ae ‘DOCTOR OF PHILOSOPHY — iy 0 ENG seat pe Sie? re: A Classification of the Cultivated Varieties of Barley A THESIS PRESENTED TO THE FACULTY OF THE GRADUATE SCHOOL OF CORNELL UNIVERSITY FOR THE DEGREE OF DOCTOR OF PHILOSOPHY BY ROY GLEN WIGGANS NH Published by the N. Y. State College of Agriculture, September 1921, Memoir 46 oo CONTENTS i PAGE See aime HeranVvestImaALOMS:.. ceo. nos ocd Sx pee ee eS 369 a SELVES LOIS poe Oa 0 a a OM he TE gd 369 INERT OINSOECIOS fy eee SNe. ticcans Gil = SLL ce 380 “TOUR as ae ene Ie Sa ee A Mee 1 381 Material used in the present classification...................0.000. 383 Belo ol the barley plant. ..... one... banda sk ho 384 ep eeerta GL OLSS jy. s tah iors kw SOS ee Ge ee SR 384 ESCH A fect 2 eR Sais rN ue ee a Se Yes ct ee Seek ee te 384 fe aimuCMaTaAChers.,..< fs. 2.04. eae oe ste Sree CO eet 385 {S12 Fa 0 Gira ey 6) F201 ae ea eA PR ea eee aa 385 Wemexthrot lastembernode.s t= (i065 peice ceed Ateneo aoe 386 NG Oer OMapOCeS... (7-02 oe lw so ae eee eee 386 oD NSE RG 0 ite ara a aR ga ee eh A 386 Eetuucot plant in early erowth a... os... ok ae ee tee 387 Hmetezence.of awns and spikes... e.c. 06 4. 5 ook ei eae 388 Micimcoes ITN UET os,» oes ch ne ee oes ony vc we be ee oe en 388 ROMANO CIM IN Se ct, 9h Tere Ae aes akg Fe ea, Set ek 388 Ree EMAC LC USS teen a. ys 3220 sea ea ieee i aa RO Ree ae 388 PED reat Aer perk, ooo aie oases Bhs Reed et SE CE 389 Brawch=hedded: barleysi).. sec. 05s. 2s i. ad woe tees 392 1 BNC ANT ER SSI ne Og en an RES 393 iheneth-ol-imemodes-of rhachis... 2 \.f2 + tse deanna: 394 Angle oF melmation Or Kerel 5. occcers sos acute wees 397 EN RETO FTO) it's" 0 Ske aR gg i RP, NESE 397 Ammeniabion OF miermodessof rhachis: 2). 0)... Saletan cen 398 Bemence HERB IATAGUCTS oth... os tas Le eek Se ees bon 8 Gaon cn weal 398 TEETH AE 2 Elle ee ine cS SY ey ee me eee Se 398 COM POSUMON 12 hc MA es oe seman ets dee er ee 399 SISO melts gee enh wast SO A re eg ee et 399 POE rng ers. easels Ma meetey os fs Ate ied oa ues eR RAS 400 (Gia TO ges SRE ge ost y WERE a we, ne Ty Oe ea 401 ie lemma, or flowerlne lume...) «5 4S .602 bc aed ee ood 403 Adherence of lemma and palea to caryopsis........... 404 Merminal ap Pen agen 2 160 he eee en, pee ee 405 Number of nerves-of lemma. <0 e465 se ee Type XI CC} Lateral’ nervesi with teeth: 22%. 36.4 ea Pk ae ee Type XII The great additions made by the investigations of these men were the discoveries of the stable characters of the rhachilla, or basal bristle, and the lateral nerves. These two discoveries marked a real advance in the matter of barley classification. Many other studies were made on minor differences in varieties, which were not found to be constant. Some years later (1906) Broili published a classification of two-rowed barleys in which he used the Sval6f system to a very large extent, adding ° Rearranged in the standard form for keys. A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 377 little to the previous work. He, however, criticized the Sval6éf investi- gators in regard to the constancy of the characters of the rhachilla and the dorsal nerves. This criticism has been shown by subsequent investi- gators to be without foundation. Beaven (1902) presented the first comprehensive classification of barleys in English. The main groups which he considered under the one species, Hordeum sativum, are as follows: A. All spikelets fertile. B. All spikelets normal. C. Spike wide with short internodes..........:.......... Hordeum hexastichum L. CC. Spike narrow with long internodes......... H. vulgare L; H. tetrastichum Keke. BB. Spikelets of median rows normal, spikelets of four lateral rows diminutive and Will COUIn END leh eee Ae ease MnP Ee Een etoag GOD ke eee peo H. intermedium Keke. Cwspike wider witht shortambernodess.. sas-ce eee s cee coe eee var. Haxtoni CC=Spikemarrow, with lone imteniodes:. 2.6 =. setae soo - s « ae le stein oes var. transiens AA. Only the median spikelets fertile. B. Four lateral rows infertile or staminate. Ce Spikewide, swith shortinternodes.. 4. 0.0.6 sl sos clecene mies H. zeocriton L. CC. Spike narrow, with long internodes. .........5........