#14 a eae iovagaes ats oa! nts hiote arate " othr ges (see ate Gitar n ‘Ae eA erat ma ait rately pe Fe terete neh ihe begs Seagate poe Arh tacs sate sie Maher ive ve hp it ne eth ost ‘a terreno bnrmiia pets arcane gre tear ethers pete. rife jah aay or £4 + reacts peor saan pelt “ys b bbe 4. hereon te" Crake worse af Dine rg. Pevnhomng abe ta p44 eanthiald eats eS neCy oars sheeely cl potions naan abet phir accent Set tare Ivrea gett eibe payabevd Sy 738 ieletat tS lpia teeta teat rere hed sis at pier oe none) 28 avinirs facet rid Wee ote ten ef aes by phe es boar pabaniadhs urate! phish geri Hoy Lo veda dex yoy apedagres May haope aee ae Breeae: yah oo y epuieaed ds bs: ferdies be tears apedt Uitree socpeedeet ) 4 ph eas ebb jreegai ne Pht A Avent yh pee arte peley ee Sppalan gaehif defing tye He eH gee spay er AA Fd pap be puis es tha ihe ab eT subsea) pha minbateapeutedrer iy See eieeha beds ipa edepe Farr amae ye TS Dying da a gtd etag vows ree aes hige phy Mpa) Pevedode fe iprge weary yr kiie vdehelecheaebcig: is at bebe pees hes tear eRe priest Jaen F jeiebehs hey Misponmine i: 4 mad eons > Pe nee { Daye te Pd eID re <9 My é App thea Hy bent Age pepe ine vet at Ariss ie) Pe erny aiage ea edrte Re nee er) avas avenge) 4 Amcnieeyey Part anadeetane Hand ett jbo atbemetie ry rdeds a rue) ag LE RUE Me peat gg phage ty Hh ooh deg ety peaeteiet 4p ; yhgeh mehr opie + eer jal begets ath pipers fer yew ats Sin saeQen oe paper Teta te ta aoa priya 20°98 ‘hanes 4 pagrus f Lp taeress yeahs). ¥ vere fe sasrapeese 49094 yep Orie ge Wart vee 4 ba poke ey Heme pupo oe eepey eet 10h fe Meee te eke + Hare ah ear daer ee mir’ plenee peSrecvsy Haat n4 meerdey ” , Pp ah pet aes 8 t awe she a yr cae + preoemts! vin a yen ee Ar aber Sponge pe feges trap ae neees | Tih vlies eriantes edi eater ae?) iisiateaneaes De erate Page My, Fh eer inecee droga ies te’ " wealitateraa tani nd paves’ BW tt hadi jal pues in pode saree Gebel mtg rye yyOee re ety Leng Wonreveemeels “ ay) cays parent way \ oo pape ee rem + SHuseeaiete + eh gps ‘ oe ean big be ging Simon aie ouacenn ah aedeangae- i aoenet Teclyrmedleeteeeh erbehe (are ees ae th walk steiseeneaten eis jitecits bey uth Fete’ br eae ster syeegniran al his At, deb bnit HbA peed band FRG £Balhethetsiail rb: eathreatie ge) 21 eaters est eae oe eet atbes ital theory ou pee raprart Hey Wade leis sae pig y Te Te dined Pafrab’: bnahdrve ne od ae emg perk wee 4rhe~ oad eho Sp hhtte {raise tA iva or eee pita: ananal arcs op papepaeingenaer it) Hanes aay “ye Sak tie ue leh ran sentir vb borte4 Malieehatsigeeene een Weriee vara pus miersorbates tata vaae seta vents oatoety eens ae daha) oa min ieee beet sph jonetie rae epi aapages: orcarpebgeeerurhel (ay | Sched peksnmerievee peers tea are ae jyaedey eae etna te peli beet os mek cAtpeed Tee ai shed ae petear! ytBoya) ‘ sienna be irae fer pat tie St RA pe Mae Sap phyreend ay ; 2 ahr oe Syregee Hi br erasie Seep Ppteaterats et $5.)ayese Pot bean ye ad EB! sSetsaset ose rei litvcuce henedtacseaspen DreTare raiesb be Teche Ler deg 1 oye oa “9542 Tistbotet set Pireten tie Pan arsenal penis stl Up EH foetus rik tug snsesetaniae ele ptm Msnraee aad Hisirast t4 ne bet elathsaanh ane a + bettie de PL b yeah pet thon held aed iat pages dower ute ene op Abepea one. phe let Cin Picante eect rerute ye po) ae bh aise mentee arte ToL EEt dalney-aeduee ene 4 ae seth dee aaa r iyvevar ata peonika by ge Braye ort tow da lebe aA rol letaeded een cas caens + pit ch ete fd LA ripltacr pq Up Bak doan a befbeege fo pr de y er, eit shi ae: es vt He Hs erate rte: tr alle hipeanne refi rain feube inane hats re ayy otavacbier oy CAMP arret Pye t Doub htcdet ade bets eames be Aeibalye ant + goiyn de ures ear toa lp by Wa08 LRire Avda phe ga aed 1 S79 Nh vers pvp) FRE cAege Chase poy eee Wrwadebeerare sane gt) sive Pebelan dep ie B297 be ODED dees BFP ih Shes petaittiee pe gan) ibeapa de yenry » jah eb " f ayehe evo} atihe ds Sala t Wt oder Avg epidt ye drpete dar arias Fell be Pad Sens hile reseeyte ae i 7” dey se: Pbedoea pe taped p20 dedade bailey pres peat ot eT TT LES pa uhL beak poataons Hien OTe Ttd i ds bipdatausid mab ees es raga 34 Be faeries Jha ipods Fibrtyeg tbe papas Ket? Lepr ag air pobdie pal sped TT baa kl here haa ath yh rif gg se edrsa doda rte eso peaetedte;) > Raced pie ees A Pabst fea ipebetean cue St delthes 41090 Feld Ac reoihed aebales fib SoU 44 vedas ashe orale yf ne pts rb seg ete FH Tens gew sepsy4 eve inp eet eat apes aa] = are peso} 388 yay hase Poy - Pri MRL a ae Higa roe Pains bei irate aby viele ie sagd: yay. selhepy eater Poteet Ad syess tobe gapepr ented ara hebl sna Ted 4 of pabeh a ned ie derive Heh bae rear ibed #4 Heacl dreds deds Hadni« pepede be bran Rast PU Werte ALP etal sete ahs re bbey apy peg rierll tae 1 i prifeai a briees bid aqeahsae cote holed Yinhah aged acpnebocnn git eerie ter’ Crit istics) abe yah Het: Vaan? ’ we Nd Meets 858 areas VOM basse dn 1oyie hs PRE DS pri see daw ss Tigre ae Bite ay fo grag eles UE imeer ert ays Te eB “ys ‘ ‘ 1420 en = Hehe 4 erin renee y) weed tiateh / } ss dedean ered opeeaueded pede ded de petaet picysnes rage OF) Fi gepe rhey oie ay ¥ 4 few tertart anions ache Agiae tit be Hadbedt ory iye #44 A eae te pe eraginety: prtegtards 4/79 oe rhe egos et say iageg serra sheds burt eae leW ) leete bbe pin eeew dylan fa finda aerive s rbamtbatey Hn aro tate) acd Abaesdrenbe see Peebrécieacherteshd a Aap oae ee sisteuet eateite SReenCaRaea par weg Oe naan ae tw On Fay ane aehetnecetoert tis Aik peel ot ae oh 1 RaaL ey Boe dis Mrpran cay Phe yen mee apn Se stechanaimre1e ie} Armee’ paecbyecernertnt ei na dejeda se paantast ad ietcteb abt Pier te alet acest ered eR toy Gata Ne LE grt Pine Pape BATS totes pe ehe Ne teaymy AP UAE AA (gt abs O2b ds Mote ret o-N Hat Oy 0 06) Pers ee ae be badetiebehesens hep: Hedeve dal: g's pede ae sateaboG 24 Depayenaye rot oh per kb od ae igtep-wudbove nepebeyerorel Ppt (rom gp WELLS LENE L LHe HARA Boner hea gh Ae Thar hpi te kets eblamiigtuahss testedey bat Pye pan an a mht op nents ott ed dre iaetet re pee ree io phates ad tiple seets srietetele sets tA paw ateh Bae Pheer emer ye ba ph aretg tent Mende sel br itasd:eryr db abanpeenebthedal Se dye EATER OL pepe TED Oe eet BALE eh a ip yg er urere ey aL durante ee tases bp aeiet res Pubmed 8 sbi tah punter t iar hitataltamnnbebaeprinmn spssace gees hgh Puen wae pat they Neb an GPA re" ee Pete be mee: be ee Wo ba gr lbw pe than ei eee ie pike a sett pe aie geaie edd (eet ribs 0 OF) stair berasbeseironts poritiinabe itary rly ben pe iy pe Sea se stand Heer eae tet be valbtihe belie ya tenet sate need he nti" tii raae 44 + Fi 8 vibes be ibeibh an) efi “tt 348 pods 1084 tela: brsebidn sree, pop r hetero ede Rebs g) eecntaee Peete Pe tae ESTOS oT eaesrd 4 jecbereocaaraesemeeeaed veep +#- Gabi metvbortsre Poi preety nan pron liye’ Mowers one a 0-004 eee AE bE | epee dhe pha ed shed aree PS phone es PLAGE LR VTLEN hae g a pacsenpt-oorse Lig lyr 7 qe phat MPeinan irae Bois + bh ptet Wee teten WRAL Whe eng Memeo hb thee neers Papeaphor fe rete genet raed bee rp Levtete Los nent bapahs 4 ’ rs LS ayes enh Pt aR Le sale Lee ped Fibs rye re fy pt Pie areca Na ale rLebeksiah pe pheeeentey Mp rte ar hrieesean eat ie hates bipkp bntartrsete th ypedtpe Pdiuntenehy beamed mee More ehd Lenore bg Ah Fig ah pan Dipwg i te aU rie thei eks ea Sea Nd pee PONT OLE TE PRESET peeves Pin venakaleebasrer teri ty tte eh 0k pee ark dy 8 pean UE REN Bh BHOERE Wh 0k Poth reel avrtanrnt wlohe hep PRED I bs poy t Sgn PEEP Pst tbe b Let phe Nate A emebrpenrrars ese pear eeorersseirerrnarre- user LPN e Loa vy veh FAeb byetgie rms song by tre Ley ene hppe ree eae urer bes apd Rtg bb se per ereeie aig he vee be Ww Dag ne ae has PPE PR ve La rn Ph Hah peat Py renee ng SP LPN PA Pomel Uo Aa ch poet Ab ahr oo hp pe heheh id bo Ler nn ene piakek hatlpe Carsetches meshed tat fed Merperereysrsy sors ense: serrriepone fet Tehri ery drier ires., PEELE RUPE be ah ae rN pevene pete oe int oy oot Piatenre Timeet ern pa tr phar bebe PP AES pa atk AAP VRP HE Ph ah ae rm aire) rch ho lg Mowied bodies siehedbdsbabeas ieties * 1h ¢ Poe Heer tet ete v4 eve 4 oy ra Woetedtseacten eae she epiny me hothe bodeveng /9ps bata Wetprine: er bh vergplarece Pe eka been ae Ad PEAS Peabo omer Aah res Ay SONOR eet ln tag : vaprperee 7207 tae os yl ed mpelite bavegen eee te tea ry fale lint shed pea $4 poirot Aa py eer hat Lpmehrurs Peet ow SOR iE renee he pee merry tee hy! LALA WAL be dpe tide an propane phe cop muatlias ehengs-s ore bre Hpupeapeneieiba Pen bana gn PRUs ed (reels eraphenphghe op Keere red Pilla use UBL ups anata pet nia peg: a 44th pop eras se ageegret™ mbes wt arytheg bee othr bee Aa A poevet 4 ana bet ar ot nae Lag ehteee iehband eaaneet Seatacart fe “a veh ae } one ». pita eee ee hancanneteiat PO? apene Lele HUGE ete ta CEPT tebe Oe a heed banda eR Ue De bathe Fere Hr 1 CAGE pL EPL we paps lek tree: a hrs et +R Ne WA eA APE CMIELY IVE Neon Or He © Prete PNAS OC EE . Lids wad npnca dad ebl at tanh url pheu/t itty olagdichymatyeebtra: bateeterees foe push were puarbmaeauenatadinnee detain teeta Nas ictal en ete lahesote pobicet pee ath sieabesesa Hoe bone t PRAGA A Pah MeL P LL EL TY Dad OPeh semrbengne gree r mE TY heme hoes buibesenetredeidedtiebenereints saps Papeireivetopeirens Pspe¥ i wadee Beene 1 het, ee beter: Maneatis bel sieelnle lene urtsirelrie rtrd ine woe ant rie iitaseceatibe bahteal voc ieareste init ips poor edadebe peste eres CiersHasieleiy rilh apa phe dobe ded rentgs he jeaeds Pl pines ance ba eongenrasies oodaheannsse hverabe rbarst vob emds oT TIE Tn ead pt re iseeael V epiariey svasuehenweasres aaivecdeies reat) ipeeeenye ite Lie ae Paris jules hee roam dakar Lad apse pat phat ibh sie Mane nete tb es nds oe aie bee greeter pe Peevaraavectaivertiect me vane pepraegeee Bete pee hee local aeet bipedal Mat Berets. “potatoes eoeies ve 4 rh wished ricpeieatt Linder ie lye ke ert saeharped terenieas Ab isininbeas teh apeye me seaoibgs Papo ead ty pees: ir She tants ek ed lade ye bette iban se othe opine) hs ry Ae aiid ser belies vin ape pet heer reese eis ie te laeas ee 4 Pb aed curses vant apes spletett ere ea ibe snk ae De eet paket we ade: yreer ae verey ih ak eee bos ort br98 ees tery enedtas hsobhepawrenaves ate faredt fale: Pe nuda pore pee 3 te tiie peprbeee oe NPP tabs Ae t ors totenie bovesethn M tak 41 ved yegidets yen attra nh ls peat tytn tate ing ad Aves aa jer Devoe ie Bed bk ced oe pee a sore iney het eho Moen! Epbenel.et Hate where ieee syegreas ie )ethers tat mide sith r ata aaHdsh fae o he Ane yo pdeget eval rtoys or) sak he Hf iearsete a stair prety ieee eg rine-t4 (4 Fe) ; bel cape teats ede gt tingen is oles mae gran se ladioh rape soaker Pe pity real eet Pee tt EE seal balhce bathe ahs Megdds CtOM Sr rrror et ein hee ead aati bd at we ebser pel Bed h hay aan hy bape eta ee ot ge 4 A496 ed ad~ led btre ete dt} mda de pe jeriedeibro rit det teave ty vite repre SHPO D parade dap epeed sped ee rel eet et atl beh Ot4e)eh 2 br cred elpt tavepe + Hi boaegot Bode pwdetews tooe he Mister sabre Outed anni PT eaLh wet oud ere 0 Tere Hey aertet 4 bier Hi aie Ehyagh rueeit " Oe ft aria etehetet Sher Hie pear.) ats Se eT iechiee sae aaean irs Neepae bao pedegeah ody a rat ved! ‘nip in Degado eb QRaTAVHN) iso) yamine ) ¢' joes agent gard it = Poss bo v4 88 be nthe bogpt OP pon ¥ Winer, tn peed ig ab! Havin hog habe tintie\ eb: pete riety tt eee ke st kee tll el Nt rte gt thy poe eeedent OF ein Joie fete Sharh = 9 eae he a8 oes Fah asthe nieaagtd sande hort wish read dane geie aieirarnait ected : at ati poelye rebar TH PRM epi it ANGRH OT Fyre Si erilerete bo 90% fitace prac nine aoe aie Boyar sis hs jablordrbs eine Chlotieat ta aE LSA AOE ret ebay alles ty Ay AQHA pOHEY ge Dey L, ese barbie ws tae Wis eae tty) defedenat finlea ks sr fis4 preyed Wane heee ini ah ahs ye berets ya rte hiner ds ve bb tal hed & : Yeager ts gs8 ae vit? VEEN leg ede FAs gehigede } Ardem be Psteehee eerie a y sega Pete peor se bry st aly irs et Thpepeae ged ys ispeas nue boa Dobe We bry pei 92 Poses hy roi hed Hind Fh Lites St eeas gh a8 ew tael ip uide seks grsete br) Pol iele 4 ped Byir ay} pane PST TAP ea gale mbi RT eI Pea ta: Sey Bal els Wear hev sures Bete abi tb yisnre te nya lone tNy ply (s Bet . yer ered potaihs faire = ye Dipeddve pe. ay ap ase’ debe dpege Fett i bene eet i be rash P pies iret Heavens 1+) eiteae 1 te ay | agegnety ede besos te ie ene PBs ae isan be Pe oat ral a pajeye penal . btaleeadad ea pts Pi ADRS oang et Me PPTerr ny mst babi heel wae ih halal atid bese e | eae tds a Rataa Ral DAh98 g- bode hana pel sitet Dept Ds Tie ajeaeg bere a age penel cued ein Ved eos pebehs Pi TW Fe pobage! Qiena yt joteoe paged minty faa Bin te + ore tejeasy 4 traeaihe fouasedey chy tetas peer siitt tt ss lolaaite We tips Pegernen an a9 4 Lee : joke vd ta Mae Aab ae het i sae ye oye srtnats a8 HE 4 air iart LL phabshadeoas pane yore Heirs bed rit 4 sae") a) eye fo Een prtq od p dew erereers Pap etate As tee $e! reiieoedy peewee > Pte htee Seyeqensene near + yeah witb ovary rap eibat jie me dieys deeds pny Hoty fred ey ih wensaiiwe Lik phaebiche bead Al NG Tenney seppererenv red Di tuis5 soe pre sare asy dior prs pEL eR Spay © snp arte ‘ See W es } et edew Feist FeGc ory are! pe ahiewt ¥ 4n@egngad) pei obeys ines ihe hetiet sere anasionet ee 20 gee tae darpribeigdeoas reeerte peyrermatryaien igety Srptara ts yews 443~ , geHervndrdete piney erty Heike drs vi Weis Wh rar ebes ety P21 be pear gene rthedee t merrere lire yar hae sone fe paed reget pty tb esbebet ten aepeie st she pee siiee gener rated Shuai Sepa ae dedear a ieee Nirea yeaa gis) sind jonwes ver sy newrrr a4 enters wee SOPOT OE Woe! 8 youre a wer wen Atl aed Pretend i eye 29 N1 oOhE hehehe igen’ Pee deer vomeunbow ove yds hope gna " VeQeed tp eet ana Seyret patna otha ery: rermnpepe ares grays prom eomn os ve paps ran oad eee PH lrg asa? ataiap eel ae ke Folge bob om ap Sipe WW Hrg ts Bhd sae ge ary Ve pry do attr yey Pr Ambe tears Podte ¥ DALI oF 09% p99! 00% meV aadon gd! PLOT BIRT OLB IeS Ga Ree es 14 Peter or rear ts armed Pores branes vader to wh Ar ee wren Bi athgewiteey EPO RL LAGARA seeerqungeat fv detere areere’t phn ie yankees ites cp ebepolar se reperg ar eit 47 4 ON Ta ebene poy Ca HED be FOU: hiro Scat ESL he nie hbereeee Hye yiieees oate ROT epee aig rks 7 Deen ETS TF yaeyatth peat pat vee ee: Shiasat eal a eneb anor phe re wt ast Le (tart ol niall abatement” aneN rent a! es ohe thar ee et es DRE Te 9erg Atm Prolesel pasptreniny shee! Teka gh ges ed Hye et gi tehdip ener Vong haces enaae. fee ig aN ashrebirasbenetied Wk, pet me erp tohenpionek my ost Mark gee He chotene elute! Oa eed cree rteteg when beim ss Maree phate poet + Laguebe igh ct ge teeth oe This ater Arheertaheh ge honan Pee rhe ree: Thtterrss, Haustrenuen pretty t ee rune dhe tetbes ph reper ne eer mat Po wap ort Mae ah oe g otha pay el pk Rack An phe wid te Aaa ating Ceeaha yp veut hes waa oyeh taytborme’ bacha bene HH Soke pacstune teheve haste) etree pencreren= ty hr durichrunaleeenuedian chgaigag! Poprbs speatesgey ® sh cick nptetatehrteeie sel gittcteket onan treet resbte bs ipebe = ert hen syesannarrey openers Sey eh shy lhec aod oaae Mesgnenel e acer fos Bes 174-05 ve brazen rasaee Wels eghedpeotes Vises ctecasepete Sarah Mahe at ataent(t i eal aoe bx peifren poihetebtrite ¥* Fet-ynpops ie fets0-0eee 2-7 B- mee pepet ie pvient-¥ar I:7 se aeretne- be8 tae bale (ee whet ‘abe tbte te) raoueeaian sit tot seit me s pronsv tere 38 arabe utele yoreseo tre sterrereehy iit 4 seh lp tale erties rere aren: brett ree ws se at bop pom pe bee nul I Dito ire a ms Wiphirmmeaphean! ribet aneset ror eds aaet tis or i Sie pout Peete pea pre Pay tes pslrosteinttra er aite oom we info eben maggie ean b pret amet v2 est ve aye oe senda 8 Slee gstares eredes Peoepe Ne hw rry rep luses aig laste pyar wont bel a eee we Oe Sere pee eee rn hat ble eet ohne jetabes ts! rae Ye be yw te sl sere May dl Persia Vriaeye ” nity sid ong oak eH ot at A lap Peg cah 240 $d ban ith gh is dshtondonpts tenes oct see Avan Teme ME Jr Deere he ye pegets iA reire 2 pee pring pobotenit Petes pring fe be ann 9 ne prey Soph ith be rosea oes fants Fehasssre seen get VI rips 2688 wen sad! we ihe e+ honene Wreereer tata Apher peri) Hrean sian deaqTige reve rows esyty Va peredeveyr eee w seated a jim peony oe pe # a PLApr ele nd om eater eesti te Sv phe genrt sae ceded toe nd I ps 4 patphinns vie Sheeatiepeers. Lampe bags i 23 10) OES 1008 e098 jarhe = ) fret babes. paige pede We Fedde ye seri eayy Aoreaya ede tent, pairs peyrete pre tarde req ona mee —eh ons tP rt) pt Onna retina pe te be pnye y lin pre atl rey pritones oat pees e pei pee ry “yay toy ict ee ” rppt nas eoaay peas dine huires 7? rhatinhteites ; soqee oie Tage iegege nm seveerh: + Peres mating ees Pgpepre /° ae. rasp on ypeat ji’ boone berss* (eas abenee TDi TT ake ne reTSe tte HF HITT desert Aree? preter et 0 toe pA he ge yeye ved eee 2G) eet NT yr psy apse reed 40). bebmwere? ye Sepe at 901 oe vannyr Dept 4 wane 1 seaeytye HART age pees at yey! Ny wire ier , * pecegeer pone ¥ ofere sagrecgege me" Pare? nL aLabdniene inet ye pow Ee POS a OF Srrettiat inate tn re at each A trveess ra gage yi: erar eat rretee ome atenetyn rnd ik hplaabe stbees a byotee 7h a) seprirangninanitae: Hate eheeinate rir jenureed reer pA sewegngt weir yee Hasse ra aah toe * aie wun geet wt oe ine #4) sh derbi 98 rie er syearuedtt le wee Mp aririny hie anpeg eri rgagi ver Sah: areas a obra Vote re att pbtpe ye i Habhenctetehis-ceee os Weacd per rae ser eet aes Ai sewer Sarat tit tor) per avecntansund POET ot see a dae Malet: remipeasategnyet mae 5 De PT0 ey Weer Aegan pean oe neagary: aTeOTy ye gege ba Peet tert hp esive nears hi eeiaprsereth basen oes ees oe es te pits +) pee Arie gwd es bY rarer ie) sine be aa tatuver the niet ovina Hh 8 peed LVS ger Fatt aeacesneaa: bepepens yeu ns pet ee esl ae abies tio hat 499 oy (Sinks Sy Eee ferret ae 67 aif 4a As OTT LT ans er aiaaeiny wigan iaia en .wiaie f 8s es ae Pee 6: geputate seb greatest iri a nas ria taed OEP UT Sena avy aoa s8 228 it eA la vasanavatriatsrers pai yi! get latins ai i eee) ae esa TL EW t fy Pure errs hl ie niet de bied eaebs CiaAbitt poddihheh ehh Lage pepe one a. cert np lar ayH eR WAM DATTA Tes ip ybe we By exchanee 4 = 1918. af . Go —= 3 ma G Pa roan Than tre . R WeCibson- Inve =a 14s al pt) ; FA far A , | i } as exer < a £ . HRS ia “ . * Ti ya 7 . ~ ‘ < ve * : . t4 ~ Ke se & “ + “ 4 a - r : g " » aa Y ; ayo be ~~ a | y ‘ ‘ . " - ~ n-th om ; vas 2 < ~ rl 7 . mt I , 1918 ¥ NIC G J t 1 “ " A rden LUME TION PAPERS a" 3 ? - f si oh: rh e ISSUED JULY 6 | Z ie : ~ BROOKLYN BOTANIC GARDEN MEMOIRS VOLUME | DEDICATION PAPERS SCIENTIFIC PAPERS PRESENTED AT THE DEDICATION OF THE - LABORATORY BUILDING AND PLANT HOUSES APRIL 19-21, 1917 wee) wile | ff & Anam ool Wo cl | | —1-4- owt — bert tT AM l\nnnmme Deere Sh ett LIBRARY NEW YORE BUTANICAL GA i | ISSUED JULY 6, 1918 BROOKLYN, N.Y., UcSvA. i _ PRESS OF THE NEW ERA PRINTING COMPANY LANCASTER, PA. ; PREFACE The papers contained in this volume were presented on April 20-— 21, 1917, either in person or by title, at the Scientific Program, which formed part of the Dedication Exercises on the occasion of the com- pletion of the Laboratory Building and Plant Houses of the Brook- lyn Botanic Garden. ORLAND E. WHITE, EpGAR W. OLIVE, ALFRED GUNDERSEN, Publication Committee iti SS ee F CONTENTS PAGE PERINSON, GEO; FE. The genus Endogoné. .. ius) s.0. 0 220. ne: I BLAKESLEE, A. F., and B. T. AvErY, Jr. A vegetative reversion ROT LOCUS ie che ib, OT ahs od Re SA ae oe 18 Britton, N. L. The flora of the American Virgin Islands..... 19 Burns, GEORGE P. Weather conditions and plant development. 119 Cook, Met T. Modern applications of botany . Sat AEDS DoncE, B.O. Studies in he genie Commnesnoren enya Nee on the distribution of the mycelium, buffer cells, and the Sermimatiom of the aecidiospore... ... 2... 6 sce eee ene ee 128 East, E. M. Intercrosses between self-sterile plants.......... 141 Harper, R. A. Binary fission and surface tension in the de: Teopmentor. the colony in. Volvox:.:. 42: posse ae 154 Harris, J. ARTHUR. Further studies on the interrelationship of morphological and physiological characters in seedlings of JE OS CUS 5 ae EE es re, 8 Ae 167 HARSHBERGER, JOHN W. American heaths and pine heaths.... 175 Hotiick, ARTHUR. Some botanical problems that paleobotany MaceienmeGit@.SONVE..... su da).4.0 He ntitege co aay eee ere eee ee 187 Howe, MarsHaty A. Further notes on the structural dimorph- ism of sexual and tetrasporic plants in the genus Galaxaura.. 191 meson ies: The, Uredinales of Oregon... .222 age see ee 198 JEFFREY, Epwarp C. Evolution by hybridization............ 298 KuNKEL, L. O. A method of obtaining abundant aaa in cultures of Macrosporium solani E. & M. eS oe ~ 306 MACFARLANE, JOHN MUIRHEAD. Synicheat an in pies structures 313 Metcatr, HAvEN. The problem of the imported plant disease as illustrated by the white pine: blister, rust.. ....2--.2)- “ane 4 227, MurriLt, WiLt1AM A. The rosy-spored A garics of North America 334 OLivE, EpGArR W. The cytological structure of Botryorhiza Hip- PO GTONCO Crane cag Rte ah ht a0 a Yb Dt a 5 38 3 an GR a ee eo 337 OstERHOUT, W. J. V. The nucleus as a center of oxidation acs 342 REED, GEORGE M. Physiological specialization of parasitic FUNG ae See circa OR us 5 :n wivtipgl an tute eee eas 348 Row.eE, W. W. Relation of marl ponds and peat bogs. . . 410 SHEAR, C.L. Pathological problems in the distribution of pecate able plant. PrOWuUCUSy wes. g- sk< cao. Cae ever ure ave nate 415 Stewart, F.C. Tubers within tubers of Solanum tuberosum... 423 ee Vi CONTENTS SHULL, GEORGE Harrison. The duplication of a leaf-lobe factor in@enhe shepiterd.s-pUEse j.'s. e.0e Ja ee eee eee eee 427 SINNOTT, EDMUND W. Isolation and specific change.......... 444 SmitH, Erwin F. .The relations of crown-gall to other over- SLOWENS tm plants. 2+. ea ee ee ee 448 Srone, GEORGE =, Contact stimulation. =... .9 40.) eee 454 Strout, A. B. Duplication and cohesion in the main axis in CLEMOT UI TMV OUS 7s bys «RAR oe Se 480 TayLor, NorMAN. A quantitative study of Raunkiaer’s growth- forms as illustrated by the 400 commonest species of Long slamel IN eVect i eer erent Gn ba ie ee 486 TRELEASE, WILLIAM... Dhe-ancient oaks) ot America. .2 2 oeeer 492 TRruE, RopNEy H., and HArvey, R. B. The absorption of cal- cium: salts by«squasi, seed linc, ar deter ie: ate 502 WHITE, ORLAND E. Inheritance studies on castor beans....... 51 THE GENUS ENDOGONE GEO. F. ATKINSON Cornell University The genus Endogone was founded by Link in 1809, and for more than a century its life history and taxonomic relationship have re- mained very obscure. Notwithstanding this obscurity in relationship and development, the structure of the mature plants is so simple and characteristic, that comparatively few species have been accredited to the genus which do not belong here. The fruit bodies are pulvinate, rounded to reniform or irregular. In size they vary from 2-4 mm. up to 2-3cm. In life relation they are saprophytes. In habitat they are hypogeous, epigeous or epixylous; and occur under or on the ground, among or underneath fallen leaves, among the rhizoids of mosses or roots of ferns, on dead wood, on sphagnum or other mosses. In color they are whitish to yellow, brown or blackish. Approximately seventeen species are known. The genus is cosmopolitan in distri- bution. Species are known from Europe (including European Russia), Ceylon, Tasmania, North America, Central America, South America and the Fiji Islands. General structure of the fruit body.—There is a peridium of slender, interwoven hyphae formed by the terminal branchlets of the hyphae which spring from the basal region, branch profusely and radiate to the periphery. The interior constitutes the “‘gleba,’’ the base or central portion of the ‘‘gleba’’ is sometimes hollow or of less density than the broad peripheral zone. The “gleba”’ is usually packed with numerous, large thick walled “resting spores,’’ oval, elliptical, or spherical in form, and yellowish, grayish or dark brown in color according to the species. These ‘‘resting spores’? are 40-100 w in diameter, are packed among the hyphae, and terminate certain branches. They have been termed spores, sporangia, ampullae, or asci, the latter term apparently having the preference, since it is employed by a majority of writers. Several large elliptical spores in an ‘‘ascus’”’ have been described in one species (Endogone pisiformis), a single large spore! in an ascus in another species (EZ. reniformis), and numerous minute spores in an ascus in several species. It is ’ 1 The spores in this species are probably the ordinary ‘resting spores,’’ the ‘“‘asci’’ of authors, for Bresadola (1896, p. 297) says that neither the asci nor their mode of dehiscence was seen. 2 1 to BROOKLYN BOTANIC GARDEN MEMOIRS doubtful, however, if true sporulation has ever been observed. In dead resting spores the contents often segregate into a number of large, rounded or elliptical bodies. In E. reniformis the single spore in an “‘ascus’’ is probably the ‘“‘resting spore’’ itself, while in the species described as having numerous minute sporidia in an ascus (often with an interrogation), the supposed sporidia are probably the numerous fat bodies conspicuous in some species. ORIGIN AND DEVELOPMENT OF THE ‘‘ RESTING SPORES’’ IN ENDOGONE LACTIFLUA BERK. Character of the mycelium.—The only important contribution to development in the genus Endogone, thus far, was made by Bucholtz in 1912, in his study of Endogone lactiflua. This species is subter- ranean and occurs in various parts of Europe. The fruit bodies range in size from 4 mm. to 2 cm. The material studied by Bucholtz was found in a plantation of Abies sibirica in Livland, Russia. The mycelium is coenocytic, profusely branched, the hyphae following a very sinuous course, but the general direction is radial and toward the periphery. It is non-septate, true cross walls being formed only in connection with the reproductive organs, though false cross walls are occasionally found in the vegetative hyphae. The hyphae vary greatly in diameter, in general becoming more slender toward the periphery of the fruit body, but irregularities in the form of swellings occur. Terminal branches on the interior are often clavate. Sack- like enlargements occur from which numerous branches often diverge. The walls of the hyphae are thick. The nuclei are minute, very numerous, and lie in a parietal layer of cytoplasm. The progametes and conjugation.—The progametes are clavate and of unequal size. They lie nearly parallel and their walls fuse at the lateral point of contact a short distance from the free ends. The nuclei in the progametes, lying near the periphery become larger, more distinct, and undergo one division. The nuclei now retreat from the terminal portion of each progamete and all except one are excluded from each gametangium by a cross wall. A few sterile nuclei some- times remain in the gametangium but soon degenerate. The origin of the selected gamete nucleus in each gametange is not known, but it is probably derived from one of the peripheral nuclei in the pro- gamete, or one of the daughter nuclei after mitosis. These gamete nuclei are much larger than the progamete nuclei, thus maintaining the nucleo-cytoplasmic relation. Each one occupies the center of its gametange. The walls at the: point of contact are now resorbed. The nucleus from the smaller gametange (antherid) migrates into the larger one (oogone). ATKINSON: THE GENUS ENDOGONE 3 Formation of the ‘resting spore’’ and simple zygocarp.—The “‘resting spore”’ in Endogone lactiflua is not formed in the immediate zygote resulting from the fusion of the two gametangia nor in the oogone, but in an outgrowth from the latter. During and immediately after fusion of the two gametangia their walls become thickened and firm, so that they can not yield to the pressure from the young growing zygote. Asa result there arises a sack-like outgrowth from the end of the oogone into which the cytoplasm from the two gametangia flows accompanied by the two gamete nuclei, the antheridial nucleus fol- lowing the oogonial nucleus. The sac-like outgrowth enlarges into an oval or broadly elliptical resting zygote. A thick, stratified, hyaline, cartilaginous wall is formed next to the primary zygote membrane, which entirely encloses the cytoplasm and other contents, thus separating them from the empty oogone. The two gamete nuclei lie side by side in the center of the zygote but do not fuse until after the resting period, except in a small variety from Germany in which the gamete nuclei fuse at once according to Bucholtz. During the growth and ripening of the zygote it becomes enveloped by slender branches which coil in a more or less spiral manner around it forming a thick cover of small cells, 2-3 cell layers deep, the walls of these cells become greatly thickened and fuse next the zygote, grading out to the thin walls of the surface. Each resting spore, or zygote, with its individual cellular envelope forms a simple fruit or simple zygocarp (zygosporocarp, as Bucholtz terms it). The fruit body of Endogone lactiflua is filled with these simple zygocarps intermingled with the mycelium, and is therefore a compound zygocarp. Germination of the ‘“‘resting spores’? has not been observed. Endogone lactiflua is the only species of the genus in which such simple zygocarps are known, 1. €.,a fruit body with a single zygote enclosed in its individual envelope. ENDOGONE SPHAGNOPHILA In July, 1916, a day or two before the close of a fungus foray organized by Mr. F. C. Stewart at his camp on Seventh Lake, in the Adirondacks, Mr. W. H. Sawyer, Jr., a member of the party, brought in some sphagnum on which were rounded, pulvinate, orange-yellow bodies resembling the plasmodiocarps of certain slime molds. A pre- liminary examination of the internal structure revealed the fact that it was not a slime mold, but apparently a phycomycete with large resting spores having a thick, hyaline, stratified, cartilaginous wall, and orange-yellow content. A pair of stalks, or suspensors, attached to one end of the resting spores in different stages of development, indicated that they had their origin in an interesting type of conjuga- tion. In the afternoon of the same day (July 31, 1916) Mr. Sawyer 4+ BROOKLYN BOTANIC GARDEN MEMOIRS and I crossed the lake and visited the same spot in order to collect more material. The dry weather during the latter part of July had lowered the water in the ravines so that this particular sphagnum moor was water-free although the ground was very soft and wet. The fruit bodies of the fungus were not very abundant, but here and there a single one was found on a sphagnum plant, rarely two or more. In nearly all cases the fruit body was attached on the upper side of the central part of the terminal rosette, or one of its radiating branches. Rarely was a fruit body found attached to one of the lower branches. Altogether some 30 or 40 fruit bodies were collected. A number were fixed in Flemming’s solution, some in Biondi’s solution and some in chrom-acetic solution. Other material was carried to Ithaca on the living sphagnum, where a few more fruit bodies were fixed. Some were kept during the winter in moist situations out of doors, and others in doors in a dried state. Finally, during the winter of 1916— 17, it was revealed to me in a semi-vision, that this fungus was a member of the interesting genus Endogone. Structure of the fruit bodies, or complex zygocarps.—The plants are 2-4 mm. in diameter, pulvinate, concave below and convex above, so that a section through the center parallel with the morphological axis is reniform. The larger plants are slightly convoluted or mildly lobed, the upper portion showing two to three broad, low convolutions. As the resting spores mature the plants are orange-yellow in color, but the pigment resides entirely in the spore content, the mycelium and spore walls being hyaline. The peridium is thin, white, and composed entirely of a dense, pliant weft of the terminal, slender branches of the radiating mycelium. The terminal branchlets are 3-5 uw in diameter at the base and taper out to a very slender point I uw or less in diameter. The walls are much thickened, so that the lumen of these narrow branchlets is nearly closed, quite so toward the apex. Many of these slender branchlets are free above the surface and give to the peridium a minutely tomentose, felt-like surface. Many of these branchlets arise very close together, and then are more or less dichotomously branched at a distance, a peculiarity often quite characteristic of the stouter internal mycelium. Internal mycelium and hold-fast.—The internal mycelium has a general radial direction from the basal depression, diverging in all directions toward the peridium. The main hyphae are 12-15 w in diameter. The branching is di- or trichotomous, or 4 to 5 or more branches arise close together, their point of origin often suggesting a ganglion-like enlargement of the parent hyphae from which the branches radiate. The course of the hyphae is more or less sinuous. ATKINSON: THE GENUS ENDOGONE 5 No cross walls have been observed, except in the progametes after copu- lation. The nuclei are minute, very numerous and lie in the peri- pheral granular cytoplasm. There is a nuclear membrane and a large nucleolus (? karyosome). The fruit bodies are quite firmly attached to the living sphagnunt leaves, but the mycelium does not appear to be parasitic, although short haustoria have been seen pene- trating the cell. The hold-fast is a rather compact lattice-like layer of mycelium forming a kind of “‘sole,’’ very closely applied to the surface of the leaves, from which here and there the short haustoria arise. The fungus is probably nourished by organic and mineral solutions carried by the sphagnum from the water of the humus substrate in the capillary stream so well provided for in the peat mosses. Conjugation of the progametes.—While there is a great resemblance in the process of conjugation and in the formation of the resting zygote of Endogone sphagnophila to the situation in E. lactiflua, the details of the process are quite different in the two species. The progamete branches lie nearly or quite parallel. In a few cases where they have been observed just prior to conjugation they do not appear to be differentiated from ordinary stout vegetative branches, except that the cytoplasm is more dense and abundant. They do not appear to be enlarged or clavate. In fact many of the vegetative branches are clavate and sometimes they are in pairs lying closely side by side, but in no case have I been able to determine with certainty that such branches are progametes. The progametes also appear to be un- differentiated before conjugation. They conjugate by lateral contact of their walls at the tip. Immediately after contact the progametes begin to swell into a clavate or fusoid form, and the wall at the point of contact is resorbed for some distance, thus forming a broad com- municating area where the cytoplasm of the two merges. During the enlargement one of the gametes frequently becomes larger than the other. The cytoplasm is very dense and fills the distal portion of the progametes, while in the proximal direction the cytoplasm is less abundant and lies chiefly in a peripheral zone next the wall. St Croix: SECHIUM EDULE (Jacq.) Sw. Cultivated for its fruit. CITRULLUS CITRULLUS (L.) Karst. [C. vulgaris Schrad.] Culti- vated for its fruit. COCCINIA CORDIFOLIA (L.) Cogn. [Cephalandra indica Naud.] Cultivated; recorded by Eggers as naturalized in shaded valleys, St. Croix. LOBELIACEAE ISOTOMA LONGIFLORA (L.) Presl. [Lobelia longiflora L.] Moist banks, fields and hillsides, St. Thomas; St. Jan; St. Croix (according to Eggers). GOODENIACEAE SCAEVOLA PLuMIERII (L.) Vahl. [Lobelia Plumierit L.] Coastal Ssandswot. Whomas st. Croix CICHORIACEAE SONCHUS OLERACEUS L. Waste and cultivated grounds, St. Thomas: St. Jan> st. Grotx. LACTUCA INTYBACEA Jacq. [Brachyramphus intybaceus DC.] Waste and cultivated grounds, St. Thomas; St. Jan; St. Croix. Lactuca sATIVA L. Cultivated for salad. AMBROSIACEAE XANTHIUM LONGIROSTRE Wallr. [X. orientale of Schlechtendal: X. macrocarpum of Eggers; X. strumarium of Millspaugh; X. echi- BRITTON: FLORA OF THE VIRGIN ISLANDS ai natum of Urban.] Waste and cultivated grounds, St. Thomas; all islands (according to Eggers). AMBROSIA CUMANENSIS H.B.K. [A. artemisiaefolia trinitensts Griseb.] Waste places, St. Croix (according to Eggers). CARDUACEAE STRUCHIUM SPARGANOPHORUM (L.) Kuntze. [Ethulia spargano- phora L.; Sparganophorus Vaillantii Crantz.] Moist grounds, St. Thomas (according to Eggers). VERNONIA SERICEA L. C. Rich. [Lepidoploa phyllostachya Cass.; Vernonia arborescens Swartziana, Lessingiana and divaricata of Eggers; V. arborescens of Schlechtendal and of Millspaugh; V. phyllostachya Gleason; Cacalia arborescens Lessingiana Kuntze.| Thickets, St. ihhomas St. Jan; St. Crom VERNONIA ALBICAULIS Pers. [V. longifolia Pers.; V. Vahliana Less.; V. thomae Benth.; V. punctata of Eggers and of Millspaugh; Cacalia thomae Kuntze; ? Conyza fruticosa of West.] Thickets, St. Miomas: St. Jans) St: Croix. VERNONIA CINEREA (L.) Less. [Conyza cinerea L.] Waste and cultivated grounds, St. Thomas; St. Croix. PIPTOCOMA RUFESCENS Cass. Thickets, Water Island, St. Thomas; Se jan. ELEPHANTOPUS MOLLIS H.B.K. [E. tomentosus of Millspaugh.] Banks, fields and hillsides, St. Thomas; St. Jan; St. Croix. PSEUDELEPHANTOPUS SPICATUS (Juss.) Rohr. ([Elephantopus spi- catus Juss.; Distreptus spicatus Rohr.] Banks, fields, hillsides and cultivated grounds, St. Thomas; St. Jan; St. Croix. AGERATUM CONYZOIDES L. [Carelia conyzoides robusta Kuntze.] Banks, fields and roadsides, St. Thomas; St. Jan; St. Croix. EUPATORIUM MACROPHYLLUM L. [Hebeclinium macrophyllum DC. | Forests, St. Thomas; St. Croix (according to West and to Eggers). EUPATORIUM ODORATUM L. [E. conyzoides Vahl.] Banks, hill- sides and thickets, St. Thomas; St. Jan; St. Croix. EUPATORIUM CUNEIFOLIUM Willd., cited by Eggers from DeCandolle (Prodr. 5: 177) as from St. Thomas, was not from our island St. Thomas. EUPATORIUM ATRIPLICIFOLIUM Lam. [E. repandum Willd.; Erig- eron atriplicifolium of Millspaugh.] Hillsides and coastal thickets, Se Phomas; St jam; St. Crom EUPATORIUM SINUATUM Lam. [E. canescens Vahl.] Rocky thick- ets, St. Thomas (according to DeCandolle); rocky hillsides, St. Jan; St; (Groix: 8 98 BROOKLYN BOTANIC GARDEN MEMOIRS EUPATORIUM TRIPLINERVE Vahl. [E.AyapanaVent.] Cultivated on ot. Croix. EUPATORIUM CAPILLIFOLIUM (Lam.) Small. [E. foeniculaceum Willd.] Cultivated on St. Croix (according to Millspaugh). MIKANIA CORDIFOLIA (L. f.) Willd. [Cacalta cordifolia L. f.; ? Eupatorium denticulatum of Schlechtendal; Mikania gonoclada DC.; Willughbaea cordifolia Kuntze; W. gonoclada Millsp.] Thickets, St. Dhomas; St: Jan; St. Croix. ERIGERON CUNEIFOLIUS DC. Grassy places and banks on the higher hills, St. Thomas; St. Jan. ERIGERON SPATHULATUS Vahl. Grassy situations, St. Thomas; Steams otuCrom LEPTILON PUSILLUM (Nutt.) Britton. [Erigeron pusillum Nutt.; Enigeron canadense of Schlechtendal and of Eggers; Leptilon canadense of Millspaugh.] Grassy places, waste and cultivated grounds, St. Thomas; /St. jan; St. Crom LEPTILON LINIFOLIUM (Willd.) Small. [Evigeron linifolium Willd. ; Conyza ambigua DC.] Waste grounds, St. Thomas. PLUCHEA PURPURASCENS (Sw.) DC. [Conyza purpurascens Sw.; P. camphorata of Millspaugh.] Wet grounds, St. Thomas; St. Croix. PLUCHEA ODORATA (L.) Cass. [Conyza odorata L.; C. carolinensis Jacq.; P. odorata normalis Kuntze.| Thickets and hillsides and in cultivated grounds, St. Thomas; St. Jan; St. Croix. BaccHARIS DIoIcA Vahl. [B. Vahlii DC.] Coastal rocks, St. Croix. EGLETES PROSTRATA (Sw.) Kuntze. [Matricaria prostrata Sw.; Pyrethrum simplicifolium Willd.; FE. domingensis Cass.] Sandy shores, St. Thomas. PTEROCAULON VIRGATUM (L.) DC. [Gnaphahium virgatum L.; Conyza virgata L.; Pluchea virgata Schl.] Hillsides and banks, St. Thomas; ot: Jans) ot. Croix. NoccA MOLLIS (Cav.) Jacq. [Lagascea mollis Jacq.) Waste grounds, St. Thomas. MELAMPODIUM DIVARICATUM (L. C. Rich.) DC. [Dysodium divar- icatum L. C. Rich.; M. paludosum H.B.K.] Ditches, St. Croix. PARTHENIUM HysTEROPHORUS L. Waste and cultivated grounds, St. Dhomas: St. Jang st. Croix: CRASSINA MULTIFLORA (L.) Kuntze. [Zinnia multiflora L.| Roadsides and banks, St. Thomas; St. Jan. CRASSINA ELEGANS (Jacq.) Kuntze. Grown in flower gardens. VERBESINA ALBA L. [Eclipia alba Hassk.; E. punctata L.; E. erecta L.]| Wet grounds, St. Thomas; St. Jan; St. Croix. _ ACANTHOSPERMUM HIspipuM DC. [A. humile of Eggers.] Waste and cultivated grounds, St. Thomas; St. Croix. te BRITTON: FLORA OF THE VIRGIN ISLANDS 22 BORRICHIA ARBORESCENS (L.) DC. [Buphthalmum arborescens L.; B. argentea DC.| Coastal rocks and sands, St. Thomas; St. Croix. WEDELIA TRILOBATA (L.) Hitche. [Szlphiwm trilobatum L.; W. carnosa L. C. Rich.] Moist grounds, St. Thomas; St. Croix. WEDELIA CALYCINA L. C. Rich. [Buphthalmum helianthoides of West.] Thickets, St. Thomas; St. Croix. WEDELIA PARVIFLORA L. C. Rich. [W. buphthalmoides of Eggers and of Millspaugh; W. affints DC.; W. acapulcensis of Schlechtendal ; W. brachycarpa of Millspaugh, St. Thomas; Sereneuwm frutescens of Kuntze.] Dry hills and thickets, St. Thomas; St. Jan. WEDELIA CRUCIANA L. C. Rich. [W. buphthalmoides of Miuill- spaugh.] Dry rocky soil, St. Croix. Endemic. ELEUTHERANTHERA RUDERALIS (Sw.) Sch. Bip. [Melampodium ruderale Sw.; Ogiera ruderalis Griseb.; Wedelia discoidea Less.] Banks, fields and waste grounds, St. Thomas; St. Jan; St. Croix. MELANTHERA CANESCENS (Kuntze) O. E. Schulz. [Amellus asper canescens Kuntze; M. Linnaei of Schlechtendal; M. deltoidea of Eggers.] Hillsides and thickets, St. Thomas. TEPION ALATUM (L.) Britton. [Verbesina alata L.] Waste and cultivated ground, St. Thomas; St. Croix. SCLEROCARPUS AFRICANUS Jacq. Waste grounds, St. Thomas. SYNEDRELLA NODIFLORA (L.) Gaertn. [Verbesina nodiflora L.; Ucacou nodiflorum Hitche.] Waste and cultivated grounds, St. Mhomas-sot. jam; ot. Croix. BIDENS PILOSA L. [Coreopsis leucantha L.; B. leucantha Willd.; B. pilosa dubia O. E. Schulz; ? B. pilosa subbiternata Kuntze.] Waste and cultivated grounds, St. Thomas; St. Jan; St. Croix. BIDENS CYNAPIIFOLIA H.B.K. [B. bipinnata of West, of Eggers and of Millspaugh.] Waste and cultivated grounds, St. Thomas; St. jan; St. Crom: Cosmos cAupATUS H.B.K. [Bidens Berteriana Spreng.| Grassy fields, banks, and in waste grounds, St. Thomas; St. Jan; St. Croix. POROPHYLLUM POROPHYLLUM (L.) Kuntze. [Cacalia Porophyllum L.; Porophyllum ellipticum Cass.| Waste grounds, St. Thomas. PEcTIS HUMIFUSA Sw. [P. serpyllifolia Pers.] Stony banks, fields, and hillsides, St. Thomas; St. Jan; St. Croix. PEctis LINIFOLIA L. [Pectis punctata Jacq.; Pectidium punctatum Less.] Rocky hillsides, banks and thickets, St. Thomas; St. Jan; St Croix: PECTIS FEBRIFUGA H. van Hall. [P. Swartziana of Borgesen and Paulsen.] Grassy places, St. Thomas; St. Croix. NEUROLAENA LOBATA (L.) R. Br. [Conyza lobata L.| Woodlands, St. Thomas. 100 BROOKLYN BOTANIC GARDEN MEMOIRS ERECHTHITES HIERACIFOLIA (L.) Raf. [Senecio hieracifolius L.; E. praealta Raf.; E. hieracifolia cacalioides of Eggers and of Kuntze.] Banks, fields, waste and cultivated grounds, St. Thomas; St. Croix. EMILIA SONCHIFOLIA (L.) DC. [Cacalia sonchifolia L.; E. sonchi- folia sagittata of Kuntze.| Banks, fields, waste and cultivated grounds, St. thomas: St. Croix: St. Jan: ; EMILIA SAGITTATA (Vahl) DC. Grown in flower gardens. CHAPTALIA NUTANS (L.) Polak. [Yussilago nutans L.; Leria nutans DC.] Woods and thickets, St. Thomas; St. Jan; St. Croix. CHRYSOGANUM DICHOTOMUM Vahl, of St. Croix, is unknown to me; it is certainly not a Chrysoganum. HELIANTHUS ANNUUS L. Grown in gardens. CHRYSANTHEMUM INDICUM L. [Pyrethrum indicum Cass.] Grown in gardens. ASTER CHINENSIS L. Grown in gardens. TAGETES PATULA L. Grown in gardens. TITHONIA TAGETIFLORA Desf. [J7. speciosa Hook.] Grown in gardens. GEORGINA VARIABILIS Willd. Grown in gardens (according to Eggers). TARCHONANTHUS CAMPHORATUS L. Cultivated on St. Croix (ac- cording to Eggers). PINACEAE THUJA ORIENTALIS L. Planted for ornament. JUNIPERUS BERMUDIANA L. Planted on St. Croix (according to West). CY CADAGCEAE CYCAS REVOLUTA Thunb. Planted for ornament. PTERIDOPHYTA CYATHEACEAE CYATHEA ARBOREA (L.) J. E. Smith. [Polypodium arboreum L.; ? C. Serra of Kuhn.] Forests, high hills, St. Thomas. POLY PODIACEAE DRYOPTERIS PATENS (Sw.) Kuntze. [Polypodium patens Sw.; Aspidium patens Sw.| Forests, high hills of St. Thomas; St. Croix. DRYOPTERIS OLIGOPHYLLA Maxon. [Polypodium invisum Sw.] St. Thomas (according to Christensen). DRYOPTERIS INCISA (Sw.) Kuntze. [Polypodium incisum Sw.] St. Croix (West, according to Eggers). BRITTON: FLORA OF THE VIRGIN ISLANDS 101 DRYOPTERIS SPRENGELIE (Kaulf.) Kuntze. [Dryoteris Balbisii Urban; Polypodium Balbisu Spreng.] St. Thomas (according to Kuhn). DRYOPTERIS SERRA (Sw.) Kuntze. [Polypodium serra Sw.; Tectaria incisa Cav.; Dryopteris serra incisa Kuhn.| St. Thomas (according to Kuhn). DRYOPTERIS MOLLIS (Jacq.) Hieron. [Aspidium molle Sw.] Forests, Signal Hill, St. Thomas; St. Jan; St. Croix (according to Millspaugh). DRYOPTERIS TETRAGONA (Sw.) Urban. [Polypodium tetragonum Sw.]| Forests, St. Thomas; St. Jan; St. Croix. DrYOPTERIS POITEANA (Bory) Urban. | Polypodium crenatum Sw., not Forst; Lastrea Poiteana Bory.] St. Thomas and St. Croix (ac- cording to Eggers); St. Croix (according to West). CYCLOPELTIS SEMICORDATA (Sw.) J. Smith. [Polypodium semi- cordatum Sw.; Aspidium semicordatum Sw.| Shaded localities, Virgin Islands (according to Eggers). GYMNOPTERIS NICOTIANIFOLIA (Sw.) Presl. [Acrostichum nicoti- anum Sw.], attributed by Swartz to St. Thomas, was probably from Porto Rico. NEPHROLEPIS EXALTATA (L.) Schott. [Polypodium exaltatum L.| Forests, St. Thomas; St. Croix. Cultivated on St. Croix (according to Millspaugh). NEPHROLEPIS RIVULARIS (Vahl) Mett. [Polypodium rivulare Vahl.| St. Thomas (according to Kuhn). NEPHROLEPIS BISERRATA (Sw.) Schott. [Aspidiwm biserratum Sw.; Aspidium acutum Schk.; Nephrolepis acuta Presl.; Aspidium punctu- latum Sw.| Forests, St. Thomas (according to Eggers); St. Croix. ODONTOSORIA ACULEATA (L.) J. Smith. [Adiantum aculeatum L.; Davallia aculeata J. E. Smith.] Pastures on high hills, St. Thomas. ODONTOSORIA CLAVATA (L.) J. Smith, is doubtfully attributed to St. Thomas by Fée. ASPLENIUM SERRATUM L. Forests, Signal Hill, St. Thomas. ASPLENIUM PUMILUM Sw. Forests and wet banks, St. Thomas; St Jan: ASPLENIUM ABSCISSUM Willd. [A. firmum Kunze.| St. Thomas (according to Grisebach). BLECHNUM OCCIDENTALE L. Banks, fields and forests, St. Thomas; Stan ot. Croix. PITYOGRAMMA SULPHUREA (Sw.) Maxon. [Gymnogramme sulphurea Desv.]. Cultivated in gardens. PITYOGRAMMA CALOMELANA (L.) Link. [Acrostichum calomelanos L.; Gymnogramme calomelanos Kaulf.; G. calomelanos pumila Eggers.| Banks, hills, walls and thickets, St. Thomas; St. Jan; St. Croix. 102 BROOKLYN BOTANIC GARDEN MEMOIRS HEMIONITIS PALMATA L. Forests, wet banks and rocky thickets, St. ihomas: sts jan; St..Croix: DORYOPTERIS PEDATA (L.) Fée. [Pteris pedata L.]| Forests and shaded banks, St. Thomas; St. Jan. CHEILANTHES MICROPHYLLA Sw. [Adiantum microphyllum Sw.| Rocky slopes, St. Thomas; St. Croix (according to West and to Eggers). ADIANTUM VILLoSUM L. Forests, St. Thomas; St. Croix. ADIANTUM LATIFOLIUM Lam. [A. denticulatum Sw.; A. inter- medium of Eggers; A. obliquum intermedium of Millspaugh.] Shaded banks, hills of St. Thomas. ADIANTUM CRISTATUM L. [? A. microphyllum of Eggers.] Hill- sides, St. Thomas. ADIANTUM TENERUM Sw. Shaded banks and ravines on high hills, St. Thomas; St. Jan (according to Eggers); St. Croix. ADIANTUM FRAGILE Sw. Thickets and walls, St. Thomas; St. Jan; St Grom ADIANTUM FARLEYENSE Moore. Cultivated on St. Croix (A. foliosum of Millspaugh). PYCNODORIA LONGIFOLIA (L.) Britton. [Pteris longifolja L.] Along rivulets in forests, St. Croix; in a water spout, St. Thomas. PTERIS BIAURITA L. St. Thomas (according to Kuhn). ANTROPHYUM LINEATUM (Sw.) Kaulf. [Hemionitis lineata Sw.] Forest, St. Peter, St. Thomas (according to Eggers). PALTONIUM LANCEOLATUM (L.) Presl. [Péteris lanceolata L.; Taenitis lanceolata R. Br.; Heteropteris lanceolata Fée.| On rocks and trees in forests, St. Thomas; all islands (according to Eggers). POLYPODIUM POLYPODIOIDES (L.) Hitche. [Acrostichum poly- podioides L.; P. incanum Sw.| On trees, St. Thomas; St. Jan; St. Croix (according to Eggers). PHLEBODIUM AUREUM (L.) J. Sm. [Polypodium aureum L.| On trees and rocks, St. Thomas; St. Jan; St. Croix. PHLEBODIUM AREOLATUM (H.&B.).J.Sm. [Polypodium areolatum H. & B.] On trees, St. Thomas; St. Jan. LEPICYSTIS PILOSELLOIDES (L.) Diels. [Polypodium piloselloides L.| In forests among rocks, Signal Hill, St. Thomas. PHYMATODES EXIGUUM (Hew.) Underw. [Polypodium exiguum Hew.; P. serbens Sw., not Forst.; P. Swartz Baker.| On trees, Bordeaux, St. Jan; St. Croix. CAMPYLONEURUM PHYLLITIDIS (L.) Presl. [Polypodium Phyllitidis L.; P. Phyllitidis repens of Eggers.] On rocks and trees in forests, St; Ehomas; St: Jans St: Crom CAMPYLONEURUM LATUM Moore. Shaded rocks, Bethania, St. Jan. BRITTON: FLORA OF THE VIRGIN ISLANDS 103 _ ACROSTICHUM AUREUM L. [Chrysodium vulgare Fée.]| Borders of marshes, St. Thomas; St. Croix. OPHIOGLOSSACEAE OPHIOGLOSSUM RETICULATUM L. Grassy places among _ rocks, Crown, St. Thomas; shaded bank, Bordeaux, St. Jan. LYCOPODIACEAE LycopoDIUM CERNUUM L. Among rocks in higher hills, St. Thomas. 4 PSILOTACEAE PsILOTUM NUDUM (L.) Griseb. [Lycopodium nudum L.; P. tri- quetrum Sw.| Shaded places among rocks, Signal Hill, St. Thomas; Bordeaux, St. Jan; Crequis, St. Croix. BRYOPHYTA MUSCI* DICRANELLA LONGIROSTRIS (Schwaegr.) Mitten. [Zvematodon longirostris Schwaegr.| St. Jan. LEUCOLOMA SERRULATUM Bridel. [L. Riedlei Besch.] On trees in wet woods, St. Thomas. OCTOBLEPHARUM ALBIDUM (L.) Hedw. [Bryum albidum L.|] On roots of Anthurium, near Caret Bay, St. Thomas; St. Croix. FISSIDENS KEGELIANUS C. Muell. [F. palmatus—of various authors, not Swartz.| On banks, St. Thomas; St. Jan; St. Croix. FISSIDENS ELEGANS Bridel. On rocks and earth, St. Thomas; St wlan. SYRRHOPODON FLAVESCENS C. Muell. On rotten wood, St. Jan. CALYMPERES RICHARDI C. Muell. . [C. Breutelii Besch.; C. hexa- gonum Besch.| On rocks and banks, St. Thomas. HYMENOSTOMUM BREUTELII (C. Muell.) Broth. [Weisia Breutelit C. Muell.; Gymnostomum Breutelu Br. & Sch.] On banks, St. Hhaeinas- St. jans2St:Croix. HyopuHiILa TorTuLA (Schwaegr.) Hampe. [Gymnostomum Tortula Schwaegr.] St. Croix. BARBULA AGRARIA (Sw.) Hedw. [Bryum agrarium Sw.] On rocks, walls and earth, St. Thomas; St. Jan; St. Croix. BARBULA CRUEGERI Lond. [Hyophila uliginosa E. G. Britton.] Bethania, St. Jan. PHASCUM SESSILE E. G. Britton. On the ground, Cowell Point and Water Island, St. Thomas. 4 Contributed by Elizabeth G. Britton. 104 BROOKLYN BOTANIC GARDEN MEMOIRS BrRyYUM CRUEGERI Hampe. Bed of stream, Tutu, St. Thomas. Sterile. PHILONOTIS SPHAEROCARPA (Sw.) Bridel. [Mnium sphaericarpum Sw.] Moist banks, high hills of St. Thomas. PHILONOTIS TENELLA (C. Muell.) Jaeger. [Bartramia tenella C. Muell.] Wet banks, St. Jan. PIREELLA CYMBIFOLIA (Sull.) Cardot. [Pilotrichum cymbifolium Sull.] On trees near Bethania, St. Jan. NECKERA DISTICHA (Sw.) Hedw. [Fontinalis disticha Sw.] On trees, rarely on rocks, St. Peter, St. Thomas. NECKERA JAMAICENSIS (Gmel.) E. G. Britton. [Hypnum jamai- cense Gmel.] On trees, Bethania, St. Jan. CALLICOSTELLA BELANGERIANA (Besch.) Jaeger. [Hookeria Bel- angeriana Besch.] On stones, Bordeaux, St. Jan. STEREOPHYLLUM LEUCOSTEGUM (Bridel) Mitten. [Leskea leuco- stega Bridel.| On wet or shaded rocks, St. Thomas; St. Jan. MITTENOTHAMNIUM DIMINUTIVUM (Hampe) E.G. Britton. [Hyp- num diminutivum Hampe.| On old wood, Bordeaux, St. Jan. TAXITHELIUM PLANUM (Bridel) Mitten. [Hypnum planum Bridel.] Wet rocks, logs and tree-roots, St. Thomas; St. Jan. SEMATOPHYLLUM ADMISTUM (Sull.) Mitten. [Hypnum admistum Sull.] Shaded banks, stones and dead wood, St. Thomas; St. Jan. HAPLOCLADIUM MICROPHYLLUM (Sw.) Broth. [Hypnum_ micro- phyllum Sw.| Shaded bank between Pearl and Bonne Resolution, St. Thomas. THUIDIUM CYMBIFOLIUM (Dz. & Mk.) Br. Jav. Shaded banks, St. Thomas. DENDROPOGON RUFESCENS Schimp., a Mexican species, has been credited to St. Thomas in Paris, Index, and accepted by Brotherus, but we have seen no specimens. HEPATICAE OF ST. CROIX, ST. JAN, ST. THOMAS AND TORTOLA® In the Synopsis Hepaticarum of Gottsche, Lindenberg and Nees von Esenbeck, published in 1844-47, three species of Hepaticae are listed from St. Croix, one from St. Jan, and one from St. Thomas. Another species, although listed from St. Kitts, was based in all prob- ability on material from St. Jan. A seventh species has been listed from St. Jan by Stephani. These seven species, which are the only ones so far reported from the islands under discussion, deserve a few words of comment. ° Contributed by Alexander W. Evans, Yale University. BRITTON: FLORA OF THE VIRGIN ISLANDS 105 The first species, Radula pallens (Sw.) Dumort., is said to have been found “‘in St. Crucis insula,” the record being based on a speci- men in the Weber herbarium. This specimen was originally referred to Jungermannia complanata L. (Radula complanata Dumort.) by Weber,® but the later determination is probably correct. The second species, Lejeunea Montagnei Gottsche, was based on material from the Mascarene Islands and is now regarded as a species of Euosmolejeunea. A specimen from St. Croix is listed in the Synopsis but is very problematical and would probably now be referred to some other species. Since the specimen in question has not been available for study, and since no later references to it are to be found in the literature, its status must be left in doubt. The third species, Lejeunea bethanica Gottsche, is based on material collected by Breutel and is said to have come from “ prope Bethaniam in Insula St. Christopheri.’”’ Many years later Stephani,’ on the basis of a specimen in the Lindenberg herbarium at Vienna, quoted the species from St. Jan, referring it to the subgenus Chetlo-Lejeunea. Still later he apparently changed his ideas regarding the habitat of the plant, citing it from St. Kitts and redescribing it under the name Cheilolejeunea bethanica Steph.2 In studying the Lejeuneae in the Lindenberg herbarium, the writer found two specimens labeled Lejeunea bethanica, both of which were collected by Breutel at Be- thania, St. Jan. One of these is very fragmentary but is apparently referable to Rectolejeunea phyllobola (Nees & Mont.) Evans; the other, which is the specimen studied by Stephani, is (in the writer’s opinion) referable to Lejeunea rather than to Cheilolejeunea. The species was originally described from a specimen in the Gottsche herbarium at Berlin, not available at the present time, and there is therefore a possibility that the actual type may have come from St. Kitts. The evidence, however, is against this view, and it seems permissible to assume that the specimen in the Lindenberg herbarium is identical with the type and that it formed a part of the same collection. Un- fortunately L. bethanica has not again been collected on either St. Jan or St. Kitts. The fourth species, Lejeunea epiphyta Gottsche, was described as ‘‘parasitans in Lej. bethanica in Insula St. Johannis prope Bethaniam (Breutel, Hb. G.).”’ This statement affords further proof that L. bethanica came from St. Jan. According to Stephani® the specimen of L. epiphyta in the Lindenberg herbarium should be referred to 6 Prodr. Hist. Musc. Hepat. 59. 1815. 7 Hedwigia 29: 86. 1890. 8 Sp. Hepat. 5: 652. 1914. 5 Hedwigia 29: 90. 1890. 106 BROOKLYN BOTANIC GARDEN MEMOIRS Lejeunea myriocarpa Nees & Mont., now Cololejeunea myriocarpa Evans. The fifth species, Anthoceros Breutelit Gottsche, was said to have been collected near Friedensthal, St. Croix. This species, in 1858, was transferred by its author'® to the genus Nofothylas, where it is still retained. The following year Milde'! showed that the type material of the species did not come from St. Croix but from the Corallberg, St. Jan. There is likewise a specimen from St. Jan in the Mitten herbarium, which is presumably a part of the original collection. The sixth species, Lejeunea linguaefolia Tayl., was found “in Insula St. Thomas (Richard in Hb. Hk. a. 1814).’”’ A specimen of this species in the Lindenberg herbarium is referred by Stephani to Brachiolejeunea corticalis (Lehm. & Lindenb.) Schiffn., and the writer would make the same disposition of a specimen in the Mitten her- barium.’ Recently, however, Stephani has apparently thrown doubt on the propriety of this reduction. In the fifth volume of his Species Hepaticarum (1912), on page 35, he includes L. linguaefolia among the species of Ptychocoleus, citing it as Pt. linguaefolius Steph., and adds that he has been unable to see the plant and that his diagnosis is simply a translation of the original description. On page 127, never- theless, he again quotes L. linguaefolia among the synonyms of B. corticalis. Since Stephani’s original reduction was based on the study of an actual specimen, the writer would regard Pt. linguaefolius as nothing more than an unnecessary synonym. The seventh and last species, Riccia Breutelii Hampe, is described as new by Stephani in the first volume of his Species Hepaticarum (1898), on page 17, the habitat being given as ‘‘ Insulae S’Kitts et S. Juan.”’ Dr. Howe informs the writer that there is some question about the identity of the St. Kitts and St. Jan plants and it is there- fore omitted from the following list, pending investigation of authentic material. In February, 1913, a botanical exploration of the islands was carried on under the auspices of the New York Botanical Garden, the Carnegie Institution of Washington and the United States National Museum.'* The Hepaticae collected by the various members of this expedition form the basis for the present report. The specimens from St. Thomas, unless otherwise noted, were collected by Mrs. Elizabeth G. Britton and Miss Delia W. Marble, those from St. Jan 10 Bot. Zeit. 16 (Anhang): 21. 1858. 1 Bot. Zeit. 17: 50. 1859. 2 Hedwigia 29: 22. 1890. 18 See Bull. Torrey Club 35: 164. 1908. it See Britton, N.(L.; Jour. NY. Bot. Gard: 14299." 1913: BRITTON: FLORA OF THE VIRGIN ISLANDS 107 and Tortola by N. L. Britton and J. A. Shafer, and those from St. Croix by J. N. Rose. The report records also two specimens collected by C. H. Ostenfeld in 1914, one on St. Thomas and one on St. Jan. 1. RicctA Brittronit M. A. Howe.!® St. THOMAS: on the ground, Water Island, N. L. Britton, E. G. Britton & J. A. Shafer 148 (a much larger plant than the original). 2. PLAGIOCHILA LUDOVICIANA Sulliv. St. JAN: on wet rocks, Bethania, 360. St. THOMAS: on rocks, St. Peter, 1453. 3. RADULA PALLENS (Sw.) Dumort. Sr. Crorx: without definite locality, collector unknown, cited in the Synopsis Hepaticarum. 4. COLOLEJEUNEA MYRIOCARPA (Nees & Mont.) Evans. Lejeunea epiphyta Gottsche. St. Crorx: without definite locality. Sr. JAN: near Bethania, J.C. Breutel. St. THOMAS: on rotten wood, near Magin’s Bay, 1317. ToRTOLA: on a rock, Road Town to High Bush, 325 m. alt., 786 (a trace only). 5. LEJEUNEA BETHANICA Gottsche. Chetlolejeunea bethanica Steph. ST. JAN: near Bethania, J. C. Breutel. 6. LEJEUNEA GLAUCESCENS Gottsche. St. THOMAS: on tree roots, Bonne Resolution, 447. 7. LEJEUNEA MINUTILOBA Evans. ST. THOMAS: St. Peter, 1251, 1254, 1255; on stones, Crown, 1365 (type). 8. LEJEUNEA PILILOBA Spruce. St. Croix: on bark of a tree, without definite locality. Sr. JAN: at base of a tree, Rosenberg, 300 m. alt., 306; on a shaded rock, Bordeaux, 350 m. alt., 57z. St. THOMAS: on ridge north of Charlotte Amalia, 406; Crown, 1365 (a trace only); on rotten wood, St. Peter, 1451. g. MICROLEJEUNEA LAETEVIRENS (Nees & Mont.) Evans. St. THOMAS: on Anthurium roots, Pearl to Bonne Resolution, 1340; on fern roots, St. Peter, 1253; on mountain behind Charlotte Amalia, C: H. Ostenfeld 77. TorTOLA: on a rock, Roadtown to High Bush, 325 m. alt., 786 (in part). 10. RECTOLEJEUNEA PHYLLOBOLA (Nees & Mont.) Evans. St. JAN: near Bethania, J. C. Breutel (specimen in the Lindenberg ‘’ Determined by Marshall A. Howe. 108 BROOKLYN BOTANIC GARDEN MEMOIRS herbarium, labeled L. bethanica); on Clusia roots, Bethania, 355. Sr. THOMAS: on roots of royal palm, Tutu, 423. ToRTOLA: on a rock, Roadtown to High Bush, 325 m. alt., 786 (in part); shaded rocks, High Bush, 375 m. alt., &75. 11. EUOSMOLEJEUNEA CLAUSA (Nees & Mont.) Evans. Sr. THomMAs: shaded bank, Pearl to Bonne Resolution, 2339; on the ground, St. Peter, 1455. 12. EUOSMOLEJEUNEA DURIUSCULA (Nees) Evans. St. JAN: shaded rocks, Bordeaux, 330 m. alt., 570. Str. THOMAS: on rocks, St. Peter, 7452; on rocks, Crown, 1454. 13. EUOSMOLEJEUNEA TRIFARIA (Nees) Schiffn. St. THomMAS: on rocks, St. Peter, 1252; on a rotten log, Crown, ASOumemalis, 107. 14. TAXILEJEUNEA OBTUSANGULA (Spruce) Evans. St. JAN: on a stone, Bordeaux, 400 m. alt., 582. 15. MASTIGOLEJEUNEA AURICULATA (Wils. & Hook.) Schiffn. St. JAN: on a tree, Bethania to Rosenberg, 243; on a wet rock, Bethania 356, 362; on loose blocks, Little Cruz Bay, C. H. Ostenfeld 391. ST. THOMAS: on a rock, St. Peter, 7256; on rocks in ravine below Tutu, 7290. 16. BRACHIOLEJEUNEA CORTICALIS (Lehm. & Lindenb.) Schiffn. Lejeunea linguaefolia Tay. Ptychocoleus linguaefolius Steph. St. THOMAS: without definite locality, L. C. Richard. 17. FRULLANIA BRASILIENSIS Raddi. Sr. JAN: on wet rocks. Bethania, 363 (mostly, specimens with- out perianths and therefore somewhat doubtful). 18. FRULLANIA KUNZEI Lehm. & Lindenb. Sr. JAN: on wet rocks, Bethania, 363 (a trace only); on rocks, Bordeaux, 300 m. alt., 550. Ig. FRULLANIA SQUARROSA (R. BI. & N.) Dumort. St. THOMAS: on rocks, ridge north of Charlotte Amalia, 407; on rocks, Bonne Resolution, 440; on rocks, Magin’s Bay to Mafolie, 1313; on rocks, Pearl to Bonne Resolution, 7338; on a stone wall, Crown, 450 m. alt., 7364. 20. NoTroTHyLAS BREUTELII Gottsche. Anthoceros Breutelit Gottsche. St. JAN: Corallberg, J. C. Breutel (type); without definite locality or collector’s name (specimen in Mitten herbarium); Bordeaux, 528. St. THOMAS: on damp earth, Nisky, N. L. Britton, LE. G. Britton & BRITTON: FLORA OF THE VIRGIN ISLANDS 109 D.W. Marble 76; hills north of Charlotte Amalia, 409; on the ground, Magin’s Bay to Mafolie, 7312. 21. ANTHOCEROS PUNCTATUS L. St. THOMAS: in a mud hole, Crown, 450 m. alt., 7363. REPORT ON THE LICHENS OF ST. THOMAS AND ST. JAN The following report is based primarily upon a study of 90 numbers of lichens from St. Thomas and St. Jan Islands; from St. Thomas, - 82 numbers collected by Dr. N.-L. Britton, Mrs. E. G. Britton, and Miss Delia W. Marble, unless otherwise stated; from St. John, 8 numbers collected by Dr. N. L. Britton and Dr. J. A. Shafer. To these have been added a few species recorded by Nylander in Flora (63: 127. 1880) and two endemic species described by Mueller- Argau. Altogether, we have represented 30 genera with 69 species and varieties, of which three species and one variety are here de- scribed as new. The discussion of the characteristics of the lichen- flora will be confined to that of St. Thomas. There is no peculiar element in the lichen-flora of St. Thomas comparable with the gelatinous rock-lichens of Mona Island (see Annals Missouri Bot. Gard. 2: 35. 1915), or with the crustose and gelatinous rock-lichens of Bermuda (see Bull. Torrey Bot. Club 43: 146-155. 1916). Such rock-lichens as occur on St. Thomas belong to widely distributed groups even where the actual species are more restricted in distribution. In fact the greater part of the lichen- flora is made up of species more or less common throughout the West Indies or even in the tropics of both hemispheres. It is probably safe to say that there are not more than half a dozen endemic species in St. Thomas. Of the three new species described in this paper, two occur elsewhere in the West Indies. The flora is comparatively rich in the variety of crustose bark-lichens, especially in the genera An- thracothecium with 5 species, Arthonia with 8 species, and Opegrapha with 5 species. It is rather surprising that only one Graphis and no Trypethelium should have been collected. On the whole the lichen- flora is rather commonplace. In the following list, stations outside of the Virgin Islands are noted for species, the range of which is limited or not well known. LICHENS: OF ST, THOMAS 1. DERMATOCARPON HEPATICUM (Ach.) Th. Fr. On soil, without definite locality, 151. 16 Contributed by Lincoln W. Riddle, Wellesley College. 110 NO BROOKLYN BOTANIC GARDEN MEMOIRS . LEPTORAPHIS EPIDERMIDIS (Ach.) Th. Fr. On bark, Bordeaux, 1381. 3. PORINA DESQUAMESCENS Fée. Without data, 1288a. 4. PoRINA NUCULA Ach. On Erythrina, St. Peter, 1444b, 1445. 5. PYRENULA LEUCOPLACA (Wallr.) Koerb. On bark, Smith’s Bay, 1276a. 6. ANTHRACOTHECIUM BREUTELI Muell. Arg. Flora 68: 339. 1885. On bark, without definite locality, collected by Breutel. En- demic. 7, ANTHRACOTHECIUM CANELLAE-ALBAE (Fée) Muell. Arg. On Cephalocereus, Smith’s Bay, 1274a; on twigs of Guettarda, Crown, 450 m. altit., 1357. South America. 8. ANTHRACOTHECIUM LIBRICOLUM (Fée) Muell. Arg. On Melicocca, Tutu, 468a; on same, Smith’s Bay, 1276, 1282; on Erythrina, Sit. Peter, 1444> ' 9. ANTHRACOTHECIUM OCHRACEOFLAVUM (Nyl.) Muell. Arg. On bark of coconut palm, near Charlotte Amalia, 489; on Trichilia, Tutu, 467. 10. ANTHRACOTHECIUM PYRENULOIDES (Mont.) Muell. Arg. On bark, IE be without definite locality, collected by Dr. Forel. Recorded by Nylander in Flora 63: 127. 1880. MELANOTHECA FOVEOLATA Muell. Arg. On bark, near Tutu, 1287. Cuba. . PARATHELIUM INDUTUM ‘1. On Cephalocereus, Smith’s Bay, Pe D Nyl. On Cephal Smith’s Bay 1274; on bark, Bordeaux, 1382 in part. Porto Rico, Colombia. . PARMENTARIA ASTROIDEA Fée. On bark, Bordeaux, 1382 in part. . ARTHONIA ANEGADENSIS Riddle Mem. New York Bot. Gard. 6: 579. 1916. On Bontia, Smith’s Bay, 1286. Anegada. . ARTHONIA CONFERTA (Fée) Nyl. On Plumeria, Tutu, 1484. . ARTHONIA GREGARIA (Weig.) Koerb. On bark, Bordeaux, 1380. . ARTHONIA OCHRACEELLA Nyl. On bark of old fallen branch, Crown, 450 m. altit., 1362. Cuba. . ARTHONIA RUBELLA (Fée) Nyl. On Melicocca, Tutu, 468b. . ARTHONIA SUBRUBELLA Nyl. Collected by Dr. Forel. Recorded by Nylander in Flora 63: 127. 1880. . ARTHONIA.™ On Guilandina, Smith’s Bay, 1281 p.p. . ARTHONIA. On Plumeria, Tutu, 463. . ARTHOTHELIUM MACROTHECUM (Fée) Mass. On mango, St. Peter, 1247. . Opegrapha acicularis Riddle sp. nov. Thallus epiphloeodes crustaceus effusus determinatus nigrolimitatus, albidus “There are included in this paper three species of Arthonia the determination of which it has not been possible to complete in the time at my disposal. L. W. R. BRITTON: FLORA OF THE VIRGIN ISLANDS 111 vel albo-cinerascens, tenuis laevis subcontinuus. Apothecia sessilia nuda ad 0.8 mm. lata, primum orbicularia suburceolata, margine crenato, dein orbicularia oblonga vel subdifformia, disco late aperto plano rugoso atrofusco, margine laciniato partim stellato-radiato nitido nigro; epithecio fusco; hymenio incolore, 120-130 uz altit.; hypothecio incolore; amphithecio sub lamina deficiente, cetero crasso nigro. Paraphyses firmae crassae ramosae haud connexae. Asci cylindrices, 8-spori. Sporae incolores aciculares rectae vel subflexuosae vel rarius contortae, 20-24- loculares, cellulis cylindricis, 60-100 x 2-3 yu. Fic. 1. Opegrapha acicularis Riddle. Vertical section of apothecium (stippling indicates dark coloration); paraphysis; ascus with two of the eight spores. On bark of Erythroxylon, Punta Aguila, Porto Rico, collected by N. L. Britton, J. F. Cowell, and Stewardson Brown, Feb. 27, 1915, no. 4682 (type!). Also, on Guilandina, Smith’s Bay, St. Thomas, 1276a, 1279; and on Coccolobis, Great Harbor Cay, Berry Islands, Bahamas, N. L. Britton and C. F. Millspaugh, no. 2545. This is a striking and distinct species belonging to the section Pleurothecium of the genus Opegrapha. Externally it is easily recog- nizable by the marked tendency for the laciniate margin of the apo- thecia to spread out in a stellate manner. The spores resemble those of Lecanactis myriadea (Fée) Zahlbr. and of Opegrapha pleistophrag- moides Nyl. But both of the species named have the black amphi- thecium complete at the base, and the spores are almost twice as thick. 24. OPEGRAPHA AGELAEA Fée. On Crescentia, Tutu, 462. Cuba, Colombia. 25. OPEGRAPHA ATRA Pers. On Guilandina, Smith’s Bay, 1278a; on Melicocca, Smith’s Bay, 1285. P12 BROOKLYN BOTANIC GARDEN MEMOIRS 26. OPEGRAPHA BONPLANDI Fée. On Melicocca, Magen’s Bay, 1310; on bark, Mariendahl Road, 1476. 27. OPEGRAPHA VULGATA Ach. On Melicocca, Tutu, 468. 28. GRAPHIS SCRIPTA (L.) Ach. On bark, Smith’s Bay, 1277. 29. PHAEOGRAPHIS INUSTA (Ach.) Muell. Arg. On Acacia, near Charlotte Amalia, 486; on Guilandina, Smith’s Bay, 1278; without locality, 464. 