Susceptibility and Resistance of Berberis and Related Genera to Puccinia Graminis 1 "

Home , Berberis vulgaris

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MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUT"ION TEST CHART NATIONAL BUREAU or STANDARDS·1963-A NATiONAL BUREAU OF SlANDARDS-196~·A - SUSCEPTIBILITY AND RESISTANCE OF BERBERIS AND RELATED GENERA TO PUCCINIA GRAMINIS 1 "

By MOSES N. J,IWINE, Pathologist, Division of Cereal Crops and Disease8,and RALPH U. COTTER, Afl80ci"te Pathologist, Division of Barberry Eradication, Bl£reat' of Plant Industry 2 • •

CONTENTS l'lIge Page Introduction...... 1 Dlscus.qion amI conclusions...... 19 Material and methods •••••••••• __ •____ •••••• 2 Summary•••.••_••______._____ 2.'; NomenclatJ)re of Berberis fmd related genen\. 2 Literature cited ___._••••__ •__ ••••_•••••_.•_. 26 Barberries susceptible to Pucdnia gramin~__ 13 ]\>[embers of Berberldaeeae apparently Im mIme from stem rust_ ••• ____ •_____ ._•••_.. 18

IN'l'RODUCTlON Nmv that it has been shown conclusively that new parasitic races and physiologic fonns of stem rust, Puccinia graminis Pers. may originate through hybridization on barberries, where the sexu.;J stage of the stem-rust fungus develops, the question of exactly what inter :mediate hosts can and do harbor this destructive parasite assumes more than usual significance. Infonna.tion regarding the suscepti bility of Berberidaceae to stem rust is rather :meagerin the literature ~ of uredinology. Consequently, many exsiccati specimens in various en herbaria jn the United States and abroad were examined for ;,rust ;; infection; and numerous sp'eci.es, v!1rieties, and. hybri~ ?f BeJ;!~eris . and related genera were artifimally moculated wIth spondia of:differ C"l ent races of P. graminis. In addition, tield observations on the z: susceptibility of barberries under natural conditions have been lli~

1 Cooperative investigations between the Bureau of Plant Industry.. U. S. Department or Agriculture, and tho State Agricultural Experiment Station of the University of Minnesota. ' 2 The writers are indebted to E. C. Stakmll!!t professor of plant pathology, University of Minnesota, nnd .agent, Division of Barberry Eradication, .tSureau of Plant Industry, U. S. Department of Agrlcul· ture, for initiating the present studies and for numerous and extended field observations and hetbarlum e)plants of the common barberry used as controls in the inoculation experiments: and to S. F. Blake, senior botanist, Division of Botany and B. Y. Morrison, Division of Foreign Plant Introduction, Bureau of Plant Industry, for helpful adVice regarding the synonymy of the plants under consideration. . 96135°-32-1 2 'J:EC:~lNICAL BULL,E'.rIN 300, U. S. DEP'l'. OF AGRICULTURE .Arnold Arboretum of Harvard UniversitYi at Highland Park, Roches ter, N. Y.i and in N. E. Hansen's barberry garden at Brookings, S. Dak. Lists of susceptible and resistant barberries, compiled as a result of these studies; have been published from time to time by the Division of Cereal Crops and Diseases.3 This bulletin supersedes all former reports and contains all the results of artificial inoculations made in the greenhouse at University Farm, St. Paul, Minn., up to the prpsent time, the direct observations made in the field and herbariol, and the reports of other investigators that appea.red in literature. MATERIAL AND METH0l1S The lar.ger part of the telial material used in the inoculation.expcl'i ments came from collections mnde in early spring in fields and mead owsof the spring-wheat area, particularly in Minnesota and Montana. When the rusted grains and grasses were gathered in the fnll the straw ""US tied up in small bundles and fastened to a wire stretched between two trees, where it was kept during the winter under approximately natural conditions. Inoculations were made on plants havrng young and tender leaflets. (.v'ig. 1.) About a dozen rlisted straws were spread on a w'e screen suspended above the plants in an incubation chamber covered with a close-fitting glass top. The germination of th~ teliospores was facilitated by sprinkling the rusted straw twice daily with tap water. The yOlmgest and most tender parts of the barberries were thus Rubjec.ted to a continuous shower of sporidia. After 72 hours the inoculated plants were removed from the moist chambers, put in a shady place for two or three days to prevent killing of the leaves by the sun, and then moved to different compartments on the green house bench. Pycnia appeared in 5 to 7 aays after date of inoculation, and aecia usually developed about 5 to 7 days later, depending on the prevailing environmental conditions. Infection resulted, at one time or another, from teliospores of all but one of the stem-rust races found in North America. The exception wac;; Puccinia gramini.y phleipratensis (Eriks. and Henn.) Stak. and Piem., which has not infected any member of tho bUJ'bcrry family thus flU' tested. NOMENCLATURE OF BERBERIS AND REr.ATED GENERA The barberry family includes a great many Fpecies, varieties, and hybrids. ThiS is particularly true of the genus Berberis. Not a few of the specific and varietal names in use are synonymous. These synonyms sometimes are puzzling and conflicting and, therefore, the barberry nomenclature has been a vexing problem to the plant path ologist and ~speci!111y to the rust student. InTable 1 only members of the Berberidaceae are given for which there is some definite infol1l1a tion, obtained either from literature or from original experiments and direct observations, regarding the relative susceptibility or resistance to stem rust. Most of the infol1l1ation regarding nomenclature was obtained from the ,v()rks of Schneider (14, 15, 16, 17) 4 and Rehder (l0, 11),

I Thefirst of these lists a , ..>enred in the following article: ST.\ KllAN, E. C., and I,EVINE, l\f. N. A P ARTUL REPilitT ON TIlE Sl'SCEPTmJl.:~Y AJIll ItE~I$TAN{,E 0)' IIERBERIS AND JlELATt:D (lENEItA TO STEA[ RUST. U.l;. Dept. Agr., Bur. Plant lntlus•. Cereal COllrier 15: 278-287. IIl'2"J. [Mimeographed.] Subsequent reports nPPeared in Inter issues of the &ame periodical. • Italic numbers in parentheses refer to Literature Citetl, p. 26. SUSC"El'TIBILITY AND RESIST.,ANC"E OF BERBERIS 3 and from the lists published in the Index Kewensis (.~, 9). Schneider was considered the final arbiter in all doubtful cases pertaining to the synonymy of Berberis, Rehder to that of Mahonia,and Index Kew ensis was the authority used for Odostemon. In certain instances reference is made to sJ?ecial studies, such a.c; that of Piper (8). In this bull!'ltin ~e!baris IS considered the valid ~eneric name for all taxononuc entities commonly classed under this genus, as well as those designated as Mahonia or Odostemon. Synonyms of doubtful validity Unless otherwise specified were omitted in this bulletin.

FIGURE i.-Young plant of Berberi3 rmlgari3ln conl1ltion to be InOCl.lated with sporidia of Puccinia I1ramini.!. X ~i While fully realizing that some of those that have been included may be inadequate and ,untenable either because of their pre-Linnaean origin or polynomial designations, it was nevertheless thought perti nent to list them on account of the possible hearing they may haye on eradication measures, quarantine regulations, etc. In taxonomic literature identical names are sometimes used for different members of Berberis. In some cases this is due to uninten tional independent publication of the same name for thoroughly dis tinct species, in others, merely to misidentification of closely related species that may have different rust reactions. For example, B. chi nensU Poir. is at least moderately susceptible to Puccinia graminis, 4 'l'ECHNICAL BULLETIN 300, U. S. DEPT. OF AGRICULTURE while B. chinensis Franch. is immune from stem rust. But B. chi nensis Franch. is merely a synonym for B. thunbergii DC.; wher~as B. chinensis Poir. is an authoritative name for a barberry species i'.'Jr which there are some 10 synonyms. In like manner, B. brachybotMJS Edgew. (B. edgeworthiana Schn.) is immune f:tom stem rust. Thus far it has not become infected as a result of artificial inoculations with P. graminis, nor has it, to the knowledge of the writers, ever been reported susceptible to stem l"Ust under natural conditions. B. brachybotrys C. Gay, on the other hand, is listed as susceptible to stem rust by both Saccardo (13) and Sydow (18). The importance, therefore, of having a precise knowledge of the identity of a given barberry is obvious. All the species, varieties, and hybrids of Berberis under considera tion in this study, and their synonyms, are listed in alphabetical order in Table 1. In each known case the literature citation shows the source of original application. The accredited names are shown in boldface tYP{J and the synonyms in italics. In Table 1, in addition to nomenclature, reference is made to the authority responsible for the classification; to the tracenble origin, or nativehabitat, of the organism in question; and to its known reaction to stem rust.

TABLE I.-Synon/plty of species and varietie!l of Berberis mul related gcnera, their traceable origin or native habitat, a·na their known react.ion to stem rust

No. Nomenclnture I .~ uth~rit~~'- -~~id: ~~~~~fn~~ .11.... -~=t ~:~~~n

1 Btrberi.! acida Gilib. (See H. vul- f (17, p. il2~) ____ Eurnsia______See B. vulgsrfs L. garls L.) : 2 B. actinocarpa,Hort. (See B. vul- j (17, p. 325) ______I do______.! Do. garJs L.) I I 3 H. aemulaus Schn.=.B. din-I (16, P.stt) ____ West Szechwan, Pyoniaonly. phnlla Schn. ! qhina.. ;' ·t n. aetnensls Pres!..______: (/~, p. 660) ____ SiCIly and Calabna, ! Moderate <0 hcs,-y. Italy. : 5 n. aggregata Schn.=B. b""i· r' {l6, p. £95) ___ _ ElISt Kansu and West 'I Very susceptihle, nor- paniculata Bean=B. geraldii Szechwan, Chinn. mal. heavy Infection. Veitch=.B. ru./gari8 aggregata' Voss. (j H. aggregata prattll Scbn.=B.; (I6,p.£l.16) ____ West Szechwan, I Liglltinfoction,P¥cnia intrgerrima Hort.=B. polV, ChinB. ; only. antha Hemsl.=B. prnUii Sehn. , i B. alba Poit. and Turp. (See B. I (J7, p. £27)--- ______See B. vulgaris alba vulgaris alba Weston.) , ! Weston. S B.lDIesuthensis Hort. - ______/______: Very susceptible. 9 B. altaica Pall. {See B. slblrlca (t~, p. 896) ____ South Siberia; north i NormallnCection. PaIL) . Mongolia.. JO B. amurensls Rupr.=E. l'uT·' (J6, p. ~9t) ____ Chihli, Cbina; Amur, ! Very susceptible, ligbt qari& Rege!=B. /JlIlgari& amltren-:. Siberia. i to hOBYY infection. 8;8 Rupr. i :I 11 B. aml£remi3 japonica Rehd.: (10, p • .~89) ____ Japan______Light to beny;- inree (See B. regeiboa Koehne.) i tion. J2 B. anguli Ztl1i& Muss. (Sl!o B. (17, p. £(8)_-_ North America______: See B. canadensis canadensis Mill.) i Mill. 1:1 B. angulosa WaIL______, (J.j, p. :1.?8) ____ .llimalayllS______._ Nonnal inCeetion. 1-1 B. aquiJo/ium Lind!. (See B. (I, p. 296; .1, Western Pacific i S~El B. dlverslfolJa dlverslfolJa Stoud.) p. 292.) Stat~; observations St~ud. made in :Michigan and Quebec. 15 B. aqulfoUum Pursh=B. 1Iu1 (1/, p. t970; 8, Idaho, Oregon, Wash- . Fruit cspecialIy heav kana Kenrney= Mahonia aqrti p. 450; 9, p. ington; British Co-' i1y rusted; leaves folium Nutt.=Odo&temon '/IU/ 164·) lumbia. very lightly In kanuBRydb. fected. 16 B. aquifoliu m faBcicuTaria Iloon. (11, p. t970) __ _ Calif., N. Max., Mex- ; See B. pinnata Lag. (See B. pinnata Lag.) leo. Ii B. arbore8CCII8 Hort. (See B. vul (/7, p. i!2~) __ .. Eumsla______See D. vulgaris L. garJsL.) 18 D. aristata DC.=B. ari.tat" (14, p . .\;;/; 17, Himalayas and Nepal, ',' Light inrectlon; pus- f/oribllnda Hk. and 'l'h.=B.. p. £04;~, p. India; Afghanistan. tules rather small at chi/ria HamiIt.=B. coriarill ! i!9~.J : times, though ollen Royle=B. degan. Hort.=B.' quite large. jloritu1I4a Lind!. I I ;.;1

