Miscellaneous Forage and Cover Crop Legumes

MISCELLANEOUS FORAGE AND COVER CROP LEGUMES

ROLAND McKEE, Senior Agronomist, A. J. PIETERS, Principal Agronomist, Division of Forage Crops and Dis- eases, Bureau of Plant Industry

X HE 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 distinguishing 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 fertilit^^ 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 tliey deserve much wider recognition and could well be used much more extensively. Improvement work with legumes other tha,n 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 iuvestigation, which will be discussed later. BREEDING WORK AND POSSIBILITIES WHILE the legumes considered in this article occupy a place secondar}^ 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 crotalarías 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 999 1000 YEARBOOK, 193 7 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 w^ill prove precisely suited for one or more such situations. ASTRAGALUS

{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 w^hich to expect a legume of agricultural value. BEGGAR WEED

{Meihomia 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 {Meihomia 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 har- vested 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. puhens (T. and G.) Vail, that has been grown in experimental plantings has growth habits that give promise of usefulness.

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 un- doubtedly be found by intensive study and developed by systematic breeding. MISCELLANEOUS FORAGE LEGUMES 1001 IJONAVIST (DolUhos IMiiU I..) Tlio hoiiavisfc, wliirli in hahits, ciildiral requirements, and uses is miK'li like tlic cowpca, has been used in Africa and southern Asia from aiiciont times, ami imiiiy varieties aro known to exist. Nowhere has improvement of the crop been attempted, although improvement with reference to nernatode and wilt resistance and other characteristics no doubt could be clbH-lcd ((ij;. 1).

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

JíUK-CLOVEK {Mi'dicaff) s¡>[>.) Many si)ecios of AJcdlc(Ujn that never are grown as cultivated crops enter into use for pasturage and arc recognized as having very great value for this puri)ose. líowevcr, tiicse liave received no attention from the plant breeder. Otlier species that are gro^vn for soil improvement and occasioiuiUy used for liay or seed have received some attention by expérimentera. In a few instances selections of distinct forms liavc been made ])y practical growers who have increased their supply of seed and distributed it locally. Natural selection resxdting from regioTud climatic differences, however, is responsible for much of the improvement in the mcdicagos. In the case of spotted bur-clover (Medicago arabica (L.) All.) at least two new forms have a])peared in the South in recent years. One of these, Aiaiiganesc bur-clo\'cr, was selected and named by A. 1002 YEARBOOK, 1937

Lee Andrews on liis place at Lafayette in eastern Alabama. The other, Early Southern bur-clover, was selected and named by A. F. Kuff at Kock Hill, S. C. Both these varieties are earlier maturing than the commercial spotted bur-clover from wliich 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

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Figure 2.—Coiled seed pods of spiny and spineless forms of spoiled bur-clover (Medicago arabica). In olher sj)ecies varialion in length of spines is not uncommon. 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). California bur-clover {Medicago hispida Gaertn.) is very variable and responds to improvement by selection. Wliere 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 Gar- den at Chico, Calif., but supplies of seed have not been mahitained and none are available commercially. The Texas Agricultural Experi- ment 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 qiiite 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 MISCELLANEOUS 1 ORAGE LEGUMES 1003 prcicroîKîc to the coiumvAvXiu i'ornis. Bluck medic (Medicago lupulina L.) is quite vuriablo iirul eould. be improved readily by selection. No work has been rei)orted OJL the ijuproveiiient of this plant.

(jlJCKI'EV {Cirv.r ariv.linatii L.) Throughout the westeiri. United Slates attempts have been made to grow the chickpeii, which is native to western Asia, but nowhere lias commercial production been suci^essfuUy established. So far as jvnown, no atteiript has Ix^on made to develop superior varieties adiipted to the United States, although it is known that the chickpea is variable and tJius presumably couhl be improved by selection.

(^\o\^ TEA {Vigna sinctisis (Tomer) Savi) A very larger nuniber of cowpea, varieties are recognized. For the most part those have been developed through several hundred years by natural hybridization and iucidental selection rather than by any planned improvement program. lîïtroduétions into the United States are recorded in the seventeenth ccTitury, but it is within the last century that specific reference is made to named varieties. In the United States in recent years considenible work hns l)een done by experiment stations in the way of bringing existing varieties together for comparative test- ing. This has resulted in a more exterisive use of superior varieties and an ehminalion of inferior ones. While selection of superior plants has thus been the priucipal mearis 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 lias 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, character- ized by its resistance to wilt and nematodes, which seriously damage most varieties. In the pj'ocess of continuous growing and natural selection through long periods of tbrui, 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 lïiiich more resistant to bean rust than re- (îcntly introdu(îed varieties, and resistance to wilt and nematode in the iron, Bra,bha.m, 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 froiïi its pa.rents, the Brabham and Groit. The origin of most of the connnercial varieties of cowpeas is unknown. The Whippoorwill has becui grown under that name since 1840, and the Iron has been known in South Carolina since 1888, but its origin is uncertain, and Jio information is available as to where tlie New Era may have come froiu. From a study of the characters of the Brabham it lias been concluded that it is a hybrid between Iron and Whippoorwill, and the Groit is regarded as a hybrid of New Era and Wliippoorwill. Improved selections from Brabham_ and Iron have been made in California aTul are being grown commercially. Ilccently a variety of unknown oj'igin was introduced into Florida 1004; YEARBOOK, 1937 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 (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

ligure 3.—(Jivtularia spectabilis, F. (J. J8096 (a), and the cuiiimerciul »train from which it was selected (6). 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 spedabUis 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 Agri- culture 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. spedabilis has been developed that ripens its seed quite unifonnly 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 MISCELLANEOUS FORAGE LEGUMES 1005 of C. intermedia and C. striata, but additional improvement is needed }3cfore these can be called superior. 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.

