDRATS AUF DIE KERN UND ZELLTHEILUNG. Jahrb. Wiss. Bot. 39: [645]-730, illus.
  (59)  Newman, I. V. 1933. STUDIES IN THE AUSTRALIAN ACACIAS. I. GENERAL INTRODUCTION. Jour. Linn. Soc. Bot. 49: 133-1438.
  (61)  Oppenheimer, H. 1934. KULTUR-PFLANZEN. Tabulae Biologicae Periodicae, Bd. 3, no. 4, pp. 349-394.
  (62)  Ranagaswami Ayyangar, G. N., and Krishnaswamy, N. 1933. A NOTE ON THE CHROMOSOME NUMBERS IN CLUSTER BEANS (CYAMOPSIS PSORALOIDES DC.). Indian Jour. Agr. Sci. 3: 934-935.
  (63)  Rau, N.S. 1929. FURTHER CONTRIBUTIONS TO THE CYTOLOGY OF SOME CROP-PLANTS OF SOUTH INDIA. Jour. Indian Bot. Soc. 8: 201-206, illus.
  (64)  Roscoe, M. V. 1927. CYTOLOGICAL STUDIES IN THE GENUS WISTERIA. Bot. Gaz. 84: 171-186, illus.
  (66)  Sakai, K. 1934. STUDIES ON THE CHROMOSOME NUMBER IN ALPINE-PLANTS. I. Japan Jour. Genetics 9: 226-230, illus. [In Japansee. English résumé, pp. 229-230.]
  (67)  Sakamura, T. 1914. STUDIEN UBER DIE KERNTEILUNG BEI VICIA cRaccA L. Bot. Mag. [Tokyo] 28: [131]}-147, illus.
  (69)  Senn, H.A. 1936. A CYTOGENETIC STUDY OF THE GENUS LaTHYRUS. Va. Acad. Sci. Proce. 1935-36: 32.
  (70)  Sethi, M. L. 1930. MICROSPOROGENESIS IN CASSIA DIDYMOBOTRYA. Jour. Indian Bot. Soc. 9: 126-139, illus.
  (71)  Simonet, M. 1932. NUMÉRATIONS CHROMOSOMIQUES DANS LES GENRES BAPTISIA, THERMOPSIS, ET LATHYRUS. Compt. Rend. Acad. Sci. [Paris] 195:738-740.
  (72)  Smet, E. DE 1914. CHROMOSOMES, PROCHROMOSOMES ET NUCLÉOLE DANS QUELQUES dicotylées. Cellule 29 [335]-377, illus.
  (73)  Strasburger, E. 1905. TYPISCHE UND ALLOTYPISCHE KERNTEILUNG. ERGEBNISSE UND By ERORTERUNGEN. Jahrb. Wiss. Bot. 42: 1-71, illus.
  (74)  Sugiura, T. 1931. A LIST OF CHROMOSOME NUMBERS IN ANGIOSPERMOUS PLANTS. Bot. Mag. [Tokyo] 45: 353-355.
  (75)  Sveshnikova, I. N. 1927. KARYLOGICAL STUDIES ON vicia. Trudy Prikl. Bot., Genetike i Selek. (Bull. Appl. Bot., Genetics, and Plant Breeding) 17 (3): [37]}+72, illus. [In Russian. English summary, pp. [63]-72.]
  (76)  ―(Sweschnikowa, I.) 1928. DIE GENESE DES KERNS IM GENUS viciA. Verhandel 5th Internati. Kong. Vererbungswiss, Berlin, Bd. 2, pp. [1415}-1421, illus.
  (77)  Tischler, G., 1921-22. ALLGEMEINE PFLANZENKARYOLOGIN. Bd. 2, illus. Berlin.
  (78)  ―1927. PFLANZLICHE CHROMOSOMEN-ZAHLEN. In Tabulae Biologicae, Bd. 4, Pa: 1-83.
  (79)  Tuschnijakowa, M.; 1935. UBER DIE CHROMOSOMEN EINIGER LUPINUS-ARTEN. züchter 7: 169-174.
  (80)  Veatch, C. 1934. CHROMOSOMES OF THE SOY BEAN. Bot. Gaz. 96: 189, illus.
  (81)  Vilmorin, R. de, and Simonet, M. 1927. NOMBRE DES CHROMOSOMES DANS LES GENRES LOBELIA, LINUM ET CHEZ QUELQUES AUTRES ESPECES VÉGÉTALES. Compt. Rend. Soc. Biol. [Paris] 96: 166-168, illus.
  (82)  Wexelsen, H. 1928. CHROMOSOME NUMBERS AND MORPHOLOGY IN TRIFOLIUM. Calif. Univ. Pubs. Agr. Sci. 2: 355-376, illus.
  (83)  Whitaker, T. W. 1934. A KARYO-SYSTEMATIC STUDY OF ROBINIA. Jour. Arnold Arboretum 15: 353-357, illus.
  (86)  Yamaha, G., and Sinoto, Y. 1925. ON THE BEHAVIOUR OF THE NUCLEOLUS IN THE SOMATIC MITOSIS OF HIGHER PLANTS, WITH MICROCHEMICAL NOTES. Bot. Mag. [Tokyo] 39: [205]-219, illus.