SUSCEPTIBILYl'¥ A,W RESISTANCE OF BERB:ERIS 5

TABLE. l.-Synonymy of 8peciell and varietie8 of Berber-i8. and related genera, .the-ir traceable origin or native habitat, and their known reaction to stt'Jm rust-Con. , --~:------·-;~:-~;t;:r!· NO,: Nomenolature I Authority mode of origin ! ltust reaction

~: B. ari&tatajlorlbunda Hk. and Th. r(~~, 'P.-46~---- -Himalay8Sand Nepal.' See B. arlst'.'tlt. DC. i (See B. arlstata DC.) I. . Indin; Afghanistan. . 20 '. B. ariuata sine-nsia Koch. (See I(17. p. tl7) ____ Caucasus______See B. chinensisPoir. i Be cblnensls Polr.) 21 B. arti~lat/l Loisel. (See B. vul- (17. p. £S4) ____ Eurasia______See B. vulgaris L. 22 B!'::!~~~ Hort______Fairly snsceptlble. 23 B. aslatl~ Roxb.=B. hl/po/eILca I (14. p. 456; 10, Himalayas and Ben- Moderately susceptl Lindl.=B. llicifolia Boxb. p. 492: 4, p. gal, India; Afghan- ble. I SOl.) Istan. 24 B. atroplLrplLrea Lauche. (See B'l (17, p. Garden form------l Very susceptible. vulgaris atropurpurea Regel) Sl6)----1 25 B. bealii Fort. (See B.Japonl~ I (11, p.I971) ___ China; Himaiayas_____ 1 See B. Japonlca (Thunb.) Spreng.) i I (Thunb.) Spreng. 26 < B. beanlana Schn..______, (16, p. 216) ____ West Szechwan, i AppareutlYimmune. ! ! I Ohina. ' 27 I Be bercmannlae Schn.=B. berg- : (16, p. 138) ____:_____do__-----______Moderately suscepti- I manniae accnthopllulla Schn. I: i ble. 28 B. bergmanniae acanthophlilla < (16, p. 138) ____ China______-' See B. bergmannlae ! Schn. (See B. bergmannlae i I I Schn • . Schn.) 1 i 29 B. brachubotrus Edgew. (See B. : (16, P.l63) ____ tl Indla______See B. edgewortb- ecigewOI-thiana Schn.) i I' lana Schn. 30 Be brlllebybotrys C. Oay______, U, p. f92). ___ -' Chllo______Normnllnfection. 31 B. bracbypoda Maxim.=B. bra- i (16, p. %85) ____ : K!lnsu, Shensi, and 1 Variable infection, chllpoda "lIIaxima Hor(..=B. vul- I j Hupeh, China. i ligbt to heavy. garis brachupoda Voss..' I I 32 B. brachupoda maximn Hort.!: (III, p. f85) ___ .1 china------See B. braebypoda (See B. bracbypoda N,llnim.) : Maxim. 33 I B. bretscbneiderl Behd., allied; (10, p. 49i) ____ North China. ______l! Normal Infection. to B. koreana Palib. I I .' 34 B. ~rtl'ipa1liculata Beau. (See B. , (16, p.295l_ .•__! Chlna______--\' Light to heavy infee aggregala Schn.) . I" '. tlon. 35 Be breVipanleuJala Schn ______-' (16, p. t96) ____ HUl?Ch, China______, Intection moder!lte. 36 B. brempes Greene. (See B~, (It, p. 816) ____ I Pacific co!lSt, Rocky ISee B.·repe!!s Lmdl. repens Lindl.) : • l\Iountams. 3; B. bUllfolla Lam.=B. dulcls, (/4, p. 14E; 4, ! Chile to Strait of 1\1a- : VeFY susceptib!e. Sweet. I p. fOt.) • gellan. l 38 B. caIIfornl!ll' Jo~on------' ______•.1 CaliL·______• "------1 Moderately resistant. 39 B. canadtIMIS Gwmp. (See B. : (/7, p. fI7) ____ : Caucasus______See B.ehlnensls Polr. eblnensls Poir.) 1 : ~O I, B. canadensis M1I1.=B. angIL- . (17, p. 907; 10, . Alleghenies, Ga., Mo __ Extremely sW!cepti /izaIM MIlSS.=B. calladensis· p. 490; 4. p. . . ble, both under (W1I1d.) Pursh= B. carolinla1/a £Ot.) greenhouse condl Sweet=B. dILmetorum /at/$simo Hons and in nature. "folio Herm.=H. fischerl Hort.= B.lolii. obveT&tHlvatia Mill.=B. latiasimo [olio canadensis Par. = B. 1nauTltanica Boer.=B. pi•• irera Raf:=B. sinernris canadt1lsis Regel=B. 8{lmis tripiicilnL3 Linn.=B. VUlgaris canadensis ,. I Alton. " : i. ' 41 j B. canadenslB (W1I1d.l Pursh. I (4,p.20£) ____ -' Alleghemes______Susceptlbl •. I (See B. canadensis Mill.)' . I ~2 B. caroli hoallghowBis Schn. (See' (16, p. 118) ____ Chlna.______: See·B. v~rnae Schn.• B. vernae Schn.) I 43 B. carolilliana Sweet. (See B. (17, p. M8l ____ Alleghenies______: Susceptible. canadensis Mill.) I. +I B. chintnais Bunge. (See B. pol- (16, p. f£7) ____ Chinll-______._! See B. polretll Schu. ntH Schn.) ; 45 B. chlnenais Franch, (See B. (17, p. ~19) ____ Japan______.. See B. tbunbergll j tbunberglIDC.): ! DC. ~6 ! ~:~~sen~':x::::}Jt'~~n~1:~~i~ (17, p. :tIB) ____ C'llucasus------_i O~~tI~~~erateIY sus Guimp.=B. guimptli Koch and BoucM=B. iberica Sweet=B. intermedin Koch=B. serotina · Lange=B. sinenBia Desr.=B. , ,pathulaia Schrad.=B. IIIllgaria · spathulala Gord. , 4i . B. chUria Hamllt. (See B. arls (14, p. 451) ----I India, Afghanistan ..__ See B. arlstata DC. t tataDO.) I 48 ' B. coerulea Hort. (See B. Vill- (17, p. 1£9) ____ . Oarden form ______See B. vulgaris vlo-' garts VioJacea Willd.) lacea Willd. 49 B. condnna HooL______(14, p. 3.91) __ ._! Hlmalay!lS ______Apparently immune. 50 I. B. coria ria Royle. (ilee B. arls (14, p. 451) ____1India, Afghanistan ___ _ Susceptible. talaDC.) 5t ' B. corui Veitch. (See B. wll (16, p.298) ____: West Szechwan, Lil\ht to heavy Infec · 80nae subcaullaJala Schn.) I China. tion. 6 TECHNICAL BULLETIN 300, U. S.bEPT; OF AGRICULTURE

TABLE 1.-Synonymy of 8pecie8 and varietie8 of Berberis tra(.eable and related genera, thflir origin or native habitat, and their known reaction to stem rU8t-'Con.

-~--~------:------"7""------''------No Nomenclature Authority Native habJ.tat or mode at origin Rust reaction

52 B.erataelfna DC_•••••••••.••••• (14, p. 657) 53 B.eretlea L_ •••• Asia Minor to Persia•• Slightly slIsceptlble. ••••_.•••••••.••••••• (14, p. 6li7) •••• Crote; Cyprus; Greece. Susceptible. 54 B. crdica Thunb. (See B. (17, p. S(9) •••• Japan_•••• thunber&fl DC.) _••••••••••• See B. th..nbl!l1lfl 55 B. dedlnata DC. Schrad•••••••••••••_ (17, p.ll0)..•. B. eanadenslsXB. Susceptible. vulp.ris. Ii6 B. deeIJnata olYPhylJa. Sehn_... (17, p. fIt) _... Hybrid 57 B. dentala rorm._•..••••• Very susceptible. Tausch. (See B. (17, p. 193) •••• B. ,ulprlsXB. 81- See B;emarlfnata emar!dnata WlIId.) blrlea. 68 B. dlapllana MaxIm.=B. WlUd. !fun· (111, p.3fl)••.• KansllandSzechwan, Modemtely susceptl· 'IIant'll8/3 Hlltch. China. 59 B.diaphanaSchn. (SeeB.aem- I ble. (l6,p.Stl)._•• West Szechwan, See B. aemuJanll ulans.Schn.) I' 00 B. dlctyopbyUa Frb.'lch•••••• Chilla. Schn. ~ ••• (16, p.lllB) •••• Yunnun, Chinn••••••. Mooarstely susceptl· 61 B. dleJslana ble. F6dde•.•••••..••••••, (16, p. 11M) ••,. Shensl and west P ycnlal Infectlon. U2 B. dlvers!folla I Szechwan, China. Steud.=R. aqui·l (I, p. e96; 4, Western Paclfio Susceptible. fo/lu111 Llndl.=,\[. diversifolia I p. t92.)...... States; observa· Sweet. 1 tlons I made In Mich· 63 B. dukb Sweet. 19an and Qllebec. (See B. bilII- ! (1.$, p. 142: 4, , Chile...... See B. buItroUal,anl. fona Lam.) f p. fBI.) tJ.I B. dumtlorum I· . .Bauhln. (See B. I (17, p. 26.$) •• __ , Eurasia••••••••••••••• SeeB.,uJprfs h vulgaris L.) , i. I 6.~ B. dumtloru1llfructu candido ! Mill. I (17, p. 2m ...... __ ...... _) See ,D. ,ulgarls alba (See B. vulprls!liba Weston.) i 611 B. dumtlort!'l1Ilatisnmofolio !. .' WestOIl. nerm. !(17, p. M'T} •••• .AlIeghemes.•••••••••.' See B. rmlgaris microphlilia i tion in the Regel. field. n B. emarginata brltzenslli schn.., (17, p. 195} •.•..•••••••••••••••••••••••I Infection ranges Crom 73 B. tmarginata latlfolia Tausch., light to heavy. I (17, p. lOS) • •••\ Hybrid roml. __ ._••.••! See (See B. emargiData Willd.) ; D emllJ'lfnata 74 B. fMc/cularis I Willd. Sims. (See B.: (4, p. £9£) ••••. CaliC., N. Mex.; Mex· I See B.plnnatal,ag. pinnata Lag.) : Ico. 75 B. fasciculala Schult. I I . (See B.! (4, p. £9.e)._ •••i •••••do.••••_.___ ••••••., Do. 76 plnnata Lag.) i i 77 :: ~::::rl~:~t~~~~';:::~:'; ~~~':~::;~~~::l ;:,r::::tlhle. densls Mill.) I :t~i~:;e~:~~·~.~~~~~- 78 R. llorlbtUldlJ Lindl. (See B.! (17, p. f04) ••••/ Nepal, India""""" arlstata DC.) : See B. arlstata DC. 79 B.loliis obvtrao-ooatis Mill. (See', (17, p. £06) ••••1Alleghenies..••.•..••• B. eanadensls Mill.) 1See B. eanadensls 80 B;franclsd-ferdlnandl I I 1 Mill. Schn•••• ' (16, p',eISi_ ••• West Szechwan, Light to heavy Ohina. I inCec· Sl B. fremontll Torr.=M. tlon. frt· (tl, p. 1971: West Texas to Utuh, Only moQerately sus- 7/lOlllii J<'tldde=O. fremonlii It, p. SIB,' Arlz., lind I Rydb. I CaliC.; I ceptlble as a general i 9, p. 164.) I Oolo.~. Nev., and mle. i New Mexico. I 82 I B.fructu albo Mill. (See B. vul- (17, p. garIs alba em"."""""""""""""j See B. vulgaris alba WestOIl.)' . Weston. 8.1 B. fructu rllbro Bobart. (See B. I (17, p. .e!4) ••.- Eurasia•••••••.••••••• vulgaris L.) ; SeeB.vulgarisL. 84 B. fuebsloldes i : Hort.=B. fU3' .•••••__ ••.•••••,•..••••••••••••••••••••• Susceptible. choidts Hort., . 85 B.fU3choidtsHort. (See B.fucb '•••.••_••••••••• B. sloides arlstataXB. vnl- I, Light to heavy inCec Hort.) : pris. tlon. 86 B. geraldii Veitch. (SeeB.allP'e-' (/6, p.1!95} •••. pta Schn.) East Kansu and west See B. aggregata 87 n; I8giana Szechwan, Chilla. Schn. Fedde=B. pube8Ctn8 (16, p. £85) ••,. Shensl and north Modemte Pampan.=B. VUluar.i8 gllglan" pycnlal in· Voss. I' Hnpeh, China. (eotlon. 88 II. g1auea=.If. glQuco. (Syn· ••••••••••••••••,...... onymy uncertain.) Susceptible. .' ......