FENUGREEK {TrigoncUa focnumgraecum 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 IJnited States fenugreek has succeeded only in California, and there it is grow^n only occasionally. At the California Agricultural Experi- ment 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. FIELD PEA (Fisurn 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 iisage 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 growler, while the field husbandman has merely grown the varieties apparently best suited to his conditions. In more recent thnes 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 suscepti- bility 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. 1006 YEARBOOK, 1937

GRASS PEA (Lathyrus sativas 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 X. sativus or related species in this way. GUAR (Cyarnopsis 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.

KmNEYVETCH {Antliyllis 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 Iddneyvetch has been attempted, and strains Tystofte No. 8 and Tystofte No. 28 when compared with commercial kidneyvetch were found to be more productive.

KUDZU-BEAN (Pueraria thunhergiann (Sieb, and Zuce.) Benlh.) 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 (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.) Ben th., has recently been introduced to agriculture. 1 Italic numbers íD parentheses refer to Literature Cited, p. 1018. MISCELLANEOUS FORAGE LEGUMES 1007 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 lowêr South, and gradually wôrked north. The character that made this spread possible wâs imdoubtedly the 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 work b}^ the late S. H. Essary, of the Tennessee Agricultural Experi- ment Station. He began his study of individual plants in 1912 and found a great variation, especially in the habit of grow^th. Among the variants was one having an erect habit and great productive capacity, which was segregated and put into experimental plantings in 1921. Tliis was introduced by the Tennessee station as Tennessee 70 and is one of the leading late-maturing varieties. Another variety with larger leaflets and ranker growth than usual wâs found b}^^ J. B. S. Norton, an explorer of the Department of Agriculture, growing wild near the city of Kobe, Japan, and W'as 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, wâs collected by Dorsett and Morse, of the Department, in eJapan in 1929 and is the earliest maturing form of Lespedeza striata. In habit it is nearly as erect as Tennessee 70. 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. T\vo 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 tlie 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 grow^th and yielding ability are like that of standard Korean, and it differs only in earliness. The late form, which is matting in grow^th habit, is still under observation. Here again the possibilities of selection have not been fully explored, and in the future better vaiieties, or varieties better suited to certain conditions, may be selected. The group to wiiich Lespedeza sericea, the perennial lespedeza, belongs consists of several species difl'ering from one another in 1008 YEARBOOK, 1937

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. júncea, 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

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. 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. 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 Turcz. 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 (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 MISCELLANEOUS FORAGE LEGUMES 1009 States none is recognized as having special value. Seed of Lotiis cormculatus 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 cyanophonc glucoside content have been reported. The elimination

Fiffire 5.—Lespedeza seruitt, showing vari;aii)ii in SWA: and liabil of growlli. I'laiUa in each row are from seed of a single mother plant. of this glucoside is desirable, since in the course of digestion it is changed to hydrocyanic acid, which is injurious to animals.

LUPINE (Lupinas 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 economi- cally 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 Kepublics, where in recent years attempts have been made to produce lupines devoid of alkaloid poisons. Both German and Russian workers report having developed strains witli little or no alkaloid, wliich they call "sweet lupines." The object 1010 YEARBOOK, 1937 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 w^ild species. These offer possibilities but as yet are too little known to justify a statement regarding results. PiGEONPEA {Cajanus indicas Spreng.) In many tropical coimtries 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, wiiere 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 Experi- ment 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. SAINÍ'OIN {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. Com- mercial sainfoin is quite variable and includes both biennial and pcT- ennial 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. SERRADELLA (Ornithopus sativas lire t.) 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 ver}^ limited. MISCELLANEOUS FORAGE LEGUMES 1011

SESBANIA (Sesbania macrocarpa Muhl.) The seed of sesbania as found in the commercial trade is gathered from wild and volunteer plants and represents the species as it is found growing naturally. The plant grows in wet or moist soils in the Southern States. No attemx)ts at improvement have been made.

SULLA {Hedysarum coronarium L.) Attempts to grow sulla in the united States have indicated that while 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 arc known to exist, few attempts at varietal improvement seem ever to have been undertaken. In New South Wales a special strain was selected for many years and tested on the experimental farms, but for other countries no work is reported. VELVETBEAN (Stizolobium spp.) The Florida velvetbean {Stizolobium deeringiamtm Bort) is one of the 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 Speclded. This variety later became known as Georgia and is the earliest maturing commercial variety now grown. In 1908 E. \Y. Miller, of Broxton, Ga., selected an early-maturing variety that was grown as Clark's velvetbean, but this proved to be so hke 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 earl}^ maturing as the Georgia but was sufhciently good to become commercialized 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 Mississipjii 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 ^^ariety seems to have been obtained that reproduces true to color. 1012 YEARBOOK, 1937 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 fnr north as central Georgia.

ligure 0.—Bush velvclbcaii. The velvetbean offers opportunity for much further improvement, and the importance of the crop suggests that such work could be done with profit. VETCH