-s:+.ss- H., distichum L. BB. Four lateral rows rudimentary and without floral organs (several Abyssinian SVATRGUIGR Ey gel eaeirisen Aero R sie Ria cite eee OEP a abn eae RL ee H. decipiens Steudel In his main divisions Beaven used these characters: fertility and width of spike, and length of rhachis internode. He used the name H. vulgare L. for H. tetrastichum Kceke., and divided the two-rowed barleys into zeocriton L. and distichum L. to correspond with the two forms of six-rowed barleys. He did not make a division of the decipiens group. In his description of varieties he used character of the spike (normal or abnormal), shape of the spike, color of the grain, terminal appendage, and character of the awn. His varietal descriptions include many recently developed varieties, especially those of Karl Hansen, which in several cases were riot yet well established. Regel (1906, 1908, 1910) differed from earlier workers (1) by basing his minor groups on races varying somewhat in environmental adaptations, (2) by recognizing only two densities, (3) by considering blue and purple as forms of white varieties, and (4) by making smooth awns a very minor character. Harlan (1914) made a study of barleys somewhat paralleling that of the Sval6f investigators. He made no classification, but gave the char- acters that are of taxonomic value a thorough study. He was able to verify the findings of many previous investigators as to the value of certain 378 Roy Gien WIGGANS characters in classification as well as in genetic studies. Probably the most important addition to the knowledge of barleys which he gave was that in regard to pigmentation, which is reviewed later, in the discussion of the morphology of the barley plant. The grouping suggested by Carleton (1916) is as follows: spontaneum, K. Koch, distichum, Linn., two-row barley. vulgare, Linn., common or nodding six-row barley. Hordeum { polystichum, Ddll hexastichum, Linn., club or erect six-row barley. intermedium, Kceke., hybrid barley. Carleton mentions one other type, Hordewm distichum deficiens, of which there are two forms, but he does not consider it as one of the main groups. Under each of the above groups Carleton separated varieties by the use of the following characters: fertility, color of the grain, shape of the spike, character of the awns, and habit of early growth. He gives only a brief consideration to the general classification, and contributes little to previous works. A little later (1918) Harlan presented a classification which is to be commended in many respects. This is given here in detail: Key to the species All spikelets fertile (6-row barley). iceramas)olvallifiorets awnediorhoodedt eee - eerie eel cicero eee vulgare L. Lemmas of lateral florets bearing neither awns nor hoods Only the central spikelets fertile (2-rowed barley). Lateral spikelets consisting of outer glumes, lemma, palet, rachilla, and usually rudiments ofitheisextialOneans acy a Seer ee aCe eat eh air oie eeae Re cucieele Reine distichon L. Lateral spikelets reduced, usually to only the outer glumes and rachilla, rarely more than one flowering glume present, and never rudiments of sexual organs....... deficiens Steud. This scheme is founded on the one character fertility, that of density being eliminated entirely. It likewise considers barley as consisting of four distinct species, on the grounds (1) that clearness is better secured by making the species a smaller unit, (2) that no group of wild plants of such wide variation is united under a single species, and (3) that there is abundant evidence that at least two parents were involved in the production of the forms now domesticated. The only difficulty in separating the main groups comes in a few cases in which the variety is more or less intermediate in character. As Harlan has pointed out, A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 379 this condition arises very seldom in the existing varieties. It may, however, cause difficulty in the future as the number of varieties increases. By the use of the adherence of the lemma and the palea to the caryopsis, the terminal appendage of the flowering glume, and the color of the grain, Harlan has described eight varieties in each of his species as follows: Key to the varieties Hordeum vulgare. Kernels hulled. Lemmas awned. Kernelsiwinitem blues onypurple.. dercr ibe cicne fetctar terre sto. a erseiac 1. pallidum. Aernelsiblackswss Serres we Sacral asa eae a a ercrerba tae hetcle et eiet ee 2. nigrum. Lemmas hooded. Kenmelsiwhites bluevior purple... 