TABLE 1.,-Synonymy oJ8pecies and varieties oj Berberis and I'!llate~ genera thf:ir traceable origin or native habitat, and their known reaction to 8tem rust-Con.

I. I Native habitat or RJ.lSt reaction NO., Nomenclature , ,I' I Autbority I mode uf.origin . 'l -'---1------.. ' ~! B. lI/aucucrns Hort. (See lB.! (14, p_.457) ____1 Hlmalayas______See B.lydum Royle. . i lyelum Royle_) i 00 ; B.glumacea Spreng. (See B. ner-I (4, p • .t9!) ____ .,'Pacific cossL______See B. nervosa Pursh. .:. I \'085 Pursh.) ! ' 91 t B. guimptli Koch and 1I0ucb6. i (17,p.£17) ____ Caucasus_____ ------ See B.~blnensls Poir. 1 (See B. eblnensis Polr.) I Infection ranging from 92 • Do baematoesr~ Wooton= M. (11, p. 1971: 9, Colorado and New light to "eav}'. I haemalocarpa liedde=O. /laem' \ p.178.) Mexico. lllocarpa Heller. 93. B. lIakooalt Hort. {See Do re- (10, p. 489) ____I Japan______------· See B. regellana Koebne. seUana Koehne.) - I !» IB.' benryaBa scbn______••______(16, p. 21Jl) ___.I West Hupeh and east Heavy pycnial' infec . - I Szecbwan, China. tion. 05 B. beteropoda Schreuk=B. Mi· 1(1~, p. 457)----1 Central Asia; Chiua.. __ Susceptib:e. eropoda coerulea Regel= B. spha- " . " erocarpa Kat. and Kir. r See B. heteropoda 00 B. heterop.,odacomtlea Regel. (See (14, p. 457) ----II--·--dO------Do beteropoda Schrenk.) . , schrenk. 97 B. heteropod(l oIllOllga Regel. (See (t4, p. 458) ____ Turkestan______See D. oblongs Schn. B. oblonp Scbn.) , ' 98 B. bookerl LeI!l.,,;,B. jamesonil' (16, p. !i5: 10, ! ~ikkim .aod Bhutan, Rather slightly SUS· Hort.=R. walllch.ana Hooker= P.495.) ,India. ceptlble. ;{!h~allichi.qna lall/olia Hk. and : 99 B. hooker! lati/olia Bean. (See B. (1O, p. 27) _____\ Java______See B. x:.ntbOllylon Hassk. 100 B.xantbOSy!oB horridaJlwgh.=B.xanthoxy-1 Hn.'lSk.) I (16, p. 27) ____ ) _____do______Po. ~H=~ I I ~~1 ~: bI~:ot~s:n~:d~0~SOO---B:-ni.\;p:.jfi6S~::=I~-indiu:.:=:::=:::::::::=:~yB~:i~n~eRoxb. asiatica Roxb.l I ' 103 B. Iberica Sweet. (See B. cbl-'\ (11, p. !J16) ____ Ii Caucasus..______See B. eblnensts Poir. nenllis Poir.} 1()4. B. Uldfolla Forst.=B. serrato- 114, p. SOIl) ____ I' Chile; Uruguay; Pata Normal infection. df!!tala Loch!. 1 ' t:0nia• 105 B ••lIcifolla Rosb. (See B. asln , (-I, p. 21Jf) _____ < Himalayas______See Do astattca Roxb. atlca Roxh.) j 106 B. inttgerrima Hort. (See B., (16, p. fOO) ___.II Chiua..______Susceptible. • anregal\la pottll Schn.) ;. 1 107 B. integerrima stenophglla Mtc ______do______InfeCf.\onH~k. relatively thosyion Hassk.) • light. 115 B. koebneana Schn______, (14, p. 814)----' Kumaon______Light to beavy infec . tion. 116 B. koreana Pallb______: (10, p. 490)----' (,hosen (Korea) _____.. Moderately suscepti ble. B. lalifolia Poit. and Turp. (8ee (17, p. 2!!S) ..__ Origin unknown ______See B. vulgaris mac 117 roearpa Jaeger. B. vulgaris maeroearpa , B.Jaeger.) latiJrsilll%/lo canadensi3 Par. •. : (17,1). £(!l)----' Alleghenies______Sea B. canadensis 118 (See D. canadenllis Mill.)! r Mill.' B.laxlflora Sehr.l!I __ ..______' (/0, p_ -I9S) ____ ~ Unknown origin but See D•. amurensls. 119 \ allied to B. amu- I B.lelebllnll Hort. ______..______': ______rensts. . Lil\ht to heavy Infec- 120 I '\ i . tlOn, I B.I",i3 Scbn. (See B.8oulleana (16, p./S7)---- China______.._ Infection rather light. 121 Sehn.) I: B. lichtrn31eillii Sehn. (See B.\ (16, p.' 214) ____ \. China, nortll.c8ntral!. See B. potanlnl 122 potaolnll Maxim.) Provinces. 1 Maxim. B. lucida Sehrad_ (See B. vul- (11, p. !i!!6) ____ Transcaucasia..------to heavy Infr.,v 123 ' LI~ht ~ garis ludda Gord.) I I tlOn. . B. Iydum Rovle=B. e/egans (1-1, p. 457: 10, 1 Himalayas______Wide range of Infec 124 I Hort.=B. eltj/ailtl3sillla Hort. = p.498.) I tion from light pyc- B. glaucescens Hort.= B. rusci· I nial to heavy secial folia Hort.=B. ulIIbtlllcala Hort. infection. =B. rulgaris tltgan31l0rt. 8 !I.'ECHNICAL BULLE!I.'IN 300, U. S. D."EP'1'. OF AGRICULTURE

TABLE l.~Synonymy of specie8 and varieties of Berberi8 and related genera, their traceable origin or native habitat, and their known reaction to stem rust-Con. -, --~~-~-""- ~--"~-----~--- -~--~---~ Native habitat or No.; Nomenclature _<\uthority mode of origin Rust reactio,l i----_. 1$ . B. Tllacrophvlla Rort. (See B. (l6, p. £7) •••-- Java______•______Susceptible: ! xantboll,lon Rassk.) 126 i B. maurltaniea Boer. (See B. (17, p. !OS). ___ AlJegbenies______: SeMeIIBI.' canadensis , eanadensls Mill.) 127 ~ B. maxima jructu Gerard. (See (17, p. 224) ____ Eurasla______: See n. vulgaris L. I B. vulprls L.) ! 128 1B. maximowie:ii Regel. (See B. (17, p. !iB1) ____ Garden form from See B. tbuubergli i tbunbel"g\l mmmowlczll seed oC Japanese maxlmowlczll 129 B.Schn.) meebanU Scbn______origin. LightSoon. to •heavY infec tion. 130 B. microph/lila Rort. (See B. (17, p. 226) ..-- Eurasla______~______See B. vulgaris L. vulprlsL.) 131 B. mieroph/llla serruta Rort. (See (17, p. 196) ____ Garden form ______See B. proVinclalls 132 B.B. mitisprol1nelalls SCh{8d ______se17ata Scbn.) (4, p. 29,'1) ______serrata Schn. Sus~ptible. 133 B; monsprli·;.8ia Rort. (See B. (17, p. 228) ____ Origin unknown ______See B. vulgaris mac vulgaris macrocarpa Jaeger.J rocarpa Jaeger. 134 B. morrisonensls Hnyata=B. (16, p. 822) ____ Taiwnn (Formosa) ____ Infection light. sp. Mayatn. 1 > 135. B. 11a1lU Greene. (See n. repens • (iI, p.1970) ___ PacIfic coast, Rocky See B. repens Llndl. • Llndl.) MountaIns. 13e IB.lIepaiensia Rort. (See B. vul- <4, p. £93) _____ Eurasia______See B. vulgaris I •. garls L.) 137 B. nepalensls (DC.) Spreng.= (10, p. ~9S; II, Indin, chIefly Rima- Susceptible. I ]'I. ?lepall/ensia DC. p. 1971.) layas. 138 ' B. nenosa Pursh=B. glvmaeea (11, p. 1971; 4, Dritish Columbia; pycnlal infection only. SIll'llng.=],I. glumQeea DC.= p.1I92; 9, p. Idaho, Calif. M. ?lervosa Nutt.=O. ?lerrasu., 164; 12, p. I Rydb. 816.) 139 ! B. 11eubertii Rort. (800 D. neu- (IS, p. 176; 10, Garden foml ______Infection range from I bertH Lem.) p. 492.) I light to heavy. 140 f n. neubertu Lem.=B. 'lleubtrlii (10, p. 49!i) ____ ' n. ,·ulgar!sXM. aqui- Do. , Hort. folium. 141 : B. neubert!! caroUnae Rort.____ (6, p. 1l26) ______Susceptible. 142 . B. neVinU "\. Gray= M. ?lmllli (9, p.164) _____ CaliL______Light pycnlnl to heavy ,· ];·edde=O. "erillii Abrams. aecial infection. 143 ! B. notabIDs Scbn ______(17, p. 204)____ Probably hybrid of n. Very susceptible. beteropoda and B. arlstata. 144 B. nutkuna Kearney. (See B. (8, P.444)----- Pacific coast ______See B. aqulfonum aqulfollum Pursh.) Pursh. 145 j D. oblonga Schn.=B. hetertopoda (14, p. 458) ____ Turkestan ______Very susceptible. ; oblonga Regel. 146 ; B. officinarum Dalechamps. (!:lee (17, p. 224) ____ EurasiB-______See B. vulgaris L. ; B. vulgaris J,.) 147 I. B. ottawensls Schn. ______(17, P.I!J!J) ____ B. vul garlsXB. Apparentlyimmnne. i . tbunbergli. 148 i B. oX/lacanlha. (See n. vulgaris (17, P.2J!4) ____ Eurasia______See D. vulgaris I,. • L.) 14.9 I B. oxgaellntha galeni 'l'itius. (See (17, p. £~4) ____ •____do______Do. i n. vulgaris J,.) ISO; B.oxllaealllhapulalaMagnol. (See (17, p. 224)·---I-----do------Do. i D. vulgaris I•.) 151 j n. parVifona SpfBb'Ue______....__ _ (J(I, p. !!96) ____ Kansu and Szechwan, Pycnial infection only. I Chiml. 152 I B. pedunculi3 racemosiB L. (See (J7, p.224) ____ Eurllsin ______See B. vulgaris J,. · B. vulgaris L.) 153 ' B. plnn3lta Lag.=B. aquifoliu11l (4, p. 292; 9, C~Iif., N. 1\lex.; Me! Snscept illle. I /aacicular/a Dean=B.jascieulariB p. 164; 1~,' lCO. I Sims=B. /aseieuluta Schult.= p. uno.) M. jaseicularia DC.=],I. &/'" '1~~ms~edde=O. jaseicu ari. r 154 B. pi.ifera Raf. (See B. cana- (17, p. 2(8) ___ -: Alleghenies______sej\e'[IIIB.. canadensis denslsMill.) 155 B. polretU Schn.=B. chinenaia (18, p. £26) ____ . Chihli and Manchu- Infection ruther light. Duoge=B. voiretii weiehangen3iB : ria, China. Schn.=B. smensis DC.=B. Bill- , en3is a1l1?1'$Iifolia Regel. 156 B. polretli laUfona Schn______!: China______Vert-susceptible. 157 B(s!,0~~~~:~~S~~.};B Scbo. (16, p. 226J ____,' _____dO______See D. polretu 8clm.

158 B.poillanlhaHemsl. (SeeB.ag~ (16,p.296) ____ West Szechwan, Infectionl1cavy. grepta prattll Schn.) China. 159 : B. potanluU Maxim.=B. lieh- (16, p. 214)---- Kansu, Shonsl, and Apparently immune. IfII3/tinii Schn.=B. 8phalerll northwestern Szech- Fedde. wan, China. 160 B. praltii Scho. (See B. aggre- (16, p.296) ____ Szecbwan, China ______Reavy infectioo. I pta prattU Soon.) SUSCEPTlBILITY AND RESISTANCE OF BERBERIS

TABLE l.-Synonymy of species and varieties of Berberis and related genera, their traceable origin or native habitat, and their known reaction to stem rust-Con.