30. CHIODECTON (Sect. Enterographa) sp. On Erythrina, St. Peter, 1445a. 31. GYROSTOMUM SCYPHULIFERUM (Ach.) Fr. On Plumeria, Tutu, 465; on Acacia, near Charlotte Amalia, 487, 494. 32. BILIMBIA CUPREA Massal. in Lotos (1856) 77. Lecidea cupreorosella Nyl. Mem. Soc. Sci. Nat. Cherb. 5: 122. 1857. Biatora cupreorosella Tuck. Syn. N. A. Lich. 2: 34. 1888. On old brick, St. Peter, 1442. Eastern United States, Europe. 33. BACIDIA ALBESCENS (Arn.) Zwackn. On Erythrina, St. Peter, 1445b. 34. CLADONIA PITYREA f. SQUAMULIFERA Wainio. On rocks, slope of Crown, 1440. 35. LEPTOGIUM CHLOROMELUM (Sw.) Nyl. On bark, Cowell Point, 103, 172; on Pisonia roots, Water Island, 153. 36. LEPTOGIUM TREMELLOIDES (L. f.) S..F. Gray. On tree-trunk, Crown, 1361. 37. LEPTOGIUM TREMELLOIDES var. CAESIUM (Ach.) Hue. On rock near Bonne Resolution, 447. 38. LECANORA CINEREOCARNEA (Eschw.) Wainio. Without data, 23a; on Guilandina, Smith’s Bay, 1281. 39. LECANORA GRANIFERA Ach. On bark, Mariendahl Road, 1476a. 40. Lecania euthallina Riddle sp. nov. Thallus crustaceus uniformis effusus crassus rimoso-areolatus, areolis 0.2—-0.4 mm. latis leviter convexis contiguis, cinereus vel sat pallide fuscescens; hypothallo nullo. Gonidia cystococcoidea. Apothecia 0.6 mm. (0.4-I1.0 mm.) lata, numerosa partim caespitosa superficialia sat elevata regularia, disco concavo castaneo vel fusco-nigricante nudo, margine proprio tenue disco concolore, margine thallino integro vel demum crenulato crasso prominente thallo concolore; epithecio fulvo; hymenio et hypothecio incolore. Asci 8-spori. Sporae incolores oblongae bilocu- lares haud placodiomorphae, 10-12 x 4-5 wu. On rock, Tutu, St. Thomas, collected by Dr. N. L. Britton, Mrs. E. G. Britton, and Miss Delia W. Marble, Feb. 8-9, 1913, no. 469 (type!). Lecania euthallina differs from L. erysibe (Ach.) Th. Fr. in the much better developed thallus (whence the specific name), it being compact, BRITTON: FLORA OF THE. VIRGIN ISLANDS 113 thicker, and more continuous. Furthermore, the apothecia are more concave, with the persistent thalline margin more conspicuous. 41. PARMELIA CETRATA f. SUBISIDIOSA Muell. Arg. On tree-trunk, Crown, 1441 in part. North Carolina, Florida, Cuba, Jamaica. 42. PARMELIA CONSPERSA (Ehrh.) Ach. On rocks, Crown, 450 m. aldtt: 50358: 43. PARMELIA LATISSIMA var. CRISTIFERA (Taylor) Hue. On tree- trunk, St. Peter, 1249; on twig, Crown, 1441 in part. 44. PARMELIA PERLATA (L.) Ach. On rocks, near Bonne Resolution, 446; on Spondias, Mandal, 1311. 45. RAMALINA COMPLANATA (Sw.) Ach. Without data, 1356a. 46. RAMALINA GRACILIS (Pers.) Nyl. On twigs of Guettarda, Crown, 450 m. altit., 1356. 47. Blastenia nigrocincta Riddle sp. nov. Thallus crustaceus arcte adnatus sat crassus, ambitu subradiato-laciniatus effiguratusque, centro rimoso-areolatus, areolis 0.4-0.8 mm. latis leviter convexa primum contiguis demum hypothallo nigro dispersis, cinereo-albescens dein fumosus aut partim luridus. Apothecia 0.3-0.5 mm. lata, superficialia dispersa vel partim caespitosa nuda, disco plano vel leviter convexa ferrugineo-aurantiaco, margine proprio sat tenue persistente nigro nitido, margine thallino nullo; excipulo externe coeruleo-nigro interne incolore; epithecio ferrugineo; hymenio incolore; hypothecio pallide fuscescente. Asci 8-spori. Sporae incolores ellipsoideae biloculares placodio- morphae, loculis poro tenue confluentibus, 12-14 x 5-6 yu. On rock, Tutu, St. Thomas, collected by Mrs. E. G. Britton and Miss Delia W. Marble, Feb. 8-9, 1913, no. 469a (type!). Also, on limestone, Montalva, Porto Rico, N. L. Britton, J. F. Cowell, and Stewardson Brown, March 2-4, 1915, no. 4810. This species is distinct in the contrasting coloration of the black margin and the orange disk of the apothecia, a character which will serve to distinguish it on the one hand from species with similar thalline characters, such as Bl. Forstroemiana (Fr.) Muell. Arg.; and on the other hand from Bl. ferruginea (Huds.) Koerb., where the disk and margin are concolorous, and from Blastenia peragrata (Fée) Muell. Arg., where the margin is black, but the disk is aeruginous- brown. 48. Caloplaca diplacia (Ach.) Riddle comb. nov. Lecanora Ach. Synop. Lich. 154. 1814. On rock, near Charlotte Amalia, 493, 495, 1485 in part. Also recorded by Nylander in Flora 63: 127. 1880. Apparently confined to the West Indies. 49. CALOPLACA MURORUM (Hoffm.) Th. Fr. On rock, near Charlotte Amalia, 1485 in part. 114 BROOKLYN BOTANIC GARDEN MEMOIRS 50. Caloplaca subsequestra (Nyl.) Riddle comb. nov. Lecanora Nyl. Flora 63: 127. 1880. On rocks, without definite locality, collected by Dr. Forel. En- demic. 51. BUELLIA DISCOLOR (Hepp) Koerb. On rock, Tutu, 469b; with- out definite locality, collected by Dr. Forel, according to Nylander (Il. c.). Europe. 52. BUELLIA PARASEMA var. AERUGINESCENS (Nyl.) Muell. Arg. On coconut near Charlotte Amalia, 489a. 53. Buellia prospersa (Nyl.) Riddle comb. nov. Lecidia Nyl. Flora 63: 127. 1880. On rocks, without definite locality, collected by Dr. Forel. En- demic. 54. PYXINE CocoEs (Sw.) Nyl. On bark, near Bonne Resolution, 445; on Melicocca, Tutu, 466. 55. PYXINE COCOES var. ENDOXANTHA Muell. Arg. On Guitlandina, Smith’s Bay, 1280; on bark, Mariendahl Road, 1475. 56. PyxtNE MEISSNERI Tuck. On coconut palm, without definite locality, collected by Dr. J. N. Rose, 3198. 57. PHYSCIA ALBA (Fée) Muell. Arg. On Erythrina, St. Peter, 1443. 58. PuysciA CAESIA (Hoffm.) Nyl. On rocks, without definite local- ity, collected by Dr: Forel. Recorded by Nyl. Flora 63: 127. 1880. 59. PHyscIA CRISPA (Pers.) Nyl. On Elaphrium, near Charlotte Amalia, 491; on roots, same locality, 492, 496; on Melicocca, Tutu, 461. 60. PHYSCIA EROSULA Nyl. Flora 63: 127. 1880. Based on material growing on rocks, St. Thomas, without definite locality, col- lected by Dr. Forel. Doubtfully distinct from the widely distributed Physcia tribacia (Ach.) Tuck. 61. PHyscia PICTA (Sw.) Nyl. On rock, St. Peter, 1260; on coconut palm, collected by Dr. J. N. Rose, 3197. 62. PHYSCIA SPECIOSA (Wulf.) Nyl. On rocks, near Charlotte Amalia, 405; on bark, St. Peter, 1248, 1250. LICHENS OF Si. ,AN I. PYRENULA MAMILLANA (Ach.) Trev. On bark of Icacorea, Bor- deaux, 597. 2. MELANOTHECA ACHARIANA Fée. On Inga, Bordeaux, 598. Cuba, Venezuela. 3. MYCOPORELLUM ELLIPTICUM Muell. Arg. Flora 72: 508. 1889. On bark, without definite locality, collected by Levier, no. 113. Endemic. BRITTON: FLORA OF THE VIRGIN ISLANDS LTS 4. ARTHONIA. On bark, Bordeaux, 577. 5. ARTHOTHELIUM MACROTHECUM (Fée) Mass. On JIcacorea, Bor- deaux, 540. 6. Graphina nitidescens (Nyl.) Riddle comb. nov. Fissurina Nyl. Lich. Japon. 108. 1890. On Nectandra, Bordeaux, 581. Florida, Cuba, Porto Rico. 7. OPEGRAPHA VULGATA Ach. On Maytenus, Little St. James Island, N. L. Britton & J. N. Rose, 1405. 8. Leptogium marginellum var. isidiosellum Riddle var. nov. Thallus isidiis tenuibus dense tectus; ceterus ut in forma typica apothecia nulla. On wet rock, road to Rosenberg, N. L. Britton & J. A. Shafer, Feb. 5-7, 1913, no. 276 (type!). The fringed apothecia being such a characteristic feature of Leptogium marginellum, it is only after some hesitation that this material has been placed here. The texture and the wrinkling of the thallus is in exact agreement, however, with typical specimens. And the relation of this variety to the species is strictly comparable with the conditions in L. tremelloides, abundantly fruiting in the tropics, and its variety caesium, with isidia but very rarely fruiting; and with L. phyllocarpum and its variety isidiosellum. 9g. PARMELIA TINCTORUM Despr. On tree-trunk, Bordeaux, 567. The following lichens are recorded in “Le Végétation des Antilles Danoises”’ by F. Bérgesen & Ove Paulsen in Revue Générale de Botanique 12: 507, 508. 1900. StictA WEIGELII (Ach.) Wainio. St. Croix; St. Thomas. GRAPHIS scRIPTA (L.) Ach. St. Croix. LECIDEA BUELLIANA Muell. Arg. St. Croix. PERTUSARIA WULFENII (DC.) Fr. St. Croix. PARMELIA PERLATA (L.) Ach. St. Thomas; St. Jan. PARMELIA TINCTORUM Despr. [P. coralloides Mey. et Flot.] St. Croix. PHYSCIA INTEGRATA Nyl. St. Jan. RINODINA sp. St. Croix. VERRUCARIA sp. St. Croix. ARTHONIA RADIATA (Pers.) Ach. St. Croix. SCHIZOXYLON sp. St. Thomas. FUNGI During our exploration of St. Thomas and St. Jan in 1913, about 25 species of fungi were obtained, and Dr. Rose collected four others on St. Croix; manuscript record has been made of these. Twenty species collected by Mr. Ricksecker on St. Croix are 116 BROOKLYN BOTANIC GARDEN MEMOIRS listed by Dr. Millspaugh in his “Flora of the Island of St. Croix,” as determined by J. B. Ellis and F. D. Kelsey.'® Thirty-one species brought by various collectors to Copenhagen, determined by E. Rostrop, are recorded by Bérgesen and Paulsen in their ‘‘ Végétation des Antilles Danoises.”’ Seven species, collected on St. Thomas during the voyage of the ‘““Challenger,’’ are listed by M. J. Berkeley in Journal of the Linnean Society 14: 352. These records duplicate each other considerably, indicating a known fungus flora of somewhat over fifty species only. Inasmuch as there must be several hundred species on the islands, a list of fungi is deferred for further mycological field work. ALGAE!® “The Marine Algae of the Danish West Indies”’ is the title of a work, now appearing in parts, in which Dr. F. Bgrgesen, of Copen- hagen, is carefully describing and adequately illustrating the seaweeds of these islands. His adoption of the English language in this work makes it immediately serviceable to American students. Volume 1, including the Chlorophyceae (Green Algae) and Phaeophyceae (Brown Algae), was published in 1913 and 1914, and, at the date of writing, the first 240 pages of Volume 2, dealing with the Rhodophyceae (Red Algae) have appeared. Other papers of importance, dealing with the algae of the Danish West Indies, are the following: Borgesen, F. A Contribution to the Knowledge of the Marine Alga Vegetation on the Coasts of the Danish West Indian Islands. Bot. Tidssk. 23: 49-57. Figs. I-4. 1900. — Et Bidrag til Kundskaben om Algevegetationen ved Kysterne af Dansk Vestindien. Bot. Tidssk. 23: 58-60. 1900. [An abstract, in Danish, of the foregoing article. ] — Contributions a la connaissance du genre Siphonocladus Schmitz. Overs. K. Danske Vidensk. Selsk. Forh. 1905: 259-291. Figs. I-13. 1905. — An Ecological and Systematic Account of the Caulerpas of the Danish West Indies. K. Danske Vidensk. Selsk. Skr. VII. 4: 337-392. Figs. 1-31. 1907. — The Dasycladaceae of the Danish West Indies. Bot. Tidsskr. 28: 271-283. Figs. 1-9. 22 My 1908. —— The Species of Avrainvillea Hitherto Found on the Shores of the Danish West Indies. Vidensk. Medd. Naturh. Foren. K@benhavn 1908: 27-44. pl. 33. Je 1908. —— Some New or Little-known West Indian Florideae. Bot. Tidssk. 30: 1-19. pls. I, 2, Figs. I-11. 23 O 1909; II. Bot. Tidssk. 30: 177-207. Figs. 1-20. 9g D 1910. —— Some Chlorophyceae from the Danish West Indies. Bot. Tidssk. 31: 127- 152. igs: 1-13; 1911; Il, Bot. Lidssk: 32:) 241-272. Figs) 1-17.) mone: '8 The new species were described in Bull. Torr. Club 24: 207-209. 1897. 8 Contributed by Dr. Marshall A. Howe. BRITTON: FLORA OF THE VIRGIN ISLANDS 117 — The Algal Vegetation of the Lagoons in the Danish West Indies. Biol. Arbejd. tilegn. Eug. Warming. 41-45. Figs. I-9. IgITI. — Two Crustaceous Brown Algae from the Danish West Indies. Nuova Notar- isia 23: 123-129. Figs. 1-3. I9gI2. — The Species of Sargassum Found along the Coasts of the Danish West Indies, with Remarks upon the Floating Forms of the Sargasso Sea. I-20. Figs. 1-8. 1914. [No. 32 of a Mindeskrift for Japetus Steenstrup. ] Cleve, Peter Theodor. Diatoms from the West Indian Archipelago. [Virgin Islands and St. Bartholomew.] Bih.*Svens. Vet. Akad. Handi. 5%: 1-22 pl. 1-5. 1878. Annot. list. Dickie, George. Marine Algae Collected at St. Thomas during the Expedition of H. M. S. “Challenger.” Jour. Linn. Soc. Bot. 14: 312-313, 17 O 1874. ist: Millspaugh, C. F. Flora of the Island of St. Croix. Field Col. Mus. Bot. 1: 441- 546. 1902. On pp. 467, 468 is a list of 17 species of marine algae, deter- mined by Professor W. G. Furlow. Vahl, M. Endeel Kryptogamiske Planter fra St. Croix. Skrivt. Naturh. Selsk. 52 20-47.) Leo2: ENDEMIC SPECIES The approximate number of species native to the islands as re- corded, excluding fungi and algae, is 1,052, as follows: SMermatophyta, ics ceds scsi eae As 890 FAUCIIGLOMIINeCAe 1 260 a. cane 8 cc.) seme aes AI | BUENO} 1 DTA Wy ner een i ee 46 Dicheness, ascents he sia ce. oo ke GS 1,052 The numbers of Spermatophyta and Pteridophyta are not likely to be increased by further exploration, but there are probably some more Bryophyta and many more lichens to be obtained. As we know the flora at the present time, the following 27 species are endemic, at least to the Virgin Island group as a whole. Valota Eggersi (Hack.) Hitche. & Chase Agave Eggersiana Trelease Peperomia myrtifolia (Vahl) A. Dietr. Pilea Richardi Urban Coccolobis Klotzchiana Meissn. Zanthoxylum thomasianum Krug & Urban Galactia Eggersu Urban Malpighia pallens Small Malpighia infestissima (Juss.) Rich. Maytenus cymosa Krug & Urban Reynosia Guama Urban Sida Eggers E. G. Baker Psidium amplexicaule Pers. Calyptranthes thomasiana Berg. 118 nat BROOKLYN BOTANIC GARDEN Eugenia sessiliflora Vahl Chrysophyllum Eggersit Pierre Forestiera Egegersiana Krug & Urban Salvia thomasiana Urban Physalis Eggersiit O. E. Schulz Solanum conocarpum L. C. Rich. Wedelia cruciana L. C. Rich. Phascum sessile E. G. Britton Anthracothecium Breutelii Muell. Arg. Lecania euthallina Riddle Caloplaca subsequestra (Nyl.) Riddle Buellia prospersa (Nyl.) Riddle Mycoporellum ellipticum Muell. Arg. The endemic elements are, then, only about 2.6 percent. of the ive flora. A few other species are almost endemic, being otherwise known only on Porto Rico or on some other neighboring island. There are a few endemic species known on Tortola, and one on Anagada, but if the native species of these two islands, additional to those of St. Thomas, St. Jan and St. Croix, were taken into account, the percentage of endemism would not be increased. Porto Rico, with a very much greater area and much higher moun- tains, has about 13 percent of its species of Spermatophyta and Pteri- dophyta endemic. MEMOIRS WEATHER CONDITIONS AND PLANT DEVELOPMENT GEORGE P. BURNS Vermont Agricultural Experiment Station The effect of weather conditions on plant development has been one of the chief problems studied during the past few years by the ecologist, the agriculturalist, the forester and in some cases by the plant physiologist. The weather, however, is a variable mixture composed chiefly of different amounts of light—direct, diffuse, white, yellow, red, etc., or darkness; moisture—precipitation, humidity, soil-moisture, etc.; heat, temperature of the air and soil; wind, etc. Each of these component parts varies within short intervals of time and each has its effect direct or indirect on the living plants. The problems of the effect of weather conditions, then, is largely a physio- logical problem and such problems should be attacked only by means of accurate experiments under controlled conditions. The ecologists have been attempting to change from the old descriptive methods in which the results of a more or less accurate study of the vegetation of a given area were published. Sometimes this study was accompanied by a few tables of meteorological data gathered from a nearby U. S. Weather Bureau station. In only a few cases were attempts made to relate these data to the descriptive part of the study and one was often at a loss to know why they were included in the publication. This type of work has served a good purpose in a preliminary way but is now outgrown. More accurate methods have been introduced by advanced workers and ecologists have adopted the plan of gathering their own data with instruments placed in the field, the attempt being made to place them under the same weather conditions as those of the plants under consideration. The largest amount of data has been collected on evaporation rates by workers with atmometers. This is probably due to the fact that these instruments are inexpensive as compared with the cost of the recording instruments necessary for collecting other data. But they lack standardization, many kinds, shapes and sizes being in use. Since no atmometer can be made to work exactly as a plant, ecologists should adopt arbitrarily one type in order that data wherever col- lected may be compared. Some ecologists have gone deeply into this phase of the work and are well equipped with field instruments record- 119 120 BROOKLYN BOTANIC GARDEN MEMOIRS ing soil temperature, air temperature, humidity, number of hours of sunshine, wind velocity, precipitation, evaporation, etc. The fundamental problem, however, presents itself after the analy- sis has been made of the elements which enter into the compound “weather.’’ It is the experimental determination of the effect of those elements, singly and collectively, as measured by the data compiled, upon the physiological activities of the plant under con- sideration. This effect can only be measured by means of accurately conducted experiments in which very expensive apparatus is used. One of our problems is so to outline the work and to set forth its fundamental importance that those in authority will be moved to purchase the ecological equipment without which these agricultural and silvicultural problems cannot be studied. In the attempt to solve the problem above outlined some workers have used a “‘plant instrument.’ A given kind of plant has been grown by the side of atmometers, etc., at stations established under different climatic conditions and an attempt has been made to interpret their effect as registered by the “plant instruments.’”’ As an illustra- tion of the attempt to interpret meteorological data in terms of plant development let us take the work dealing with temperature. One method contemplates the subtraction of a constant from the tempera- tures recorded and considers that thermometric degrees in excess of this constant are available for purposes of plant development. A second method seeks to express growth-rate in terms of the velocities of chemical reactions. A third—the physiological method—attempts to take into account the optimum and maximum temperatures as related to plant growth, and the attempt has been made to develop one formula which will express the combined effect of rainfall, evapora- tion and temperature on plant growth. This represents but little more than an attempt to show what might be done if we had sufficient experimental data on the reaction of plants to the complex conditions known as the weather. Much of this work has been based on averages—averages for a month, a year, or a number of years. We read that a large amount of data assists in ‘‘smoothing out the curve” or that the “spas- modically jerky graph may be smoothed.’ It is certainly true that in some cases the curve should not be smoothed out, because it is the spasmodic graph that shows sudden changes and the extremes. The burden of this paper is to show that, in some cases at least, averages for long periods are of little value as compared to the importance of the data obtained for certain critical periods in the conditions of the environment as shown by the ‘‘spasmodic graph.” Data collected for a short period in the summer may be very important, but are by BURNS: WEATHER CONDITIONS AND PLANT DEVELOPMENT 121 no means as valuable as those gathered by recording instruments during long periods of time. These latter data are valuable, however, primarily because they cover critical periods during which the environ- mental conditions are most severe for plant development. The limit- ing factor is not the average for the long period, but the maximum or minimum for any factor or group of factors during certain critical short periods of the longer season under consideration. This may be made more clear by illustration. The effect of shade on the develop- ment of white pine seedlings was under study. Lath shades known as “full-shade’’ and “half-shade’’ were used. It was found that germination took place sooner and that larger numbers of seedlings were produced in the “no-shade’”’ bed than in those partly or fully shaded. The temperature of the soil was the controlling factor. The average soil temperatures computed from readings recorded every two hours with a Friez machine for the 24 days during the germination period were “‘full-shade”’ 47° F., “half-shade’’ 46.8° F., ‘‘no-shade”’ 49° F. These differences are too slight to have been responsible for the observed differences in germination. If we look at averages only and shut our eyes to the daily fluctuations we would conclude that soil temperature was not the controlling factor. When, however, the records were examined for extremes it was found that on certain days temperature variations occurred of as much as 20° F. as between the soils of the different beds, the soil of the ‘‘no-shade’’ bed reaching 73° F. It is easy to believe that such differences may constitute a controlling factor, in view of the fact that Atterberg has shown that this temperature is about the optimum for germination of these seeds. One evening the nursery foreman reported that every seedling in our nursery was dead. Examination showed that the white pine leaves which the day before had been a beautiful green were brown and apparently dead. A closer study showed that the ends of the leaves including about one third of the leaf were dead. However, very little of this leaf browning occurred in another nursery located near a river bank, protected from the prevailing wind and on a richer soil. In the upper nursery where the browning occurred we had a number of recording instruments but unfortunately none in the lower nursery. The records show: that three slight showers and one heavy rain (1.28 inches) had fallen just previous to the appearance of the trouble; that for five days the sun had shown from nine to twelve hours daily; that a very heavy wind blew for three days before, especially while the sun was shining; that the humidity dropped daily below 50 percent, one day reaching 35 percent; that the air temperature was usually below 75° F.; that slightly protected areas in the upper nursery showed less damage than did the rest of the nursery. 122 BROOKLYN BOTANIC GARDEN MEMOIRS It is of course impossible with any degree of certainty to determine from these data why the trees in one nursery suffered severely, whereas those in the other nearby location were but slightly affected. How- ever, if one were to hazard a guess he might say that it was due to excessive transpiration, the chief immediate factor being differences in wind velocity. A similar result has been obtained experimentally when trees which have been shaded were suddenly exposed to sun and wind. The next day they showed “tip-burn”’ of the pathologists, or a ‘physiological disease’? whatever that may be. One fact is clear. The death of the leaves was not due to the average conditions prevailing during the summer. During the winter months the average soil temperatures in the nursery for depths of three, six and twelve inches were 35.8° F., 37.9° F., and 38.5° F. Each figure is the average of 2,100 readings taken every two hours from the record made by a Friez machine. They show little differences in temperature at the various depths given. When, however, the record is examined for critical periods it is found that probably the most important season was that from March 28 to April 18. During this period of 21 days the soil three inches deep froze and thawed sixteen times, at six inches, nine times and at twelve inches four times. Similar data collected in the adjacent forest showed that the soil both at six and at twelve inches thawed only once. We have no experimental data which determine the physio- logical meaning of these facts, but it is easy to surmise that in studying the effect of soil temperatures on plants we will not go far afield if we study carefully conditions obtaining during these critical periods as well as indeed full more than the general averages for the entire winter. Numerous examples could be given to show that averages extending over long periods for humidity, sunshine, wind, air temperature, etc., not only explain little but, on the other hand, conceal the essential facts. In all study of the relation of weather conditions to the develop- ment of plants the importance of critical periods in the environment must be taken into consideration. MODERN APPLICATIONS OF BOTANY MEL T. COOK New Jersey Agricultural Experiment Station It is very doubtful if any science is so thoroughly misunderstood by the public as the science of botany. To the average layman it is usually a study of flowers which usually involves harmless collections, classifications and mysterious Latin names; a study for the faddist; a study without applications of any value whatever. It is strange that a subject dealing with organisms upon which we are dependent for practically all of our food, clothing and fuel, a large part of the material for building and the manufacture of useful implements of various kinds, and most of our drug products should be so misunder- stood. Yet, even the educated layman knows more about the Panama canal than he does about wheat, more about flying machines than he does about potatoes, and more about the Woolworth building than he does about cabbage. The names of great and near great military leaders, statesmen, ministers, physicians, architects, theatrical stars, ball players and pugilists are familiar to those millions, while very few can name a single person who has contributed to the feeding and clothing of mankind. In fact, few people, even among the educated classes, realize that agriculture, horticulture and forestry are in reality specialized branches of botany. A brief statement of the early history of the subject may offer an ex- planation of this anomalous position of our science. Botany had its rise in the development of the medical professions, in the efforts of the practitioner to determine the uses of plants in the art of healing. This resulted in the study of local flora of a number of the most advanced countries and also the search for plants in foreign countries. Very naturally, the great number of species of plants forced these early students to formulate some system of classification whereby their materials might be catalogued. With their increasing knowledge of these species, it became necessary to devise new systems until finally this phase of the subject became all important. In the meantime, the medical profession gradually discontinued the use of the less important of the medicinal plants for those that were most easily obtained, most economical in preparation and most efficacious in use. A little later, we find the physician studying the crude drug and 123 124 BROOKLYN BOTANIC GARDEN MEMOIRS . a little later the prepared drug and paying little or no attention to its origin. Thus the two professions developed along diverging lines. In the meantime, the invention and development of the microscope opened new and interesting fields to the botanists as well as to other Scientists and also resulted in the rise of bacteriology, which has had such a marked influence on many lines of work, especially medicine. At the same time the scientific study of agriculture was beginning to attract attention but, unfortunately, it is not an outgrowth of botany. Chemistry became the first sponsor for this new field of research and the first directors of many of our American agricultural experiment stations were chemists; they studied the soils and de- veloped formulas for fertilizers—for what? To make plants grow, to increase plant production, and thus the problem of plant growth was taken by the chemists instead of the botanists. Horticulture was very closely associated with botany and the developments of horticulture and botany were combined in many of our agricultural colleges. In many cases these soon came to be known as departments of horticulture, the botany becoming a vanish- ing factor; but in later years botany has re-entered these colleges as an independent, but in many cases a secondary subject. In those agricultural colleges in which botany has had a continuous existence, the lines of research were by no means the same. In some cases, they studied weeds and devised methods for their control; in others, they co-operated with the horticulturists in the study, introduction and improvement of valuable food and fiber plants; in others they studied the causes and methods of controlling plant diseases, but in many cases the second phase of the subject was quickly taken over by the now independent departments of horticulture. It is impossible to tell just what the result would have been if our botanists of a quarter of a century ago had been as energetic in the development of the applied side of botany as the chemists were in the development of the applied side of chemistry. But it is reason- able to suppose that the results would have been similar, and that we would have today, not only the applied phases of botany, but we would also have far more workers on technical problems. The future of botany in America is brighter that at any time in its history. It is a recognized subject in our universities, in arts and in agricultural colleges. It is recognized, both as a cultural subject of great value and interest and as a science with a direct bearing on the affairs of mankind. The botany of today means not only tax- onomy, morphology, cytology and physiology as purely scholastic subjects but all in their relation to applied plant physiology, plant breeding and plant pathology with a direct bearing on horticulture, agronomy and forestry. On COOK: MODERN APPLICATIONS OF BOTANY 12 Plant growth is no longer a problem for chemists but for the plant physiologist, who is trained not only in botany, but in chemistry, physics and geology. Plant physiology has outgrown the expectations of its most enthusiastic devotees of a decade ago, and no one can foretell its future. It will doubtless result in important changes in agricultural methods. Plant breeding, along the lines of artificial selection, is very old; in fact, it must have originated with the first steps in civilization. Many of our valuable economic plants were selected, grown and used by man before the beginning of written history and many im- proved varieties have been developed by self-taught, practical workers, men of great natural endowments and keen powers of observation. However it is none the less true that they are the products of the workings of natural laws and that a knowledge of these laws enables the present generations to work more rapidly than their ancestors. Many of our modern plant breeders are very properly more interested in researches leading to a knowledge of these laws than in their appli- cation. A law fully established and well understood will very soon be utilized by those interested in increased production. But the breeder should not loose sight of the very great value of plant breeding to agriculture. The final and true standard of measure of the value of any science must be in terms of its contributions to the welfare of mankind. Plant pathology is one of the last of these branches of applied botany to be considered. It had its rise in the taxonomic study of fungi, many of which were recognized as the causes of plant diseases. Therefore, this study very naturally led to the study of methods of control. Indefinite and uncertain methods for the control of plant diseases have been used from time to time for more than a century. But a lack of definite knowledge of the causes and the physiology of these diseases and the actions of the remedies made the results very uncertain and very soon led to their disuse. Modern plant pathology had its beginning in the works of de Bary and Berkeley, but did not make much progress until the latter part of the last century. The progress during the last decade has been rapid and has emphasized the necessity of many lines of study, such as a more thorough knowledge of the life history and taxonomy of the parasites, a knowledge of the physiological factors influencing both host and parasite and a knowledge of the physiological effects of the fungicides. It is also extremely important that we make extensive investigations on that ever increasing number of diseases which cannot at this time be attributed to any definite organism. The prosecution of these lines of investigations means more in- 126 BROOKLYN BOTANIC GARDEN MEMOIRS tensive researches in the taxonomy, morphology and physiology of the fungi and other organisms that cause diseases; in the morphology and physiology of the flowering plants; and plant breeding. The directing ideal in physiology and plant breeding must be the improve- ment of the plant for economic purposes, the development of resistance to disease and the increase in plant production. Many phases of plant pathology are practically untouched. The greatest advancement has been made in the study of the diseases of orchard fruits, much has been done in the study of cereals, shade and forest trees, and certain truck crops, such as potatoes. While much work still remains to be done on the diseases of these crops, much more is necessary on miscellaneous truck crops and on ornamentals. The fact that truck and ornamental crops are grown under glass presents new and complicated problems of the greatest economic im portance. : Many people, even botanists, have the idea that all phases of applied botany must be restricted to agricultural colleges. This is an unfortunate error which tends to broaden the gap between the botany on one side and horticulture, agronomy, forestry, etc., on the other. Only recently, a well-known government plant pathologist told the speaker that he had no great difficulty in securing young men trained in plant pathology but that, unfortunately, many of them were not trained in botany. Applied botany is in very great need of workers who have a thorough fundamental training in botany plus a specialized training in applied botany. Much of this work can be done to an advantage in our universities provided the proper viewpoint can be obtained. I use the term ‘‘viewpoint’’ guardedly, for while it is true that many of our workers in applied botany are poorly trained in fundamental botany, it is also true that many of our uni- versity men are about as well fitted for applied botany as the students of Hebrew. It has been said that no one can apply a science unless he has ‘learned the science, but it is equally true that some learn a science that cannot be applied. The suggested ‘applications in some technical papers compare very favorably with the comic sheet in the Sunday papers. But the few lines of work indicated in this paper do not include all that are open to the botanists. Many of the manufacturing industries are needing, and will need for years to come, many men trained in botany and biochemistry. Some time ago the writer was asked to recommend such a man to make investigations on cellulose. Failing to find such a man, the company employed a chemist. The manufacture of rubber is another industry in which the services of a properly trained botanist can be very useful. And there are many COOK: MODERN APPLICATIONS OF BOTANY 127 other lines of work too numerous to mention in the short time avail- able for this paper. Furthermore, in the very near future, America may be called upon to furnish botanical workers for the world. A prominent London journal has already called attention to the necessity for the development of the agricultural resources of Great Britain’s colonies, and admitted that the workers must come from America. The great resources of South America are practically undeveloped. Thus far, those countries have called on European countries for most of their workers, but in the near future they will probably turn to this country. With the close of the great international war, now in pro- gress, the United States will probably become the great education center of the world, but we must give educational work largely along industrial lines. Are the American botanists prepared to meet the new demands? STUDIES IN THE GENUS GYMNOSPORANGIUM—I. NOTES ON THE DISTRIBUTION OF THE MYCELIUM, BUFFER CELLS, AND THE GERMINATION OF THE AECIDIOSPORE B: ©: DODGE Columbia University ‘The sporophytic mycelium of different species of Gymnosporangium exerts an influence in connection with the growth of the tissues of the cedar hosts which is manifested in a variety of ways. It is not clear just why one species will cause the formation of a rather fleshy gall, while another species will lead to the development of a witch’s-broom or a hard, woody burl. A study of the interrelationships of host and parasite, especially the more intimate association of the hyphae and the host cells may help to solve some of these interesting questions. I wish to report briefly at this time the results of some studies made to determine: (1) the distribution of the mycelium in an infected leaf; (2) the possibility of its spread from leaf to stem; (3) the degree to which it spreads up and down; (4) its distribution in wood, phloem and cortex; (5) the distribution of haustoria. Four-nucleated aecidiospore germ-tubes of Gymnosporangium transformans, and the formation of buffer cells in the teleutospore sori of G. fraternum and G. transformans will be noted. Farlow! determined the general distribution of the mycelium in the host for a number of American Gymnosporangia and described with considerable clearness the primary effects of the parasites on the tissues of the host plants. He found that the burls on Chamaecyparis infected by G. biseptatum are probably the result of a stimulation of the cambium by the hyphae mainly distributed in the cambium region. There appeared to be very little in the nature of a deleterious effect of the parasite on this host. It was evident to him, however, that the actual presence of mycelium in a given tissue is not necessary to account for distortions or abnormalities. In G. Ellisii the fungus interferes with the normal growth of the host, producing proliferations and swellings of the stems and branches. This may be due to a dis- turbance in the nutritive processes, the primary cause of which may ‘Farlow, W. G. The Gymnosporangia or Cedar Apples of the United States. Am. Mem. Boston Soc. Nat. Hisc. 1-38. pl. 1, 2. 1880. 128 DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 129 lie at some distance. Farlow noted that the hyphae of this species are exceptionally large and that the brown mycelium runs down into the wood and along the medullary rays and also makes other brown patches extending some distance in circular areas between the annual rings. The greater part of the mycelium is found near the cambium and large masses of it are collected at points in the bark in prepara- tion for the formation of sori. Wornle? made an extensive study of the relationships of host and parasite in nine species, and his report published in a forestry journal furnishes a valuable contribution on the subject. He en- deavored to determine the particular tissues with which the mycelia are associated and stated his conclusions with considerable positive- ness. G. Juniperinum was of special interest to him inasmuch as he supposed that the sori found on leaves as well as those on small twigs belong to the species that produces larger sori on the main stems. He could see that the mycelium in an infected leaf was connected with that from a small twig. He also learned that the leaf form is _ per- ennial. Four successive cork callus formations were found in one case, showing that for four years a sorus had been developed at the same point on the leaf. In the stem-inhabiting type he found that the mycelium is present in the wood as well as in the bast and cortex. Radially placed strands of parenchyma accompanied by mycelium are common in the wood; ‘“‘Schlafende Augen”’ he calls them. Hyphae are intercellular, and he noted in some cases the presence of haustoria. Although Wornle was not himself clear regarding the relationship of the three forms of the rust which he called G. Juniperinum, he was inclined to believe that the fungus gains entrance through the leaves, the mycelium later running down the twigs and into the main stem, where it becomes firmly established. Fischer? has shown that this was a false assumption since W6érnle was dealing with at least two species, but the accuracy of Wérnle’s observations is not ques- tioned. The mycelium of G. clavariaeforme, according to Wo6rnle, is not present in the wood, although considerable transformation of tracheid tissue is to be seen in infected stems; arcs and sectors of this tissue are replaced by parenchymatous cells. He found, however, no mycelium in such areas. As the mycelium is generally distributed in the cortex and bast, he assumes that the cambium is in some way stimulated to develop more than a normal amount of wood cells, some 2 Wornle, P. Anatomische Untersuchung der durch Gymnosporangium-Arten hervorgerufenen Missbildungen. Forst. Nat. Zeits. 3: 68-84, 129-172. 1894. 5Fischer, E. Studien zur Biologie von Gymnosporangium juniperinum. Zeits. Bot. 1: 683-714. f. I-8. 1909; 2: 753-764. I9I0. 10 130 BROOKLYN BOTANIC GARDEN MEMOIRS of which are inhibited in their growth, lacking bordered pits and having thin walls, that is, are more in the nature of parenchyma. Wornle was especially fortunate in having the opportunity for con- sultation with Hartig and Tubeuf in his work, but he was handicapped in studying American species by being restricted to a limited number of dried specimens. He agrees in the main with Farlow’s account of the location of the mycelium in G. biseptatum and G. Ellisit, although he makes no mention of Farlow’s work. He concludes further that the mycelium of G. biseptatum is intercellular and is entirely absent in the wood. The tracheids are somewhat irregular and have thinner walls than ordinarily. He found that the hyphae of G. Ellisti are about 8 w in diameter and are present in the wood, bast and cortex. The brown hyphae are associated with brownish cells which together make easily recognizable patches. The mycelium here also is strictly intercellular. His study of a three year old stem of red cedar infected with G. clavipes disclosed the fact that the tissues of the host are only slightly affected. The mycelium is distributed not only beneath the sorus but in the whole periphery of the twig, especially in the bast region. The wood is entirely free from the fungus. Wo6rnle pre- dicted that this rust must develop sori one year after inoculation because in this three-year-old stem he found traces of two former sori, one above the other. Harshberger’s account* of the relationships of hyphae and host cells deserves special consideration, inasmuch as it does not agree in certain important particulars with the statements made by Farlow and Wornle. He finds that in G. biseptatum the mycelium is quite generally present in the wood region where he states the hyphae are for the most part strictly intracellular. They run down through the lumen of a tracheid, pass out through bordered pits, enter an adjacent tracheid, or move over to medullary ray cells which they penetrate and thus become established where they receive nour- ishment sufficient to maintain their perennial growth as the wood of the burl increases in diameter. He believes that the hypha actually in the lumen of the cambium cell is responsible for the stimulation of this cell to produce abnormal amounts of wood! He describes and figures these intracellular hyphae in much detail, especially the hyphae in longitudinal sections of wood. The explanations accom- panying his figures leave no doubt of Harshberger’s opinion regarding the identification and location of intracellular hyphae. He lays much stress on the presence of “plugged’’ tracheids. He believes that they are caused by the mycelium with which they are generally asso- 4 Harshberger, J. W. Two Fungous Diseases of the White Cedar. Proc. Acad. Nat. Sci. Philadelphia 1902: 461-504. pl. 22, 23. DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 131 ciated. Hyphae may sometimes be intercellular. Swollen and nodular hyphae are not infrequent. Harshberger questions whether haustoria are ever present. It is well known from the work of Hartig and others that hyphae of wood-destroying fungi are capable of boring through lignified cell walls. Such fungi obtain their nourishment by activities leading to the disorganization of wood cells. The rusts are highly parasitic and haustoria play an important part in their nutrition. It would be interesting to find that such trunk parasites as G. biseptatum and G. Ellisii are more like the common heart rot fungi than they are like other rusts where the hyphae crowd in between the cells or mass in the intercellular spaces. My own observations do not support several statements made by Harshberger. (GYMNOSPORANGIUM ELLISII I have succeeded in infecting Chamaecyparis by spraying potted cedars with aecidiospores of G. Ellisii (G. myricatum). Several cedars naturally infected and bearing brooms of different ages have also been grown in pots, so that I have had an abundance of material in all stages of growth for study. The sorus usually matures about twenty-one months after inocu- lation. Where young leafy branches have been infected we find that the sorus may break out either in the leaf axil or through the leaf itself. At this time there is very little distortion of the twig. The primordium of the axial sorus is partly in the tissue at the base of the leaf and partly in the stem cortex beneath. Where the sorus emerges through the leaf we find that there is an increase in the number of mesophyll cells and the sorus primordium is not far below the epidermis. Strands of hyphae can be traced down to the short vein and into the woody portion of the stem. Serial sections show that the mycelium does not travel up and down the stem very rapidly; in some cases only one or two cm. in the first two years. Where a rapidly growing main stem is infected the hyphae run as much as five cm. in the same time. Trunks thirty years old have been cut and one such shows traces of mycelium for a vertical distance of only about ten cm., although the fungus had been active during the life of the tree, thirty years. Sections taken from various parts of a small artificially infected plant bearing a dozen potential witches’ brooms show that each broom will be the result of a separate infection. The mycelium does not enter at one point and spread through the entire plant. However, if the original infection should be at the growing point of the main stem a broom is formed that permanently dwarfs the plant. The mycelium invades every tissue except the cork. It is found in patches in all of the annual rings, and is espe- 132 BROOKLYN BOTANIC GARDEN MEMOIRS cially abundant along some of the medullary rays. It is not evenly distributed. The hyphae seem to travel in fascicles and they are everywhere intercellular. Sections of the wood show that there are strands of parenchyma that, from appearances, would seem to be burrowing through the wood, thrusting the tracheids aside as though endowed with great power. These same parenchyma strands are also found in the cortex. They run in almost every direction. Hyphae are always associated with them. Tracheids in infected areas of the wood are considerably modified. The walls are thinner, the cells are prismatic and in many cases have failed to develop bordered pits. The walls of such cells frequently appear to be broken down or crushed in and partially disorganized. It may very well be that the fungus has some power to disorganize lignified cell walls. Wherever hyphae occupy the lumen of a cell it is likely to have been the result of such mass action. There is no boring through the walls nor entering tracheids through bordered pits. The ‘“Schlafende Augen,’ or parenchyma strands, in the cortex and along the line of medullary rays in the wood as well as the patches of abnormal or partially developed tracheid tissue are the result of the stimuli proceeding from hyphae that were nearby at the time this tissue was being de- veloped. It is difficult to understand how a cambium cell harboring a hypha could divide at all, or how a tracheid could change its form once it has become lignified. The cambium reacts in such a way as to cut off by the excessive development of tracheids certain fascicles of hyphae and thus check the radial and longitudinal advances of the fungus. The apparently isolated patches of mycelium found in the heart wood are nevertheless quite generally connected above or below with some radially placed strand that ultimately reaches the cortex. This may be the main reason why one finds living hyphae deeply imbedded beneath several rings of wood. Haustoria may occasionally be found in cells of the cortex medullary rays, but they are not abundant. Some of these haustoria are bi- nucleated. There seems to be no question that Wo6rnle was right in stating that the hyphae of G. Ellisii are intercellular. GYMNOSPORANGIUM BISEPTATUM Harshberger and Wornle disagree on a second important point in their studies of G. biseptatum. This relates to the presence or absence of mycelium in the wood cylinder of the cedar. I have as yet been unable to infect the cedar with this species. I have studied specimens naturally infected and especially one from a DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM § 133 small plant which I was able to transplant and grow in the greenhouse. In 1915 this small burl bore two sori. The same burl bore six sori in 1916. The branch was six years old when cut. The mycelium spreads quite evenly through the cortex and is especially abundant beneath a sorus where we find one or two large haustoria in nearly every cortex cell. The medullary ray cells of the cortex are likewise attacked and the mycelium penetrates down to the cambium. The walls of the tracheids are somewhat thicker than usual, in this respect differing from the specimens examined by Wornle. I have been unable to find any intracellular hyphae, and in this six-year-old branch there are certainly no hyphae inside of the cambium ring, that is, in the wood cylinder, such as Harshberger describes. The most striking feature about this fungus is the great abundance of large haustoria found in nearly every cell of the cortex in the vicinity of a sorus. GYMNOSPORANGIUM CLAVIPES The red cedar may be infected with G. clavipes without difficulty by spraying with aecidiospores. Plowright® states that it takes two years for G. clavariaeforme to mature sori, but Tubeuf*® found that sori developed one year after inoculation of the juniper. My experi- ence with G. clavipes may serve to explain this discrepancy. On August I, 1915, a small cedar was inoculated with Gymno- sporangium clavipes. A few sori appeared in 1916 on what was, in 1915, the growing region of the main stem. In 1917 sori burst out quite generally over the plant. ‘The question has arisen: Is it possible that from the original point of infection of 1915 the mycelium ran down the main stem out into the branches where further sori formed in 1917? Inspection showed that the sori were not evenly scattered along the branches, but appeared in groups with intervening spaces of some length between, varying from one to several cm. Serial sections of some of the smaller branches made at points between groups of sori do not show the presence of mycelium. For several inches near the top of the main stem the sori are so close together that mycelium appears to be continuous. It is noteworthy, however, that there are no sori on those parts of the plant that have grown since the plant was inoculated in August, 1915. The mycelium is intercellular and lies for the most part well out in the cortex just beneath the cork, some hyphal ends even pushing in between the inner cork cells. It may require only one year for full development at the 5 Plowright, C. B. British Uredineae and Ustilagineae. 1893. 6 Tubeuf, C. Mitteilungen iiber einige Pflanzenkrankheiten. Zeitschr. Pflanzenkr. 3: 201-205. 1893. 134 BROOKLYN BOTANIC GARDEN MEMOIRS growing point where abundant food is available, or even take two years in regions less favorably located. The characteristically binucleated haustoria are of large size and are easily demonstrated. I have examined several stems three years old but do not find that the mycelium spreads out through the entire cortex and into the bast as described by Wornle. G. clavipes brings about less increase in development of wood tissue than G. biseptatum, but this may be due to the fact that the mycelium does not approach the cambium as closely. This species may develop strictly foliicolous sori in which case the mycelium is very limited in extent. The sorus is then not deep seated. Haustoria can be found in epidermal cells. (GYMNOSPORANGIUM TRANSFORMANS I have previously reported’ that two leaf-inhabiting species of Gym- nosporangium can be distinguished on Chamaecyparis. The account of the cultures in support of this statement is being published in another paper. For convenience I shall call one form G. transformans. Its aecidial form is Roestelia transformans on Aronia. Gymnosporangium fraternum is an appropriate name for the second leaf form which infects Amelanchier. The aecidium is very similar to that of Roestelia Botryapites; | am not prepared to prove that it is R. Botryapites. A cytological examination of cedar leaves infected with G. transformans and G. fraternum reveals further characteristics by which they may be distinguished. If we section a leaf of Chamaecyparis infected with G. transformans, we find that the mycelium is especially abundant in the large inter- cellular spaces of the spongy mesophyll and the hyphae push in be- tween the cells of the palisade on all sides. No hyphae are to be found in the epidermis. If the section includes the short vein of the leaf we see that hyphae are prevented in some way from entering the vascular tissue. There is an irregular row of large cells surrounding the vein of the leaf. These pericycle(?) cells do not normally form a_ per- fectly closed ring; it is occasionally broken by smaller supporting cells. In regions where the hyphae reach the large cells one can find, here and there, that they have been penetrated by one or two haustoria. Such infected cells are about one third larger than usual. The cytoplasm is rather dense, including considerable stored food and takes the gentian violet stain somewhat deeply. The nuclei appear to be quite normal. MHaustoria are also occasionally found in mesophyll and palisade cells. The cells of the mycelium are bi- nucleated and the nuclei stand out very clearly, especially where 7 Dodge, B. O. Report on further cultures of Gymnosporangia. Paper read at the December meeting of the Botanical Society of America, New York, 1916. DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 135 leaves have been fixed at a time when the sorus is fully matured. The mycelium is confined to the leaf bearing the sorus and does not ordi- narily invade the stem at any point along the line of attachment. The large cells surrounding the leaf vein appear to prevent the my- celium from entering the phloem of the stem. At the point where the sorus is to be developed, we find a well- defined pseudo-parenchyma, the cell walls taking the orange stain. The upper cells of this pseudo-parenchyma are somewhat enlarged and elongated. These are likewise binucleated. They soon begin to swell, lose their cytoplasm, and the nuclei degenerate. In order to show these upper cells in this condition the material must be fixed at the earliest possible time that an infected leaf can be distinguished, Fic. 1. Section of a sorus of G. transformans on a leaf of the southern white cedar at the narrowest portion of the young sorus. The epidermis is broken up On either side, only traces of the cuticle and fragments of the cell walls being visible. At the center epidermal cells are still visible. A number of buffer cells in various stages of degeneration can be seen, and binucleated teleutospore buds growing through the buffer cells are common. that is, when a spot appears as a slight, waxy, translucent, light orange blister. In such cases the epidermis may not have been ruptured - and fixation of the mycelium is not apt to be of the best, unless the leaf is cut through. The upper cells mentioned become mere bladdery sacs and during this process of swelling the inner walls of the epidermal cells, and hypodermal cells when present, are broken down either by enzyme action or by actual pressure, and the heavily cutinized epi- dermis is lifted up and split open (Text-fig. 1). The splitting usually occurs in a line along one side of the leaf, but very often the split runs longitudinally down through the middle. Sometimes two sori develop side by side on the same leaf. The bladdery cells evidently function as buffer cells to disrupt the epidermis. These buffer cells perhaps represent simply the first series of teleutospore mother cells 136 BROOKLYN BOTANIC GARDEN MEMOIRS that are sacrificed in order that the epidermis may be broken open (Text-fig. 2). The true basal cells now grow out through the buffer cells, forming a club-shaped bud which soon becomes binucleated. These nuclei divide and the stalk is cut off; later three pairs of nuclei can be seen and the wall is formed between the two cells of the teleuto- spore (Text-fig. 3). Quite mature spores can be found along the line where the epidermis first ruptures and all stages in their development can be seen further back. Fic. 2. A portion of a sorus of G. transformans showing two rather pointed buffer cells forcing epidermal cells aside, fragments of the walls of the epidermal cells lie just above the buffer cells at the right. Four young teleutospore buds are visible. Fic. 3. Teleutospore of G. transformans. The gametophytic stage on Aronia may sometimes attack the young stem, giving rise to an irregular herbaceous gall, which becomes covered with horn-like projections from which the aecidia arise. If such a plant is kept in the greenhouse all winter, aecidiospores will continue to be formed in some cases for several months after the leaves have fallen from the plant. These spores are regularly bi- nucleated and possess 7 or 8 germ pores irregularly distributed (PI. I, Fig. 1). The spores germinated on agar or water frequently form a swollen pouch near the tip of the germ tube. The nuclei of the spore push through the germ pore, apparently one closely following the other (Pl. I, Fig. 3). At the next stage we find two nuclei lying in the germ tube just outside of the pore (PI. I, Fig. 4). These nuclei then migrate further out into the tube and come to lie in the swollen pouch (Pl. I, Fig. 5) where they presumably divide conjugately, since many cases have been observed where there were four nuclei, in pairs, lodged in this portion of the tube (Fig.'6). The germ tube now elongates rapidly and branches freely at the tip (Fig. 8). The four nuclei move forward and may lie along the tube in a row in the wider portion at the end or they may be distributed, one nucleus in each branch, or DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 137 two lying further back, the other two occupying separate branches. A cell wall is finally laid down, cutting off the outer portion of the germ tube containing the nuclei and most of the cytoplasm (Figs. 8-10). This curious method of germination is in no sense similar to the development of a promycelium, though four nuclei are pro- duced in each case. It may be that it is fairly common among the rusts as Sappin-Trouffy has pointed out.’ Whether or not it is pos- sible to find an appropriate artificial medium for the development of the mycelium of a rust in artificial cultures, it would seem that such cases as these afford at least a starting point. Four-nucleated germ tubes are the rule in these cultures, but fully developed tubes with only two nuclei are not difficult to find (Fig. 11). In some of my cultures in which the petiole of a leaf had been infected at the junction with the blade, it was found that the winter bud was larger than usual. The mycelium must have run down the petiole and become established in the bud. When such plants were put in the cold frame over winter and taken out in the spring, these buds developed small leaves which at once became evenly covered with spermogonia and later were transformed into large galls from which aecidia developed quite normally. In some cases the mycelium seems to penetrate into the tissues of the stem where a spindle-shaped swelling or burl is formed. In the following spring a green gall bursts out through the cork, forming a nodular swelling outside and from this spermogonia and aecidia are produced. I have had several cases in which Roestelia transformans has survived the winter and developed aecidia the following spring The same is true in my cultures of R. Botryapites. In October, 1915, winter buds of six Amelanchiers showed signs of being infected. All of these survived the winter and developed spermogonia and ripened aecidiospores in the month of June, which is several months earlier than they can be found in nature. These are not cases where the formation of an aecidium has simply been delayed. On the con- trary, an entirely new crop of spermogonia arises from newly formed tissue, new gall growth, and we find the aecidia developing as in normal cases of infection with sporidia. (GYMNOSPORANGIUM FRATERNUM The buffer cells in the teleutospore sori of G. fraternum are much more striking in appearance, forming as they do a perfectly even palisade layer that frequently extends entirely across the sorus without interruption (Text-fig. 4). This is a very characteristic feature of 8 Sappin-Trouffy, P. Recherches histologiques sur la famille les Urédinées. Le Botaniste 5: 59-244. f. I-69. 1 D 1896. 138 BROOKLYN BOTANIC GARDEN MEMOIRS the rust. The difference in the shape of the buffer cells of G. trans- formans and G. fraternum corresponds roughly with that between the teleutospores; they are comparatively long in the latter species. On January 30, 1917 a potted plant naturally infected with G. fraternum was taken from the cold frame and examined. Several leaves showed by the presence of slight yellowish spots that they would develop sori. Sections of one leaf cut on this date showed Fic. 4. Section of a leaf of the white cedar infected with G. fralernum show- ing the layer of buffer cells at the time when the epidermis has been quite completely disorganized. At the right a large palisade cell in the process of disorganization, but the nucleus is still visible and a haustorium is present. that the pseudo-parenchyma or teleutospore primordium was well marked. The buffer cells were mostly without granular contents and nuclei (Text-fig. 5, 4). In a few cases fragments of the degenerated nuclei could be seen. By February 2, a sorus taken from the leaf of the same plant showed great numbers of teleutospore buds in the 2- and 4-nucleated stages (Text-fig. 5, C). A few buds had 6 nuclei, and the stalk cells of these had been cut off. Buffer-cell walls were just visible as narrow irregular lines showing most distinctly at the base of the young teleutospore. Two days later, February 4, cross-walls had been formed in many spores but nuclear fusion had DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM 139 not occurred (Text-fig. 5, D). Remnants of buffer-cell walls were now difficult to find. A mature spore is shown in Text-figure 5, EF. The mycelium of G. fraternum penetrates through the leaf in every direction. The mesophyll cells are usually somewhat enlarged and are packed rather closely together with small intercellular spaces. Haustoria are quite abundant in such cells. The cells surrounding the vein are especially affected. They appear to be filled with minute granules and as many as eight or ten haustoria can be found ina single cell, the more common number being two to four. A complete ring of these large cells is formed. This is due to an increase in number as well as their larger size. This, taken in connection with the increase in the mesophyll tissue, gives the leaf a slightly thicker and more compact appearance. The mycelium does not invade the Fic. 5. a, pseudoparenchyma with buffer cells; 6, 2-nucleated stage of the young teleutospore; c, 4-nucleated stage; d, 6-nucleated stage with cross walls; e, small spore after nuclear fusion, vascular tissue even to the extent of penetrating the phloem of the stem. Both G. transformans and G. fraternum are capable of producing sori two or three years in succession, the latter may produce a sorus even after the leaf has apparently died. While haustoria are more numerous and attack the individual cells more vigorously in the case of the latter species, G. transformans seems in some way to be more destructive, as infected leaves more frequently die after maturity of the first sorus. Of the two species, G. fraternum is clearly the more nearly related to G. biseptatum, both from the nature of their teleuto- spores and the similarity of the aecidia in the two species. If the mycelium of G. fraternum is ever able to push in beyond the large cells surrounding a vein and get into the central cylinder of the stem, therefore nearer the cambium, we should look for a stimulus such as 140 BROOKLYN BOTANIC GARDEN MEMOIRS might lead to the formation of a greater amount of wood tissue such as we find in the burl of G. biseptatum. G. fraternum has almost con- stantly 2-celled teleutospores, 3-celled spores are exceedingly rare. In G. biseptatum 3- and 4-celled spores predominate. I should be highly gratified to learn that the change in environment from the leaf to the stem, or more exactly from cortex to phloem, by the fungus could bring about such a decided change in the structure of its spores. EXPLANATION OF PLATE I Gymnosporangium transformans Stages in the Germination of the Aecidiospores. XX 750 Fic. 1. A binucleated spore. Fic. 2. The germ tube has pushed out, the nuclei are still within the spore and do not show appreciable change in form. Fic. 3. One nucleus is crowding through the germ pore, the other lies beneath. The ‘‘pouch”’ is formed just back of the tip of the germ tube. Fic. 4. Both nuclei have escaped from the spore and lie just outside of the germ pore. Fic. 5. Two nuclei lie in the expanded portion of the germ tube. Fic. 6. Four nuclei are plainly visible in the ‘“‘pouch.’”’ The germ tube has not begun the second stage of its growth. Fic. 7. Tip end of fully developed germ tube showing four nuclei in a row. Fics. 8 and g. Other fully developed germ tubes showing a more pronounced type of branching. Fic. 10. The germ tube has made about the maximum growth of which it is capable under artificial conditions, a cross wall cuts off the main portion of the granular cytoplasm at the forward end of the germ tube. VoLumeE |, PLATE I. BROOKLYN BOTANIC GARDEN MEMOIRS. DopGE: GYMNOSPORANGIUM TRANSFORMANS INTERCROSSES BETWEEN SELF-STERILE PLANTS E. Ma EAST Bussey Institution, Harvard University The fact that self-fertilization is practically impossible in certain hermaphroditic plants, although both the pollen and the ovules are functional in crosses, has been known since the time of K6lreuter (1760-1765). The oddity of the phenomenon has been a lure for al- most every hybridist from that time forward. As in the case of most other genetical problems, however, our knowledge of its cause and meaning remained in status quo from the time of Darwin until Men- delian days. Indeed when the writer began his investigations on the subject in 1910, the only considerable post-Darwinian work had been done by a zoologist (Morgan, 1904) on the self-sterile ascidian, Ciona intestinalis. Since 1910 botanical papers have appeared by Correns (1912), Compton (1913) and Stout (1916), but these investigations will not be discussed here, as it is proposed to treat in this paper only certain phases of the work carried on by the author and his associates! during the past seven years, leaving critical review for another place. For our purpose it seems essential only to present a hasty sketch of the subject as left by Darwin. In addition to the utilization of most of the previous and the con- temporaneous work, Darwin (1876) carried out several investigations of his own on the five self-sterile species, Eschscholtzia californica, Abutilon darwinti, Senecio cruentus, Reseda odorata and Reseda lutea. Darwin’s first important result was that the expression of self- sterility in Eschscholtzia californica and Abutilon darwinti was influ- enced by changes in external conditions. Six generations of Esch- scholtzia californica had been found to be completely sterile in southern Brazil by Fritz Miiller (1868, 1873). As English plants were self- fertile, Darwin obtained from Miiller seed of Brazilian plants of known self-sterility. The plants which they produced in England, while not wholly self-fertile, tended toward self-fertility, which fact Darwin attributed to the lower English temperature. A second generation of seedlings proved to be still more self-fertile. Conversely, seed of English stock was somewhat self-sterile the first season and one plant ' The author desires to make grateful acknowledgment to Dr. O. E. White and Dr. J. B. Park for their painstaking aid in this work. Without it, the numerous experiments undertaken could not have been completed. 