2a.c. «acct ss es eels a os seinen: 3. horsfordianum. ermal smn La eke tec yr crea et oniecdontne ayer acen a leearonensta anes ccaxe yore anateyotapst 4. atrum. Kernels naked. Lemmas awned. KWennelsiwhites ple, .0r) Purple es 2.54. ole fey one a eiett yee) vehicle 5. coeleste. ene salslstekestc om ieee conden enaye orth cictts ec pet oun ca teane: sooty Shea searege 6. duplinigrum. Lemmas hooded. iKemelsswhittesbluesor purples je. co ochre) Seteisbe sis st are eee oe elel ale 7. trifurcatum. MERI ELS MOLL Ketan cecha a stort hiss Rissa ste ae tera ce een aiedem sc oyauehane’ a sreteteiG 8. aethiops. Hordeum intermedium. Kernels hulled. Lemma of central floret awned or awnless. Kernelsawihites blue or purples > jis cota seein astoittene © Generel s 9. haxtoni. terre logo lerc Keane rs catego pees casket tence thet oie Stale 2 chek eueier ences cecum 10. mortoni. Lemma of central floret hooded. Mernelsawinites blues or purple...c.: soo. Saadies eek eels tose aie 11. subcornutum. Gnmnelap pre kere wes aye pad cosy sun 2s sone Rise om ere Pema a nes Ie 12. atricornutum. Kernels naked. Lemma of central floret awned or awnless. iKermelsawhitesbluevor purpleikc > Sd st..0 O_O OREO SX [eus9y oO 10/09 axytds jo odeyg soAIoU [ei07e'T YIMOIS Jo qqey Ayre = sUM®B qjoours 10 YSnoy BTNeU AyisueqT pousr9ey Jo 10J0D esvpuedde [euruise 7, BuIUIE] JO aoueleypVy AVY IET suryuel pesodoig oyids jo epnjzny ayids JO U}PIM Avisusq suM®B qjoours Io ysnoy peus9y JO 10]0D auIn[s 193nQ Jouslsy Jo 10]0D asepuedde [eururia J, BUIUIOT JO aousley py AWTIOT SI6I UuvlIeTT YIMOIS Jo yqey Ape ase puodde [eurursey, ayids jo advys uleid jo 10]JOD A} 10 9T6T u0jJIPIBD ByoRyy souin[s I23nO SUM®B yjoours Io ysnoy urIvis jo 10[0g Buty JO aoualaypy 9epousejUt jo yysueT ayids JO U7PIM AMIE cO6T usAvog SUME yqoours Io ysnoy Bey, jousey, jo aseg ————— | eee axtds Se OO EN eyids JO UFPIM 9pousequt jo yjsueT AUTO ules jo 10[0D BuUIUIaT JO ously py souln[s z91ng esepuodde [Bulut J, 668T 319419} V saAJou [e1907e'T Sera [eus98y jo edeyg APTI T €68T UO pue® 6881 plBBs199 NT ayids jo 1OJOD sje[a yids jo Juour -o0UBlIY aytds jo adeyg AVIA S88T S804 sTpoRyy [aur8y yjoours Jo ysnoy eyids jo ysueT [eus9y jo ysueT aytds Jo 10]0D soumn[s 123NQ BUIUIA] JO aouday py syuei IO SMOL jo Joquinny asevpuedde [eutursa,y, AUTO S38T ayouloO yy IT or BUIUIA] JO aoual0ypY Ayisueqd AWE T SI8T seqqnyg BUIUIZ] JO aouslay pV Ayisueqd ; AYO SPLT snhevuury yusy SAY BIGH, 40 NOILONULSNOQ FHL NI SHOLVOILSHAN] SQOIUVA AM SUTLOVYVHO+dO ASQ LNGUAKGIG] AHL ONIMOHY Gav], AUVAWAG A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 383 5. The classification of cultivated varieties by Beaven, wherein the treatment of the main groups was superior to that of Kérnicke. 6. The discovery by Neergaard of the stability of the long- and the short-haired rhachillas, and the smooth and the barbed lateral nerves. 7. The researches of Harlan in regard to pigmentation in barleys. 8. The reclassification of barleys by Harlan wherein the main groups were established on the basis of fertility alone. The relative weight given the different characters by the various investigators are summarized in the table on the opposite page. It is seen that there is a very marked lack of agreement in the use of the various characters for classification purposes, with the one exception of fertility, which is used as a basis of separation for the main groups or species in every case but one. MATERIAL USED IN THE PRESENT CLASSIFICATION In the present study 627 specimens were under observation, many of which were alike in name and in all observable morphological characters. These so-called varieties were largely collected in 1915 by Professor E. G. Montgomery, of Cornell University, for the purpose of classification. The most valuable individual collections were those obtained from the California Experiment Station at Berkeley, California, the E. Clemens Horst Company of San Francisco, and the Okonomisch-botanische Garten of Halle University in Germany. From these three sources alone an aggregate of 475 specimens were obtained, including all the types of any economic importance and practically all the rarer types. This original collection has been enlarged by additions from the Department of Plant Breeding at Cornell University, from the Office of Foreign Seed and Plant Introductions and the Office of Cereal Investigation of the United States Bureau of Plant Industry, from the state agricultural experiment stations, particularly those of Virginia and Wisconsin, and from other minor sources, all of which have aided materially in completing the collection. The collection has been in the hands of the writer since 1915 and has been grown each year in rod rows, one foot apart. Since yield was not a factor in the present study, the rate of planting was adjusted in all cases so that good development of individual plants might be obtained. 