~~r---'-'-~::::=-- "~--'Auth:ri~y- rN~!d: ~r~~ior - Rust reaction ___~______-I______rl______'I______

161 . B. pronndaJIs serrata Scbn.= (11, p.196)_" __ Garden form ______Modoratelnfectton. , B. micmphyUu 8

19.1 'I' B. spina aeida aice oxuaruntha" (17, p. 2£4) ____ \ Eurasia______See B. vulgaris L. Dodon. (See B. vulgaris L.) : ; 194 B. apillis trlplicibu~ L. (See B. (17, p. J10B and! E u r a 5 i a; Nor t h Susceptible. canadellsis Mill. and B. vul-, £24·) i America. l garisH i ...... 195 '. B. 81apJlana Soon. (See B. wU- (16, ,.p, 298) ____1 CWDn______. Light aecral mfectlon. \ sOllae stapHana Sclln.) ; ; 1116 ' B •.,ttllOphulln Hance. (See B.' (16, p.137)---- __ .._<10__•______See ,B. soulleana 1 Houl\l'ana Schn.) I' . Schn. 96135°-32--2 , '.J;','

10 T.F.CHNICAL BULLE'l'IN 300, U. S. DEPr.OF AGRICULTURE

TABLE l.-Sytl()nymy ()f species and varietics ()f Berberis and related genera, tllcir traceQble origin or nati!16 habitat, and their kn()wn reacti()n i() stem rU8t~Con.

:r- Nomcnclaturo . ---l-A::;:-'l- ·~o !lBbitat,or~-~-::::::I~ _'1 ! 1,__nJ_O_d_o_o_'_o_rig_in __ I:_--,-,-______11171 B • ..rocoullalato Schn. (See B'r (16,P.1!98) ____/ West Szechwan, Light tQheavy aeclal 1fibonaesnbeauUalataSchn.) China. lniectlon. 198 B. 8WasOJI Buckl.=B. tTlfolia (1, p. e06~ 9, Tex. to MoxIco______Light pycnla! to med. Schult.=M. 8Wa.!tfli l'eddecO'j p.118.) f 8wa8evi Reller. ! erute oeclal infection. 190 B. tblbetfea Schn.=B. DU/qari81 (16, p. 2£.0 ____ West Szechwan, tMbtlica Voss. ! ('hinn. Light to heavy inCec 200 B. tllunb,eTlliana Schult. (See B. ! (17, p. i19) ____ Japll!!______tion. tbllnberlil DC.) . See 101. thunberglJ 201 11. t,bun~~1iI B. clainellJli8 (17, p. i19: lof, ---__ DC. Dq.= do______Apparently Immune Frnnch.=B. cuttea Thunb.=B. p.464.) under field condi 8i71ernri8 Koch = B. 8intllJli! all tions as well as in the glU/ifol/a Mats.=B.tllunbergiana greenhOuse. Schult.=B. vulgari8 Thunb. I 202 B. tbunbergll auopurpurea 11______Garden form______Apparently immune. Rort. 203 B.thullbtrgli daw30nilBean. (See (11, p. £21) ____ 1' Japan______See B. tbunbt!rlil B. thunberwU minor ReM.) " minor ReM. 204 1B. tb~nberlll maxlmowlesll! (11, p. 22/) ____ , Garden (orm Crom Apparentlr immlmo , Scbn.=B. maximowlczi! Regel. I ! seed oC Japanese under a! condItions. i origin. 205 I11. tbunbergil minor ReM.=B. (17, p. 22J) ____j Japan.______• tAu'nbeTg;1 daw30nii Bean. I Do. 206 B. trifolia Schult. (See B.8wa (1,p.296)--___; Tex. to Mexico______I seJi Buckl.) t See 11. swase;1 Buckl. 207 B. trlfoliota Rartw. (See B. trl (11, P.1971) ___I' _____ dO______folJolata Morie.) See 11. trlfoHolata 208 B. ~follolat& Morie.=:B.. trio (II, p. /971; I, dO______1\10ric. !_____ Light pycnla! tei mod fo/mla Hartw. = .t[. trifol.o/ata p. 206: 9, p. erate secla) InCection; Fedde=O. tri[olio/aIU8 Heller. 178.) . rarely heavy InCec tlon. 209 '1: B. umbeData WaIL______(14. p.455; 10, Himalayas______.. _:_. i LIght to heavy iniec p·493.) : 210! B. umbtUi<:ata Hort. (See B. (l4, p •.m).-__;_____do______tlon. ()'dum Royle.) , See B. Iyelum Royle. 211 B. vanBeatil Schn______(17, p. £3O)_' __ B. nitebllXB. vul I j Light to .heavy Infec i garls. tion. 212 I B. vernae Scbn.=B. caroli hoa.·1 (16, p. 218J ____I West Kansu snd west Light to medium In. , Uhotn&ia Schn.=B. illtegerrima I SzeehwaIi, Chins. I, uenophulla Maxim. i fectlon. 213 '" B. violacea Polt. nnd Turp. (See I (17, p. 229) ____ , Garden COrID ______See B. vulgaris lio B. vulgaris liolacea Willd.) I . lacea Willd. 2H ! B. viridis HorL______.'______•______• __ : _____ Light to hoo\'y inCec ' . ' l' tion. 215 B. vulgaris L.=B. acida GUib.= 1' (4, p. £92; 17, r Eurasia; escaped from Extremely snsceptlble , B. aetinocarpo Rort.=B. arbor- 1 p. 211.'-££5.) culture anI! natural under nearly all con. , f3cen& Hort.=B. articulata ized In North dlUons. , Loisel. = B. dumetorum Bauhin "\merica. =B. fntetu Tubro Bobart=B. ITritabi//8 Sa!lsbury=B. maximo fruet" Gernrd = B. microphulla lIort.=R. lIepole1l3i8 Ho,t.=B. ojJicillamm Dalechamlls= 8. o.ruacalltha=R. ru:yacalltha qal. rlli Titius=R. oruacallthaPlltala i\lagtlol. =B. pedunculi3 raulllo, ~13 Linn.=B. racemifer(l folii. ciliatis HalIer=B. racem03a rllbra Slokes= B. 3anguinalenta Hort. =B.3errulata Rnl.=B. 3pecio3a lIort.=B. 3f)ina aeida &ioB orua. caT/tha Dodon.=B. &pin/s trip. licibl/8 Linn.=B. vrugari. arbor tleenl Koch= B. tlulgari8 eana. densi! Dlppel=B. vulgaris eu rrllgari6 Bri!luet=B. oulgari. hderophulla Wierzbicld= B. oul. uarls maxima Hort.=B. vulgari~ 'lRicropll!llla Kooh=B. oulgarl. normali3 Dippel=B. oulgari. lIormali8 tupiea RegeJ=B. oul. garia (lblongata Koch= B. vt'/gari., oboraJo-oblonga Schrad.·=B. Vllf garis rubra Aiton= B. vrtlgari. terotina Jaeger=B. oulgari3 SiD(' erf3pin!l" Camerarius=R. tlul. gari.. 8ptCi03a Jaeger=B. vu/. garia tupica Fiori=B. olllgari, tvpica 81lbrotrtnda Schu.'= R. ruluo quae, eI o.rVI}eanlha pulala 1 l\Iagool. SUE\CEprmILITY A~"DRESISTANCm OF BERBERIS 11

T AlJLE 1.-Synonymy of speci" and varieties of Berberis mid, related genera, their 'raceal;le~'igin or native habitat, ~nd their knoWft reaction to stem rust-Con. ,

NO.! Nomenclature -I Authority Native",l)abitatmode or origin or "I. Rust rCllction I

216 B.~laarl'Regel. (BooB.amur (16, p.I91)-___ Chitin, Clilna; Amor, Ses B. amurenllfs elllla Rnpr;) Siberia. RUIlf. 217 B. vulgar" Thunb. (&.e B. {tt, p. 484)- ___ Japtm..______Bee B. thunbeqli thunbeqll DC.) DO., " 218, B. rmlgar" d fruit oiold SIl3Ch. {/7, p. S£9) ____ Probsbly hybrid _____ ~ See B. vulgaris vlo (See B. yulpds vloJnee" laeea.WiIld. Wllld.) 219 B. rmlgaru aggregala VOSS. (See (18 p B95) , West China.. ' See B. a""resata Schn. 220 B.B·y~~t~~eston=a. (11: p: tB7)~] ______.... ~~~~~~~~~~ susceptible. alba 1'oit. and Turp.... B. clume- I . torum truetu candido MIll.=B. ,I fruetu albo MIIl.= B. wIgar" !laoa Schmd.=B. rmlgarla fruelu j I albo Klrchn.=B. wlgari.leuco- i I (arpo Kuntze=B. wlgar" fIOr 1IIal" leucocarpa Regll!. t 221 B.r.ulgarilr amuren.l. Rupr. (See I' (16, p. S01)._••: Chlhli, CWna; Amnr, i See B. amurcnsls B. amurensls Rupr.) Siberia. 1 Rupr. " 222 B. wlgarilr apprena Hort. (See (10, p • .j89). ___ __ .. ______--___! See B. Yul,garls as- B. v~lPris asperDla Don.) I • perw;:"Dan._ 223 B. pulgan.arboruttm Koch. (See I(11, p. S!l4) ____ , Eurasia______See B.vjilgarls,u. B. vulprls L.) j 224 11. vulgaris a8peJ'ma Don=B. (10, p. 489) ______•___ Susceptible. ouIgarl3 awella Hort. ! 225 B. yulprlS atrol,3urlJurea Regel (17, p. Zt6) ____ \! Garden form ______Light to heavy Infec =B. alropurpurta Lauche=B. tion when artificially ~rpurta Koch=B. llU/garia t inoculated as well as {o1il3 alr~rpurel8 SCh,ueb.=B. I under natuml condi rulgari. {oli" purpurti. Koch= tions. B. pu/garf3 1Iormali. alropur purea Regel=B. rulgar" pur- _ . l• purta Bertin. : 226\' B.lIulgarilr brachltpOOa Voss. (See: (16, p. £85)-.-- China•• ______See B. br3Chypoda B. brachypoda Maxim.)' ! Maxim. 227 : Rr.ulgarI8callademi.! Alton. (See I (17, p. !08}----i Alleghenies_ .. ______See B. canadensis : B. canadensIs Mill.) ! I • Mm. 228 ~ B. rulgar" canaden8i8 Dippel. (17, p. ,U~) ____ !' Eurasla______See B. vulprls L. ; (See B. vulprls L.) " 229 ! R llU/garis Cllanocarpa Kuntze. I (17, p. fB9) ____ i Garden rorm ______See B. vulgaris vlo i (See B. vulgarIS vloIaeea t: lacca Willd. , Willd.) ': 230 I' B. IIulgarls elegam Hort. (See B. I (14, p. 457)---- Hlmalayas______See B.lyclum Royle. Iyclum Royle.) I 231 I RlIulgariaemarginala Gord. (See (17, p. 19!J) ___ -'I Hybrid______susce,ptible. t B. emarpnata Willd.) 232 f B. rulgari. etU1ulgari3 Briquet. (17, p. £15) ___ -. Eurasia______._._____\ See B. ywprls L. : (See B. vulgaris L.) i 233 B. vulgari. flava Schmd. (Boo B' (17, p. !lm____ 1. ______..__ , See B. vulgariS alba i vulgaris aIbQ Weston.) Weston. 234 : B. rulgarf3 f(,lii3 alropllrpltTti8 (17, p. U6) ____ : Garden form ______See B.vulgarls atro , Schueb. (See B. vulgaris.l Jlurpurea Regei • • tropurpurca Rege!.) I , 2:15 R vulgaris foilis purpurti8 Koch. ! (17, p. Uu) ______dO. ______! Do. , (See B.vUJprIsatropurpurea ! Regel.), . 236 'I' B. puwarf3 (metu albo Kirchn. i (17, p. 2m ______, See B. vulgaris alba , (See B. yutprls alba Weston.) , ; Weston. 23; I B. vulgaris fructu marim,o Horne- (17, p. ilI8):__ -' Origin unknown------l See B. vulgaris ma l mann. (See B. yulprls ma-: croearoalaeger. I erocarpa Jaeger.) I 238 R rulgar., gi/glana Voss. (See B. (16, p. B85) ____ china.______SeeB.sDglanaFedde. gUliana Fedde.) I 239 B. pulgari. hderophlllla Wlerz- (17, p. BI4) ____1 Eurasia______..____ See B. vulgaris L. hicki. (See B. vtilprls L.)· j , 240 B. vulgaris Japanica Regel. (See (10, p. 489)----1 Japan______" Susceptihle. B. re~ana Koehne.) . 241 B. w/garis uacocarpo Kuntze. (17, p.IB7)--_-,------See B. vulgaris alb. (See B. yulprls alba Weston.), Weston. 242 Bi~~!~J~cld. Gord.=B. (17, p. Si6) ____ ; Tmnscaucasia__ ------i See B.lacida Schmd. 243 B. yuigarlo lutea Don=B. rul- (II}, p. 489) ______I Susceptible. garl.uanthocarpo Rort.! I ! 2-H B • .,ulorls macroearv.a Jaeger I (17,p. Se8) ____ : Origin unknown ______. Do. =B. lati{oliePolt. and Turp.=B. ; i I momptlien.ia Hort.=B. rulgari. : ' fruelu mO%imo Hornemann. I ; 245 i R Im/garia maxima Rort. (See (17, p. Sl6)----1 Eurasia..______-'I' See B. vulgaris L. t B. vulK1!rls J,:) _ '. 246 B. pulgarr, lI11crophulla Koch. (17, P.1!l4) ____ ,' _____dO______Do. I (See B. vulprls L.) 12 ~J'ECHNICAL BULLETIN 300, U. S. DEPT. OF AGRICUL'rURE