141 142 BROOKLYN BOTANIC GARDEN MEMOIRS wholly self-sterile the second season, when grown in Brazil. One may assume, I think, arguing from data of similar character, that this progressive result was not due to actual inheritance of an acquired character but rather to the fact that the first generation in each case passed a portion of its life cycle in the original environment. Similar results were obtained in the case of Abutilon darwinit, which though self-sterile in its native Brazil, became moderately self- fertile late in the first flowering season in Darwin’s greenhouse. Darwin made more detailed experiments on Senecio cruentus, Reseda odorata and Reseda lutea and found, as he believed, that each plant though self-sterile was cross-fertile with every other plant. His pollination experiments with Senecio cruentus and Reseda lutea were so inadequate that they may be omitted from consideration; it was really his experiments on Reseda odorata that were thought te establish the fact of complete cross-fertility. DARWIN’S EXPERIMENTS ON Reseda odorata IN 1868 Male Parents Ae panG pn pe) ee Aa cS eae. | BuleealeS: Alcala | g 0 | — — — — _— gC ie aS. Ee Es | 3 eo | ee | en, ee OD Nase |S fete || Sea 30 St JB TR. WS 1 eS | oO jew | | F | S45 G | Ss: Only sixteen cross matings were made, however, and this is not sufficient to prove the point, as is shown by one of our own experi- ments, where I13I cross-matings were made with only 4 cases of cross- sterility. From the fertile cross-pollinations Darwin raised four plants in 1869. Three of these proved to be self-fertile and one self- sterile. Six more plants were grown in 1870. Of these, two were almost self-sterile and four were almost completely self-fertile. The former produced altogether five seeds from self-pollinations, and the resulting plants proved to be self-sterile like their parents. These varied results Darwin attributed to a difference in inherited sexual constitution, but it seems to me that this conclusion should be ques- tioned. Our own results have proved conclusively that toward the EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 143 very last of the flowering season? self-sterile plants may sometimes become somewhat self-fertile. Darwin’s (1876, p. 346) general conclusions are as follows: ‘Finally, the most interesting point in regard to self-sterile plants is the evidence which they afford of the advantage, or rather the necessity, of some degree or kind of differentiation in the sexual elements, in order that they should unite and give birth to a new being. It was ascertained that the five plants of Reseda odorata which were selected by chance could be perfectly fertilised by pollen taken from any one of them, but not by their own pollen; and a few additional trials were made with some other individuals, which I have not thought worth recording. So again, Hildebrand and Fritz Miiller frequently speak of self-sterile plants being fertile with the pollen of any other individual; and if there had been any exception to the rule, these could hardly have escaped their observation and my own. We may therefore confidently assert that a self-sterile plant can be fertilised by the pollen of any one out of a thousand or ten thousand individuals of the same species, but not by its own. Now it is obvi- ously impossible that the sexual organs and elements of every indi- vidual can have been specialised with respect to every other indi- vidual. But there is no difficulty in believing that the sexual elements of each differ slightly in the same diversified manner as do their external characters; and it has often been remarked that no two individuals are absolutely alike. Therefore we can hardly avoid the conclusion that differences of an analogous and indefinite nature in the reproductive system are sufficient to excite the mutual action of the sexual elements, and that unless there be such differentiation fertility fails.” One cannot but admire these inductions Darwin has so cleverly drawn from such meager data, nevertheless one cannot accept them today just as they stand. The reasons for this statement will be seen more clearly when our own data have been presented, but a brief can be submitted with only the support of the work known to Darwin. In the first place, the seemingly contradictory results that were obtained in the experiments on Reseda odorata are not necessarily con- fusing. As reported, self-sterile plants produced varying ratios of self-sterile and self-fertile plants. Unfortunately, the progeny of the self-fertile plants was not followed. If it has been, the problem might have been more easily solved, for, in all probability, the daughter plants would have been self-sterile. It is my own belief, however, that the answer can be read in the casual remarks dropped by Darwin in the midst of his careful descriptions, remarks to which he paid little attention. Darwin found that both Eschscholtzia california and Abutilon darwinti, though self-sterile in Brazil tended to become self- fertile in England,—especially late in the flowering season. Now 2 Cf. Darwin’s observation on Abutilon darwinu. 11 144 BROOKLYN BOTANIC GARDEN MEMOIRS these facts together with that mentioned above regarding the in- constancy of the results obtained from planting the seed of self-sterile plants, may be interpreted by the assumption that he was dealing entirely with fluctuations in all of the five species investigated. These species genetically were wholly self-sterile. The tendency toward self-fertility was due to conditions. In other words, these plants genetically self-sterile needed conditions conducive to a fine healthy growth to bring out their self-sterility. In the lower temperature of England, at a time of decline (the last of the flowering season), they became phenotypically somewhat self-fertile. In the light of my own experiences, I believe we can reconstruct a picture of Darwin’s experi- ments on Reseda odorata with considerable confidence. He isolated the plants that he desired to test under nets; then came pressure of other work, and the data were not collected until the plants had ceased flowering. At that time capsules were found beneath the nets, and this seemed to prove at least a partial self-fertility. But instead of this procedure, suppose that successive self-pollinations had been made throughout the season. The presumption is that the plants would have been declared to be self-sterile with the same remark added which he jotted down in the case of Abutilon darwint, viz., they ‘‘became moderately self-fertile late in their flowering season.” Again, Darwin found no cross-sterility in the plants tested, and concluded that a self-sterile plant can be fertilized with the pollen of any one of a thousand or ten thousand individuals of the same spe- cies. Such a conclusion was less cautious than was Darwin’s wont for it was made from a total personal experience of some twenty-odd cross- matings only, unless his records are extremely incomplete. Indeed this conclusion must have been somewhat of a surprise to himself since he states that ‘“‘it is obvious impossible that the sexual organs and elements of every individual can have been specialized with respect to every other individual.’’ He surmounted this difficulty by assum- ing that the sexual elements of each plant differ slightly in the same manner as their external characteristics, and that this slight difference is sufficient to excite the mutual action of the sex elements necessary in order to have fertilization ensue. The kernel in this conclusion, that differences in the reproductive systems of two self-sterile plants are necessary in order to promote cross-fertilization, is so similar to that to which the writer has been forced after seven years of rather intensive work as to be uncanny, for it seems to have been reached in spite of rather than because of the data at hand. This feeling of surprise at Darwin’s clairvoyancy may seem affected, since he was usually in advance of his time, but it is a fact perhaps worth men- tioning as a confession of omission that the writer reached his con- EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 145 clusions as the outgrowth of work on heterozygosis and did not refer to Darwin’s view until recently. Be this as it may, a short com- parison of Darwin’s main induction with the facts from which it came will, I think, show a real reason for wonderment. He believed in universal cross-fertility of self-sterile plants, his basis being the small number of cross-fertilizations made by Hildebrand, Miiller and him- self; although Robertson Munro (1868), with whose work he was familiar, had found cross-sterility in Passiflora alata, and even the works of Hildebrand and Miiller as published leave the matter in doubt. Now how much more reasonable the general induction mentioned above seems if one assumes (1) that self-sterile plants breed true for self-sterility but may show a slight degree of self-fertility as a fluctuation under certain conditions, (2) that a variable but limited number of germinal “‘factors’’ influence the success of matings, cross- fertilization being possible only when two plants differ in these effective factors, and (3) that when two plants have the same effective factorial composition, cross-sterility of the same type as self-sterility exists. This is what we believe our own work has shown, as we shall try to demonstrate. Emphasis must first be laid upon the fact that the behavior of self-sterile plants among themselves and the relation between self- fertile and self-sterile plants are distinct problems. Compton (1913) found the relation between self-fertile and self-sterile plants of Reseda odorata to be that of a simple Mendelian monohybrid with self-fertility dominant. The same relation appears to hold in crosses between the self-fertile species Nicotiana langsdorffii and the two self-sterile species with which our work has been done, Nicotiana forgetiana and Nicotiana alata. There is some single differential between self-fertility and self- sterility. Given the proper composition a plant breeds true for self- sterility. The behavior of self-sterile plants among themselves therefore must be considered separately. Our work, as stated before, has been done with the two self-sterile species, Nicotiana forgetiana and Nicotiana alata, and largely with crosses between these species. Both of these species are affected in their manifestation of self-sterility by certain environmental changes, Nicotiana alata much more than Nicotiana forgetiana. Self-sterility is determined by the inheritance received, but it can develop fully only under environmental conditions which promote a normal healthy growth, and during the period of intense flowering. Toward the end of the flowering period, especially under conditions adverse to vegetative growth, self-sterility sometimes shows a marked and rather sudden decline. A few seeds, or even a well-developed seed capsule may then be obtained. This is not a common occurrence; indeed, it 146 BROOKLYN BOTANIC GARDEN MEMOIRS is rare, but it is a possibility. Three cases of seed production out of over three hundred plants tested have been observed in Nicotiana forgetiana. A considerably higher percentage of fertility has been marked in Nicotiana alata. Self-sterility can be restored in such plants, however, if they are allowed to go through a period of rest and are then, by proper treatment, brought into vigorous flower again. This is not the whole evidence that this occasional end-season fertility is a pseudo-fertility brought about by external conditions— a fluctuation. Three generations of Nicotiana alata plants have been grown from selfed seed produced by end-season fertility without the occurrence of a single plant which behaved in every way like a truly self-fertile individual. This phenomenon, therefore, while teaching us to test self-sterility only during the main part of the flowering season, has shown that there is no reason why fusion between gametes produced by a self-sterile plant may not occur provided the male generative nucleus enters the embryo sac. Such unions may take place without affecting the self-sterility of the progeny. What is then the difference in behavior that makes a cross-poilina- tion effect fertilization while a self-pollination produces nothing? What occurs is this: After a self-pollination the pollen grains germinate and the tubes pass down the style at such a slow even rate that they reach only about half way to the ovary before the flower wilts and falls off; while the pollen tubes after a cross-pollination, though starting at the same rate as the others, grow faster and faster until fertilization is effected in four days or less. The curve of distance traversed plotted against time is in the case of the self-pollination nearly a straight line, while in the case of the cross-pollination it simulates that of an autocatalytic reaction. From these facts it seems reasonable to suppose that the secre- tions in the style offer a stimulus to pollen tubes from other plants rather than an impediment to the development of tubes from pollen of the same plant. And we believe that this stimulus is in some way caused by certain effective differences in the factorial composition characterizing two compatible plants and that if two plants do not have these effective differences in factorial composition they are by the same token cross-sterile with each other. It is clear that this assumption presumes that the pollen grains matured by a given plant behave as if they are sporophytic as regards that part of their con- stitution that affects self-sterility and cross-sterility. The pollen grains of any plant may carry many different hereditary factors, they may even carry several different factors which function in controlling the success or failure of particular cross-matings in the next generation, but in their own action on the stigmas of other plants they behave EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 147 as if each carried the composition of the mother plant from which it came. In other words, as far as its action in fertilization is concerned, a pollen grain partakes of the character of its mother plant and is like its sisters; as far as the hereditary characters carried on to the next generation are concerned, sister pollen grains may differ both from their mother and from each other. A part of our evidence on these points we shall present. For further details the reader isreferred to a forthcoming paper in Genetics.’ The first experiment to which attention is called is an inbreeding experiment performed on a cross between Nicotiana forgetiana and Nicotiana alata. If sister plants are mated in successive generations after an original mating Aa & Aa, by Mendelian recombination there results a gradual approach to 1/2 AA, 1/2aa ando Aa. Expectation of homozygosis in successive matings is 1/2, 5/8, 11/16, 24/32 --- I (Jennings, 1916). If, therefore, plants of like constitution as far as effective factors are concerned are cross-sterile with each other, cross-sterility should become more and more apparent in generations succeeding F.. To test this possibility, a comparatively small number of cross-matings was made on the Fy», F3, Fy and F; generations. In the F, generation, out of 131 intercrosses on 20 plants only 4 were unsuccessful. The percentage of unsuccessful matings increased from this time on, until in the F; generation about 21 percent of the cross- matings tried on 20 plants were impossible to make. In this experiment as well as in all others, results showed that reciprocal crosses were alike in their compatibility. If two plants were fertile together, they were fertile reciprocally; if two plants were incompatible, they were incompatible reciprocally. This is proof of the sporophytic behavior of the factors affecting the behavior of self-sterile plants. The two crosses to be described next are reciprocals made with the same two individuals. Made with Nicotiana alata and Nicotiana forgetiana as parents, they are in a sense repetitions of the cross just described, but it is hardly probable that they duplicate it. Both of these species must consist of plants which differ among themselves in the factors which affect self-sterility, hence any crosses in which different individuals are used may show different results. All of the individuals resulting from this cross were grown in a greenhouse as potted plants. The F; generation came into blossom during the latter part of the winter. Conditions were extraordinarily favorable for growth and the pollinations were all made while the plants were vigorous, hence scarcely any trouble arose over classi- fication of the results through end-season pseudo-fertility. ’This paper has since appeared. See ‘‘Studies on Self-sterility I. The Behavior of Self-sterile Plants.’’ Genetics 2: 505-609. 1917. 148 BROOKLYN BOTANIC GARDEN MEMOIRS Our study was made on a population of 53 plants. Pedigree numbers from 0 to 39 inclusive represent the cross NV. alata < N. for- getiana; pedigree numbers 40 to 52 inclusive represent cross JN. for- getiana X N. alata. Each plant was selfed one or more times, and all proved abso- lutely self-sterile. Further each plant was back-crossed with pollen from a single plant of each of the parent species with complete success in every case. The plants used in this case were not the individuals that entered into the cross, however, for unfortunately these were not available. ABE Be RESULT OF MATINGS ON F; PLANTS 0 TO 39 N. alata X N. forgetana and on Plants 41 to 52 N. forgetsana X N. alata Ped. No. Fertile with Ped. No. Sterile withPed. No. Ob n aes AA AGRI ci pein othe ae: aera ecun eke Bee eicnees ate ee 22, 34, 38, 49 Tet oe. Bi IR AN Os At sewn eee eis Sec de Rand neon coetheady ees 8 DP ave is AS US PAT RAAR (5 2 lucwrrce aes ciel hstneusvnders Bay sue yraaeich ave 072223 Bide teeter ZO MAC DOOD OIG fia tendaie a. crave tyes a Skene Care 4, 6, 18, 41, 46 Bieta ts DIO PLO AMA ate c tee ware PRR oras ota tene ie ate shaeesied chon 18 Ce ae. eae 2EBIGs ON LOVMS WAG in am ae cece State cmereeera us 8, 44 ORFs se LOS AS YA Ae Pere eed oN pom csketenate go eu edaee eee 3, 4, 18, 40 Leaner DMS OD GAA eee re rich ones eet OS os Der eas eens re 18, 46 to) tise Bratt OHO EO NSO AOs A Om se mie sence tet oy oer he 5, 44 Oe. Be Osea aS Qahota ie Mh ex ate Atanas Charset ie epee ey 2 LO W285 27S TOMATO ASO SESMAO SAA cic acu-t: eremeeietsusascrce Meas is ski seks =k 2 23, 24, 27,134 040 1G Giessen re Pee 2S 2 LO AAA AO 1) a eee OW TOR 22 AGE fe sr chepeio pera tra isi ie Oats egy 2 6, 18, 46, 52 1g ts Oe BIRO SES AALWA ORM rr iNe iene canta oeaaiees, Sevencnete sneysneet: AO), UGS Ay eu DA tn ears INS PRCLOPTRCNe tatc tena ty ol Dt Deen SIE ED Ghee omelet enor oc 10, 34 TIS Miegtrstate Li), UO Ape iS a2 Oye sar on Sesat certs adele atte otis 9,,13,. 14; 23; 44 TOS si Ie eta Rojeay eh Oe lok is AE OP Ae oe eld ao o 729 ope cae LAs TOTO S20 422552 Ores eee eessi wie RL ee 16, 26, 44 NO meee: APO prrits Ae Der aula olen AWLS cone aaceasomaupoasc 3, 46 LO Perc 1 by eae Paeg Po VyeV Wee Vitis Oo Ea OR PG OD! BIOS AD iO Gi os 18 2O one ee Pri OF, iO}, Is}, Ail, A, Ay, 2105 MO, AY Bech oo ono ee 43 eM Wh ah ate Ae el2 TOES A Gira. Bae ees AER te ecco cs 2.10; 22925, 27a PPARs Ee T2P AD PAAR ey ae bares yee eS ys We CME Pe pares ee 14, 23, 24, 36, 48 7c RE ee Bile hopes ciae! oe ophtentencgs Oe esa ee cere ga Ie Ok ours 9, 10, 37, 48 2) eee BOP 205 2ZOW2O NAAM SS ae ricme ate rar ees TO) 22, 23 9gOnai Dre wane Seis Bird AAO OL ders eco eRe Rat ants 2) O} 23,27, DOs cu ORES 55225232 54 OM Siasieaet ash ieeenn ee eee 28, 29, 44 DT RE RCAAE BAD S432 AA WA Oleh sper c nis ae ean BIS cee te er ap 2G) 30) 24949 ZA HORE eh D852 LOT sa Ohad Obs muebetets om yer ee we deine cco hk 8,26, 2044 710 ie ean Pip lily heh Ae Aah Ail IS Clo). eyo Shy) ably vl a Soa 5, 20, 28, 30, 44 2 Ober S59 205,33 AAS AS MAO OR ae tebe od ae ares Oy 2 2227, oso eee 22 BD iD sce thd ROE aA ist SNe Mg EPO hats saa os 8, 29, 36, 44 BO Nei crs Oi 27 1231205.3 Oma Ane Sand Ain ened ny ee arses eer 18, 33, 46 gaee Bh S: LGW2a 2 TAG ec men es ave ator taper. Jobat, fe, | 18, 32 Bary Se ce. PR Pay Mee V:Ingy Leena, at ben Pres ct Aue a MNi ae ot Ec Oe NON23y desi BG rosters 3°19, SLO 2D s OMA nS TAs Rete me eee 8 ZOOM oh ise fo aie Ye Se I Kop eI noe A int a ie (eae 10, 23 Bye ier 30 ;:42'°43, AAs A Gieiacten aoe eae ees 0; 109722, 2353430 EAST:. INTERCROSSES BETWEEN SELF-STERILE PLANTS 149 Fol oh OtaaO DOs ONFAD WAZ IAG ae -s) is sien chepencontsia a) 6 bi a) stein 2A 377: BIS NoaTIOEAE Oy Aiko ey StB ORS cin b oie Bebe. a det cael s.edausienae enensy News gal ee 18, 40, 42 Ore tere oe: ZA EAA GEA Her © Emtec tonsioct cske seal cei ercener serene 6, 33, 46 70 Se er ie OWA AA eA OMEN as eh Sar avele acer eee eee ee 33, 40, 46 AD Nees 2O AA ae ani ery Rah ag mec O re Oct PEI 39, 41, 45 ABS ae: 5, 27, 33, 38, 39, 40, 42, 44, 46, 51 AAC sins 10, 14, 23, 34, 45 ABP tes OMG ACE Oteeees Me ata n ie a a) Ae Hs ehsicr cnc) ots ees 46, 52 AOR to NOM22 AT AAR ES Te Ret y koe eisks cia the hehe Sie sora She ee 52 A hoe, ct 2 OAD AA EA Se Onno 527 estas cis Sit ieseisbnerneiy oe 38 BSi So shs. 3 AO MA WAGGA Ob ata, Went as ttiae a aeieas ov Plcleueps wteis ace ena Sees LO; 23) 245.27, 34 AQm ys er AQAA WAS Wenner hitrts, catnccnass © oxsat ot & ake sacle OF9) 275345 47 SOs, sais OAS Os aeRO a cae eis Shoe ee, Melted cicy «SoS wthaud:S-Soe tn ote ON 27. 27 Cy late eee One 23503 O PAG A Oy 15 Ole rare carn hel ceversietes cmleyes 8, 29 52 eis tacks LOCO 20 wae G Lamon mele cieices 6h duro 2:4 Breis Gee ea 354, O18, 41, 45; 46 The numerous cross-matings made are shown in Table 1. There were 103 reciprocal matings. Of these 100 gave duplicate results, 39 pairs being fertile and 61 sterile. The three which did not check are: 2 X 3, sterile, 1 pollination 3 X 2, fertile, 1 pollination x 52, fertile, 1 pollination x 6, sterile, I pollination X 21, fertile, 1 pollination X 37, sterile, 1 pollination classed as fertile, classed as sterile, classed as sterile. Since but one pollination was made in each of these cases we have made our decision as to fertility or sterility by a consideration of the circumstantial evidence. The behavior of these plants in other crosses shows conclusively that 3 should be fertile with 2, 6 sterile with 52, and 21 sterile with 37. They have been classed accordingly. That this grouping is correct is further shown by the fact that the mating 3 X 2 (classed fertile) was made at the height of the flowering season, while the matings 6 X 52 and 37 X 21 (classed sterile) were re- spectively the last and next to the last matings made on those plants. In spite of the fact that plants 0-39 are from cross N. alata & N. forgetiana, and plants 40-52 are from cross N. forgetiana * N. alata, they behave as one family in intercrosses. The entire population can be grouped into 6 classes in which there is interclass fertility and intraclass sterility. The following explanation may be necessary to make it clear just how Table II was obtained from Table I. Table I shows all of the matings, but in the form given it is not easy to see at a glance every combination in which a particular plant was used, both as male and as female. It was necessary, therefore, to make a new table, in which the pedigree numbers in the column at the left were tabled as males, and the pedigree numbers in the columns headed “Fertile matings’ and “Sterile matings’’ were tabled as females. 150 BROOKLYN BOTANIC GARDEN - MEMOIRS Thus plant 2, used as a female, was fertile with pollen from plants 4, 18, 41, 44 and 52, and sterile with plants 9, 22 and 23; but pollen from plant 2 was fertile on plants 1, 3, 4, 5, 7, II, 18, 20, 28 and 29, and sterile on plants 9, 10, 13, 25 and 27. It is clear, therefore, that instead of the 8 matings on plant 2 that Table I appears to show, there are really 21, the 3 reciprocals of course being counted but once. These tables were combined for analysis. In the interest of economy of space only one is shown, however, since the second can easily be made from the first. The four exceptions in this huge set of matings are in reality negligible. Matings 15 X 44 and 31 X 36 were sterile, though they do not belong to the same class. Plant 15 was sterile to 4 plants of Class A and fertile to 2 plants of Class B, 3 plants of Class C, and to the isolated individuals forming classes D and F. It is unquestionably a member of Class A. Plant 44 was sterile to 7 individuals in Class C and fertile to 17 plants of Class A, 12 plants of Class B and to the singletons forming classes D, E and F. This evidence places it un- mistakably as a member of Class C. Plant 31 is also a member of Class C as evidenced by 3 sterile matings within that class and by fertile matings with t plant of Class A and 3 plants of Class B. Plant 36 is like plant 15 thrown into Class A by its sterility with 3 others of that class, and by its fertility with 3 individuals of Class B, with 2 of Class C, and with the lone plant of Class D. In view of this evidence and the fact that in these two matings but one pollination was made in each case, they are much more likely to be errors of record or of technique than true exceptions to our classification. The other two exceptions, matings 45 X 18 and 33 X 46, were fertile where from the evidence of numerous other matings they should have been sterile. Here again but one pollination was made in each case; and, coincidence though it may be, each pollination was the last mating made on that particular plant. What is more probable than that this is a pseudo-fertility appearing during the wane of the flower- ing season of the two mother plants, No. 45 and No. 33? Six groups appear in Table II, but there is proof of the existence of only five. Groups A, B, C, D and F are definitely established. Plant 11, on the other hand, is an isolated individual rather than a class. It does not belong to groups A, B or C; but unfortunately it was not crossed either with Class D (plant 20) or with Class E (plant 43), hence one cannot say that it does not fall into one or the other of these two classes. In the three large groups the distribution of individuals is 22, 16 and 12. About all that can be said about the type of this distribution is that the classes are not of equal size. On the other hand, it is EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 151 interesting to note that the plants of both cross No. 2 and cross No. 3 fell into the three groups as if they were samples of the same popula- tion. There were 40 plants of Cross No. I, and 13 plants of the TABLE: It PLANTS OF F; GENERATION OF RECIPROCAL CROSS BETWEEN JN. forgetiana AND N. alata, GROUPED IN ACCORDANCE WITH THEIR BEHAVIOR IN INTERCROSSES Plants 0-39 are products of the cross; plants 40-52 are products of its reciprocal . . Cases Fertile in Group Cases Sterile in Group Group Ped. No. | Boe | ao ee Nom ms ms B GE D E F lew | | | HH [MORO OOne Ono, CHOP OrORS | | = ee | | | | CODCDDODCODOOOODOOOCOODODOOOWMNINHPOWHTNMNOUNHO OPBUNOW OF [ROR IOBRORO Orel | | | el | _ | ° | | | 4 4 ios) rs SONOS ONO: O07 oO | | | | [tah tek | | | ber [ere | = [0 .0) al _ COC MOWWAMWBWAWHHBNHABRADGCOCDODCODCIDCODCDCDCACCDCOOGCGOGCGC8G80 80 1a | Se | ee ee |] | | xe) 1) {ly ee} | Iss (on _ I | | |W} te) + | MEPBROHHOODOOOHOOH OOOO OWWAUMUWUNWUBENNAAWUNNANN AH COOCKWNNNWWRWWWO KHER NMNHHHNNWNHP MOUS UUMNWHWW DAU! | os) oS) DwWOnNNHANAUFP HP ANNWBWNHF WOOO 0 ° | o ° Wich OnOmOMOnOnore) O10) C11O © C10 0 OO OO OO Or OO OOOO O oO HF OOO Oo O96 | 152 BROOKLYN BOTANIC GARDEN MEMOIRS TABLE I1—Continued Cases Fertile in Group Cases Sterile in Group Group | Ped. No. | | BA Cal | ee eal ie on aie ee D| Baie | | iN Cee: 16 5 | © I I - | oO One fo) (e) - 1G) Ata\t (2 eO I - | - O Onis 0 |) | 26 G20 r|/-/]- o |) S07 )4 > 0, See 28 Oleh sO oe a oO 0} 4) | = | = 2 9 6 | oO = = = O Or 4 = = = 31 I By Oro ee I O-le3 -/|-|]- ee Me eee me eee Os coelp se ees = | = 44 Ty | 125| 90 I I I I @:| 7 || 0: | ouimias Vio 3 Sigma ae amet Coa I - O One -}|o]- De epee ZOOM O UE sOr ly orcleas oO I - oO ONE Co) 0; | ‘Oral ne 43 | SS ae (er. 0") so. Eos! eo.4|\ om cms | Ora ee II CMI f Reel - = (0) oO o| Oo - | - O reciprocal, Cross No. 2. In the classes A, B and C the proportions were 18, 10, 10 and 4, 6, 2 respectively. This similar behavior of the progeny of reciprocals seems to us strong corroboratory evidence in favor of the conclusion that reciprocal crosses always behave in like manner as regards self-sterility. The study on this family is but one of several that have been made. but we believe that the data on it alone show unmistakably that the behavior of self-sterile plants in intercrosses is governed by a relatively small number of factors which act through pollen as if the pollen grain possessed the characters of the sporophyte from which it came, and that the gametes of plants having like constitutions as re- gards effective factors are incompatible in the sense that they do not make a normal pollen-tube growth and hence do not reach the ovary in time for fusion to occur. This interpretation shows both why plants are self-sterile and cross-sterile. It accords completely with the fact that a population of plants may be divided into groups on the basis of their mating proclivities and that each member of any group is cross-sterile with every other individual of that group although it is fertile with every individual of every other group. These assumptions being true, it ought to be possible by con- tinuous self-fertilization, utilizing end-season pseudo-fertility, to obtain ultimately a population in which every individual possesses the same effective self-sterility factors. In such a population all of the plants will not only be self-sterile, but will be cross-sterile. Sucha population has been obtained. EAST: INTERCROSSES BETWEEN SELF-STERILE PLANTS 153 REFERENCES CITED Compton, R. H. Phenomena and Problems of Self-sterility. New Phytologist 12: 197-206. 1913. Correns, C. Selbststerilitat und Individualstoffe. Festschr. d. mat.-nat. Gesell. zur 84. Versamml. deutsch. Naturforscher u. Arzte Miinster i.W. pp. 1-32. I9I2. Darwin, Chas. Effects of Cross- and Self-fertilisation in the Vegetable Kingdom. Ed. 2, 1878. N.Y. D.Appleton. 1876. Hildebrand, F. Bastardierungs Versuche an Orchideen. Bot. Ztg. 23: 245-249. 1865. — Ueber die Nothwendigkeit der Insektenhilfe bei der Befruchtung von Cory- dalis cava. Jahrb. wiss. Bot. 5: 359-363. 1866. — Ueber die Bestaubungsvorrichtungen bei den Fumariaceen. Jahrb. wiss. Bot. 7: 423. 1869. Jennings, H.S. The Numerical Results of Diverse Systems of Breeding. Genetics I: 53-89. 1916. Kélreuter, J. G. Vorlaufige Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen, nebst Fortsetzungen I, 2 u. 3. Pp. 1-266. Ostwald’s Klassiker, Nr. 41. Leipzig: Engelmann. 1761-6. Morgan, T. H. Some Further Experiments on Self-fertilization in Ciona. Biol. Bull. 8. 313-330. 1904. Miiller, Fritz. Notizen iiber die Geschlechtsverhaltnisse brasilianischer Pflanzen. Bot. Ztg. 26: 113-116. 1868. — Bestdiubungsversuche an Abutilon-Arten. Jen. Ztschr. f. Naturwiss. 7: 22-45, 441-450. 1873. Munro, Robertson. On the Reproduction and Cross-fertilization of Passifloras. Bot. Soc. Edin. 9: 399-402. 1868. Stout, A. B. Self- and Cross-pollinations in Cichortum intybus with Reference to Sterility. Mem. N. Y. Bot. Gard. 6: 333-454. 1916. BINARY FISSION AND SURFACE TENSION IN THE DEVELOPMENT OF THE COLONY IN VOLVOX Ro A. “HARPER Columbia University In Volvox we have an incipient metaphyte with a many-celled body of definitely organized form and the fundamental differentiation of soma and germ cells fully established. Klein’s (’89, ’90) argument that Volvox differs from true metaphytes in that the differentiation of the germ cells does not take place until after cell division is com- pleted is not very illuminating. The germ cells do not appear in Oedogonium until a considerable series of undifferentiated cells have been formed. Falkenberg’s comparison of the entire colony to a zoosporangium also overlooks the very important fact that in the development of the Volvox colony growth regularly alternates with cell division (at least after the first few divisions) just as it does in the development of the soma of one of the higher metaphytes. Vol- vox is frequently referred to as one of the best known algae. There is general agreement as to the order of cell divisions in the formation of the colonies, both for the egg and the asexual germ cells. The litera- ture has been frequently summarized. To proceed further with the study of Volvox from the standpoint of evolution and morphogenesis, we may compare it with such a simple coenobe as Gonium. In addition to their more obvious struc- tural characteristics there are two essential differences between the colonies in such forms as Gonium and Volvox. First, the adhesion between the daughter cells is much more firm in the latter. In Gonium, as all observers testify, the adult colonies break up into their component cells with the greatest readiness. I have figured such broken-up colonies (’12, Pl. V, Fig. 23). Slight pressure, change in the chemical composition of the medium in which they are, etc., lead to almost explosive separations. One marked difficulty in getting good photographs of the colonies is due to their tendency to go to pieces. On the other hand, Volvox shows almost no tendency to separate into its component cells. The colonies may be crushed into formless masses without isolating a single cell and I know of no chemi- cal or other stimulus which will cause their cells to fly apart as do those of Gonium. 154 On Sat HARPER: BINARY FISSION AND SURFACE TENSION 1 The second difference lies in the very fundamental fact that, as noted, in Volvox the germ cell grows to relatively large size before dividing and the daughter cells grow in size between the successive cell divisions. This is a very long step toward the full metaphytic habit in ontogeny. It marks a return to the habit of the simple protophyte like the bacteria and the appearance of a new point of departure in the development of the morphogenesis of a metaphytic plant body out of the primitive habit of reproduction by swarm- spores which is seen in Chlamydomonas and Sphaerella. In these protophytes, the cell having reached maturity forms from four to eight swarmspores by rather rapidly succeeding divisions of the mother cell. They escape by breaking of the mother cell wall and then as free individuals proceed to grow to the size of the parent. Swarmspore formation in Chlamydomonas and Sphaerella is a step beyond the conditions in Euglena, for example, where in ordinary reproduction each cell division is followed at once by the individualiza- tion of the daughter cells and their independence as separate organisms. In these particulars we may distinguish three steps in the evolution of the metaphyte from the typical protophyte. r. Cell division, in simple, direct alternation with growth, reproduction and individualization, practically simultaneous and _ identical processes. Euglena. 2. Cell divisions at unequal intervals, reproduction multiple and in alternation with growth. Individualization delayed. Chlamy- domonas, Sphaerella. 3. Cell division and growth in direct alternation, reproduction multiple and individualization delayed by intercalation of a true em- bryonic period. Volvox. In Volvox individualization is already in essence the complex process of differentiation and maturing which we find in the highest plants and animals. In Gonium individualization of the daughter colony, as I have shown in a former paper (’12), is accompanied merely by certain gliding movements of the cells upon each other by which an approximation to a least surface configuration is achieved so far as is possible for sixteen ovoid cells arranged in a flat plate. In Volvox, with the retention of multiple or colony reproduction as in Sphaerella, we have growth intercalated again between each successive cell division and also a specialization in function between germ and somatic cells. Complete individualization is delayed till the colony has become very many-celled. What may be called a pseudo-growth comparable to the elongation of the cells just back of the root tip by absorption of water and the formation of large central vacuoles is also represented in Volvox by the formation from the cell 156 BROOKLYN BOTANIC GARDEN MEMOIRS walls, as Meyer has most fully described, of the large masses of slime by which the protoplasts in the adult colony are surrounded. This slime, about whose nature Cohn, Klebs, Blochmann, and others have differed so widely, is, as is now generally recognized, the gelatinized cell wall comparable to the secondary thickenings in collenchymatous tissues through which extend the broad strands which provide for the again much disputed intercellular protoplasmic connections which are so conspicuous in the adult colonies. The firm adhesion of the daughter cells to each other and the re- establishment of the primitive cell division-growth rhythm are two further conditions to be reckoned with in the development of the colony of Volvox as compared with that of Goniwm. We may turn now to the morphogenetic processes involved in the reproduction of Volvox. Braun (’75) first clearly recognized the “division by the wheel-forming type”’ in Eudorina as distinguished from the ordinary successive bipartitions at right angles in Palmella, etc. Braun refers the readjustments of rounding up and rearrange- ment of the cells in forming the globular colony to the pressure of the developing slime envelopes. It has not been sufficiently emphasized that in the two-cell stage and in the four-cell stage, as in the cleavage of the animal egg, the halves and the quadrants respectively will tend by surface tension to round up and give us a plate-shaped instead of a globular mass. In Eudorina Goebel shows that the four cells tend to round up and are shortened. He further represents them as tending to remain at one end of the mother-cell cavity and to adhere to the surface of the mother-cell wall. This leads to a divergence of their major axes and (82, p. 36, Fig. 17) gives already at this stage a polar opening. Over- ton (’89, Taf. II, Fig. 10, a, 6, c) figures evidence of this divergence of the four cells and it has been observed by others. The third division by the wheel type, or radial type, gives us with the rounding up of the cells a disk consisting of four interior and four peripheral cells alternating with them, the familiar cross figure. The four inner cells may appear much larger and are commonly so figured. Overton (’89, Taf. II, Fig. 12, a and b) has shown very clearly that the apparent relative size of the central and peripheral cells varies with the level at which they are observed. The peripheral cells have slipped out of the plane of the central four so that the group of eight is already markedly concave. Two factors are pri- marily concerned in this displacement. First, the fact that we have by binary fission in two planes at right angles a group of eight rounded bodies which can form no stable least surface configuration in one plane and, second, the disk-shaped group formed by the rounding HARPER: BINARY FISSION AND SURFACE TENSION 157 up of the eight cells by surface tension tends to conform to the shape of the cavity of the mother cell. The whole is an expression of the incompatibility of the principles of surface tension and binary fission complicated still further by the rather firm adhesion of the cells to each other. If division had produced seven instead of eight cells and if they were free to adjust their interrelations in accord with their capacity to achieve a position in which their pressure relations were as nearly as possible mutually compensatory, we might have the typical least surface group of one surrounded by six in one plane. If ten cells were produced by division and if, as in Hydrodictyon, the rounded form of the mother cell were a dominating factor we might get one cell surrounded by five in the form of a saucer conforming to the curved surface of the mother cell. A further series of five added on the margin of the saucer and the figure could be closed by the re- maining cell. If the eight cells produced by binary fission were free as in Pediastrum, we might get a group like the typical eight-celled colony of P. Boryanum with two inversely bilaterally symmetrically placed central cells and two groups of three peripheral cells also inversely bilaterally symmetrically placed with reference to each other (’16). Pressure of the mother cell might make the group slightly concave. With eight cells produced by bipartition from four strongly ad- herent mother cells and themselves rather firmly adherent the familiar concave cross figure is the best approximation to a least surface configuration. I have noted that in Volvox there is growth of the daughter cells intercalated between the divisions. This is very slight at first. In the early stages, as has been generally noted, the mass of the young colony seems little larger than that of the mother cell. In the preparda- tion for the third division, however, there is a marked elongation of the four cells. In the two-celled stage the halves appear symmetrical, or one cell may be slightly oblique (Fig. 1, Pl. II). In the four-celled stage the sectors at first appear quite symmetrical and uniform in appearance (Figs. 2, 3, Pl. II) but with the preparation for the third division a char- acteristic change in the form of the cells is observed. This growth period intercalated between the cell divisions is an essentially differ- entiating metaphytic character and makes possible in Volvox as in higher types the formation of the indeterminately large and many- celled colony as contrasted with the fewer-celled colonies of Goniwm, Pediastrum, etc., in which the cell-division stages are sharply separated from the cell-growth stages. The growth in the four-celled stage of Volvox is quite specific in that it is not a mere swelling of the cell in 158 BROOKLYN BOTANIC GARDEN MEMOIRS all its dimensions. It results rather in a characteristic elongation in one axis of the cell, the axes of elongation of the four cells tending to be tangential to the general outline of the four-celled group. Klein figures this elongation of the cells very clearly for Eudorina in the four- and again in the eight-celled stage (’88, Taf. VI, Fig. 61, 63). Overton shows it more crudely (89, Taf. II, Fig. 10) and I have been able to photograph it for one of the four cells (Fig. 4, Pl. II). It may take place successively rather than simultaneously in the four cells and apparently proceeds in either direction around the group. Biitschli’s figure (’83, Taf. XIV, 1 g.) shows rather crudely the resulting arrange- ment of the cells just after the third division. The division seems to be nearly simultaneous in all four cells and the wheel-formed group of eight results. This characteristic growth and elongation of the cells at this stage leads naturally to the oft-noted fact that the plane of the third division cuts that of the second obliquely rather than at right angles. A determining factor is, however, obviously the tendency to bisect the elongated cell at right angles to its major axis as well as the direct relation between the second and third cleavage planes. The elongation of the cells during division reminds one at once of the familiar elongation of the egg cell and other free globular cells at the time when the bipolar karyokinetic spindle figure is at its climax of development. We have no good figures of karyokinetic division in Volvox but Overton’s figure from a 200-celled colony (89, Taf. III, 18) shows telophase stages with the cells all elongated and the spindles in every case in the long axes of the cells. It seems obvious that such a cell form in division implies a spindle figure with polar asters and justifies the assumption that the same internal forces are operating in the elongation of the Volvox cell as in the dividing egg. Typical polar asters may be expected to be found at such a stage as that shown in Overton’s figure, like those shown by Swingle (97) for a corresponding stage of division in Sphacelaria. We may conclude then that the adhesion of the four mother cells makes it necessary that the movement of material preparatory to the production of two equivalent rounded daughter cells should take place upon their free surfaces and the result is the characteristic bulging and elongation of the four cells during division. That this change of form is associated with the production of the karyokinetic figure with two polar asters seems clear from the figures of division in other alga cells with centrosomes. In any one of the cells of the four- celled group (Fig. 3, Pl. II), for example, if the third spindle figure has its axis 90° from that of the second division and in the same plane it is obvious that one of the asters will not have space for its full expression and if the adhesion of the quadrants is strong the yielding will be on HARPER: BINARY FISSION AND SURFACE TENSION 159 the free surface of the cell and tend to give us the oblong cell form shown in the figures. That elongation precedes or accompanies divi- sion is indicated by the number of oblong cells in the older colony shown in figure 7, plate II. If the four cells should remain together and flattened upon each other the successive divisions at right angles would give very variously shaped and far from rounded daughter cells, as shown in Biitschli’s diagram (’83, Fig. 1). In effect, as viewed from the pole, the eight cells come to lie in an up-and-down zigzag line instead of forming an in-and-out zigzag, thus enabling each cell to remain as nearly as possible isodiametric without reducing the compactness of the group. It has been generally agreed that each of the eight cells of the wheel figure divide to give a sixteen-celled stage and that the spherical or ellipsoidal form of the colony may be achieved (as it obviously is in sixteen-celled colonies of Eudorina) in this sixteen-celled stage. Goroschankin (’75) held that the 16- and even the 32-celled stage in Exudorina is a plate-shaped disk and that the transformation to a globular form came rather suddenly with the gelatinizing of the cell walls. He has not been followed in this view by later writers, though they have very generally been inclined to accept his account of the order of the cell divisions. Braun (’75), Biitschli (83), Overton (’89), Klein (’90), and others hold that the spherical or rounded form of the colony is practically achieved by the sixteen-celled stage. The fourth division is then a binary fission of all eight of the cells giving the sixteen-celled stage. The radial elongation of the four central cells on their free surfaces is followed naturally according to Hertwig’s law by their transverse division. (See Klein’s figure of Eudorina, ’90, Taf. V, Fig. 63.) The four peripheral cells of the eight-cell stage also elongate before dividing, just as in the four-cell stage and divide transversely, giving the sixteen-celled group. In Gonium the cells glide upon each other so as to form a series of groups of three with the central square opening, as I have pointed out elsewhere (12). The greater adhesion of the four central cells in Volvox and the elongation of the cells before division prevent the Gonium configuration. The same situation develops as in the pre- ceding third bipartition. There are eight new cells formed and these tend to more than fill the space on the margin of the curved disk of eight. Such a group is in very unstable equilibrium. The rounded shape of the mother-cell cavity influences the direction in which the cells glide upon each other, and adhesion tends to develop the com- pact groups of three. Of the eight new marginal daughter cells the four coming from the original group of four form a part of the equa- 12 160 BROOKLYN BOTANIC GARDEN MEMOIRS torial series, while the four coming from the peripheral four cells of the eight-celled stage (Text-fig. 2) form a new polar group of four whose cells alternate with those of the original polar group. Henfrey (’56) described the sixteen-celled colony of Eudorina as consisting of two polar groups of four and an equatorial circle of eight cells. Biitschli (’83) describes the sixteen-celled colony as consisting of four upper central cells, a ring of eight equatorial cells and four lower cells which alternate with the upper and gives a figure (Taf. XLV, Fig. 1, 7). Overton’s (’89) description of the arrangement of the cells in the eight-celled stage is essentially like that of Biitschhy, though as Meyer notes, he ascribes the origin of the central cavity of the colony to the divergence of the cells in the four-celled stage and the subsequent hollowing out of the cells toward the center of the I tN Fics. I AND 2. For explanation see text. colony. Neither of these authors gives any very clear account of the arrangement of the eight equatorial cells or their relationship to the polar groups. It is not easy to obtain satisfactory photographs of these sixteen- celled stages since, as noted, they are practically spherical and the number of cells is so small that no characteristic groups can be ob- tained in one focus. From a study of a large number of cases and photographs of various views I have been able to obtain data for the model made of marbles held together with wax and shown in two views in text-figures I and 2. Text-figure I shows the anterior pole of the young colony with the four cells forming a square, as has been observed by all students of the group. The opposite posterior pole of the colony would show a similar group of four, each cell alternating with the four at the posterior pole. HARPER: BINARY FISSION AND SURFACE TENSION 161 These are the relations of the two groups of four, as described by Biitschli and Overton. As noted, it is with reference to the position of the eight equatorial cells that clear description has been lacking. Text- figure 2 shows an equatorial view of the model and it is obvious at once that a least surface configuration requires the eight equatorial cells to form a zigzag belt, each cell alternating with two cells of the polar groups of four. With this arrangement the colony is made up of eight groups of three cells, the most compact arrangement possible, each of the four cells of each polar group appearing in two of the groups of three. The groups of three are so placed with reference to each other as also to form the groups of four seen in both the polar and equatorial views. The whole colony would consist of eight of these groups of four, since each of the eight equatorial cells is a member of three such groups and each cell of the polar group is a member of two such groups. The whole forms as perfect an approximation to a least surface configuration as can be achieved by sixteen cells arranged in as nearly as possible a spherical group. Such a configuration may be regarded, it seems to me, as the type configuration and illustrates the inter- action of surface tension, adhesion and binary fission as morphogenetic factors. In the actual colonies the cells are, of course, flattened upon each other and, as I have pointed out, elongate during division. As the photograph (Fig. 7, Pl. Il) shows, they are quite variable in both form and size. As is shown clearly by Klein also in all figures of the so- called polar openings, phialopores, the posterior group of four do not form any such definite square group as is maintained by the anterior group of four, yet the tendency to the formation of the groups shown in the model is, it seems to me, obvious. The abundance of pentagons and hexagons in the cell outlines of the adult colonies is good evidence of a tendency to the most compact configuration possible. I have not attempted to make the model with any great accuracy. We lack data as to the relative efficiency of adhesion and surface tension in the cells, which would be necessary for the exact determina- tion of their interrelations. The actual relations of fission, adhesion and surface tension in the processes just described are perhaps brought out more clearly in the two diagrams (Text-figs. 3 and 4). If the protoplasm of the mother cell were a mass which grows merely by imbibition and swelling, and if the cutting up of the mass into cells were merely a secondary phenomenon following the principle of the rectangular intersection of the cleavage planes and surface tension, as. Hofmeister (’67), Sachs (’78), and other critics of the cell theory have assumed, we should expect a configuration of the cells like that shown in text-figure 4. This is the sort of configuration Magnus 162 BROOKLYN BOTANIC GARDEN MEMOIRS (13) has obtained in his very interesting and suggestive experiment with paraffine wax cooling over mercury, a configuration determined entirely by the molecular forces operating in the system. It would be possible to form a sphere out of these nineteen units, but there is nothing in their number to favor the change from the discoid to the spherical grouping. To make a hollow or solid sphere out of such a group would involve very fundamental rearrangements. On the other hand, if we make a diagram of the arrangement of the sixteen cells produced by binary fission, assuming for the sake of simplicity that each pair of daughter cells instead of remaining flattened upon each other with the resulting lateral displacement (Klein, Taf. VI, Figs. 61-63) rounds up completely after the third division and that the four cells first formed remain fixed by adhesion, we get the 3 4 Fics. 3 and 4. For explanation see text. configuration shown in text-figure 3. Here it is obvious that as con- trasted with the arrangement in text-figure 4 it is a simple matter for the group to become cup-shaped and spherical simply by folding in the radial series and that the four outermost cells will come together ina group of four with its members alternating with those of the original group of four. The group produced by binary fission in two planes with elongation of the cells upon their free surfaces and strong adhesion tends naturally, especially in the cavity of a spherical mother cell, to produce a globular young colony at the sixteen-celled stage. As noted, all authors agree in maintaining that all the cells of the colony divide up to the sixteen-cell stage when the spherical form is practically complete. The later stages have not been followed. HARPER: BINARY FISSION AND SURFACE TENSION 163 Biitschli regards it as an open question whether all the cells divide in the later stages. It is, however, obvious that successive bipartition of all the cells is the natural method of maintaining the globular form already achieved. Any excess or deficiency of the number of divi- sions in any considerable group of cells would manifest itself at once as a bulge or depression in the surface of the expanding sphere unless compensatory divisions elsewhere and far-reaching, gliding move- ments of the cells among one another were possible. There is no evidence either of the occurrence or the possibility of such movements. As has been many times observed, the daughters of the original group of four can be recognized late in the life of the colony in their original positions with respect to each other and the colony as a whole. Oltmanns (04) follows Goebel (’82) and Goroschankin (’75) in asserting that this original polar group of four cells does not divide after the sixteen-cell stage but does not give any very positive evidence on the point. Overton (’89, Taf. III, Fig. 18) in a colony of about 200 cells shows a group of eight cells, six of which are dividing. It is, of course, obvious that a failure of the original group of four to con- tinue dividing after the sixteen-celled stage would not prevent the maintenance of the rounded form of the colony in case there were compensating divisions in the adjacent cells. It is interesting to note that Kirchner (’79) finds the development of the colony from the fertilized egg of V. aureus essentially like that of the asexual germ cell. His figures give some indication of the elongation of the cells before division and he describes the cup-shaped form of the colony in the eight-cell stage. It seems to me that we are justified in concluding that Volvox, though showing deep-seated specialization of somatic and germ cells in which it contrasts markedly with Eudorina, Pandorina and Gonium, still like them shows vegetative totipotency and equivalence of its cells in the growth of the colony. This is an important consideration in view of the question as to the origin of the differentiation of germ and somatic cells which is so conspicuous in the adult colony of Vol- vox, and entirely lacking in the simpler members of the series. This differentiation is such that the germ cells are distributed solely in a single half or three fourths of the colony, the remaining portions re- maining persistently somatic-sterile. The fertile area of the colony is regularly the posterior half or three fourths as the colony swims. It would be of great theoretical interest in this earliest appearance of the differentiation of soma and germ plasm if it could be shown that the cells bearing the germ plasm were different in cell lineage, age since last division, relative maturity as indicated by total number of divisions undergone, or in any other way, from the remaining cells of 164 BROOKLYN BOTANIC GARDEN MEMOIRS the colony which show no capacity for reproduction and apparently undergo senile degeneration. If, for example, as is so commonly and loosely stated in textbooks, the colony were formed by marginal growth and cell division, forming first a curved plate and finally a sphere, so that the cells at one pole would be ontogenetically younger than those at the other, we might expect this to be the basis for differentiation of germ plasm and soma. The evidence is, however, that all the cells of the adult colony are of the same generation and ontogenetically equivalent. The difference in their behavior is to be sought, then, in their relative environment and internal development as the colony grows. Their position in the posterior portion as the colony swims and around the pole which is nearest the point of connection between daughter colony and mother colony are obvious epigenetic factors in their environment which may be of significance. The distribution of the parthenogonidia at relatively equal intervals may be due to diffusion phenomena affecting nutrition directly or as stimuli, the whole com- plex perhaps suggesting analogy with Liesegang phenomena. The attempt to differentiate the eight daughter colonies commonly formed in asexual colonies of V. globator as descendants of the eight cells produced by the third division seems to me wholly artificial. This third division is not essentially different from the other divisions. In V. aureus also the number of daughter colonies varies. Meyer (’95) notes that the protoplasmic connecting strands between the cells are more numerous in this region of the germ cells than in the anterior part of the colony. They are especially well developed between the germ cells and the sterile cells as Janet’s diagrams show so strikingly (’12, Fig. 4). It is also in this region, as noted, that the young colony maintains its connection with the mother colony through protoplasmic strands from the cells around the posterior polar opening which connect with the adjacent cells of the parent as shown by Overton (’89, Taf. III, Fig. 16), and Janet (12, Fig. 1). The germ cells are borne then in that region of the colony which up to birth was most directly connected with the mother colony and perhaps received from it a large amount of food materials in the early growth stages. The antheridia of V. globator form the so-called packets of anthero- zoids consisting of bundles of sixteen to thirty-two gametes. These are formed by binary fission of the mother cell in two planes. The eight-cell stage shows the wheel figure. The cells instead of forming a globular colony ordinarily form a flat plate like the simpler Goniwm. Whether this is really due to a tendency to recapitulation retained in the sexual germ-cell formation or whether it is due to the elongation HARPER: BINARY FISSION AND SURFACE TENSION 165 of the male gametes as they form, is not clear, though it is generally agreed that the cell arrangement in the bundle of male gametes is homologous with that of the young vegetative colony. Klein (’97) describes colonies of antherozoids in the form of hollow spheres. Stein (’78) had also observed them. Chodat (’02) reports plate-shaped individuals of Pandorina. These cases indicate how closely the plate-shaped and the spherical colonies are related physi- cally. The globular antherozoid bundles in Volvox may well be regarded as tending toward the vegetative and away from the sexual condition. That such changes should involve changes in surface tension and adhesion is quite conceivable, as the elongation of the antherozoid cell body must of course be a factor in determining the form of colony. The further study of the formation of these abnormal male colonies, especially, should throw light on the whole series of morphogenetic problems here involved. The facts as known are certainly quite in harmony with the view that such presumably easily influenced factors as adhesion and surface tension, combined with the more fundamental and ever-present incompatibility between the principles of binary fission and least surfaces, may be of determining significance in the transition from the plate-shaped to the three- dimensional globular form of colony with all its evolutionary sig- nificance. INDEX OF LITERATURE Cohn, F. Untersuchungen ueber die Entwickelungsgeschichte der mikroskop. Algen u. Pilze. Nova acta Leop.-Carol. 24 (1): Ior. 1854. Henfrey, A. Notes on Some Fresh-water Confervoid Algae. Trans. Mic. Soc. London, N.S., IV, p. 49. 1856. Hofmeister. Die Lehre von der Pflanzenzelle. Leipzig. 1867. Braun, Al. Bemerkungen zu Cohn’s Schrift ueber Volvox. Sitz.-Ber. d. Ges. naturf. Freunde zu Berlin, Bot. Z. 190. 1875. Braun, Al. Ueber einige Volvocineen. Sitz.-Ber. d. Ges. naturf. Freunde, Bot. Z. L895 1875. Cohn, F. Die Entwickelungsgeschichte der Gattung Volvox. Beitr. z. Biologie der Pflanzen 3: 93. 1875. Goroschankin. Die Genesis bei den Palmellaceen. Versuch einer vergl. Morpho- logie der Volvocineae. Nachr. d. Kais. Ges. f. Naturw. usw. Moskau, 16. 1875. Sachs, J. Ueber die anordnung der Zellen im jiingsten Pflanzenteilen. 1878. Stein, Fr. v. Organismus der Infusionstiere. 3: I. 1878. Kirchner. Ueber die Entwickelungsgeschichte von Volvox minor. Cohn’s Beitrage 3:95. 1879. Goebel, K. Grundziige der Systematik und speziellen Pflanzenmorphologie. 1882. Falkenberg. Die Algen. Schenk’s Handbuch, p. 284. 1882. Biitschli. Protozoa. Bronn’s Klassen u. Ordn. d. Tierreichs. 1. 1883. Klebs, G. Organization einiger Flagellatengruppen und ihre Beziehungen zu Algen und Infusorien. Arb. d. bot. Inst. Tiibingen 1: 339. 1883. 166 BROOKLYN BOTANIC GARDEN MEMOIRS Klein, L. Morphologische und biologische Studien ueber die Gattung Volvox. Pringsh. Jahrb. 20: 133. 1889. Klein, L. Neue Beitrage zur Kenntnis der Gattung Volvox. Ber. d. d. bot. Ges. VeRA2» losg: Overton, E. Beitrag zur Kenntnis der Gattung Volvox. Bot. Centralbl. 39: 65. 1889. Klein, L. Vergl. Untersuchungen ueber Morphologie und Biologie der Fortpflanzung bei der Gattung Volvox. Ber. d. naturf. Ges. zu Freiburg i. B. 5: 1890. Meyer, Arthur. Ueber den Bau von Volvox aureus Ehrb. und V. globator Ehrb. Bot. Centralbl. 63: 225. 1895. Swingle, W. T. Zur Kenntnis der Kern- u. Zellteilung bei den Sphacilariaceen. Prings. Jahrb. 30: 299. 1895. Kofoid, L. A. Plankton Studies. II. On Pleodorina illinoisensis, a New Species from the Plankton of the Illinois River. Bull. Il. State Lab. Nat. Hist. Ro 27g 1tskoys- R. Chodat. Algues vertes de la Suisse. Berne. 1902. Oltmanns. Morphologie u. Biologie der Algen, Jena, 1904. Harper, R. A. The Structure and Development of the Colony in Gonium. Trans. Am: Mic.)So0e! 31: 1655 1912. Janet, C. Le Volvox. Limoges, 1912. Magnus, W. Ueber zellenférmige Selbst-differenzierungen aus flussiger Materie. Ber. d. d. bot. Ges. 31: 290. 1913. Harper, R. A. On the nature of types in Pediastrum. Mem. N. Y. Bot. Gard. 6: Olay LOLG: DESCRIPTION OF PLATE II All figures of asexual reproduction in Volvox taken with Zeiss apochrom, ob- jectives and the compensating eye-pieces. Fic. 1. Two-celled stage of young colony. X about 600. Fic. 2. Second division just completed. XX about 600. Fic. 3. Four-celled stage of young colony. X about 350. Fic. 4. Four-cell stage, one cell elongating preparatory to third division. < about 350. Fic. 5. Ejight-cell stage showing the wheel or radial arrangement of the cells. x about 350. Fic. 6. Samestageaslast. The young colony photographed after being teased out of the mother cell. X about 400. Fic. 7. Young colony not yet set free from mother colony and cell walls not yet gelatinized.. X about 400. VOLUME I, PLATE Il. BROOKLYN BOTANIC GARDEN MEMOIRS. BINARY FISSION IN VOLVOX HARPER: FURTHER STUDIES ON THE INTERRELATIONSHIP OF MORPHOLOGICAL AND PHYSIOLOGICAL CHARACTERS IN SEEDLINGS OF PHASEOLUS! J. ARTHUR HARRIS Station for Experimental Evolution, Cold Spring Harbor, N. Y. INTRODUCTORY REMARKS In a series of papers published during the past several years I have emphasized the importance of investigations of the relationship be- tween the morphological and the physiological characteristics of the organ and of the organism. The structural variations of the organs of which the organism is made up are the resultant of intrinsic and extrinsic factors—of heredity and environment, or of nature and nurture. Morphogenetic processes must, therefore, be investigated by physiological methods, and be interpreted in physiological, and ultimately in physical and chemical, terms. The purpose of this paper is to supplement and extend the results of an earlier study? in which it was shown that in bean seedlings char- acterized by certain morphological variations from type, the develop- ment of primordial leaf tissue is less than in normal controls grown under conditions as nearly as possible identical. The data then available indicated that a reduction of the volume of primordial leaf tissue is associated with abnormalities of all the types studied, but that the type of variation influences, in some degree, the amount of reduction. In these first experiments the conclusions were based on primordial leaves only. The use of such leaves has the obvious disadvantage that they are completely formed in the seed, and undergo merely an enormous expansion (and an undetermined amount of differentiation) in the 1 Studies on the Correlation between Morphological and Physiological Char- acters, V. Studies I-IV of the series are to be found in Genetics I: 185-196. I916; 2eeto—2i2. OL 72 a2o2—-2008) TOL 7: 2 Harris, J. Arthur. Studies on the Correlation of Morphological and Physio- logical Characters: The Development of the Primordial Leaves in Teratological Bean Seedlings. Genetics 1: 185-196. Ig16. 