384 Roy GLEN WIGGANS MORPHOLOGY OF THE BARLEY PLANT | The first thing necessary in making a key for a group of plants is a detailed study of the morphological characters of the plants. This study is necessary for two reasons — to acquaint the investigator with the plants with which he expects to work, and to learn the characters that are not in- fluenced by environmental conditions. These are the characters that must be given the important places in the classification. Therefore a considera- tion of the morphological characters is of much importance in this paper. The various discussions herein not only include a description of the individual characters, but also consider their value taxonomically and the use made of them in the present and in previous classifications. However, the relative taxonomic value of the various characters is not given any weight in the arrangement of this section. The morphological characters as discussed herein are divided into three groups and are treated according to the following order: gross characters, spike characters, and spikelet characters. GROSS CHARACTERS The gross characters of barley, which include color, shape, and size of the leaves, number and size of culms, roots, and some other characters, are the least valuable among the three groups of characters from a taxonomic standpoint. The differences in these characters are usually not sufficient to warrant taxonomic divisions. Furthermore, they are the most variable of all the characters under different environmental conditions. Their chief value is to be found in varietal descriptions. Foliage The foliage of barley varieties presents a rather wide range of variations, all of which are difficult not only to describe but also to recognize. Varia- tions occur in color, length, width, and number of leaves. Kérnicke (1885), in his varietal descriptions, used four shades of green — bright, dark, bluish, and yellowish. Difficulties immediately arise in such descriptive terms, because the personal factor is too great in describing or recognizing such a character for it to have much value. Other investi- gators have recognized difficulties in using such characters, and as a eee A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 385 consequence have used color of foliage with much precaution. The length and the width of the leaves are variables which, like color, are largely dependent on environment and are limited in their use in the same way. For them to be of any value, accurate statistical work must be done. As a rule, when significant differences are found, it is between the larger groups, where no such detailed observations are necessary for the distinction. The most marked variation in leaves is between the spring and the winter barleys, which are discussed later. The variation in the number, exclusive of the basal leaves, is identical with the number of nodes, which is discussed in connection with culm characters. Culm characters The culms of barley. vary in several characters, but, in general, greater variations are produced by different environments than exist between closely related forms. For this reason these variations are, like the variations in the foliage, of minor importance in classification. Their only value is in varietal description. Such characters as height of the plant, number of culms to the plant, diameter of the culms, thickness of the culm wall, number of nodes, and length of the last internode, have been used more or less extensively in descriptions. K6rnicke (1885, 1895, and 1909) made use of many varying vegetative characters in descriptive work. Height of plant The height of certain varieties in a particular locality varies as much as 100 per cent and for this reason may be of importance locally, but these same varieties grown in another region may have a reverse relationship in regard to height. This has been well shown by Harlan, who in 1911 selected thirteen pedigreed barleys representing a wide range of types and planted them at four widely separated points. He found a marked regional response For example, Odessa was short and unpromising in Minnesota and little better in California, but was very tall and vigorous in both Montana and North Dakota. The Abyssinian varieties, on the contrary, grew well in California but were very short elsewhere, as is seen from the accompanying table taken from Harlan (1914): 386 Roy GLEN WIGGANS INFLUENCE OF GEOGRAPHICAL LOCATION ON THE LENGTH OF THE CuLmM IN 13 REPRESENTATIVE SELECTIONS OF BARLEY GROWN av Four WiprELy SEPARATED POINTS, THE SELECTIONS BEING ARRANGED IN THE ORDER OF THEIR Heicut at EAcH PorINnt | St. Paul, Williston, Moccasin, Chico, Minnesota North Dakota Montana California Hordeum vulgare..... Senvianise soe cscs Odessa oss ie eevee S. P. I. No. 20375 Oderbrucker......... Odessa) an aera ne Hordeum vulgare.... . Oderbrucker Meine hariae ei re ete Hordeum vulgare..... SUEpPTISeH ser erate Abyssinian SUM Mt ae eracracreer Ship Geo ouoaeces Sova. pos Gobobact Servian IPEINIGESS erssrea ial) ee Oderbrucker........ S/S AIAN Tae osian oil ch e Smyrna SUEPLISe se eiolee Manchuria......... S. P. I. No. 20375...| Manchuria SeLVIAM no ackeeee SUMMIta selene Kitzing, 6-rowed....| Summit SP Noe 20375 ea | SULpIISer mene tae Man chiiniatesess sear Odessa Kitzing, 2-rowed..... Kitzing, 6-rowed....| Oderbrucker........ Kitzing, 6-rowed Kitzing, 6-rowed..... Sh 24 IIE Gs AUB all Smomsneeh, Sako bec hcce Princess Abyssinian eee Princess) 1s ee ee ae (PAY SSinIaAn see eee Kitzing, 2-rowed SHAW MT. ay Go oekebwls ae Abyssinian......... Kitzing, 2-rowed....| Surprise Odessa chiectetstrmt ikatzine+2-rowed: -.-)|/ te tincCessae ae crete Hordeum vulgare Length of last internode The relation of the spike to the leaf sheath depends entirely on the length of the last internode — the one on which the spike is borne. The shorter the last internode, the less the spike will be exserted. The failure of the spike to be exserted from the leaf sheath has been used repeatedly in describing such barleys as the Smyrna and Princess varieties. The fact that this character occurs in the same varieties in widely differing localities is evidence enough that it is a true varietal character, but there is considerable variation within the variety. It is, however, characteristic enough in a few instances to determine a variety. In the present classification the length of the last internode is used in varietal descriptions. Number of nodes The number of nodes varies from three to seven in different varieties, but in all cases there is sufficient variation within the varieties to cause an overlapping, thus making the character uncertain. The number of leaves to a culm is identical with the number of nodes and consequently varies in the same way. Roots So far as it was determined, there are no varietal differences of roots that can be employed in classification. The ratio between tops and roots A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 387 may vary to some extent between varieties, but similar variations occur within the variety as a result of local conditions. Consequently, root characters are of no value either for classification or for descriptive purposes. Habit of plant in early growth The habit of the plant in early growth is very important because it is by this character that spring and winter varieties are separated. The difference appears both in the number and the attitude of the culms and in the number of leaves. The ordinary spring varieties of barley have a small number of culms which stand erect at all stages of growth, and only a few basal leaves. Winter varieties, on the other hand, have a large number of culms and leaves which are more or less decumbent during the early part of the development of the plant. It is at this stage that such varieties pass the winter period. In the spring, when new growth begins, a few of the many culms elongate, producing the flowering stalks and the grain of the plant, while the others remain undeveloped and sooner or later disappear as a vital part of the plant. If these same varieties are seeded in the spring, a large tuft of leaves and very short culms are produced early in the season. The plants will remain at this stage for a considerable period before the flowering stalks are produced; in some cases, in fact, the flowering stalks entirely fail to appear, while those that do appear are usually infertile or produce very little grain. For this reason, it seems that a dormant period is needed for the proper development of these varieties. An intermediate condition, which may be called semz-erect, is also found in some varieties, in which the number of culms and leaves is above the average for barleys and the culms tend to spread out to some extent. All of such varieties are, however, spring varieties. This character can be used only in varietal description. The distinction between winter barleys and spring barleys was made by Kornicke (1885) and by Carleton (1916) in describing varieties, but has never been used as an important distinction between large groups of varieties. In the present classification the habit of the plants in early growth is used only in minor separations. 