{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. atropurpúrea Desf., V. calcarata Desf., V. monantha Desf., V. dasycarpa Ten., and V. angustijolia 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 imiform 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 MISCELLANEOUS FORAGE LEGUMES 1013 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 improvemtrnt 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 l)een 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 wdll 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 of 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 commercialized. 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 T'. villosa imported from central Europe under the name hairy vetch is smooth vetch and j-esembles tlie smooth vetch variety grown in Oregon. Of the common vet(;h. {V. saMva) selections that have been developed, a wliite-ilowered variety, F. C. 02830, that is somewhat superior in vigor and winter hardiness to commercial Oregon common vetch and was selected by Sohoth in 1915, is perhaps outstanding. The white-ilowxr character offers a ready means of identification in the field and will enable the grower to keep his seed pure. In the Xetherlands, 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. GENETIC STUDIES IN MISCELLANEOUS LEGUMES'^ 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 of 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 (7^. muîigo L.) and the mung bean (P. aureus Roxb.) are 2 This section is wriííeu primarily for students or others professionalîv interested in breeding or genetics. 1014 YEARBOOK, 1937 usually self-pollinated, although the mode of anthesis in many cases would permit of cross-pollination. Unpublished observation of legume plantings in the United States indicates that Crotalaria, Viciay 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 i. 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-fer tile. No sign of hybridiz- ing has been observed, though the species have been grow^n 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 cowpca {Vigna sinensis (Torner) Savi), chickpea {Cicei- arietinum L.), adzuki bean (Phaseolus angularis (Willd.) Wight), bonavist (Dolichos lablab L.), horsebean (ViciaJaba L.), pigeonpea {Cajanus indiens Spreng.), lupine (Lupinus spp.), peanut (Arachis hypogaea L.), velvetbean {Stizolo- bium deeringianum Bort), and field pea (Pisum arvense L.).

CHICKPEA (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 w^ere 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 (IS) relating to color and number of flowers. In maldng artificial hybrids it was found that in order to prevent accidental crossing emasculation should be eft'ected the evening of the second day pre- ceding fertilization. The flowers open naturally on bright days be- t\yeen 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. MISCELLANEOUS FORAGE LEGUMES 1013

(4) 111 the absence of B the flower is white whether P is j)resent or absent. (5) Greenness in the standard is devek)j:)ed in the absence of the factor W. Greenness is therefore recessive to nongreen. (6) Singleness depends upon a factor S and is dominant to doubleness.

COWPEA (Figna sinensis (Torncr) 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 dcflnitely 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, (r. Dotting; converts Holstein spots into numerous small ones. II. Holstein type of seed-coat spotting. /. Eye with indefinite margiu. L. Longitudinal furrowing of the surface of the seed. A^. Presence of anthocyanin i:>igment factor. P. Purple seed coat. R. Red seed coat. (This is the general factor for color, the absence of which determines white seed coat, wliite iloNvers, and a'osence 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. Tlie eight factors By I\ N^ P, Ry TF, L\ and X, either singly or in combination, give rise to the ten distinguishable seed-coat colors, purple, black, dull black, blue, coiFce, maroon, buff, red, piiilc, and wiiite. The factors P, £?, Z), T, and F and probably S belong to a linked group. The factors Z), JT, and F restrict the dis- tribution of color pigments in the seed coat. Three independent eye factors 7, 77, and Ey 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 /?, there is a tendency for the seeds to be small, weak, or abortive. The flower color has been found to be correlated with tlie coloration in the seed coat, joints, i^eduncles, stipules, and petioles {16^ pp. 282-283). Experiments reported by Haigli and Lochrie {6) indicate a progressive variation with age of a simple Mendelian ratio in the cowpea. The results in the Fg cultures from successive days of flowering showed an orderly variation in the simple Mendelian 3:1 ratio, an 1016 YEARBOOK, 19S7 excess of récessives 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 (5), 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.

BONAVIST {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 deter- minate and indeterminate growth the segregating ratio in the Fa 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 cft'ect except in the presence of the other, when it converts white flower into purple, and brown seed into black, aud causes pigmentation of the nodal region.

LUPINE {Lupinus spp.) Burlingame (2) reports studies of Lujyinus 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 heterozygou s 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 allelomorpliic to that for dark-blue and not improbably constitute a system of multiple allelomorphs. Mutations are frequent, some are already known to he 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 (^, p. 447). Hallqvist (7) studied seven diflerent types of flower color and five types of seed color in Lupinus angustijolius L. His conclusions were as follows (7, j;. SUY 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, v/hich suggested that the homozygous nonhairy might be lethal. MISCELLANEOUS FORAGE LEGUMES 1017

ADZUKI BEAN {Phaseolus angularis (Willd.) Wight) Kakizaki (1^), 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 y a factor for black-spotting of seed- coats, and its recessive allelomorph s, respectively, and hardly any crossing over occurs between them. The / factor, which intensifies the purple color of the stem, and its recessive allelomorph i are also very closely linked with the B factor, which produces a blackish-brown color of the ripe pods, and 6, its recessive allelomorph, and this linkage is also very close, so that crossing over hardly ever occurs between them. The PS linkage group is independent of the I-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),

nORSEBE4N (Vicia Jaba L.) The horsobean 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 typo of which albino and normal are the homozygous types. Sirks (16) in a study of quantitative inheritance in Viciajaba presented evidence to indicate that quantitative factors do exist.