T AllLE. I.-Synonymy. of cpeeies and varieties of Berberis andrelaled lle1ter~ their tracliable origin or native habitat, and their known reaction lostem rU8t-,IJOn. .

I Native habitat or N°'I Nomenclature Authority mode of OrigLl Rust reaction ------·l-~------I------I------p. 24'i B. />!llgariB mierophlllla Regel. (17, 199) ____ Hybrid origin______c See B. emarginata (See :8. emarglnata Willd.) WlJld. 248 B. vulgaris nigra Don_.______(10, p. 489) ____ Probably hybrld------Susceptible. 249 B. vulgarla tlOrmali8 DIppel. (See (11, p. gS~) ____ Eurns;a. ______Sell B. vulgarisL. B. vulgaris L.) 250 B_ vulgari.! 'Ilormal~ atropurpur~a I U7, p. ££0)---- Garden form ______!se.e B. vulgariS auo- Regel. (See B. vulgaris stro purpurea Regel. purpurelf! Regel.) . 251 I' B. lIu/garia 'Ilormalis. leucocarpa (J7, p. sm ____ ------·1 See B. vulgaris alba Regel. (See B. vulgaris alba I Weston.. . Weston.) i 252 B. vulgaris 'IIormat!s typiel: Rage\. (17, p. !i£-I) ____ Eurasla______1 See B. vulgaris L. (See B. vulgaris L.) . i 253 B. lIulgar~ oblongata Koch. (See (17, p.1124) ______dO______, Do. B. vulJpll'ls L.) 2M IB. vulgarIs obovato-oblonga Schrad. (17, p. 21-1) ______do______i Do. (See B. vulgaris L.) I 255 B. /IU.'oor!a purpuno Bertin. (Jr, p. i£6) ____ GRmen form ______! J,fght to h~avy infee- (See B. vulgaris atropurpurea I tlon. 256 B~v':t~~~i8 rubra Alton. (See B. (17, p. ~.e~) ____I Eurasla______' See B. vuli;arls L. , vulgaris L.) , :l5i ! B •. vulgaris ,anguino/en/a Hort. (17, p. 2£5 and Garden (orw_-- ______1 Very suscept.ible. (probably the same as B. sall- i 2£9.) 1 I gltlllo/etlta Hort.) (See B. vul- 1 I I II!m'Is L. and B. VUlgaris vJo- I " I racea WllId.) ( 2581 B. IlUlgari. serotlna Jaeger. (See \ (J7, p. 224) ____ Eurasia ______, See B. vulgaris L. B. vulgarIs L.) . 259 B. vulgaris sbeyaJle Hort______1______1' Light to medium in fection. 260 B. vulgaris 8ire crtsllinu., Carner· , (17, p. 224) ----J Eurasla______See B. vulgaris L. arius. (See B. vulgaris L.) , 261 B. vulgaris spathula/a Gord. (See, (17, p. 217)- ___ Caucasu8..______Susceptible. B.eblnensls Polr.) . ' 262 B. IlUlgariB 8pecio.a Jaeger. (Sea (17, p. 224)---- Eurasia------l See B. vuHgarls I" B. vulgaris L.) 263 B. lJ"ulgaris 8tapfiona Voss. (See i (16, p.1!98) ____ China______" See B.1vIIsonae stall 26! B.Bn:~~:::!'&~~~-~~I~~~~--,------I------______; Li~r.~oS~e~vy idee ~1i5 B. vulgaris thibetica Voss. (See B. (16, 'p. .e£4)---- China______, See B. tWbetJca Schn. tbibetlea Srhn.) , . • I· 260 B. vulgaris tgpiea Fiori. (See B. (J7, p. J!24) ____1Eurasia______,,_. See B •. vlDl~rls L. vulgaris I •.), , ~'67 . B. vulgariatupieo 8ubrotuneR Schn. ! (J7, 1'. £20) ______do______Do. I See B. vulgarls L.)· I ~'6S I B. vulgaris vtolaeea ,YiUd.=B. (17, p. 2!!9l ____ Garden (orm prob-' Suscept.ihlc. • coerulta Rort.=B. 8anguinea ably hybrl d be . Kocll=B. 8anguinolenta lIort.= tween IIJ. vulgaris B. violaeea Poit. and Turp.=8. and B. eblnensls. rulgaria dfruit molet Spach=B. vulgaris cllanoearpa Kuntze. 26U B. vulgari. wi180mae Voss. (See! (J6,p.297) ____ West Szechwan, 18ee ·B. wll80na·e B. wJlsonae Hems!.) \ China, HemsL 270 B. vulgari. xanthocarpa Hort. (10, p. 489) ______ISee B. vulgarlslutea (See B. vulgaris lutea Don.) Don. 271 B. pulga quae et oXl/aeanlha pulata (17, p. J!!!4) ____ Eurasia______See B. Vlwgarls L. Magno\. (800 B. vulgariS L.) 2i2 B .•r.allichiana Hooker. (See D. (1IJ, p. 26) _____ India______,' See B. booker! Lorn. bookerl Lorn.) 27a B. wallichia'1l4 Lindl. (See B. (l6, p. S1'} _____ lava_____... ______.. f See B. xanthoxy)on xanthox),lon Bassk.) I HassI>. 2i4 B. lOallichia1!a latifolia Hk. and (16, p.!!O) _____ India.______See B. bookerl Lorn. Th. (See B. bookerl Lorn.) 1, 275 B. wallichiana xanthCJXvlon Schn. (16, p. m----- Ja.a______See B. llantboxylon ; (See B. xantboxylon Hassk.) Hn...

T_'\BLE l.-SytiOnymy of sllecies and varieties of Berberis and related genera, their traceable origill or native habitat, and their known reaction to 8tem t·1t8t.-Con.

:Nativc habitat or No·1 Nomenclature I Authority I mode of origin Rust reaction -1------,---1-----1---- 182 B. xaUlthux),lun Hnssk.=British Columbia to See B. nervosa Pursh. vosa Pursb.) : , Calif. I 291 N. haemcUocarpa Fedde. (See B'I' (ll, p. 1971) ___ ' Colo. aud N. Mex_____ , See B. baematooarpa baematocarpa Wooton.), ( . Wooton. 292 .1I.japonicaDC. (SeeB.Japon-' (11, p.I97/) ___ 1 Cbinr.; Himalayas _____j' See B. JailOnlca lea ('rbunb.) Spreng.)' i , (Thunb.) Spreng. ~!93 M. mpaulensi8 DC. (See B. \' (10, p. 493; 11, I' India, chielly Hima- ! See B. nepalenBls nepaIcnsJs (DC.) Spreng.) p.I971.) layos. , Spreng. ~.1 .ll. lIerv08iJ Nutt. (See B. ner- i (11, p.I97I) ___ , British Columbia, j BeeB. nervosa Pursb. yosa Pursh.) ! i Idabo, Calif. ' 295 1 .ll. neoilli; Fedde. (See E. ne- ,______• Calif______, See B. nennl! A. I vln,I.A. Gray.) I '.. I Gray•. 296 I. M. pmnata Fedde. (See B. pin.. ' (11, p.I970) ___ CahC. to Mexlt'o ______: Susceptible. nata Lag.) 2971 M. repen. Don. (See B. repen,s ; (11, p.,19'i'O) ___ British Columbia to IApparently immune. I Lllld!.). 1 , N. Mex, ., . 298 1 M. 8wa8ey. Fedde. (See B. sw'a- (I, p. f96) _____ Tex. to M6.'(loo______, Moderately susceptl i se),1 Buckl.) ble. 299 ' M. Irifoliolala Fndde. (See B.,' (II, p.I971; I, ,_____dO______Sligbtly to moderately trlfoUolata Moric.) p. 1196.) : susceptible. I 300 Odo.temon aqui[oliu1R Rydb. (See; (9, p. 164,' 12, : Alberta to CaUL_____ See B. repens Lind!. B. ropens Lmd!.) : p. S16.) 301 O. ja&ciculari8 Abrams. (SCl3 B. i (9, p. 164)-----, Celif. to Mexico ______1! See B. plnnata Lag. ) p1nnat~.Lng.) " : 1 30'. O. jremontll Rydb. (See;S. fre- , (9, p. 164; 12, Nev. to Tex ______, See B.fremontu Torr. montll Torr.) , p. SI6.) :1031 O. haematocarpus HeUer. (See~. (9, p. 178) _____ Colo. and N. Mex---- See B. haematoearpa haematoearpa Wooton). : Wooton. , . 3M O. 'lIer~OSU8 Rydb. (See B. ner- t (9, p. 164; IS, Britisb Columbia; See B. nervosa Parsh. vosa Pursh.) p.516.) Idaho, Calif. l' 305 O. 7ItOillii Abrams. (Se-a E. ne.. (9, p. 164) _____ Cali!.______See B. nevlnll A. nnll A. Gray.) Gray. 306 O. 'nutkanu8 Rydb. (Se.e D. aqul- (9, p. 164) _____ Pacific coast ______' See B. aqulfoUum foUum Pursb.) Pursh. 307 O. reptns Cockerell. (See B. re- ' (9, p.164) ___ .. Alberta to CalIL ____ .' Apparentlyimmune. pens Lind!.) ; :iOS O. 8lUa8tlli Heller. (See B. swa- i (9, p.17[f) _____ Tell. to Mexico ______' Slightly to moderately se)'l Buck!.), I susceptible. 309 O. lri/ollo/atus Reller. (See B•• (9, p. 178) ______do______' See B. trlfoUoJata _I trirOU~~~_t~"!v~~r~~ _M_o_r_ic_._____

BARBERRIES SUSCEPTIBLE TO PUCCINIA GRAMn~IS As stated in the introduction, infonnation regarding susce:ptibility was obtained from three sourc~a, namely, results of artificial mocula tions in the greenhouse, observations of natural infections in the field and of herbarilIDl material, and a study of records in the literature. In Table 2 those species, varieties, and garden fonns are listed for which definite kno\vl~dge regarding susceptibility to stem rnst is 14 TECHNICAL BULLETIN 300, 11. S. DEM'. QF AGRICULTURE available. The results of original inoculation experiments and direct observations, as well as references to literature, are recorded in sepa rllte columns.