388 Roy GLEN WIGGANS Emergence of awns and spikes The date at which the head appears is a note that has been generally taken by all barley breeders as well as by those studying varietal differences. More recently this has been replaced by a note on the date of appearance of the awns. Harlan (1914) showed that the latter is more nearly accurate and easier to obtain than the former. This has proved to be true in the present investigation. The only objection to using the date of the emergence of the awns, instead of the date of the appearance of the head, is that hooded and awnless varieties cannot be compared with awned varieties. This is not a serious objection, however, as this character can be used only to distinguish strains within a variety. The relationship — of strains in this regard varies with the locality. In the few cases of awnless and hooded varieties, it is necessary to use the date of the appear- ance of the spike as the distinguishing character. The date of appearance of either the awn or the spike is so variable that it has no taxonomic value. Time of maturity Ordinarily the time of maturity is correlated with the date of emergence of the awns, but, as is the case with most other correlated characters, some exceptions occur. This character is of value at times, not only in distinguishing different varieties or strains in a given environment, but also in detecting mixtures in the field. A pure strain will mature all spikes within a very few days. This character is much more reliable when used in connection with early-maturing than with late-maturing varieties. Late-maturing varieties are often ripened abnormally by unfavorable weather conditions. This character is used in the present classification only in distinguishing strains otherwise similar, and in varietal descriptions. Production The yield of varieties, although in a given region varying from very small to very large, cannot be employed for classification purposes because it is almost wholly dependent on environment. Consequently, production has no place in the present investigation. SPIKE CHARACTERS The characters of the spike, including variations in fertility, density, and rhachis, are far more important from the standpoint of classification A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 389 than the gross characters already discussed. Some of these spike char- acters which are the most conspicuous and the easiest to recognize, have been used in all previous barley classifications. In’ most instances either density or fertility has been given first place in the formation of groups. The differences in the characters of the spike are great and have been found to be constant under different environmental conditions. Because of these facts they have proved to be of much taxonomic value, and are so considered in the present classification. Fertility Barley varieties, as stated above, have been divided into groups according to the fertility of the spikelets, by all investigators who have worked on their classification. The first pre-Linnaean divisions were made on this character when only two groups were recognized, the two-rowed and the six-rowed. Linnaeus likewise followed this scheme, but used in connection with it the density of the spike, thereby making four groups or species. In 1885 Steudel (cited by Kornicke, 1885) fully described a third degree of fertility, which he designated as deficiens. K6rnicke (1885) recognized a fourth degree of fertility in his group, known as intermedium. Thus, four definite stages of fertility have been recognized as important in the formation of groups in the case of barley. In order to understand the variation in fertility, it is necessary first to know the structure of the barley head. Barley, in common with all other members of the genus Hordeum, produces three single-flowered spikelets at each node of the flattened rhachis. Structurally the spikelets are very similar, each having two outer glumes, a lemma, and a palea, which inclose the sexual organs. In all cultivated barleys, the central spikelet of the series of three at each internode is always fertile. The variation in fertility is found only in the side spikelets, which present four stages of fertility, as has already been stated. These different con- ditions of fertility are as follows: (1) all three spikelets equally fertile, with the lemmas of each projected into a terminal appendage, either an awn or a hood as the case may be, and with the kernels of the side spikelets almost as large as the kernels of the medium spikelets (fig. 51, A and B, and fig. 69, H. vulgare); (2) all three spikelets fertile, but the lemmas of 390 Roy GLEN WIGGANS B Cc D Fig. 51. THE STRUCTURE OF SIX-ROWED BARLEYS A, Dorsal view of perfect condition of fertility, where all spikelets are equally fertile and awned; B, ventral view of same; C, dorsal view of second condition of fertility, where side spikelets are fertile but about one-half the