167 168 BROOKLYN BOTANIC GARDEN’ MEMOIRS germination of the seed and the development of the plantlet to the stage at which measurements were made. Since the development of the primordial leaves during the germina- tion and establishment of the seedling is relatively great, it seemed quite legitimate to use the weight of green tissue produced by these leaves as a measure of the physiological capacity of seedlings of various types. The fact that these leaves are differentiated in the seed, does, however, constitute a valid objection against their use as a measure of the physiological capacity of the seedling. For such purposes a constant based upon some organ developed later in the life of the individual is desirable. One of the purposes of this paper is to present the results of deter- minations upon a later developed organ. The one chosen is the first trifoliate leaf. This leaf was used because groups of plants of a higher degree of uniformity can be selected at the time of maturity of this leaf than at any later stage in the development of the plant, and because the first compound leaf reaches a degree of maturity sufficient for the purposes of the present study before the primordial leaves are too old to be used for a series of determinations. It is, therefore, possible to repeat, at a slightly later stage of development of the plant, the determinations made on the primordial leaves in the first study as a basis of comparison with the work already done and with the series of constants to be obtained for the first compound leaves of the same plants. In the first investigation the green weight of the leaf tissue served as the fundamental measurement. In addition to this character certain measurements on the sap properties were also made. In the study of the saps some difficulties were encountered, and it seemed most desirable to discontinue that phase of the study temporarily and to carry out determination of dry weight and water content instead. These new measurements have, therefore, been added to these for green weight. * MATERIALS AND METHODS The materials upon which this study is based are the same as those previously employed—a mixture of slightly different strains of navy beans. The seeds which were germinated in the fall and winter months of 1916 were grown in field cultures in 1915. Seeds from individual plants were germinated in sand. In sorting, the morphologically aberrant seedlings were laid aside with a normal plant to serve as a check for each abnormal. An abnormal and a control seedling from the same seed plant and germinated in the same HARRIS: INTERRELATIONSHIP IN PHASEOLUS 169 seed flat were potted side by side in a three inch pot and allowed to grow to the proper stage of maturity under conditions as favorable as we were able to give them. Before the samples were taken, the plants were carefully inspected and all pairs, one member of which had died, had been injured or which showed in its subsequent development any abnormality in addi- tion to these specified were discarded. Note that there was no direct selection for the characters of the abnormal plantlets in this process, since both abnormal and control were discarded if either was unsuited for the purposes of the experiment. There probably was a fairly stringent imdzrect selection, since the death rate and the mutilation rate of the variant individuals was probably greater than that of the normals. Thus more pairs were probably discarded because of an injury to or the death of the ab- normal member of the pair than because of the death or injury of a normal member. The probability that the materials were somewhat selected before the physiological measurements discussed in this paper were carried out renders the findings of greater significance than they would otherwise be. After the pairs of seedlings had grown until the first compound leaf had attained its full size, and the second compound leaf was developing, but before the primordial leaves had materially deteri- orated, samples of leaves were taken by nipping off the laminae only, or the laminae and the single petiolule of the terminal leaflet in the case of the compound leaf. These samples of tissues, each from 100 plants, were enclosed in flasks, weighed, and dried to constant weight in a bath surrounded by boiling water. Thus the technique employed was exceedingly simple. Because of the size of the samples dealt with, the relative infrequency of the abnormalities, and the large number which had to be discarded, the routine has been excessively laborious. For example, the weighings of the 23 samples and checks discussed in the present paper involve 13,800 leaves gathered from 4,600 plants which were secured by germinating and classifying nearly half a million seedlings. The structural variation in the bean seedling which is probably the simplest, and the most frequent, is a slight vertical separation of the cotyledons which are normally sensibly opposite in insertion. The amount of the separation is difficult to express quantitatively, since it is in some degree dependent upon the length of the axis. In our studies of seedling variation in Phaseolus, three grades of separation of the cotyledons have been recognized. The line of demarcation between these grades is a quite arbitrary one. This is also true of 170 BROOKLYN BOTANIC GARDEN MEMOIRS the line between ‘‘normal,’’ and ‘“‘abnormal”’ as applied to the dis- tinction between plants which have cotyledons inserted on the same level and those which have one of the pair sensibly higher on the axis than the other. “Slightly but distinctly separated,’ has been the descriptive term used in our classification schedules. The cotyledons range in position from those which are just perceptibly not inserted on the same level to those which are perhaps two or three or four milli- meters apart. So imperceptible is the line of distinction between nor- mal and abnormal plants that in the classification of the seedlings frequent discussions arose concerning the normality or abnormality of individual plants. In the present paper I am considering only the simplest type of abnormality. This course has been followed for two reasons. First, the proof of the existence of a physiological differentiation associated with a very slight structural variation is of far greater in- terest than the demonstration of measurable physiological differentia- tion associated with great morphological variation. Second, other types of abnormality with which I have dealt are so difficult to secure in satisfactorily large series that the number of samples as yet avail- able is not sufficient to justify detailed comparisons between the different types of abnormality. I hope ultimately to be able to meet these difficulties. For the present the one type of structural devia- tion dealt with serves to illustrate the method and one phase of the results of the investigations. : PRESENTATION OF DATA Consider first of all the green weight of the organs selected. The average green weight of the primordial and of the first com- pound leaves for plants which are normal except for slight separation of their cotyledons is shown in Table I. With one single exception, the average weight of the primordial leaves of the normal plants is higher than that of the abnormal plants. — In the single exception to the rule, the difference is small in amount. The average weight of the first compound leaf produced by abnormal plants of this class is in every case but one lower than the weight produced by the sensibly normal individuals. The exception to the rule is the same sample as in the case of the primordial leaves. The average weight of primordial leaf tissue in the abnormal plants is .5873, the average weight for normal plants is .6680, and the average difference —.0807. The differences in mean weights range in the individual samples from +.0074 to —.1286. For the first compound leaf of the same plants the average weight of the tissues from abnormal individuals is .4797, from a normal plant it is .5610, while the average HARRIS: INTERRELATIONSHIP IN PHASEOLUS 171 difference between the sample and the control is —.0813. The differ- ences in average weight vary from +.0368 to —.2492. TABLE [| Mean Green Weight per Plant of Primordial Leaves and of First Compound Leaf Primordial Leaves ] | | First Compound Leaf Sample Abnormal 2 .6034 35 5648 36 5951 39 =| -5619 40 | .6096 4I | .6068 42 .5879 43 .6222 46 .5956 47 .7058 48 | .6389 49 | .5902 53 -5402 54 +5720 56 -5380 61 .5193 64 5853 65 5747 66 .5886 7C .6853 71 -5639 72 5565 73 5933 Control Difference | .7096 —.1062 .6767 —~ 1 LTO .6361 —.0410 16277 —.0658 .7052 —.0956 -7304 —.1236 -O141 — .0262 -7508 —.1286 -7160 — E204: .6984 |! +.0074 272 — .0883 16674 20772 5990 | —.0588 .6530 | —.0810 .5921 | —.0541 .5827 — .0634 .7052 —.1199 26928") =. 1108 .6790 | —.0904 7000)" =—-02T3 .6059 | —.0420 6744 | —.1179 .6140 —.1107 | Percentage Difference 15.0 16.5 Abnormal 5132 +5444 5931 .5160 ‘5179 -4877 .4712 .5008 -4645 .5841 -5593 .4960 -4491 .4091 +3994 -4443 +4530 .4402 .5246 -4794 4132 -3799 3933 +5929 .6188 .6254 -5549 .6138 .6140 .7204 .6115 .6019 +5473 | .6395 .5851 -4948 4547 .4646 4811 .5848 ‘5717 .5960 .4998 -4534 .4882 .4887 Control Difference poe SoEse | —-0797 | 13.4 0744" || “352-0 | —.0323 5.2 —.0389 7.0 | —.0959 15.6 (Sp —=s1263 20.6 | —.2492 34.6 | ——LtO7 18.1 —— lay Aa ee 2eO +.0368 | 6.7 —.0802 | 12.5 —.0891 15.2 —.0457 | 9.2 — .0456 10.0 | —.CG652 | 14.0 = O8 089 |e f.0 —.1318 | 22.5 SES ty |) 25.0 —.0714 | 12.0 — .0204 4.1 — .0402 8.9 —.1083 22):2 — :0954 19.5 If these differences be reduced to percentages by using the weight of the normal plants as a base, as shown in the final columns of each section of the tables, it appears that the primordial leaves of the morphologically aberrant plants are from 3.0 to 18.0 percent lighter «than the leaves of the normal plants in the 22 samples in which this relationship between the two types of plants holds for the primordial leaves. Thus the percentages are highly variable. the 23 determinations is 11.95 percent. The average for In the case of the first com- pound leaves, the percentage reduction ranges from 4.1 to 34.6 with an average of 14.06 in the 23 samples. Note that the percentage shows that the difference between the abnormal and the control sample is far less in the case of the single exception, sample 47, than it is in Thus it is only 1.1 as compared with an average the average series. value of 11.95 for the primordial leaves and only 6.7 as compared with the average of 14.06 percent in the compound leaves. I now turn to a consideration of dry weight. 172 BROOKLYN BOTANIC GARDEN MEMOIRS The primordial leaves of the abnormal plants in which the two cotyledons are slightly separated are, as shown in Table II, lighter TABLE II Mean Dry Weight per Plant of Primordial Leaves and of First Compound Leaf Primordial Leaves First Compound Leaf SOUS | : Percentage : | Percentage | Abnormal | Control Difference | Difference} Abnormal | Control | Difference | Difference 2 OLAS Ee O53)740 |) =. 0002 Lau .0442 | .0517 | —.0075 | 14.5 35 | .0366 | .0483 —.O117 | 24.2 .0465 10530 |) 20065) |peress BO) 0422 |) 50457; |) 0035 |" 17-7 .0476 | 0499 | —.002 4.6 39 ~+| .0409 | .0467 —.0058 12.4 .0430 | .0470 | —.o040! 8.5 40 0438 | .O5II 0073, 14:3 .O415 0496 | —.0081 | 16:3 4I | .0431 .0526 »| —.0095 LSet .0406 | 0494 | —.0088 | 1728 2 .0416 | .0504 | —.0088 17.5 .0383 0519, | —.0136 | 26% 43 0429 | .0532 —.0103 19.4 0391 | .0493 | —.O102 20.7 46 .0408 | .O501 — .0093 18.6 .0400 | .0492. | —.0092 | 18.7 47 .0442 | .0446 — .0004. 9 0442 | .0433 =-<0009) | 228 ASH y\esc0420) |) 0464) 15-0044 5) 1955 0444 | .0525 —.0081 | 15.4 AQw |) -OZ8L i) LO4260)|/—— 0055 12.6 0397 | .0472 —.0075 | 15.9 53 0365 | .0410 | —.0045 11.0 0399 | .0427 | —.0028 | 6.6 54 .0384 0445 | —=20061 4|| 13-7 .0339 | .0412 —.0073 Wy 56 | .0349 | .04g1 —.0142 ; 28.9 .0331 | .0395 —.0064 ! 162 61 0356 | .0402 —.0046 11.4 .0383 .0417 —.co34 | 8.2 64) | .6354 | And, with less assurance, it may 5 Mar. Alg. Dan. W. I. 2: 95-99. f. 102-104. 1916. VOLUME |, PLATE III. BROOKLYN BOTANIC GARDEN MEmoIRS. (ANTHERIDIAL, SECTION GALAXAURA MARGINATA (ELL. & SOLAND.) LAMOUR. Howe: ““VEPRECULAE. ’) BROOKLYN BOTANIC GARDEN MEmoiRS VOLUME |, PLATE IV. HOWE: 1. GALAXAURA MARGINATA (ELL. & SOLAND.) LAMourR. (TETRASPORIC, SECTION *“* BRACHYCLADIA.’’) 2. GALAXAURA FLAGELLIFORMIS KJELLM. (SECTION ‘‘ RHODURA.”’) 3. GALAXAURA SQUALIDA KJELLM. (SECTION ‘*‘ MICROTHOE.’’) HOWE: DIMORPHISM IN GALAXAURA 197 be surmised that Galaxaura oblongata (Ell. & Sol.) Lamour.® has its tetrasporic condition in G. comans Kjellm. And just as the line of demarkation between Galaxaura oblongata and G. cylindrica seems a little uncertain and arbitrary, so also is the line of separation between G. comans and G. lapidescens. Where there is so much difference in habit as there is between the ‘“ Rhodura’’ forms on the one hand and the ‘“Microthoé” and ‘“‘Eugalaxaura’’ forms on the other, there is manifestly more need for a cultural demonstration of their correlation as alternating generations than there is in the case of the Cameratae- Spissae and Brachycladia-Vepreculae groups, where the two phases have the same outward appearance. But while experimental demonstration or further observations in the field may be desirable for a precise correlation of the ‘“‘Rhodura”’ forms, the existing evidence that these ‘‘Rhodura”’ forms represent tetrasporic phases of ‘‘ Micro- thoé”’ and ‘“ Eugalaxaura’’ forms seems convincing. EXPLANATION OF PLATES III AND IV PLATE III Photograph of a formalin-preserved antheridial specimen of Galaxaura marginata (Ell. & Soland.) Lamour., representing Kjellman’s section ‘‘Vepreculae.’’ Speci- men from San Juan, Porto Rico (Howe 2304); natural size. PLATE IV Fic. 1. Photograph of a formalin-preserved tetrasporic specimen of Galaxaura marginata (Ell. & Soland.) Lamour., representing the section “ Brachycladia.”’ Specimen from San Juan, Porto Rico (Howe 2304); natural size. Fic. 2. Photograph of a formalin-preserved specimen representing a form of Galaxaura flagelliformis Kjellm. and belonging in Kjellman’s section ‘‘ Rhodura.”’ Specimen from Santurce, Porto Rico (Howe 1859a); natural size. Fic. 3. Photograph of a formalin-preserved specimen of Galaxaura squalida Kjellm. Specimen from Santurce, Porto Rico (Howe 1859b); natural size. 6 Galaxaura fragilis of Kjellman and of Borgesen; not Dichotomaria fragilis Lamarck, the type specimen of which in herb. Mus. Paris. appears to have the structure of the ‘‘Spissae’’ group. THE UREDINALES OF OREGON! HH. S. JACKSON Purdue University Since no account of the rusts of any of the states bordering on the Pacific coast is available for reference by the students of the flora of that region, it has seemed desirable to bring together in the form of an annotated list the results of a study of the species occurring in Oregon, which has extended, intermittently, over a period of eight years. The account is the result of a study begun by the author in 1909 at the Oregon Agricultural College and finally prepared in the form here presented at the Purdue University Agricultural Experiment Station. On account of the great diversity of conditions the State of Oregon presents a wonderful field for work in any phase of botanical study. The area of the State is approximately 96,000 square miles, an area considerably greater than that included in all the New England states combined. The great range of climatic and topographical features existing in the State offers favorable conditions for the development of a flora not only large in number of species, but very diverse in char- acter. Within the confines of the State is to be found a range in alti- tude from sea level to perpetually snow-capped mountains. The annual rainfall varies from over 80 inches in some localities to below 10 inches in others, resulting in the development of a flora almost tropical in its luxuriance on the one hand, and one having many of the characteristics of a desert on the other. The diversity of the Phanerogamic flora which has developed under these conditions offers an especially attractive field for the student of the parasitic fungi. Many species of all groups are to be found especially in those portions of the State having a heavy annual rainfall. Rusts are found in great profusion in all sections of the State. The first collections of this group made by the writer were accumu- lated in connection with an effort to obtain general material for class use. The greater part of the collections have been made in connection with local excursions, the primary object of which was recreation. 1Contribution from the Botanical Department of Purdue University Agri- cultural Experiment Station. 198 JACKSON: UREDINALES OF OREGON 199 Many of the collections made by the writer at localities outside of Benton County have been picked up in spare moments on trips taken in connection with Experiment Station or Extension Service duties. During 1914 and 1915, however, a number of special excursions were made primarily for collecting this group of fungi. In addition to those made by the writer, several hundred collections made by his former associates, assistants and students at the Oregon Agricultural College are included. The greater number of these were collected by Prof. H. P. Barss, Mr. F. D. Bailey and Mr. G. B. Posey. To these have been added a considerable number of records obtained from miscellaneous sources. Several of these were obtained from the herbarium of the New York Botanical Garden, and of the National Museum. A considerable number are in the Arthur Herbarium at the Purdue University Agricultural Experiment Station. The greater number of these were obtained originally from phanerogamic speci- mens mainly collected. and distributed by pioneer botanists of the region, particularly W. C. Cusick, Thomas Howell, J. B. Lieberg and E. P. Sheldon, E. R. Lake and others. A few collections were made in Oregon by Dr. David Griffiths and associates, most of which were distributed in his ‘‘West American Fungi.” Mr. E. Bartholomew collected at a few localities in Oregon in 1915,and distributed the specimens in the exsiccati, ‘‘ Fungi Colum- biani’’ and ‘‘ North American Uredinales,”’ which he edits. A number of specimens of rusts, the records of which were obtained mainly from the Arthur Herbarium, were made by Moses Craig, at one time botanist at the Oregon Agricultural College. It is evident that he made quite an extensive collection of rusts in Oregon, but the location of his collection at the present time is unknown to the writer. One of the most interesting collections which it has been the privilege of the writer to examine was made by Dr. J. R. Weir, mainly in the northeastern and southwestern parts of the State. This col- lection consists of about 130 numbers and was sent to this laboratory for study in 1915 and 1916. Another interesting collection of about 30 numbers was made by Dr. E. P. Meinecke in southwestern Oregon and forwarded to the writer for study. Approximately thirteen hundred collections have been examined in the preparation, of this account and are listed in the following pages. By far the greater number of these, about one thousand, were made in western Oregon, including the Cascade Mountains. Of this number about six hundred were made in the Willamette Valley, four hundred having been collected in Benton County, mostly in the vicinity of Corvallis. Two hundred and fifty are listed from the Cascade moun- tain region, most of the collections having been made in the vicinity 200 BROOKLYN. BOTANIC GARDEN MEMOIRS of Mt. Jefferson or in Hood River County. About sixty collections are recorded from the counties bordering on the coast, only a few of which are from the southern coast counties. Less than three hundred collections have been made in eastern Oregon, almost half of which are from the mountainous region of the northeastern part. It will be seen from the above summary of the distribution of the collections recorded that only a very small portion of the State has been explored for this group of fungi. Much remains to be done, particularly in southwestern Oregon and in eastern Oregon. The mountains of the Coast Range have been explored only in Benton County, where several collections have been made on Mary’s Peak. The coast counties also offer a rich field for the collector. The region around Corvallis in Benton County is the only portion of the State that can be said to have been carefully explored for rusts. Corvallis is situated at the junction of the Willamette River and Mary’s River and lies in the edge of the foothills of the Coast Range. In spite of the fact that the exploration so far conducted is in- adequate to furnish a very accurate idea of the rust flora of the State, records for 220 species are brought together in the account which follows. These occur on about 500 different host plants. In connec- tion with the study of the collections 10 species have been found which are believed to be new to science, 8 of which are described in the following pages. A number of European species not previously recorded for North America have also been collected in the State. A large number of unrecorded host plants have been encountered. The number of species of economic importance found in the State is worthy of mention. All of the grain rusts recorded for North America with the exception of the corn rust (Puccinia Sorghi) are known to occur in the State, including the recently discovered Puccinia glumarum. All of the rusts troublesome to florist’s crops in the greenhouse, including Puccinia Antirrhini, are known to occur. The Pacific coast rust of pears and quinces promises to become of con- siderable economic importance. The large number of forest-tree rusts found in the State offers an especially attractive field for investi- gation. Many heteroecious species are known to occur whose life history is not yet determined.. The rusts occurring on Salix are especially in need of investigation. In the notes given in connection with the account of the species which follows, an effort has been made to summarize the available information on the life history as shown by any culture work which may have been conducted either by American or European authors. Notes on the distribution in North America are given whenever it was considered of sufficient importance. The genera are listed alpha- JACKSON: UREDINALES OF OREGON 201 betically under each family and the species similarly under the genus. The host plants are also arranged alphabetically under each species. No attempt has been made to give a full list of synonyms. In general, however, sufficient synonymy is given to show the origin of the specific name used as well as any names which have been in general use. For convenience of reference the specific nomenclature used is that in use in this laboratory, and in general conforms to that used in the ‘‘ North American Flora.’’ For similar reasons the generic nomen- clature follows that proposed by Dr. Arthur, for use in the “ North American Flora,’ except that Melampsora, Puccinia and Uromyces, are retained. In order to follow this system consistently it has been found necessary to establish a number of new combinations. Under each host is given a list of the specimens examined, with locality, county, date and collector, followed by the collector’s number. Numbers between 1000 and 3450 refer to collections in the Oregon Agricultural College Herbarium, if made by members of the botanical staff or by students. If no collector is given it may generally be assumed that the collection was made by the writer. In order to prevent unnecessary repetition, Benton County is not cited following. collections made at Corvallis and Philomath. Similarly Hood River County is implied for all collections from Hood River or Mt. Hood and all collections made at Portland are from Multnomah County. The writer is under great obligations to all those who have contrib- uted specimens for study and especially to those whose names have been mentioned previously. He is also greatly indebted to those botanists who have given so generously of their time in determining host plants. Dr. A. S. Hitchcock and Mrs. Agnes Chase have named most of the grasses. Dr. Theo. Holm and Dr. K. K. Mackenzie have each determined a considerable number of species of Carex. Dr. C. R. Ball has named most of the willows. Dr. F. V. Coville and Dr. K. M. Wiegand have each determined several specimens of Juncus. Mr. Paul Standley has determined a large number of specimens from miscellaneous families. Dr. J. C. Arthur and his former associates very kindly determined a considerable number of specimens of the rusts occurring on grasses and sedges, and verified the determinations of others, which the writer sent from Oregon at various times during the period when the col- lections were being made. The greater part of the collections recorded, however, have been worked over since the writer took charge of the work of this laboratory. During this period it has been his privilege to be able to consult freely with Dr. Arthur and to have the unre- stricted use of the collections, catalogues, and manuscript notes on the rusts, which have been accumulated at the Purdue University 202 BROOKLYN BOTANIC GARDEN MEMOIRS Agricultural Experiment Station during the many fruitful years of Dr. Arthur’s administration of the department of botany. Without this assistance the preparation of this account in the form presented would not have been possible and the writer takes great pleasure in acknowledging his indebtedness to Dr. Arthur and to the various assistants in this laboratory for any help which they may have given. COLEOSPORIACEAE 1. Coleosporium Adenocaulonis sp. nov. O.and I. Pyenia and aecia unknown. II. Uredinia hypophyllous, few, scattered on conspicuous angular yellowish spots, small, 0.I-0.2 mm. across, early naked, orange yellow fading to whitish, becoming pulverulent, ruptured epidermis con- spicuous; urediniospores globoid to ellipsoid, 18-24 by 23-26 un, wall light golden brown or colorless, 2—3 w thick, prominently and moder- ately verrucose; pores indistinct. III. Telia unknown. On CARDUACEAE: Adenocaulon bicolor Hook.—Corvallis, Sept. 20, 1914, 1540. This species is very inconspicuous, developing very small sori on the under side of the leaves on yellowish spots. 2. COLEOSPORIUM MADIAE Cooke, Grevillea 7: 102. 1879. On CARDUACEAE: II, III. Mada citriodora Greene—Mary’s Peak, Benton Co., Aug. 15, 1914, 1514. Madia exigua (Sm.) Greene—Corvallis, July 29, 1914, 1475; Philomath, Aug. 15, 1914, 15106. Madia glomerata Hook.—Corvallis, Aug., 1889, E. R. Lake, July, 1O10, 7750, July 29, 19015, 3241; Portland, Aug. 24, 1015, Essbar tholomew, 5964 (Barth. Fungi Columb. 4970). Madia racemosa (Nutt.) T. & G.—Corvallis, July, 1910, rz60, Sept. 12, 1910; 7925; Wren, Benton Co:, June 26; 1914, 13 70jmneme 1322, 1326; Blk City, Lincoln Co., Aug. 20, 1914, 2535; Philomach May I0, 1914, 3246. Madia ramosa Piper—Corvallis, July 29, 1914, 1470. Madia sativa Molina—Corvallis, Aug. 12, 1910, 11763, July 29, 1914, 1474. The aecial connection of this very common species has not been demonstrated by cultures and no field observations have been made in Oregon. Judging from distributional data, however, it seems probable that Peridermium californicum Arth. & Kern may be gen- etically connected. JACKSON: UREDINALES OF OREGON 203 From field observations made by the writer it is evident that in western Oregon this species overwinters in the uredinial stage. 3. COLEOSPORIUM OCCIDENTALE Arth. North American Flora 7: 94. 1907. On CARDUACEAE: IT. Senecio triangularis Hook.—Mary’s Peak, Benton Co., Aug. 15, 1914, 1518. This species is known otherwise only from the type collection made in Falcon Valley, Washington, on S. hydrophiloides Rydb., by W. N. Suksdorf in 1900. The aecial connection is not known and no clues are available. The aecia, in common with other species of Coleosporium whose life history is known, should be looked for on the leaves of Pinus sp. The above collection, however, was made in a region where no pines exist in a radius of several miles. It is probable that this species, in common with some other members of the genus, is capable of being carried over the winter in the uredinial stage. The only pine-leaf-inhabiting Peridermium known to the writer in the present range of this species is P. montanum Arth. & Kern, which has been shown to be genetically connected with a Coleosporium on Aster and Solidago, referred to C. Solidaginis. 4. COLEOSPORIUM SOLIDAGINIS (Schw.) Thiim. Bull. Torrey Club 6: 296. 1878. Uredo Solidaginis Schw. Schr. Nat. Ges. Leipzig 1: 70. 1822. Peridermium acicolum Und. & Earle, Bull. Torrey Club 23: 400. 1896. Peridermium montanum Arth. & Kern, Bull. Torrey Club 33: 413. 1906. On PINACEAE: I. Pinus contorta Dougl.—North slope Mt. Hood, Aug. 7, 1914, 1670. On CARDUACEAE: II, III. Aster conspicuus Lindl.—Hilgard, Union Co., July 10, 1914, 1532; Austin, Grant Co., Aug. 1915, J. R. Weir, 759. Aster Cusecku Gray ?—Corvallis, Sept. 21, 1914, 1548. Aster Douglastt Lindl.—Hood River, Aug. 26, 1915, E. Bartholo- mew, 5972 (Barth. Fungi Columb. 4grz); Corvallis, June 29, 1914, G. B. Posey, 1370. Aster foliaceus frondeus Gray—Hood River, July 22, 1915, 3137; Clatskanie, Columbia Co., May 20, 1914, F. D. Bailey, 2564, Oct. 29, 1914, 2531; Corvallis, June 29, 1914, G. H. Godfrey, 1307. Aster Hallit Gray—Corvallis, July 29, 1914, 1471; Wren, Benton @o., july 26; 1914;-F7 16. 204 BROOKLYN BOTANIC GARDEN MEMOIRS Solidago caurina Piper—North slope Mt. Hood, Aug. 7, 1914, 1005. Solidago elongata Nutt.—Corvallis, July 29, 1915, 3244; Scotts, 7 miles N. of Fort Klamath, Klamath Co., Sept. 20; 1912) Bee. Meinecke, Cr D 7. Solidago missouriensis Gray?’—Sumpter, Baker Co., Aug. 21, 1915, Jie Weir207- Solidago tolmieana Gray ?—Hood River, July 23, 1915, 3254. The life history of this species was first demonstrated by Clinton (Science N. S. 25: 289. 1907; Ann. Rep. Conn. Exp. Sta. 1906: 320. 1907; 1907: 375. 1908). He successfully infected Solidago rugosa with aeciospores of Peridermium acicolum on Pinus rigida. The single collection of aecia listed above (1610) agrees with the description of P. montanum Arth. & Kern and was collected in the immediate vicinity of Solidago caurina (1605). The possibility of genetic relationship was made note of at that time. Hedgcock (Mycologia 4: 144. 1912; Phytopath. 3: 16. 1913) has also made similar observations and more recently (Phytopath. 6: 65. 1916) has cultured this Peridermium successfully on Aster conspicuus, using aecial material on Pinus contorta collected in Montana. Weir and Hubert (Phytopath. 6: 68. 1916) working independently from Hedg- cock, with similar aecial material, have also demonstrated by cultures that this Peridermium has its uredinia on both Aster and Solidago, having obtained infection on A. laevis geyert, S. canadensis and S. Missouriensis. Sydow (Monographia Ured. 3: 621. 1915) suggests that the form on Aster in North America is different from C. Solidaginis on Solidago and should either be united with the Asiatic C. Asterum (Diet.) Syd. or that it represents an unrecognized species having a different Pert- dermium as its aecial form. The culture work of Weir and Hubert (1. c.), however, shows that P. montanum is genetically connected with uredinia on both Aster and Solidago and does not lend support to Sydow’s view. . While the two species of Peridermium included here are widely separated as to range and are morphologically distinguishable, it seems best until further culture work is conducted to recognize but one American species. UREDINACEAE 5. CALYPTOSPORA COLUMNARIS (Alb. & Schw.) Kiihn; Rab.-Wint. Fungi Eur. 3521. 1886. (Hedwigia 26: 28. 1887.) Aecidium columnare Alb. & Schw. Consp. Fung. 121. 1805. Calyptospora Geoppertiana Kiihn, Hedwigia 8: 81. 1869. JACKSON: UREDINALES OF OREGON 205 ON PINACEAE: I. Altes grandis Lindl.—Scottsburg, Lane Co., Sept. I, 1914, G. G. Hedgecock, 20270. Abies magnifica A. Murr.—Road to Crater Lake, Union Creek, Camp Grant, Klamath Co., Sept. 23, 1913, E. P. Meinecke, Cr D 20. ON VACCINIACEAE: IIT. Vaccinium macrophyllum (Hook.) Piper—Austin, Grant Co., June, 1913, J. R. Weir, 25; Sumpter, Baker Co., June, 1913, J. R. Weir, 24; Silver Creek, Josephine Co., July 28, 1913, E. P. Meinecke, Sz (D6) Dr. Vaccinium myrtilloides S. Wats.—Road to Crater Lake, Union Creek to Camp Grant, Klamath-Co., Sept. 23, 1913, E. P. Meinecke, CD) To: Vaccinium ovalifolium Smith—Larch Mt., Multnomah Co., Aug. 1910, 1156; North slope Mt. Hood, Aug. 7, 1914, 1608. Vaccinium ovatum Pursh—Dothan, Douglass Co., Sept. 8, 1914, G. B. Posey, 1932; Waldo, Josephine Co., Sept. 5, 1916, J. R. Weir, 280; Oregon, April 19-31, 1911, H. D. House. Vaccinium parviflorum Smith—Whitewater Ranger Station, near Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 1750. Vaccinium scoparium Lieb.2-—Mary’s Peak, Benton Co., Aug. 15, 1914, 1284. Specimens of aecia collected in various parts of North America on Abies balsamea, A. concolor and A. lasiocarpa are now referred to this species in the Arthur herbarium. The life history was first demonstrated by Hartig (Allg. Forst.- u. Jagdzeitg. 289. 1880), who conducted culture investigations using aecia on Abies pectinata and telia on Vaccinium Vitis-idaea. He obtained successful infection in both directions. Other European investigators, notably Dr. G. Winter, have amply confirmed these _results. (Klebahn, Die Wirtsw. Rostpilze 391. 1904.) In America, Arthur (Mycologia 2: 231. 1910) was the first to culture this species and succeeded in obtaining aecia on Abies Frasert following exposure to infection from telia on’ Vaccinium pennsyl- vanicum sent by W. P. Fraser from Nova Scotia. Later in the same year Fraser made the first field collection of aecia on Abies balsamea (Science 30: 814. 1909) and later (Mycol. 4: 177. 1912; 6: 27. 1914) confirmed Arthur’s work by obtaining infection on Abies bal- samea from telia on Vaccinium pennsylvanicum. 6. CHRYSOMYXA WEIRII Jackson, Phytopath. 7: 353. 1917. On PINACEAE: Picea Engelmanii Parry—Whitman Nat. Forest, Oregon, July 17, LOE, Pa ks Weir, 27-7; 206 BROOKLYN BOTANIC GARDEN MEMOIRS This species differs from C. Abtetis in the narrower, somewhat smaller spores which do not long remain in chains but soon break apart. No evidence of germination has been seen in any of the col- lections. This is the only American representative of the genus as restricted by Arthur. (Result Sci. Congr. Bot. Vienne 338. 1906.) It is known to the writer otherwise only from single collections from British Columbia and Idaho. It is doubtless not uncommon in the northwest. , 7. CRONARTIUM FILAMENTOSUM (PK.) Hedgcock, Phytopath. 2: 177. IQI2. Peridermium filamentosum Pk. Bot. Gaz. 7: 56. 1882. Uredo coleosporioides Dietel & Holway, Erythea 1: 247. 1893. Peridermium stalactiforme Arth. & Kern, Bull. Torrey Club 33: 419. 1906. Cronartium coleosporioides Arth. N. Am. Flora 7: 123. 1907. On PINACEAE: I. Pinus contorta Dougl.—Scotts, Anna Creek, Klamath Co., May 23, 1912, E. P. Meinecke, used for inoculation on Castilleja miniata; Gold Center, June 20, 1914, H. F. Wilson, 7856; North slope Mt. Hood, elev. 3,000-4,000 ft.; Aug. 7, 1914, 3332; Sumpter, Baker Co., May, 1916, J. R. Weir. On SCROPHULARIACEAE: II, III. Castilleja sp.—North slope Mt. Hood, 3,000-4,000 ft., Aug. 7, 1914, 1612, 1615 (collected near 3332); Ashland Toll House, Jackson Cos, Sept. 27; 1913, E: P..Meimnecke, Cr D 22. Hedgcock (I. c.) was the first to publish a record of connection of Peridermium filamentosum with a Cronartium on Castilleja by cul- tures. He considered this distinct, however, from Cronartium coleo- sporioides, which Meinecke had cultured in 1911 (Phytopath. 3: 167— 168. 1913) and shown to have for its aecial form P. stalactiforme. Meinecke’s culture material was collected in Klamath Co., Oregon. Further culture work has been carried on by Weir and Hubert (Jour. Agr. Research 5: 781-785. 1916) in which it is shown that the gall-forming Peridermium on Pinus contorta which has previously been commonly referred to P. Harknessit Moore is but a form of P. filamentosum. All the records of the aecial stage given above are of the gall- forming type. The Hood River material was collected in the im- mediate vicinity of the telial form on Castilleja. 8. CRONARTIUM PYRIFORME (Pk.) Hedgc. & Long, Alt. Stage Peri- dermium pyriforme 3, 1914. Peridermium pyriforme Peck, Bull. Torrey Club 6: 13. 1875. JACKSON: UREDINALES OF OREGON 207 Cronartium Comandrae Peck, Bull. Torrey Club 11: 50. 1884. Peridermium Betheli Hedge. & Long, Phytopath. 3: 251. 1913. On PINACEAE: I. Pinus ponderosa Dougl.—Hood River Co., May 10, 1910, 3333; Sumpter, Baker Co., May, 1916, J. R. Weir. On SANTALACEAE: II, III. Comandra umbellata (L.) Nutt.—Corvallis, June 20, 1909, E. R. Lake, 3068, July 24, 1914, 2510, Road to Ashland toll house, Jackson Go, cept. 27, 1913, E. P: Meinecke; Cr D°23; Hood River Co:, June 20, 1914, 1995, July 22, 1915, 3143; Dufur, Wasco Co., June 30, 1914, 1337; Indian Creek, Malheur Co., Sept. 16, 1897, E. P. Sheldon, 8934. The collection of aecia on Pinus ponderosa made at Hood River consisted of a large fusiform gall at the base of the trunk of a young tree about 2 inches in diameter. The gall entirely encircled the tree which was noticeably stunted from the effects of the parasite. The foliage also showed a distinct yellow cast. The life history of this common and widespread species was first demonstrated by Hedgcock and Long (Il. c.). They succeeded in obtaining the development of uredinia on Comandra umbellata by exposing them to infection from aecia on Pinus ponderosa collected in Washington and California and on Pinus pungens from Pennsylvania. In a later publication the authors (Bull. U. S. Dept. Agr. 247: 1-20. 1915) discuss the economic importance of this fungus as a disease of pines and record in detail the results of extensive culture work. g. HyALopsora Aspipiotus (Peck) Magn. Ber. Deuts. Bot. Ges. 19: 582. I9gol. Uredo Aspidiotus Pk. Ann. Rept. N. Y. State Mus. 24: 88. 1872. ON POLYPODIACEAE: Phegopteris Dryopteris (L.) Fée—Austin, Grant Co., Aug. 1915, J. R. Weir, 764. 10. HYALOPSORA LAEVIUSCULA (D. & H.) Arth. North Am. Flora 7: LEZ: 1907. Uredo laeviuscula Dietel & Holway, Erythea 2: 127. 1894. ON POLYPODIACEAE: Polypodium occidentale (Hook.) Maxon—Vicinity of Mt. Jefferson, July 27, 1907, E. R. Lake, 2508; Corvallis, March 25, 1915, G. B. Posey, 2626; Hood River Co., May 16, 1915, 3042; Bridal Veil, Multnomah Co., May 18, 1915, 3025. Polystichum munitum (Kaulf.) Presl., Mary’s Peak, Benton Co., Apr. 23, 1915, G: B. Posey, 3041. The urediniospores in this species are smooth in all collections as shown by very careful examination with the oil immersion objective. 15 208 BROOKLYN BOTANIC GARDEN MEMOIRS This species has not previously been recorded on the latter host so far as the writer is aware. 11. HyaALopsora Potypopit (DC.) Magn. Ber. Deuts. Bot. Ges. 19: hoe. TOOT. Uredo Polypodii DC. Fl. Fr. 6: 81. 1815. ON POLYPODIACEAE: Filix fragilis (L.) Underw.—Road to Lost Lake, Hood River Co., July 24, 1915, 3024. 12. MELAMPSORA sp. Il. Uredinia amphigenous, chiefly epiphyllous, scattered or occa- sionally gregarious, round, 0.5-I1 mm., early naked, somewhat pulverulent, orange fading to yellowish, ruptured epidermis not con- spicuous;. uredospores ellipsoid or obovoid, 15-19 by 21-24 u, wall colorless, uniformly 2.5—3 uw in thickness, moderately to closely verru- cose-echinulate; paraphyses numerous, chiefly peripheral, clavate or occasionally capitate, 18-26 by 45-70 yu, wall colorless, usually uni- formly 1-2 wu thick, occasionally thickened at apex to 4 up. ON SALICACEAE: Populus alba .—Sheridan, Yamhill Co., July 7, 1914, He Pe Barss, 1935; Cottage Grove, Lane Co., July 17, 1914, 1033; Phuilo- math, July 20, 1915, 3300. The only other American collection on this host known to the writer is one in the Arthur herbarium, collected by E. Bethel, Aug. 7, 1913, at San Jose, Cal. These specimens differ from all other North American collections on Populus. It seems most probable that this is an introduced European species. Only uredinia are present in American collections and it is quite impossible to assign it to any known species without telial material. A description of the uredinial stage drawn up from the Oregon collections is given for the benefit of those who may have occasion to study this form. The Oregon collections were all made from low, rapidly growing water sprouts. 13. MELAMPSORA ALBERTENSIS Arth. Bull. Torrey Club 33: 517. 1906. Caeoma occidentalis Arth. Bull. Torrey Club 34: 591. 1907. On PiNACESE: I. Pseudotsuga mucronata (Raf.) Suds.—Southeast Mt. Jefferson, Linn Co., July 3, 1914, i: DL Bailey,.z2541; Sumpter, BakersGe; July,.20, 1913, |. Re Weu2754 1 Convallis, June. too: The life history of this species has been studied by Arthur (Myco- logia 4: 29 and 59, 1912), who obtained infection resulting in pycnia JACKSON: UREDINALES OF OREGON - 209 and aecia on Pseudotsuga by exposing the foliage to infection from germinating telia on Populus tremuloides collected in Colorado. Out of four trials, three were successful. No infection was obtained on Larix: It is noteworthy in this connection that all of the northwestern collections have larger spores than specimens from Colorado. The former show spores 20-28 by 24-32 u while the average of the latter are 16-20 by 19-26 wp. The culture work was conducted with Colorado material, nearly if not all of which was collected in immediate associa- tion with M. albertensis on P. tremuloides. The type of Caeoma occidentale, on the other hand, was collected in British Columbia and has larger spores. It seems entirely possible that the northwestern collections may represent a different species and have genetic relation- ship with some form on Populus other than M. albertensis. 14. MELAMPSORA ARCTICA Rostr. Medd. Grénland 3: 535. 1888. ON SALICACEAE: Salix Bebbiana Sarg.—Sumpter, Baker Co., Aug. 1915, J. R. Weir, 167. Salix fendleriana And.—Sumpter, Baker Co., June, 1913, J. R. Weir, S. Salix lutea Nutt.—Sumpter, Baker Co., June, 1913, J. R. Weir, 4. Salix sitchensis Sanson—Dothan, Douglass Co., Sept. 8, 1914, G. B. Posey, 3342. Salix sp.—Scott’s, 7 miles from Fort Klamath, Klamath Co., Sept. 20, 1913, E. P. Meinecke, Cr D 6. It is with considerable hesitation that the above collections have been referred to this species. Only those collections which have small, rather thin-walled uredospores, accompanied by an abundance of thin-walled, clavate paraphyses, are included. Fraser, working with material collected in Nova Scotia (Mycol. 4: 187. 1912; 5: 238. 1913), has made a cultural study of this species. He succeeded in obtaining infection on Abies balsamea with production of pycnia and aecia following exposure to germinating telia from Salix discolor. 15. MELAMPSORA BIGELOW! Thiim. Mitth. Forstl. Vers. Oest. 2: 37. 1879. On PINACEAE: I. Larix occidentalis Nutt.—Hood River Co., elevation 4000°, July 23, 1915, 3305, 3305. On SALICACEAE: II, III. Salix Bebbiana Sarg.—Austin, Grant Co., Aug. 1915, J. R. Weir, 162, 163; Sumpter, Baker Co., July 19, 1913, J. R. Weir, 272. 210 BROOKLYN BOTANIC GARDEN MEMOIRS Salix cordata Muhl.—Sumpter, Baker Co., Aug. 1915, J. R. Weir, TO: Salix Piperi Bebb.—Philomath, Oct. 29, 1911, 3346; The Dalles, Wasco Co., Aug. 26, 1915, E. Bartholomew (Barth. Fungi Columb. 4730). Salix pseudocordata Anders.—Hilgard, Union Co., July 10, 1914, 1530. Salix scoulerrana Barr.—Corvallis, Sept. 19, 1910, 17165; St. Johns, Multnomah Co., June 23, 1915, W. E. Lawrence, 3347; Austin, Grant Co:, Aug. 1915, J. R. Weir, 765; Portland, Aug. 24, tors Be Bartholomew (Barth., N. Am. Ured. 1417). Salix sp.—Calamity, Aug. 1901, Griffiths & Morris (Griffiths, W. Amer. Fungi 347); Crater Lake, Klamath Co., Sept. 22, 1913, E. P- Mieinecke, Cr Pk D (2) 13; Hood River Co., May 14, 1914 a5@0; Aug. 5, 1914, 1483, 1484; Beaverton, Washington Co., July 15, 1914, F. D. Bailey, 1507; Austin, Grant Co., Aug. 25, 1915, J. R. Weir, 262. The above specimens are tentatively assigned to this species. There are quite certainly not less than four species of Melampsora on Salix in North America. The characteristics by which they may be separated in the uredinial stages are not well worked out at the present time. The larger spored forms have been included here under M. Bigelow. Arthur (Jour. Myc. 11: 60. 1905) first established the fact that this rust has its aecia on Larix. He succeeded in infecting Larix decidua in two trials, by inoculating with basidiospores from germi- nating telia on Salix amygdaloides collected in Wisconsin. This result was later confirmed (Jour. Myc. 13: 194. 1907) with telial material collected in Indiana. Weir and Hubert (Phytopath. 6: 372. 1916) have succeeded in obtaining infection of this species from Salix bebbiana Sarg. collected in Montana on Larix occidentalis, and from SS. cordata mackenzieana collected in Idaho on Larix europea. The same authors (Phytopath. 7: 109. 1917) have recently repeated the work with the last-named species of Salix and obtained infection with development of pycnia and aecia on both L. occidentalis and L. europea. 16. Melampsora confluens (Pers.) comb. nov. Uredo confluens Pers. Obs. Myc. 1: 98. 1796. ON GROSSULARIACEAE: I. Ribes lacustre (Pers.) Poir.—Philomath, May 3, 1913, F. D. Bailey, 1107. ONS SALICACEAR: 1 TIT. Salix argophylla Nutt.—Freewater, Umatilla Co., June 17, 1913, F. D. Bailey, 1764, Aug. 12, 1915, F. D. Bailey, 3324. JACKSON: UREDINALES OF OREGON 211 Salix scouleriana Barr.—Cascade Locks, Hood River Co., Aug. fi 1O10,, 7276, Myrtle Creek, Douglass.Co., June 9, 1914, F: D. Bailey, 3345; Hood River Co., July 23, 1915, 3343, 3306; Ashland, Jackson Co., Sept. 10, 1914, 3340, 334I. Salix sp.—Scott’s, 7 miles north of Fort Klamath, Klamath Co., Sept. 20, 1913, E. P. Meinecke, Cr D 4, Cr D 9; Austin, Grant Co., Aug. 25, 1915, J. R. Weir, 263; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 266; White Pine, Baker Co., July 20, 1913, J. R. Weir, 270; Unity, Baker Co., Aug. 1915, J. R. Weir, 278. It is impossible to assign with any degree of certainty the collec- tions which should be referred to this combination. Only those col- lections having small, rather thick-walled spores, accompanied by an abundance of capitate, thick-walled paraphyses, are included. It is possible that some of the collections included under M. Bigelowii should be referred here. No culture work has been conducted in America. A summary of European work has been made by Klebahn (Die Wirtsw. Rostpilze 424. 1904). In addition to the above, aecia have been collected in America on Ribes saxosum from Utah, R. vallicola, Colorado, and R. lacustre, British Columbia. 17. MELAmpsorA LInI (Pers.) Desmaz. Pl. Crypt. (Fasc. 41) 2049. 1850. Uredo miniata Lint Pers. Syn. Fung. 216. 1901. On LINACEAE: Linum Lewisit Pursh—Blue Mts., 7,000-8,000 ft., eastern Oregon, 1897 (from phanerogamic specimen in Gray Herb. Harvard Univ.); Hermiston, Umatilla Co., May 12, 1915, 2664. Arthur (Jour. Myc. 13: 207. 1907) has shown this species to be autoecious. He sowed basidiospores from Linum usitatissimum on the same host and on L. Lewisii and obtained the development of pycnia and aecia. 18. MELAMPSORA OCCIDENTALIS Jackson, Phytopath. 7: 354. 1917. On SALICACEAE: II, ITI. Populus trichocarpa Nutt.—Corvallis, Sept. 1909, 1069, Oct. 15, 1912, 1024 (type), March 12, 1916, G. B. Posey; Trail to Sulphur Springs, Benton Co., Nov. 2, 1914, 3369; Scott’s, N. of Fort Klamath, Klamath Co., Sept. 20, 1913, E. P. Meinecke, CrD2; Clatskanie, Columbia Co., Oct-6, 1914, F..D. Bailey, 3358, Oct: 20, ro1g, F. D. Bailey, 3306; Sumpter, Baker Co., Aug. 21, 1915, J. R. Weir, 265; Medical Springs, Union Co., Aug. 1913, J. R. Weir, 117. This species differs from all other species of Melampsora on Populus IMD BROOKLYN BOTANIC GARDEN MEMOIRS in the large size of the urediniospores which are only slightly flattened and are evenly verrucose-echinulate. The teliospores are much longer than those of IV. Medusae and are thickened at the apex. The charac- ter of the telial sori suggests that this species may be closely allied to M. albertensis. The sori are much larger as are also both uredinio- and teliospores. This species may be the same as that recently cultured by Weir and Hubert (Phytopath. 7: 108. 1917), who used telial material from P. trichocarpa referred to M. Medusae and obtained successful infec- tion on Larix europea and L. occidentalis. The actual material used for infection and the aecia resulting have not been seen by the writer but telial material sent by Dr. Weir from Montana agrees with the form described above. Aecia from the same locality on L. occidentalis agree in general with aecia of Melampsora Medusae and M. Bigelowit. The walls of the aeciospores are however somewhat thinner, I-2 y, and considerably thickened on opposite sides to 3-5 uw. They measure 17-19 by 19-26 pw. Additional culture work, and a careful comparison of the resulting aecia with those of M. Medusae would be desirable. In any case, the morphological characters of the uredinial and telial stages are considered sufficient to warrant separation. 19. Melampsora Piscariae sp. nov. O.and I. Pycnia and aecia unknown. II. Uredinia hypophyllous, scattered, rounded, 0.3-0.5 mm. across, early naked, somewhat pulverulent, orange yellow fading to whitish, ruptured epidermis conspicuous; urediniospores globoid to ellipsoid, 14-16 by 16-19 uw; wall colorless, 1.5—2 w in thickness, finely and closely verrucose-echinulate; paraphyses numerous, intermixed with the spores, capitate, smooth or with an occasional conical echinu- late marking, 32-64 4 long; heads 12-18 yu broad, wall uniformly thick, 2:5-4 pz: III. Telia not seen. ON EUPHORBIACEAE: Piscaria setigera (Hook.) Piper (Eremocarpus setigerus Benth.)— Corvallis, Sept. 20, 1914, 3308, type. Known only from the type locality. This species is referred to the genus Melampsora with considerable confidence in spite of the absence of telia, on account of the structure of the sorus, the character of the spores, and the presence of scattered capitate paraphyses. 20. MELAMPSORELLA ELATINA (A. & S.) Arthur, N. Amer. Flora 7: [Li \TOO7: Aecidium elatinum Alb. & Schw. Consp. Fung. 121. 1805. JACKSON: UREDINALES OF OREGON 213 Melampsorella Cerastit (Pers.) Schroet. Krypt. Flor. Schles. 3!: 2060.1 E887. On PINACEAE: I. Abies grandis Lindl.—Mary’s River, west of Wren, Benton Co., Aug. 2, 1914, 1297; Sumpter,-Baker Co., July 20, 1913, J. R. Weir, 270. Abies lasitocarpa (Hook.) Nutt.—Crater Lake, Klamath Co., Sept. 9, 1916, J. R. Weir, 209; Sumpter, Baker Co., July, 1913, J. R. Weir, 274. On CARYOPHYLLACEAE: II, III. Cerastium vulgatum L.—Corvallis, May 1, 1915, 2667. Cerastium viscosum L.—Corvallis, June 28, 1915, 3079. Stellaria borealis Bigel.—Corvallis, April 5, 1914, 7287. This rust, which is doubtless common throughout the state, is remarkable in that both stages develop from a perennial mycelium. The aecial stage forms large or small witches’ brooms on the branches of various species of Abies, each leaf of which bears the conspicuous aecia in two rows on the under surface of the leaves. The life history was first worked out by Fischer (Zeitschr. fiir Pflanzenkr. II: 321. 1901) and has been amply confirmed by other European investigators. A summary of this work has been made by Klebahn (Die Wirtsw. Rostpilze 397. 1904). In America Arthur (Mycol. 4: 58. 1912), using aecial material on Abies lasiocarpa collected in Colorado, has succeeded in obtaining infection resulting in uredinia on Cerastium oreophilum. 21. MELAMPSOROPSIS PIPERIANA Arthur, N. Amer. Flora 7: 120. 1907. ON ERICACEAE: Rhododendron Californicum Hook.—Newport, Lincoln Co., June 2, 1892, A. Isabel Mulford (Specimen in Herb. N. Y. Bot. Gard. and in Herb. J. C.. Arthur), May 16, 1914, G. H. Godfrey, 1280; Larch Mt., Multnomah Co., Aug. 1910, rz18; Parmelia Lake, near Mt. Jefferson, July 3, 1914, F. D. Bailey, 1939; Trail to Hanging Valley, near Mt. Jefferson, Aug. 11, 1914, H. P. Barss & G. B. Posey, 1623. 22. MELAMpPSOROPSIS PyROLAE (DC.) Arth. Résult Sci. Congr. Bot. Vienne 338. 1906. Aecidium (?) Pirolae DC. FI. Fr. 6: 99. 1815. Aecidium conorum Piceae Reess, Abh. Nat. Ges. Halle 11: 102. 1869. Chrysomyxa Pirolae Rostr. Bot. Centr. 5: 127. 1881. Peridermium conorum Piceae Arth. & Kern, Bull. Torrey Club 33: AZESS tI COG: 214 BROOKLYN BOTANIC GARDEN MEMOIRS On PINACEAE: O, I. Picea Engelmanit Parry—Sumpter, Baker Co., Sept. 25, 1909, G. G. Hedgcock, 1916. On Pyro.aceaE: II, III. Pyrola secunda L.—North slope Mt. Hood, Aug. 7, 1914, 1607; Trail to Elk Meadows, Hood River Co., July 23, 1915, 3061; Columbia Highway, Multnomah Co., Aug. 19, 1916, J. R. Weir, 269. Pyrola sp.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R Weir, 268. The genetic relation of this species with Peridermium conorum Piceae was first suggested by Rostrup (I. c.).. So far as the writer is aware the first culture work confirming this observation was made by Fraser (Mycol. 4: 183. 1912), who succeeded in obtaining infection resulting in pycnia and aecia on the cones of Picea mariana and P. canadensis, following exposure to germinating telia on Pyrola americana and P. elliptica. 23. Milesia Polystichii Wineland n. sp. O. and I. Unknown. II. Uredinia hypophyllous, scattered, roundish, 0.2-0.3 mm. across, bullate, brownish yellow, tradily dehiscent by a central pore, peridium well developed, cells above polygonal, approximately isodiametric, diameter about 7 pu, cells at the sides elongated to 21 yu, outer walls 2-2.5 4, inner walls 2.5-3 mu; urediniospores obovoid, ellipsoid, or oblong, 18-23 by 26-35 u, wall colorless, 1.5-2.5 in thickness, strongly and sparsely echinulate, pores obscure. III. Telia unknown. ON POLYPODIACEAE: Polystichum munitum (Kaulf.) Presl.—Grant’s Pass, Josephine Co., Sept. 5, 1916, J. R. Weir, 260 (type). This species was separated from material referred to Hyalopsora laeviuscula in the writer’s herbarium by Miss Grace O. Wineland who has been studying the fern rusts of North America in this laboratory. 24. PUCCINIASTRUM ABIETI-CHAMAENERII Kleb. Prings. Jahrb. f. Wiss. Bot. 34: 387. 1900. On PINACEAE: I. Abies grandis Lind|.—Dee, Hood River Co., July 23, 1915, 3355. Abies lasiocarpa Nutt.—North slope Mt. Hood, 4,500 ft., Aug. 9, 1914, 3205. On ONAGRACEAE: II, ITI. Chamaenerion angustifolium (L.) Scop.—Bonneville, Multnomah Co., Aug. 11, 1910, 1075; Garden Home, Multnomah Co., Aug. 1911, 1990; Southwest slope Mt. Jefferson, July 3, 1914, F. D. Bailey, JACKSON: UREDINALES OF OREGON 215 3247; Odell, Hood River Co., Aug. 5, 1914, 1618; Crater Lake, Klamath Co., Sept. 21, 1913, E. P. Meinecke, Cr Pk D (2) 2; Portland, Aug. 24, 1915, E. Bartholomew (Barth., N. Am. Ured. 1482). This species is separated from P. pustulatum largely on the basis of culture investigations. All of the culture work has been conducted with the above host species or other members of the same genus or section of Epilobium. European investigators have amply demon- strated the connection of this form with aecia on Abies (Klebahn, Die Wirtsw. Rostpilze 393. 1904). In America, Fraser, working in Nova Scotia (Mycol. 4: 176. 1912), was the first to conduct culture experiments. He obtained, in three trials, the development of aecia on Abies balsamea, from sowings with teliosporic material from C. angustifolium. With the aecia thus obtained he sowed back to Chamaenerion and obtained uredinia. Weir & Hubert (Phytopath. 6: 373. 1916) conducted similar work with Idaho material and suc- ceeded in obtaining the development of aecia on Abies lasiocarpa. The aecia thus obtained were sown back on the telial host (Phytopath. 7: 109. 1917), with the result that uredinia were developed in abundance. 25. PuUCCINIASTRUM GALII (Link) Fischer, Ured. d. Schweiz 471. 1904. Caeoma Galu Link, in Willd. Sp. Pl. 6?: 21. 1825. ON RusiAceEseE: II. Galium triflorum Michx.—Corvallis, April 29, 1914, F. D. Bailey, 1992, May 9, 1914, 1991, July 5, 1914, H. P. Barss, 1996, June 29, 1914, G. B. Posey, 13173, July 10, 1915, 3104; Oregon City, Clackamus Co., Aug. 20, 1915, E. Bartholomew, 5934 (Barth. N. Am. Ured. , 1679); Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 287. This species is known to the writer from North America only from the above collections and a specimen collected by J. W. Macoun in British Columbia in 1915, one from Palmer Lake, Colorado, Sept. 6, 1913, by E. Bethel, both in the Arthur Herbarium, and a collection made by Dr. H. Fitzpatrick and the writer at Michigan Hollow Swamp near Ithaca, N. Y., July,1916. All are on G. triflorum. ‘The species is evidently common in western Oregon and if aecia are developed, doubtless occur on Abies grandis. Field observations made by the writer, however, would indicate that this rust winters over in the uredinial stage. 26. PUCCINIASTRUM GOODYERAE (Tranz.) Arth. N. Am. Fl. 7: 105. 1907. Uredo Goodyerae Tranz. Trudi S. Peterb. Obshch. Est. Otd. Bot. 235/255) L693: 216 BROOKLYN BOTANIC GARDEN MEMOIRS ON ORCHIDACEAE: II. Goodyera Menzies (Lindl.) Morong.—Parmelia Lake, West slope Mt. Jefferson, July 3, 1914, F. D. Bailey, 7627; North slope Mt. Hood, Aug. 9, 1914, 1620. 27. PUCCINIASTRUM MyrtTILLI (Schum.) Arth. Résult. Sci. Congr. Bot. Vienne 337. 1906. Aecidium ? Myrtilla Schum. Enum. PI. Saell. 2: 227. 1803. Pucciniastrum Vacciniorum (DC.) Dietel, in E. & P. Nat. Pfl. Teer Aen OO. ON VACCINIACEAE: II. Oxycoccus macrocarpus (Ait.) Pursh—Astoria, Clatsop Co., Aug. 1916, G. M. Darrow, comm. C. L. Shear 2905. Vaccinium caespitosum Michx.—Mary’s Peak, Benton Co., Aug. 15, 1914, 1517, 1520. Vaccinium macrophyllum (Hook.) Piper—Whitewater Ranger Station, West slope Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 3314; Ashland, Jackson Co., Sept. 10, 1914, 33106. Vaccinium ovatifolium Sm.—Whitewater Ranger Station, West slope Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 3375. Vaccinium sp.—North slope Mt. Hood, 4,000 ft., Aug. 7, 1914, 1600, 1609; Sucker Creek, Josephine Co., July 27, 1913, E. P. Mein- ecke, Sz (D6) D3. Clinton (Rep. Conn. Agr. ‘Exp. Sta. I909-IQIO: 719. I9QI1) was the first to show that the aecial stage of this species occurred on Tsuga canadensis. He successfully infected Gaylussacia baccata by sowing with aeciospores from Jsuga, resulting in the development of the typical uredinia of this species. Fraser in 1912 (Mycol. 5: 237. 1913) confirmed Clinton’s work by obtaining the development of aecia on the leaves of Tsuga canadensis following sowings from teliosporic material on Vaccinium canadense. The same author in 1913 (Mycol. 6: 27. 1914) obtained aecia on Tsuga canadensis following sowing of teliosporic material from Gay- lussacia resinosa. The aecia developed in these experiments are similar to those of Peridermium Peckti, but may represent an unde- scribed form. No aecia collected in the west have been referred to this species though they doubtless occur on Abies or Tsuga heterophylla. 28. PUCCINIASTRUM PUSTULATUM (Pers.) Dietel, in E. & P. Nat. Pfl. Pett A7 ee Teor. Uredo pustulata Pers. Syn. Fung. 219. {801. ON ONAGRACEAE: Epilobium adenocaulon Haussk.—Corvallis, Oct. 29, 1911, F. D. JACKSON: UREDINALES OF OREGON wa Batley, ni77.NoOv.4; 1O1l, F. D. Bailey, 7777, June 18, to14, F. DD. Bailey, 3218, July 29, 1914, 1450; Hilgard, Union Co., July 10, 1914, 1533, 1535, Glendale, Douglass Co., July 17, 1914, 1505, North slope Mt. Hood, Aug. 7, 1914, 1488; Whitewater Ranger Station, West slope Mt. Jefferson, H. P. Barss & G. B. Posey, 3219; Ashland, Jackson Co., Sept. 10, 1914, 3221. Epilobhium brevistylum Barbey—Corvallis, July 14, 1914, G. B. Posey, 3220. No successful culture work has been conducted with this form, as here interpreted, either in Europe or America. Aecia doubtless occur on species of Adbzes. From field observations it is quite evident that in western Oregon at least this species overwinters in the uredinial stage. 29. PUCCINIASTRUM PYROLAE (Pers.) Dietel, in E. & P. Nat. Pfl. res 47.) 1807. Aecidium Pyrolae Pers. Gmel. Syst. Nat. 2: 1473. I79I. Uredo Chimaphilae Peck, Ann. Rep. N. Y. State Mus. 46: 33. 1893. ON PYROLACEAE: Chimaphila umbellata (L.) Nutt. (C. occidentalis Rydb.)—Spencer Creek, Klamath Co., 5,000 ft., July 10, 1903, E. B. Copeland, 3774 (Sydow, Ured. 1795); Whitewater Creek along trail to Hanging Valley, Mt. Jefferson, Aug. 11, 1914, H. P. Barss & G. B. Posey, 1908; North slope Mt. Hood, Aug. 7, 1914, 1614. Pyrola secunda L., Klamath Co., July 10, 1903, E. B. Copeland (Sydow, Ured. 17971). 30. PUCCINIASTRUM SPARSUM (Wint.) E. Fischer, Beitr. Krypt. Schweiz 2?: 469. 1904. Melampsora sparsa Wint. in Rab. Krypt. Fl. 11: 245. 1881. ON ERICACEAE: Arbutus Menziesit Pursh—Myrtle Creek, Douglass Co., June 8, 1914, F. D. Bailey, 7837; Glendale, Douglass Co., July 17, 1914, 12985; Ashland, Jackson Co., Sept. 10, 1914, 1838; Corvallis, April, I9Q1I, 3374; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 244. Arctostaphylos Manzanita Parry—Grant’s Pass, Josephine Co., Sepinj 1o16;, |. KR .Weir, 245, 2247- Arctostaphylos nevadensis A. Gray—Northwest slope Mt. Jefferson, Auge 14, 1014, Hi. Ps Barss: & Gs B. Posey, 3290: No aecial collections have been referred to this species in America. Fischer (Cent. fiir Bakt. 46: 333. 1916) has cultured this species. He used germinating telial material on Arctostaphylos alpina and sowed 218 BROOKLYN BOTANIC GARDEN MEMOIRS on species of Abies and Picea, obtaining the development of pycnia and aecia on Picea excelsa. This is the only case in which culture investigations have shown the aecia of Pucciniastrum to occur on Picea. 31. UREDINOPSIS COPELANDII Sydow, Ann. Myc. 2: 34. Feb. 1904. Uredinopsis Atkinsoni Magn. Hedwigia 43: 123. Mar. 1904. Peridermium balsameum Peck, Rept. N. Y. State Mus. 27: 104. 1875= op-p: On PINACEAE: I. Abies grandis Lind|.—Trail to Sulphur Springs, Corvallis, Benton Co,.; Nov. 7, 1914,,3330; Corvallis, Feb: 2; 1914, 3300. Abies nobilis Lindl.—Mary’s Peak, Benton Co., Feb. 7, 1914, F. D. Bailey, 3337, Aug. 15, 1914, 3334; 33351 3383073330: On PotypopiAcEséE: II, III. ’ Athyrium cyclosorum Rupr.—Hoover, Linn Co., Aug. 19, 1914, H. P. Barss & G. B. Posey, 3032; Mary’s Peak, Benton Co., Aug. 15, 1914, 3033, 3034; Elk City, Lincoln ‘Co:, Aug: 20; 1o14tez0 ar Hood River Co., July 23, 1915, 303z; Grant’s Pass, Josephine Co., Sept. 5, 1916, J. R. Weir, 258; Yaquina, Lincoln Co., July 17, 1915, O35: There seems to be no good reason for separating U. Copelandu Sydow from U. Atkinsonwi. All gradations in the length of the beak of the urediniospores are found on the above collections. Most of the material has urediniospores with long beaks and some of the collec- tions show spores with both long and short beaks. Field observations as well as a study of morphological characters would support the view that the aecia commonly referred to Pert- dermium balsameum occurring on Abies grandis and A. nobilis in western Oregon are genetically connected. Fraser (Mycol. 5: 236. 1913) has cultured U. Atkinsonu by sowing aeciospores of Peridermium balsameum on Aspidium Thelepteris followed sparingly by the development of uredinia. 32. UREDINOPSIS PTERIDIS Dietel & Holway, Ber. Deuts. Bot. Ges. 132: 13312 $5895: Aecidium pseudo-balsameum Diet. & Holw. Erythea 7: 98. 1899. Peridermium pseudo-balsameum Arth. & Kern, Bull. Torrey Club 332430: 1906: On PrInAcEAE: I. Abies amabilis (Loud.) Forb.—Whitewater Creek, near Mt. Jeffer- son, Aug. 11, 1994, H.P.Barss & G. B. Posey, 3204: Abies grandis Lindl.—Corvallis, Aug. 1910, 3299, May 8, 1909, comm. Clarence D. Learn, April 29, 1914, F. D. Bailey, 3303; Ump- JACKSON: UREDINALES OF OREGON FAIS) qua Nat. Forest, near Diston, Lane Co., Oct. 27, 1909, Geo. G. Hedg- cock; Philomath, Jan. 6, 1914, 3298; Wren, Benton Co., Aug. 3, 1914, 3296; Ashland, Jackson Co., Sept. I0, 1914, 3297; Mary’s Peak, Benton Co., Feb. 7, 1914, G. H. Godfrey, 3307, Aug. 15, 1914, 3302; N. slope Mt. Hood, 4,000 ft., Aug. 9, 1914, 1616. On PotypopiAcEAE: II, III. Pteridium aquilinum pubescens Underw.—Corvallis, Sept. 1909, 1142, July, 1910, roS2, Aug. 1910, 1058, Oct. 6, 1914, 3109; Bonneville, Multnomah Co., Aug. II, 1910, 1076; Scappose, Columbia Co., iuly 25, 19o1t, 71067; North slope Mt. Hood, Aug. 9; 19014, 7677; Ashland, Jackson Co., Sept. 10, 1914, 1993; Portland, Aug. 24, 1915, E. Bartholomew (Barth. N. Am. Ured. 1485); Grant’s Pass, Josephine Co, Sept: 5, 1916, J. R» Weir, 250. This species in all its stages is very common in western Oregon and the association of the infected aecial and telial hosts is everywhere apparent. From field observations made by the writer and others it has been assumed that Peridermium pseudo-balsameum was the aecial stage of this species. Recently Weir and Hubert (Am. Jour. Bot. 4: 328-332. 1917) have conducted cultures showing the genetic con- nection of this species with aecia on Abies grandis. The authors evidently do not consider the aecia identical with P. pseudo-balsameum. The description which they give, however, agrees very well with the type of that species. According to the writer’s present interpretation, there are two closely related species of Peridermium on Abies grandis in western Oregon. One is to be referred to P. balsameum and is presumably genetically connected with Uredinopsis Copelandiu (cf. 31). The other is P. pseudo-balsameum and is genetically connected with the species under discussion. The walls of the aeciospores in P. balsameum are considerably thinner than those of P. pseudo-balsameum. In the former they are I-1.5 uw while in the latter they are 2—2.5 yw in thickness. PUCCINIACEAE 33. EARLEA SPECIOSA (Fr.) Arth. Résult Sci. Congr. Bot. Vienne 341. 1906. Aregma speciosa Fr. Syst. Myc. 3: 496. 1832. Phragmidium speciosum Cooke, Grevillea 3: 171. 1875. ON ROSACEAE: Rosa gymnocarpa Nutt.?—I, Austin, Grant Co., Aug. 1915, J. R. Weir, 188. 220 BROOKLYN BOTANIC GARDEN MEMOIRS 34. GYMNOSPORANGIUM BETHELI Kern, Bull.- Torrey Club 34: 459. 1907. Roestelia Bethelt Kern, Bull. Torrey Club 34: 461. 10907. On MALACEAE: I. Crataegus Douglas Lindl.—Joseph, Wallowa Co., Aug. 19, 1899, C. L. Shear (Ellis & Ev. Fungi Columb. 1480). ON JUNIPERACEAE: III. Juniperus occidentalis Hook.—Whitney, Baker Co., Aug. 1915, PRA Welt e570: Another specimen on Crataegus sp. indet. from eastern Oregon (ex herb. Ellis) bearing no date is in the Arthur herbarium and has been examined by the writer. The life history of this species was first demonstrated by Arthur (Jour. Myc. 14: 23. 1908) and later repeatedly confirmed. Telia are otherwise known only on Juniperus scopulorum from Colorado, Idaho and Montana. Aecia are known to occur only on Crataegus sp. in the Rocky Mt. region and in eastern Oregon and Washington. 35. GYMNOSPORANGIUM BLASDALEANUM (Dietel & Holway) Kern, BulliNeye Bot Gard.7 2487. Srorn. Aecidium Blasdaleanum Dietel & Holway, Erythea 3: 77. 18095. Gymnosporangium Libocedrt Kern, Bull. Torrey Club 35: 509. 1908. On MALaAceaeE: I. Amelanchier florida Lindl.—Eugene, Lane Co., July 11, 1914, G. B. Posey, 3270; “Albany, Linn Co:,; June 11, 1913, D. W23Rare baugh, 37717; Cottage ‘Grove, Lane Co., June 13, 1913, 3706; qiume 20, 1013, ©. &. Stewart, 391777; Webanon; Linn Co:, Aug. 2 ene EF. DW. Bailey, 3774," Grater Lake, Klamath Co., 7,000 ft.) Semis 1913, 0H. . Memecke, "Cr Pe D air; Jackson Co., July, 19145 hee Reimer, 7791; Between Albany and Lebanon, Linn Co., June 13, 1913, C. E. Roberts, 1788; Lost Prairie, Sept. 1891, M. Craig; Halsey, Linn Co., June 9, 1913, 3170; Corvallis, July 29, 1915, 3150; Ashland, Jackson Co., Sept. 10, 1914, 3047; N. W. Mt. Jefferson, Whitewater station, Aug. 17, 1914, Hi. BP. Barss & G. B. Posey, 3043, Augwmeasy 1916, HP. Basss: Crataegus Douglas Lind|.—Halsey, Linn Co., June 9, 1913, 3214; Albany, Linn Co., D. W. Brumbaugh, 3212; Cottage Grove, Lane €o,, May 21, 1013, 4700, June 14, 1913, 3200; Eugene, Laney@er May 8, 1913, 3173. Cydonia japonica Pers.—Eugene, Lane Co., June, 1914, G. H. Godfrey. Cydonia vulgaris L., Halsey—Linn Co., June 9, 1913, 3166; Irving, Lane Co., 1913,.Comm. S. J. Quigley, 18712; Creswell, Lane Co; —- JACKSON: UREDINALES OF OREGON Dg May 5, 1913, Comm. K. V. Miller, 7873; Eugene, Lane Co., Aug. 1912, 1084; Talent, Jackson Co., May 18, 1916, F. C. Reimer. Pyrus baccata Linn.—Lorane Valley, Lane Co., May, 1915, C. E. Stewart, 3387. Pyrus commums L.—Kerby, Josephine Co., June 1, 1899, Comm. E. F. Meissner, 1845; Brownsville, Linn Co., May 24, 1913, D. W. Brumbaugh, rgz1; Eugene, Lane Co., May 8, 1913, 3172, April 22 1915, 2620. Pyrus diversifoia Bong. (P. rivularis Dougl.)—Cottage Grove, izane'Co., June 13, 1913; 3275; 3211. Pyrus toensis (Wood) Bailey—Cottage Grove, Lane Co., June 13, 1913, 2654, Eugene, Lane Co., May 21, 1913, 3270. iPpyrus malus .— Eugene; Lane Co:, July 10; 1913; J. O.. Holt; bgo7, Cottage Grove, Lane Co:., May 23, 1913, 'C.. E. Stewart, ror, June 20, 1915, C. E. Stewart, 887. Sorbus aucuparia Linn.—Cottage Grove, Lane Co., June 13, 1913, aio, Mugene, Lane Co., June 1, 1914, G. H. Godfrey, 3222: Sorbus hybrida Linn.—Cottage Grove, Lane Co., May 21, 1913, © nm7o, \ime 13, 1913, 3107: ON JUNIPERACEAE: Libocedrus decurrens Yorr.