'l'.-I.lILE 2.-Species, varieties, and hybrid.~ of barbeN;" known to be ?lIOTe or le8s 81'8 ce1)lible to stem Tl'St 1

Source or Inrorl':1ltlon

Orlginsl data

! ltosnlts oC urtlJlclw inocnlatk'n with moos of ! Puccinia gmmhil~ ~' I 01 I ogros· . '.. <;ill , DerberJR specJes, varJetil's, find I tJdis ovcllsepone secullS tnttcl i: i hybrJcls ------~ il!!Lltemturercfer. ; I~ b ~ ~ a .g, enoos reporting ~ 1 I' ~ =: a =: a =: § =: a =: ai susceptlbllJty &.. , O'J oS =:!l oS:3 .sa:9 :: ~:9 -E: ., , ',d "S -a m ~l e..; ~ a :i ~ 0. ~ ... ~ tC\ '2! -c.s 81:.2 fj1:~ i k.:2 ~ -S ~ 8'f OJ, :.s!!i:.9a::.s ::.s=::.s!~ ~ ! 0"3 co <:) '3 to 0 '3 0 ,.. CQ 0 __ to.c; ~I .8~:.811~tJ~!i!!.!~f!-g g: ~~~~g~§g~~~~~~~¥ ~ zi~zi~zi~Zi~zi~~ , . -l------~r----- 3 i aemuJans Schn...... 1 O? 1 O? 1 O? 9 1 L 2 O? __ I______.~ ... ____. 4 i aetnensls PresL___..__ ..__.... --•.-- 1 O? ...... 29 11 H 6 5 M + (l.J,18) 5/agsrepta Schn....__ ..__...... __ • ...... 2 2 H 8 O? ..., ____ ....____.... 6 aggrepta prattJl Scbn...... __... __.... __ ... __ • 10 2 L 1 O? _,."...... __• __ .. 8 alesuthensls Hort...______...... 1 O? IlL 13 10 H 1 1 L +'...... 9 al/aica Pall. (=slblt'lea PaIJ.)__ ..... '" ..... '" .. '" ...... _ .....' (13,6) 10 • amurensls Rupr...... 3 2 H ...... 86M 36 32 H +1 (13,18,6,5) 11 al1;df;1~~~~~~:~::-...... _. :.... ,.. '" 4 1 L 1 O? .h...... 13 augulosa WalL...... ~ ...... '" '" ...... '" ...... +-1...... 15 aqulfollum Pmsh...... 1 0 t 10 O? 3 O? 92 35 L 39 3 L +, (13,18,6,2,6,7) 18 arlstata DC....____...... """ 7 6 MIl L 14 1 L 3 O? +1 (13, 18, 6,)!, 7) 22 uvenslsHort...... IlL 4 4 H IlL ..I...... 23 aslatJea Roxb...... '" ..... '" ..... """ 2 1 L 1 1 L +1 ([8) 24 at=~~:p~~~~:'k~~:...... + (6) 27 bergmaon!ae Schn...... 1 O?...... 3 I M 6 O! ..,...... 30 braebybotrYIi C. Oay...... 1 O?...... 6 2 M 3 0 t: , -:1 (13, I~) 31 braebypoda Maxim...... 1 O? 187M 9 2 CLJ ~ (,) 33 bretsllbnelderl Rehd...... __...... '" .. '" '" ...... '" +/'...... 35 brevlpanleuJata Scbn...... 1 1 L ...... 33M 11 11 M _ ...... 37 b:ulfolla !"'m...... '" ...... '" + (18,7) 38 ealIfornleaIepson...... _...... 2 O? ...... 9 5 L 2 O? _...... 40 eanadensls MilL...... 5 5 H 1 O? ...... 14 13 H 39 34 M +/ ([, 7) 41 cUlIademis Pursh. (=cana- I 43 1ca~~~~~oot..·(;;eaii.:· ...... - ... '" ...... ~...... -. -.- "I (13, 18, 6, I) densls Mf1l:} ...... _..._...... ',-- ...... -...... +1 (1",18) 46' ebinensls POlr._...... _ ...... 80M 6 O? - (7) : I::~8Jeitc~~~~o1~:8~~· .. ~ ..... "..-...... -....., . (3) I eaullalata Schn.)...... 1 O? 1 O? .. '"... 8 O? 6 3 M _ ...... _~._. =;= 56~ I deellnata=:r~~~~:::::==:=::::= oXfP.bylJa Scbn.':....:= '''''_::: ==: ::1 :=: 1 :::L ...== :::_.... ==: ==~7 ==~ 7 =~= H ==~ 5 ==~ 4 H +~I====::=:=:=:=iii..._...... 58 !dlapbana Manm...... '" ..... '" '" ...... 6 6 l'tI _ ...... 60 dleJ)'opbylJa Froncb...... "', 8 O? 4 3 M + (7) 6l , dlehlana Fedde...... _...... _...... 16 1 L 6 O? _ ...... 62 1dlversifoUa St~ud...... _...... '.' ...... '" '" ...... •. (1) 67 durobliveosls Schn...... 6 4 M 21M,+ ...... it emarglnata Wll\d...... _ ...... I O?...... "'1'" 5 ~ M ...... + (2) 72 ! amarglnata brltzeusisSchn...... 3 L J 1 H + ...... 76' rendieriA. Gray...... __ " • __•__.. __ ...... __ ...... 14 12 H 4 4 M + (1, .'a) 1 See footnote 3, p. 2. " 0,

SUSOEPTlBILI':l'Y AND R'ESISTANOE OF BERBERIS 15

TABLE2,~Species, parieties, and hybrids of barberry ~nownto be l'1iOr:e or'lesSltltil~ ceptible to stem TUst-Continued ~ ,

Source 01 Information

Origillllldata

,------"1-1------ 17 fillCAerf Hort. ,( - eanadenala' • MiIl.)______------___ ------II (tS) 8180 fremontilf'randad.fel'dlnandi Torr______Schn____ --2 --2- ---- L 2- --- 1 --- L --__ ---______--- .!10 112 1\1 M --;;0 --- 4 --- L +- ______------ _ Sl fnebllehles Hort______-______-______+ ______85 flUdolduH{lrt.(=fneluioides Hort.)______2 2 II ______;; ___ 16 14 M 4 4 L + : ______87 ~&lana Foode______,-- ___ --- 1 O? 1 0 1 17 3 L 3 O? - ______88 iIIIuea....______j__ ------__ ------___ --- ______-- _____ ~ ___ "~ (18,6) 112 IiHmatoearpa Wootou______--- IlL ______21 15 1\1 6 M - ______'j,i henr)'ana Schu______.i._ --ii ------.~ 1 L 1 O? - ______=_ 95 beteroDOda Schrenk______l 1 u? 1 O? ______lOlL 2 O? + (t8, &) 101!18 h:rbrldasenatahookeil Lem, ______Hort______1__ ___------___ --__ ___--- ___--- __-- ___------___ 197 41 II L ---J ---O? --- +______------ _ 104 Jllelfolla FOlst______--_ --- __ --- ___ -______2 2 II _____.. ___ + (18,6) 106 iRtt3trrimaHort. (=aanpta 110 ia~~~_~~~~~~~~~~~~~~r ~~~ ~~~ ~: ~~: ~:: ~: ::: ~:: :~~ ~~~ ~~~ ~~~ :~~ ~~~ :______~~, mt~ra'IIta:;~rF.(~~.iitiioi;.iiin-i------18 1 L 17 O? ------ 115 k.:~~n.sCiin::==~=====:J= =:= ::: == === :== == =:= =:= 1~ ~ i --2 -i -i,; == =:===::::=:::: mr:~~J:~::=::=::=::::==I== ::: =:: == =:: =:: == =:= === __ ~ __: _~ ::::= ::: +:::::::::::::::: 120 ' Idec:JlnU Hort__ J 1 L 6 6 ¥ 14 14 M + (7) 121 ------1------123 i~d~S~h;:~~~~r ------11 6 ,J 1 O? ------ ' ludda Gord.)______!__ ------5 3 1\1 2 1 L -______•______12.. l:vdum ROYle ______,l o? 2 2 L ______15 101\1 6 1 L + (6,~, 7) 125 'Tb~J~~ H~~)--~:~~~-._- ____•______:______(13) l~ ~=~~~i;i;~:=:::::!:: ::: ::: :: ::: ::: :: ::: ::: --~ --~ ; ::: ::: ::: ~ ~~~~~~~~~:~:5~~~ 13; nepalenslll (Thunb.) Spreng_. ------•. ------'I (/8; 8) i~ =s:: Pl~~:-(;:iieiibertiq------.------4 1 L 1 O? --,------Lem.)______, 1 ] M ______4 4 H IlLI+"I (IS, 18, t) 141 nenllerto earoI1nae Hort_____!______. (6) 142 nennO A. Gmy ______;______1 0 1 ______11 5 M 1 1 L !__[" ______m153 =i:l~~:::::::::::?:Plnnata Lag______+_ :::--_ ---::: =~_____ =:~ .__:~: ::__ :::::______11~52? f ::~7 ::~O? ~: --~II:::===:=::=::::(I) }~ ==:~~OIii-Scliii:::::::::'::·::: ::: :: ::: ::= == :=: ::= ~ i II i g ~ : ::::::::::=::::: :: :~~~!~~u£;=-I------3 1 II ------.. pronndaJIsprattU Schn·) BetTata______Scbn____ I__ ---___ ---___ --__ ---___ ---___ -.__ ---___ ---___ 4J O? 2 H ---7 --- 7 -- M -______------ _ 161 j t~I =:W~~~~i~~/ --~ -:- -: --~ ~~ -~ --~ -~- -~: -== -~ -~~ -~ ~ +l------~~~ , I ,loIaeea 'VUd.) ------1-- -__ .__ ._ -- ____ -______8 1 L - __ --- ___ -..------______172 , Btrfd~j.~~~~:--~:~~~-~~~-! I' O? IlL ______104M 2 O,? ..1. ______16 'l'ECffNICAL BUi:ILE'J'lN 300, U. s. DEP'J,'. 01" AGRICi,-'::TURF!

TABLE 2.-Species, varieties, anfi hybrids of barberry known to be more or less 8US • ceptible to stC11l rust-Continued