size of median spikelets and not awned; D, ventral view of same the side spikelets without terminal appendages, ending either in a point or bluntly, and the kernels of the side spikelets about one-half the size of the kernels of the median spikelets (fig. 51, C and D); (8) the side spikelets infertile, but possessing rudimentary sexual organs and all the structural parts of the fertile spikelets except the terminal appendages of the lemma (fig. 52, A and B); (4) the side spikelets infertile, without rudimentary sexual organs and with all structural parts very much reduced. In some cases, only the two outer glumes and a rudimentary rhachilla remain as evidence of the side spikelet (fig. 52, C and D). The relative position and size of the spikelets in the various types is diagram- matically shown in figure 53. These four conditions of fertility, previously recognized, have remained practically stable since they were first described. For this reason, and because they have been constantly used in all early classifications, they A CLASSIFICATION OF THE CULTIVATED VARIETIES OF BARLEY 391 are of the greatest taxonomic value. Some intermediates between these stages have been described from time to time by different authors, especially K6rnicke (1885). Most of these which were the results of crosses have proved to be unstable and have gone out of existence as varieties. Crosses between two of the above-named conditions of fertility give in the second generation practically all steps between the two parent types, but these intermediates have proved heterozygous in future generations and have broken up in a similar manner to the original first- generation cross. In the present classification, these four conditions of fertility are used as the first and most important character in the subdivision of cultivated barleys, for three reasons: (1) stability in all environmental conditions; (2) ease of recognition; and (3) weight given by all earlier investigators. A B Cc D Fic. 52. THE STRUCTURE OF TWO-ROWED BARLEYS A, Dorsal view of third condition of fertility, where side spikelets are infertile but possess all structural parts except terminal appendage of lemma; B, ventral view of same; C, dorsal view of fourth condition of fertility, where side spikelets are very rudimentary; D, ventral view of same 392 Roy GuLen WIGGANS a \ ty ) D E F Fig. 53. DIAGRAMMATIC SKETCH OF CROSS SECTIONS OF SPIKES OF VARIOUS TYPES OF BARLEY A, H. vulgare (dense form); B, H. vulgare (lax form); C, H. intermediwm; D, H. distichon (dense form); E, H. distichon (lax form); F, H. deficiens Branch-headed barleys One other structure sometimes arises in barleys, which has caused descriptions to be written of what were termed seven-rowed, eight-rowed, or nine-rowed barleys. Such a condition, which might be called a stage of fertility, is brought about by repetitions of the three spikelets at a node. In other words, instead of one series of three spikelets at each node of the rhachis, there are three series (fig. 54). In the case of the two-rowed barleys, if this occurs regularly a six-rowed barley would be produced, but instead of only three spikelets at one internode, all of which were fertile, there would be three fertile and six infertile spikelets. If the same thing were to happen in the case of the six-rowed barleys, an eighteen- rowed barley would result. At each node of the rhachis nine kernels would appear. No instances have been reported in which there was such a replication at every node. The change usually comes at or near the base of the spike, and at only a small number of nodes. 438 (HNO <2, PARAS re 439 UBRTNG ITER Ae eRe ON Ane 438, 439, 441 CAD LT GN aE Se 424 [GD [REDO TRO C DO re 424 Bayerische Landgersté. .... 0.2. i6.s.080- 438 SLEEK CSSIE EN oN wane on calcd ches +. eae ete 427 CLL eS Saeed A Og, = 422, 424 MESENUSIN ASSETS oils aS o kale sats a « Eee 439 [EGG IRGES SS ipl ay oa ee A | VP 421 Bestehorns Diamant...........+.e.-.. 441 PSESTE MONS INUISETS oo osc. ee sad ale = db lerae'e 441 BiaAckentinesse’.\ sstetie ww bok) ees 428 SACK MOUIIMICL: oa. 5 os ons on 6h eee 426 LACK WO-TOWEUs. 7 fs. «cy ct ehelel stole stale) 423 Schwarze Zweizeilige Gerste..........-. 449 ISCOULISCHLE PAULI re sera iota aoa, setuanel tere) ote 439 Scottische Perlchevaliergerste........... 439 eoitish Chevalier. <0 )-. sic ces we wee ote 439 Scottish Lothian Chevalier...........: -, 438 Scottish Lothian Standwell............. 