—Eugene, Lane Co., Feb. 28, 1913, 3213, Feb. 21, 1914, F. D. Bailey, 1675, Mar. 20, 1914, 3070; Breiten- bush Hot Springs, Marion Co., Mar. 27, 1915, E. A. Hartley, 2621; Cottage Grove, Lane Co., Mar. 8, 1914, C. E. Stewart, 1588; Ashland, Waekcon.Co:, Sept: 10, 1914, 17530; Corvallis, Mar-.30, Tors, J. G. Corsaut, gor; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 185. This species is very common in its aecial stage on all the native members of the Malaceae as well as most of the cultivated fruits and ornamental plants belonging to this family which may occur in the range of the incense cedar. There is considerable evidence also that the disease is gradually spreading beyond the natural range of the telial host. Observations made at Corvallis support this view. Previous to 1915 no specimens of this species had been collected in Benton County though careful search had been made many times. The incense cedar does not occur naturally in that locality but is frequently planted for ornament. There are several fine examples on the campus at the Oregon Agr. College. In 1915 a very sparing in- fection of the aecial stage was found in the vicinity on native hosts only, and the cedar trees on the campus were found to be sparingly infected. Reports of the occurrence on quince have come from Salem, far north of the natural range of the incense cedar. The writer believes that the disease will gradually spread throughout the Wil- ’ 222 BROOKLYN BOTANIC GARDEN MEMOIRS lamette Valley on the incense cedars planted for ornament and, in certain cases, will become a serious menace to cultivated pears and quinces. The life history of the species was first worked out by Arthur (Mycol. 1: 252. 1909; 4: 57. 1912). He succeeded in showing that aecia occurred on Crataegus and Amelanchier. ‘The telial material used for the cultures was collected at Eugene, Oregon, by Prof. A. R. Sweetzer. The writer has also studied this species in some detail (Phytopath. 4: 261-269. 1914; Ore. Expt. Sta. Biennial Crop Pest Rep. II: 204-212. 1915) and has reported the results of culture work and field observations. Similar work is also briefly reported by O’Gara (Science N. S. 39: 620-621. 1914). The previous records of the occurrence of this species on Malus floribundus Siebold and Sorbus sambucifolia Roem. made by the writer (1. c.) should be corrected to read Pyrus toensis and S. aucuparia respectively. For the correct determination of these hosts the writer is indebted to Prof. W. W. Eggleston. This species has since been successfully cultured in the greenhouse on Pyrus sinensis by Prof. H. P. Barss. The writer, using aecial material on quince, the result of infection experiments made in 1914, has obtained sparing infection on Libocedrus resulting in telia which matured in February 1915. The trees were kept in the greenhouse at the Oregon Agr. College. 36. GYMNOSPORANGIUM HARKNESSIANUM (Ell. & Ev.) Kern, Bull. Ne Y Bot Gandy 440- “Tort: Roestelia Harknessiana Ell. & Ev. Kern, Bull. Torrey Club 34: A462: 1907. On MALACEAE: I. Amelanchter alnifolia Nutt.—Redmond, Crook Co., July 2, 1914, 1303, Fort Rock, Lake €o,, Oct. 10, 1915, Wendover, 3375. This very interesting species has otherwise been reported only from northern California. The telia form is unknown. In the col- lection made by the writer listed above, the aecia occurred only on fruits and twigs. There was every evidence that this fungus is perennial. Some of the specimens show fresh aecia on branches having four annual rings, surrounding or extending from cankered areas bearing evidence of old aecial cups. There is slight hypertrophy. The branches are frequently girdled and killed. 37. GYMNOSPORANGIUM JUNIPERINUM (L.) Mart. Fl. Crypt. Erlang. Ba2, LSi7: Tremella juntperina L. Sp. Pl. 1157. 1753. JACKSON: UREDINALES OF OREGON 223 On MaAtacese_E: I. Sorbus occidentalis (S. Wats.) Greene—North slope Mt. Jefferson, along trail to Hanging Valley, Aug. 15, 1914, H. P. Barss & G. B. Posey, 7395; Columbia Highway, Multnomah Co., Aug. 19, 1916, J. R. Weir, 279. On JUNIPERACEAE: ITI. Juniperus sibirica Burg.—North slope Mt. Jefferson, Aug. 26, Eto. il. P..Barss, 3300. The genetic connection of the forms of this alpine species has been abundantly demonstrated by European investigators, first by Hartig (Lerb. Baum-Kr. 133. 1882), and later by many others. Arthur in r911 (Mycol. 4: 57. 1912), using telial material from J. stbirica collected in Colorado, succeeded in obtaining infection result- ing in pycnia only on Sorbus americana. The species is known in America only from the Rocky and Cascade Mountains of the United States and Canada. 38. GYMNOSPORANGIUM JUVENESCENS Kern, Bull. N. Y. Bot. Gard. V=445. IOUL. On Mataceae: I. Amelanchier sp.—Hurricane Creek, Wallowa Co., July 24, 1897, E. P. Sheldon, 8622. ON JUNIPERACEAE: III. Juniperus scopulorum Sarg.—White Pine, Baker Co., Aug. 1915, J. R. Weir, 1609. This species causes witches’ brooms on the telial host somewhat similar to the eastern G. nidus-avis Thax. Arthur has repeatedly cultured it, showing that the aecia occur on Amelanchier and Sorbus (Jour. Myc. 13: 203. 1907; 14: 18. 1908; Mycol. 1: 239. 1909; 4: 195. I9I12). 39. GYMNOSPORANGIUM KERNIANUM Bethel, Mycologia 3: 157. I9II. ON JUNIPERACESE: III. Juniperus occidentalis Hook. ia, Crook Co., July 2, 1914, 1392, May 15, 1915, 3390. The above specimens are somewhat doubtfully referred to this species. The witches’ brooms are large and open, sometimes reaching 2-3 feet in diameter. The teliospores are somewhat more tapering at the apex than is typical for the species and average shorter and somewhat narrower, 19-22 by 45-65 u. The only aecia collected in the vicinity are properly referred to G. Harknessianum. There was no very direct field evidence, and unless the above collections repre- sent an undescribed form there is little possibility that the two can be genetically connected. Arthur (Mycol. 4:62. 1912) has cultured 16 224 BROOKLYN BOTANIC GARDEN MEMOIRS G. Kernianum on Amelanchier but obtained the development of pycnia only. Field observations and collections by Bethel in Colorado indicate strongly that the aecial stage occurs on Amelanchier. The Roestelia, however, is quite different from R. Harknessiana. 40. GYMNOSPORANGIUM KOREAENSE (P. Henn.) Jackson, Jour. Agr. Research 5: 1006. I916. Roestelia koreaensis P. Henn. in Warburg, Monsunia 1: 5. 1899. Gymnosporangium asiaticum Miyabe, Bot. Mag. Tokyo 17: 34. 1903. (Hyponym.) Gymnosporangium Haraeanum Syd. Ann. Myc. 10: 405. IgI12. Gymnosporangium chinense Long, Jour. Agr. Research I: 353. 1914. ON MALACEAE: I. Pyrus sinensis Lind|.—Portland (Orient), June 11, 1914, 2066. ON JUNIPERACEAE: III. Juniperus chinensis L.—Portland (Orient), March 29, 1915, 2668. This species has been shown by the writer (Il. c.) to have been established at Portland (Orient), Oregon, on trees imported from Japan. It has been cultured on Pyrus sinensis and Cydonia vulgaris. 41. GYMNOSPORANGIUM NELSONI Arth. Bull. Torrey Club 28: 665. I9Ol. ON JUNIPERACEAE: III. Juniperus occidentalis Nutt.—Austin, Grant Co., April, 1916, J. R. Weir, 257. Juniperus scopulorum Sarg.—Whitman Nat. Forest, Aug. 1915, J. R. Weir, 766. This species causes conspicuous galls on the branches of Juniperus. The aecial stage has been collected on Amelanchier, Cydonia, Pera- phyllum, Pyrus and Sorbus. Arthur (Mycol. 4: 61. 1912; 7: 78. 1915) has conducted cul- tures, using telial material from Colorado. Weir & Hubert (Phyto- path. 7: 109. 1917) have recently confirmed these results, using material collected in Montana, on J. communis and J. scopulorum. 42. GYMNOSPORANGIUM NOOTKATENSIS (Trel.) Arth. Am. Jour. Bot. 35 44. O16. Uredo nootkatensis Trelease, Alaska Harr. Exped. 5: 36. 1904. Uredo Chamaecyparidis-nutkaensis Tubeuf, Nat. Zeits. Forst.- Landw.:2: "ol. 19r4: ON JUNIPERACEAE: Chamaecyparis nootkatensis (Lamb.) Spach—North slope Mt. Jefferson, trail to Hanging Valley, Aug. 15, 1914, H. P. Barss & G. B. JACKSON: UREDINALES OF OREGON 225 Posey, 1394; Whitewater Ranger station, Aug. 28, 1916, H. P. Barss; Foot of Mt. Jefferson, Aug. 28, 1916, H. P. Barss. The material collected by Barss and Posey in 1914 contained teliospores in the uredinia and forms the basis of the transfer of the very interesting and much discussed Uredo nootkatensis to Gymno- sporangium. A full account of the history of this species has been given by Arthur (I. c.). In the collections of 1916 made in the same locality by Prof. Barss, teliospores were found in great abundance with the uredinia, and in many sori predominated. The uredinio- spores were germinated in this laboratory and the germ tubes found to develop in the usual way for urediniospores. 43. GYMNOSPORANGIUM SorBI (Arth.) Kern, Bull. N. Y. Bot. Gard. 7: A338; \1GED: Aecidium Sorbi Arth. Bull. Torrey Club 33: 521. 1906. On MALACcEAE: I. Sorbus occidentalis (S. Wats.) Greene—Whitewater Ranger station, Mt. Jefferson, Aug. 28, 1916, H. P. Barss. There is little doubt that the suggestion of the genetic relationship of this species with Gymnosporangium nootkatensis (cf. 42) originally made by Kern (Science 31: 833. 1910) and later re-affirmed by Arthur (Am. Jour. Bot. 3: 43-44. 1916) will prove to be correct. The above collection extends the range of the aecia to correspond exactly with the range of the known collections of uredinia and is the most southern record. It seems best, however, for the purpose of this list to retain the above name till actual cultures confirming the prediction have been made. 44. GYMNOSPORANGIUM TUBULATUM Kern, Bull. N. Y. Bot. Gard. 7: SE clOLT. Roestelia tubulata Kern; in M. E. Jones, Bull. Univ. Mont. 61: 64. 1910. On MALAcgEAE: I. Crataegus Douglasii Lindl.—Minam River, Wallowa Co., Oct. 5, 1897, E. P. Sheldon, 9061; Wallowa Nat. Forest, Sept. 28, 1910, G. G. Hedgcock, 1944. The above collections were found in the Arthur herbarium at the Purdue University Experiment Station. The specimens show chiefly foliage infection, though the first-mentioned collection also includes infected fruit. Weir (Phytopath. 5: 218. 1915) has recently demonstrated by cultures that the telia, which were previously unknown, occur on the twigs of Juniperus scopulorum forming irregularly lobed galls. Telia 226 BROOKLYN BOTANIC GARDEN MEMOIRS have been collected only in Idaho and western Montana. Weir and Hubert in 1916 (Phytopath. 7: 109. 1917) have confirmed the above results. : 45. KUNKELIA NITENS (Schw.) Arth. Bot. Gaz. 63: 504. 1917. Aecidium nitens Schw. Schrift. Nat. Ges. Leipzig 1: 69. 1822. ON ROSACEAE: Rubus nigrobaccus Bailey—Freewater, Umatilla Co., June 27, TOI3 4h. De Baileya Tmo. Rubus vitifolius Cham. & Schlecht. (cult. loganberry)—LaGrand, Union Co: July.20; 1914. C. Cate’ 1S5 im. Kunkel’s results (Bull. Torrey Club 40: 361. 10913; 43: 559: 1916; Amer: Jour. Bot. 1: 37. 1914) indicate that two rustcnom Rubus, both commonly referred to Gymnocontia interstitialis or Caeoma nitens, occur in North America, one a short-cycled form having the morphology of a Caeoma, the other a brachy-form with caeomoid aecia and telia of the type of Puccinia (P. Peckiana Howe). Arthur (I. c.) has recently based the genus Kunkelia on the short-cycled form. The inclusion of the Oregon collections under Kunkelia follows the disposition made of them by Arthur. 46. NYSSOPSORA ECHINATA (Lev.) Arth. Result. Sci. Congr. Bot. Vienne 342. 1906. Triphragmium echinatum Lev. Ann. Sci. Nat. III. 9: 247. 1848. ON UMBELLIFERAE: Ligusticum Cusicki Coult. & Rose—Steins Mts., Harney Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 340). Ligusticum purpureum Coult. & Rose—North slope Mt. Jefferson, Nuss, 1Olde ld. PsBarssicc G. Bs Posey, 2520: 47. PHRAGMIDIUM DISCIFLORUM (Tode) J. F. James, Contr. U. S. INat. Herbs3% 2765 1805; Ascophora disciflora Tode, Fungi Meckl. 1: 16. 1790. ON ROSACEAE: Rosa sp. cult:—Empire, Coos Co., Oct..2, 1911, comm. JeR Brown, 3754; Portland, May 10, 1914, comm. W. C. Dietz, 3756; Eugene, Lane Co., June 1, 1914, G. H. Godfrey, 3147; Sutherlin, Douglass Co., March 9, 1915, comm. Gladys Franz, 2577. 48. PHRAGMIDIUM IMITANS Arth. N. Am. Flora 7: 165. 1912. ON ROSACEAE: Rubus leucodermis Dougl., Philomath, May 10, 1914, 7830. Rubus neglectus Pk., Ore. Agr. Coll. Pathologium, Corvallis, July 80) TOT5)4027- Rubus strigosus Michx.—Stream banks, Eastern Oregon, 4,000—- 5,000 ft. elev., July, 1897, W. C. Cusick, 1729. JACKSON: UREDINALES OF OREGON 227 49. PHRAGMIDIUM IVESIAE Sydow, Ann. Myc. 1: 329. 1903. Phragmidium affine Sydow, Ann. Myc. 2: 29. 1904. On ROSACEAE: Potentilla blaschkeana Turcz.—Philomath, June 20, 1910, 1503; Austin, Grant Co., June, 1913, J. R. Weir, 147; Baker Co., June, 1913, leek Weir, 77; Sumpter, Baker €o., June; 1913, J. R. Weir; 3; Hil- gard, Union Co., July 10, 1914, 1534. Potentilla glomerata A. Nels.—Andrews, Harney Co., Aug. 1901, Griffiths & Morris (Griffiths, West Am. Fungi 5770). Potentilla gracilis Dougl.—Corvallis, June 20, 1909, E. R. Lake, 1499, July 29, 1914, 1477; Wren, Benton Co., June 26, 1914, 1323. Potentilla sp.—Corvallis, June, 1910, zrz0, 3149, Aug. 1911, F. D. Bailey, 2071. 50. PHRAGMIDIUM JONEsII Dietel, Hedwigia 44: 128. 1905. ON ROSACEAE: Ivesta Batleyt S. Wats.—Steins Mts., Harney Co., July 27, 1898, W. C. Cusick, Phan. Herb. 1967. (From specimen in herb. Field Museum 108727.) 51. PHRAGMIDIUM MONTIVAGUM Arth. Torreya 9: 24. 1909. ON ROSACEAE: Rosa gymnocarpa Nutt.—North slope Mt. Hood, Aug. 9, 1914, 1478; Bank of Minam River, Union Co., alt. 5,100 ft., Oct. 4, 1897, E. P. Sheldon, 9053. Rosa pisocarpa Gray ?—Hilgard, Union Co., July 10, 1914, 1537. Rosa sp.—Trail Creek Cafion, Wallowa Co., May 18, 1897, E. P. Sheldon, 8073; Corvallis, May 1, 1914, 1466, April 25, 1915, G. B. Posey & C. M. Schearer, 3753, April 28, 1915, 3151; North slope Mt. Hood, Aug. 7, 1914, 7619; Mouth of Salmonberry River, Tillamook Co., July 17, 1915, G. VanGundia, 3089. 52. PHRAGMIDIUM OCCIDENTALE Arth.; Earle, in Greene, Pl. Baker. 2:3, TOOL: On ROSACEAE: Rubus parviflorus Nutt.—Wallowa Lake, Wallowa Co., Aug. 1899, C. L. Shear, 952 (Griffiths, W. Am. Fungi 329); Jackson Co., July 9, 1903, E. B. Copeland (Sydow, Ured. 1788); Mt. Hood, Aug. 31, 1901, E. W. D. Holway, Aug. 7, 1914, 1636; Glen Brook, Benton Co., Aug. 1909, 1719; Trail to Hanging Valley, Mt. Jefferson, H. P. Barss & G. B. Posey, 1785; Mary’s Peak, Benton Co., Aug. 15, 1914, 1285; Elk City, Lincoln Co., Aug. 20, 1914, 1626; Dothan, Douglass Co., Sept. 8, 1914, 7930; Corvallis, May 4, 1915, 3059; Unity, Baker Co., Aue. TOUS. ). ke. Weir. 242, Austm, Grantu€o:, Ate, sora. )ael. Weir, 238. 228 BROOKLYN BOTANIC GARDEN MEMOIRS 53. PHRAGMIDIUM POTENTILLAE (Pers.) P. Karst. Bidr. Finl. Nat. Folk 31: 49. 18709. Puccinia Potentillae Pers. Syn. Fung. 229. 1801. On ROSACEAE: Potentilla aracnoides Lehm.—Austin, Grant Co., Aug. 1915, J. R. Weir, 161. Potentilla Hippiana Lehm.—Austin, Grant Co., Aug. 1915, J. R. Weir, 158. 54. PHRAGMIDIUM ROSAE-ACICULARIS Liro, Bidr. Finl. Nat. Folk. 65: 428. 1908. ON ROSACEAE: Rosa nutkana Presl.—Bridal Veil, Multnomah Co., May 18, 1915, 3348; Edge of woods on Minam River, Union Co., Aug. 11, 1897, E. P. Sheldon, 5667. Rosa sp.—Corvallis, July 28, 1914, 3146. 55. PHRAGMIDIUM ROSAE-CALIFORNICAE Dietel, Hedwigia 44: 125. 1905. ON ROSACEAE: Rosa gymnocarpa Nutt.—Corvallis, July 29, 1914, H. P. Barss, 1469; Mary's Peak; Benton Co., Aug: 15, 1014, 7512, 1510, agen Ashland, Jackson Co., Sept. 10, 1914, 3084. Rosa nutkana Presl.—Corvallis, July 29, 1914, 1473; Portland, Aug. 23, 1915, E. Bartholomew, 5950 (Barth. N. Amer. Ured. 7626); Hood River, Aug. 26, 1915, E. Bartholomew, 5973 (Barth. Fungi Columb: 4632); > Austin, Grant Co:; Aug: 1915, J. R. Weimmagges Bend, Crook Co., Sept. 11, 1916, J. R. Weir? 202. Rosa pisocarpa Gray—Corvallis, April 5, 1914, 1523. Rosa sp.—Bonneville, Multnomah Co., Aug. II, I910, 1072, Cogvallis, Aug: 10, Torts 4752, Spring. 1ro14, H.C. Gilbertiemes Eugene, Lane Co., July 11, 1914, G. B. Posey, 1467; Ashland, Jackson Co., Sept. 30, 1914, 3350; Whitewater Creek, near Mt. Jefferson, Aug. Pi pLOLA pls be wbarss.cc (Gab. eosey. 3302. 56. PILEOLARIA TOXICODENDRI (Berk. & Rav.) Arth. N. Am. Flora 7; 147. 1907. Suh Uromyces Toxicodendri Berk. & Rav. Grevillea 3:56. 1874. Pileolaria brevipes Berk: & Rav. Grevillea 3: 58. 1874. On ANACARDIACEAE: Rhus diversiloba T. & G.—Corvallis, April 29, 1914, F. D. Bailey, 1831; Grant’s Pass, Josephine Co., Sept. 5, 1916, J. R. Weir, 256; Jim Creek, Wallowa Co., June 14, 1897, E. P. Sheldon, 6270. 57. POLYTHELIS FUSCA (Pers.) Arth. Résult. Sci. Cong. Bot. Vienne 341. 1906. Aecidium fuscum Pers., in Gmel. Syst. Nat. 2: 1473. 1791. JACKSON: UREDINALES OF OREGON 229 ON RANUNCULACEAE: Anemone oregana A. Gray—Mary’s Peak, Benton On May 23, IQI5, 3030. Anemone quinquefolia L.?—North slope Mt. Hood, Aug. 9, 1914, 1621. 58. Puccinia ABSINTHII (Hedw. f.) DC. Fl. Fr. 6: 56. 1815. Uredo (Puccinia) Artemisit Hedw. f.; DC. in Lam. Encycl. Meth. Bot. 8: 245. 1808. Puccinia similis E. & E. Bull. Torrey Club 25: 508. 1808. On CARDUACEAE: Artemisia dracunculoides Pursh—Sherman, Sherman Co., July 1, 1914, 2671, May 16, 1915, 2672. Artemisia frigida Willd.—Bend, Creole Cor Sept tiieeT OG) iia kk: Weir, 272. Artemisia ludoviciana Nutt.—Eastern Oregon, Aug. 1914, H. F. Wilson, 3321; Eugene, Lane Co., July 20, 1914, F. D. Bailey, 1504; Portland, Aug. 21, 1915, E. Bartholomew, 5939 (Barth. Fungi Columb. 5048); Sumpter, Baker Co., June, 1913, J. R. Weir,.9z; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 249; Hood River Co., July 2IETOTS, 3158: Artemisia rigida (Nutt.) A. Gray—Eastern Oregon, 3,500 ft. altitude, Sept. 1900, W. C. Cusick, 2504; Lost Valley, Wheeler Co., Sept. 9, 1894, J. B. Lieburg, 858. Artemisia tridentata Nutt.—Redmond, Crook Co., Sep D5. O1s; Kirk Whited, 31787, July 2, 1914, 2555; Sherman, Sherman Co., July 1, 1914, 1938; Umatilla, Umatilla Co., July 11, 1914, 1293, May ER tons, 3036,) Park, Union :Co:,, Oct, 9; 1897, E~ Ps Sheldon; 9773: This species, presumably a brachy-form, though no pycnia have yet been observed, is not to be confused with any other species on this host genus. The only other species recognized in North America is P. conferta (cf. 90) which is a micro-form. 59. Puccinia abundans (Pk.) comb. nov. Aecidium abundans Pk. Bot. Gaz. 3: 34. 1878. Puccinia Crandallii Pam. & Hume, Proc. Dav. Acad. Sci. 7: 250. 1899. Puccinia Kreageri Ricker, Jour. Myc. Ir: 114. 1905. ON CAPRIFOLIACEAE: I. Symphoricarpos albus (1..) Blake—Head of Applegate Creek, Jackson Co., July 29, 1913, Bs. P. Meinecke, Cr D (2) 5; Bridal Veil, Multnomah Co., May 18, 1915, 3054, Mary’s Peak, Benton Co., May 21, 1915, 3036; Hilgard, Union Co., July 9, 1914, 2546; Hood River, May 14, 1914, 2566, July 21, 1915, 3063; Springbrook, Yamhill 230 BROOKLYN BOTANIC GARDEN MEMOIRS Co., May, 14, 1914, F. D. Bailey, 2567; Philomath, April 26, 1914, 2572; Corvallis, April 28, 1915, 2612; Grant’s Pass, Josephine Co., Sept. 2, 1916, J. Re Wein, Zor On Poaceae: II and III. Festuca confinis Vasey (Poa Kingit S. Wats.)—Steins Mts., Harney Co:, July 2; 1896, J. B: Weiberg, 2025. Festuca idahoensis Elmer—Hilgard, Union Co., July 10, 1914, 1358, 1362; Redmond, Crook Co., July 2, 1914, 1424, 1430 Festuca rubra L.—Hilgard, Union Co., July 10, 1914, oe Mary’ s Peak, Benton Co., Aug. 15, 1914, 1571, 1573; Newport, Lincoln Co., July 18.1915, 3207. Festuca subulata Trin.—Ashland, Jackson Co., Sept. 10, 1914, 1563. The connection between this common western form on Festuca with Aecidium abundans was shown by Arthur in 1910 (Mycologia 4: 27. 1912). In three trials, using telial material on F. confinis, col- lected in Colorado and Utah, infection resulting in pycnia and aecia on Symphoricarpos racemosus was obtained. 60. PucciInta ACETOSAE (Schum.) Koern. Hedwigia 15: 184. 1876. Uredo Acetosae Schum. Enum. PI. Saell. 2: 231. 1803. ON POLYGONACEAE: Rumex acetosella L.—Maples Station, Tillamook Co., Sept. 15, 1915, F. D. Bailey, 3102; Corvallis, Oct. 19, 1915, G. B. Posey, 3000. This species has been recorded previously from North America only from Florida on R. hastatulus (Holway, North Am. Ured. 1: 35. 1906). Specimens on that host referred to this species are in the Arthur herbarium also from S. Carolina and Massachusetts and on R. acetosella from Massachusetts, Florida, New York and Indiana. All the specimens bear uredinia only. It is possible that some or all of the material should be referred to Uromyces Acetosae Schroet., as the two species are indistinguishable in the uredinial stage. 61. PUCCINIA AMBIGUA (Alb. & Schw.) Lagerh., in Bubak, Sitz. Ver. Bohm. Ges. Wiss. 1898, 28: 14. 1808. Aecidium Galit ambiguum Alb. & Schw. Consp. Fung. 116. 1805. Puccinia difformis Kunze, Myc. Hefte 1: 71. 1817. Allodus ambigua Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. On RUBIACEAE: Galium aparine L.—Wren, Benton Co., June 26, 1914, 1330; Ashland, Jackson Co., Sept. 10, 1914, 3090. This species possesses aecia and telia only in the life cycle. It has been studied by Bubak (1. c.) who found that primary aecia were followed by secondary aecia. Later Trebaux (Flora 81: 394-404. 1895) repeated this observation and conducted culture work con- JACKSON: UREDINALES OF OREGON 2351 firming Bubak’s contention. This species should not be confused with P. punctata Lk. (cf. 156) which occurs on the same host from this region. 62. PUCCINIA ANGELICAE (Schum.) Fckl. Symb. Myc. 52. 1869. (Not P. Angelicae E.& E. 1884.) Uredo Angelicae Schum. Enum. PI. Saell. 2: 233. 1803. Puccima Archangelicae Blytt, Christiania Vid. Selsk. Forhandl. Nos.6:.51. . 1896: Bullaria Angelicae Arth. Résult Sci. Congr. Bot. Vienne 346. 1906. On UMBELLIFERAE: Angelica genufleca Nutt.—Woodburn, Clackamas Co., Sept. 1885, Thomas Howell. Angelica Lyalliat Wats.?—Larch Mt., Multnomah Co., Aug. Io, 1910, 2613. This species is evidently rare in North America having been reported otherwise only from a single collection from Washington on A. genuflexa and one from New York on A. atropurpurea. It is a brachy-form though pycnia have not been seen in American collec- tions. This species has smooth teliospores and is easily separable from Puccinia Ellisit (cf. 98) on the same hosts from our region, which has verrucose spores. 63. PUCCINIA ANOMALA Rost. Thiimen, Flora 1877: 92. 1877. Puccinia straminis simplex WKoern. Land. u. Forstw. Zeit. no. 50. 1865. Puccinia Hordes Otth. Mitt. Nat. Ges. Bern. 1870: 114. 1871. (Not P. Hordei Fckl. 1860.) Puccinia simplex Erikss. & Henn. Getreideroste 238. 1896. (Not P. simplex Peck. 1881.) Aecidium Ornithogalum Bubak, Ann. Myc. 3: 223. 1905. On Poaceae: II, III. Hordeum montanense Schribn.—Corvallis, July 26, 1914, 1414. Hordeum murinum L.—Corvallis, July 8, 1914, G. B. Posey, 1354. Hordeum nodosum L.—Corvallis, July 26, 1914, 3257. Hordeum vulgare L.—Corvallis, July 6, 1914, 1683, Aug. 13, 1914, 1091, 1708. This, the leaf rust of barley, is evidently very common in Oregon, much more so than the collections listed above would indicate. It is evidently not abundant in America except on the Pacific coast. In the Arthur herbarium, specimens on wild barleys are represented only from Oregon, California and Utah. On the cultivated barley speci- mens are at hand only from California, Iowa and Wisconsin. It is D352 BROOKLYN BOTANIC GARDEN MEMOIRS evidently spreading into the eastern United States as the writer col- lected it in August, 1916, at Ithaca and Savanna, New York. Tranzschel has shown that this rust in Russia has its aecia on Ornithogalum umbellatum and O. narbonense (Mycol. Cent. 4: 70. 1914). 64. PuccINIA ANTIRRHINI Diet. & Holw. Hedwigia 36: 298. 1897. ON SCROPHULARIACEAE: Antirrhinum majus L.—Portland, Aug. I909, comm. Charles Ladd, roSo, Aug. 28, 1914, comm: P.-C. Sehmeir, 79r4; Salem Marion Co., July, 1911, comm. Mrs. Lord, 1127; Corvallis, June 26, LOI2 O55). AUS LOD. TO25e The snapdragon rust is very common in Oregon both in gardens and in the greenhouse. For a long time it was known to occur only in California. As snapdragons came to be used more commonly in greenhouse culture the rust has gradually spread through the distri- bution of cuttings, till at the present time it is known to occur in most of the central and eastern states. 65. PUCCINTA/ARNIGADIS Pk: Bot. "Gaz. 0: 227. 1681. On CARDUACEAE: Arnica cordifolia Hook.—Near Aneroid Lake, July 1, 1899, II, ER: Lake yi707.. A very distinct species having minutely verrucose teliospores, not thickened at the apex, and is known only from the Rocky moun- tain and Pacific coast regions. 66. PUCCINIA ASARINA Kunze, in Kunze & Schmidt, Myk. 1: 7o. 1817. Puccinia Asari Link in Willd. Sp. Pl. 67: 68. 1825. Dicaeoma asarinum Kuntze, Rev. Gen. Pl. 3: 467. 18098. On ARISTOLOCHIACEAE: Asarum caudatum Lind|.—Portland, Aug. 30, 1915, E. Bartholo- mew, 5977 (Barth. Fungi Columb. 4840). This micro-form is known from North America on the above host, otherwise only from California, Idaho and Washington. 67. PUCCINIA ASPERIFOLII (Pers.) Wettst. Verh. Zool.-Bot. Ges. Wien 35: 541. “1885: Aecidium asperifolu Pers. Obs. Myc. 1: 97. 1796. Puccinia dispersa Erikss. Zeitsch. f. Pflanzenkr. 4: 257. 1894. On POACEAE: Secale cereale L.—Hood River, June 19, 1914, 1402; Corvallis, July 28, 1914, 1682; Bend, Crook Co., Sept. 11, 1916, J. R. Weir, 243. The leaf rust of rye is evidently common throughout the state. JACKSON: UREDINALES OF OREGON 233 This species has its aecia on species of Anchusa and Lycopsis in Europe as was first shown by De Bary (Monatsber. k. Akad. d. Wiss. Berlin 211. 1866). No aecia referable to this species have been found in America, but Arthur (Mycologia 1: 236. 1909) obtained the develop- ment of pycnia on Lycopsis arvensis secured from Europe, following exposure to germinating telia on rye collected in Indiana. This culture indicates that the European and American rusts are identical. 68. PUCCINIA ASPERIOR E. & E. Bull. Washb. Lab. 1: 3. 1884. Puccinia oregonensis Earle, Bull. N. Y. Bot. Gard. 2: 349. 1902. Allodus oregonensis Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. Allodus aspertor Orton, Mem. N. Y. Bot. Gard. 6: 193. 1916. On UMBELLIFERAE: Leptotaenia dissecta Nutt.—Corvallis, June and July, 1898, M. Craig, April 14, 1899, M. Craig, June, 1910, 2614, March 24, 1914, G. B. Posey,2665; Mary’s River near Wren, Benton Co., June 5, 1915, 2673. This is one of the most common and conspicuous of rusts, prob- ably widely distributed throughout western Oregon. The type of P. oregonensis, the second collection listed, was described as on Sanicula bipinnata, which is clearly an error for the above host as was first pointed out by Holway (N. Am. Ured. 14: 93. 1913). 69. PucciniA AsTERUM (Schw.) Kern, Mycologia 9: 224. I917. Aecidium asterum Schw. Schrift. Nat. Ges. Leipzig 1: 67. 1822. Aecidium Solidaginis Schw. Schrift. Nat. Ges. Leipzig 1: 68. 1822. Caeoma asteratum Link in Willd. Sp. Pl. 6: 51. 1825. Caeoma (Aecidium) erigeronatum Schw. Trans. Am. Phil. Soc. II. A292), “1832. Puccinia extensicola Plowr. Brit. Ured. Ustil. 181. 1889. Puccinia Caricis-Erigerontis Arth. Jour. Myc. 8: 53. 1902. Puccinia Caricis-Asteris Arth. Jour. Myc. 8: 54. 1902. Puccinia Caricis-Solidaginis Arth. Bot. Gaz. 35: 21. 1903. On CARDUACEASE: I. Aster sp.—Philomath, May to, 1914, 1309, 3066; Hilgard, Union Co., July 10, 1914, 3056; Corvallis, May 9, 1914, 3367, 3368, April 31, 1915, W. E. Lawrence, 3048; Hood River, May 14, 1914, 3022; Sumpter, Baker Co., June 1913, J. R. Weir, 85. Erigeron speciosus DC.—Near Whitewater ranger station, Mt. Jefferson, Aug. 16, 1914, H. P. Barss & G. B. Posey, 3292. Euthamia occidentalis Nutt.—Mary’s River, Benton Co., June 1898, M. Craig. 234 BROOKLYN BOTANIC GARDEN’ MEMOIRS ON CYPERACEAE: IJ and III. Carex athrostachya Olney—Philomath, May 10, 1914, 3286, Cor- vallis, June 29, 1914, G. B. Posey, 1333. Carex canescens L.—Hood River, Aug. 5, 1914, 3005. Carex Deweyana Schw.—Glendale, Douglass Co., Aug. 17, 1914, 1409; Elk City, Lincoln Co., Aug. 20, 1914, 1381, 1383; Philomath, May 10, 1914, 3284; Trail to Sulphur Springs, Benton Co., Nov. 3, 1912, 3288; Corvallis, Apr. 29, 1914, F. D. Bailey, 3283, May 19, 1913, 1193. Carex festiva Dewey—Newport, Lincoln Co., July 18, 1915, 3279. Carex Goodenowu J. Gay (C. vulgaris E. Fr.)—Hood River, June 20, 1914, 1405. Carex phyllomanica W. Boot?—Grant’s Pass, Josephine Co., Sept. A eLOLOn UA ak- VN eligneeOr Carex praegracilis Boot (C. marcida Boot)—Corvallis, May 9, LOM 32072 Carex scoparia Schk.—Corvallis, June 24, 1914, F. D. Bailey, 1380. Carex stipata Muhl.—Sherwood, Washington Co., July Io, 1914, F. D. Bailey, 17355; Hood River, May 14, 1914, 3014; Hubbard, Clackamas Co., May 27, 1914, 3011; Portland, May Io, 1914, F. D. Bailey, 3003; Corvallis, Aug. 10, 1910, 1789, May 9, 1914, 3285, July 29, 1915, 3281; Eddyville, Lincoln Co., Aug. 8, 1915, Hoerner, 3350. Carex straminea Willd.—Hood River, July 24, 1915, 3280. Carex subfusca W. Boot.—Corvallis, July 29, 1914, 1444. Carex sp.—Ashland, Jackson Co., Sept. 10, 1914, 3008; Corvallis, July 29, 1914, 1442; Cottage Grove, Lane Co., July 14, 1914, 1350; Philomath, Jan. 6, 1914, r78o. In 1901 Arthur (Qiour: Myc: 8:54. 1902) first bégan culture work showing that aecia which occur commonly on Aster, Solidago and related hosts are genetically connected with uredinia and telia on various species of Carex. The culture work conducted by Arthur is extensive and extends over a period of years from 1901-1914. In this series of culture work aecia have been produced on various species of Aster, Solidago, Erigeron, Leptilon and Euthamia, using telia from Dulichium and from many species of Carex from various parts of North America our. Miyec. 854° 1902; Ir: 58.> 19055 "12eaSs 906; 14) 13. 1908> JBot-sGazs 35) 15, 21; 1903; Mycolimeeaae 1900} 22224. “1010; “Ales, 16.) oi2; 72770, 81. TOTS) ey ase 1911 (Mycol. 4: 181. 1912) confirms Arthur’s results in part by successfully infecting Aster acuminatus using telial material from Carex trisperma L. JACKSON: UREDINALES OF OREGON 235 This study has also shown that the species as here considered is a composite form made up of several distinct physiological races. The species is separable from all other American species of Puccinia on Carex by the presence of two pores in the upper part of the rather small (12-19 by 16-23 w) urediniospores and the medium-sized (12—20 by 35-50 u) teliospores. The aecia of this species should not be confused with those of P. stipae (cf. 166), which occur on the same generic hosts and other Carduaceae in the west. In P. Asterum the aecia are cupulate, the peridium conspicuous, the spores globoid, nearly colorless and small (11-15 by 13-18 uw). InP. stipae the aecia occur on hypertrophied areas, the individual cups are gall-like and open by a central aperture, the peridium is evanescent, the spores are cinnamon brown, globoid, large (21-26 by 22-29 nw). The aecia of the latter species have not been found in Oregon, but doubtless occur abundantly in the eastern part of the state. 70. PucctntA ASTERIS Duby, Bot. Gall. 2: 888. 1830. ON CARDUACEAE: Aster conspicuus Lindl.—Hilgard, Union Co., July 10, 1914, 7538; Crater Lake, Klamath Co., Sept. 3, 1916, J. R. Weir, 782. Aster sp.—Corvallis, Aug. 10, 1911, F. D. Bailey, 1775, May 1, 1915, 3050; Austin, Grant Co., June 1913, J. R. Weir, zo2. 71. PUCCINIA ATRO-FUSCA (Dudley & Thompson) Holway, Jour. Myc. TOr+228: 1904" Uromyces atro-fuscus Dudley & Thompson, Jour. Myc. 10: 55. 1904. ON CYPERACEAE: Carex Douglasit Boot.—Enterprise, Wallowa Co., July 24, 1897, E. P. Sheldon (from Phan. spec. S634). This species possesses amphispores which were first mistaken for the teliospores of a Uromyces. It may be distinguished from other Carex rusts by the presence of the amphispores together with the normal urediniospores, the latter are 20-26 uw long and thin walled, 1.5-2 wu thick. The aecial connection is unknown. It is known only from the Rocky mountain and Pacific coast regions. 72. PUCCINIA BALSAMORRHIZAE Pk. Bull. Torrey Club 11: 49. 1884. Trichobasis Balsamorrhizae Pk. Bot. Gaz. 6: 276. 1881. On CARDUACEAE: Balsamorrhiza deltoidea Nutt.—Corvallis, July 29, 1914, 1472; Hermiston, Umatilla Co., May 12, 1915, 2663. Balsamorrhiza sagittata (Pursh) Nutt.—Hood River, Aug. 11, 1909, 3186; Durfur, Wasco Co., June 19, 1914, 1530. 236 BROOKLYN BOTANIC GARDEN MEMOIRS 73. PUCCINIA BICOLOR Ell. & Fv. Bull. Torrey Club 27: 572. 1900. ON CICHORIACEAE: Hieracium cinereum Howell—Hood River, July 22, 1915, 3325. Hieracium Scouleri Hook.—White Pine, Baker Co., June 1913, J. R. Weir, 7. Hieracium sp.—Durfur, Wasco Co., June 30, 1914, 1338. This very distinct micro-form is known otherwise only from the type collection made at Waitsburg, Wash., May 7, 1900, on H. Scoulert, by R. M. Horner and distributed in E. & E. Fungi Col. 1570. 7A, PUCCINIA BISTORTAE (Strauss) WCF br. 6: 6r. cons, Uredo Polygoni Bistortae Strauss, Ann. Wett. Ges. 2: 103. 1870. ON POLYGONACEAE: Polygonum imbricatum Nutt.—Oregon?, Aug. 1880, Thos. Howell. Polygonum Newberryt Small—Crater Lake, Klamath Co., Sept. 22, 1913, E. P. Meinecke, Cr Pk D (2) 14; N. slope Mt. Jefferson, 2,600 ft., Marion Co., Aug. 16, 1914, H. P. Barss & G. B. Posey, a709- Aus. 27, ro16, Hs P. Barss, 3395. This species may be distinguished from other North American Polygonum rusts by the medium-sized teliospores (16-21 by 26-35 yu) with wall of uniform thickness, without hyaline umbo. It is not known elsewhere in North America on the first mentioned host and otherwise only from Washington on P. Newberryi. Aecia are un- known. 75. PUCCINIA BLASDALEI Diet. & Holw. Erythea 1: 248. 1893. ON ALLIACEAE: Allium attenuifolium Kellog—Corvallis, June 2, 1915, C. E. Owens, 2681. Allium acuminatum Hook.—Hood River Co., June 10, 1917, Leroy Childs. This rust may be distinguished from other Allium rusts by the strongly developed stroma in the telial sori, and the tendency to form confluent telia covering large areas on the stems and leaves. The teliospores are large (16-26 by 40-61 y), thickened to 4-10 yp at the apex. 76. PUCCINIA CALOCHORTI Pk. Bot. Gaz. 6: 228. 1881. Allodus Calochorti Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. On LILIACEAE: Calochortus macrocarpus Dougl.—Redmond, Crook Co., July 21, 1912, Kirk Whited, 3782; Hills near Malheur River, Harney Co., June 6, 1901, W. C. Cusick, 2544; Powder River Mts., Baker Co., Aug. 1896, C. V. Piper, 2460. JACKSON: UREDINALES OF OREGON 23iF All of the above specimens were secured from phanerogamic specimens, the first from the herbarium of the Oregon Agr. College, the others from the herbarium of the N. Y. Botanical Garden. The species is an opsis-form. 77. PUCCINIA CAMPANULAE Carm. Smith’s English Flora 5: 365. 1826. Puccinia Campanulae Fckl. Sym. Myc. 53. 1869. ON CAMPANULACEAE: Campanula Scoulert Hook.—Mary’s Peak, Benton Co., Aug. 15, 1914, 2559; ‘Hood River, July 24, 1914, 3023. A comparison of the above collections with European material shows that the rust is identical and should be referred as above. This is a micro-form unrecorded in America so far as the writer is aware, and known otherwise from North America only from collections made by the writer and others, on C. rotundifolia at Fall Creek, Ithaca, New York. 78. PUCCINIA CHELONIS Diet. & Holw. Hedwigia 36: 297. 1897. ON SCROPHULARIACEAE : Chelone nemorosa Doug|.—Mt. Hood, Sept. 1, 1901, E. W. D. Holway. A micro-form known otherwise only from Washington on the same host. 79. PUCCINIA CHRYSANTHEMI Roze. Bull. Soc. Myc. Fr.1'7:92. 1900. ON CARDUACEAE: Chrysanthemum sinense Sabine—Portland, Nov. 1914, W. H. Dunham, 19806. The above collection from a greenhouse is the only collection we have seen from Oregon. It is doubtless not infrequent in greenhouses throughout the state. The life history is unknown. This rust is evidently a native of Japan, having been introduced into America and Europe where it has become widespread on cultivated chrys- anthemums. 80. Puccinta CicHortit (DC.) Bell, in Kickx. Fl. Fland. 2:65. 1867. Uredo Cichoru DC. Fl. Fr. 6: 74. 1815. ON CICHORIACEAE: Cichorium tntybus L.—Corvallis, Sept. 21, 1914, G. B. Posey, 7931. 81. PuccintA CIcuTAE Lasch, Klotsch. Herb. viv. myc. No. 787. 1845. Puccinia Cicutae Thiim. Bull. Soc. Imp. des Nat. Moscow 52: 136. 1877. 238 BROOKLYN BOTANIC GARDEN MEMOIRS On UMBELLIFERAE: Cicuta occidentalis Greene?—Klamath Falls, Klamath Co., Sept. 8, 1916, J. R. Weir, 223. Cicuta sp.—Eastern Oregon, June, 1885, T. Howell. The last-mentioned specimen is from the herbarium of W. G. Farlow. It is marked on Peucedanum. ‘The host is clearly Cicuta sp. 82. PUCCINIA CIRCAEAE Pers. Roemer’s Neues Mag. 1: 119. 1794. ON ONAGRACEAE: Circaea pacifica Asch. & Magn.—West of Noon station, Benton Co., Aug. 8, 1914, H. P: Barss, 7206; ‘Hood River; July 24516805 3062; Near Mary’s Peak, Benton Co., Aug. 15, 1914, 3263; Sumpter, Baker Co., July 16, 1913, J. R. Weir, 790. 83. Puccinta Cirsit Lasch, in Rabh. Fungi Eur. No. 89. 1859. Puccinia inclusa Syd. Monog. Ured. 1: 56. 1902. On CARDUACEAE: Cirsium americanum (Gray) Robinson—Wren, Benton Co., June 26, 1914, 1332; Corvallis, May 20, 1915, 3242. Cirsium edule Nutt.—Elk City, Lincoln Co., Aug. 20, 1914, 2520. Cirsium undulatum (Nutt.) Spreng.—Sherman, Sherman Co., July 1, 1914, 1966. . 84. PUCCINIA CLAYTONIATA (Schw.) Pk. Bull. N. Y. State Mus. 6: 226... L899; Caeoma (Aecidium) claytoniata Schw. Trans. Am. Phil. Soc. II. Ace204, 1832. Puccinia Mariae-Wilsoni G. W. Clinton; Peck, Bull. Buff. Soc. Nat. Scirt: 6659-1873; Allodus claytoniata Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. On PORTULACACEAE: Claytonia, lanceolata Pursh?—Austin, Grant Co., May, 1916, J. R. Weir, 206. 85. PucciniA CLematipis (DC.) Lagerh. Tromsé Mus. Aarsh. 17: 47. 1895. Aecidium Clematidis DC. FI. Fr. 2: 243. 1805. Aecidium Aquilegiae Pers. Icon. pict. IV. 58. 1806. Puccinia tomipara Trel. Trans. Wis. Acad. Sci. 6: 127. 1885. Puccinia Agropyri E. & E. Jour. Myc. 7: 131. 1892. Puccinia cinerea Arth. Bull. Torrey Club 34: 583. 1907. Puccinia alternans Arth. Mycol. 1: 248. 1909. Puccinia obliterata Arth. Mycol. 1: 250. 1909. JACKSON: UREDINALES OF OREGON 239 ON RANUNCULACEAE: I. Aquilegia formosa Fish.—Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 2573; Hood River, May 14, 1914, 2565, May 9, 1915, 3040; Bridal Veil, Multnomah Co., May 18, 1915, 3051. Aquilegia truncata Fisch. & Mey.—Pokegama, Jackson Co., July 9, 1903, E. B. Copeland (Sydow, Ured. 1767), E. B. Copeland, 3711 (Rocky Mt. Herb. 45896). Clematis Drummondu T. & G.—Freewater, Umatilla Co., July 10, 1914, 2562. Clematis hirsutissima Pursh (C. Douglasii Hook.)—Austin, Grant Go sjuly, 1014; J. RK. Weir, 783: , Clematis ligusticifolia Nutt.—Corvallis, Linn Co., Sept. 2, 1914, F. D. Bailey, 2563, Benton Co., May 4, 1915, 3307. Thalictrum occidentale A. Gray—Corvallis, May 4, 1912, 1147, July 4, 1914, G. B. Posey, 3067, May 4, 1915, 3270; Wren, Benton So;\une'26, 1914, 73370; White Pine, July, 1913, J. R.. Weir, 753; Austin, Grant Co., Aug. 1915, J. R. Weir, 205. Wms oxcean: DT: III: Agropyron dasystachyum (Hook.) Vasey—Redmond, Crook Co., July 2, 1914, 1432. Agropyron lanceolatum Scribn. & Sm.—Redmond, Crook Co., july2, 1914, 7427. Agropyron spicatum (Pursh) Rydb.—Wren, Benton Co., June 26, LOI4, £320. Bromus carinatus Hook. & Arn.—Newberg, Yamhill Co., June 8, 1913, F. D. Bailey, 1797; Portland, July 10, 1905, J. J. Davis, Aug. 23, 1915, E. Bartholomew, (Barth. Fungi Columb. 4846); Corvallis, Sept. 10, 1914, 1577; Philomath, May 10, 1914, 3193; Hood River, May 14, 1914, 1587, 1588, 1593; Grant’s Pass, Josephine Co., Sept. 3, nore, [~R. Weir; 207. Bromus carinatus californicus Shear—Philomath, Jan. 6, 1914, 1148. Bromus grandis (Shear) Hitche.—Corvallis, June 4, 1914, 7384. Bromus hordeaceus L.—Portland, May 21, 1914, 1582. Bromus hordeaceus leptostachys Beck.—Hood River, May 20, 1914, 1585; Springbrook, Yamhill Co., May 14, 1914, F. D. Bailey, 7594, 1595: | Bromus marginatus Nees.—Hood River, Aug. 6, 1914, 1559, July 27, 1915, 3192; -Corvallis, June 24, 1914, G. B. Posey, 1389, June 29, 1914, G. H. Godfrey, 1312; Rose City Park, Portland, Jan. 9, 1914, 1198; Redmond, Crook Co., July 2, 1914, 1422; Hilgard, Union Co., July 10, 1914, 1365. Bromus secalinus L.—Cottage Grove, Lane Co., July 14, I914, F252: 17 240 BROOKLYN BOTANIC GARDEN MEMOIRS Bromus tectorum L.—Clatskanie, Columbia Co., May 20, 1914, he Da balley. 1507, Bromus villosus Forsk.—Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 14006. Bromus vulgaris Shear—Ashland, Jackson Co., Sept. 10, 1914, 1569; Corvallis, Feb. 14, 1914, 3261, July 4, 1914, G. B. Posey, 1478, July 29, 1914, 1443; trail to Sulphur Springs, Benton Co., Nov. 3, 1914, 3195; Mary’s Peak, Benton Co., Sept. 15, 1914, 1574. Elymus condensatus Presl.—Albany, Linn Co., Aug. 1907, D. Griffiths. Elymus glaucus Buckl.—Wren, Benton Co., June 26, 1914, 2321, 1331; Ashland, Jackson Co., Sept. 10, 1914, 1562, 1564; N.slope Mt. Hood, Aug. 7, 1914, 1556; Mary’s Peak, Benton Co., Aug. 15, 1914, 1575; The Dalles, Wasco Co., July 1, 1914, 1347; Garden Home, Multnomah Co., July 20, 1915, 3202; Hood River, June 20, 1914, 1403, Aug. 5, 1914, 3204; Corvallis, Feb. 14, 1914, 3262; June 20, 1914, G. B. Posey, 1304,"G. H.. Godfrey, 7305, July 20; 1914, 0m: 1440. Elymus triticoides Buckl.—Columbia River, near mouth of De- schuttes River, Sherman Co., July 29, 1914, M. E. Peck. Poa ampla Merrill—Hood River, July 22, 1915, 3259. Puccinella Nuttalliiana (Schult.) Hitche.—Grand Ronde Valley, Union Co., July, 1914. Sitanion californicum J. G. Smith—N. slope Mt. Jefferson, Aug. 12.1044, He Ps Barss. 7500: Sitanion glabrum J. G. Smith—Umatilla, Umatilla Cg., July 11, IQ14, 1370. Sitanion jubatum J. G. Smith—Redmond, Crook Co., July 2, 1914, 1428. Sitaniton velutinum Piper—Hood River, July 22, 1915, 3255; Hermiston, Umatilla Co., May 12, 1915, 3179. This common subepidermal species, as here considered, includes nearly if not all the forms having aecia on Ranunculaceous hosts. Dietel (Oesterr. bot. Zeitschr. 42; 261. 1892) was apparently the first to culture this species. Klebahn (Die Wirtsw. Rostpilze 292. 1904) has presented a summary of Dietel’s work together with that of other European investigators. In America, Arthur has conducted extensive culture work beginning in 1904, using telial material from various parts of the country, on five different genera of grasses representing ten species, and has success- fully cultured them on five genera of Ranunculaceae. His work indi- cates the presence of a number of well-marked races. (Jour. Myc. 11: 62. 1905,13:197. 1907,14:15. 1908; Mycologia 1: 246, 248, 249. 1909; 2.225.. TOTO, 47540 1OQE2.7 - 7302.) LOLS, 6: 162s lomas) JACKSON: UREDINALES OF OREGON 241 This species may be distinguished from other grass rusts having the telia long covered by the epidermis primarily on the urediniospore characters. They are not accompanied by paraphyses, the wall is moderately thin, 1.5—2 uw, pale yellow to cinnamon brown, and the pores are scattered. The telia may or may not be surrounded by stromal hyphae and are rather narrow, 13-24 wu. 86. PuccINIA CLINTONII Peck, Rept. N. Y. State Mus. 28:61. 1876. ON SCROPHULARIACEAE: Pedicularis bracteosa Benth.?—N. slope Mt. Jefferson, Aug. 16, 1914, H. P. Barss & G. B. Posey,-2545. Pedicularis racemosa Dougl.—N. slope Mt. Jefferson, 7,000 ft., Awe. 13, 1914, H. P. Barss, 2544. 87. Puccinia Cnicit-Mart. Fl. Mosq. 226. 1817. Puccinia Cuirsii-lanceolatt Schroet., Cohn, Krypt. Fl. Schl. 3}: 3°72, 1887. On CARDUACEAE: Cirsium lanceolatum (L.) Scop.—Corvallis, Oct. 21, 1911, F. D. Bailey, 1963, Mar. 6, 1914, G. H. Godfrey & F. D. Bailey, 1965; Elk Gity, Lincoln Co., Aug. 20, 1914, 1964; The Dalles, Wasco Co., July 1, 1914, 1334; Portland, Aug. 21, 1915, E. Bartholomew (Barth. Fungi Columb. 5053). 88. PuccINIA COMANDRAE Pk. Bull. Torrey Club 11: 49. 1884. ON SANTALACEAE: Comandra umbellata (L.) Nutt.—Dufur, Wasco Co., July 30, 1914, 2504; Hood River, May 18, 1915, 2660, July 22, 1915, 3141. This micro-form, found commonly in the Rocky Mt. and Pacific coast states, possesses teliospores having similar morphological char- acters to those of the heteroecious rust P. pustulata (Curt.) Arth., which has aecia on Comandra and uredinia and telia on Andropogon. A number of such correlations between micro-forms and the telia of heteroecious forms whose aecia occur on the same host have been pointed out by Travelbee (Proc. Ind. Acad. Sci. 1914: 233. 1915) among species occurring in North America. Dietel (in Engler & Prantl, Die Nat. Pflanzenf. 11**: 69. 1897) was apparently the first to point out this sort of correlation between P. mesneriana Thiim. and P. coronata (P. Rhamnt). 89. PUCCINIA COMMUTATA Sydow, Monog. Ured. 1: 201. 1902. Allodus commutata Arth. Résult. Sci. Congr. Bot. Vienne 345. 1906. ON VALERIANACEAE: _ Valeriana occidentalis Heller—Hilgard, Union Co., July 10, 1914, 1541. 242 BROOKLYN BOTANIC GARDEN MEMOIRS go. PUCCINIA CONFERTA Diet. & Holw. Erythea 1: 250. 1893. Puccinia recondita Diet. & Holw. Erythea 2: 128. 1894. ON CARDUACEAE: Artemisia ludoviciana Nutt.—Corvallis, Sept. 2, 1914, F. D. Bailey, 2592, Sept. 4, TOA 2500: A micro-form in which the teliospores resemble quite closely those of P. Absinthit (cf. 58), a brachy-form also common in the west. It is probable that this should be considered a correlated form. 91. PUCCINIA CONVOLVULI (Pers.) Cast. Obs. 1: 16. 1842. Uredo Betae Convolvuli Pers. Syn. Fung. 221. 1801. ON CONVOLVULACEAE: Convolvulus atriplicifolius (Hallier f.) House—Central Point, Jack- son Co., Oct. 6, 1914, M. P. Henderson, 7949; Grant’s Pass, Josephine Co; Sept. 4; 1916, | sk Weir. 227: 92. PUCCINIA CREPIDIS-ACUMINATAE Sydow, Oestr. Zeitschr. 51: 27. IQOl. ON CICHORIACEAE: Crepis gracilis (D. C. Eaton) Rydberg—Baker City, Baker Co., July, 1913; > Ra Weir 75-7: 93. PuccINIA CyAnlI (Schleich.) Pass. Rabh. Fungi Eur. No. 1767. 1874. Uredo Cyani Schleich. Pl. Helv. 95. ON CARDUACEAE: Centaurea Cyanus L.—Corvallis, June, 1913, 7745, April 8, 1914, 2557, July. 20, 1914, 2552, July. 4, 1914, G. B.Posey, 3106; Orenear Washington Co., April 2, 1915, 3060. 94. Puccin1A DEBARYANA Thiim. Flora 58: 364. 1875. Puccinia compacta DeBary, Bot. Zeit. 16: 83. 1858. (Not P. compacta Berk. 1855.) On RANUNCULACEAE: Anemone Drummondu Wats.—Mt. Hood, 7,000 ft., foot of Eliott Glacier, Sept. 1, 1901, E. W. D. Holway. 95. Puccinté DENTARIAE (Alb. & Schw.) Fuckel, Symb. Mycol. Nachtr a 47513878. Uredo Dentariae Alb. & Schw. Consp. Fung. 129. 1805. ON CRUCIFERAE: Dentaria tenella Pursh—Corvallis, April 5, 1914, 7288. A micro-form occurring on the petioles and blades of the basal leaves causing considerable distortion. So far as the writer is aware this species is known from North America only from the above col- lection. JACKSON: UREDINALES OF OREGON 243 96. PUCCINIA DICHELOSTEMMAE D. & H. Erythea 3: 78. 1895. Allodus Dichelostemmae Orton, Mem. N. Y. Bot. Gard. 6: 183. Ig16. ON ALLIACEAE: Hookera pulchella Salisb. (Brodiaea congesta Smith)—Dallas, Polk Co., March 20, 1900, W. N. Suksdorf (Barth. N. Am. Ured. 1541); Corvallis, April 28, 1915, 2611, May 1, 1915, 2069; E. of Wren Station, Benton Co., April 17, 1915, 2678. The first-mentioned collection bears aecia and a few telia. It is probable that they belong together. The last specimen mentioned consists of aecia only. The others bear telia only. The two stages rarely occur together. This species may be separated from all other species of Puccinia occurring on Alliaceae by the very large, broad teliospores (38-45 by 43-58 uw) having smooth walls 5—7 wu thick. 97. PucctiniA DouGtasit Ell. & Ev. Proc. Phil. Acad. 1893: 152. 1893. Puccinia Richardsontui Sydow, Monog. Ured. 1: 317. 1902. Allodus Douglasu Orton, Mem. N. Y. Bot. Gard. 6: 198. I916. ON POLEMONIACEAE: Phlox condensata (A. Gray) E. Nels.—N. slope Mt. Hood, 7,000 ft., mug, 7, 1914, III ,-7404,.1, (2024. Phlox diffusa Hook.—N. slope Mt. Hood, 7,000 ft., Aug. 7, 1914, 1602, 1603; Sept. 1, 1901, E. W. D. Holway. 98. PucciniA ELxiist DeToni, in Sacc. Syll. Fung. 7: 651. 1888. Puccinia Angelicae E. & E. Bull. Wash. Lab. 1: 3. 1884. (Not P. Angelicae Fckl. 1869.) Puccinia Bakeriana Arth. Bull. Torrey Club 31: 3. 1904. On UMBELLIFERAE: Angelica genufleca Nutt.—Corvallis, Sept. 7, 1901, E. R. Lake, 14090. Otherwise known only from Idaho and Washington on the above host, and from California on A. tomentosa. It is doubtless a brachy- form, though no pycnia have been found. The teliospores are closely and finely verrucose, a character which enables one to distinguish this species easily from P. Angelicae (cf. 62). 99. PUCCINIA EPILOBII-TETRAGONI (DC.) Wint. in Rabenh. Krypt. 1 Ben ene: 8 A iota he Uredo vagans a Epilobit-tetragont DC. FI. Fr. 2: 238. 1805. Puccinia Gayophyti Billings, in King, Geol. Expl. 4oth Par. 5: 414. 1871. Puccinia Oenotherae Vize, Grevillea 5: 109. 1877. Puccinia Boisduvaliae Pk. Bot. Gaz. 7: 45. 1882. 244 BROOKLYN BOTANIC GARDEN MEMOIRS Puccinia Clarkiae Pk. Bull. Torrey Club 11: 49. 1884. Puccimia glabella Holw. N. Am. Ured. 1: 76. 1907. ON ONAGRACEAE: Boisduvalia densifolia (Lindl.) Wats.—Minam River, Wallowa Co., Oct. 2, 1897, E. P. Sheldon, 9049; Corvallis, July, 1910, 1776, Aug. 10, 1911, 1123, Sept. 20, 1914, 1546; Calapooya Valley, Douglas Co., July 24, 1899, M. A. Barber (Rocky Mt. Herb. 40989); Grant's Pass, Josephine Co., Sept. 2, 3, 1916, J. R. Weir, 222, 251. Boisduvalia glabella (Nutt.) Walp.—Burns, Harney Co., Aug. Igot, Griffiths & Morris (Griffiths, W. Am. Fungi 385). Boisduvalia stricta (A. Gray) Greene—Corvallis, Aug. 13, 1914, 1492; Medford, Jackson Co., June 26, 1915,.G. B. Posey) g2z¢e Wimer, Jackson Co., July 22, 1892, E. W. Hammond, 149 (Rocky Mt. Herb. 486096). Clarkia pulchella Pursh—Hilgard, Union Co., July 10, 1914, 1520. Epilobium minutum Lind|l.—Corvallis, Aug. 15, 1909, I170. Epilobium paniculatum Nutt.—Cole’s Creek, Wallowa Co., June 10, 1897, E. P. Sheldon, 8263 (Rocky Mt. Herb. 70411); Hood River, May 14, 1914, 1510, May 16, 1915, 3271; Hilgard, Union Co., July 10, 1914, 7530, 1544; Ontario, Mahheur Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 383); Corvallis, Aug. 1910, 3065, Aug. 10, 1911, F. D. Bailey, 1774; The Dalles, Wasco Co., June 19, 1914, 3107; N. slope Mt. Hood, Aug. 7, 1914, 1491; Near Cascade Locks, Hood River Co., Aug. 11, 1910, 1073; Philomath, May 26, 1914, 3351, April 21, 1899, Moses Craig; Klamath Falls, Klamath Co: Voepiac, 1016. jek y Weir 2252 Gayophytum ramossissimum T. & G.—Redmond, Crook Co., July I, 1914, 2536; Hood River, July 23, 1915, 3272; Farewell Bend, Crook Co., July 15, 1894, J. B. Lieberg, 435 (Rocky Mt. Herb. 66228). Godetia amoena (Lihm.) Lilja.—Corvallis, July, 1910, 7775, Wren, Benton Co., June 26, 1914, 1327. Sphaerostigma Boothi (Dougl.) Walp.—Muddy Station, John Day Valley, May 12, 1885, Thomas Howell. Sphaerostigma dentatum (Cav.) Walp.—Pleasant Creek, near Wimer, Jackson Co., April 23, 1889, E. W. Hammond, 143. As here considered, this species includes all the long-cycled autoe- cious forms occurring on Onagraceae. The treatment follows the disposition made of them by Bisby in his recent admirable discussion of the Onagraceous rusts (Amer. Jour. Bot. 3: 538. 1916). 100. PUCCINIA EPIPHYLLA (L.) Wettst. Verhl. Zool.-Bot. Ges. Wein. 35: 541. 1885. Lycoperdon epiphyllum L. Sp. Pl. 1653. 1753. Aecidium Tussilaginis Pers. in Gmel. Syst. Nat. 2: 1473. 1791. JACKSON: UREDINALES OF OREGON 245 Puccinia Poarum Niels. Bot. Tidsskr. II. 3: 26. 1877. On POACEAE: Poa ampla Merrill—Hood River, May 14, 1914, 159TI. Poa annua L.—Hood River, July 22, 1915, 31906. Poa macrantha Vasey—Newport, Lincoln Co., June 20, 1915, 3123. Poa pratensis L.—Corvallis, May 19, 1913, F. D. Bailey, z194, March 29, 1914, G. B. Posey, 3126, April 29, 1914, F. D. Bailey, 3125, June 29, 1914, G. B. Posey, 1311; Philomath, May 10, 1914, 3124; Hood River, May 14, 1914, 1586, 1592; The Dalles, Wasco Go.,-)uly: 1, ror, 7302; 'N. slope .Mt.. Hood, “Aug. 7; 1954; 1557; Ashland, Jackson Co., Sept. 10, 1914, 1565; Kamela, Union Co., July 22, 1915, M. E. Peck; Klamath Falls, Klamath Co., Sept. 8, 1916, J. R. Weir, 224, 239. Poa triflora Gilib.—Klamath Falls, Klamath Co., Sept. 8, 1916, J. R. Weir, 239a. Poa sp.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 228, 229; Klamath Falls, Klamath Co., Sept. 8, 1916, J. R. Weir, 218; Austin, Grant Co., Aug. 1915, J. R. Weir, 194. This rust is especially common in western Oregon particularly on blue grass. Only uredinia are known in the above collections as is the common condition except in those made in the far north or at high elevations. Nielsen (Bot. Tidsskr. 2: 26. 1877) was the first to show the relation between this rust and Aecidium Tussilaginis Gmel. He suc- ceeded in infecting P. annua, P. trivialis, P. nemoralis, P. fertilis and P. pratensis by sowing aeciospores from Tussilago farfara. He in- fected the aecial host by sowing with teliospores from P. annua. Additional observations and culture work have been recorded by various European authors, which have been summarized by Klebahn (Die Wirtsw. Rostp. 290. 1904). 101. PuccINIA ERIOPHORI Thiim. Bull. Soc. Imp. Nat. Moscow 55: 208. 1880. Aecidium Ligulariae Thiim. Nov. Giorn. Bot. Ital. 12: 196. 1880. Aecidium Cinerariae Rostr. Overs. Kong. Dansk. Vid. Selsk. Forh. Koph. 1884-5: 17. 1884. On CARDUACEAE: I. Senecio ductaris Piper—Alpine meadow, E. Mt. Hood, 5,000 ft., July 23, 1915, 3320. ON CYPERACEAE: III. Eriophorum polystachyon L.—Alpine meadow, E. Mt. Hood, 5,000 SE, July-23- 1015, 7327- In the alpine meadow where the above collections were made the Aecidium was very abundant and in fine condition. A search was 246 BROOKLYN BOTANIC GARDEN MEMOIRS made for overwintered telia on Cyperaceous hosts associated with the Senecio and the only rust found was on very much weathered leaves which have been determined by comparison of the microscopic struc- ture as above indicated. Tranzschel (Beitr. Biol. Ured. III: 4. 1907), working in Russia, was the first to culture this species. He used telial material on Eriophorum angustifolium to successfully infect Ligularia sibirica and Senecio paluster. In America, Arthur (Mycol. 8: 131. 1916), using aecial material from New York on Senecio aureus, obtained successful infection result- ing in uredinia and telia on E. viridi-carinatum. 102. Puccinia Eriophyllii sp. nov. O.and I. Pyenia and aecia unknown. II. Uredinia amphigenous and _ caulicolous, scattered, small, roundish, 0.3-0.6 mm. across, early naked, pulverulent, pulvinate, chestnut brown, surrounding epidermis not conspicuous; uredinio- spores globoid, obovoid or oblong, 21-27 by 26-29 u, wall chestnut brown, 2-3 » thick, moderately and finely echinulate, pores 2, approxi- mately equatorial. Ill. Telia amphigenous, scattered, small, round, 0.3-0.6 mm. across, early naked, compact, pulvinate, blackish brown, ruptured epidermis not conspicuous; teliospores ellipsoid to oblong, 18-21 by 26-30 wu, apex and base rounded, not constricted, wall chestnut brown, 2-2.5 w thick, uniform, minutely and obscurely verrucose; pedicel colorless, deciduous. ON CARDUACEAE: Eriophyllum lanatum (Pursh) Forbes—Wren, Benton Co., June 26, 1914, 1319 (type). Eriophyllum leucophyllum (DC.) Rydberg—Redmond, Crook Co., July 1, 1914, 30853. Distinguished from other species of Puccinia on related hosts by the very small teliospores. 103. Puccinia Fendleri (Tracy & Earle) comb. nov. Aecidium Fendleri Tracy & Earle, in Green, Pl. Baker 1:17. 1901. Puccinia Koeleriae Arth. Mycologia 1: 247. 1909. On BERBERIDACEAE: I. Berberis aquifolium Pursh—Corvallis, May 12, 1914, 1276, June 7, 1908, J. C. Bridwell, 3389. Berberis nervosa Pursh—Hilgard, Union Co., July 10, 1914, 2568. ON POACEAE: Koeleria cristata (L.) Pers.—Hilgard, Union Co., July 10, 1914, TOS: JACKSON: UREDINALES OF OREGON 247 The aecia of this species have often been confused with those of P. graminis (cf. 151). Arthur in 1908 (Mycol. 1: 246. 1907), using telial material from Koeleria cristata from Colorado, obtained, as a result of infection experiments, the development of aecia on Berberis aquifolium. 104. PUCCINIA GEMELLA Diet. & Holway, in Sydow’s Monog. Ured. 1: 541. 1903. ON RANUNCULACEAE: Caltha biflora DC.—N. slope Mt. Jefferson, 6,000 ft., Aug. 13, 1914, H. P. Barss & G. B. Posey, 1624; foot of Mt. Jefferson, 5,000 ft., Aus. 28, 1916, H. P: Barss, 3407. A micro-form, differing from P. Treleasiana Pazsch., which occurs in the Rocky Mt. region on Caltha sp., in the smooth spores. 105. PuccINnIA GENTIANAE (Strauss) Link, in Willd. Sp. Pl. 67: 73. 1825. Uredo Gentianae Strauss, Ann. Wett. Ges. 2: 102. I810. ON GENTIANACEAE: Gentiana oregana Engelm.—Sumpter, Baker Co., July 16, 1913, J: Re Weir, ror. 106. PucciniA GILIAE Hark. Bull. Cal. Acad. 1: 34. 1884. ON POLEMONIACEAE: Navarettia tntertexta (Benth.) Hook.—Corvallis, Aug. 1898, Moses Craig, July, 1910, 7727; Umpqua Valley, Douglass Co., June, 1887, Thomas Howell, 7835; Hood River, Aug. 17, 1888, L. F. Henderson (673), 1120. This is a hemi-form distinct from P. plumbaria (cf. 150) which is an opsis-form. In P. Giliae the telia are early naked and the spores smooth. In P. plumbaria the telia are long covered by the cinereous epidermis and the spores are finely and closely verrucose. 107. PUCCINIA GLUMARUM (Schmidt.) Erikss. & Henn. Zeits. Pflan- Zenkts 4207. 1804: Uredo glumarum Schmidt. Allg. Oekon. Fl. 1: 27. 1827. Puccinia neglecta West. Bull. Soc. Bot. Belg. 2: 248. 1863. Trichobasis glumarum Leyv.; Cooke, Myc. Fung. 208. 1865. ON POACEAE: Elymus glaucus Buckl.—Hood River Co., May 14, 1914, 1590, 1590, 1597, July 23, 1915, 3199. Hordeum Gussoneanum Parl.—Corvallis, June 4, 1914, F. D. Bailey, 1385. Hordeum vulgare L.—Moro, Sherman Co., June 11, 1915, F. K Ravn and A. G. Johnson. 248 BROOKLYN BOTANIC GARDEN MEMOIRS Sitanion hystrix (Nutt.) J. S. Smith—Redmond, Crook Co., July I, 1914, 1423, 1429. Sitanion jubatum J. G. Smith—Ashland, Jackson Co., June 7, 1916, H. B. Humphrey; Klamath Falls, Klamath Co., Sept. 11, 1916, Jo Ro Weir? 232: Triticum aestivum L.—Moro,.Sherman Co., June 25, 1915, D. E. Stephens, 3371, 3379; Medford, Jackson Co., June 8, 1915, F. K. Ravn, A. G. Johnson, 3370. Triticum compactum Host.—Moro, Sherman Co., June II, 1915, F. K. Ravn & A. G. Johnson (Barth. Fungi Columb. 4756); June 25, 1915, D. E. Stephens, 3373, 3378, 3380, 3385. Triticum diococcum L.—Moro, Sherman Co., June 11, 1915, F. K. Ravn & A. G. Johnson. Triticum vulgare L. (Collective)—Corvallis, June 10, 1915, 2676, 2679, 2680, June 12, 1915, 3134; Hood River, July 22, 1915, 3742; Moro, Sherman Co., June II, 1915, F. K. Ravn, A. G. Johnson, M. A. Carleton. This very important wheat rust was first found in the United States, May 21, 1915, at Sacaton, Arizona, by Dr. F. K. Ravn, the eminent Danish cereal pathologist, who at that time was making a tour of investigation of cereal diseases in company with pathologists of the Department of Agriculture (Carleton, Science N. S. 42: 58. 1916). A few weeks later the rust was found by Dr. Ravn and party at Medford and Corvallis and later was detected at Moro, Oregon. There is every evidence that this rust has been present in the western states for some years. Several collections reported above were made in 1914 but were confused with P. Clematidis. The writer is indebted to Dr. H. B. Humphrey for examining most of the collections and for the detection of several specimens belonging here, previously referred to other species. 108. PUCCINIA GRANULISPORA Ell. & Gall.; Ellis & Ever. Bull. Torrey Glib. 22761-1805; On ALLIACEAE: Allium nevit Wats.?-—Austin, Grant Co., Aug. 1915, J. R. Weir, 204. 109. PUCCINIA GROSSULARIAE (Schum.) Lagerh. Troms¢. Mus. Aarsh. 17: 60. 1895. Aecidium Grossulariae Schum. Pl. Enum. Saell. 2: 223. 1803. Puccinia Pringsheimiana Kleb. Zeits. fiir Pflanzenkr. 4: 194. 1894. Puccima Magnusivi WKleb. Zeits. fiir Pflanzenkr. 5: 79. 1895. Puccinia albiperidium Arth. Jour. Myc. 8: 53. 1902. Puccinia unitporula Orton, Mycol. 4: 201. 1912. JACKSON: UREDINALES OF OREGON 249 On GROSSULARIACEASE: I. Ribes divaricatum Dougl.—Bridal Veil, Multnomah Co., May 18, 1915, 3252; Hilgard, Union Co., July 10, 1914, 30071. Ribes lacustre (Pers.) Poir.—N. slope Mt. Hood, Aug. 7, 1914, 2561. Ribes sanguineum Pursh—Bridal Veil, Multnomah Co., May 18, 1915, 3253- Ribes sp.—Philomath, April 26, 1914, 2571; Corvallis, April 11, 1915, 3045- On CYPERACEAE: II, III. Carex festiva Dewey—Hilgard, Union Co., July 10, 1914, 1360. Carex Goodenowu J. Gay—Hilgard, Union Co., July 10, 1914, 1359. Carex Kelloggu W. Boot—Portland, Aug. 21, 1915, E. Bartholomew 5941 (Barth. Fungi Columb. 4962). Carex magnifica Dewey—Clatsop Co., Nov. 7, 1913, 1195. Carex mertensui Prescott—Mt. Hood, Aug. 7, 1914, 3004. Carex monitle Tuckerm.—Clatskanie, Columbia Co., May 20, 1914, B.D; Bailey, 3073: Carex nebraskensis Dewey—Hilgard, Union Co., July Io, 1914, F205. Carex phyllomanica W. Boot?—Klamath Falls, Klamath Co., Sept. 8; 1916, J. R. Weir, 252. Carex spectabilis Dewey—W. slope Mt. Jefferson, July 3, 1914, F. D. Bailey, 1417; Vicinity Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 3007. Carex sp.—In open meadow along Minum River, Wallowa Co., Aug. 20, 1897, E. P. Sheldon, 87571. This common form having aecia on Ribes sp. was first cultured by Klebahn in 1892. The species has since been extensively studied by the culture method in both Europe and America (Klebahn, Die Wirtsw. Rostp. 295-302. 1904) under various names. In America, Arthur began culture work in 1901 and has reported the results of numerous cultures (Jour. Myc. 8: 53. 1902; I0: II. EQOA; EL: 59. “1905; 12: 65: 1906; 13: 196. 40077 14712. 1008; Mycol. 4-132. 1912; 72> 67; TO15; 7:78: TOES). he species is doubtless represented by several biological strains and further culture work will need to be conducted in order to determine their limits. Considerable confusion has resulted on account of the variable number and position of the germ pores in the urediniospores. 110. PUCCINIA GRUMOSA Syd. & Holw. in Sydow, Monog. Ured. 1: 645.” 1003. ON LILIACEAE: Stenanthium occidentale A. Gray—Bridal Veil, Multnomah Co., May 18, 1915, 2670; Hood River, July 24, 1915, 3082. 250 BROOKLYN BOTANIC GARDEN MEMOIRS This species, described from a collection on Zygadenus elegans made by Professor Holway at Banff, Alberta, has been previously known only from the original collection. The above collections clearly belong here and add a new host. The only other collection of Puccinia on Stenanthium known to the writer is one obtained by him in January, 1917, on a phanerogamic specimen of S. gramineum col- lected in Georgia by A. W. Curtis, now in the herbarium of the New York Botanical Garden. This has been referred to P. atropuncta, a species known only from east of the Rocky mountains on related hosts. 111. PUCCINIA HARKNESSI Vize, Grevillea 7: 11. 1878. Puccinia cladophila Pk. Bot. Gaz. 4: 127. 1879. ON CICHORIACEAE: Lygodesmia juncea (Pursh) D. Don—Denio, Harney Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 396c). Ptiloria paniculata (Nutt.) Green—Sherman, Sherman Co., July 1, 1914, 2535. 112. Puccinia Helianthi-mollis (Schw.) comb. nov. Aecidium Heliantht-mollis Schw. Schrift. d. Nat. Ges. Leipzig 1: 68. 1822. Puccima Hehantht Schw. Schrift. d. Nat. Ges. Leipzig 1: 73. 1822. On CARDUACEAE: Helianthus annuus L.—Sherman, Sherman Co., July I, 1914, 2525; Corvallis, Aug. 1910, F. D. Bailey, 7729; Umatilla, Umatilla Co., July 113 1914, 7268: 113. PuccINIA HEMIZONIAE Ell. & Tracy, Jour. Myc. 7: 43. 1891. ON CARDUACEAE: Hemizonia truncata (DC.) Gray—Grant’s Pass, Josephine Co., July 12, 1887, Thos. Howell. Lagophylla ramossissima Nutt.—Grant’s Pass, Josephine Co., Sept. 2, 1916, J. R. Weir, 220. 114. PuccINIA HEUCHERAE (Schw.) Diet. Ber. der Deutsch. Bot. Ges. 03942. 751891: Uredo Heucherae Schw. Schrift. Nat. Ges. Leipzig 1: 71. 1822. Puccinia Tiarellae B. & C. Grevillea 3: 53. 1874. Puccima spreia Pk. Rep. N. Y. State Mus. 29: 67. 1878. Puccinia congregata E. & H. Bull. Calif. Acad. Sci. 1: 26. 1884. ON SAXIFRAGACEAE: Heuchera micrantha Dougl.—Hood River, Feb. 1, 1915, 3266; Mary’s River, Corvallis, June 5, 1915, 2674; Ashland, Jackson Co., Sept Ol MoOi4, erga: JACKSON: UREDINALES OF OREGON 251 Leptaxis Menziesti (Pursh) Raf.—Hood River, July 24, 10915, 3318, 3359. Mitella Brewert Gray?—N. slope Mt. Jefferson, Aug. 27, 1916, Ee. Barss; 3307. Mitella ovalis Greene—Mary’s Peak, Benton Co., May 23, 1915, 3037- Mitella sp.—N. slope Mt. Jefferson, 8,000 ft., Aug. 8, 1914, H. P. Barss & G. B. Posey, 2529. Tellima grandiflora (Pursh) Dougl.—Corvallis, July 15, 1910, mie. Apr.o, £014, 3075; Austin, Grant Co. Aug. 19055, J. R.. Weir, 274. Tiarella untfoliata Hook.—Bridal Veil, Multnomah Co., Aug. 11, 1910, 1070; Ashland, Jackson Co., Sept. 10, 1914, 3028. 115. Puccinia hieraciata (Schw.) comb. nov. Caeoma (Aecidium) hieraciatum Schw. Trans. Am. Phil. Soc. II. A 292. - 1832> Puccinia patruelis Arth. Mycol. 1: 245. 1909. On CypERACEAE: II, ITI. Carex praegracilis Boott (C. marcida Boott)—Ontario, Malheur Co., Aug. 1901, Griffiths & Morris; Andrews, Harney Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 339a). This species shows a distribution from coast to coast and has aecia on Cichoriaceous hosts. Arthur (I. c.) has conducted one successful culture and obtained the development of pycnia and aecia on Agoseris glauca following sowings of teliosporic material on Carex pratensis from Colorado. Other aecia having a similar morphology, including a rare form on Hieracium collected by Schweinitz, on which the present name is based, are properly referred here. No aecial collections have been made in Oregon. 116. PucciniA HiERAct (Schum.) Mart. Fl. Mosq. 226. 1812. Uredo Hieracui Schum. Enum. PI. Saell. 2: 232. 1803. Puccinia sejuncta Syd. Ann. Myc. 1: 236. 1903. ON CICHORIACEAE: Meracium albiflorum Hook.—Hood River, May 16, 1915, 3312, ful: 22, TOL5;-3 327. Hieracium cinereum Howell—Hood River, July 22, 1915, 3324. Hieracium gracile Hook.—N. slope Mt. Jefferson, Aug. 6, 1914, li. P. Barss,. 2547. ; Hieracium scouleri Hook.—Austin, Grant Co., Aug. 1915, J. R. Weir, 156. Mieracium sp.—Hilgard, Union Co., July 9, 1914, 3379; Austin, Grant Co., Aug. 1915, J. R. Weir, 273; Klamath Falls, Klamath Co., sept. 3, 1906), J. R. Weir, 270. 252 BROOKLYN BOTANIC GARDEN MEMOIRS This species may occur on the same plants with Aecidium colum- biense (cf. 215) which is doubtless the aecial stage of some heteroecious rust not yet determined.. Sydow (I. c.) has described P. sejuncta based on such a mixture. 117. PuccintA HOLBOELLII (Hornem.) Rostr. Middelser om Groen- land 3: 534. 1888. Aecidium Holboellit Hornem. Fl. Dan. 37: 11. 1840. Puccinia Barbareae Cooke, Grevillea 8: 34. 1879. The type of P. Barbareae was described as on a “‘Cruciferous plant’’ from Oregon, Dr. Lyall 6z. The data on the type collection at the Kew Herbarium reads “Oregon Boundary Commission, Ft. Coville to Rocky Mts. 1861, Dr. Lyall 67.’ Since Ft. Coville is in north- eastern Washington there would seem to be little chance of this col- lection having been made within the state of Oregon. However, since it has been recorded from our limits both in the original descrip- tion and by Holway (N. Am. Ured. 1: 45. 1906) it is included here with the above explanation. The species undoubtedly does occur in eastern Oregon as the range includes all the surrounding states. 118. PUCCINIA HOLCINA Erikss. Ann. Sci. Nat. 9: 274. 1899. On POACEAE: Holcus lanatus L.—Corvallis, June 10, 1915, 2678, June 12, 1915, 3113; Toledo, Lincoln.Co., July 19, 1915, 3776; Yaquina, Lincoln Col; July-17; 1015, 3777, Salem, Marion Go. .May 1, 19 1T4-n@aene Godfrey, 317178; Portland, Aug. 19, 1915, E. Bartholomew (Barth. Fungi Columb. 4852). Evidently a common rust in western Oregon, otherwise known from North America from a few other collections made on the Pacific coast from California to Vancouver Island, B. C., and from a single locality along the eastern coast in Massachusetts. This species may be easily separated from P. Rhamnz (cf. 159) in the telial stage by the evenly thickened apices of the teliospores. The urediniospores of the two rusts are similar. Those of the present species are somewhat larger and globoid, 19-24 by 23-27 u, while in P. Rhamnzi they are globoid or broadly ellipsoid, 16-20 by 18-24 wu. The aecial connection is unknown. The rust has evidently been introduced from Europe and is able to maintain itself by over-wintering urediniospores. 119. PuccintiA HyPocHoERIpIs Oud. Nederl. Kruidk. Arch. II, 1: 175. Dove. On CARDUACEAE: Hypochaeris radicata L.—Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 2543. JACKSON: UREDINALES OF OREGON it) Sat W 120. Puccinia insperata sp. nov. O. Pycnia not seen. I. Aecia chiefly hypophyllous and_ petiolicolous; in crowded groups on yellowish spots 2—3 mm. across; cupulate, 0.2—.25 mm. broad; peridium whitish, margin recurved, lacerate; peridial cells rhombic, 19-27 by 35-45 u, overlapping, outer wall I-1.5 w thick, inner wall 3-4 4 thick, verrucose; aeciospores globoid or broadly ellipsoid, 15-19 by 19-23 p, wall colorless, 1-1.5 uw thick, finely and closely verrucose. II. Uredinia amphigenous, scattered, round, 0.2-0.5 mm. across, tardily naked, pulverulent, pulvinate, cinnamon brown, ruptured epidermis conspicuous; urediniospores subglobose or broadly ellip- soid, occasionally obovate, 19-21 by 23-29 uw; wall cinnamon brown, 1.5-2 uw thick, minutely and closely echinulate, pores 2—3, scattered. Ill. Telia amphigenous and petiolicolous, scattered, round, 0.2- 0.8 mm. across, tardily naked, pulvinate, becoming pulverulent, blackish brown, ruptured epidermis conspicuous; teliospores ellipsoid or broadly obovoid, occasionally somewhat irregular, 16-20 by 23-32 u, rounded at both ends, scarcely or not at all constricted, wall chestnut brown, 1.5-2 uw thick, uniform, smooth; pedicel colorless, deciduous. ON CICHORIACEAE: Nabalus hastatus (Less) Heller—Hood River, May 16, 1915, 2662, July 24, 1915, 3265, type. A very distinct species separable from the eastern P. orbicula Pk. by the smooth teliospores and the presence of a definite peridium in the aecidium. The two collections were made at the same spot. The first shows aecia unaccompanied by pycnia, associated with telia chiefly on the petioles, suggesting strongly an opsis-form. The second collection, however, shows scattered uredinia and telia with a few old aecia. ii. PuccintA Iris (DC.) Wallr in Rabenh.. Krypt. Fl. in3 23: 1844. Uredo Indis DC. Encycl. 