Sour~"e o[ !n[ormatlon

Original data

: Results of artificial inoculation with races of Puccinia graminis '" ---~--~--.---~----I] agros- c Berberis spe~ies, 'l'lll'letle~, lind tldls u\·elllle poon seeIllls trltlcl t. hybrids !------, EJ,lteraturerei~r. I)· 11 $ a b $ ;1 encesreporting o I ~:: ~:3 te:: 0'1 == 0'1 == a susoopt,."jlIty ~ 12: "_s S ~ ~ 5 en ~ E tIJ ] 5 ~ ~ :a f ~ =~c.=~c.-='tc.-~c.-t)~.! "E. .i:~~i:o.;.~-E.s~-t!.!~~~~ .... ~~ .:.Egj,::.sgj:::.s::s::.e:g:::'::::sJ: S ~a:~a:~'3:~:;:::a:'O §'h ~~~~~~~-§~~-§~~§~;-~ ... E9I. ... E ... 8 ... 8 ... ." t: § :,) ~ ;3 ~ 1~:3 ~ = ';:1 ~ ~ UJ _ ~__ _~ rZ :: ~ rZ ~I~ c2 ~ ~I.!. ~ z c2 2 -~'ll-_____ ,176 IiIblrlea P.lIL______--1------+, (13,18 6,5) i~ , ~~~::gu ~~~C-{~ebiiieiisb------1 --- --1---·------r- 4)' 2 L 2 2 H +------:----- Potr.) ______-T------,---'------j--- 8 Ii MI ___ ------+1 (18 1) 185 suuUeana 8cl1l1______-- -__1___ 1 0/' t -- ---r-- 151 3 M 2 0·/ ______• __ 188 species 355 Furrer______-- ___ 1__ -- - __ --_ -- ---J-- IlL ------t------:__:::=__ i: :ra:~I!- ~.ln'. 5~wiiSOnae- J °l? --1------___t___ 2 1 L------1--- --. ------ 197 sJ.:t:~I'l:i:: s~~Jn:--(:;Wii:- -- --r- -- --r------1--- ~ L I o(? +------ 6OnaesubeauUalJltaSchn.}. ______1 0, t ____.'___ 6 4 M J o;? + ______198 811'_11 BllckL______• 1 0l? all[ L -- --+-- 26 20 ~I 4 4 L __ (1,7) 199 thlbetiea Selln_: ______1 11 II -;; _71 ___ -- ___1___ ~ 3 M 2 2 L ------______208 tr!folJoIata MOflC______• -- ___ j___ _ 2 L ______12 12 1\1 2:! 21 L _. (l) 209 umbellata WnlL_____ •___•___ 11 1 L 1 O,·f. ____ --_ 19 12 H a H ._ (3) ~1l ,anfleetH Scbn ______-T--,------__ - __ -- ___:--_ ~ 4 H 2 1 1\1,______.12 vernae Schn______--i------a 1M 2 1 1\11- ---______~}~ I::~l:'f::=:::=:=::=:::=::::: i:fiil-ii30 -iii!-M -OI--sIIi a:J ~ ~ 2:!~I,:zoll1 ir. +-ris;is,-i;;-i,=5: 2:!0 !vulpr1s alba Weston______-r-l------r--!+ ------!:-:~,-:~ nl i yulgarls asperma Don______--,------___1___ --1---'--- -.------" ------r-- +------ !!25 I yulprlsatropurpurea itegeL ------22M ---I--- -- + ______~1 !j~~~~m:~~~~~l1d.r~~~~- ______;____J. ______+______.______r + ______2~O I i!:j~IK::!~~~~-~I~-~=~------l-- __.1..__.1.._ ------1------,---1--- +------ ~~ ~::.=I~~~~pa-juege;=: ::,==: :::: ::: ::: :: ::: =:: ::= ::: ::: :::j::: ::::j: ::::::::::::::=: ~ ~~;.:~~~:::~~~:~~- -r- --r ------r-I------r--r-- +------ 257 ,,~~r~·)-iri;,giijiio-lriii;i-Ii;ri~- 11 1 L 1-- --T------I 1 I! l\t III 9, H + ------.----. (~yulprls ttolaeea WlUd.) __ -- __• _____ - __ -______•______---J---'--- + ______.______i!; ~~s~~~=I~grt-o;;r(C -r-- ---;-- ---( 1 1 Ml 5 4/ M ---;---1------..------ '. (=chlncnllis POlr.)------.. --- --T------.--- 26i \ yu]prlssuJeataHort..------__ ~ ______J 1 L -_ -__ .-----J--'1 1 II 17.17/' Mt'++ ----------.______26i1; yuJprIs ttolacea Willd______.1 ______I .r ______1______• __ __ J ______+ ____ .______:ml , utIsonae HemsL______11 1 L 12 O,! -- --- ___I\I 6 M 13, 1 1>1'------.----- 279 j ~:~~Hort~::_:--~'~~~:~~~~- --t--- ___ ._ ---t--- _. ______1 O? I,lL ______•____ _ 280 . wllsonae VUf. "j,'lrelly" nort._ -- .--- __ J __ ------___I 2 1 L 1 0l? ------ :!81 ' wIIsonanar. "Splll'k1er" Hort_I__I___ ---i_=_---I-=-~= ~-=~: II 0l? -- -~------NOTE.-Expiunation of symbols.-&latirt 31UctplibilUj/: L=ligl1t Infection, M=moderate infection, H""heavy infection, ?=reaction doubtflll; field observations: <+)=normal infection, (-)=no infection de~ted. In the case of artificial inoculations, relative susceptibility is ex pressed in the tenns light (L), moderate (M), and heavy (H) infec tion. When inoculations have thus far failed to produce infection the ),eaction is recorded as inconclusive, or doubtful, and symbolized SUSCEPTUHLI'fY AND RESIs'rANeE OF BERBERIS 17 by a question mark (?). Further experiments ,,,ill be necessary to prove either by positive results that the plantr. in question are sus ceptible, or by numerous negatiye results that they are immune. "'here direct field or herLrarium obserntions were made, the plus (+)

a a

A b

FIGt:RE 2.~lllf"ction oC PI/ceil/in gramini. on B,·rheris. X 2; A.-Light inCection oC P. gramini., trifici on B.Jremol/tii: a, 1')'<'lIia; b, R<,rin. n.-\'pry light infection oC P. gramini., avenae on B. triroliolata: u, l>ycnin; b, ucciu

sign illdictltes that the plltiltS WNe normally rw;ted, the minus sign (-) that no infection WtlS de.tl.'cted. • It will be 81.'1.'11 from 'ruble. 2 and from Figlll'cs 2 to S, indusiYe, thut barherry plants differ in their reactions both as to rust races und to environmental conditions. The same barberry plant may 18 TECHNICAL BULLETIN 300, U. s. DEPT. OF AGRICULTURE react differently to the same rust under differing conditions. It may also vary in its reactions to different rusts under the same 'Condit,ions. Two varieties or spedes of barberry may also react similarly to one race of rust but may differ in their reaction to another race. Sus ceptibility in the barberry, therefore, is relative and is subject to biotic and climatic variations.

MEMBERS OF BERBERIDAf!EAE APPARENTLY IMMUNE FROM STEM RUST ill Table 3 are included those members of the barberry family which have not become infected after repeated artificial inoculations with the different parasitic races of stem rust and have never been found rusted in nature, although some of them had been under close observation for many consecutive years. While there can be no reasonable doubt that certain of the species and varieties recorded in Table a are immune, cert,ain others may in the future prove to be susceptible to some race or form of P. graminis. This may be par ticularly true of those species, like B. ottawensis Schn., 01' B. Potaninii Maxim., which thus far have been subjected to onlyaJimited number of tests. Yet it should be remembered that in all cases B. mdgaris L. was used as a check, and in each instance the latter became in fected while the others did not.

TABT,E 3.-Genera and species of Berberidaceae which did not ru.~t as a r.,csult oj repeated artificial inGculations with sporidia of different varieties of Puccinia graminis and which nel'cr have been found rusted in nature

ci Unsuccessful Inocu .g Z lation trlal$ with .. • Hosts tested Pucclniagraminls :5l ~'" varieties ~g 0 .. l>. I--;---;---;--;---I~ ~ I e ~ <: .. l>. :5l ::I ill <: 0 Name Synonym <: >. .;~~~~~~ UJ ~~E&~!:~ --:------:,------I-- . No. ;.;:- No. No. No. ~ Nio 26 49 ~~r:::::~~k'._~:~~_::::='==:====:==:==~::::::::::::=:::::::: g :::: :::: :::: U:::r--3 68 B. edpworthlana schn______1B. brachvbotrU8 Edgew______18 ______18 ______147 159 t ~~'rir:l~~f~~=::::::::: -:B:iichitmitiiii(Schn:::B:;piiQltra- 1~ :::: :::: i g--d::::: Fedde. I 168 B. repens LindL______B. brtl1ipt8 Greene, B. nana 53 ______19 34 5 Greene, M. bretlipe& Rehd.• M. I reptns Don, O. aquifolium I Rydb., O. reptn& CierI. I ~'OI B. tbunbergll DC______B. chint7l&18 Fmnch., B. crttica 32 ____ 4 ____ 21 7 12 Thunb., B. rinemis Kocb, B. dnends angmtlfolia Mats., B. thunbergiana Schult.=B. vulgaris Thanb. I 202 I B. tbunbugll atropur- ______; ______, ______310 12 ____ 14 248 36 ____ _ pureaHort. :lOt B. tbunbergll maxImo- B. maximowiczii RegeL ______47 ____ 13 ____ 24 10 i 3 wlezll Schn. 205 B. tbunbertlll minor Rehd __ B. thunbergii dalo8onii Bean______i a ______3 ____ , 2 Caulopbyllum tbaUe- ______35 ____ 11 ____ 5 ]9,____ _ DlpbyDdae)'DI08atroIdea Mi~h". Michx______4 ____ 1 ____ 2 1 I,____ _ PodopbyUum~etrersonla dlpbyUapeJtatumPers ____L__ ------______221 ______. ____------____ ----____ 2 !:-----____ _

From the results obtained it would seem safe to conclude that at least the following species are resistant to stem rust: Berberis repens SUSCEPTIBILITY AND RESISTANCE OF BERBERIS 19

Lindl. (=]{. Tepens Don, O. repens Cockerell), and 1). thunbergii DC., with some of its horticultural varieties. It may also be safe, perhaps, to add to this group Oaulophyllum thalictl'oides Mich.'\:. (Blue cohosh), Diphylleia C'ljmosa Mich.,:. (umbrellaleaf), Jeifersonia diphyllaL. Pers. (twinleaf), and Podophyllum peltatum L. (mayapple). Although these plants have not been very extensively studied, the <:mtire absence of rust .in the limited number of observations and experiments made indicate immunity. Moreover, their rather distant taxonomic relationship to the genus Berberis makes it quite doubtful that they are' ataH susceptibleto P'uccinia graminis.

mSCUSSION AND CON CLUSIONS

Because 01 the impor tance of the barberry, not onJyas a carrier but also as a breeder of new and pos sibly dangerou'3 rust forms, a knowledge of the range of susceptible taxonomic en,. tities becomes absolutely necessary. One of thegreat A - ] est difficulties encountered is the multiplicity of syno nyms. Some of the species of Berberis are very poorly defined, and it is almost im possible to distinguish them without knowing their cul tural and possible genetic. origin. There are several garden forms, a consider ahle number of hybrids, and mll.ny closely related vari eties j most of these are uni formly classified under the genus Berberis, while some of themi\.re grouped inter-' 6 changeaUy either as Ber beris, 2"rttthonia, or Odos- FIOUlIE 3,-Modcrntc infwtion of Puccinia gramini& agrOllti. temon. In this bulletin the di3 on Berberi! brcvipaniculata: a, Pycnis; b, Ilccia. X 2 members of all these three genera are included under the genus Berberis. A knowled~e of the authority for the different nomenclatural designtttions IS of pttramount importttnce. Two or more plants entirely different morpholo&ically and physiologically may have been ~ri.Yen the sttme name by d.ifferent authors, e. g., Berberis vulgaris. There are at lettst three distinct plants described in literature under this name. Thus, B. '1xulgaris L. is the aceredited nttme for the common or European barberryj B~ 1lUlgaris Regel is a different species of barberry and is a synonym for B. amurensis Rupr.j and B. t"Ulgaris 20 TECHNICAL BULLETIN 300, U. S. DEPI'. OF AGRICULTURE

!"13 r '~. _

1 j

", . \

FIG uas 4.-Species of Berberis Infected with parasitic races of Puccinia gramini8. X 2. A.-Bu btris VIIlgaris infected with P. graminu pone. (Noto the long aecial cups.) B.-B. vulgaris Infected with P. gramillis agrostidis. (Note the relative large size of the cluster cup.) C.-B. $ieboldii Inft,<:ted with P. gramin/uualis. (Note l,'ngth of aeclal cups) SUS~EPTIBILITY A,;ND RESISTANCE OF BERBERIS 21 Thunb. is synonymous with, B. thunberfJ:ii DC. In view of the fact that the first two are exti'emely susceptlble to stem rust, and the last one immune from stem rust, the necessity for includin~ thes.uthor's name in the complete name of the plant described is obVlous. Unfor tunately, the authority is not always known, and for this reason the author's name is omitted ih a few cases cited in the tables of, this bulletin.

,0·

FIGUlIlJ G.-l'ycnitll infection on Berberi3 a.ialiro inoculated with Puccinia gra1lli7li. tritici. Tho, pycnia died, precluding tho formation of aecia. X 3, Informll,tion regarding the susceptibility of different barberries was obtained from a study of the literature on this subject, from extensive field observations and the examination of herbarium ma terial, and from numerous artificial inoculations in the greenhouse. Of the sb: races of stem rust prevalent on this continent, namely, Puccinia gramini.s tritici Eriks. and Renn., P. graminis avenae Eriks. and Renn., P. graminis secalis Eriks. and Renn., P. graminis poae Eriks. and Henn., P. graminis agrostidis Eriks., and P. graminis 22 'l'ECHNICAL BULLETIN 300, U. S. DEP'l'. OF AGRICULTURE phleipratensis (Eriks. and Benn.) Stak. and Piem., all except the last named produced normal infection at one time or another on most of the barberries tested. Not all the barbeny hosts that harbor stem rust are under all con ditions equally susceptible to tlllS fungus. FO.r instance, the aecia of P. graminis tritici and P. graminis avenae that developed on Berberis jremontii Torr. and B. trijoliolata Moric., respectively (fig. 2), and those of P. graminis agrostidi.s on B. brevipaniculata. ~Schn. (fig. 3) were considet' "-'1 ably smaller and readi ly distinguishabl~ from those produced by P. graminis poae and P. graminis .agrostidis on B. vulgaNs L., or by P. grmninis secalis on B. sieboldii Miq. (fig. 4), even when inoculated at the same time and cultured under the same conditions. TIlls would indicate that although species like B. jremontii and B. trifoU olata may be infected with different races of P.graminis, they are really only slightly con gemal,orin effect some what resistant, to the stem-rust fungus. In nature these species are seldomif ever found in fected with stem rust. Some other barberry species, like B. aemu lans Schn. 01' B. asiatica Roxb., thus far re sponded to artificial in oculations only to the . i, extent of the produc tion of pycnia which F1GUR.: fl.-Sprouts of Baberi.• vulvari. infected with Puccinia died prior to the for museduramini. by Stcali. hen \'Y showingInfection. distortion X 3i of stems,leaves, and nodes, nlatl'on of aeCl·a. (Fl'g. 5.) Still other species and Yarieties, such as B. canadensis Mill., B. declinata oX7JphyllaSclm., and B. umbellata 'Vall., as a rule, checked closely with the reaction of B. vulgaris L. (Figs. 6, 7, and 8.) Although in general those species that resemble the European bar berry are susceptible to stem rust, and others resembling the Japanese barberry are resistant, it should be borne in mind that there is no absolute correlation between taxonomic characters and rust reaction. For example, hybrids between Berberis vulgaris L. and B. thunbergii DC. that haye nearly all the taxonomic chul'l1cters of B. thunbergii may SUSCEPTIBILITY AND RESISTANCE OF BERBERIS 23 be rather'susceptible. In the same way B. neubertii Lem., which is a hybrid between B. ag:uifolium Pursh and B. m.iLgaris L. and has some of the characters of the Mahonia, exceeds the latter in its suscepti bility to rust.