VELVETBEAN (Stizotobium spp.) Several species of Stizolohium 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 arc S, deeringianum Bort X S, pachylobium Piper and Tracy, iS. deeringianum X S. niveum, and S, deeringianum X S. hasj00, 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. 1018 YEARBOOK, 1937

PiGEONPEA {Cajanus indicas Spreng.) Inheritance studies with the pigeonpea carried on at the Hawaiian Agricultural Experiment Station have been reported by Krauss (Ï4j V' IS), According to his findings— * * * red flower standards are doTninant 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 glaV^rous; large, flat pods are dominant over small round pods, and large seeds over small seeds. Four and five seeded pods arc 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 differ- entiating 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 indiens 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 w^as 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 closel^^ linked with maroon color of the pod. LITERATURE CITED (1) BELLING, J. 1911. SECOND GENERATION OF THE CROSS BETWEEN VELVET AND LYON BEANS, ria. Agr. lOxpt. Sta. Rcpt. 1911: Ixxii-civ, illiia. (2) BURLINGAME, IJ. L. 1921. VARIATION AND HEREDITY IN LtiPiNiTS. Amer. Nat. 55: 427-448, illus. (3) DARLINGTON, D. C. 1929. VARIEGATION AND ALBINISM IN VICIA FAB A. Jour. Gorietics 21: [1611-168, illus. (4) DAVE, B. B. 1934. INHERITANCE OF CHARACTERS IN CAJANUS INDICUS. Indian Joiir. Agr. Sei. 4: 674-691, illus. (5) GREB, R. J. 1935, TWO GENETICALLY DIFFERENT ROOTLET TYPES IN THE LUPINE. JOUR. Heredity 26: 503-504, illus, (6) HAIGH, J. C, and LOCHRIE, J. V. 1929. INVESTIGATION OF A MENDELIAN RATIO IN VIGNA SINENSIS BY A CONSIDERATION OF THE PROGENY FROM SUCCESSIVE DAILY CROSSES. Ann. Bot. [London] 43: [783]-803. (7) HALLQVIST, C. 1921. THE INHERITANCE OF THE FLOWER COLOUR AND THE SEED COLOUR IN LUPiNUs ANGUSTiFOLius. Hereditas 2: [299]-363, illus. MISCELLANEOUS FORAGE LEGUMES 1019

(8) HARLAND, S. C. 1920. INHERITANCE IN DOLICHOS LABLAB., L., PART I. JoUl*. Geiictics 10: 219-226. (9) HOFMANN, F. W. 1926. HYBRID VIGOR IN COW PEAS. Jour. Heredity, 17: 209-211, illus. (10) HoAVARD, A., HOWARD, G. L. C, and KHAN, A. II. 1915. SOME VARIETIES OF INDIAN GRAM (CICER ARIETINUM L.). ludia 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. (KHESARl, 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 JIEAN. 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. Hawaü Agr. lOxpt. Sta. Bull. f)4, 46 pp., illus. (15) SlRKS, M. J. 1929. .GROWTH AND INHERITANCE OF LEAF DIMENSIONS IN THE BROAD- BEAN (VICIA FAB A L.). K. Akad. Wetensch. Amsterdam, Proc. Sec. Sei. 32: 1066-1084. (16) SPILLMAN, W. J., and SANDO, W. J. 1930. MENDELIAN FACTORS IN THE COWPEA (VIGNA SPECIES). Mich. Acad. Sei., Arts, Letters, and Papers 11: 249-283, illus. APPENDIX

TABLE ].—Chromosome numbers in legumes

Chroniosoiüe iiuiTibers ^ Kcft'niiico XaiTie 1 110. 2n

Acacia arabica Willd ±52, ±104 Hm), (m A, baileyana F. Mucll (r,9, 60) A. cyanophylla Lindl 26 (26, m) A. dealbata Link___ 26 {26, 28) A. dec'urrens Willd 26 (27,28) A. dermatophyUa F. JNJuüll. [Bcutk.] 13 26 (to) A. ebúrnea Willd dz52, ±104 (27,28) A. farnesiana AVilld ±52, ±104 (26,28) A. hórrida Willd ±52, ±104 (26,28) A. longifolia Willd _ 26 (27,28) A. lophantha 24 (U) A. nilotica ±52, ±104 (26) A. podalyriaefolia A. Cunn __ 20 (26,28) A. saligna Wendl._ 26 (27) A. scorpioides A. Chev., var. adstringens (Schum. and Thon.) A. Chcv. 52,104, 208 (27,28) A. scorpioides A. Chev., var. nilotica Bent h ±52, ±104 (28) A. scorpioides A. Chev., var. pubescens Benth ±52, ±104 (27,28) Aeschynomene indica L 20 (U) Amorpha californica Natt 10 (48) A.fruticosa L 20 (48) Do- 40 (48) A. microphylla Pursh 10 (48) Amphicarpa monoica (L.) Ell 10 (//) AnthyUis alpesfris Kit 12 (7) A. barba-jovis L 14 (7) A. gerrardi L_ __ 16 (7) Í 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. 2 Letters following numbers denote number of times verified by other authors. o=l; d=2; c=3; f=4; g=5; h=G; j=8; k=9; s=17; t=18. 3 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. 1020 YEARBOOK, 1937