441 SHAISGITLET Ties citi eae BG AO Oe EDC 438 SEICCUONAEN eieto clean che era's Sn clef stare 446 SClECHON 259 A eseeler estes nse ie oes 427 SRIECHOMOOS eye ceraels eres isle/o/a sieneta terete 430 PAGE SO CRONTINIS) 5 BGs coe GUAT CO EOE 440 SELCCHOMIGOTS: con, ha ae ate meets ae 443 SelectioniGlamcecees ceca ee oes 443 SelechiomiOl Geary teria eicie terest orators 443 Selection G2 es. cores ele che wrote wares 446 SelectiomiG2Z een cok ae se Sik oes oe 446 SElECHONIG25 Rete sae eee cae eco ee 447 SCIECHOMNIOZON eee eee cisine faerie 447 SelectiomiG aetna Cadi ge cn ro ne ee 446 Short six-rowed) Winter. +=... acme cece 426 PSA RLV OP ote ahah AMER a eae 423 ISULVET RANG Nise aioe a 2 sent aces 423(2), 439 SMIVIMA eee eee scree ene uae SO ee 4 SLUT Sa sid open ODO ORO BO OOROONEe 424, 439 TCU TION O LE oan ahaa ctchetaei. aie: eherece aisiaya eae 439 PSH CTRL I2) CAR NEGRO OI anne Cine oe 424 South African Cape Early.......... 423, 424 South African Golden Grain............ 439 South African Six-rowed.............- 424 SOULRUILUSSLON Fao ene ee 423(2) SVU Or SUN. VEU S «oie pap aLabonoe 422 SEE LEU SOLON aay o seve oat eros 421 SP snlerd0 3205 0 ee Sora tectslnetsavere 433 SERPS TSOC LD ie oe PAN ye anortnis aloteresa ls 423 SRE IBLOCU Sic ear Seis ate ee tee oe 423 SERED TOOK OE «Ae ce arte triers oeere setae 423 (50) 2rd eG Re intact ee Oe ne 443 SP STAI Sere oct a ortetere 446 Sap BEADS Gee yer cra tore sits nearest 429 S. P. I. 41157. 429 Se RELL D Siertay Wa tates ete ae aevelelsieias 422 SEPIA USO crete a cence sl eratorsiove 426 SATS LLG OUR Aa stoe sh tsioterens sic eaters 422 Sel il Ole ack ees etre avah efovevet tere 422 [Sopal 28 Game A LIU) aie cera Seer A a a en ee AN 443 Spiegelgerste aus Utund..........++++. 441 ISEQTUOAD CLL ee eee EN Morapn: frac oehel cae, ae oko 441 SUED HO sis RSA OOE COE AES EE DU TOO So 438 SUCCESS Hitter eerie eal cto ater ere iaes 427 ISUICCOSS ee ate nade erouche, ee Arana erste) shore 427 IS ULCLESS) DCONOLESS aU ariel aioe Sisiohsialiers 427 ISLS TUT ENS Bae ROR eea A ice SA Ae eae ear 438 Svalof Chevaliergerste..........+eee00- 440 Siralloy, JECHIGNGs 65 a noob boob enous aoc 438 (SUGANO) ETEMCCSE ys lade crcl = «ala a. 6 tele (eie =e 438 SUQLOI SW HUTUUS ei Netiaes. scalelele le ols\c siete 442 SUG OfpSUSUCMNGIS ers 212 aie <)-) eickenetetilel iets 441 ISUGLO eS OMUNEE este sate acco) <=) sicheisl« crete 441 Sia A ORO Or 441 Swedish Chevalier IT...........ce.e0-- 440 IS QUEAES IAG OLUm a ra oka fa, isrne ciavereioiens 438 SiatIS 0 GOULGI., ohn oo COS OA DODUOOOD OC 438 Swedish Oland...........-- Patel o/ chee 438(2) MSILEGIS ie EMINCESS Ite raiete) a clels ciel srcievere eters 438 456 PAGE Swedish Six-rowed............-+-+00- 423 Swedish Uplands. sa 558 soe, pace ae oboe 442 SWISS omens eR ane ae ee 424, 425 IS OOSSSUMIE Bon bab bobo sen sobous or 423, 438 Syllatrge ee Nae teas eee ae er eee 440 SUNTAN re ES se 438 TASMANIAN ots Sita reas hehe sicke EIS Re 442 Tasmanian Batiledore...............- 439 TasMantanyl deal s<-5> Ae ett-eeke ere ee 442 TCL ti ere rst ee es. eee a ee 425 Renkaneeroce rene nee SS iD Oa 425 Tan OWA ese sco Merete eR CET ate 422 AD GUMIPH Sk So se Bk Gee teas aa 425 TRUMISUATU SE eine ous © etonle ates ca eiiegte 422, 424 Y Mid 0S) DAs Bioko SSI St eed aocrcraeit Shae 439 MOAT), JOKE OSS S 6 od donnenncaGe 439 TP ICUS WPA CS ILEURET Mea tonsie ick rece 424 SA SOS ZANOTTI owncoaae gn sodcaoGes 423 LARA AN POON Dis on ep gon souecca%ose 439 Turkish (Rhodes to Sea of Marmora).... 424 TurkishySmiycnass 0. .0- sachs eee 438 Turkish (Smyrna Highland)........... 438 Turkish Smyrna Highland............ 439 Turkisk Smyrna Lowland............. 425 Turkishe Syrians 3).2.:.-3.jc 6 oc setoeke sos 440 Roy GLEN WIGGANS PAGE Two-rowed Blacks". ics 2.208 ale ven 438 UngarischesHanna: pence eee 438 Unterfrankische Zuchtaus-schussgerste.... 433 Uplands oto ac ne ees ae eee 442 Utah’ Winters. 22 56 cb oe oe eee 425 Utah Winten. .25 ceive a: cence 3 ee 425 Vermont ‘Champion. 2.2. +2... eee 438 Vaolacewmiz oo oecdncicis ¢. 00-00 eee eee 428 Virginia, Black Hulless..) mice see 428 Vargun@siicoded seo. see eee 427 Varginid Selection 1.5 sats acvaereer ete 441 VorgunianSclection/2 see eee 441 Virginia Seleciton-64 ..5 2:2 as. ceeeee 427 Vanginia SelectioniG47~ 22 26 seer 441 WebbssBartloselen.. fom s sis ctoiacee eee 441 Wessling 3.95 ties 78. oon 439 Wessling’s Trownengerste.............- 440 White Hulless.. 5.2. a3. eee eee 429 Wisconsin... 003 20.302 Sono ee 423 Wasconsin' Gir. oneness oe ee 423 Wasconsmv, Pedigree’. .c0 2 ie eee 423 Wasconsim Winter... ..2 2. scan see 424 Wustermarsch Wintergerste............ 422 Verlag: can. obs oe ten eee Renee 424 ZEiNesS Viered hs. 53.0% 5, ca San BA pee 438 ar = (coe eS 7 ae r # Ms, vt ve Fee | f re a v. a 7 iy i m * Os, i re i “s LIBRARY oF a