8: 224. 1808. ON IRIDACEAE: Iris tenax Dougl.—Corvallis, June 24, 1914, F. D. Bailey, 1343; Wren, Benton Co., July 22, 1914, 1473, Ashland, Jackson Co., Sept. IO, 1914, 1904. 122. PuCCINIA JONESII Pk. Bot. Gaz. 6: 226. 1881. Allodus Jones Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. ON UMBELLIFERAE: Peucedanum triternata (Pursh) Nutt.—Austin, Grant Co., Aug. 1915, J. R. Weir, 149, 157. 254 BROOKLYN BOTANIC GARDEN MEMOIRS 123. Pucctnia Licustici Ell. & Ev. Bull. Torrey Club 22: 263. 1895. Puccinia luteobasis Ell. & Ev. Bull. Torrey Club 24: 457. 1897. ON UMBELLIFERAE: Ligusticum aptifolium (Nutt.) Gray—Corvallis, 1911, 7166, April 15, 1913, EF. D. Bailey, 3050, March 23, 1914, (G. B: Posey, gaame March 22, 1914, G. H. Godfrey, 2549; Orenco, Washington Co., Junei22)- 1013, 3070: 124. PUCCINIA LUXURIOSA Sydow, Monog. Ured. 1: 812. 1904. Puccinia tosta luxurians Arth. Bull. Torrey Club 29: 229. 1902. ON POACEAE: Sporobolus airoides Torr.—Andrews, Harney Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 304). Bethel (Phytopath. 7: 93. 1917) has reported successful cultures of this rust on Sarcobatus vermiculatus, amply supported by field observations. Arthur (Mycol. 1: 234. 1909) has infected that host with teliospores of P. subnitens (cf. 167) from Nevada. Bethel, however, failed to obtain infection on any of the aecial hosts for that species with teliospores from Sporobolus airoides in Colorado. It is possible that the two forms represent biological races of the same species and should be united. 125. Puccinta MAJANTHAE (Schum.) Arth. & Holw. Bull. Lab. Nat. Hist. Univ. Iowa 5: 188. Igot. Aecidium Majanthae Schum. Enum. PI. Saell. 2: 224. 1803. Puccinia sessilis Schneider, in Schréter Abh. Schles. Ges. 49: 19. 1870. On POACEAE: Phalaris arundinacea L.—Beulah, Malheur Co., Aug. tIgo1, Griffiths & Morris (Griffiths, W. Am. Fungi 26a). No culture work has been reported in America. In Europe the aecia have been found on Allium, Arum, Convallaria and various other related hosts. Various names have been given to the different bio- logical forms (Klebahn, Die Wirtsw. Rostp. 263-272. 1904; Sydow, Monog. Ured. 1: 776-784. 1904). Aecia on Smilacina, Polygonatum, Maianthemum and Uvularia from the central and eastern United States doubtless belong here in whole or in part. 126. PuccINIA MALVACEARUM Bert. Gay’s Hist. de Chile 8:43. 1852. On MALVACEAE: Abutilon ? sp. (cultivated shrub)—Corvallis, May, 1914, W. E. Lawrence, 3364, Nov. 17, 1914, W. E. Lawrence, 3363, Jan. 12, 1915, 2627. Althaea ficifolia Cav.—The Dalles, Wasco Co., Aug. 25, 1915, E. Bartholomew (Barth. Fungi Columb. 4758, N. Am. Ured. 1559). JACKSON: UREDINALES OF OREGON 255 Althaea rosea Cav.—Corvallis, April 27, 1907, E. R. Lake, 1299, Aug. 1909, 1068, Sept. I910, ror4, Jan. 12, 1915, W. E. Lawrence, 2627; Gibbon, Umatilla Co., June 5, 1911, comm. 1850; Klamath Falls, Klamath Co., Sept. 19, 1911, 1031; Salem, Marion Co., May 2, 1913, comm. 7866; Svenson, Clatsop Co., June 5, 1913, comm. 1895; Grant’s Pass, Josephine Co., July, 1913, 1926; Dayton, Yamhill Co., April 6, 1914, comm. 1479; Lents, Clackamas Co., April 27, 1914, comm. 1936; Ione, Morrow Co., June 26, 1914, comm. 1937; The Dalles, Wasco Co., July 1, 1914, 1342; Hillsboro, Washington Co., July 26, 1914, 7738; Medford, Jackson Co., March 31, 1915, 2556. Malva rotundifolia L.—Corvallis, Apr. 12, 1909, J. C. Bridwell, 3388, Aug. 30, 1913, 1741, Feb. 14, 1914, 1181; The Dalles, Wasco Gor, july 1, 1914, 1303. Malva ? sp.—New Pine Creek, Lake Co., July, 1910, comm. 3376. 127. Pucctnta McCLatcHIEANA Diet. & Holw. Erythea 2: 127. 1894. ON CYPERACEAE: Scirpus microcarpus Presl.—Elgin, Union Co., Aug. 19, 1897, E. P. Sheldon, $735; Beulah, Malheur Co., Aug. 1901, Griffiths & Morris (Griffiths, W. Am. Fungi 348); Glenbrook, Benton Co., Aug. 1909, 1190; Hubbard, Marion Co., May 27, 1914, 2518; Gresham, Multnomah Co., June 6, 1914, F. D. Bailey, 2516; Tualatin, Washing- ton Co., July 10, 1914, F. D. Bailey, 1356; Hood River, Aug. 5, 1914, 2520; Mary’s Peak, Benton Co., Aug. 15, 1914, 2517; Orenco, Wash- ington Co., April 2, 1915, 3386; Yaquina, Lincoln Co., July 20, 1915, aziz toledo, Lincoln Co., July 19, 1914, 3072. 128. PUCCINIA MELANCONOIDES Ell. & Hark. Bull. Calif. Acad. Sci. Toys P1oO4. Allodus melanconioides Arth. Result Sci. Congr. Bot. Vienne 345. 19006. ON PRIMULACEAE: Dodecatheon latifolium (Hook.) Piper—Hills, N. W. Corvallis, April 5, 1914, 1290, May I, 1915, 3049, April 13, 1912, F. D. Bailey, root; Moist Woods (Corvallis?), April, 1897, Moses Craig. 129. PuccInIA MENTHAE Pers. Syn. Fungi 227. 1801. On LABIATAE: Mentha canadensis L.—Corvallis, Aug. 10, 1910, 1167, Nov. 4, EOrieereze:. (Clatsicanie, Columbia :Co:, Oct.6, 1914, ©. >) Bailey, 3090; Portland, Aug. 21, 1915, E. Bartholomew (Barth. Fungi Columb. 4908). Mentha canadensis lanata Piper—Toledo, Lincoln Co., Sept 25, 1o1r, FE. D: Bailey, 1762. 18 256 BROOKLYN BOTANIC GARDEN MEMOIRS Mentha piperata L.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 255. Mentha spicata L.—Hood River, Aug. 6, 1914, 1482; Elk City, Lincoln Co., Aug. 20, 1914, 3217; Portland, April, 1914, comm. 3208; The Dalles, Wasco Co., July I, 1914, 7340. 130. PUCCINIA MESOMEGALA B. & C. in Peck, Rep. N. Y. State Mus. 25 ¢ Lite UO73; Dicaeoma mesomegalum Kuntze, Rev. Gen. Pl. 3: 469. 18098. ON CONVALLARIACEAE: Clintonia uniflora Kunth.—Mt. Hood, Sept. 1, 1901, E. W. D. Holway, 1016, Road to Mt. Hood, Aug. 7, 1914, 1601; Klamath Co., Oct. 7, 1903, E. B. Copeland (Sydow, Ured. 1776); Bridal Veil, Multnomah Co., Aug. 11, 1910, 1079; Parkdale, Hood River Co., March 20, 1915, L. Childs, 3788; Sumpter, Baker Co., July 16, 1913, J. R. Weir, 276. This very distinct micro-form is very common in the mountains of the northwestern states on the above host, and on C. borealis in the northern tier of states from New Hampshire to Minnesota and in Canada. 131. PUCCINIA MICROMERIAE Dudley & Thomp. Jour. Myc. ro: 54. 1904. On LABIATAE: Micromeria chamissonis (Benth.) Greene (M. Douglasii Benth.)— Mary’s Peak, Benton Co., June 20, 1910, 3163; Corvallis, June, 1910, 1157, May 4, 1912, F. D. Bailey, 1734; Philomath, April 20, 1912, FD, Bailey, 7736; Jan. 6, 1914, 7754; ‘Bugene, Lane Co.,, July, 1914, G. B. Posey, 1292; N. slope Mt. Hood, Aug. 7, 1914, 2560; Washington Co., July—Aug. 1897, Moses Craig; Grant’s Pass, Jose- phine Co., Sept. 3, 1916, J. R. Weir, 252. 132. PUCCINIA MICROSORA K6rn.; Fuckel, Fungi Rhenani 2637. 1874. ON CYPERACEAE: Carex mirata Dem.—Clatsop, Clatsop Co., Nov. 7, 1913, 1190. Otherwise known only locally from the eastern United States. The aecial stage is unknown. 133. Puccinta MILLEFOLII Fckl. Sym. Myc. 55. 1869. On CARDUACEAE: Achillea millefolium L.—Philomath, May 10, 1914, 1834; Yaquina River, Elk City, Lincoln Co., Aug. 20, 1914, 1625; Hood River, July 22 TOUS 270: This micro-form, evidently introduced from Europe, is otherwise JACKSON: UREDINALES OF OREGON 257 known only from a few collections made in California and single collections from Montana and New Mexico. 134. PUCCINIA MONARDELLAE Dudley & Thomp., Jour. Myc. ro: 53. 1904. ON LABIATAE: Monardella odoratissima (Benth.) Greene—Hilgard, Union Co., July 10, 1914, 7531. Monardella villosa Benth.—Ashland, Jackson Co., Sept. 10, 1914, 2542; Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 2534. 135. PUCCINIA MONOICA (Pk.) Arth. Mycologia 4: 61. 1912. Aecidium monoicum Peck, Bot. Gaz. 4: 230. 1879. On PoaceseE: III. Koeleria cristata (L.) Pers.—Austin, Grant Co., Aug. 1915, J. R. Weir, 237. This species has aecia on Arabis as has been shown by Arthur (Mycol. 4:59. 1912; 7:75. 1915). No aecial collections have been seen from Oregon but this stage undoubtedly occurs in the eastern part of the state. The collection cited under P. Holboellii (cf. 117) was said to have aecia on a part of the original specimen which un- doubtedly are to be referred here rather than to A. Barbareae DC. as was done by Vize in the original notice of P. Barbareae. 136. PUCCINIA MONTANENSIS Ellis, Jour. Myc. 7: 274. 1883. Aecidium Hydrophylli Pk. Bull. Buff. Soc. 1: 68. 1873. Aecidium Phaceliae Pk. Bull. Torrey Club 11: 50. 1884. Aecidium Mertensiae Arth. Bull. Torrey Club 31: 6. 1904. ON HypDROPHYLLACEAE: I. Hydrophyllum albifrons Heller—Corvallis, April 5, 1914, 1289; Mary’s Peak, Benton Co., May 21, 1915, 3029. Hydrophyllum capitatum Dougl.—Near Crystal Lake, Corvallis, May 20, 1899, E. R. Lake, z6o00. Hydrophyllum tenuipes Heller—Corvallis, April, 1910, 1111, 3105; April 15, 1912, F. D. Bailey, 1724. Hydrophyllum sp.—Horse Creek Canton, Wallowa Co., May 14, 1897, E. P. Sheldon, Sogo. Mertensia laevigata Piper—Jefferson Lake, Marion Co., Aug. 1892, Moses Craig; Parmelia Lake, Cascade Mts., July 2, 1914, J. H. Corsaut, 2554. ON BORAGINACEAE: I. Phacelia heterophylla Pursh—Philomath, April 20, 1912, 1168. Phacelia leucophylla Torr.—Austin, Grant Co., Aug. 1915, J. R. Werir, 155. 258 BROOKLYN BOTANIC GARDEN MEMOIRS ONSPOASCEAR: 1) TIT. Elymus glaucus Buckl.—Glendale, Douglass Co., July 17, 1914, 1347. The aecial stage of this rust is very common in western Oregon particularly on Hydrophyllum. The uredinial and telial stages are doubtless much more common than the single record above would indicate. The species is very difficult to separate in the uredinial stage from P. Clematidis (cf. 85) and it is probable that some of the collections referred to that species belong here. Arthur (Mycol. 8: 139. 1916) sowed aecia from Hydrophyllum capitatum on Agropyron tenerum and Elymus virginicus. On the for- mer uredinia and telia developed, and on the latter a few uredinia only. This is the only successful culture with this species, though aecia on other Hydrophyllaceae and on Boraginaceae are referred here on morphological grounds. ; 137. PUCCINIA MUTABILIS Ellis & Gal. Jour. Myc. 5: 67. 1889. On ALLIACEAE: Allium Geyert Wats.—Blue Mts., July 5, 1897, W. C. Cusick, 1627. The writer is indebted to Professor Holway for the specimen on which this record is based. 138. PUCCINIA MADIAE Syd. Monog. Ured. I: 121. 1902. ON CARDUACEAE: Madia elegans Don.—Corvallis, June, 1910, 2679. Madia glomerata Hook.—Corvallis, Aug. 1899, E. R. Lake. Madia sp.—Hood River, June 20, 1914, 3349. This species is very close to, and possibly identical with, P. Hemi- zontae (cf. 113). 139. PUCCINIA OBSCURA Schroet., Pass. Nuov. Giorn. Bot. Ital. I, 02256) 13877. Aecidium Bellidis Thiim. Fungi Austr. 635. hyponym. 1873. Puccinia Bellidis Lagerh. Bol. Soc. Broter. 8: 134. 1890. On JUNCACEAE: Juncoides parviflorum (Ehrh.) Coville—Ashland, Jackson Co., Sept, 10, Lord, 2570;sJbend wrook iCo., Sept: 11, TO1L6, Rem 200; Ukiah, Umatilla Co., Aug. 21, 1903, M. A. Crosby. No culture work has been conducted in America. Plowright (Jour. Linn. Soc. Lond. 20: 511. 1884) has shown the aecia to be A. Bellidis, having cultured the species in both directions. Other Euro- pean workers have confirmed Plowright’s results (Klebahn, Die Wirtsw. Rostp. 317. 1904). JACKSON: UREDINALES OF OREGON 259 140. PUCCINIA OBTECTA Pk. Bull. Buff. Soc. Nat. Hist. 1: 66. 1873. Aecidium compositarum Bidentis Burrill; DeToni in Sacc. Syll. Fung. 7: 799. 1888. ON CYPERACEAE: Scirpus americanus Pers. (S. pungens Vahl.)—Westfall, Malheur Co., Aug. 1901, Griffiths & Carter (Griffiths, W. Am. Fungi 353). Arthur (Jour. Myc. 14: 20. 1908) has cultured this species on Bidens. Using telia on S. americanus from Indiana successful infec- tion resulting in pycnia and aecia was obtained on B. frondosa and B. connata. Aecia have not been collected west of the Rocky Mt. region. The above collection was issued as P. canaliculata, which is now interpreted as occurring only on Cyperus having aecia on Xan- thium and is unknown in Oregon. 141. Puccinia Ortonii sp. nov. O. Pycnia few, imperfectly known. I. Aecia chiefly hypophyllous, gregarious, in roundish or elongated groups 6-8 mm. across, short cupulate, 0.2-0.3 mm. in diameter; peridium yellowish, the margin erose; peridial cells oblong or rhombic, 20-26 by 29-35 u, slightly overlapping, the outer wall finely striate, 8-10 uw thick, the inner verrucose or slightly tuberculate, 4-6 wu thick; aeciospores globoid or broadly ellipsoid, 18-19 by 19-24 yu, wall color- less, I-1.5 uw thick, very closely and finely verrucose. II. Uredinia amphigenous, scattered, round, 0.5-I mm. across, tardily naked, cinnamon brown, cinereous when covered, ruptured epidermis conspicuous, pulverulent; urediniospores broadly ellipsoid (or when young obovoid), 19-26 by 23-32 uw, wall cinnamon brown, 2-3 uw thick, moderately and very minutely and obscurely echinulate; pores 3-5, scattered. III. Telia amphigenous, scattered, round, 0.2-I mm. across, tardily naked, ruptured epidermis conspicuous, chestnut brown, cinereous when covered, pulvinate, somewhat pulverulent; teliospores broadly and somewhat angularly ellipsoid, 18-26 by 30-42 uw, rounded at both ends, slightly or not constricted at the septum, wall chestnut brown, 1.5-2.5 w thick, uniform, smooth, with hyaline papilla over pore of apical cell which is usually at the apex but occasionally placed laterally, pore of lower cell varying in position from near the pedicel to the septum; pedicel deciduous, colorless. ON PRIMULACEAE: Dodecatheon Hendersonu leptophylla Suks.—Lake of the Woods, Cascade Range, Aug. 1892, Moses Craig. The above collection bears aecia and uredinia only. This species differs from P. melancoides (cf. 128) in the presence of uredinia in the life cycle. It is to be regarded as a correlated form with that species. 260 BROOKLYN BOTANIC GARDEN MEMOIRS The species is dedicated to Prof. C. R. Orton who was the first to separate the material from the opsis-form. The following is a list of the specimens from other localities in the Arthur herbarium. Dodecatheon alpinum Greene—Susanville, California, 5,000 ft., June 30, 1897, II, 111i, M. E. Jones; Bluff Lake, San Bernardino Mts., California, 7,400 ft., Sept. 1895, III, Miss Nora Pettibone, 2653; Mt. Eddy, Siskiyou Co., California, Sept. 7, 1903, i, 1, III, E. B. Copeland, (Sydow, Ured. 1774, type). Dodecatheon Jeffreyi Van Houtte—South of Sitka, Alaska, ui, ITI, Aug. 29, 1916, J. P. Anderson, 337; Vancouver Island, British Colum- bia, Aug. 26, 1908, 11, III, E. W. D. Holway. Dodecatheon (tetrandrum Suks.?)—Mt. Adams, Washington, 6,000-— 7,000 ft., Aug. 31, 1886, W. N. Suksdorf (Barth. N. Am. Ured. 554,1457). 142. PUCCINIA OxyRIAE Fckl. Symb. Nachtr. 3: 14. 1875, ON POLYGONACEAE: Oxynia digyna (L.) Hill—Strawberry Mt., 8,000 ft., Grant Co., Septy2, 1913, WE, Lawrence, 1m: Evidently a rather rare species represented in the Arthur her- barium otherwise only by single collections from Colorado, Utah, Idaho, Alberta and British Columbia on the above host. 143. Puccinia PALMERI D. & H. Erythea 7: 98. 1899. (Not Aecidium Palmert And. 1891.) Allodus Palmeri Orton, Mem. N. Y. Bot. Gard. 6: 202. 1916. (Not A. Palmert Arth. 1906.) ON SCROPHULARIACEAE: Pentstemon Menziesit Hook.—Near Mt. Jefferson, Linn Co., July 3, 1914, F. D. Bailey, 3046; Mt. Hood, 6,0c0 ft., Sept. 1, 1901, E. W. D. Holway; Horse Creek, Wallowa Co., June 24, 1897, E. P. Sheldon, S368. This is an opsis-form common in the Rocky Mt. and north Pacific states. Dietel & Holway (1. c.) based their combination on Aecidium Palmert Anderson. This Aecidium, as has been recently determined by Orton, working in this laboratory, is the aecial stage of the heter- oecious rust P. Andropogonis Schw. and not the aecia of this species. While the name P. Palmeri D. & H. has been misapplied in this way it seems best to retain it for this species, particularly since the telia described apply to this fungus. 144. PUCCINIA PARKERAE Diet. & Holw. Erythea 3: 78. 1895. ON SAXIFRAGACEAE: Ribes lacustre (Pers.) Poir.—Whitewater Ranger Station, near Mt. Jefferson, Aug. 28, 1916, H. P. Barss, 3395; Hood River, July 23 1OR5, 7076. JACKSON: UREDINALES OF OREGON 261 This short-cycled form possesses teliospores which closely resemble the telia of the Carex rusts having aecia on Ribes (cf. P. Grossulariae, 109) as has been pointed out by Holway (N. Am. Ured. 1: 53. 1906). It is not to be confused with the rust having a similar life history in the eastern United States, P. Ribis DC. The latter has verrucose spores, while in the one under discussion the spores are smooth. 145. PUCCINIA PATTERSONIANA Arth. Bull. Torrey Club 33:29. 1906. ON POACEAE: Agropyron spicatum (Pursh) Rydb.—Dufur, Wasco Co., June 19, 1914, 71398; Hilgard, Union Co., July 10, 1914, 1364; Grant's Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 208. The aecial form of this distinct heteroecious rust is unknown. 146. PucciniA Peck (DeToni) Kellerm. Jour. Myc. 8: 20. | 1902. Aecidium Oenotherae Pk. Ann. Rep. N. Y. State Mus. 23: 60. 1873. (Not P. Oenotherae Vize, 1877.) Aecidium Peckit DeToni, in Saccardo Syll. Fung. 7: 790. 1888. ? Puccinia ludibunda E. & E. Proc. Phil. Acad. 1893: 153. 1893. ON CYPERACEAE: Carex Hookeriana Dewey—Springbrook, Yamhill Co., May 14, 1914, F. D. Bailey, 30175; Whitewater Station near Mt. Jefferson, Aug. 12, 1914, H. P. Barss & G. B. Posey, 3006. The above collections are referred somewhat doubtfully to this species as no aecia have been collected in the Pacific northwest. The aecia occur on Onagraceae as was first shown by Kellerman (1. c.) and later by Arthur (Bot. Gaz. 33:13. 1903; Jour. Myc. 8: 52. 1OO2 ETA 55. LOO; £2405. 19065) 13; 195. Too7eeMiycol:. 13233. HOOO;) 2-222) “TO1O;- 4255: “1o12): 147. PuccINIA PENTASTEMONIS Pk. Bull. Torrey Club 12: 35. 1885. ON SCROPHULARIACEAE: Pentstemon diffusus Dougl.—Bridal Veil, Multnomah Co., May 18, EGI5, 3207. 148. PUCCINIA PIMPINELLAE (Str.) Mart. Fl. Mosq. Ed. II: 226. 1817. Uredo Pimpinellae Strauss, Wett. Ann. 2: 102. 1810. Puccinia Osmorrhizae Cke. & Peck, in Peck, Rep. N. Y. State Mus. 2073. SIS 78- Puccinia trifoliata E. & E. Bull. Torrey Club 22: 58. 1895. ON UMBELLIFERAE: Osmorrhiza brevipes (Coulter & Rose) Suks.—Corvallis, May 4, 1912, F. D. Bailey, 2550, July 3, 1914, G. B. Posey, 1998, Apr. 28, 1915, 3313; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 196; Austin, Grant Co., Aug. 1915, J. R. Weir, 780. 262 BROOKLYN BOTANIC GARDEN MEMOIRS Osmorrhiza divaricata Nutt.—Jackson Co., Sept. 7, 1903, E. B. Copeland (Sydow, Ured. 1778); Mt. Hood, Hood River Co., Aug. 7, 1914, 3087. Osmorrhiza Liebergit (Coulter & Rose) Suks.—North of Mt. Jefferson, Aug. 28, 1916, H. P. Barss, 3400. Osmorrhiza occidentalis Nutt.—Paisley, Lake Co., Aug. 1914, J. C. Elder, 3783. 149. PucciniA PIPERI Ricker, Jour. Myc. 11: 114. 1905. On PoacEAE: II and III. Festuca pacifica Piper—Eight Dollar Mt., Oregon, June 12, 1904, C. V. Piper, 6502, type. This species is evidently rare, since it is known only from the type locality listed above and from two localities in California. The life history is unknown. 150. PUCCINIA PLUMBARIA Pk. Bot. Gaz. 6: 238. 1881. Aecidium Giliae Pk. Bot. Gaz. 4: 230. 1879. (Not P. Giliae Hark. 1884.) Allodus Giliae Orton, Mem. N. Y. Bot. Gard. 6: I99. 1916. ON POLEMONIACEAE: Gilia gracilis (Dougl.) Hook.—Mary’s Peak, Benton Co., May 21, 1915, 3310; Hood River, May 14, 1914, 1526, 2514, June 9, 1915, 3274. Phlox speciosa Pursh—Sherman, Sherman Co., ‘July 1, 1914, 2515. I51. PUCCINIA POCULIFORMIS (Jacq.) Wettst. Verhl. Zool. Bot. Ges. Wein 35: 544. 1885. Lycoperdon poculiforme Jacq. Coll. Austr. 1: 122. 1786. Aecidium Berberidis Pers. in Gmel. Syst. Nat. 2: 1473. 1791. Puccinia graminis Pers. Neues Mag. Bot. 1: 119. 1794. Puccinia Phlei-pratensis Erikss. & Henn. Zeit. f. Pflanzenkr. 4: 140. 18094. Uredo quinqueporula Arth. & Fromme, Torreya 15: 265. I915. On Poaceae: II, III. A gropyron dasystachyum (Hook.) Vasey—Sherman Sta. O. W.R.R. & Nav. Co., Sherman Co., July 1, 1914, 1420. A grostis exarata Trin.—Ashland, Jackson Co., Sept. 10, 1914, 1567. Alopecuris californicus Vasey—Corvallis, Sept. 21, 1914, 1552. Avena fatua glabrata Peterm.—Philomath, Jan. 16, 1914, 7138. Avena satwa L.—Corvallis, Aug. 12, I91I1, 3761, July 6, tor4, 1667, Aug. 13, 1914, 1661, Aug. 9, 1915, G. H. Godfrey & F. D. Bailey, 3135; Cottage Grove, Lane Co., July 14, 1914, 1674; Pleasant Hill, EaneiCo., Augi7. T9lg.be De bavlevaumnso: JACKSON: UREDINALES OF OREGON 263 Beckmannia erucaeformis (L.) Host.—Corvallis, Aug. 8, 1915, 30206¢a. Dactylis glomerata L.—Cottage Grove, Lane Co., July 14, 1914, 1670; Corvallis, July 6, 1914, 1666; Clatsop, Clatsop Co., Nov. 3, 1913, 1155; Philomath, Jan. 6, 1914, 1084. Deschampsia elongata (Hook.) Munro—Glendale, Douglass Co., July 17, 1914, 7410; Ashland, Jackson Co., Sept. 10, 1914, 1566, 1568; Wren, Benton Co., June 26, 1914, 7329; Garden Home, Multnomah Moenlialy 20, TOTS, 3757; 3156. Elymus glaucus Buckl.—Philomath, Jan. I, 1914, 11752; Bend, Crook Co., Sept. 11, ro1r6, J. R. Weir, 277. Festuca elatior L.—Wren, Benton Co., June 26, 1914, 1324; Cottage Grove, Lane Co., July 14, 1914, 1348; Corvallis, July 29, 1915, 31790. Festuca megalura Nutt.—Corvallis, June 24, 1914, 1390, Aug. 13, 1914, 3191. Festuca myuras L.—Cottage Grove, Lane Co., July 14, 1914, 1351. Festuca pacifica Piper—Corvallis, July 6, 1914, 1434, July 29, 1914, 1410. Hierochloe macrophylla Thurb.—Glendale, Douglass Co., July 17, 1914, 1471 (type of Uredo quinqueporula). Hordeum distichon L.—Corvallis, Aug. 20, 1915, H. P. Barss, 3784. Hordeum vulgare L.—Corvallis, Oct. 5, 1914, 3165. Lolium multiflorum Lam.—Cottage Grove, Lane Co., July 14, 1914, 7349; Corvallis, Sept. 20, 1914, 1551; Tualatin, Washington Cory |uly-10,.1914, PF: D. Bailey, 1357. Lolium subulatum Vis.—Corvallis, March 22, 1915, 3277. Phleum pratense L.—Cottage Grove, Lane Co., July 14, 1914, 1668; Hood River, June 19, 1914, 1665; Springbrook, Yamhill Co., June 22 tO De Bailey, 3770; Briton, Lincoln Co; July 17, 1915; VanGundia, 3086, 3057; Corvallis, Aug. 13, 1914, 1662; Philomath, Janek, 1604 7757> Gresham, Multnomah Co., Aug:7, 1913, Bo D: Bailey, 1740; Sumpter, Baker Co., June, 1913, J. R. Weir, 97; Grant’s Pass, Josephine Co., Sept. 2, 1916, J. R. Weir, 246. Triticum compactum Host.—Moro, Sherman Co., June II, 1915, F. K. Ravn & A. G. Johnson. Triticum vulgare L.—Corvallis, July 29, 1914, 1684; Hood River, AUS. 5, 1o14, 216090; Ashland, Jackson’ Co., Aug. 28, 1913; 1725; Union, Union Co., Aug. 13, 1915, F. D. Bailey, 3733; Albany, Linn Co., Aug. 22, 1900, E. B. Townsend, 3384. Since the classic researches of DeBary, who first demonstrated heteroecism in rusts by showing that this species has aecia on Berberis, this rust has received more attention on the part of investigators than any other species. (Klebahn, Die Wirtsw. Rostp. 205-235. 1904.) 264 BROOKLYN BOTANIC GARDEN MEMOIRS In America the most important work has been conducted by Carleton (Div. Veg. Phys. & Path. U.S. D. A. Bull. 16. -1899; Bug: Pl; Industry, U. S. D2 A. Bull.63. 1904); Arthur Jour, Mycex sage LQOD SV PU 5 7.9 1905; 122L7-. “19005 1325108... 19075414 = TOs Mycol. 2: 227. 1910; 4:18. 1912); Freeman & Johnson (Bur. PI. Ind. U.S. D. Ay Bull. 216; 1011); Stakman (Minn, Exp:Staseulk 138. 1914; Jour. Agr. Research 4: 193-199. 1915) and Stakman and Piemeisel (Jour. Agr. Research 6: 813-816. 1916; 10: 429-495. I9I7). In Oregon the rust is apparently not as important on wheat and other grains as it is in the spring wheat districts east of the Rocky mountains. No aecial collections have been made. A number of unrecorded hosts appear in the above list. Since the publication of Uredo quinqueporula by Arthur and Fromme (I. c.), telia have been found on a duplicate specimen which determines that that species is properly referred here. The number of pores in the uredospores of that collection is unusual for this rust, the usual number being 4, and in the absence of telia was considered of sufficient impor- tance to separate it as a distinct species. 152. PUCCINIA POLYGONI-ALPINI Cruchet & Mayor, Bull. Herb. Bois 8: 245. 1908. ON POLYGONACEAE: Rumex paucifolius Nutt.—Crater Lake, Klamath Co., Sept. 9, 1OTo, J Re Weir 253: This specimen is referred to the above species on account of the hyaline umbo covering the pore of the apical cell in the teliospore. The species is described from material on Polygonum alpinum from Europe with which our material closely agrees. A collection on that host from Idaho is also to be referred here. The rust is unlike any other recorded on Rumex. The only other collection recorded on this host from North America is the one on which P. uniformis Pammel & Hume from Wyoming was based, which Holway (N. Am. Ured. 1: 36. 1906) considers to be on Polygonum sp. and refers to P. Bistortae. 153. PUCCINIA POLYGONI-AMPHIBII Pers. Syn. Fung. 227. 1801. Aecidium Geranit maculati Schw. Schr. Nat. Ges. Leipzig 1: 67. 1822. ON POLYGONACEAE: ‘ Polygonum amphibium L.—Brandt’s Ranch, Wallowa Valley, Aug. 26, 1897, E. P. Sheldon, 8972; The Dalles, Wasco Co., Aug. 25, 1915, E. Bartholomew (Barth. N. Am. Ured. 1566). Polygonum Muhlenbergii S. Wats. (P. emersum Britt.)—The Dalles, Wasco Co., Aug. 26, 1915, E. Bartholomew (Barth. Fungi Columb. JACKSON: UREDINALES OF OREGON 265 4762); Portland, Aug. 21, 1915, E. Bartholomew (Barth. Fungi Columb. 4567). Polygonum pennsylvanicum L.—Corvallis, Sept. 20, 1914, 1547; Clatskanie, Columbia Co., Oct. 10, 1914, F. D. Bailey, 1944. No collections of aecia referable to this species have been made west of the Rocky mountains. Tranzschel (Centr. f. Bakt. II, 11: 106. 1903) was the first to show that this species has aecia on Gera- nium. Arthur working with American material has confirmed Tranzschel’s results (Jour. Myc. 11: 59. 1905;12: 18. 1906). 154. PUCCINIA PORPHYROGENITA Curt.; DeToni in Sacc. Syll. 7: 703. 1888. Puccinia porphyrogenita Curt. in Thiim. Myc. Univ. 545 (hypo- nym). 1876. Puccinia acuminata Pk. Rep. N. Y. State Mus. 23: 57. 1872. (Not P. acuminata Fckl. 1869.) ON CORNACEAE: Cornus canadensis L.—Near Mt. Jefferson, Aug. 1892, Moses Craig; Larch Mt., Multnomah Co., Aug. 11, 1910, 1078; South Mt. Jefferson, Linn Co., July 3, 1914, F. D. Bailey, 7840; Mt. Hood, Aug. 7, 1914, 1604; Trail to Hanging Valley, Mt. Jefferson, Aug. 11, 1914, H. P. Barss & G. B. Posey, 1622. 155. PUCCINIA PROCERA Diet. & Holw. Erythea 1: 249. 1893. On POACEAE: Elymus arenicola Schrib. & Smith—Umatilla, Umatilla Co., May II, 1915, 3200, 3201, July 11, 1914, 1374, 1375; Sherman Sta. O. W. R. R. & Nav. Co., Sherman Co., July 1, 1914, 1421. This species is distinguishable from other forms on Elymus by the large urediniospores, 26-32 by 32-48 uw. The aecial connection is unknown. 156. PUCCINIA puNcTATA Link, Ges. Nat. Freunde Berlin Mag. 7: 30. 1816. On RUBIACEAE: Galium aparine L.—Hood River, July 24, 1915, 3225; Ashland, Jackson Co., Sept. 10, 1914, 3239; Corvallis, May 1, 1915, 3148. Galium asperrimum A. Gray—Big Canon, Wallowa Co., Aug. 24, 1897, E. P. Sheldon, 8774. Galium triflorum Michx.—Mary’s Peak, Benton Co., Aug. 15, 1914, 1513, 1515; Elk City, Lincoln Co., Aug. 20, 1914, 2528; Hood River, July 24, 1915, 3224. Galium sp.—Philomath, April 20, 1914, F. D. Bailey, 2570; Cor- vallis, April 8, 1914, 7524; Dufur, Wasco Co., June 30, 1914, 1335. 266 BROOKLYN BOTANIC GARDEN MEMOIRS 157. PUCCINIA PYGMAEA Erikss. Fungi Par. Scand. 9: 449. 1895. ON POACEAE: Calamagrostis aleutica Bong.—Newport, Lincoln Co., July 18, 1915, 3204, Aug. 30, 1914, 1579. The above collections show uredinia only. The aecial connection is unknown. 158. PUCCINIA RECEDENS Syd. Monog. Ured. 1: 146. 1902 ON CARDUACEAE: Senecio harfordi Greenman—Bridal Veil, Multnomah Co., May 13, TOT5303275- Senecio sp.—Hilgard, Union Co., July 10, 1914, 1542. 159. PuccINIA RHAMNI (Pers.) Wettst. Verhl. Zool.-Bot. Ges. Wein 35: 545. 1885. Aecidium Rhamni Pers. in Gmel. Syst. Nat. 2: 1472. 1791. Puccinia coronata Corda, Icones I: 6. 1837. ON RHAMNACEAE: I. Rhamnus purshiana DC.—Corvallis, July 5, 1911, F. D. Bailey, T1135, May.0, 1914; 7627, May 12, 1914, 1277, July 5, Tor4sheeee Barss, 1940; Hood River, May 14, 1914, 1278; Clatskanie, Columbia Co., May 20, roi4, F. D. Bailey, r2é7; Cottage Grove, Lanes@ae May I, 1915, C. E. Stewart, 3058. On Poaceae: II, III. Agrostis alba L.—Toledo, Lincoln Co., July 19, 1915, 3779. Agrostis alba maritima Meyer—Philomath, Jan. 6, 1914, 1749, 1150. Agrostis exarata Trin.—Corvallis, Feb. 14, 1914, 3098, June 29, 1914, G. B. Posey, 1300, 1306, Sept. 20, 1914, 7553, Sept. 5, 19Res 1576, Wee, 1915, G. B. Posey; Hood River, June 19, 1914; en: Philomath, June 26, 1914, 1344; Eugene, Lane Co., July 11, 1914, G. B. Posey, 1377; Newport, Lincoln Co., July 18, 1915, 3720; Garden Home, Multnomah Co., July 20, 1915, 3121; Portland, Aug. 21, 1915, E. Bartholomew, 5943. Agrostis foliosa Vasey—Yaquina, Lincoln Co., July 17, 1915, 3722. Agrostis longiligula Hitchc.—Jetty, Lincoln Co., July 19, I915, VanGundia, 3256. Agrostis microphylla Steud.—Wren, Benton Co., June 26, 1914, 1314; Cottage Grove, Lane Co., July 14, 1914, 1353; Corvallis, July 6, 1914, F. D. Bailey, 7436. Avena sativa L.—Newport, Lincoln Co., July 18, 1915, 3132; Briton, Lincoln Co., July 19, 1915, G. VanGundia, 3088; Marshfield, Coos Co., July, 1916, C. E. Owens; Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 3162. Calamagrostis canadensis (Michx.) Beauv.—Clatskanie, Columbia Co., May 20, 1914, F. D. Bailey, 758o. JACKSON: UREDINALES OF OREGON 267 Calamagrostis hyperborea Lange—Clatsop, Clatsop Co., Nov. 7, IQII, II00. Festuca elatior L.—Elk City, Lincoln Co., Aug. 20, 1914, 1380. Festuca subulata Trin.—Ashland, Jackson Co., Sept. 10, I914, 1501; Mary’s Peak, Benton Co., Aug. 15, 1914, 1572; Elk City, Lincoln Co., Aug. 20, 1914, 1382. Holcus lanatus L.—Canby, Clackamas Co., July 21, 1911, 1782; Philomath, Jan. 6, I914, 3115, 1133; Cottage Grove, Lane Co., July 14, 1914, 3714; Mouth of Salmonsberry River, Tillamook Co., July 17, 1915, G. VanGundia, 3085, 3129; Yaquina, Lincoln Co., July 17, 1915, 3112; Jetty, Lincoln Co., July 19, 1915, G. VanGundia, 3130; Eddyville, Lincoln Co., Aug. 10, 1915, Hoerner, 3090; Grant’s Pass, Josephine Co., Sept. 2, 1916, J. R. Weir, 227; Portland, Jan. 9, 1914, 71739; Elk City, Lincoln Co., Aug. 20, 1914, 1370. Lolium multiflorum Lam.—Near Gray Station, Linn Co., July 4, 1914, 1419; Corvallis, July 6, 1914, 1435, Sept. 20, 1914, 1554; New- port, Lincoln Co., July 21, 1915, 3128. Lolium perenne L.—Corvallis, Aug. 3, 1914, 1412, Sept. I, 1914, Qi27. Panicularia elata Nash—Clatskanie, Columbia Co., Aug. I1, 1913, F: ®. Bailey, rro5. Panicularia pauciflora (Presl.) Kuntze—Orenco, Washington Co., June 13, 1914, 1388; Neah-Kah-Nie Mt., Tillamook Co., Sept. 17, 1915, F. D. Bailey, 3258; Portland, Aug. 21, 1915, E. Bartholomew, 5942 (Barth. Fungi Columb. 4973). This coronate-spored grass rust is evidently very common through- out western Oregon on native grasses. It is, however, not common in the Willamette valley on oats. All of the collections on that host are from near the sea coast. DeBary (Monat. Akad. Wiss. 211. 1866) was the first to conduct cultures indicating the genetic connection with aecia on Frangula and Rhamnus in Europe. Since that time many European investi- gators have conducted culture experiments (Klebahn, Die Wirtsw. Rostp. 254-262. 1904). In America this species has been cultured by Arthur (Bull. Lab. Nat. Hist. State Univ. Iowa 4: 398. 1898; Jour. Myc. 11:58. 1905; Mycol. 4: 18. 1912) and Carleton (Div. Veg. Phys. & Path. U. S. Dept. Agr. 16: 48. 1899; Bur. Pl. Industry, U. S. Dept. Agr. Bull. 63:15. 1904). The only culture made with Pacific coast material was made in 1916 in this laboratory under the writer’s direction, using telial material on Agrostis exarata sent to the writer from Corvallis by G. B. Posey. This was used to inoculate Rhamnus Purshiana, with 268 BROOKLYN BOTANIC GARDEN MEMOIRS the development of pycnia and aecia. This host is the only one on which aecia have been collected in Oregon, and they are very abundant, as the number of collections indicates. 160. Puccinia Romanzoffiae sp. nov. O. Pycnia not seen. III. Telia chiefly hypophyllous and petiolicolous, crowded on con- fluent groups, 0.5-I mm. across or covering extensive areas on the petioles, early naked, pulverulent, chestnut brown, ruptured epi- dermis noticeable; teliospores somewhat irregularly ellipsoid or oblong, 19-24 by 34-42 uw, rounded above and below, not or scarcely con- stricted; wall chestnut brown, 2-3 » thick, marked by large sparsely distributed irregular tubercles, thickened at apex by a low sub-hyaline umbo to 4-5 p, pore of lower cell at septum similarly thickened; pedicel colorless, short deciduous. ON HyDROPHYLLACEAE: Romanzoffia sitchensis Bong.—Mt. Jefferson, 8,ooo0 ft., Aug. 14, 19o04, HP: sBarss, 25905 ty pe: This species is distinguished from other species on this family of hosts by the character of the markings of the teliospore. In P. Hydrophylli Pk. the teliospores are closely and finely verrucose while in P. Phaceliae Syd. & Holw. they are smooth. The character of the teliospores in the latter species suggests a correlation with P. mon- tanensis (cf. 136) which has aecia on Phacelia and other members of the family Hydrophyllaceae. 161. PUCCINIA RUBEFACIENS Johans. Bot. Centr. 28: 394. 1886. ON RUBIACEAE: Galium boreale L.—Hilgard, Union Co., July I0, 1914, 1540; Austin, Grant Co., Aug. 1916, J. R. Weir, 247. The teliospores of this micro-form are very similar in shape and size to those of the opsis-form P. ambigua (cf. 61) and the eu-form P. punctata (cf. 156). These three species on Galium doubtless repre- sent a series of correlated forms. 162. PUCCINIA RUGOSA Billings, King’s Rep. 4oth Par. 914. 1871. (Not P. rugosa Speg. 1886.) Puccinia Troximontis Pk. Bot. Gaz. 6: 227. 1881. Puccinia Columbiensts E. & E. Proc. Phil. Acad. 1893: 153. 1893. ON CICHORIACEAE: Agoseris laciniata (Nutt.) Green—Corvallis, July 10, 1915, 3215. 163. PUCCINIA SAXIFRAGAE Schlecht. Fl. Berol. 2: 134. 1824. Puccinia curtipes Howe, Bull. Torrey Club 5:3. 1874. JACKSON: UREDINALES OF OREGON 269 ON SAXIFRAGACEAE: Saxifraga Marshall Greene—Hood River, May 16, 1915, 3268; Mary’s River, E. of Wren, Benton Co., April 17, 1915, 2617. Saxifraga odontoloma Piper—Corvallis, May 1, 1915, 3260. 164. PUCCINIA SHERARDIANA Koern. Hedw. 16: 19. 1877. Puccinia Malvastri Peck, Bull. Torrey Club 12: 35. 1885. On MALVACEAE: Sidalcea virgata Howell—Corvallis, May 31, 1892, A. T. Mulford, Reape. 12, 1912, F. D. Bailey, 3354, June 23, 1913, F. D. Bailey, 7126, Apne o, 1914, 3352, Apr. 29, 1914, G.. B. Posey, 3353, Apr. 30, 1915, 3071; Newburg, Yamhill Co., Apr. 9, 1915, F. D. Bailey, 3072. 165. PUCCINIA SIDALCEAE Holw. N. Am. Ured. 1: 67. 1907. On MALVACEAE: Sidalcea oregana Gray—Klamath Co., July 10, 1903, E. B. Cope- land, type. This collection was distributed as P. Sphaeralceae E. & E. in Sydow’s Uredineen 1782. 166. PuccINIA STIPAE Arth. Bull. Iowa Agr. College Dept. Bot. 1884: 160. 1884. On Poaceae: II and III. Stipa comata Trin. & Rupr.—Umatilla, Umatilla Co., July 11, 1914, 1369, May I1, 1915, 3205; Hermiston, Umatilla Co., May 12, 1915, 3200. This species has aecia on various genera of Carduaceae including Aster, Solidago, Grindelia and Senecio, as has been shown by Arthur Wout Nive. It: 63. 1905; Mycol.. 4: 19: 1912, 72 72:- I915)- No aecial collections have been made in Oregon, though that stage is doubtless not uncommon in the eastern part of the state (cf. 69). 167. PUCCINIA SUBNITENS Dietel, Erythea 3: 81. 1895. ? Aecidium Sarcobati Pk. Bot. Gaz. 6: 240. 1881. ON CHENOPODIACEAE: I. Sarcobatus vermiculatus (Hook.) Torr.—Eastern Oregon, Aug. 1902, D. Griffiths (Vestergren, Micro. Rar. Sel. 552). On Poaceae: III. Distichlis spicata (L.) Greene—LaGrand, Union Co., March, 1915, C. C. Cate, 3278; Umatilla, Umatilla Co., July 11, 1914, 1367, 1373; Moro, Sherman Co., Aug. 4, 1914, C. R. Ball, 1856. This remarkable species has aecia on a large number of hosts in the Polygonaceae, Chenopodiaceae, Amaranthaceae, Cruciferae, etc. as was first shown by Arthur (Bot. Gaz. 35: 19. 1903; Jour. Myc. Fi 54: HIOOs L216. +1906, 13: 197; 1907; 142 15. . 19085. Mycol. 270 BROOKLYN BOTANIC GARDEN MEMOIRS I: 234. 1900, 2: 225. 1910,'4: 18. 1912). Bethel (Phytopathasg 92-94. 1917) has also conducted very extensive cultures and proven the genetic connection with aecia on many hosts. In 1915, Arthur. (Mycol. 8: 135. 1916), using telial material sent by the writer collected by Mr. C. C. Cate at LaGrand,- Ore., obtained the development of aecia on Chenopodium album. This is the only culture made with material from the Pacific coast. The aecia on Sarcobatus are included here on the strength of culture work conducted by Arthur, in which he obtained aecia on that host using telial material on Distichlis from Nevada. The matter is complicated by the fact that Bethel (1. c.) finds that the aecia on this host in Colorado go to P. luxuriosa (cf. 124) and P. subnitens can not be made to infect Sarcobatus. It is possible that the two species repre- sent closely related biological forms. 168. PUCCINIA SYMPHORICARPI. Hark. Bull. Calif. Acad. Sci. 1: 35. 1884. ON CAPRIFOLIACEAE: Symphoricarpos albus (L.) Blake—Corvallis, Oct. 17, 1909, F. L. Griffin, 7047, July, 1910, 1086, Aug. 1910, 1050, May 4, 1912, 1087, May 19, 1913, F. D. Bailey, 7158; Sheridan, Yamhill Co., July 7, 1914, H. P. Barss, 1291, July 8, 1916, H. P. Barss, 3396; North slope Mt. Hood, Aug. 7, 1914, z611; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 234; Hood River, July 24, 1915, 3064. This micro-form is exceedingly abundant in western Oregon. This species is morphologically correlated with the telial stage of P. abun- dans (cf. 59), which has aecia on the same host, as has been pointed out by Travelbee (Proc. Ind. Acad. Sci. 1914: 233. 1915). 169. PuccintiA TARAXACI (Reb.) Plowr. British Ured. & Ustil. 186. 1889. Puccinia Phaseolt var. Taraxaci Reb. Fl. Noem. 356. 1804. ON CICHORIACEAE: Taraxacum Taraxacum (L.) Karst.—Corvallis, June, 1910, 1117; Bonneville, Multnomah Co., Aug. 10, 1910, 3078; Ashland, Jackson Co., Sept. 10, 1914, 3074; The Dalles, Wasco Co., July 1, 1914, 3073; Talent, Jackson Co., June 22, 1915, G. B. Posey, 3077; Newburg, Yamhill Co., Apr. 13, 1914, F. D. Bailey, 3076. 170. Pucctnta ToumMeyi Syd. in Sacc. Syll. Fung. 16: 299. Feb. 1902. Puccinia circinans Ell. & Ev. Bull. Torrey Club 27: 61. 1900. (Not P. circinans Fckl. 1869 or Dietel 1897.) Puccimia chasmatis Ell. & Ev. Jour. Myc. 8: 15. .May, 1902. JACKSON: UREDINALES OF OREGON yA ON SCROPHULARIACEAE: Penistemon sp.—Canyon City, Grant Co., Aug. 26, 1914, W. E. Lawrence, 3185. 171. PUCCINIA TRAUTVETTERIAE Syd. & Holw. in Sydow, Monogr. Wred= 5552. 1903. ON RANUNCULACEAE: Trautvetteria grandis Nutt.—S. W. slope Mt. Hood, July 23, 1915, os L. This interesting micro-form, known only from a few collections from the mountains of the northwestern states, has also been reported from Japan. 172. PUCCINIA TRITICINA Erikss. Ann. Sci. Nat. VIII. 9: 270. 1899. ON POACEAE: Triticum aestivum L.—Hood River, June 20, 1914, 1399. Triticum ovatum Rasp.—Myrtle Creek, Douglass Co., June 9, poral. 1). Bailey, 7407. Triticum vulgare L.—Lebanon, Linn Co., Aug. 2, 1913, F. D. Bailey, 1126; Myrtle Creek, Douglass Co., June 9, 1914, F. D. Bailey, 1941; Cottage Grove, Lane Co., July 14, 1914, 1676; Corvallis, July 6, 1914, B50. 3100; july 10, 1914, F. D. Bailey, 1677, July 20, 1914, 1685; Bend, Crook Co., Sept. 11,, 1916, J. R. Weir, 207: This, the common leaf rust of wheat, is very abundant in western Oregon. The life history is unknown. In morphological characters, it resembles closely the forms on native grasses commonly referred to P. rubigo-vera, most of which are now included in P. Clematidis (cf. 85). 173. PUCCINIA UNIVERSALIS Arth. Jour. Myc. 11: 21. 1908. Aecidium Dracuncult Thiim. Bull. Soc. Nat. Moscow. 58: 212. 1878. (Not P. Dracuncult Auers. 1850.) On CARDUACEAE: I. Artemisia sp.—White Pine,Baker Co., June, 1913, J. R. Weir, 720. On CYPERACEAE: IJ, III. Carex multicaulis Bailey—Grant’s Pass, Josephine Co., May 5, 1887, Thomas Howell. - Carex praegracilis W. Boott. (C. marcida Boott.)—Redmond, Crook Co., July 2, 1914, 1425. Carex Rossii Boott.—Hood River, July 23, 1915, 3289. Carex umbellata Schk.—Hood River, July 23, 1915, 3282. This species has aecia on Artemisia as has been shown by Arthur (Jour. Myc. 14: 21. - 1908; Mycol. 2: 224. I910, 4: 16. I912). 174. PUCCINIA URTICATA (Lk.) Kern, Mycol. 9: 214. 1917. Aecidium Urticae Schum. Enum. PI. Saell. 2: 222. 1803. Caeoma urticatum Link, in Willd. Sp. Pl. 6: 62.. 1825. 19 Die BROOKLYN BOTANIC GARDEN MEMOIRS Puccimia Urticae Lagerh. Mitt. Bad. Ver. 2: 72. 1889. (Not P. Urticae Barcl. 1887.) Puccinia Garrett Arth. Bull. Torrey Club 32: 41. 1905. On URTICACEAE: I. Urtica Lyalla S. Wats.—Philomath, April 26, 1914, 1529, May Io, 1914, 2569; Corvallis, May 1, 1915, 3052. On CYPERACEAE: I], III. Carex Barbarae Dewey (C. laciniata Boott.)—Grant’s Pass, Josephine:Go., Sept. 3;,1916;-]. Ik. Weir; 250. Carex magnifica Dewey—Newberg, Yamhill Co., April 13, 1914, F. D. Bailey, 3009; Clatskanie, Columbia Co., May 20, 1914, F. D Bailey, 3002; Neah-Kah-Nie Mt., Tillamook Co., Sept. 17, 1915, PDE Barley23357- Carex nebraskensis Dewey—Andrews, Harney Co., Aug. I9go1, Griffiths & Morris (Griffiths, W. Am. Fungi 339). Carex rostrata Stokes (C. utriculata Boott.)—Clatskanie, Columbia Co., May 20, 1914, F: D. Bailey, 30710; Redmond, Crook Co;, july 1914, 1433. Carex sp.—Hood River, May 14, 1914, 3027. The connection of this Carex rust with aecia on Urtica was first shown by Magnus in 1872 (Vehr. Bot. var. Prov. Brandb. 14: 11. 1872). Many other European investigators have confirmed Magnus results (Klebahn, Die Wirtsw. Rostp. 293. 1904). In America Arthur has conducted numerous successful culture experiments with this species.(Bot. Gaz. 29: 270. 1900; 35: 16. 19033 > Jour: /Miye: 82527) 1902312 152 L906; 14. iia eee Mycol. 2: 223. 1910; 4:17. 1912). Kellerman has also conducted successful culture experiments (Jour. Myc. 9: 9. 1903). None of the culture work, however, has been conducted with Pacific coast material. 175. PUCCINIA VERATRI Duby, Bot. Gall. 2: 890. 1830. Puccinia Veratri Clinton, in Peck, Rep. N. Y. State Mus. 27: 103. 1875. ON ONAGRACEAE: I. Epilobium sp.—Parkdale, Hood River Co., May 14, 1914, 511. ONC DIMACEAS: T1LS0LI: Veratrum californicum WDurand—The Meadows, Wallowa Co., Aug. 18, 1897, E. P. Sheldon, 8774. Veratrum viride Ait.—Calloway Station, Benton Co., June 28, 1901, E. R. Lake, 1731; Parkdale, Hood River Co., May 14, 1914, 1279; Hilgard, Union Co., July 10, 1914, 1934. The aecial collection is referred here with some confidence. It was made in the immediate vicinity of Veratrum plants showing fresh JACKSON: UREDINALES OF OREGON Om Oo uredinia. The aecia were somewhat old and no uredinia were found on other Epilobium plants in the vicinity. Tranzschel (Ann. Myc. 7: 182. 1909) established the connection of aecia on Epilobium with P. Veratri, obtaining his clew from the close morphological resemblance of the teliospores of this species to those of P. Epilobii DC. Bisby (Am. Jour. Bot. 3: 527-561. 1916) has pointed out the morpho- logical similarity of this species with Uromyces plumbarius (cf. 201), P. Epilobu, P. Epilobu-tetragoni (cf. 99) and P. Epilobi-Fletscheri. 176. PUCCINIA VIOLAE (Schum.) DC. FI. Fr. 6: 62. 1815. Aecidium Violae Schum. Enum. PI. Saell. 2: 224. 1803. ON VIOLACEAE: Viola adunca J. F. Smith—Mary’s Peak, Benton Co., May 21, IQI5, 3223. Viola glabella Nutt.—Corvallis, Linn Co., April 16, 1912, ro8T; Corvallis; May 19, 1912, F. D. Bailey, 3708, April 29, 1914, F. D. Bailey, 3164, July 14, 1914, H. P. Barss, 2548; Hood River, May 14, 1914, 3197; Portland, Aug. 30, 1915, E. Bartholomew, 5978 (Barth. N. Am. Ured. 1677); Sumpter, Baker Co., July 16, 1913, J. R. Weir, 186; Mary’s Peak, Benton Co., Aug. 15, 1914, 2547. Viola nephrophylla Greene—Hilgard, Union Co., July 10, 1914, 2557- Viola rugulosa Greene—Horse Creek Canyon, Wallowa Co., June 4, 1897, E. P. Sheldon, 8258. Viola sp.—N. slope Mt. Hood, Aug. 7, 1914, 2553. 177. PUCCINIA WULFENIAE Diet. & Holw. Erythea 3: 79. 1895. Puccinia Syntheridis Ell. & Ev. Bull. Torrey Club 27: 61. Igoo. ON SCROPHULARIACEAE: i Synthyris rotundifolia Gray—Philomath, April 20, 1912, 1146; Corvallis, April 8, 1914,:1286. 178. TRANZSCHELIA PUNCTATA (Pers.) Arth. Résult Sci. Congr. Bot. Vienne 340. 1906. Aecidium punctatum Pers. Ann. Bot. Usteri 20: 135. 1796. Puccima Pruni-spinosae Pers. Syn. Fung. 226. 1801. ON ROSACEAE: Amygdalus Persica L.—Kiger’s Island, Benton Co., Oct. 5, 1913, C. M. Scherer, 7825. Prunus domestica L. (Italian Prune)—Salem, Marion Co., Aug. 1909; 7062; Yambhill’ Co; Sept. 9); 1911;-1040;~ Corvallis» Oct. 20, 1914, Gy By. Posey, p50: This is not an uncommon disease of the prune, though apparently doing little damage. It is less common on the peach. No aecial collections have been made in the northwest. 274 BROOKLYN BOTANIC GARDEN MEMOIRS Tranzschel (Trans. Bot. Acad. St. Petersb. 11: 67-69. 1905) was the first to culture this species showing that aecia occur on Anemone. In America Arthur (Jour. Myc. 12: 19. 1906; 13: 199. 1907) has shown that the aecia on Hepatica common in the eastern United States are genetically connected. 179. UROMYCES AEMULUS Arth. Bull. Torrey Club 38: 373. 1911. Nigredo aemula Arth. N. Amer. Flora 7: 241. 1912. On ALLIACEAE: Allium validum S. Wats.—Paisley, Lake Co., Aug. 1914, J. S. Elder, 7087. 180. UROMYCES AMOENUS Syd. Ann. Myc. 4: 28. 1906. On CARDUACEAE: Anaphalis margaritacea occidentalis Greene—Hood River, July 23, 1915, 3243; Crater Lake, Klamath Co., Sept. 9, 1916, J. R. Weir, 235. Anaphalis margaritacea subalpina Gray?—N. slope Mt. Hood, Aug: 75 1914, 7673. 181. URoMYCES ARMERIAE (Schlechtd.) Lev. Ann. Sci. Nat. III, 8: 375. 1847. Caeoma Armeriae Schlechtd. Fl. Berol. 2: 126. 1824. ON PLUMBAGINACEAE: Statice armeria L.—Newport, Lincoln Co., May 16, 1914, C. E. Owens, 1999, July 18, 1915, 3078. This species differs from U. Limonu in the shorter, broader telio- spores and the short mostly deciduous pedicel. The first collection mentioned bears aecia accompanied by uredinia, the second, uredinia and telia only. The rust is abundant on a cliff near the seashore. So far as we are aware this is the first record of this species in America. 182. Uromyces Beckmanniae sp. nov. O and I. Pyenia and aecia unknown. II. Uredinia amphigenous, scattered, elliptical, 0.5—-0.8 mm. long, soon naked, pulverulent, cinnamon brown, ruptured epidermis notice- able; paraphyses none; urediniospores globoid or broadly ellipsoid, 19-24 by 23-30 uw, wall colorless or pale yellow, 2—2.5 uw thick, finely verrucose-echinulate, pores 8-10, scattered. Ill. Telia amphigenous and culmicolous, scattered or crowded, oblong, 0.4-0.7 mm. across, often confluent to form crusts or lines, tardily naked, blackish brown; teliospores obovoid or ellipsoid, angular, 20-26 by 29-40 wu, apex rounded or angular, narrowed below; wall chestnut brown, I-2 yu thick, smooth, but showing distinct longi- tudinal ridges, apex thickened, 3-6 yu, pedicel colorless or slightly tinted next to the spore, equalling the spore or usually deciduous. JACKSON: UREDINALES OF OREGON 275 ON POACEAE: Beckmannia erucaeformis (L.) Host.—Corvallis, Sept. 21, I9gII, 1183; south Mary’s River, Sept. 30, 1914, 3144, Oak Creek, July 29, 1915, 3145 type, Aug. 14, 1915, 3026, May, 1916, G. B. Posey. Evidently the most common rust in Oregon on this host. It differs from U. Hordei, which is in general a southern form not known on the Pacific coast, in the larger teliospores which show distinct longi- tudinal ridges. From U. Jacksonii (cf. 192) this species differs in the thickened apices of the teliospores. No clue is available as to the aecial host. The rust is difficult to separate from Puccinia Rhamni (cf. 159) in the uredinial stage. 183. Uromyces BropIEAE Ell. & Hark., Harkness, Bull. Cal. Acad. Ser. 1:25. 1584: Uromycopsis Brodieae Arth. Result Sci. Congr. Bot. Vienne 345. 1906. On ALLIACEAE: Brodiaea sp.—Corvallis, May 4, 1912, F. D. Bailey, 3304, April 25, 1915, 2625. This opsis-form is evidently common in western Oregon. The rust is usually found attacking the tips of the leaves early in the spring. Aecia usually predominate, the telia being inconspicuous and easily overlooked. 184. UROMYCES CARNEUS (Nees) Hariot, Jour. de Bot. 7: 376. 1893. Aecidium carneum Nees; Funk. Krypt. Gew. Ficktelgeb. 25: 4. 1818. Uromyces lapponica Lagerh. Bot. Nat. 1890: 274. 1890. Uromycopsis lapponica Arth. Result Sci. Congr. Bot. Vienne 345. 1906. On LEGUMINOSAE: Astragalus sp.—Austin, Grant Co., Aug. 1915, J. R. Weir, 268. 185. UROMYCES CARYOPHYLLINA (Schrank.) Wint. in Rab. Krypt. FI. TAG, TSS. Lycoperdon caryophyllinum Schrank. Baier. Fl. 2: 668. 1789. Nigredo caryophyllina Arth. N. Am. Flora 7: 246. 1912. On CARYOPHYLLACEAE: Dianthus Caryophyllus L.—Portland, Sept. 30, 1912, F. D. Bailey, 1089, Dec. 19, 1912, F. D. Bailey, 1744; Corvallis, Dec. 1910, 3781. 186. Uromyces FABAE (Pers.) DeBary, Ann. Sci. Nat. IV, 20: 80. 1863. Uredo Fabae Pers. Neues Mag. Bot. I: 93. 1794. Nigredo Fabae Arth. N. Am. Flora 7: 251. 1912. 276 BROOKLYN BOTANIC GARDEN MEMOIRS ON LEGUMINOSAE: Lathyrus obovatus (Torr.) White?—Sumpter, Baker Co., July 16, 1916, J. R. Weir, 792; Austin, Grant Co., Aug. 1915, J. R. Weir, 797. Lathyrus oregonensis White—Andrews, Harney Co., Aug. Igo1, Griffiths & Morris (Griffiths, West. Am. Fungi 349a); Spencer Creek, Klamath Co., July 10, 1903, E. B. Copeland (Syd. Ured. 1764). Lathyrus pauciflorus Fern.2-—Klamath Falls, Klamath Co., Sept. 8, 1916; J. R. Weir, 203. Lathyrus polyphyllous Nutt.—Mt. Hood, Aug. 31, 1901, E. W. D. Holway. Lathyrus sulphureus Brewer—Corvallis, May 9, 1914, 3226; Ashland, Jackson Co., Sept. 10, 1914, 3320. Lathyrus sp.—Scotts, 7 mi. N. of Fort Klamath, Klamath Co., Sept. 20, 1913, E. P. Meinecke, Cr D 10; Glendale, Douglass Co., July 17, 1914, 1506; N. Mt. Hood, Aug. 7, 1914, 1490, Aug. 9, 1914, 1486; Whitewater Forest Station, Aug. 12, 1914, H. P. Barss, 3248; Garden Home, Multnomah Co., July 20, 1915, 3249. Vicia americana Muhl.—N. slope Mt. Hood, Aug. 7, 1914, 1489, Aug. 9, 1914, 1485; Corvallis, Sept. 21, 1914, 1545. Vicia linearis (Nutt.) Greene—Mary’s Peak, Benton Co., June 20, 1910, 1501; Philomath, June 20, 1910, 1502; Newberg, Yamhill Co., April 13, 1914, 1525; Springbrook, Yamhill Co., June 25, 1914, F. D. Bailey, 3238. Vicia truncata Nutt.—Bonneville, Multnomah Co., Aug. 10, Ig10, 1179; Hood River, May 14, 1914, 1527. Victa sp.—Dothan, Douglass Co., Sept. 8, 1914, G. B. Posey, 1550. 187. UROMYCES FALLENS (Desmaz.) Kern, Phytopath. 1: 6. 1911. Uredo fallens Desmaz. Pl. Crypt. 1325. 1843. Nigredo fallens Arth. N. Am. Flora 7: 254. Ig12. On LEGUMINOSAE: Trifolium pratense L.—Mary’s Peak, Benton Co., June, 1910, 3004; Springbrook, Yamhill Co., June 22, 1914, G. B. Posey, 3234; Corvallis; July “15, -to14G. Bs Posey 3003; Oct. 26, 1914 Ga Posey, 1983; Parkdale, Hood River Co., Aug. 5, 1914, 3701; Portland, Aug. 23, 1915, E. Bartholomew (Barth. Fungi Columb. 4788). 188. UROMYCES HETERODERMUS Syd. Ann. Mycol. 4: 29. 1906. On LILIACEAE: Erythronium parviflorum (Wats.) Goodding—Corvallis, April 13, 1912, F. D. Bailey, 1774. A short-cycle form not uncommon in the Rocky Mt. and Pacific coast regions. JACKSON: UREDINALES OF OREGON DE. 189. Uromyces Hotwayt Lagerh. Hedwigia 28: 108. 1899. Uromyces Lalu G. W. Clinton; Peck, Ann. Rep. N. Y. State Mus. 27: 103. 1875. (Not U. luli Kunze. 1873.) Nigredo Lalu Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. On LILIACEAE: Lilium parviflorum (Hook.) Holzinger—Wren, Benton Co., July, 1911, W. E. Lawrence, 1144; Hood River, May 9, 1915, 3044; May 16, 1915, 2661; Bridal Veil, Multnomah Co., May 18, 1915, 2659; Portland, June 21, 1915, 3060. 190. UROMYCES HyYPERICI-FRONDOSI (Schw.) Arth. Bull. Minn. Acad. WattsSGl. 272015. 0 E883. Aecidium Hyperici-frondosit Schw. Schr. Nat. Ges. Leipzig 1: 68. 1822. Nigredo Hyperici-frondosi Arth. Résult Sci. Bot. Vienne 344. 1906. ON HyYPERICACEAE: * Hypericum Scoulert Hook.—Corvallis, June 24, 1914, F. D. Bailey, 1628, July 29, 1914, 1476; Hood River, June 20, 1914, 3372. This species has not been previously reported west of the Missis- sippi valley. 191. UROMYCES INTRICATUS Cooke, Grevillea 7: 3. 1878. | Uromyces Eriogoni Ell. & Hark.; Harkness, Bull. Cal. Acad. Sci. Es 29. 1884: Nigredo intricata Arth. N. Am. Flora 7: 244. 1912. ON POLYGONACEAE: Eriogonum compositum Dougl.—The Dalles, Wasco Co., July 3, 1914, 1300; Hood River, July 22, 1915, 3140. Eriogonum microthecum Nutt.—Redmond, Crook Co., July 2, 1914, 2537: Enogonum stellatum Benth.—Wren, Benton Co., June 26, 1914, 1320; Hilgard, Union Co., July 10, 1914, 1439. Eriogonum umbellatum Torr.—Mt. Hood, 7,000 ft. elev., Sept. 1, 1901, E. W. D. Holway, 6,500 ft., Aug. 9, 1914, 1481, 1493. Enogonum vimineum Dougl.—Elgin, Union Co., Aug. 15, 1899, C. L. Shear (Ell. & Ev. Fungi Columb. 1470). Eriogonum virgatum Benth.—Grant’s Pass, Josephine Co., July 13, 1887, Thomas Howell, from Phanerogamic specimen in the her- barium Missouri Bot. Gard. Eriogonum sp.—Waloupi Canyon, Wallow Co., Aug. 18, 1897, E. P. Sheldon, 8727; Hermiston, Umatilla Co., May 12, 1915, 3039, 3250; Hood River Co., July 22, 1915, 3140. 278 BROOKLYN BOTANIC GARDEN MEMOIRS 192. URoMYCcES JACKSONII Arth. & Fromme, Torreya 15: 260. I9I15. ON POACEAE: Agrostis Halli Vasey—Corvallis, Sept. 4, 1914, 1576. Agrostis maritima Lam.—Hood River Co., Aug. 26, 1915, E. Bartholomew, 5971 (Barth. Fungi Columb. 4992). Deschampsia caespitosa (L.) Beauv.—Toledo, Lincoln Co., July 19, 1915, 3104. Deschampsia elongata (Hook.) Munro—Orenco, Washington Co., June 13, 1914, 2396; Corvallis, July 6, 1914, 2658, type, July 209, 1914, 1438; Glendale, Douglass Co., Aug. 17, 1914, 1408. Hordeum jubatum L.—Umatilla, Umatilla Co., July 11, 1914, 1376. Hordeum nodosum L.—Portland, May 21, 1914, F. D. Bailey, 7583. In addition to the above collections this species is now recognized on Agrostis pallens from California and Muhlenbergia Lemmoni from Arizona and New Mexico. Collections have also been made on Deschampsia elongata in Washington and on Hordeum nodosum in Washington and California. 193. Uromyces Junci (Desmaz.) L. Tul. Ann. Sci. Nat. IV, 2: 146. 1854. Puccinia Junct Desmaz. Pl. Crypt. 81. 1825. Nigredo Junci Arth. N. Am. Flora 7: 238. 1912. ON CARDUACEAE: I. Arnica cordifolia Hook.—Austin, Grant Co., Aug. 1915, J. R. Weir, 190. On JuncaAceAE: IJ, III. Juncus balticus Willd.—Redmond, Crook Co., July 2, 1914, 1426; Umatilla, Umatilla~Co., July 11, 1914, 7372; Toledo, Lincoln €or July 19, 1915, 3391. This species develops its aecia on various Carduaceae. In Europe cultures have been conducted by various authors, according to Klebahn (Die Wirtsw. Rostp. 329. 1904), showing that the aecia occur on Pulicaria dysenterica (Inula dysenterica). In America Arthur (Mycol. 4: 22. I912, 7:77. 1915) has shown by culture experiments that aecia occur on Carduus and Ambrosia. The aecia on Arnica are referred here on morphological grounds. 194. UROMYCES JUNCI-EFFUSI Sydow, Monog. Ured. 2: 290. 1910. Puccinia Junci Schw. Trans. Am. Phil. Soc. II. 4: 295. 1832. (Not P. Junci Desmaz. 1825.) Uromyces effusus Arth. Jour. Myc. 13: 193. 1907. (Not U. effusus DeToni. 1888.) Nigredo Junci-effust Arth. N. Am. Flora 7: 239. 1912. JACKSON: UREDINALES OF OREGON 279 ON JUNCACEAE: Juncus Bolandert Engelm.—Ashland, Jackson Co., Sept. 10, 1914, 2a 2G: Juncus ensifolius Wikstr.—Minum River, Wallowa Co., Aug. I1, 1897, E. P. Sheldon, Aug. 20, 1897, 8751a; Corvallis, Aug. 10, I9ITI, 1188, July 29, 1914, 2522; Philomath, Oct. 28, 1911, 7184, 1185, 1186, jan.-6; 1o14, 1700; Clatsop,. Clatsop Co., Nov. 7; 1913, 7799; Hood River Co., Aug. 5, 1914, 2521; Ashland, Jackson Co., Sept. 10, 1914, 2524; Newport, Lincoln Co., July 17, 1915, 3394. Juncus Mertensianus Bong.—Big Creek, Waldport, Lincoln Co., Aue. 23, 1915, F. D. Bailey, 3361. Juncus orthophyllus Cov.—Silver Lake, Lake Co., (?) Aug. 20, 1891, J. B. Lieburg, from Phan. spec. in N. Y. Bot. Gard. 765. Juncus oxymeris Engelm.—St. Johns, Multnomah Co., July 28, 1902, E. P. Sheldon, from Phanerogamic specimen in National Museum IIO1Q. This species has not been connected with any aecial form. From field observations made by the writer in Oregon it seems probable that the aecia are to be looked for on Aster. 195. Uromyces Lupini Berk. & Curt. Proc. Am. Acad. 4: 126. 1858. Nigredo Lupini Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. ON LEGUMINOSAE: Lupinus laxiflorus Dougl.—Garden Home, Multnomah Co., Aug. 1909, 1828, July 20, 1915, 3240. Lupinus rivularis Dougl.—Springbrook, Yamhill Co., May 14, 1914, F. D. Bailey, 1528. Lupinus sp.—Mt. Hood, Aug. 31, 1901, E. W. D. Holway, Aug. 9, 1914, 3227; Bonneville, Multnomah Co., Aug. II, 1910, 1074, 1088; Philomath, May 10, 1914, 3108; Springbrook, Yamhill Co., June 22, 1914, F. D. Bailey, 3111; Jetty, Lincoln Co.,-July 19, 1915, VanGundia, 3131; Newport, Lincoln Co., July 20, 1915, 3264; Hood River, July 23, 1915, 3020; Eddyville, Lincoln Co., Aug. 9, 1915, Hoerner, 3750. 196. Uromyces MEpIcAGINIs Pass. in Thiim. Herb. Myc. Oecon. 156. 1874. Nigredo Medicaginis Arth. N. Am. Flora 7: 256. 1912. On LEGUMINOSAE: Medicago lupulina L.—Albany, Linn Co., Aug. 1907, David Griffiths; Medford, Jackson Co., June 26, 1915, G. B. Posey, 3057. The aecia of this species in Europe have been shown by Schroeter (Krypt. Fl. Schl. 3!: 306. 1887) and by Trebaux (Ann. Myc. ro: 74. 1912) to occur on various species of Euphorbia. 280 BROOKLYN BOTANIC GARDEN MEMOIRS No aecia in America have been found which can be referred to this species. There is, however, no evidence at present available for be- lieving the American species different from the European. 197. UROMYCES MINIMUS Davis, Bot. Gaz. 19: 415. 1894. Nigredo minima Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. ON POACEAE: Muhlenbergia comata (Thurnb.) Benth.—Wallowa Valley, Wallowa Co., July 28, 1900, Wm. C. Cusick. Muhlenbergia racemosa (Michx.) B.S.P.—Wallowa Valley, Wal- lowa Co., July 28, 1900, Wm. C. Cusick. 198. UROMYCES OBLONGA Vize, Grev. 5: I10. 1877. Uromyces minor Schrét. Krypt. F. Schles. 3!: 310. 1887. Uromycopsis minor Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. On LEGUMINOSAE: Trifolium albopurpureum T. & G.—Corvallis, E. R. Lake, 32209. Trifolium dubium Sibth.—Corvallis, Apr. 10, 1914, F. D. Bailey, 1522; Orenco, Washington ‘Co., June 13, 1914, 3226; * Yaquina, Lincoln Co., July 17, 20, 1915, 3100; Gerlinger, June 22, 1914, G. B. Posey; 323.2- Trifolium eriocephalum Nutt.—Corvallis, July, 1910, 1989. Trifolium Halla Howell—Corvallis, June 6, 1899, E. R. Lake, 3232, May. 12, 19003, A. Hi. Post, 3230: Trifolium microdon H. & A.—Corvallis, May 11, 1907, E. R. Lake, 1408. Trifolium oliganthum Steud.—Corvallis, May 11, 1914, F. D. Bailey, AE hehe Trifolium procumbens L.—Corvallis, June 22, 1901, L. C. M., 7633, July, 1910,7170- Trifolium tridentatum Lindl.—Philomath, June 24, I914, 1345; Corvallis, May 28,, 1903, E> oR: Lake; 3230, May 11, 1914; 2 Bailey, 3235. This opsis-form is very common on native Trifolium sp. The original collection by Harkness was reported as occurring on “ Bur cloves”? now considered an error for Trifolium. 199. UROMYCES OCCIDENTALIS Diet. Hedwigia Beibl. 42: 98. 1903. Nigredo occidentalis Arth. N. Am. Flora 7: 252. 1912. ON LEGUMINOSAE: Lupinus sp.—Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 248. 200. UROMYCES PERIGYNIUS Halsted, Jour. Myc. 5: 11. 1889. Nigredo perigynia Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. JACKSON: UREDINALES OF OREGON 281 ON CYPERACEAE: Carex arthrostachya Olney—Corvallis, July, 1910, 7791, 1102. This species is morphologically indistinguishable from P. Asterum (cf. 69) in all spore stages except in the possession of one-celled telio- spores. Like that species the aecia occur on Aster and Solidago. The genetic connection was established by Arthur in repeated experiments Clone: Myce 10: 16. 1904; Mycol: 4:21. 1012, 7:75. 1915, 7:.83- 1915). Fraser (Mycol. 4: 181. 1912) has confirmed Arthur’s results in part. The above collection is the only one so far known from the Pacific coast. 201. UROMYCES PLUMBARIUS Peck, Bot. Gaz. 4: 127. 1906. Nigredo plumbaria Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. ON ONAGRACEAE: Pachylophus marginatus (Nutt.) Rydb.—Snake River, E. Oregon, June 3, 1901, W. C. Cusick (Phan. spec. 2542). Pachylophus montanus (Nutt.) A. Nels.—Crooked River, Crook Co., July 3, 1901, W. C. Cusick (Phan. spec. 2633). This species is correlated in morphological characters with P. Epilobu-tetragoni (cf. 99). 202. UROMYCES POLYGONI (Pers.) Fuckel, Symb. Myc. 64. 1869. Puccinia Polygoni Pers. Neues Mag. Bot. 1: 119. 1794. Nigredo Polygoni Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. ON POLYGONACEAE: Polygonum aviculare L.—Corvallis, Sept. 4, 1914, 1929; Clatskanie, Columbia Co., Oct. 6, 1914, F. D. Bailey, 1945; Medford, Jackson Co., June 20, 1915, G. B. Posey, 3055. 203. Uromyces porosus (Peck) comb. nov. Aecidium porosum Peck, Bot. Gaz. 3: 37. 1878. Uromyces coloradensis Ellis & Ev. Erythea 1: 204. 1893. Uromycopsis porosa Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. On LEGUMINOSAE: Vicia americana Muhl.—Orenco, Washington Co., April 23, 1915, Rerron, (anes ste, Ti 9237. The two collections were made at the same spot on different dates. The first consists of aecia only and the second of telia only. 204. UROMYCES PROEMINENS (DC.) Pass.; Rabh. Fungi Eur. 1795. 1873. Uredo proeminens DC. FI. Fr. 2: 235. 1805. 282 BROOKLYN BOTANIC GARDEN MEMOIRS Nigredo proeminens Arth. N. Am. Flora 7: 259. 1912. ON EUPHORBIACEAE: Euphorbia glyptosperma Engelm.—Wasco Co., July 23, 1885, W. N. Suksdorf (from Phan. spec. in N. Y. Bot. Gard.). Euphorbia oregonensis Millsp.—Horse Creek Canyon, Wallowa Co., May 20, 1897, E. P. Sheldon, S715. 205. UROMYCES PUNCTATUS Schrét. Abh. Schles. Ges. 48: 10. 1870. Nigredo punctata Arth. N. Am. Flora 7: 253. 1912. ON LEGUMINOSAE: Astragalus Purshit Dougl.—Austin, Grant Co., Aug. 1915, J. R. Welr, 236. The aecia of this species have been shown by European authors to occur on Euphorbia cyparissias. No aecial collections have been made in America. 206. UROMYCES ScirPiI (Cast.) Burrill, Bot. Gaz. 9: 188. 1884. Uredo Scirpi Cast. Cat. Pl. Mars. 214. 1845. Nigredo Scirpt Arth. Résult Sci. Congr. Bot. Vienne 344. 1906. ON CYPERACEAE: Scirpus paludosus A. Nels.—Waldport, Lincoln Co., Aug. 23, 1915, F. D. Bailey, 3323. This species was first shown by Dietel (Hedwigia 29: 149. 1890) to have its aecia on Sium latifolium and Hippurus vulgaris. Other investigators have added other Umbelliferous hosts to the list. In ‘America Arthur (jour: Myc. 13: 199: “1907; “14: 17. 4908; Mycol. 1: 237. 1909) has shown that Cicuta maculata is an aecial host. Fraser (Mycol. 4: 178. 1912) has confirmed Arthur’s work. Aecia on other hosts are properly referred here on morphological grounds. The species can doubtless be separated into a number of biological forms when more extensive culture work has been conducted. 207. UROMYCES SILPHII (Burrill) Arth. Jour. Myc. 13: 202. 1907. Aecidium compositarum Silphit Burrill; DeToni in Sacc. Syll. Fung. 7: 798. 1888. Uromyces Junci-tenuis Sydow, Monog. Ured. 2: 289. IgI0. Nigredo Silphi Arth. N. Am. Flora 7: 239. 1912. ON JUNCACEAE: Juncus occidentalis (Cov.) Wieg.—Corvallis, Aug. 10, I911, 1187, June 24, 1914, F. D. Bailey, 1387, July 29, 1914, 1445; Philomath, Jan. 6, 1914, 1108, May 10, 1914, 3393; Hood River, July 24, 1915, 3392. Arthur (Jour. Myc. 13: 202. 1907; 14:17. 1908) has shown that this common species has its aecia on Silphium. Using telial material on J. tenuis from Indiana, West Virginia and Nebraska, five successful JACKSON: UREDINALES OF OREGON 283 infections of Silphium perfoliatum were obtained, all of which resulted in the development of pycnia and aecia. The aecia on Silphium have been collected, so far as known to the writer, only in the Mississippi Valley from Ohio to Wisconsin, Kansas and Missouri, on three species of Silphium. The range of the telial collections referred here, how- ever, is much greater including nearly the entire United States and Canada except the south Pacific slope. It seems probable that some plants other than Silphium, at present unrecognized, also serve as aecial hosts for this species. 208. UROMYCES SOLIDAGINIS (Sommf.) Niessl, Verh. Natur.-Ver. Brum: 10: 163. 1872: Caeoma Solidaginis Sommerf. Suppl. Fl. Lapp. 234. 1826. Telospora Solidaginis Arth. Résult Sci. Congr. Bot. Vienne 346. 1906. ON CARDUACEAE: Solidago sp.—Dufur, Wasco Co., June 30, 1914, 1336. This is the only micro-Uromyces occurring in both Europe and America. This species shows a morphological correlation with P. Asteris (cf. 70). 209. UROMYCES SPRAGUEAE Hark. Bull. Calif. Acad. Sci. 1:36. 1884. Uromycopsis Spragueae Arth. Résult Sci. Congr. Bot. Vienne 345. 1906. ON PORTULACEAE: Calyptridium roseum Wats.?—Crater Lake, Klamath Co., 7,000 ft., Sept. 22, 1913, E. P. Meinecke, CrPRD (2) 11. Spraguea multiceps Howell—Strawberry Mt., Grant Co., Sept. 2, 1913, W. E. Lawrence, 1177. 210. UROMYCES SUBSTRIATUS Sydow, Ann. Myc. 4: 30. 1906. Nigredo substriata Arth. N. Am. Flora 7: 253. I9g12. On LEGUMINOSAE: Lupinus sp.—Austin, Grant Co., Aug. 1915, J. R. Weir, 150. 211. Uromyces TriroLi (Hedw. f.) Lev. Ann. Sci. Nat. III, 8: 371. 1847. Puma Trfolu Wedw.t.; DC. Fl. Fr. 2: 225. 1805. Nigredo Trifolit Arth. Result Sci. Congr. Bot. Vienne 344. 1906. ON LEGUMINOSAE: Trifolium hybridum L.—Corvallis, Aug. 1909, 3092; Hood River Co., May 14, 1914, 3o9r, Aug. 5, 1914, 3095; Portland, June 11, 1914, 3233; Springbrook, Yamhill Co., June 22, 1914, G. B. Posey, 3103; Garden Home, Multnomah Co., July 15, 1914, F. D. Bailey, 1508; Grant’s Pass, Josephine Co., Sept. 3, 1916, J. R. Weir, 277. 284 BROOKLYN BOTANIC GARDEN MEMOIRS 212. UROPYXIS SANGUINEA (Peck) Arth. N. Am. Flora 7: 155. 1907. Uromyces sanguineus Peck, Bot. Gaz. 4: 128. 1879. Puccinia mirabilissima Peck, Bot. Gaz. 6: 226. 1881. ON BERBERIDACEAE: Berberis aquifolium Pursh—Sauvies Island, Multnomah Co., Apr., 1882, Joseph Howell; Siskiyou, Jackson Co., May 31, 1894, E. W. D. Holway (Barth. N. Am. Ured. 1400); Corvallis, March 26, 1908, C. C. Cate, 3382, April 13, 1908, F. L. Griffin, 39383, March 14, wens: 1137; Philomath, Jan. 1, 1914, 1753; Tualetin, Washington Co., March 25, 1915, F. D. Bailey, 2616; Grant’s Pass, Josephine Co., Sept. 3, 1916, J..R. Weir, 18&2a. Berberis pumila Greene, Pokegama, Klamath Co., July 9, 1903, E. B. Copeland (Sydow, Ured. 1777; Baker, Pacific Coast Fungi 3708). FoRM GENERA 213. AECIDIUM ALLENII Clinton in Peck, Rep. N. Y. State Mus. 24: O23.) G72. On ELAEAGNACEAE: Lepargyrea canadensis (L.) Greene—Strawberry Mt., Grant Co., Sept. 2, 1913, W. E. Lawrence, 1173; Sumpter, Baker Co., June, 1913, J. Re Weir;.6;Gold Center, July, 1914,,H.F. Wilson, 7622: 214. AECIDIUM COLLINSIAE Ell. & Ev. Bull. Washb. Lab. 1: 4. 1884. Aecidium Tonellae D. & H. Erythea 3: 77. 1895. ON SCROPHULARIACEAE: Collinsia parviflora Lind|.—Philomath, April 20, 1912, 1169. This species is evidently an heteroecious form known otherwise only from Washington on the above host and on C. Rattoni and C. tenella. It is apparently distinct from P. Collinsiae P. Henn. (Hedwigia 37: 269. 1898) as stated by Hennings. . The aecia of the latter, judging from the description, arise from a limited mycelium. There is no evidence of telia in any of the collections of A. Collinsiae ex- amined. P. Collinsiae has evidently been collected but once and material is not available for examination. 215. AECIDIUM COLUMBIENSE Ell. & Ev. Erythea 1: 206. 1893. On CICHORIACEAE: Mieracium albiflorum Hook.—Hood River, road to Lost Lake, May 16, 1915, 3245. Meracium sp.—Bridal Veil, Multnomah icon May 18, 1915, 3291. The aecia arise from a distributed mycelium and are not followed by any other stage. Puccinia Hieracitt may however occur on the iS) CO On JACKSON: UREDINALES OF OREGON same plants and even on the same leaves. Sydow (Ann. Myc. 1: 326. 1903) described P. sejuncta on such a mixture. 216. AECIDIUM DELPHINI Barth. Jour. Myc. 8: 173. 1902. Aecidium Batesianum Barth. in E. & E. Fungi Col. 1901. 1904. ON RANUNCULACEAE: Delphinium depauperatum Nutt.—Mary’s Peak, Benton Co., May 21 kO05, 3210. Delphinium sp.—Corvallis, April 11, 1915, 2615; Redmond, Gxook €o:, May 15, 1915, 3327. This species is possibly identical with aecia on other Ranuncu- laceous hosts referred to P. Clematidis (cf. 85). For purposes of this list it is retained as a separate form as no cultures have been conducted. 217. AECIDIUM GRAEBNERIANUM Henn. Hedwigia 37: 273. 18098. Aecidium Alaskanum Trelease, Harr. Alaska Exp. 5: 37. 1904. On ORCHIDACEAE: Limnorchis dilatata (Pursh) Rydb.