TABLE 1.—Chromosome numbers in legumes—Continued

Chromosome numbers Xanie Reference no. 2n

A. maritima Sehweigg 12 (7) A. tetraphylla L 16 (7) A. vulneraria fi 12 (U) Arachis hypogaea L 20e 40e (44) A. hypogaea (Spanish and small Japtm peaimts). 10 05, pp. 338-342) A. namhyguare 40 {37) A. prostrata Benth. var. rasteiro ±40 im A. rasteiro Chevalier (?) . .. 40 iS8) Astragalus alopecurioides L US) A. altaicus Bunge (4) A. baeticus JJ 8 L (47) A. candidissimus Led 10 (4) A. echinus DC' ._ 64 (4) A. edulis Dur (48) A. exscapus B. transsilvanîcus A. and G. ■ A. transsilvanicus Biirth. (4) A.falcatus Lam 8 (47) A. galegiformis L . 8 (47) A. hamosus L 24 im Do 48 (4) A. hypoglottis L 16 (4) A. massiliensis Lam 16 (48) . A. membranaceus Fisch. [Bungo] . 16 (4) yi. mollis . m) A. monspessulanus L U7) A. secundus'DC • (66) A. sesameus L (47) Do 16 (4) A. sieversianus VnW 16 (4) A. siriicus L . _ . _. (44) Do . 16 (74) A. transsilvanicus 16 (4) A. vuipinus VVilld (47) Baptisia australis R. Br (71) B. australis R. Br. var, exalta Suoot (ó) B. sulphurea Engolm (7î) B. tinctoria R. Br (■5) Biserrula pelecinus L 8 (47) Cajanus indicus Spreng 11 (65) Calophaca wolgarica Fisch .. 8 (47) Canavalia ensiformis DC 11 (44) C glaàiata DC 22 (7) Caragana arborescens Lam _ _ _ 16c (7) C.frutescens DC 32 (4) Carmichaelia australis R. Br _ .. ... 1Ô (48) Cassia didymobotnja ... 11 28 (70) C.dimidiata . _.. 16 (74) C. fistula.. 77, pp. 541- 623. (7. leschenaultiana DC_ 24 (U) C. mi mosoides L ,16 (44) C. occidentalis L 13 (56) C. purpurea Roxb 10 (31) C sophera L 12 (44) C. tomentosa L 12c (36) C. tora L_. 13 (16) Cercis sitiquastruifi. (13) Cicer arietinum L_... (17) Do. 14e US) Do_ 16 (55) C. kabulium 16 (17) Clitoriaternatea L 16 (7) (Mutea arborescens L 16 (4) C.halevica Lam as) C. media Willd. (C, arborescens L. X C. orientalis Lam.).. (48) C. orientalis Lam (4S) Crotalaria alata Ham (44) C.anagyraides B.. B. K (44) C. arenada Benth (se) C. júncea^ 16 (2) C. obovata G. Don (32) C. retusa Jj 'Î6" (44) C. usaramoensis Back. [Baker]. (44) C. valetonii Back (44) * Approximately. MISCELLANEOUS FORAGE LEGUMES 1021

TART.E 1.—Chromosome numbers in legumes—Continued

Chromosome numbers lleference Name no.

CrjamoDsis vscraloides DC 14 (6S) Cytisus canariensis O. Kiintze.. 40 (fí) C. nigricans L (81) Do. 48 (5) C. purpureus 48 (73) C.scovarnis Link 24c (U) C. scovarius (U) C. sessUifolius L (â) JDesmodhim grandißorum (Walt.) DC. 11 01) D.perpesium DC.5 11 (U) JDolkhos bifloras (6S) D. lablab L ]]c (78) Do. 22 (41) Do 24 (6S) D. lubia Forsk 22 (7) D. înultiflorus 24 (ó8) Z). niloticus Del 22 (7) IJ. orwöiws Wall 22 (7) Dorcynium herbaceum VilL. 14 (7) D. hirsutum Ser 14 (7) Z>. rectum Ser 14 (7) D. suffruticosum Vill 14 (7) Erytbriria cHsta-galli L 44 (7) Galega offidnalis L (47) Do. 10 (4) 0. Orientalis Lam U7) Genista fcrox Poir 48 (5) G. pilosa L 21 (5) G. sagittalis L 44(42-45) (6) G. tinctoria v. angustifolia Ledeb. 48 (Ô) G. trîangularîs Kit 48(48-50) (ó) Glycine gracilis Skvortzow 40 (U) G. hispida-. 38 (SO) G. hispidaMax 20 40 (U) G. soja Sieb, and Zuce 20c 40d (44) G, soja var. akasaya 38 (S6) G. ussuriensis 40 (7) Glycyrrhiza áspera Pali 16 (4) G. echinata L (47) G. uralensis Fisch (4) Hedysarum clongatum Fisch, var. albiflorum Ledcb. (66) Ilymenocarmis circinnat us Síwñ (7) Indigofera asvera Perr (S2) /. decora Lindl 48 (4) 7. diphylla 8 (S2) /. gerardiana Wall _-. .. 2-1 (47) I. kirilowi Maxim 8 (44) 7. pam^om Heyne 7 (32) 7. pseudotinctoria Matsum 8 (44) I. pseudotincforia __ (74) 7. suffruticosa Mili .._. 10 (44) 7. sessiliflora DC 16 (Sé) 7. viscosa Lam 8 (Sé) Laburnum adami 48 (73) L. alpinum Griseb _ 48(£0) (0) L. vulgäre... 48 (73) Lathyrus angulatus L_- (71) L. annuusJj (71) L. avhaca L__ — (lé) L. articiUatus L 7c 14c (14) L. cícera L 14 US) L, cirrhosus Ser (71) Lathyrus clymenum L-. (7í) Do.. .14 (53) L. crassipes Gillies 14 (69) L. dumetorum Philippi 14 (69) L. ensifolius Bad— (71) L. grandiflorus Sibth. and Sra.. (14) L. heterophyllus IJ (71) L. hirsutus L (71) L. latifolius L (84) L. macropus Gillies (69) L. magellanicus'Ldjn. (63) L. maritimus Bigel T 7c (44) * Approxnnately. « Apparently name is wrong. 1022 YEARBOOK, 1937