FIGUu£ 7.-Individuallcn( o( Berberi! vulgar!a infected with Puccinia graminia stcalia. X 5 Particular attention should be called to the fact that the small trailing Mahonia. (B. repens Lind!.), so common in the mountains and foothills of the Western States in the barberry-eradication area, is evidently immune from stem rust. All attempts to infect it through artifici~ inoculations have failed. Neither has it been 24 'l'ECHl'IICAL BULLEUN 300, U. S. 1>EP'l'. OF AORICUV.PURE

found infected in nature. Bolley and Pritchard (2, p. 641) list B. repens among the species susceptible tC', Puccinia {}T'aminis, taking Saccardo (13) as their authority. But Saccardo, contrary to custom, states that B. l'epens Lind!. harbors Aecidium berberidis Gmel., and it is not at aU clear whether he had in mind P. graminis or some other

Fi!lt:RE S.-Whorl olienfiNS oC Berberis vulgari. InCected wil,h Puccinia gram/nis &teal/s. Note also aecia on stcm ·bo\·c. X 6 rust. To the write1'S' knowledge the.l~ is no other reference in literature that the small :Mahonia is susct:~tible to stem rust. TIle tall Mahonia (B. aquifolium Pursh) is at least moderately susceptible to stem rust. It has been infected in the greenhouse as a result of artificial inoculations and has been found rusted under natural con SUSCEPTIBILITY AND RESISTANCE OF BERBERIS 25 ditions. Futhermore, according to Doctor Stakman, there are several badly rusted specimens of this MSl.honia in the cryptogamic herbarium of the University of Berlin, Germany. Heavy infection on B. aqwijoli1tln, especially on the fruit, h~s been reported also from Czechoslovakia. Jaczewski (5) reported rusted tall Mahonia in Russia. SUMMARY Thel'e are slightly more than 100 established species, varieties, and hybrids of Berberis, regarding whose l'eaction to stem rust there is some definite knowledge. A number of these tf.£Xonomic entities have an average of three synonyms each. Some of the species of Berberis are rather ill defined: and it is almost i:I:npossible to dis tinguish them without knowing their cultural and genetic origin. In the present bulletin, plants that are sometimes classified as Mahonia or Odostemon are grouped under the genus Berberis. Different authors have in some cases applied the same name to entirely different plants. Frequently these plants differ not only in botanical characteristics but also in reaction to rust. It is therefore extremely important in studying and discussing barberry plants to have the authority appended to the scientific names under review. Information re~arding susceptibility was obtained from three sources: (1) ReSUlts of artificial inoculations in the greenhouse; (2) observations of natural infections in the field and examination of herbarium material; and (3) studies of the published reports of other illvestigll GOt'S. The artificial inoculations were made with sporidia of all the stenH~ust races known in North America, and ~ection was obtained with all except the timothy rust. Most of the field obser vations were made in the Arnold Arboretum at Harvard University and in Hi~bland PaTk, Rochester, N. Y. Exsiccati specimens were examined In various herbaria in this country and abroad. Apprmdmately 90 per cent of the accredited members within the genus Berberis, as recognized in this investigation, were susceptible in varying degrees to one or more of the parasitic races of P'Ucc-inia nram.i'm:s. The variation in sllsceptibility depended on the constancy of biotic and clinlatic factors, sllch as physiolpgic specialization of the rust pathogene, genetic nature and age of the barberry host, prevail ing atmospheric temperature, relative humidity, light intensity, etc. Only a dozen species and varieties of Berberis seem to be immune from stem rust. Some of these may, as It result of further and more extensive c:-..-perimentation, prove to be snsceptible. There seems to be little doubt, however, that B, thunbergii DC., and B. repens Lindl. are actually immune. In general, species resemblin~ the common barberry are susceptible to stem rust, and those resembling the .Tapanese barberry are reSIstant or immune. But there are some notable exceptions. It should, there fore, be borne in mind that there is no absolute correlation between gross morphological and taxonomic characters and rust reaction. This applies also to plllnts commonly referred to as Mahonia. vVhile B. repens Lindl., the small trailing Mahonia, evidently is immune from Puccinia gra:minis, the tall Mahonia, B. aquijolium, undoubtedly is susceptible, as proved both by artificial inoculations and direct observations. 26'l'ECHNICAL BULLETIN 300, U. S. DEptl'. 0]' AGRICULTURE

LITERATURE CITED (1) ARTHUR, J. C. 1920-1926. DICAEOMA POCULlFORME. In North American Flora, v. 7, p. 295-298, 775. (2) DOLLEY, B. L., and PRITCHARD, F. J. 1906. RUST PROBLEMS: FACTS, OBSERVATlONSAND THEORIES. POSSIBLE MEAN/! OF CONTROL. N. Dak. Agr. Bxpt. Sta. BuI. 68, p. [607] 672, illus. (3) BUTLER, E. J., and HAYMAN, J. M. 1906. INDIAN WHEAT RUSTS. India Dept. Agr. Mem., Bot. Ser. 1 (2) : 1-52, illus. (3a) DURRELL, L. W., and LUNGREN, E. A. 1926. BERBERIS FENDLERI, AN ALTERNAT1~ HOaT Ot' PUCCINIA GRAl\IlNIS TRITICI. Phytopathology 16: 23'1-235. (4) BOOKER, J. D., and JACKSON, B. D. 1893. BERBERIS. In Index Kewensis Plant-arum Phancrogamarum, t. 1, p. 292-293. Oxonii. (5) ,L\C7.EWSKI, A. VON 1910. aTUDIEN" UBER DAS VERHALTEN DEfI. SCHWAR7.ROSTES DES GETREIDEN IN RUSSLAND. Ztschr. Pfill.nzclilkrank•. 20: [321}-359, illUB. (6) KLEBAHN. H. 1904. DIE WIRTWECHSELNDEN ROSTPU:,ZE; VERSUCH EINER GESAMTDAR BTELLUNG IHRER BIOLOGIBCHEN Vl!JRHALTNIBBE. 447 p., illus. Berlin. (7) MELANDER, L. W., and CRAIGIE, J. H. 1927. NATURE OF RESISTANCE OF DERBERIS SPP. TO PUCCINIA GRAMINIS. Phytopathology 17: 95-114,. illus. . (8) PIPER, C. V. 1922. THE. IDENTIFICATION 01<' BERBERIS AQUIFOLIUl\[ AND BERBERIS REl'ENS. U. S. Nat!. Mus. Cuntrib. U. S. Natl. Herbarium 20; 437-451, illus. (9) PRAIN, D. 1913-1921. ODOSTEMON. In Index Kewensis Plantarum Pllanerogama rum Sup. 4: 164; 5: 178. Oxollii. (10) REHDER, A. 1914. BERBERIS. In Bailey, L. H., The Standard Cyclopeditl. of Hor,ti (11) culture, v. 1, p. 488-493. New York.

1916. MAHONIA. In Bailey, L. H'I The Standard Cyclopedia of Horti culture, v. 4, p. 1970-1971, illus. New York. (12) RYDBERG, P. A. 1917. FLORA OF THE ROCKY MOUNTAINS AND ADJACENT PLAINS, COLORADO, UTAH, WYOMING, IDAHO, MONTANA, SASKATCHEWAN,ALBERTA, AND NEIGHBORIN(' PARTS OF NEBRASKA, SOUTH DAKOTA, ·NORTH DAKOTA, AND BRITISH COLUMBIA. 1110 p. New York. (13) SACCARDO, P. A. 1898. BERBERIS. In his Sylloge Fungorum ••• v. 13, p. 174-177. Patavii. (14) SCIINEIDEH, C. K. 1905. DIE GATTUNG BERBERIS (EUBERBERIS). VORARBEITEN FUR EINE MONOGRAPHIE. BuI. Herbier Boissier (2) 5: 33-48, 133-148, (15) 391-403, 449-464, 655-670, 800-831. 1908. WEITERE BEITRAGE ZUR KENNTNIS DER GATTUNG BERBERIS (EUBER (16) ___ BERIS). BuI. Herbier Boissier (2) 8: 192-204, 258-266. 1916-1918. WEITERE BEITRAGE :/IUR KENNTNIS DER CHINESISCHEN ARTEN DER GATTUNG BERBERIS (EUDERBERIS). ,5sterr. Bot. Ztschr. 66: [313}-326, 1916; 67: 15-32, 135-146, 213-228, (17) 284-300, 1918. 1923. NOTES ON HYBRID BERBERIS AND SOME OTHER GARDEN FORMS. Jour. Arnold Arboretum 4: [193]-232. (18) SYDOW, P., and SYDOW, H. 1904. PUCCINlA. ORAMINIS. In Monographia Uredinearum ••• v. 1, p. 692-698. Lipsiae. (19) WATERHOUSE, W. L. 1921. ON TUE PRODUCTION IN AUSTRALIA OF THE AECIDlAL STAGE OF PUCCINIA GRAMINIS PERS. Roy. Soc. N. S. Wales, Jour. and Proc. 55: 278-288, illus. " '

ORGANIZATION. OFTHE..iNITED STATES- ))EPARTME~ ,OF AGRICULTUREo , -WBENTIiIS PUBIJCATION WAS LAST PlUNTEU ','

Secretary of Agriculture______-.__ - ARTHUR M. HYDE. AsBi8tant'Se'(:retary ______-:-'.cR W. ;DUNLAP.: . . D(redor of Scientific Work~------,:,.---..:,,--- A.F, W:OQDS, Diredor of Reg'l4latory Work __ --~-~_-- ~_~..:'" -wAJ.Tlffi G~ CAMPBELL. Director of &tension Work ______~ C. W.WA.BBURTON. Director of Pers,!nne~ and Business Adminis- W. W. STOCKBERGER• . trahan. Diredor of InJormation______M. S. EISENHOW.ER; Solicitor______,E. L. MARSHALL. Weather Bureati. ______"______CHARJ.ES F. MARVIN, Chief. BU1'eau of Animal Indu8try ______JOHN R. MOHL:Ell, Chief. ' Bureauo! Dairy Industry ______O. E. REED, Chief. Bur~au of Plant Industry ______WILLIAM A., TAYLOR, Chief. Forf;st Service_-----~~----- ______R. Y. STuART, Chief. Bureau of Chew-istry andSoils______,_ H. G. KNIGHT, Chief. Etmiau of Entomology______C. L. MARLATT, Chief. Bureau of Biological Survey ______PAUL G., REDINGTON, Chief. Bureau of Public Roads ______THOMAS H. MACDONALD, Chief. Bureau of Agricultural Engineering ______S. H. MCCRORY, Chief. Bureau of Agricultural Economics ______NILS A. OLSEN, Chief. Bureau of HmneEconomics ______LOUISE STANLEY, Chief. Plant Quarantine and Control Administration_ LEE A. STRONG, Chief. Grain Futures Administration ______J. W. T. DUVEL, Chief. Food and Drug Administration ______WALTER G. CAllPBELL, Director of Regulatory Work, in Charge. Office of &periment Statioll.8 ______JAMES T. JARDINE, Chief. Office of Cooperative Exle7lshn lI'ork ______Library______C. B. ,SMITH, Chief. CLAIDBEL R. BARNETT, Librarian.

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