—Horse Lake, Cascade Mts., Aue, to00, J. C. Bridwell, 3322. This unconnected Aecidium is doubtless heteroecious since no other stages have been found following the aecia on any of the col- lections examined. The species is known otherwise only from Alaska and in the mountains of British Columbia, Montana and California. 218. PERIDERMIUM COLORADENSE (Diet.) Arth. & Kern, Bull. Torrey Club 33: 426. 1906. ON PINACEAE: Picea Engelmanui Parry—Whitman National Forest, Wallowa Co., Juty,,1913, J. R. Weir, 277. This species forms large witches’ brooms. 219. PERIDERMIUM ORNAMENTALE Arth. Bull. Torrey Club 28: 665. 19Ol. ON PINACEAE: . Abies concolor (Gord.) Parry J. R. Weir, 745. Abies nobilis Lind|.—Larch Mt., Multnomah Co., Aug., 1910, 3293. White Pine, Baker Co., June, 1913, 220. Uredo Phoradendri sp. nov. O. Pycnia not seen. II. Uredinia amphigenous, gregarious, not crowded, spots not conspicuous, punctate; rounded or slightly elongated, 0.4-0.8 mm. across, tardily naked, somewhat pulverulent, bright orange, dehiscent by an elongate or irregular fissure of the epidermis, ruptured epidermis conspicuous and persistent; peridium membranous, at first hemi- spherical, remaining closely adherent to the ruptured epidermis, 286 BROOKLYN BOTANIC GARDEN MEMOIRS made up of colorless isodiametric cells, 14-19 wu across, sometimes somewhat rhomboidal, smooth, wall 1-1.5 4 thick; urediniospores ellipsoid or obovoid, 17-33 by 26-32 u, wall colorless, 1.5—2.5 uw thick, very closely and minutely echinulate, pores very indistinct, 10-12, scattered. III. Telia unknown. On LORANTHACEAE: Phoradendron villosum Nutt.—Corvallis, Sept. 21, 1915, C. E. Owens, 3377 type. INDEX TO SPECIES Figures following the names refer to the species numbers in the list. Synonyms are in italics. New names and combinations are in bold-face type. Aecidium abundans 59 Alaskana 217 Allenii 213 Aquilegiae 85 aspertfolit 67 Asterum 69 Barbarae 135 Batesianum 216 Bellidis 139 Berberidis 151 Blasdaleanum 35 carneum 184 Cinerariae 101 Clematidis 85 columbiense 116, 215 columnare 5 Collinsiae 214 compositarum Bidentis 140 compositarum Silphit 207 conorum Piceae 22 Delphinii 216 Dracunculi 173 elatinum 20 Fendleri 103 fuscum 57 Gali ambigum 61 Geranit-maculati 153 Giliae 150 Graeberianum 217 Grossulariae 109 Helianthi-mollis 112 Holboellit 117 Hydrophylli 136 Hyperici-frondost 190 Ligulariae 101 Majanthae 125 Mertensiae 136 monoicum 135 Myrtillt 27 nilens 45 Oenotherae 146 Ornithogalum 63 Palmeri 143 Peckit 146 Phaceliae 136 Pirolae 22 porosum 203 pseudo-balsameum 32 punctatum 17 Pyrolae 29 Rhamnt 159 Sarcobatt 167 Solidaginis 69 Sorbt 43 Tonellae 214 Tussilaginis 100 Urticae 174 Violae 176 Allodus ambigua 61 aspertor 68 Calochorti 76 claytoniata 84 commutata 89 Dichelostemmae 96 Douglasit 97 Giliae 150 Jonesit 122 melancontoides 128 oregonensis 68 Palmeri 143 Aregma speciosa 33 Ascophora disciflora 47 JACKSON: UREDINALES OF OREGON Bullaria A ngelicae 62 Caeoma Armeriae 181 asteratum 69 claytontata 84 erigeronatum 69 Galit 25 hieraciatum 115 nitens 45 occidentalis 13 Solidaginits 208 urticatum 174 Calyptospora columnaris 5 Geoppertiana 5 Chrysomyxa Abietis 6 Pirolae 22 Weirii 6 Coleosporium Adenocaulonis | asterum 4 Madiae 2 occidentale 3 Solidaginis 4 Cronartium coleosporiodes 7 Comandrae 8 filamentosum 7 pyriforme 8 Dicaeoma asarinum 66 mesomegalum 130 Earlea speciosa 33 Gymnoconia interstitialis 45 Gymnosporangium dsiaticum 40 Betheli 34 Blasdaleanum 35 chinense 40 Haraeanum 40 Harknessianum 36, 39 juniperinum 37 juvenescens 38 Kernianum 39 koreaense 40 Libocedri 35 Nelsoni 41 nootkatensis 42, 43 Sorbi 43 tubulatum 44 Hyalopsora Aspidiotus 9 laeviuscula 10, 23 Polypodii 11 20 Kunkelia nitens 45 Lycoperdon caryophyllinum 185 epiphyllum 100 poculiforme 151 Melampsora albertensis 13, 18 arctica 14 Bigelowii 15, 16 confluens 16 Lini 17 Medusae 18 occidentalis 18 Piscariae 19 Sparsa 30 sp. I2 Melampsorella Cerastii 20 elatina 20 Melampsoropsis Piperiana 21 Pyrolae 22 Milesia Polystichii 23 Nigredo aemula 179 caryophyllina 185 Fabae 186 fallens 187 Hyperici-frondosi 190 tntricata 191 Junct 193 ‘Junci-effusi 194 Lilit 189 Lupini 195 Medicaginis 196 minima 197 occidentalis 199 perigynia 200 plumbaria 201 Polygoni 202 proeminens 204 punctata 205 Scirpt 206 Silphit 207 substriata 210 Trifolit 211 Nyssopsora echinata 46 Peridermium acicolum 4 balsameum 31, 32 Betheli 8 californicum 2 coloradense 218 conorum Piceae 22 287 288 BROOKLYN BOTANIC GARDEN MEMOIRS Peridermium filamentosum 7 Harknessii 7 montanum 4 ornamentale 219 Peckii 27 pseudo-balsameum 32 pbyriforme 8 Stalactiforme 7 Phragmidium affine 49 disciflorum 47 imitans 48 Ivesiae 49 Jonesii 50 montivagum 51 occidentale 52 Potentillae 53 Rosae-acicularis 54 Rosae-californicae 55 Spectosum 33 Pileolaria brevipes 56 Toxicodendri 56 Polythelis fusca 57 Puccinia Absinthii 58, 90 abundans 59, 168 Acetosae 60 acuminata 154 Agropyri 85 albiperidium 109 alternans 85 ambigua 61, 161 Andropogonis 143 Angelicae 62, 98 anomala 63 Antirrhini 64 Archangelicae 62 arnicalis 65 Asari 66 asarina 66 asperifolii 67 asperior 68 Angelicae 98 Asteris 70, 208 ° atro-fusca 71 Asterum 69, 200 atropunctata IIo Bakeriana 98 Balsamorrhizae 72 Barbareae 117, 135 Bellidis 139 bicolor 73 Bistortae 74 Blasdalei 75 Puccinia Botsduvaliae 99 Calochorti 76 Campanulae 77 canaliculata 140 Caricts-A steris 69 Caricis-Erigerontis 60 Caricis-Solidaginis 69 chasmatis 170 Chelonis 78 Chrysanthemi 79 Cichorii 80 Cicutae 81 cinerea 85 circaeae 82 circinans 170 Cirsii 83 Cirsit-lanceolati 87 cladophila 111 Clarkiae 99 claytoniata 84 Clematidis 85, 107, 136, 172, 216 Clintonii 86 Cnici 87 Columbiensis 162 Collinsiae 214 Comandrae 88 compacta 94. commutata 89 conferta 58, 90 congregata 114 Convolvuli 91 coronata 88, 159 Crandallit 59 Crepidis-acuminatae 92 curtipes 163 Cyani 93 DeBaryana 94 Dentariae 95 Dichelostemmae 96 difformis 61 dis persa 67 Douglasii 97 Dracunculi 173 Ellisi 62, 98 Epilobii 175 Epilobii-Fleischeri 175 Epilobii-tetragoni 99, 175, 201 Eriophori ror epiphylla 100 Eriophyllii 102 extensicola 69 Fendleri 103 JACKSON: UREDINALES OF OREGON Puccinia Garrettii 174 Gayophyti 99 Gentianae 105 gemella 104 Giliae 106 glabella 99 glumarum 107 graminis 103, 151 granulispora 108 Grossulariae 109, 144 grumosa IIO Harknessi ITI Heliantht 112 Helianthi-mollis 112 Hemizoniae 113 Heucherae 114 hieraciata 115 Hieracii 116, 215 Holboellii 117, 135 holcina 118 Hordet 63 Hydrophylli 160 Hypochoeridis 119 inclusa 83 insperata 120 Iridis 121 Koeleriae 103 Jonesii 122 Junct Desmaz. 193 Junct Schw. 194 Kreagerit 59 Ligustici 123 ludibunda 146 luteobasis 123 luxuriosa 124, 167 Madiae 138 Maegnusit 109 Majanthae 125 Malvacearum 126 Malvastri 164 Mariae-Wilsoni 84 McClatchieana 127 melanconoides 128, 141 Menthae 129 mesneriana 88 mesomegala 130 micromeriae 131 microsora 132 Millefolii 133 mirabilissima 212 monardellae 134 monoica 135 Puccinia montanensis 136 mutabilis 137 neglecta 107 obliterata 85 obscura 139 obtecta 140 Oenotherae 99 orbicula 120 oregonensts 68 Ortonii 141 Osmorrhizae 148 Oxyriae 142 Palmeri 143 Parkerae 144 patruelis 115 Pattersoniana 145 Peckiana 45 Peckii 146 Pentastemonis 147 Phaceliae 160 Phaseoli var. Taraxact 169 Phlei-pratensts 151 Pimpinellae 148 Piperi 149 plumbaria 106, 150 Poarum 100 poculiformis 151 Polygont 202 Polygoni-alpini 152 Polygoni-amphibii 153 porphyrogenita 154 Potentillae 53 Pringsheimiana 109 procera 155 Pruni-spinosae 178 punctata 61, 156, 161 pustulata 88 pygmaea 157 recedens 158 recondita 90 Rhamni 88, 118, 159 Ribis 144 Richardsontu 97 Romanzoffiae 160 rubefaciens 161 rubigo-vera 172 rugosa 162 Saxifragae 163 sejuncla 116, 215 sessilis 125 Sherardiana 164 Sidalceae 165 289 290 Puccinia similis 58 simplex 63 Sphaeralceae 165 Spreta 114 Stipae 166 straminis simplex 63 subnitens 124, 167 : Symphoricarpi 168 Syntheridis 177 Taraxaci 169 Tiarellae 114 tomtpara 85 tosta luxurians 124 Toumeyi 170 Trautvetteriae 171 Treleasiana 104 trifoliata 148 Trifolit 211 triticina 172 Troximontis 162 uniporula 109 universalis 173 Urticae 174 Urticatum 174 Veratri 175 Violae 176 Wulfeniae 177 Pucciniastrum Abieti-Chamaenerii 24 Galii 25 Goodyerae 26 Myrtilli 27 pustulatum 24, 28 Pyrolae 29 sparsum 30 Vacciniorum 27 Roestelia Betheli 34 Harknessiana 36 koreaensts 40 tubulata 44 Telospora Solidaginis 208 Tranzschelia punctata 178 Tremella juniperina 37 Trichobasis Balsamorrhizae 72 glumarum 107 Triphragmium echinatum 46 Uredinopsis Atkinsonw 31 Copelandi 31, 32 Pteridis 32 BROOKLYN BOTANIC GARDEN MEMOIRS Uredo Acetosae 60 Angelicae 62 Artemisit 58 Aspidiotus 9 Betae Convolvuli 91 Chamaecyparidis-nutkaensis 42 Chimaphilae 29 Cichorit 80 coleosporiodes 7 confluens 16 Cyant 93 Dentariae 95 Fabae 186 fallens 187 Gentianae 105 glumarum 107 Goodyerae 26 Heucherae 114 Hieracti 116 Tridis 121 laeviuscula 10 miniata Lint 17 nootkatensis 42 Phoradendri 220 Pimpitnellae 148 Polygoni Bistortae 74 Polypodi 11 proeminens 204 pustulata 28 quinqueporula 151 Scirpt 206 Solidaginis 4 vagans a Epilobii-tetragoni 99 Uromyces Acetosae 60 aemulus 179 amoenus 180 Armeriae 181 atro-fuscus 71 Beckmanniae 182 Brodieae 183 carneus 184 caryophyllinus 185 coloradensts 203 effusus 194 Eriogoni 191 Fabae 186 fallens 187 heterodermus 188 Holwayi 189 Hordei 182 Hyperici-frondosi 190 intricatus I9I JACKSON: UREDINALES OF OREGON Uromyces Jacksonii 182, 192 Junci 193 Junci-effusi 194 Junci-tenuis 207 lapponica 184 Lilit 189 Lupini 195 Medicaginis 196 minimus 197 minor 198 oblonga 198 occidentalis 199 perigynius 200 plumbarius 175, 201 Polygoni 202 porosus 203 Abies amabilis 32 balsamea 5, 14, 24 concolor 5, 219 Fraseri 5 grandis 5, 20, 24, 25, 31, 32 lasiocarpa 5, 20, 24 magnifica 5 nobilis 31, 219 pectinata 5 Abutilon? sp. 126 Achillea millefolium 133 Adenocaulon bicolor 1 Agoseris glauca 115 laciniata 162 Agropyron dasystachyum 85, 151 lanceolatum 85 spicatum 85, 145 tenerum 136 Agrostis alba 159 alba maritima 159 exarata I51, 159 foliosa 159 Hallii 192 longiligula 159 maritima 192 microphylla 159 pallens 192 Alopecuris californicus 151 Althaea ficifolia 126 rosea 126 Alliaceae 75, 96, 108, 137, 179, 183 Allium acuminatum 75 attenuifolium 75 Uromyces proeminens 204 punctatus 205 sanguineus 212 Scirpi 206 Silphii 207 Solidaginis 208 Spragueae 209 substriatus 210 Toxicodendri 56 Trifolii 211 Uromycopsis Brodieae 183 lapponica 184 minor 198 porosa 203 Spragueae 209 Uropyxis sanguinea 212 Host INDEX Allium Geyeri 137 nevil 108 validum 179 Amaranthaceae 167 Amelanchier alnifolia 36 florida 35 Amygdalus Persica 17 Anacardiaceae 56 291 Anaphalis margaritacea occidentalis 180 margaritacea subalpina? 180 Anemone Drummondii 94 oregana 57 quinquefolia 57 Angelica atropurpurea 62 genuflexa 62, 98 Lyallii 62 tomentosa 98 Antirrhinum majus 64 Aquilegia formosa 85 truncata 85 Arabis sp. 135 Arbutus Menziesii 30 Arctostaphylos alpina 30 manzanita 30 nevadensis 30 Aristolochiaceae 66 Arnica cordifolia 65, 193 Artemisia dracunculoides 58 frigida 58 ludoviciana 58, 90 rigida 58 tridentata 58 Asarum caudatum 66 292 BROOKLYN BOTANIC GARDEN MEMOIRS Aspidium Thelepteris 31 IQ, 133) 138) 158, 172) 1s80;mocs Aster acuminatus 69 208 conspicuus 4, 70 Carex athrostachya 69, 200 Cuseckii 4 Barbarae 174 Douglasii 4 canescens 69 folicaeus frondeus 4 Deweyana 69 Halli 4 Douglasii 71 laevis geyeri 4 festiva 69, 109 Astragalus sp. 184 Goodenowii 69, 109 Purshii 205 Hookeriana 146 Athyrium cyclosorum 31 Kelloggii 109 Avena fatua glabrata 151 laciniata 174 sativa I51, 159 magnifica 109, 174 marcida 69, 115, 173 Balsamorrhiza deltoidea 72 mertensii 109 sagittata 72 mirata 132 Beckmannia erucaeformis 151, 182 monile 109 Berberidaceae 103, 212 multicaulis 173 Berberis aquifolium 103, 212 nebraskensis 109, 174 nervosa 103 phyllomanica 109 pumila 212 praegracilis 69, 115, 173 Bidens connata 140 pratensis 115 frondosa 140 rostrata 174 Boisduvalia densifolia 99 Rossii 173 glabella 99 scoparia 69 stricta 99 spectabilis 109 Boraginaceae 136 stipata 69 Brodiaea sp. 183 straminis 69 congesta 96 subfusca 69 Bromus carinatus 85 trisperma 69 carinatus californicus 85 umbellata 173 grandis 85 utriculata 174 hordeaceus 85 vulgaris 69 hordeaceus leptostachys 85 Caryophyllaceae 20, 185 marginatus 85 Castilleja miniata 7 secalinus 85 Centaurea cyanus 93 tectorum 85 Cerastium oreophilum 20 villosus 85 viscosum 20 vulgaris 85 vulgatum 20 Chamaecyparis nootkatensis 42 Calamagrostis aleutica 157 Chamaenerion angustifolium 24 canadensis 159 Chleone nemorosa 78 hyperborea 159 Chenopodiaceae 167 Calochortus macrocarpus 76 Chenopodium album 167 Caltha biflora 104 Chimaphila occidentalis 29 Calyptridium roseum 209 umbellata 29 Campanulaceae 77 Chrysanthemum sinense 79 Campanula rotundifolia 77 Cichoriaceae 73, 80, 92, III, I16, 120, Scouleri 77 162, 169, 215 Caprifoliaceae 59, 168 Cichorium intybus 80 Carduaceae I, 2, 3, 4, 58, 65, 69, 70, 72, Cicuta maculata 206 7ONSSs1 879190) 93) LOL wO2) Lia anna. occidentalis 81 JACKSON: UREDINALES OF OREGON Circaea pacifica 82 Cirsium americanum 83 edule 83 lanceolatum 87 undulatum 83 Clarkia pulchella 99 Claytonia lanceolata 84 Clematis Drummondii 85 Douglasii 85 hirsutissima 85 ligusticifolia 85 Clintonia uniflora 130 borealis 130 Collinsia parviflora 214 Rattoni 214 tenella 214 Comandra umbellata 8, 88 Convallariaceae 130 Convolvulaceae 91 Convolvulus atriplicifolius 91 Cornaceae 154 Cornus canadensis 154 Crataegus Douglasii 34, 35, 44 Crepis gracilis 92 Cruciferae 95, 167 Cydonia vulgaris 35, 40 japonica 35 Cyperaceae 69, 71, IOI, 109, I15, 127, 132, 140, 146, 173, 174, 200, 206 Dactylis glomerata I51 Delphinium depauperatum 216 Dentaria tenella 95 Deschampsia caespitosa 192 elongata I5I, 192 Dianthus caryophyllus 185 Distichlis spicata 167 Dodecatheon alpinum 141 Hendersonii 141 Jeffreyi 141 latifolium 12? tetrandrum? 141 Elaeagnaceae 213 Elymus arenicola 155 condensatus 85 glaucus 85, 107, 136, I5I triticoides 85 virginicus 136 Epilobium adenocaulon 28 brevistylum 28 minutum 99 225 Epilobium paniculatum 99 Ericaceae 21, 30 Erigeron speciosus 69 Eriogonum compositum I9gI microthecum I9gI stellatum I9gI umbellatum 191 vimineum I9I virgatum 191 Eriophorum angustifolium 101 polystachyon Ior viridi-carinatum I01 Eriophyllum lanatum 102 leucophyllum 102 Eremocarpus setigerus 19 Erythronium parviflorum 188 Euphorbia cyparissias 205 glyptosperma 204 oregonensis 204 Euphorbiaceae 19, 204 Euthamia occidentalis 69 Festuca confinis 59 elatior 151, 159 idahoensis 59 megalura I51 myuras I51 pacifica 149, I51 rubra 59 subulata 59, 159 Filix fragilis 11 Galium aparine 61, 156 asperrinum 156 boreale 161 triflorum 25, 156 Gaylussacia baccata 27 resinosa 27 Gayophytum ramosissimum 99 Gentiana oregana 105 Gentianaceae 105 Gilia gracilis 150 Godetia amoena 99 Goodyera Menziesii 26 Grindelia sp. 166 Grossulariaceae 16, 109 Helianthus annuus 112 Hemizonia truncata 113 Heuchera micrantha 114 Hieracium albiflorum 116, 215 cinereum 73, 116 294 BROOKLYN BOTANIC GARDEN MEMOIRS Hieracium gracile 116 Scouleri 73, 116 Hierochloe macrophylla 151 Hippurus vulgaris 206 Holcus lanatus 118, 159 Hordeum distichon 151 Gussoneanum 107 jubatum 192 montanense 63 murinum 63 nodosum 63, 192 vulgare 63, 107, 151 Hookera pulchella 96 Hydrophyllaceae 136, 160 Hydrophyllum albifrons 136 capitatum 136 tenuipes 136 Hyperiaceae 190 Hypericum Scouleri 190 Hypochaeris radicata 119 Inula dysenterica 193 Iridaceae 121 Iris tenax 121 Ivesia Baileyi 50 Juncaceae 139, 193, 194, 207 Juncoides parviflorum 139 Juncus balticus 193 Bolanderi 194 ensiformis 194 mertensianus 194 occidentalis 207 orthophyllus 194 oxymeris 194 tenuis 207 Juniperaceae 34, 35, 37, 38, 39, 40, 41, 42 Juniperus chinensis 40 communis 41 occidentalis 34, 39, 41 scopulorum 34, 38, 41, 44 sibirica 37 Koeleria cristata 103, 135 Labiatae 129, 131, 134 Lagophylla ramosissima I12 Larix decidua 15 europea 15 occidentalis 15 Lathyrus obovatus 186 oregonensis 186 Lathyrus pauciflorus 186 polyphyllus 186 sulphureus 186 Leguminosae 184, 186, 187, 195, 196, 198, 199, 203, 205, 210, 211 Lepargyrea canadensis 213 Leptaxis Menziesii 114 Leptotaenia dissecta 68 Libocedrus decurrens 35 Ligularia sibirica 101 Ligusticum apiifolium 123 Cusickii 46 purpureum 46 Liliaceae 76, 110, 175, 188, 189 Lilium parviflorum 189 Limnorchis dilatata 217 Linaceae 17 Linum Lewisii 17 usitatissimum 17 Lolium multiflorum 151, 159 perenne 159 subulatum I51 Loranthaceae 220 Lupinus 199, 210 laxiflorus 195 rivularis 195 Lycopsis arvensis 67 Lygodesmia juncea III Madia citriodora 2 elegans 138 exigua 2 glomerata 2, 138 racemosa 2 ramosa 2 sativa 2 Malaceae 34, 35, 36, 37, 38, 49, 43, 44 Malus floribundus 35 Malva rotundifolia 126 Malvaceae 126, 164, 165 Medicago lupulina 196 Mentha canadensis 129 canadensis lanata 129 piperata 129 spicata 129 Mertensia laevigata 136 Micromeria chamissonis 131 Douglass 131 Mitella Breweri 114 ovalis 114 Monardella odoratissima 134 villosa 134 JACKSON: UREDINALES OF OREGON Muhlenbergia comata 197 Lemmoni 192 racemosa 197 Nabalus hastatus 120 Navarettia intertexta 106 Onagraceae 24, 28, 82, 99, 175, 201 Orchidaceae 26, 217 Ornithogalum umbellatum 63 Narbonense 63 Osmorrhiza brevipes 148 divaricata 148 Liebergii 148 occidentalis 148 Oxycoccus macrocarpus 27 Oxyria digyna 142 Pachylophus marginatus 201 montanus 201 Panicularia elata 159 pauciflora 159 Pedicularis bracteosa 86 racemosa 86 Pentstemon sp. 170 diffusus 147 Menziesii 143 Peucedanum sp. 81 triternata 122 Phalaris arundinacea 125 Phacelia heterophylla 136 leucophylla 136 Phegopteris Dryopteris 9 Phleum pratense I51 Phlox condensata 97 diffusa 97 speciosa 150 Phoradendron villosum 220 Picea canadensis 22 Engelmanii 6, 22, 218 excelsa 30 mariana 22 Pinaceae 45-5,.6; 7,-8,.13,-15, 20; 22, 24, 31, 32, 218, 219 Pinus contorta 4, 7 ponderosa 8 pungens 8 rigida 4 Piscaria setigera 19 Plumbaginaceae 181 Poa ampla 85, 100 annua 100 295 Poa fertilis 100 Kingu 59 macrantha 100 nemoralis 100 pratensis 100 triflora 100 trivialis 100 Poaceae 59, 63, 67, 85, 103, 107, 118, 124, 125, 135, 136, 145, 149) ‘151, 155521575 259, LOO 167)/°172,< 182: 192, 197 Polemoniaceae 97, 106, 150 Polygonaceae 60, 74, 142, 152, 153, 167, IQI, 202 Polygonum alpinum 153 amphibium 153 aviculare 202 emersum 153 imbricatum 74 Muhlenbergii 153 Newberryi 74 pennsylvanicum 153 Polypodiaceae 9, 10, II, 23, 31, 32 Polypodium occidentale 10 Polystichum munitum 10, 23 Populus acuminata 18 alba 12 angustifolia 18 balsamifera 18 candicans 18 tremuloides 13 trichocarpa 18 Portulaceae 84, 209 Potentilla aracnoides 53 blaschkeana 49 glomerata 49 gracilis 49 Hippiana 53 Primulaceae 128, 141 Prunus domestica 178 Pseudotsuga mucronata 13 Pteridium aquilinum pubescens 32 Ptiloria paniculata III Puccinellia nuttalliana 85 Pulicaria dysenterica 193 Pyrus baccata 35 communis 35 diversifolia 35 ioensis 35 malus 35 rivularis 35 sinensis 35, 40 296 BROOKLYN BOTANIC GARDEN MEMOIRS Pyrola americana 22 Scirpus microcarpus 127 \ elliptica 22, paludosus 206 secunda 22, 29 pungens 140 Pyrolaceae 22, 29 Scrophulariaceae 7, 64, 78, 86, 143, 147, 170, 177,20: Ranunculaceae 57, 85, 94, 104, 171, 216 Secale cereale 67 Rhamnaceae 159 Senecio aureus IOI Rhamnus purshiana 159 ductaris 101 Rhododendron californicum 21 harfordii 158 Rhus diversiloba 56 hydrophiloides 3 Ribes divaricatum 109 palustre Io1 lacustre 16, 109, 144 triangularis 3 sanguineum 109 Sidalcea oregana 165 saxosum 16 virgata 164 vallicola 16 Silphium perfoliatum 207 Romanzoffia sitchensis 160 Sitanion californicum 85 Rosa gymnocarpa 33, 51, 55 glabrum 85 nutkana 54, 55 hystrix 107 pisocarpa 51, 55 jubatum 85, 107 Rosaceae 33, 45, 47, 48, 49, 50, 51, 52, velutinum 85 53, 54, 55, 100, 178 Sium latifolium 206 Rubiaceae 25, 61, 156, 161 Solidago sp. 208 Rubus leucodermis 48 canadensis 4 neglectus 48 caurina 4 nigrobaccus 45 elongata 4 parviflorus 52 missouriensis 4 strigosus 48 rugosa 4 vitifolius 45 tolmieana 4 Rumex acetosella 60 Sorbus americana 37 hastatulus 60 aucuparia 35 paucifolius 152 hybrida 35 occidentalis 37, 43 Salicaceae 12, 14, 15, 16, 18 sambucifolia 35 Salix amygdaloides 15 Sphaerostigma Boothii 99 argophylla 16 dentatum 99 Bebbiana 14, 15 Sporobolus airoides 124 corda mackenzieana 15 Spraguea multiceps 209 cordata 15 Statice Armeria 181 discolor 14 Stellaria borealis 20 fendleriana 14 Stenanthium gramineum 110 lutea 14 occidentale 110 Piperi 15 Stipa comata 166 pseudocordata 15 Symphoricarpos albus 59, 168 scouleriana 15, 16 racemosus 59 sitchensis 14 Synthyris rotundifolia 177 Sanicula bipinnata 68 Santalaceae 8, 88 Taraxacum Taraxacum 169 Sarcobatus vermiculatus 124, 167 Tellima grandiflora 114 Saxifraga Marshallii 163 Thalictrum occidentale 85 odontoloma 163 Tiarella unifoliata 114. Saxifragaceae 114, 144, 163 Trautvetteria grandis 171 Scirpus americanus 140 Trifolium albopurpureum 198 JACKSON: UREDINALES OF OREGON Trifolium dubium 198 eriocephalum 198 Hallii 198 hybridum 211 microdon 198 oliganthum 198 pratense 187 procumbens 198 tridentatum 198 Triticum aestivum 107, 172 compactum 107, I51 diococcum 107 ovatum 172 vulgare 107, 151, 172 Tsuga canadensis 27 heterophylla 27 Tussilago farfara 100 Umbelliferae 46, 62, 68, 81, 98, 122, 123, 148 Urtica Lyallii 174 Urticaceae 174 Vacciniaceae 5, 27 Vaccinium caespitosum 27 Vaccinium canadense 27 macrophyllum 5, 27 myrtilloides 5 ovalifolium 5, 27 ovatum 5 parviflorum 5 pennsylvanicum 5 scoparium 5 Vitis-idaea 5 Valeriana occidentalis 89 Valerianaceae 89 Veratrum californicum 175 viride 175 Vicia americana 186, 203 linearis 186 truncata 186 Viola adunca 176 glabella 176 nephrophylla 176 rugulosa 176 Violaceae 176 Xanthium sp. 140 Zygadenus elegans 110 EVOLUTION BY HYBRIDIZATION EDWARD C. JEFFREY Harvard University Not long ago we were told that the investigation of the problems of evolution had left behind the inexact if broad phase of study in the field and had now entered upon the more accurate and satisfactory stage of quantitative elaboration under laboratory conditions. Leay- ing aside the question whether whatever exactitude in connection with this tendency has not been more than offset by a corresponding narrowness of outlook, it is now quite apposite to inquire if the experi- mental methods of the physiologist are in reality in the position to supply final light upon the fundamental problems of evolution. The judging of living beings by what they do rather than by what they are, has made notable progress in recent years. We are often told for example that an organ is the tool of a function and consequently should be defined by its performance rather than by its organization. I need not point out the essential fallacy of the physiological definition of an organ cited above. It obviously breaks down the moment it is used on any wide range of facts. Perhaps the most striking illustration of depending overmuch upon physiological data is supplied in connection with present investi- gations upon the all important question of the origin of species. It is practically universally assumed in genetical studies, that the capacity to breed true under exacting experimental conditions is the most reliable criterion of good species. It has for example assumed that breeding results obtained with Oenothera and Drosophila are of funda- mental importance for the science of biology. By those of us who have neither been intoxicated with the cult of the evening primrose nor bowed the knee in the temple of the god of flies, this conclusion will in general be held undemonstrated. We must obviously know a good deal more about the antecedents of those forms which have been raised in recent years to the dignity of veritable biological touch- stones, before we can admit the validity of the far-reaching con- clusions drawn from their genetical behavior. The question of the origin of the species is as much with us ae as it was at the time of the publication of Darwin’s epoch-making work. Darwin himself ultimately ventured no explanation of the causes of the changes concerned in the formation of new species, but 298 JEFFREY: EVOLUTION BY HYBRIDIZATION 299 contented himself mainly with pointing out that a general process of variation has been going on from age to age in matter endowed with life. He emphasized the fact that the struggle for existence on the one hand and the selection exercised by environment on the other provided an important directing influence upon the development of new species of plants and animals. In recent years a doctrine of old standing has been revived, namely the hypothesis of mutation. It has been maintained that new forms or elementary species arise spon- taneously from formerly existing species. This doctrine has been particularly advanced by the activities of the Dutch physiologist De Vries and his disciples in this and other countries. It is a general observation in connection with the activities of the lower organisms that in the process of their often extremely active development they give rise to inhibiting substances. In the case of the common yeast for example we have the formation of alcohol, which finally, by a high degree of concentration in fermenting sugary solutions, brings the activity of the yeast organism to a close. It is of interest to note in this connection that it is precisely in Holland that scientific opposition to the mutation hypothesis of De Vries has recently appeared. To Dr. Lotsy we owe a recent volume on Evolution by Means of Hybrid- ization, which attacks the mutation hypothesis at its very base through the contrary hypothesis that all changes in living matter are due to crossing or hybridization and are not the consequence of spontaneous internal or mutational phenomena. The author argues that since hybrids are notoriously variable all variability must be due to hybridism. This appears to be reasoning in a vicious circle. Clearly the most definite evidence in regard to hybridism as the cause of new species should be demanded before the possibility of the appearance of new types in this manner can be admitted. We fortunately have extremely good testimony on this subject from the earlier investigations published by Kerner in Austria and Brainerd in this country. Kerner in his well-known Pflanzenleben as well as in an earlier publication in the Oesterreiche Botanische Zeitung has brought forward much evidence as to the origin of new species as the result of hybridization in the mountainous regions of eastern central Europe, where the floras of the Pontic, Mediterranean and Baltic areas meet. It is impossible within the time at my disposal to make more than a very brief reference to the results reached by this writer. He has made it clear that the members of different floras are very apt indeed to produce new species by hybridization in nature and that these species, where they are advantageously equipped as compared with the parent forms, flourish within the same region. In case they have qualities which enable them to live where the parental species 300 BROOKLYN BOTANIC GARDEN MEMOIRS are not able to survive successfully they are found to the exclusion of one or both of these. This last conclusion upsets the conventional assumption that hybrids can only exist where their originating species occur side by side. It is clear from the general results of the highly important systematic and geographical investigations of Kerner that new species may appear as the result of spontaneous hybridization. The more recent evidence supplied by the investigations of Brainerd upon the violets and certain Rosaceae point equally positively in the same direction. This author has made it clear that a number of recognized species of Viola and Rubus are in reality hybrids in their origin. A particularly interesting result reached by Dr. Brainerd is that these hybrid species may become absolutely fixed in spite of their mode of origin and respond not only to recognized systematic but also to genetical criteria for species. It is too often assumed at the present time that the best criteria of species are physiological. On this basis the capacity to breed true in cultures and to produce offspring which comply with the tests of genetical analysis is regarded as of paramount importance. Since many known hybrids comply equally with recognized species with these tests it has become clear that what a plant does in cultures can- not be accepted as an infallible evidence of its antecedents. Where physiological criteria fail, we turn to the more constant ones furnished by morphological characters. It has been recognized for nearly a century that sterility is often a marked feature of hybrids, especially when they result from the crossing of somewhat incompatible species. The causes of incompatibility are apparently unknown as often species more different in their external characteristics and more widely separated in geographical range can be crossed with greater success than those nearly related on the evidence of external features and geographic coincidence. For example the horse and the zebra produce fertile hybrids, while the horse and the ass, when crossed, give rise ordinarily to infertile mules. Similarly our common canoe, yellow and black birches, which often grow side by side without hybridizing all apparently cross with a considerable degree of readiness with the more isolated shrubby birch of swamps, Betula pumila, according to the investigations of Jack and Rosendahl in this country. Hybrids may present in the case of plants a number of interesting morphological characteristics. For example we frequently find a high degree of imperfection in their gametic cells, male and female, with the emphasis of sterility nearly always on the male. This feature is often so marked that it is impossible to fertilize a hybrid with its own pollen, even when the ovules present a considerable degree of fertility. The morphological imperfection in pollen grains can obvi- JEFFREY: EVOLUTION BY HYBRIDIZATION 301 ously be most easily estimated and varies in proportion from a small percent to complete sterility. The sterility resulting from hybrid- ization should not be confused, as sometimes happens, with sterility arising from purely physiological causes. For example the common horse radish, Lilium bulbiferum and L. candidum, under ordinary conditions do not set seed, by reason of the fact that the assimilates are too strongly determined to the subterranean parts to permit of the necessary materials being set free for the formation of seeds. It has been found however that the girdling the top of the subterranean stem in the horse radish or cutting off the flowering axis in the case of the lilies, brings about the formation of normal seeds. Similarly very marked climatic change or subjection to starvation or other extremely unfavorable physiological conditions results in the degeneracy of reproductive as well as other parts. Conditions like these are, how- ever, very easily distinguished from the sterility normally resulting from hybridization. Sterility in hybrids is of particular interest from the genetical standpoint because it more or less completely upsets the expectancy of Mendelian ratios in cultures of the offspring of species hybrids. This is doubtless one of the causes why the Mendelians have in general manifested so little interest in the genetical study of hybrids between natural species. Obviously however if the crossing of species in nature is a common cause of the origin of new species this part of the evolu- tionary field cannot be safely neglected if we are to reach any broad and permanently valid conclusions as the modus operandi of the origin of species. Another feature in the organization of hybrids is the frequent in- crease in the typical generic chromosome number as a consequence of crossing. For example we find in the much crossed oriental species of Chrysanthemum a number of chromosomes in the gametic nuclear divisions varying from 9 (the normal) to 18, 27, 36, and even 45, in other words, two, three, four and five times the normal gametic number. Similarly in another compositaceous genus Dahlia we find in the species D. coronata sixteen chromosomata in gametophytic divisions while in the hybrids between D. variabilis and D. coccinea thirty-two chromosomes have been enumerated. One further example will point the situation. In the monotypic Liriodendron and in certain species of Magnolia, nineteen gametophytic chromosomes have been counted, while in M. soulangeana (suspected of hybrid origin) as well as M. Yulan and M. grandiflora twice that number or more of chromo- somes have been observed. If we contrast the situation in these examples with that presented by the genera Pinus and Lilium, which are not at all prone to hybridism, we note a curious contrast. In the 302 BROOKLYN BOTANIC GARDEN MEMOIRS species of the pine or lily, the chromosomes are always of the same number and do not vary as in the examples cited above, as occurring in connection with hybridism. Frequently the addition of chromo- somes under hybrid conditions is not simply a doubling, tripling, etc., of the original number but a mere arithmetical addition. Among the vascular cryptogams and the mosses similar cases of multiplication of the normal number of chromosomes are frequently found in species growing in very damp situations or actually in the water and in which hybridization is accordingly favored. Having enumerated a few of the morphological characteristics of hybrids we are now in the position to apply the facts elucidated to the case of the Oenotheras, which have been brought particularly into prominence in connection with the mutation hypothesis of De Vries. In the so-called mutants of Oenothera lamarckiana as well as in that species itself, we find all the cited stigmata of hybridism as presented by incompatible species, namely a high degree of sterility, amounting in some cases to complete impotency, particularly in the male gametic cells, failure to segregate in accordance with Mendelian ratios and the multiplication of the number of chromosomes beyond the normal gametophytic number seven, or sporophytic fourteen. In O. gigas the gametophytic chromosomes are 14 instead of the normal seven, while in O. semigigas there are 21 in the sporophyte instead of the normal fourteen and in O. lata, O. semilata and O. rubricalyx, fifteen. Similar conditions have been described in many other species of Oenothera and their so-called mutants. It accordingly appears abundantly clear when morphological considerations are brought into court as well as the physiological data derived from experimental breeding that the genus Oenothera is obviously affected by contamina- tion through hybridization in its various species and their so-called mutants. The conclusion may accordingly be drawn that so far as the genus Oenothera throws light upon the origin of new species at all, it vouches rather for the multiplication of species as a consequence of hybridization than for their appearance as a result of the mysterious process of mutation. In conclusion we may turn to an objection which has been raised by De Vries and other mutationists to the interpretation of morpho- logical sterility as an evidence of hybridization. It has been claimed that this feature is an accompaniment of mutation. It is most un- fortunate for the mutationists that a phenomenon so generally recog- nized as associated with the crossing of species should at the same time occur in mutating forms. The burden of proof that such forms are not of hybrid origin distinctly lies with the mutationists. We have however positive evidence that this is not a possible interpre- JEFFREY: EVOLUTION BY HYBRIDIZATION 303 tation. In monotypic genera such as Ginkgo, Liriodendron, Calla, Spathyema, etc., the pollen grains under normal physiological condi- tions of development are all alike and perfect. In Fig. 1, Plate V, is shown the pollen of Zannichellia palustris, a species isolated in our northern North American flora. It is clear that the grains are strik- ingly uniform and are all well developed. For comparison with the genus just mentioned which has very few species and consequently cannot be considered as highly variable, let us take the common pondweed Potamogeton, of which there are very many species. Fig. 2 shows the situation in the large genus just mentioned. The cells are not by any means all perfectly developed and are conspicuously char- acterized by extreme variations in size. It might be maintained on this basis of the illustrations furnished in Figs. 1 and 2 alone that variability in size of pollen grains is associated with the multiplication of species or in other words with the phenomenon of mutation. Against this view in the forms under discussion may properly be urged the fact that many natural hybrids between species of Potamogeton are known which manifest the usual morphological features of such forms. A clearer elucidation of the situation is furnished by the conditions in large genera, where a number of the species coincide both in geo- graphical distribution and in the time of flowering. As a first illus- tration in this connection may be taken the genus Rubus, which has recently been investigated by Dr. Hoar. Fig. 4 shows the condition of the contents of the anther in R. villosus. Clearly the pollen varies greatly in size and perfection of development. A similar condition has been described by the author just cited in a large number of the species of Rubus. The general situation might be interpreted in view of the very numerous and at the same time very variable species of Rubus as an argument for the correlation of mutation and _ pollen sterility. When however the facts in species of the genus, which are in some manner isolated, are examined quite a different light is thrown on the subject. Fig. 3, Plate V, illustrates the pollen of R. odoratus, the flowering raspberry, which opens its blossoms at a considerably later period than the mass of the species of the genus. Care has been taken to include as large a number as possible of the grains in the field -of view. It is obvious that the variation in size and frequent defective development of unisolated species of Rubus, are conspicuous by their absence. If irregularities in the development of the contents of the anthers were a feature correlated with mutation in the genus Rubus then we ought to find it equally present in isolated and unisolated species. Since that is not the case, the natural inference is that the sterility present in the pollen of species subject to hybrid contamina- 21 304 BROOKLYN BOTANIC GARDEN MEMOIRS tion by reason of their coincidence of flowering periods, is actually the result of previous specific crossing. This view of the matter is strongly confirmed by the fact that the investigations of Brainerd and others on that genus have revealed a large number of natural hybrids. For a parallel case we may now turn to the genus Ranunculus. If any of the species which flower in the early summer are examined, such as R. acris, R. repens, R. aquatilis, R. Cymbalaria, a considerable proportion of imperfect pollen is usually present and frequently the grains vary extremely in size. This situation is shown for R. acris in Fig. 6, Plate V. Obviously there is a great range of size in the grains and some are imperfect. This condition is most naturally interpreted as a consequence of previous hybridization. Fig. 5, Plate V, illustrates the condition of the pollen in R. rhomboideus, a species which flowers in the very early spring long before the other species of the genus have opened their blossoms. The numerous grains shown in the illustration are clearly well developed and somewhat uniform in size. In the species under consideration as well as in R. odoratus, perfection in development of pollen is clearly correlated with isolation from possibility of contamination with other species. In view of the facts supplied in the above instances, which might be almost indefinitely multiplied in representatives of other angio- spermous families, it appears clear that the obvious interpretation of pollen sterility is to be adopted, namely as an indication (where it occurs under normal conditions of growth) not of mutability but of previous hybridization. The large situation which is so _ briefly illustrated by the accompanying photographs, indicates the necessity of bringing morphological criteria relating to the cytology and develop- ment of the gametic cells (pollen and embryo sacs) into court, as well as the data derived from genetical behavior, in speculations in regard to the origin of species. It seems clear from the evidence supplied on the systematic and phytogeographical sides on the one hand and that from the standpoint of morphology on the other, that the crossing of species in nature is an extremely common cause of the multiplication of species. It is further obvious that physiological and genetical criteria must not be given greater weight than the more reliable ones supplied by actual history and by morphological structure, in the all-important biological question of the origin of the species. It is finally apparent that the genetical status of the Oenotheras is so dubious that they cannot be brought into court to furnish decisive evidence in favor of the muta- tion hypothesis of De Vries. It may be added in conclusion that the multiplication of species by hybridization does not by any means invalidate the Darwinian hypothesis but merely supplies an additional BROOKLYN BOTANIC GARDEN Memoirs. Volume I, PLATE V. JEFFREY: EVOLUTION BY HYBRIDIZATION JEFFREY: EVOLUTION BY HYBRIDIZATION 305 agency for the formation of species. It appears moreover logically impossible to regard hybridization as the universal and sole cause of the appearance of new species, as has been,recently maintained by Lotsy in his Evolution by Means of Hybridization, since the original species must have come into existence by some other means than by hybridization. The adaptation of the floral structures of the Angio- sperms to cross fertilization, emphasized many years ago by the Austrian botanist Kerner is doubtless of significance in connection with the ever-increasing volume of evidence for the wide occurrence of natural hybrids in this large and successful group of seed-plants which have to so notable a degree furnished the facts for the existing general biological theories. DESCRIPTION OF PLATE V Fic. 1. Pollen of Zannichellia palustris, showing great uniformity in a species unable to hybridize. X 400. Fic. 2. Pollen of Potamogeton diversifolius, showing great diversity of size and development in pollen of a species subject to hyridization. X 400. Fic. 3. Pollen of Rubus odoratus, a species which flowers late and consequently is not subject to crossing. X 400. Fic. 4. Pollen of Rubus villosus, a species flowering with a number of others and consequently subject to hybridization. X 400. Fic. 5. Pollen of Ranunculus rhomboideus, showing uniformity in a species exempt from crossing by early date of flowering. XX 4oo. Fic. 6. Pollen of Ranunculus acris, a species flowering at the same time as a number of others and consequently exposed to hybridization. X 400. A METHOD OF OBTAINING ABUNDANT SPORULA- TION IN CULTURES OF MACROSPORIUM SOLANI E. & M. L.. O. KUNKEL Bureau of Plant Indusiry, U. S. Department of Agriculture Although the early blight fungus, Macrosporium solani, often fruits abundantly when growing as a parasite on potato leaves, it usually does not bear very many spores when grown in pure culture. Jones (4) reports that some of his cultures when old fruited rather freely. Jones and Grout (5) state, however, in their technical de- scription of the organism as Alternaria solani (E. & M.) Sorauer that it sporulates “sparsely in pure cultures.”’ Enough spores may be obtained by growing it in the ordinary way on culture media to test its parasitism to the potato plant. Galloway (3) performed this experiment as early as 1893 and Jones (4) repeated it a few years later. Nevertheless, the failure to obtain spores in quantity from pure cultures has made it impossible to perform ex- tended infection experiments with this important parasite. The writer found M. solani doing considerable damage in the potato fields of Aroostook County, Maine, last August, and in the hope of obtaining a strain of this fungus that would fruit abundantly in pure cultures, a considerable number of isolations were made. Cultures were in each case made from single spores. The organism was isolated from fifty-four different potato plants selected at random in half a dozen potato fields in the vicinity of Presque Isle, Maine. All of these single-spore strains were grown on a number of different culture media, including potato agar, string-bean agar, prune agar, and glucose agar. The several strains showed considerable differences in the appearance of their growth in culture, but none of them pro- duced more than an occasional spore on any of the media tested. In a former paper the writer (6) has described a method of retarding the growth of Monilia sitophila (Mont.) Sacc. by lowering the vapor tension of the atmosphere above pure cultures. It was recalled that by checking the mycelial growth in this way the fungus could be made to fruit more abundantly than when grown in a moist atmosphere. In the hope that this method might serve to induce sporulation, cultures of MW. solani were subjected to like treatment. More spores 306 KUNKEL: SPORULATION IN MACROSPORIUM SOLANI 307 were obtained in these cultures than when the atmosphere above the fungus growth was allowed to remain near saturation. It was found, however, that this method of treatment would not bring about very abundant sporulation in the case of the early blight organism. A number of other methods were tested, and the idea of wounding the mycelium was finally hit upon. This seems to be the stimulus neces- sary to bring about abundant sporulation in cultures of M. solant. It will fruit profusely on any of the media above mentioned if the mycelium is thoroughly wounded after the culture is two or three days old and has made a good start. So long as the mycelium is undisturbed it grows very vigorously through and over the surface of most culture media. In these cultures very few conidiophores are produced and very few spores are developed. Undisturbed cultures often fail to Fic. 1. A wounded hypha bearing conidiophores of Macrosporium solani. X 650. Fic. 2. A wounded hypha of Macrosporium solani showing the production of conidiophores near the point at which the hypha was cut. goo. bear a single spore. If, however, the radiating mycelial strands are severed at the proper stage in the life of the culture, thousands of conidiophores, each bearing a spore, will develop from the cells of these hyphae. The wounding may be accomplished by scraping the surface of the culture with a sterile scalpel or even with a strong platinum needle. The more thoroughly it is done, the greater will be the quantity of spores produced. Conidiophores arise abundantly near the point at which the mycelium is broken. Sometimes each consecutive cell of the mycelium for a considerable distance produces one or more conidiophores. Such a series of fruiting cells is shown in Fig. 1. At the distal end of some 308 BROOKLYN BOTANIC GARDEN MEMOIRS of these conidiophores may be seen a scar. This indicates the point at which the spore was attacked. The production of conidiophores near a wound is shown in Fig. 2. In Fig. 2 may be seen a portion of a cell that was killed when the hypha was cut. Each of the photo- micrographs (Figs. 3 and 4) shows portions of the surface of two different string-bean agar cultures of the early blight fungus. These photographs indicate the abundance of spore production when the mycelium is properly wounded. Many thousands of spores may Fic. 3. Fruiting culture of Macrosporium solani. This photograph of the surface of a string-bean agar culture shows the abundance of spore production when the mycelium is properly wounded. X 15. thus be obtained from a single Petri dish culture. Spores produced in this way were sprayed with an atomizer onto potato plants growing in a greenhouse. They produced good infection not only on the older leaves but on young leaves as well. The method of stimulating spore production by wounding was tested out on each of the single spore strains isolated from the potato KUNKEL: SPORULATION IN MACROSPORIUM SOLANI 309 fields of Maine. They all responded in like manner, producing spores in great numbers. The method has also been applied to other Macro- sporiums that do not fruit readily in culture. M. tomato, Cooke, a parasite of the tomato and M. daturae, Fautr., a parasite of the jimson weed, Datura Stramonium L. respond in the same way to the wound- stimulus. Although M. daturae fruits quite freely without this stimulus, the number of spores produced in any culture can be greatly increased by wounding. MM. tomato fruits even more sparingly than Fic. 4. Fruiting culture of Macrosporrum solani. This picture shows the spores more highly magnified and indicates their abundance in a wounded string- bean agar culture. XX 250. M. solant in unwounded cultures. When thoroughly wounded it bears spores in great numbers. The abundant sporulation of M. solani in culture makes more easy the study of its fruiting stages. The successive steps in the development of conidiophores and spores can easily be observed. The mature conidiophores are always several-celled. They may arise singly or in whorls, as is shown in Fig. 2. The typical conidio- 310 BROOKLYN BOTANIC GARDEN MEMOIRS phore bears a single spore. Successive production of spores on the same conidiophore, such as has been described by Miyabe (7) for M. parasiticum Thiim., has not been observed for M. solani. Occa- sionally the spores are borne in chains of two. This may occur quite frequently when the atmosphere above the culture is saturated with water vapor and only a few conidiophores are being produced by a vigorously growing mycelium. It occurs very seldom or not at all in cultures that are sporulating abundantly. The occurrence of spores in chains of two is exceptional for M. solani. The writer has never observed longer chains and is inclined to the view that Duggar (1) is correct in leaving this fungus in the genus Macrosporium rather than to put it into the closely related genus Alternaria as Jones and Grout (5) have done. Duggar’s objection that the catenulate method of spore production has not been seen except in artificial culture does not hold, however, for the writer has more than once observed the spores in chains of two on the potato leaf. M. daturae when grown under very moist conditions also produces its spores in chains of two. This fungus in rare instances even produces spores in chains of three. Under more normal conditions, however, the spores are always borne singly. As one finds it growing on its host or observes it in pure cul- ture, the catenulate method of spore production is exceptional, and it seems doubtful whether any useful purpose would be served by transferring it to the genus Alternaria. The writer has examined many fruiting cultures of the tomato parasite, but has never observed it producing spores in chains even under very moist conditions. Besides usually producing their spores singly, both the early blight fungus and the jimson-weed fungus show other characteristics which seem to put them with the Macrosporiums. They both possess a coarser mycelium than is usual for the genus Alternaria. Their spores are larger and under most conditions are produced in smaller numbers than is common for an Alternaria. Moreover, their spores are normally attenuated into a beak similar to that on the spores of other species of Macrosporium, such as M. catalpae, E. & M., M. cucumerinum, E. & E., M. caudatum, C. & E., M. concentricum, Winter, M. brassicae, Berk., M. porri, Ellis, and M. tomato, Cooke. M. caricinum, one of the four species mentioned by Fries (2) at the time he established the genus, bears spores which are, according to his description, attenuated at both ends. No such beaks are to be observed on spores that are borne in chains. Since M. solani, M. tomato and M. daturae are parasitic on closely related plants, they have by some authors been considered identical. Sorauer (9) seems to have had this notion and Duggar (1) states that M. solani ‘“‘is found not only upon the potato but also upon tomatoes KUNKEL: SPORULATION IN MACROSPORIUM SOLANI my) | and upon the jimson-weed (Datura stramonium).” By growing the Macrosporiums obtained from these three hosts side by side in pure culture it is easy to observe that they are by no means alike. Not only are they different culturally, but the spores they produce are quite unlike morphologically. The spores of M. daturae have an attenuated beak that is very much longer than the beak on the spores of M. solani or M. tomato. The beak on the spores of M. tomato is finer than the beak on the spores of the other two species. The mycelium of M. tomato is also finer than the mycelium of the other two forms. The spores of M. daturae are larger and the spores of M. tomato are smaller than those of WM. solani. On such media as string- bean agar and glucose agar the three fungi show wide differences. M. solani produces a gray felty growth on string-bean-agar and usually colors it red. No such color is to be observed in the case of M. tomato and M. daturae on the same substratum. The growth of M. solani on glucose agar is a rusty gray color. Colonies of M. tomato and M. daturae on this same medium are blue in color. M. daturae on many different media produces colonies showing marked zonation, such as is not to be observed with the other two forms. These three parasites are unlike in so many different ways that the writer believes they should be considered separate species, rather than strains of a single species. It is, of course, probable that either fungus may infect more than one host. Sorauer (8) reports that he was able to infect tomato leaves with the Macrosporium from the potato. This does not prove, however, that the three hosts as they grow in nature are attacked by one and the same fungus. An examina- tion of the spores of the three fungi when grown side by side in wounded cultures brings evidence that this is not the case. The method of obtaining abundant sporulation in cultures of M. solani here described makes possible more extended infection-experi- ments than have hitherto been undertaken. It is believed that it will also be of service in any study of the systematic relationships of the genus Macrosporium. The principle involved in the response of the early blight fungus to a wound-stimulus is one well known to science. Conditions unfavorable to vegetative growth often lead to fruiting in unprolific plants. The orchardist recognizes this truth when he prunes his trees or feeds them with fertilizers poor in nitrogen compounds. BIBLIOGRAPHY 1. Duggar, B. M. Fungous Diseases of Plants. Ginn and Co., New York, 1909. 2. Eries, H. (Syst. Myc: 32373. 1829. 3. Galloway, B. T. The Macrosporium Potato Disease. Agric. Science 7: 370- 382. 18093. 312 BROOKLYN BOTANIC GARDEN MEMOIRS aN . Jones, L. R. Potato Blights. Vt. Agric. Exp. Sta. R.g: 88. 1896. . Jones, L. R. and Grout, A. J. Notes on Two Species of Alternaria. Bul. Tor. Club 24: 254-258. 1897. 6. Kunkel, L. O. Physical and Chemical Factors Influencing the Toxicity of Inorganic Salts to Monilia sitophila (Mont.) Sacc. Bul. Tor. Club 41: 265- 293. I914. 7. Miyabe, K. On the Life History of Macrosporium parasiticum Thiim. Ann. Bot. 3: 1-26. 1889. 8. Sorauer, P. Auftreten einer dem amerikanischen “ Early Blight ’’ entsprech- enden Krankheit an den deutschen Kartoffeln. Zeits. f. Pflkr. 6: 1-0. 1896. 9. —— Handbuch der Pflanzenkrankheiten. 2: 455, Berlin, 1908. uo SYNCHRONISM IN PLANT STRUCTURES JOHN MUIRHEAD MACFARLANE Unwwersity of Pennsylvania In all departments of botanical inquiry it is becoming increasingly evident that wide observation and exactness of record are indispens- able, if we are to reach wide and exact conclusions as to plant life. So the carefully tabulated experiments of Koelreuter, Gaertner, Herbert, Darwin, Mendel, Vilmorin and others regarding plant cross- ing during the past century were the appropriate starting points for the more extended and exact results that are now being secured by plant breeders. The characteristic also of exact and correlated behavior on the part of plant organisms powerfully impressed the writer as he advanced in his studies of parent and hybrid types, from 1889 onward. Not the least striking of his results were those bearing on the relation of plants to environal atmospheric agents or stimuli, such as light, heat and water supply.t. So alike as regards constitutional vigor and period of blooming as for chemical nature, color, and odor of hybrids, it was concluded that each detail was more or less exactly between that of the parents; ‘‘while some vary to a greater or less degree from one or other parent.’’ Impressed, therefore, by such conditions, the writer has observed closely, during a period of twenty-five years, the action of those environal agents which we speak of collectively as climatic conditions, and the reaction of plant parts to these agents, with a view to determining how exactly each plant organism is corre- lated with its environment. This line of inquiry has received con- siderable attention during the past seventy-five years, under the term ‘‘phytophenology.”’ But the study, as well as the results se- cured, have been very largely ignored by botanists, or even ridiculed by some as yielding no conclusions of value. We would emphatically assert that few lines of investigation will compare with this if the studies are prosecuted in exact manner, and are planned so as to cover a definite field. The present communication may be suggestive in connection with future possible developments at such an experimental institution as the Brooklyn Botanic Garden, which has already had so successful a history under its able director. 1 Gard. Chron. 93: 753. 1891. Trans. Roy. Soc. Edin. 37: 255. 1892. 313 314 BROOKLYN BOTANIC GARDEN MEMOIRS Plant variation as due to varying environal stimuli is a phenomenon witnessed everywhere around us. The difference in size and color between similar plants growing in shade and sunshine; the difference in growth and habit between plants exposed to moist rich soil and to light dry sand; the difference in time of leafing, blooming, and fruiting between plants situated at lower and higher levels, are facts that are familiar to all. But the fundamental causes of such differences, as well as the important conclusions to be drawn, have hitherto been too much overlooked. Even the records of leafing, the blooming and fruiting of flowering plants, the shedding of spores by pteridophytic and bryophytic species, or the conjugation period of algoid and fungoid types have often been given in most haphazard, or totally misleading, manner in many of our local floras and manuals. We desire, therefore, to inquire how far such great seasonal con- ditions as the above can be reduced to exact limits, and if possible to ascertain what fundamental principles underlie their expression. The writer selects first the blooming period of higher plants as a phenome- non that all can witness and verify to greater or less degree in daily life. Given that some one locality is chosen where a certain number of individuals of a species are exposed to as exactly like environal con- ditions as possible, it may then be asked how nearly synchronous may the blooming periods be amongst these, and how correctly can we define these for any region. In illustration, the following may be cited from amongst many others that the writer has watched 2 Neglecting the wayward skunk-cabbage—that nevertheless can be reduced to system—the first plant to bloom each season is the silver maple. This year (1917) hundreds of trees opened many flowers synchronously on March 11, instead of on the 13th, as is averagely the case. -Furthermore, the opening took place about nine to eleven A.M. Favored by bright suns the expansion continued upward along the branches, as is averagely the case, for a period of nine days, and by this time the earliest flowers were beginning to push out their green fruits. If we compare now the same trees for previous years it may be said that during 1912, and as a result of continued snows and frosts, the unfolding occurred with equal abundance and exactness on March 17. In 1913 a remarkable record was made. The weeks of fall weather during 1912 were balmy and mild, and even at times warm. Asa result many heat units over the average were absorbed by the trees and caused precocious though unobserved preparation for spring unfolding in 1913. And here we would emphasize again, con- trary to views previously expressed by many, that record must be kept of environal conditions continuously throughout years, if true * The results recorded are given for West Philadelphia unless otherwise stated. MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 315 results are to be secured as to the action of environmental agents. To attempt to start the record from such an artificial period as the first of any year is to set an arbitrary limit to the continuity of changes and activity in vegetation. The short-lived snow and slight frosts of late November started the apparently needful winter maturation of tissues, and_ this was followed by an almost continuous series of genial days until the third week of January. So on the morning of January 19 abundant first flowers expanded on all observed trees, but a cold wave on the 21st split the flowering period in half, and only on March 2, with the advent of a bright day and warm sun did the opening again proceed until March 10. This striking result had not been paralleled through a previous period of at least thirty-five years. As a contrast, in 1914 ex- pansion occurred only on March 16, owing to the frosts and late tem- peratures occurring throughout February and on to March 15. In the neighborhood of Wayne, Pa., with an elevation of 475 feet, with greater exposure to cold winds and less influenced by the heat of a great city, the opening did not take place until the 22d of March. Records like the above that extend over more than twenty years would suggest that floral expansion is not a somewhat haphazard and irregular event, but is rather an exact reaction of an organism to definite and cumulated environal actions or stimuli. If such be true, we should expect it to extend probably throughout flowering plants as a group. Partial proof is subjoined. The red or swamp maple (Acer rubrum) each year succeeds the silver species in blooming at an average interval of twelve days. This year, eleven trees, observed in like locality, all opened on the morning of March 26, while the climax of blooming was reached on the 4th of April. - According to the valuable statistics secured by Dr. Mackay and his committee of observers, it may be instructive here to point out that the same species in Nova Scotia has expanded averagely on May 5, or 41 days later than in the Philadelphia region. The white poplar (Populus alba) is of exceptional interest from the standpoint of the present communication. Staminate catkins an- nually mature, and lengthen synchronously, amid like environment on a definite day, and the shedding of abundant pollen proceeds for one or at most two days. Thereafter the catkins soon shrivel and within a week have mostly fallen. The average blooming date is April 7, but this year, stimulated by the warmth of mid-March days the tassels suddenly lengthened on March 28. Pollen was completely shed by the 29th, and sidewalks over wide areas were covered with fallen tassels by the 2d of April. But though a comparatively rare 316 BROOKLYN BOTANIC GARDEN MEMOIRS tree, the pistillate plant of this dioecious species has been proved to mature its stigmas synchronously with the shedding of pollen from the staminate trees. The yearly variations from the above dates vary according to environal—mainly heat—stimuli. Thus in 1912, after a cold February and early March, succeeding favorable days caused complete pollen dispersal on the 27th and in part on the 28th. During the precocious springtime of 1913, the flowering occurred suddenly and uniformly on the morning of March 18, while in the suc- ceeding year like expansion took place on March 31. The Carolina poplar (Populus deltoides) is like the last species specially abundant in the staminate trees, rare in the pistillate ones. Its average bloom- ing period is April 16. In 1913 its behavior was arresting in the sud- den and exact procedure shown. Here, these trees under similar environment lengthened their catkins fully and started to disperse abundant pollen on March 24, between 9 and 10 A.M. of a bright warm day. The young catkins were almost emptied by 5 P.M., only a few of the smaller terminal flowers still retaining a quantity. By next day, scant remnants could alone be secured. But on that day four trees which grew on wind-swept and shaded street corners were found only beginning to dehisce. In 1914, owing to prolonged cool periods and warmer ones alter- nating, blooming occurred on April 6. Many catkins matured only on the 7th. During the present year, the action and reaction of environal energy and of organismal tissue have been most suggestive. For stimulated by the bright warm suns of March 31 and April 1 ‘some catkins lengthened gradually during these and succeeding days, and began to discharge pollen extensively on April 4. But rather low temperatures on that day and those succeeding, accompanied often by rain, prolonged reaction fitfully until the roth of the month. So we learn from such statistics, that have frequently been verified for the above, as well as for other species, that if a sufficient environal stimulus act quickly and continuously, an extensive synchronous blossoming may ensue, that is completed within a few hours each year. On the other hand, if temperature units be more gradually expended, and specially if such be combined with wetting conditions that prevent establishment of tissue tension in anthers, pollen discharge may be prolonged over a considerable period. The alder (Alnus incana) that averagely lengthens and opens its catkins on March 24, and that shows crimson papillose stigmatic surfaces in exactly synchronous manner, was in like state on the 25th, but in 1913, March 12 was the date. Pollen dispersal is usually com- pletely effected in three or at most four days, this lengthened period being due to differences in position and so in time maturation of the catkins on each twig. MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 317 The above are all shrubby or arborescent forms, and so are more directly subject to changes of temperature than are species that perennate by subterranean parts. The energizing factors are less complex in the former; while in the latter, warmth, moisture, porosity and chemical composition of the soil, become highly important govern- ing factors. So the digestion and transfer of the reserve foods in rhizomes, corms and bulbs is effected more gradually and the flowering period is usually more prolonged, though the exactly synchronous unfolding of the first blooms is as striking as in any of the shrubs or trees. Of our three commonest spring flowers, Hepatica triloba, Sanguinaria canadensis and Claytonia virginiana, the two first aver- agely appear in bloom on April 9, and open successive flowers for a period of 18 days in Hepatica, 10 to 12 days in Sanguinaria and about 25 days in Claytonia. Claytonia appears in bloom on April 12, aver- agely. But in 1913, our woods showed a sudden unfolding for the first two on March 18 and of the last on March 20 or 21. Another herbaceous plant deserves special notice here as illustrating an interesting phase in synchrony, namely, the dandelion. Like the English daisy (Bellis perennis), this is a hardy plant which retains wintergreen leaves and shelters amid close grassy sod. So very slight changes of temperature in winter will cause both of these to unfold their earliest flower-heads in apparently regular manner, specially if growing in sheltered sunny places. But such by no means represents the first exact growth period for the season, which for the dandelion occurs averagely on April 23. Then, instead of the scant or occasional heads of earlier date, our lawns show a sudden yellow coloring by 9 A.M. that is continued for almost a month thereafter, as successive heads expand and as the florets in each successively open. During 1913, the behavior was noteworthy. For lawns were abundantly yellowed over from January 16 to 20. But all suddenly closed and were destroyed by frosts that succeeded from January 21 to March 2. Then came a warm stimulating March, with the result that from the 15th of the month onward dandelions were abundant. In connection with his graduate class on the Gymnospermia, the writer became interested from 1898 onward in the behavior of the Japanese ginkgo tree (G. biloba), at first only with the aim of securing appropriate material, but as the years passed the phenological relation became of equal interest. Two large staminate trees grow near the historic old Hamilton Mansion, adjoining the University Botanic Garden. These suddenly and synchronously lengthened their catkins in 1898 on the morning of May 2, and when visited on the succeeding day few were still polleniferous. This suggested to the writer a closer study of the subject from the standpoint of individual and species 318 BROOKLYN BOTANIC GARDEN MEMOIRS behavior. During the previous two years he had secured scant supplies of good seeds from a large pistillate tree fronting the old Jones Home at 65th and Callowhill Sts. Throughout the summer of 1898 he examined it from time to time, and noted that the seeds ma- tured wholly on the southeast side, though no staminate tree was then known to exist for miles around. In the succeeding spring the two staminate trees matured between 9 and 10 A.M. on May 5, and by the 6th were equally free of pollen as before. But the possible synchronous relation of these to the pistillate tree was now determined. For, accompanied by one of his students and laden with staminate branches, a visit to the latter tree was made on the 5th. A ladder was secured and examination of the small green ovules on the branches clearly revealed that each was exuding a shining viscous droplet for pollen entanglement. The staminate branches were hung over the western side of the tree and then shaken. As summer advanced the abundant maturing ovules alike supplied wealth of material for study, and by their structure showed that perfect pollination had been effected. Continued study of the above trees in succeeding years showed that synchronous maturation and rapid pollen discharge annually took place as early as April 20 in 1913 and as late as May 22 in 1904, according to the stimulating amount of heat units, of rain condition and of soil moisture. In time also as the above statistics became known, the writer learned of staminate trees on the grounds of Girard College, and Laurel Hill Cemetery, which doubtless had con- tributed pollen to the pistillate tree, widely removed from them, during previous years. : The peach (Prunus persica) deserves attention as being an intro- duced woody plant, that bears attractive flowers and valuable eco- nomic fruit. Trees under like environment open averagely on April 22 and then in considerable numbers. Climax of blooming is reached five days thereafter and within another four days the flowers have all fallen. But in 1913 a sudden wealth of bloom appeared on April 1. It need scarcely be added here that a synchronous activity amongst pollinating bees was a feature of the event. The common field daisy, that like the dandelion and English daisy, are all European and introduced weeds, differs markedly nevertheless from the other two in that it shows no unseasonable flowers through- out the winter months and does not even unfold as a harbinger of spring. For averagely a sudden wealth of flower heads expands on May 24, and for a month thereafter added heads appear in what may be—did we only know accurately enough—regulated succession. Almost exactly a month after the last, the first flowers of the intro- duced moth mullen (Verbascum blattaria) come into bloom on June MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 319 26, and in wonderfully exact and graded succession, with definite time interval between each, later flowers open along the elongating axis. But just three days before, or on June 23, the central flowers of the cymes on our common Indian bean (Catalpa bignonioides) expand, and for about 16 days thereafter successive blooms open in exact ratio, if weather conditions are favorable. But a decided retardation may occur, if cold winds and wetting rains interfere. We would draw attention now to some cases of synchrony in floral parts. From the time of Linnaeus onward descriptions of floral clocks have been frequent, and no matter what value we attach to such, the very device points to a surprisingly exact time during the twenty-four hours when the blooms of each species open. But the maturation and opening of each flower, as well as the behavior of such parts as the stamens and styles of it, may vary according to the degree of thermic energy, or lumic energy, or both that act on these. Such variation seems to explain apparently contradictory results that have been recorded by different botanists. Thus Kerner’s attractive state- ments® regarding Silene nutans and its successive maturation of one row of stamens in each flower, on one evening, of a second row on a succeeding evening, and of the styles on the third, have called forth adverse comment along with the statement that no such exact suc- cession occurs. But first about twelve years ago along the Trafoi Ravine in the Tirol, and nearly seven years ago by Morgarten Field in Switzerland, the writer was able to prove that both statements might apply according to environal conditions. For when the days and nights in August are warm, with clear sky overhead, the exact succession noted by Kerner can readily be traced. But if the days, and even: more the nights, be cold, raw and at times accompanied by rains, then a nearly simultaneous maturation of all ten stamens and at times even of the styles may occur. _ By careful observation, however, during warm dry days we were able to determine an even more exact and synchronous behavior of the ten stamens than that noted above. For on the first evening from about 6 to 7.30 the anthers of the earlier 5 stamens matured in circular succession with a clear time interval between each, and then were ready to scatter pollen; while on the succeeding evening, the second five ripened similarly. Such again calls to mind the time period shown in maturation of the five anthers on the stamens of the giant cow parsnip (Heracleum lanatum). It was found even that the dropping of the anthers in Silene mentioned by Kerner takes place not together, or irregularly, but in correlated succession if environal stimuli are favor- 3 Nat. Hist. of Pl. 2, p. 154. 