TABLE 1.—Chromosome numbers in legumes—Continued

Chromosome mimbers Name Reference 2n

niger 14c {Í5) nigrivalis A. B iirkart 14 m) nissolia IJ (71) numidicus Batt (71) ochroleucus Hook 14 (69) ochrus DC 14 (ÍS) odoratus L 14d (84) pnlustris (69) pannonicus L. fGarcke] 14 (53) paranensis A. Burkart 14 (69) parodHA. Burkart 14 (69) pratensis L 14 (53) puhescens Hook, and Arnot (71) Quadrimarginatus Bory and Chaiib. (69) rotundifolius W illd (71) sativus L 14 (U) sessifolius Hook, and Arnot 14 setifolius 1J sylvestrislj (71) Do... (Ó3) L. aphaericus Rotz. (71) L. tingitanus L (53) L tuherosus L. (21) L. venosus Muhl..- 28 (69) L. vernus Bornh (71) Do.. 'iîc" (68) Lens esculenta Moench.. (78) Do. "Í4f' (68) Lespedeza bicolor Turcz .. (44) L. cyrtobotrya Miq (44) L. daurica Schindl (11) L. homoloba Nakai (44) L. sericea Bonth (11) L. sieboldi Miq .. (44) L. stipulacea Maxim 20 (11) L. tomentosa Siebold 20 (11) L. variegata Cambess 18 (11) Lotus angustissimus L 12 (7) L. corniculatus IJ 12c (40) L. corniculatus L. var. alpestris Lamotto. 24 (7) L. corniculatus \j. vnr.japonicus Regel... (44) L. crHicus L 28 (7) L.. cytisoides L 14 (7) L. filicaulis Dur 12 (7) L.hispidus Des,t 24 (7) L. ornithopodiodes L __ 14 (7) X. requicni Mauri 14 (7) Jj. siliguosus L 14 (7) L. tetragonolobus L 14 (7) L. uliginosus Schkiihr 14 (7) Ijupiñus albicoccineus -_ 48 (79) IJ, albus L oO (79) Do-. 4 40 (7^) angustifolius L 20 (85) Do 24 (44) .Do 4ÜC (61) barkeri Lindl TÁ)C (5) densiflorus Benth 48 (79) douglasii Agar 48 (79) elegans H. B. and K, T. H 48 (79) hartwegii Lindl 48-.W (79) hirsutus L. var. rnicranthus BoivSS-, 50 (79) iuteus L J.23 (78) Do .__. 24 (44) Do 44-4.5 (33) Do (61) micrnMhus Dougl 48 (79) mufabilis S wect 24 (85) Do 42 (54) Do 48 (79) nanus Dougl 48 (79) ornatus Dougl 48 (79) pilosas L. [Murr.] 42 (79) polyphyllus Lindl (il) ' Approximately. MISCELLANEOUS FORAGE LEGUMES 1023

TAIîLE ].—Chromosome nunihars in legumes—Coiilinueil

! Chromosome i numbers Ileference Name L no. \ n \ 2/2 1 \ _. _

IJ. polyphyllufs lAinl] __ _ -. __. -. __ .- 48 (79) L. puhescensUcnth _ - ..- ¡ i 48 (79) IJ. subcdrnosusîïooi: . __ ■ _ . . 36 (79) L. Sfjr.r.ulp.nfMít .. __ . ; 48 (79) L. varius'Li - - __... .._ 48 0^') L. venustiis'VWm. ._ 48 (?'.9) Medkagoapiculata\^\\\ú .. . 16 (29,28) M. arabica All ]6c (m M. arbórea L 32 (29,28) M.carstiensisV^u\L.. ._. .. 16 (2S) M. ciliaris Krook _ 16d (29, 28) Äi. cor onaía l^Gi^r __ ._ . ... ._. ._ 16 (23) M.denticvlataWiMú . 16 (29, 28) M.disdformis^^C _. . 16 (29,28) M.dzawakheficaTioráz .. _. _...... _._ _. . 16 (II) JV/. echinus DC 16c (25, 28) M.falcatal. _ . _. __.. ._ îfV (78) Do-— 16 (2S) Do 32(1 (29,28) M.gerardiWñlást.nnáKn ..... 16 (29, 28) M.§luHnosaM. Bich-.. . 32 an M.kelix^Vim -. H) (25,28) ^'I. hemici/cla Groi>íih. _ . ... 32 (tl) M.hisjMa _ . Me (2S) MA'niertezta\\'ú\ .. 16c (2'3) M.laciniata'M\]\ _ . .. 16d (29,28) jV/. lappacea Dosr 16 (29, 28) M.liUoralislihoáe _ . 16c (29,28) M.lupulinalj.- ...... __ ...... " " 8c (78) Do._ _ _._...... 16d (2Ö, 28) M. lupulinatvpicaXJThñn . . 32 (Ö) M.maculataWiUá 1« (25,28) 16 (29,28) M. media Pcrs 32,35 (23) M. mínima L . . 16c (25,28) M.murex\Yülú ... . ]6d (29,28) M. muricata (L.) All 16 (23) M. nigra Krock- _ _. _ _ __. 16 (29.28) M. obscura ÜGiz __. _ __. _ ._ 16,17 or 18 (23) M.olivaeformis Guss 16 (29, 28) M. orbicularis AU. _ ... __ ... . 16d (25,28) M. ovaiis urban (syn. Trigonella ovalis Boiss.) . 32 (6) M. pentacycla DO 1 . 3" 16 (29, 28) M.platycarpa {'l^.)Tràul\--.. (78) Do 16c (23) M. radíala L. (syu. Triqonella radiata Boiss.) .. 16 (G) M.rtqidulaDQ ." 16 (25,28) M.rigidula {IJ.) Dü?i>T..^ ._ .__ . 14 (23) A/^. roíaí a Boiss- ...... 16 (23) A'f. rugosa Desr_ _ 32 (23) M. rutheiiica TY2í\Ií\^ _ .. . 16 (23) ■ M. sativa Jj _ _ ._ _ 16d (78) Do-- 32íí (19) M.scvtellataMUl.. . _ ...... 32c (23,28) M.so/cirolnDiiby.... ._...... _ 16 (23) M. sphaerocarpa\lerU)].- _ . ._ _ _ 16 (25, 28) M. tenoreana Her _ _ 16 (25,28) M. tomata Mill 16 (25, 28) M.tríbuloidesDeí¡,r... _ .._.... 16 (29, 28) M.truncatula Q'àertn 16 (25,28) M. tuberculata Willd 16d (29, 28) M.'t7irb^nata^^'\l[d.. . _ 16c (29, 28) Meiîlotus alba Mecí. [Desr.(?)] 8(1 16g (19) M. dentata Pers_ 16 (6) M. indica A\L^^ 16c (23) M. itálica (Í,.) Lam I 16 (^) M. mdllotus indica A. and G. (syn. M. parvifiora .Dcsf.) . .. J . 16 (^) M. messanensis AW __. . ■' 16 (8) M. neaüolitana Tf^A^. ('i!.vr\. M. nracilrs DC^. ^ i _ 16 (6) ÎV/. officinalis 1 8 (W) Hn 1 iöe (19) ^f. segetalis Ser _...... _'. 16 (8) M. speciosa Dur ._. .__ . . J . 16 (0) M. suaveolens háb- __...... j 16 (6) M. sulcata Desf _. 16d (23) 1024 YEARBOOK, 193 7

ÏATtLE 1.—Chromosome numbers in legumes—Continued

Chromosome numbers Reference Xame no.

M. táurica Ser 10 (ß) M. wolgica Poir lOc (6) Millettiajapónica A. Gray.. (44) Mimosa púdica L 2t (44) Onohrychis crista-galli liam. 14 (14) 0. viciaefolia Scop (Í2) Do- 22 Í6t) Ononis alopecuroides L 32 (6) O. biflora Dcsf 32 ifi) O. fruticosa L 32 Ui) 0. hircina Jacq _ 32 (30) (G) O. natrix L 32 (6) 0. ornithopodiodes L 32 (6') O. reclinata L 64 (6') O. rotundifotia L 32 (6') O. spinosa L 32 (30) (fí) 0. viscosa L 32 (Ö) Ornifhopus sativvs Tirot If) (44) Oxyiropis halieri Biingc 16 (4) O. rishiriensis Matsum * -)3 (6'6') O. nraknsis VoAl. [DC] 16 (4) O. mfjinaf.a Fisch 10 (4) Packyrhizus angulatus .. (65) Paroclietus communis if) (m Pliaseotus aconitifolius Jacq _ 22 (7) P. acutifolius A. Gray 22 (41) P. angularis ^YiM. [(Willd.) W. F. Wiglit]. 22 (41) 7\ aureus Hoxb 22 (4t) P. capensis Tlujub 22 (7) y^. chrysantkos Sav 11 22 (m P. Lunatus L 11 U-i) Do. 22 (41) P. multifloTus Willd. (0) Do 22c, (41) P. mumjo IJ 22 Í41) Do.. 24 {6S) P. nigerrimus Juss 22 (7\ P. radiatus L i ..-. (4^) Do...- I 22c (^ri Do -.1 24 (63) P. irilobus ' 22 (41) P. rvlgarisjj . i (44) Do.. 22g (4n Piptanthus nepalensis Sweet. 18 (5) Pis um arvense Jj (85) Do Ï4 m P. elalius Bicb.-_ 14 (51) P.fulrum SibUi. [SJbth. and Sm.]. 14 (51) P. hvmile Boiss. [Boiss. and ÍsToó].. 14 151) P. jomardi Schrank 14 (51) P. sativum L 14t (8) Psoralea bituminosa L (48) Do.. * 2ÔC (47) P.glandulosaJj.- Í20 (47) P, macrostachya 20 (48) P. palaesf.ina L-...... Í20 (47) Rhynchosia pkaseoloides DC_ 22 (7) Robinia boyntonii AsJio (83) R. fertUis Asho 10 (83) R. hartwigii Kochne JO (83) R. hispida L _ In (83) Do-. 30 (48) R. kalscyi Hiitohins 10 (83) R. Inxurians (Dieck) Schneid. 10 (8S) R. psevdoacacia L 10 (47) Do 22 (4) R. viscosaYent (83) Securigera coronilla DC 12 (7) Sesba nia acuLeata Pcrs (-Í-V) Soja hispida Mönch 40 (41) Ä. 'mfl.r, lllini variety 20 40 (80) Sophora nngusüfotium Sieb, and Zuce 9 (44) S. chinensis G. [G. Don] 2S (5) ^ Approximate. MISCELLANEOUS FORAGE LEGUMES 1025

TABLE 1.— Chromosome numbers in legumes—Gonliniied

Chromosome numbers Name Reference no.

A. dnvidii Kon 16 (ô) Sophora flavescens Ait 18 (5) S. japónica L 28 (5) S. moorcroftiana Bcjiili 16 (49) Spariium junceum L ;-52) (5) Sivainsonia galefjifolia R. Br. var. albiflora Lindl—_. lo iW Tephrosia hookeriana Wit. and A IG (U) Thermopsis alterniflora Regel [Regel and Schmalh.] (ß) T. montana ISiUtt {71) Do- 18 (5) Trifolium albopurpureum T. and G- 16 (82) 7\ aiexandrinum 16 (8ê) T. alpestre L (1) Do- 16 (4^) T. ambùjuum M. B. 16 (4ê) T. angustifolium L__ 14 (4ê) T.arvense L (1) Do. 14 (4^) T. badium (1) T. campestre ._, T. cUîolalum Benth. {T. ciliatum Nutt.). 16 (8ê) T. dkhotomum H. and A .__ 32 (82) T.filiorme L ... 14 (42) T.fragiferum L (1) Do- 10 (42) T.fucatumJAnûl. 16 (82) T. glomeratum (1) Do.. 16 («) T. hybridum L. Do.. 'iöd T. incarnatum JJ. % Do- 14d (42) Do 16 (61) T.lappacewm IJ (1) Do 16 (42) T. lupinaster L 48 (42) T. maritimum lluús. 16 T. medium L % Do- *8() (42) T. mkrocephalum Pursli. 16 (82) T. minus (1) Do (82) T. montanum L H'O (1) Do- 16 (42) T. obtusiflorum Hook.. 16 (82) T. ochroleucum (0 2\ pannonicum Jacq.-. '4S-4Ü (1) Do.. (42) Trifolium parviflorvm KJirJi. 16 T. pratense L iif J)o. (52) Do 14d (42) Do (24) (61) T. procumbens li 14 (42) T. reflexum L 16 (82) T. repens L <12 (52) Do 14 Do. If) ai) Do 16 (20) Do 32d (42) Do (24) 28 (61) T. resupinatum L (1) Do 16 (42) T. rubens L 16 (42) T. scabrum L 16 (42) T. spadiceum L 14 (42) T. sguarrosum L 14 (.42) T. subterraneum 16 (82) T. thalii T. lumens Stov .-_ 16 (42)(^>^ T. variegatumJsutt 16 (82) T. wormskoeldii Lehm 48(?) (82) Trigonella balansae Boiss. [Boiss. and Reut.-]- 16 (76) T. calliceras Fisch. 16 (76) * Approximate. 1026 YEARBOOK, 1937

TABLE 1.—Chromosome numbers in legumes—Conliniied

Chromosome numbers Reference Name no.

T. coerulea (L.) Ser IG (S3) T. corniculata L 16 (76) T. crética (L.) l>esr lf')C (23) T. foenumgraecum L -•'v,r'V\ ]f)C (23) T. glomerata Ilort. (syn. I\f.cdicago brachycarpa í isch.). - 10 (6) T. melUotus coerulea A. and G. (syn. Meiilotus coerulea 3Jesf.)- 16 (6) T. rnonspeliaca L Í0 (B) T.polycerata L 28 (6) T. sínttía L. (syn. 7". canceUata DQSL) 16 (5) U/i?^: europaeus L Ü6 (5) U. Tiaww-s Forst 40 (5) V.parvifl.orusVouYV - - - (5) Vicia alpe6tr\s S\.%\ú\. (Stev.) -- (34) V. amoena Fisch (78) Do. 24 (75) V. amphicarpa L ].() (76) V.angustifolia L.... . 12d (75) V. atropurpúrea Desf. (75) 1)0. I4d (6S) V. aurantia Boiss. 14 (SA) V. biihynica 1/ 14d (7o) V.calcarata Desf.. 14 (34) V. cracca L (5c 12c (67) Do. 14 (76) Do. (78) Do- 28 (75) V. dasycarpa l'en.. 14c (75) V. disperma DC — 14d (75) V. dumetorum 14 (84) V. erviformis Boiss. 14 (34) V. ervilia Willci 14d (75) V.fabaJj (50) 'Do 12k (57) Do 14 (22) V. gracilis Lois (78) Do- (75) V. grandiflora Scop. (78) Do- 14c (75) V. hirsuta S. F. Gray .- i4d (75) V. hybrida L 12c (75) V. hyrcanica Fisch, and Mey. 12 (34) V. iathyroides IJ 12 (34) V. lutea L (78) Do 14c (75) V. macrocarpa (78) Do. 10 (34) Do 12 (75) V. monantha Desf.. (78) Do 14d (75) V. musquinez Bosc. 14 (34) V. narbonensis L... (78) Do. 14d (75) V. or obus DC (78) Do 12c (75) V.pannonica Cranlz. 12c (75) Do. (78) V. peregrina L. (78) Do". 12 (H) Do- 14 (75) V. picta Fisch, and Moy. (78) Do (75) V. pisiformis (34) V. pseudocracca Bortol... (75) Do 14c (68) T^ pseudoorobus 12 (68) Do 14 (34) V. pyrenaica Pourr 14 (34) Vicia salira L 6d (68) Do (44) Do 12g (39) V. sepium L Hd (75) V. serratifolia Jacq (78) Do 14c (75) V. sicula GQSS 14 (34) V. silvática L (78) Do...- (75) MISCELLANEOUS FORAGE LEGUMES 1027

TABLE 1.—Chromosome numbers in legumes—Continued

Chromosome numbers Name Reference no. 2n

V. tenuifolia Rolh. m 24c 05) V. tetrasperma Moench. 7c (78) Bo- "Í4c (75) V. unijuga A. Br. 6 (78) Do 12 (ß8) 1)0- (U) Do- 12c (7ô) Do. . 24 (S9) V. villosa Roth.. (78) 1)0- 14c (76) Vigna catjang (lîurm.) "VValp.. 22 (41) Do- 24 (63) V. glabra Savi 22 (7) V. owtthue'fim Vog 22 (7) V. sefiQuipedalis A. I. IMerters [F. Agcaoili] (U) V. sinensis Endl (U) V. ungukulata (L.) Walp 22 (40 V. vcxillaia Benth 22 (7) Wistaria bTachybotrysS\e\). and Zuec (40) W. floribunda DC (40) W.frutesceiis (L.) Ppir (64) W. macrostachya ISiult (64) W. muliijuga Van Ilout.te {JV. chintnsis var. mulfijuga ITook,)- (4)^ W. sinensis Sweet (64) W. venusta Kolider and Wilson (64)

REFERENCES FOR CHROMOSOXME NUMBERS

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