22, 320 BROOKLYN BOTANIC GARDEN MEMOIRS able. But here again this is duplicated by Heracleum and other Umbellifers, as well as by many Araliads. The writer can never forget his first knowledge obtained for the almost exactly synchronous floral expansion in Oenothera grandiflora. Two enthusiastic botanical lady friends had cultivated many of these handsome plants along with numerous other species nearly twenty- three years ago. He was asked to pay a visit about 7 P.M. on a mid- July evening. Chairs were set out amongst a group of the Oenotheras and he was asked to watch and listen. From 7.15 to 7.50 a constant succession of “‘pfuffs’’ was heard, that indicated the bursting of the sepals and unfolding of the petals, which rapidly took place before one’s eye. From the condition where scarcely a flower was open to begin with, to that seen at 7.50 the change was striking, for now the plants were gay with large expanded blossoms. Six years afterward the writer was carried by train from Botzen to Meran in the Tirol toward 7 P.M. and running for miles by the banks of the Adige River he witnessed the same synchronous series of events for plants of Oenothera that had been introduced there. In connection with genetical studies increasing attention has been paid during the past fifteen years to the behavior of varieties and hybrids. But extremely little has been published as to the relative period of flowering, fruiting or like phenological conditions for each parent and for the hybrid. But a very wide field for exact observa- tion is here awaiting study. The writer has drawn attention to some results and has since accumulated others. Thus, the relative pro- duction or not in the wild state of hybrid Sarracenias is almost wholly determined by the synchronous or asynchronous relation of the flowers. So the scarcity of wild hybrids of S. rubra with other species is in part due to difference in locality, but in large measure to later blooming period of that species. Under cultivation by placing plants in greenhouses of different temperatures a synchronous blooming can be effected, and such striking hybrids as S. Popet and S. Chelsonii represent the progeny. In such cases then an exact expenditure or retardation of definite heat units effects a synchrony that in their natural environment does not exist. The practical application of such methods in the prosecution of hybridization experiments will ensure success where failure might otherwise result. The writer has watched with interest the phenological behavior of a wild hybrid between Myrica cerifera and M. carolinensis, that his former graduate student, Dr. Youngken, has described under the name of M. Macfarlanet. The first of these is a narrow-leaved ever- green shrub or low tree, that has its northern limit in New Jersey round the mouth of the Great Egg Harbor River, and there the rather MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 321 dull deep-green leaves are quite green even in mid-April of each suc- ceeding year alongside other and deciduous vegetation. These ever- green leaves defoliate in May or early June after the young leaves and flowers have well expanded. The species moreover invariably occurs only where facing sea breezes or along the edges of ocean inlets, as observed along a stretch of the New Jersey coast-line from the mouth of the Great Egg Harbor River to Cape May Point, and is invariably a swamp-loving plant. Myrica carolinensis—the common Waxberry or Bayberry—is a deciduous species of much wider range and greater hardihood, which drops its elliptic obovate shiny leaves by the end of November, at latest, and whose bare twigs throughout the winter show only the small protruding staminate catkins on one plant and the even smaller pistillate buds on another. The species occurs often many miles removed from the influence of ocean winds or brackish water and nearly always in dry sandy soil. The hybrid—M. Macfarlanei—is of semi-evergreen habit. © It grows frequently interspersed with both parents along the area of the New Jersey coast already named and doubtless will be recognized south- ward to Florida. Its lanceolate leaves of rather shining aspect remain green to the end of March, or only become in part brown and fall during April. More extensive and exact studies made as to the occasional retention of leaves on low young shoots of M. carolinensis into mid-winter and the retention of the evergreen leaves on M. cerifera to an even later date than the writer has indicated, may yet make our knowledge much more perfect regarding this striking hybrid and the possible synchrony of floral events, as well as leaf duration on individuals of the parent and of the hybrid. It might be added that the hybrid inhabits soil areas which are fairly intermediate between the swamps of the one species and the dry sandy soil of the other. If we consider now germination of seeds, equally suggestive syn- chronous procedure is observed. Only two, amongst many studied, need be mentioned as having recently been closely examined side by side with each other. The seeds of the little annual Floerkea pro- serpinacoides germinated this year in immense quantities over several moist shrubby valleys on March 17. The radicle had protruded and the cotyledons had become swollen on March 25, the first or trifoliate leaf was uniformly mature on April 7 and the second leaf was unfolding by Aprilt1. In contrast, subterranean seeds of Amphicarpaea monoica alongside the above, were still dormant on April 11, but on April 14 many had simultaneously begun to germinate. Moreover, but in line with all of the above, the annual trans- formations that occur in our woodlands in spring and throughout DL BROOKLYN BOTANIC GARDEN MEMOIRS summer, when compared from year to year, are as exact in relation to time and energy expenditure as are the flowering periods. Thus, if we compare the vegetative growth of the Yellow Adder’s Tongue during 1913 with that of the present year the tips of the leaves were simul- taneously emerging from the ground over wide areas of a valley on March 27, while this year they appeared on the 11th of April. In both cases these leaves were 23 inches high 3 days thereafter, and so comparatively suddenly transformed wide woodland areas from a bare unclothed aspect into rich showy brown-green verdure. If we consider now a few naked eye details that depend on definite histological changes, it may be said that botanists are aware that for any given species of shrub or tree a fairly definite period arrives when easing and separation of the epidermis along the stem is effected, after cork formation has replaced it functionally. Some species show this change in the latter part of the first year, many in the early part of the second, while others may be delayed until the following autumn or even later. A more pronounced though related occurrence is seen annually in the oriental plane. The extensive flakes of old dull-gray cork start to separate synchronously on the average about June 28 and so reveal the white younger cork underneath with increasing effect during the next few days, but variation as to date of this event may occur from year to year according to environal stimuli. Oppor- tunity has twice occurred for comparing this with the behavior of the same species round Kew Gardens, England, and there a like change starts on July 29. This comparative result agrees closely with other data obtained as to floral maturation. Closely related again to the above studies is one that has scarcely been touched in this country, but which has been investigated by Hoffmann-Ihne in their observations at Giessen. This is an exact comparison of flowering periods according to longitude and latitude, particularly the former. Exceptional facilities exist for the prose- cution of such an inquiry in this country, for were thirty or forty observational stations established under competent workers, and the whole correlated at a central office, valuable results would accrue after a period of ten to twelve years. A feature of interest here is that a considerable number of plants of the eastern seaboard extend their range from central or northern Florida to Newfoundland or even Labrador. One of these which the writer has shortly referred to elsewhere’ is our native pitcher plant (Sarracenia purpurea). In northern Florida, as for example round Ponce de Leon, it starts to bloom in the last week of March and continues until April 10. In the Charleston region, as at Summerville, it is averagely five days later; 4 Engler’s Pflanzenreich, vol. 4 (1908), p. 23. MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 323 round Wilmington, N. C., it begins on April 18; in central New Jersey, it opens on the 20th of May; in northern New York and Minnesota it blooms from the 14th to the 26th of June. In eastern central Maine the period is from the 8th to the 20th of July, while in Labrador—the northern limit of the species—it finishes in mid-August. Thus a period of fully five months is represented, and a longitudinal area of about 2,000 miles is covered, in the floral maturation of this one species. In connection with such records, and probably due to the more gradual and even expenditure of environal heat stimuli is the much more extended floral period of species in the cool north than in the warm south. Thus while the double Crimson Rambler and Dorothy Perkins roses show floral attractiveness from June 10 to June 25 averagely round Philadelphia, on eastern Mt. Desert the period extends from July 15 to August 30. Were the valuable records, inaugurated in 1892 for Canada by Mackay, to be linked up with like records from widely distributed stations in this country, and were all to be correlated with tempera- ture or thermotactic and moisture or hydrotactic stimuli, as has in part been done by the Canadian observers, a most valuable foundation for the establishment of facts regarding the action of definite environal stimuli would be made. A very wide field for exact study, still left practically untouched, is the observation and recording of sporangial ripening and spore dissemination in pteridophytic and bryophytic genera and species. One or two references need only be made here. For years the writer was puzzled to know when spore-dissemination took place in the sensitive fern (Onoclea sensibilis). Though the green sporophylls shot up in late July and became greenish-brown in autumn, opening of the modified pinnae and dissemination of spores clearly did not take place before winter. Passing through a swampy patch of this on March 24 of five years ago his clothes became browned over with the shed product. Subsequent study has shown that this event occurs averagely on March 25, and in any one patch or locality with surprising synchronous exactness. Like observations should be made for Onoclea Struthiopteris. The sudden and simultaneous elongation of the sporophores and the subsequent rupture of the sporangia in such hepatics as Pellia endiviaefolia is familiar to all in mid-April, but we still lack exact day and hour records through succeeding years for the entire group of scale mosses. The predicable manner in which, when fresh horse manure is placed under bell jars in the now familiar laboratory experiment with Pilobolus, an abundant crop of the black sporangia is shot forth on a 324 BROOKLYN BOTANIC GARDEN MEMOIRS. morning after a definite number of days of growth, is as indicative for that fungus as is the previous growth of Mucor on the same medium, within a shorter period. What conclusions, it may now be asked, can be drawn from data such as the few above given? Time, space, energy and matter are the four great interrelated phenomena of the world, as of the universe generally. Not a few physicists now question the existence of the last of these, but inert and mobile ether particles as focal centers and pathways for ‘tubes of energy’’ seem to be helpful—even necessary requirements. For by ° their gradual aggregation under increasing condensations of energy we can explain the origin of the elements, and equally the compounds of these. But the fundamentally important consideration is how, when, and to what extent in given times, do definite tubes of energy distrib- ute themselves. In the foregoing pages a set of simple facts has been recorded that any average observer might accumulate. But the real value of many of them has been overlooked, because we have not fully realized the significance of the causes that bring them about. For in the past we have largely viewed biological phenomena as static or semistatic exhi- bitions of so much material substance. But we have in great measure failed to realize that matter as such is physically passive or inert, and that the fundamental moving, transforming, upbuilding, and dis- integrating agency in all of the above phenomena of phytophenology consists in definite expenditures of definite amounts of energy along definite: material pathways. Or to use Faraday’s phrase as applied to inorganic changes, we are dealing with “tubes of energy”’ that are distributed along definite material pathways, at stated climatic periods, and that are marvelously exact for any one species, or any one organ of a species. In the process these tubes of energy are exactly expended so as to stimulate the inert material particles to take up water, to digest or metabolize reserve products, to convey the metabolized products to definite cells or cell walls, to build these up into new material linkages or combinations, and in the process to effect growth of leaves, opening of flowers, dehiscence of anthers or of sporangia, maturation of ovules and extrusion therefrom or from some accessory part at exactly appropriate time of viscous entangling secretions that strand the pollen grains, and that in time aid in the germination of these; or again that start like initial changes in dormant seeds, once so many units of heat, moisture, and oxygen have cumulated as summated tubes of energy after a definite period of time; or that develop new cells or transform older ones, so as to effect shedding of epidermis or MACFARLANE: SYNCHRONISM IN PLANT STRUCTURES 325 peeling of bark on twigs of trees and shrubs at exact time; or that effect passage of food material from elaborating to storing centers, and from the latter in turn to young developing organs. Even such irregular and delayed occurrences as already noted re- garding the flowering during the present season of the Carolina poplar, or during 1913 of the silver maple (A. saccharinum) represent fitful and prolonged results due to weakened or cancelled tubes of energy- stimulation expended over an extended period, as compared with the normal succession of events that may be consummated within a few hours on a definite day or days. So we might summarize as follows our conclusions drawn from study of phenological and related events: 1. For any one locality, under like environal surroundings, the average annual period of seed-germination, leaf-formation or unfolding, first period of blooming, dissemination of pollen, and other responses by flowering plants, seem to be synchronous often toa day, and even to certain hours of one day. 2. In monoecious and dioecious flowering plants, under like en- vironment, all evidence tends to indicate that maturation of compli- mental floral organs is effected in exactly synchronous relation, and so abundant pollination usually ensues. 3. A like principle apparently applies to the maturation and dispersal of spores and organs of conjugation. 4. The behavior of plant hybrids strongly suggests that each is a blended combination of parental characters as to period of leafing and defoliation, of blooming and pollination, capacity for climatic resist- ance and other phenomena. So each shows synchronous behavior in its organs, that is a mean—all environal factors being considered— between those of the parents. 5. The principle, advocated by the writer for the past six years, of environal action and organismal reaction, seems to hold true in the organic as in the inorganic world, and only needs to be amplified and demonstrated by increasingly accurate and extended observations on plants over wide areas. 6. In the evolution of all plants and of all plant parts, the funda- mental and important consideration is the exact distribution of lines or ‘“‘tubes of energy” (Faraday) along otherwise inert material path- ways; the lines of inflowing energy constituting stimulation actions, the lines of outflowing energy constituting reactions on the part of the organism. 7. Such actions and reactions show an optimum, as well as a maxi- mum and minimum of interrelation. The optimum for the indi- viduals of each species, and for the organs of this, under like environ- ment, often constitutes a phase-relation that recalls like phase-rela- tions amongst inorganic bodies. ws) iT) ON BROOKLYN BOTANIC GARDEN MEMOIRS 8. The material constituents of each plant organism—in them- selves inert—constitute the gauge or measure by which tubes of in- flowing or stimulant energy, and outflowing or reaction energy can best be estimated. The structure of all plant organisms, therefore, is a cumulated expression of the continued flow of definite tubes of energy, and the resultant placing, in definite and orderly manner, of otherwise inert material particles, when moved by such streams of energy. THE PROBLEM OF THE IMPORTED PLANT DISEASE AS ILLUSTRATED BY THE WHITE PINE BLISTER RUST HAVEN METCALF Bureau of Plant Industry, U. S. Department of Agriculture Within the last few weeks the civilized world has stood aghast at the stories told by survivors of the devastation wrought by the German army in its retreat from Northern France. Particularly schrecklich are the stories and pictures showing the rows of fruit trees cut down without being utilized for fuel, apparently with no purpose other than wanton destructiveness. But if we pause to think we realize that the Germans did not do the worst that they could have done. The fruit trees are cut down, to be sure, but others can be planted in their places. We may expect in future years to see the old orchards completely rehabilitated. But let us suppose that instead of simply cutting down trees in Europe, any enemy should see fit to leave them standing but thoroughly incoulated with diseases which would not only destroy the present stand of trees but would prevent their future profitable culture in the same territory. To take an extreme example, suppose that some malignant person or nation should see fit to introduce into Europe from America, the Colorado beetle, the pear blight, the chest- nut bark disease, and the citrus canker. Surely the world would stand aghast at this if its significance was generally realized, because this would not only destroy or seriously damage the present generation of plants concerned but would tax the agricultural resources of Europe with a perpetual burden. No more fiendish blow to the economic resources of a country could be imagined. Yet this is exactly what we have allowed foreign nations to do to us without resistance, through our lax laws regarding the importation of live plants, or rather through our virtual absence of laws on this subject. We have permitted our country to be invaded by not one but many—perhaps hundreds—of diseases and pests which constitute a permanent tax upon our agri- cultural and forest resources, and up to the present time we have made scarcely any serious effort to prevent further invasion. The Bureau of Entomology has issued a large bulletin entitled “A Manual of Dangerous Insects Likely to be Introduced in the United States Through Importations.’”’ I understand that this 327 328 BROOKLYN BOTANIC GARDEN MEMOIRS bulletin lists about 2,700 such insects and that it also lists 130 important insects which have already been introduced into the United States. Ifa thorough canvass was made of foreign literature I have no doubt that as many plant diseases could be located and described which are likely to be introduced into the United States, and many of them produce as much devastation as the chestnut bark disease, the asparagus rust, the potato blight, the citrus canker, or many other diseases that could be named. Unfortunately, we have at present no corresponding manual of plant diseases that are likely to be introduced into the United States. : There has never been a time when the danger from imported diseases and pests was so great as now. Commerce in living plants has in recent years extended to the ends of the earth. I have in mind one nursery company which makes a specialty of novelties from the Orient. This company is distributing throughout the United States plant material from all parts of Asia. Most of the things that they bring in are woody plants, many related to our American species, and on account of our comparative ignorance of the botany and zoology of the Orient we have no idea what diseases and pests are coming in with Oriental material. The San Jose scale, the chestnut blight, and the citrus canker are only a part of those that have come in already. Not only is commerce being carried on with countries from which hitherto there have been only scattering importations of live plant material, but material brought in now is much more miscellaneous and reaches this country in a much shorter time. There is at present a limited amount of port inspection but too limited to be efficient and the canker diseases and many insects can not be detected by any sort of inspection. The roots of plants imported with earth about them can not even be inspected and such plants constitute a par- ticularly dangerous class of imported material. The white pine blister rust (caused by Cronartium ribicola Fischer), which I am here considering as a fairly typical example of the imported disease, has long been known in Europe. It apparently originated in Asia and spread in Europe upon Pinus cembra. When the American white pine (Pinus strobus) was introduced into Europe it proved sub- ject to the disease. The first authenticated record of importation of white pine transplants from Europe to the United States dates back only to 1899. From that time until prohibited by law such importa- tion was extensive, as such transplants could be imported more cheaply than they could be grown in America. There was, furthermore, a prevalent belief among nurserymen that white pine seedlings could not be successfully grown in America, a belief which has since been proved erroneous. METCALF: WHITE PINE BLISTER RUST 329 The disease was first positively reported in America in 1906, on Ribes. No notice appears to have been taken of this warning. In 1909 enormous quantities of diseased pine nursery stock were im- ported. Probably 95 percent of all diseased seedlings imported into America came from a single nursery, that of J. Hein’s Séhne at Halsten- bek, Germany. This nursery, on account of its use of Ribes hedges, was curiously well adapted to distribute the disease. In June, 1909, a meeting was held in New York City of pathologists and foresters of New England and the Middle Atlantic States, at which a further alarm was definitely sounded. With one exception, all states repre- sented discouraged importation of white pine from that time, but commercial nurseries continued to import extensively until such importation was made illegal in 1912. Unfortunately, no studies of the white pine blister rust have been made in Europe by any American investigator, but if European accounts of the behavior of the disease can be trusted, the disease has apparently spread more rapidly and with greater virulence in New England than it did in Europe. Prob- ably the new climatic and host relations are more favorable to the disease. In any case the problem of invasion presented by this disease makes an interesting study. The black currants, especially the cultivated varieties, are particularly subject to the disease and in areas of scattering infection are reliable indicators of its presence. It was hoped by many that the disease might prove to be only one of nursery stock and reproduction, but at several points in New England, New York and Minnesota, it is attacking large trees. On Ribes the disease was in 1916 generally prevalent throughout New England, which means that the actual infection of pine is much more general than is obvious at present. Inspection of nursery stock for blister rust is largely futile since the rust often incubates in pine tissue for many years before becoming apparent by distorting growth or fruiting. According to Ravn this incubation period may be as long as twenty years. The control of the disease in America presents three separate problems: First. West of the Mississippi River. In this territory the disease is not known to occur, but undoubtedly has been shipped in on nursery stock of either pine or Ribes. If it has not been carried into this territory on nursery stock already, there is little possibility of its ever getting in by natural means. During the coming season an extensive survey will be made of these states to determine whether the disease is or is not present. If the disease should once become established under western forest conditions, its control would be hopeless. All 5-needle pines of this area, including the very valuable sugar pine and 330 BROOKLYN BOTANIC GARDEN MEMOIRS western white pine, are subject to the disease, and wild Ribes of many species are abundant. No species or variety of Ribes yet tested is immune to the disease. Second. From the Mississippi River to the Hudson River. There is an area about 30 miles square in Minnesota and Wisconsin north- west of St. Paul which is now known to be heavily infected. Probably more infection will be found in Minnesota. In Michigan, Indiana, Ohio, Pennsylvania, and New Jersey the disease has been found in a few nurseries and plantations and is believed to have been eradicated at these points. In New York west of the Hudson River it has been found in both nurseries and plantations and largely eradicated, but on account of the extensive planting of pine nursery stock in the Adirondacks heavy infections are to be anticipated there. — In general, the commercial currant-growing sections, such as the region from Rochester to Buffalo, may be expected to soon show general infection. In Canada the Niagara Peninsula is already generally infected, and at least scattering infections occur elsewhere in Ontario. This infec- tion is of course a serious menace to Michigan. Third. East of the Hudson River. Here infection is so general that the only hope of successful growing of white pine in the future lies in the elimination of the alternate host of the disease; namely, Ribes. Whether such elimination can be made at a sufficiently low cost to be profitable remains to be seen. Probably in localities where Ribes occur sparsely, as in Connecticut and Rhode Island, a great deal can be accomplished. However, in many sections of rough country, where wild Ribes are too prevalent to be profitably eradicated, white pine growing may be expected to become impracticable. Throughout any section where the blister rust becomes prevalent, the effect is to make the white pine a cultivated plant; that is, it can not be profitably grown, or perhaps not grown at all, unless the ground is kept free from Ribes by artificial means. There are certain difficulties which stand in the way of any general campaign of disease control which involves wholesale eradication of diseased and susceptible plants. It remains to be seen whether these difficulties are or are not insurmountable. 1. There is in the United States no central authority to act in any matter involving destruction of diseased plant material or pre- cautionary destruction of that which is not diseased. Whatever destruction is undertaken must be undertaken in each state under the separate legal authority of that state. The state laws are not uni- form. In some states they are adequate and well supported by public sentiment, in others they are wholly inadequate and apply only to special cases. In general there are few states in which the laws are METCALF: WHITE PINE BLISTER RUST 3611 not so flexible but that a single unconvinced or cantankerous indi- vidual can nullify the work of an entire community. 2. The laws governing plant eradication are administered in different states by various officers, but in the majority of states by the state nursery inspector, who is nearly always an entomologist. Some of these entomologists are thoroughly trained in plant diseases and fully appreciate their significance. Others have little knowledge of them and less interest. All of these officers are overworked already. In spite of the activities of plant pathologists, there is profound popular ignorance as to the nature and significance of plant diseases and especially of the dangerous qualities of newly imported diseases. The general public is far better informed regarding “bugs”’ than regarding fungi and as a matter of fact the average man considers that plant diseases are caused by “‘bugs.”’ 3. In the case of the blister rust, there is no single interest or centralized affiliation of interests whose securities are menaced by the disease. The white pine industry is diffused over a wide territory and in the hands of many separate individuals and organizations. What is everybody’s business is likely to be nobody’s business. If the white pine industry, like the redwood industry, for example, or like the citrus industry, were in the hands of a few people or consti- tuted the dominant business in certain areas, the control of the disease would be much simpler. 4. Up to the present time there has been no adequate quarantine against the disease, either state or national. This difficulty, however, will shortly cease to exist as many states have recently declared quarantines and on June 1, possibly earlier, an adequate national quarantine will go into effect. 5. Finally, we have a very serious consideration which is applicable to all undertakings at the present time. The nation is at war. The young men who would ordinarily be employed in an eradication campaign will soon be drawn away into military work or into the various lines of industry which bear directly upon the conduct of the war. The majority of the persons employed in this particular eradica- tion campaign can only be employed from the middle of April to the first of November. Necessarily, men will be loath to accept such temporary employment when they can in other lines secure permanent employment at an equally. high or higher wage. In any case, the work if successfully prosecuted or indeed if prosecuted at all will involve much larger expenditures for wages than in normal times. What then is the outlook for the control of the white pine blister rust? It may be expected that the future course of the disease will be much like that of the gypsy and browntail moths; that is, the dis- 332 BROOKLYN BOTANIC GARDEN MEMOIRS ease can be controlled for a long period of years in those localities where infection is not general but in areas of general infection the control will be only local and the efficiency of this local control will largely depend upon whether the white pine as a crop is of sufficient value to sustain the added expense of the eradication of Ribes. It is to be hoped that any infections found west of the Mississippi River will be scattering and small, for, as has already been stated, if the disease once becomes established under western forest conditions its control even on a local basis will be absolutely out of the question. Two facts are always in favor of the control of this disease: (1) The disease has two hosts and can not pass from pine to pine. (2) The disease is a comparatively slow one; that is, slow as compared with such a disease as the chestnut blight. If at any future time in a completely infected locality the increasing value of the white pine or a change in industrial conditions makes local control profitable such control can be undertaken regardless of the prevalence of the disease at the time, since wherever Ribes can be thoroughly eradicated healthy pine stock can be grown and will not take the disease from the already diseased pines. The entire blister-rust problem is, however, but one phase of a larger problem, which may be stated as follows: does free trade in plant diseases and insect pests pay? Is it an economically sound national policy? Is the entire 1mporting nursery business worth as much to the country as the damage which it has already caused? Not a single plant disease or insect pest that has once become estab- lished in this country has been eradicated or, in the present state of knowledge, is ever likely to be. No matter how well controlled, it remains in every case a permanent tax against our economic resources. Even if we succeed in controlling the white pine blister rust we may be absolutely certain that other diseases and pests are being intro- duced which will be just as serious, for we know definitely that the undesirable plant immigrants are not yet all here. It is much more important to safeguard the country against further invasions of this kind than to control this or any other disease or pest that has already been carelessly permitted to establish itself. It isa matter of common knowledge, which I scarcely need to repeat here, that the countries of Europe, and even ends of the earth like Tasmania and South Africa, have long since protected themselves against the importation of diseases and pests either by prohibition of entry of nursery stock, or by exclusion of large classes of such stock. The United States is far behind in this matter. The future danger is far greater than the present. The most dangerous class of nursery stock is the ornamental trees and shrubs, VOLUME |, PLATE VI. BROOKLYN BOTANIC GARDEN MEMOIRS. gow AT igh iar; “eRe 4a. a. eae: WR ye TSS Yer METCALF: WHITE PINE BLISTER RUST BROOKLYN BOTANIC GARDEN MEmoirs. VOLUME I, PLATE VII. METCALF: WHITE PINE BLISTER RUST METCALF: WHITE PINE BLISTER RUST 533 on account of the great number of species represented, and the widely various parts of the earth from which such stock comes. No section of the country is likely to suffer more from this source than Long Island and the general vicinity of New York City. May I express the hope that this Botanic Garden, already a leader in local and national sentiment in horticultural and botanical affairs, will take a position in this matter which will help to bring about the suppression of this very dangerous traffic? EXPLANATION OF PLATES VI AND VII PLATE VI. A native white pine (Pinus strobus) in thick stand, completely girdled by a blister rust canker. Kittery Point, Me. (Photograph by Mr. W. S. Carpenter, of the New York State Conservation Commission.) PLATE VII. A young native white pine (Pinus strobus), completely girdled and showing several infections of blister rust on trunk and branches. Kittery Point, Me. (Photograph by Mr. W. S. Carpenter, of the New York State Conservation Com- mission. ) THE ROSY-SPORED AGARICS OF NORTH AMERICA WILLIAM A. MURRILL New York Botanical Garden This subtribe of the gill-fungi is characterized by rosy or rosy- ochraceous spores and is well represented by the common and widely distributed species, Pluteus cervinus. ‘The North American species may be grouped in ten genera, distinguished by the following key: Subtribe PLUTEANAE Pileus irregular, dimidiate or resupinate. I. CLAUDOPUS. Pileus regular, sometimes eccentric in Pleuropus. Volva and annulus wanting. Stipe cartilaginous. Margin of pileus incurved when young. Lamellae decurrent. 2, ECCILIAY Lamellae adnate or adnexed. 3. LEPTONIELLA. Margin of pileus straight and appressed when young; lamellae free or adnexed. 4. NOLANEA. Stipe fleshy. Lamellae decurrent, rarely varying to adnate. 5. PLEUROPUS. Lamellae sinuate or adnexed. Spores not angular, rosy-ochraceous in mass. 6. LEPISTA. Spores angular, rose-colored in mass. 7, ENTOLOMA. Lamellae free. 8. PLUTEUS. Volva wanting, annulus present. g. CHAMAEOTA. Volva present, annulus wanting. 10. VOLVARIOPSIS. A few of these generic names may be unfamiliar to you, such as Leptoniella for Leptonia, Pleuropus for Clitopilus, and Volvariopsis for Volvaria, but these changes have been required by the rules of nomenclature. The time at my disposal will not permit more than a hasty sum- mary of the North American species belonging to these genera. A fuller treatment may be found in North American Flora, volume 10, part 2, to be issued shortly. I. CLAUDOPUS Claudopus nidulans, the best known species, occurs throughout Canada and the United States; C. avellaneus is known from Oregon only; and there are no species reported from tropical North America. 334 MURRILL: ROSY-SPORED AGARICS 335 The other species of the genus occur in the eastern United States, from the Atlantic seaboard to the Rocky Mountains. 2. ECCILIA The best known species is E. atrides, of the eastern United States. Three species are confined to the Pacific coast, and four to tropical North America. 3. LEPTONIELLA Leptoniella serrulata, characterized by the black, serrulate edges of the lamellae, is the best known species. Four are confined to the Pacific coast and seven to tropical North America. The rest occur in the eastern United States. 4. NOLANEA The two best known species are N. conica and N. mammosa, the latter distributed throughout temperate North America but not generally recognized by mycologists. One species is confined to the Pacific coast and three to tropical North America. 5. PLEUROPUS This genus contains many edible species, among them P. prunulus, P. orcellus, and P. abortivus, the last readily distinguished by the peculiar aborted hymenophores. Two species are confined to the Pacific coast and one to tropical North America. 6. LEPISTA One of the best edible species we have is L. personata, better known as Tricholoma personatum. ‘This species and L. tarda occur through- out temperate North America. Two other species are confined to the eastern United States, and there are none known from tropical North America. 7. ENTOLOMA This also is a temperate genus, the only species (E. Murrait) reaching tropical North America being found in the high mountains of Jamaica. Four species are confined to the Pacific coast. The best known species in the eastern United States are: E. strictius, E. Grayanum, E. sericeum, E. rhodopolium, E. Murra, and E. salmoneum. Because of the very poisonous European species, E. lividum, the mem- bers of this genus have been largely avoided by mycophagists. 23 336 BROOKLYN BOTANIC GARDEN MEMOIRS 8. PLUTEUS The best known species are P. leoninus and P. cervinus, which are widely distributed. Five species are confined to the Pacific coast and fifteen to tropical North America. g. CHAMAEOTA Only one species, C. mammillata, has been known in this country and this only from Michigan. 10. VOLVARIOPSIS The species of this genus are apt to be widely distributed, as is the case with many fungi which inhabit manure. The best known species are probably V. bombycina, V. speciosa, V. volvacea, and V. pusilla. Four species are confined to tropical North America. GENERAL SUMMARY Genera Oe See ner cons New Species Total Claudopust saree oe 4 4 I 9 FE CGUALG Sa OTE ae I 15 9 25 WGC PLONI EL Oacenece ese she I 28 14 43 INOLG ICG Pres cae teeters ees I 16 II 28 IPICULG DUS an eee 2 23 5 30 JED Ie SMG Eanes Faceted fc 3 I oO 4 FEN OLOMNO ee een eee 2 277, 34 63 JANDUATIS Ms ol Paid SRSA Oe 2 25 30 57 ChaMacciar meee eee to) I I 2 VOlUOnOPStS aoa ie ay 6 13 2 21 DOr. pe cee ets aeis s Suskere 22 153 107 | 282 THE CYTOLOGICAL STRUCTURE OF BOTRYORHIZA HIPPOCRATEAE EDGAR W. OLIVE Brooklyn Botanic Garden This species of rust, occurring on the host Hippocratea volubilis L., was first described by the author, in collaboration with Professor H. H. Whetzel, from material collected in Porto Rico in 1916.! It is there recorded as a somewhat peculiar form, though somewhat like a lepto-Uromyces, with only one spore form in its life-cycle. The fol- lowing diagnosis is there published:? ““O. Pycnia wanting (probably not formed). III. Telia mostly hypophyllous but sometimes amphi- genous or caulicolous, generally from a localized mycelium, sometimes from a systemic invasion affecting entire young shoots; localized sori densely crowded in more or less orbicular or irregularly shaped, somewhat hypertrophied pulvinate areas, I mm.—I cm. or more across, the affected areas yellowish when young, when older becoming whitish due to the germination of the spores; in older leaves often killing affected spots, which turn brown, the resultant rounded, swollen dead areas then bearing a striking resemblance to certain insect galls. “Telia pulverulent, erumpent, from a definite, superficial, ure- dinoid hymenium which arises just under the epidermis, without . peridium; teliospores uninucleate, borne singly at the end of pedicels which arise from a binucleate mycelium 13-14 by 18-24 yu, thin-walled, oval, with a rounded apical protuberance, germinating apically at maturity to produce each a long, cross-septate basidium (promy- celium) bearing 4 basidiospores (sporidia), similar in shape to the teliospores and 8 by II—I2 up. “Vegetative mycelium composed of coarse intercellular hyphae, made up of binucleated cells, some of which send large botryose, or irregularly shaped, haustoria into adjacent cells.” The generic name, Botryorhiza, is, in fact, derived from the botryose character of the haustoria, a striking feature which, so far as I am aware, is possessed rarely if at all by other rusts. It is, however, 1 Endophyllum-like rusts of Porto Rico. Amer. Journ. Botany. 4: 44-52. pls. Z—2: » IOL7. ZN LGs De Aye 337 338 BROOKLYN BOTANIC GARDEN MEMOIRS pointed out in the above mentioned paper, that certain smuts (as Doassantia deformans, e. g.) also possess botryose haustoria.® The 18 figures in Plate VIII show fairly clearly the salient features of the cytological structure and development of Botryorhiza, with the exception of the sexual fusions, which have not as yet been found. Undoubtedly, however, these fusions and the consequent transition from the uninucleate condition initiated in the germinating pro- mycelium to the binucleate condition prevalent in the vegetative mycelium, must take place early in the development of the latter. Two hypophyllous sori are shown in Fig. 1, the one at the right a very young one pushing through a stoma. The mycelium and hymenial hyphae are seen to be composed of binucleate cells, as is also the case of the young spores. Two of the peculiar botryose haus- toria are shown at the lower portion of the figure, nearly filling the host cells. Fig. 2 shows a portion of the coarse, branched, inter- cellular mycelium, with some its binucleate cells. This mycelium varies from about 5 to 7 win diameter. One cell is shown with four nuclei, evidently a result of a recent conjugate division. The con- spicuous thickenings drawn along the edges of the hypha are colored red in the preparation with Flemming’s triple stain; their mode of origin and significance still remain to be solved. Figs. 3-7 show five varying views of the large botryose haustoria. The narrow isthmus connecting the enlarged haustorium with the extra-cellular mycelium is clearly shown in each case, as is also the interesting fact that the haustorium, even in those cases in which the host cell is almost entirely filled by it, pushes in as it grows the plasma membrane of the host protoplasm. Strictly speaking, therefore, in no case is the haustorium really inside the host protoplasm. Fig. 3 is a section of a young haustorium showing this invagination of the host cytoplasm. Four nuclei of the rust are also shown in this prepara- tion. In Fig. 4 an older haustorium showing its peculiar botryose swellings, has pushed up into an unusually dense mass of host cyto- plasm, now shrunken away from the invaginated haustorium. Figs. 5, 6, and 7 show almost equally clearly this phenomenon of invagina- tion of the host cytoplasm. In Fig. 5, particularly, the haustorium is seen to almost fill the host cell. An idea of the large size of the haus- toria may be gained from the fact that while the cells of the leaf of the host Hippocratea measure from 15 to 20 u in diameter, those of the fungous haustoria range from about Io to 14 w in diameter. Figs. 8 to 14 show various stages in the formation and germination of the teliospores. Fig. 8 is of a young sporiferous hypha showing two ’Lutman, B. F. Some contributions to the life history and cytology of the smuts. Trans. Wis. Acad. Sci. 16: 1191-1244. 1910. See his Figs. 44 and 45. OLIVE: STRUCTURE OF BOTRYORHIZA HIPPOCRATEAE 339 pairs of conjugate nuclei, those of the stalk and also those of the young spore, which is shown in process of abstriction. As in other cases of cell-division among fungi and algae, a ring-formed constriction grows in from the periphery, thus cutting off in this instance the binucleate spore from the binucleate stalk cell. Fig. 9 is a drawing of such a young spore, showing the two nuclei; in Fig. 10, the two nuclei have fused, though it is evident from the presence of the two nucleoles that this fusion has only recently taken place. Figs. 11 and 12 represent mature spores, each borne on binucleate stalks, the latter figure showing the apical protuberance so characteristic of most spores. That this protuberance is the primordium of the apical germ-tube is apparent from a perusal of Figs. 13 and 14. These spores, as is stated above, germinate at once on maturity. In Fig. 14, the hetero- typic nuclear division is proceeding; this, however, is so poorly stained in the preparation, that it is not possible to make out the details of the process. Figs. 15, 16, and 17 show the basidia, or promycelia, each composed of four uninucleate cells, which result from the germination of the teliospores. In Fig. 15 appears a type of germination which apparently results from growth of the basidium in a very damp situa- tion; the four cells of the basidia in such cases often break apart and function independently. Fig. 16 shows the more usual type of germination, in which each of the four cells sends out a branch, to bear finally at the tip of each a single uninucleate basidiospore on a sterigma. Two of these oval basidiospores are shown in Fig. 18. GENERAL DISCUSSION It will be seen from these figures and the accompanying description that while Botryorhiza undoubtedly resembles a short-cycled lepto- Uromyces in the one-celled character of its teliospores, it is sufficiently distinct in other respects to justify its being placed in a new genus. Some of these differences are as follows: the walls of the teliospores in Botryorhiza are thin instead of thickened, as is usual in Uromyces; they are colorless, instead of brown or otherwise colored, as in Uro- myces; there are no germ-pores in the walls of the spores of Botryorhiza, whereas the teliospores of Uromyces are characterized by one or more. Finally, the possession of such strikingly large, botryose haustoria, so characteristic of Botryorhiza, is, in my opinion, a very distinctive feature. . Apparently the more usual type of rust haustorium, so far as our few studies on the subject have revealed, is that of an irregular, branching hypha. Atkinson‘ has, however, figured the haustoria of Uromyces caryphyllinus as somewhat irregular and botryose in form; 4 College Botany. Henry Holt & Co. P. 87. 340 BROOKLYN BOTANIC GARDEN MEMOIRS and Pole Evans® in his careful investigation of the histology of nine species of cereal rusts has shown that while the young haustoria of various species may be small and sac-like, or even “ hammer-headed”’ in others, the prevailing type of mature haustoria in these cereal rusts seems to be the cylindrical or branched form. Attention was called in the earlier description of Botryorhiza to the fact that Lutman had figured botryose haustoria in Doassantia and it was there suggested that this Porto Rican rust might have some other features in common with smuts. But the fact that the sporifer- ous hyphae are sent out through a stoma or through the ruptured epidermis before the spores themselves are cut off from their tips, and, further, that there is produced in Botryorhiza (and apparently in rusts in general) a definite, superficial hymential layer from which the spores arise constitutes two essential points of difference from the smuts. The latter, as Lutman has clearly emphasized,° have their spores pro- duced either from a group of deeply imbedded multinucleate hyphae, which break up directly into spores (in the Ustilaginaceae) or from the tips of the side or main branches of the prevailingly binucleate hyphae (as in the Tilletiaceae). It has also been brought out in the description of the characteristics of Botryorhiza that the mycelium is composed of a branching system of very coarse hyphal threads. These hyphae measure from 5 to 7 in diameter. In my own work on various rusts, I had never before met with such a coarse mycelium. Pole Evans,’ however, has called attention to the fact that the mycelial threads of Puccinia glumarum reach the relatively enormous size of 10 to 19 u in diameter; also Dodge in his paper in this Memoir® has noted that Farlow® and Wornle!® have found the hyphae in Gymnosporangium Ellisit to be exceptionally large, being, according to the latter author, about 8 pu in diameter. 5 The cereal rusts. I. The development of their uredo mycelia. Annals of Botany 21: 441-446. 1907. OES Cho (Os WAI Og Coy Ds AUS 8 See p. 128. °The gymnosporangia of the United States. Ann. Mem. Boston Soc. Nat. Hist. 1880: 1-38. 10 Anatomische Untersuchung der durch Gymnosporangium-Arten hervor- gerufenen Missbildungen. Forst. Nat. Zeits. 3: 68-84; 129-172. 1894. BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME |, PLATE VIII. art Sie Lat) AEN O OLIVE: CYTOLOGICAL STRUCTURE OF BOTRYORHIZA OLIVE: STRUCTURE OF BOTRYORHIZA HIPPOCRATEAE 341 EXPLANATION OF PLATE VII All drawings have been made with the camera lucida, and with various combina- tions of Zeiss apochromatic lenses. Except where otherwise noted, the magnification has been 1,000 diameters. Botryorhiza Hippocrateae Whetzel & Olive Fic. 1. Two young sori, showing the hymenial layer, composed of binucleate cells, and method of spore formation. At the right a young sorus pushing through a stoma. Somewhat diagrammatic. XX 500. Fic. 2. Branching hypha, showing the binucleate cells, whose walls in some places show peculiar thickenings. XX 500. Fic. 3. Young haustorium, containing 4 nuclei. Note the invagination of the host cytoplasm. Fic. 4. Another haustorium, pushing into a mass of dense, granular host protoplasm. Fic. 5. A fully mature haustorium, showing its botryose lobings. Fics. 6 and 7. Haustoria in partial section; showing the pushing in of host protoplasm. Fic. 8. A young hypha from the hymenium, showing the constricting wall cutting off stalk from spore. % 1,500. Fic. 9. A young spore, with two nuclei. Fic. 10. A young spore in which the two nuclei have just fused, as evidenced by the presence of two nucleoles. Fics. 11 and 12. Mature teliospores, showing binucleate stalks. Fic. 13. A spore showing young germ-tube. Fic. 14. Another spore, with growing basidium, or promycelium, The nucleus is in metaphase of the heterotypic division; but the preparation is poorly stained. Fics. 15,16 and 17. Three basidia, which have divided into the characteristic 4 cells. Fig. 16 shows the branches, each of which will bear ultimately a single basidiospore. Fic. 18. Two basidiospores, showing the uninucleate condition. THE NUCLEUS AS A CENTER OF OXIDATION W. J. V. OSTERHOUT Harvard University In 1897 Spitzer! reported that nucleoproteins extracted from certain animal tissues possess the same oxidizing power as the tissues themselves. The idea that the nucleus is a center of oxidation was advocated by Loeb? who pointed out that it would explain why cells deprived of their nuclei are unable to live for a long time or to regen- erate missing parts. R. Lillie* sought to obtain direct experimental evidence by applying to the cell reagents which become colored on oxidation. The reagent chiefly employed was a mixture of alpha naphthol and paraphenylene diamine which yields upon oxidation a deep purple dye, indophenol. The oxidation takes place slowly on exposure to air, but is greatly accelerated in the presences of living cells or extracts of living tissues. Lillie found that in certain tissues of the frog (especially liver, kidney and leucocytes) the colored oxidation products were deposited in and about the nucleus, especially at the surface of contact between nucleus and cytoplasm. Wherry! applied methyl green to Amoeba and reported that it was oxidized in the cytoplasm but not in the nucleus. Schultze’ applied a number of stains to plant and animal cells and found that they were oxidized in the cytoplasm but not in the nucleus. Unna‘® has investigated a large number of cases by the use of leuco- methylene blue and has reached the conclusion that the nucleus is a center of oxidation. Unna’s theories have been criticized by Oppen- heimer’ and by Schneider.’ Mathews? has come to the conclusion that the nucleus is directly concerned in oxidation. 1 Pfliiger’s Archiv. 67: 615. 1897. 2 Archiv. f. Entwickelungsmechanik der Organismen 8: 689. 1899. 8’ American Jour. of Physiology 7: 412. 1902. 4Wherry, E. T. Science, N. S. 37: 908. 1913. 5 Schultze, W. H. Verhandl. d. deutsch. path. Ges. 16: 161. I913. 6Unna, P.G. Archiv. f. mikr. Anat. 78. 1911. Godoletz, P. und Unna. P. jun. Berlin, klin. Wochenschrift 49: 1134. 1912. Unna, P. G. und Godoletz, L. Oppenheimer’s Handb. d. Biochem. Erganzungsband. 1913. S. 327. 7 Oppenheimer, C. Die Fermente und ihre Wirkung 2: 790, 810. 1913. 8 Schneider, H. Zeit. wiss. Mikr. 31: 478. 1914. ° Mathews, A. P. Physiological Chemistry p. 180. 1915. 342 OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 343 Warburg’? found that it was possible to isolate the nuclei from erythrocytes of birds (by freezing and thawing) and that such nuclei consumed oxygen about as rapidly as the normal cells. While this indicates that the nucleus is the principal agent in oxidation other experiments of Warburg have been interpreted to indicate that oxida- tion is practically confined to the surface of the cell.!! In these experi- ments” it was found that NaOH greatly increased oxidation in the sea-urchin egg but did not penetrate sufficiently to cause a change of color in eggs stained with neutral red. In a later paper R. Lillie” comes to the conclusion that rapid oxidation occurs at, the surface of the cell as well as at the surface of the nucleus. This conclusion is based upon a study of the indophenol reaction in the corpuscles of frog’s blood. The use of the indophenol reaction may encounter an objection on the ground that the result may depend to a considerable extent on the manner in which the reagent penetrates. If the oxidizing substances of the cell are largely concentrated in the nucleus those which are present in the cytoplasm will first meet the reagent at the cell surface and may produce at that point a deposit of granules of indophenol. In the same manner the oxidizing substances which are retained within the nucleus will first meet the reagent at the surface of the nucleus and produce a deposit in that region. If, therefore, the indophenol reaction shows a higher oxidative activity in the nucleus it may doubtless be depended on, since its error presumably lies in the opposite direction. But if it indicates a marked oxidative activity at the surface of the cell (or at internal surfaces, including that of the nucleus) we must be cautious in drawing conclusions. It would seem that more reliable evidence can be obtained by investigating cases where it is not necessary that the reagent should penetrate from without owing to the fact that the cell itself produces substances which become colored on oxidation. The writer has investigated a case of this kind. The plant chosen was the Indian Pipe, Monotropa unifiora, which is extremely well suited to such investigations because the colorless cells contain a‘ chromo- gen which oxidizes and darkens very rapidly upon injury. An addi- 10 Warburg, O. Zeit. f. physiol. chem. 70: 413. IQIO-II. This interpretation is by no means necessary. Cf. Loeb and Wasteneys, Jour. of Biochemistry 14: 459. 1913; 21: 153. 1915; also, Osterhout, ibid. 19: 335. 1914. Owing to the buffer action of protoplasm and to the presence of pigment the penetration of a small amount of alkali is not easily detected. 2 Warburg, O. Zeit. f. physiol. chem. 66: 305. 1910. Biochem. Zeit. 29: 414. IgIo. 13 Jour. of Biol. Chem. 15: 237. 1913. 344 BROOKLYN BOTANIC GARDEN MEMOIRS tional advantage is that the leaves are so thin and transparent that they may be placed under the microscope and the details of cell struc- ture studied with care before the cells are injured or treated with reagents. In a typical leaf cell the cytoplasm is transparent and nearly color- less, with a few granules, while the nucleus is only slightly less trans- parent and as a rule shows a few granules and a nucleolus. When a leaf is mounted in a drop of water under a cover glass the cells remain unchanged in appearance for hours. If an intact portion of the leaf is cut or crushed the cells in the neighborhood of the injury soon change their appearance. In the course of five or ten minutes the nuclei of the cells nearest the injury assume a more granular (or vacuolated) appearance and soon begin to darken. The darkening does not begin at the surface but appears to take place almost simultaneously throughout the whole mass of the nucleus. Not until the nucleus has become very dark (so as to stand out very conspicuously when the preparation is viewed under the low power of the microscope) does the cytoplasm begin to darken per- -ceptibly. It may be several hours after the nucleus has darkened before a change of color can be perceived in the cytoplasm. (This is also true where the thickness of the cytoplasm has been increased by plasmolysis so as to be as great as that of the nucleus.) The darkening of the cytoplasm does not seem to be more rapid at the surface than elsewhere. That the darkening is due to oxidation is shown by several facts. Among these the following may be mentioned. 1. A microscope slide is smeared with vaseline, a leaf is laid upon the vaseline and more vaseline is carefully placed upon the leaf. A small splinter of glass (from a broken slide) is placed on the leaf and another slide is gently pressed upon it, so as to spread the vaseline and bring the glass splinter close to the leaf without injuring the latter. Care should be taken that any air bubbles which may be included in the vaseline are not in contact with the leaf in the neighborhood of the splinter of glass. The leaf is left over night in order that the oxygen present in the intercellular spaces (or adhering to the surface of the leaf) may be used up by respiration. On the following morning the upper slide is pressed down with sufficient force to drive the splinter into the leaf and crush it. It is then placed on the stage of the microscope and kept under observation. It is found that while some darkening occurs it is at first largely confined to the drops of juice forced out of the leaf by the crushing (the juice seems to spread along the fibro-vascular bundles in some cases). The darkening of the nucleus and cytoplasm is usually much slower than in air (especially with fresh leaves). OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 345 The darkening which occurs is due in part to free oxygen left in the leaf and in part to oxygen in compounds from which it can be split off for the oxidation of the chromogen (analogous to anaérobic respiration). 2. If leaves are torn in two or crushed at once, dropped into boiling water, 0.1 M HCl, 0.1 M NaOH, 0.1 M KCN, or 3 percent H2O: the darkening does not occur. ‘These agents are inhibitors of oxidation in living tissues. Hydrogen peroxide may inhibit at high concentration, but accelerate at low concentrations. In NaOH and KCN the leaf becomes pale yellow: this seems to be due to the action of hydroxyl ions. 3. The chromogen may be extracted by placing stems in 0.1 M NaOH in a bottle completely filled (so as to exclude air) and tightly stoppered (with a glass stopper coated with vaseline). The solution becomes pale yellow (or slightly reddish) and may be kept in this condition for months. On opening the bottle and pouring out the solution into a shallow dish it at once becomes red as the result of oxidation. The behavior seems to be analogous to that of pyrogallol, which is easily oxidized by the air in alkaline solution, but not in neutral solution except under the influence of oxidases (from plants or animals) or other catalyzers. That the darkening of the nucleus is due to oxidation taking place in the nucleus itself and not to the taking up by the nucleus of a stain produced in the cytoplasm or vacuoles is shown by the following experiment. Plants were ground in a mortar and allowed to stand until they became black. The juice was squeezed out and centrifuged, giving an inky fluid. In this were placed pieces of leaves which had been treated with 0.1 KCN and afterward with water. ‘The solution was allowed to stand until it became concentrated by evaporation: it then appeared black. It was found that where the nuclei had been squeezed out of the cut cells by the knife they had taken up some stain but not more than the cytoplasm. In cells which were merely cut open there was little or no staining of the nucleus. We must therefore conclude that oxidation occurs more rapidly in the nucleus than elsewhere in the cell. The only way to escape this conclusion would be by assuming that at the moment of injury there is a sudden migration into the nucleus of some or all of the substances necessary for the oxidation. This is not only very improbable from a theoretical standpoint, but observation shows that it can not be the case, for in this migration the substances would mingle and produce the pigment either outside the nucleus or at its surface before any pigment appeared in the interior of the nucleus. Observation of the nucleus shows that the pigment appears as soon in the interior of the nucleus as at its surface. 346 BROOKLYN BOTANIC GARDEN MEMOIRS We may therefore conclude that the substances necessary for oxidation do not suddenly migrate into the nucleus at the moment of injury, but that they must exist there before the cell is injured. We may ask why the nucleus does not become darkened in the normal condition of the cell. The investigations of several workers have made it probable that the pigments produced by oxidation under normal conditions are at once reduced, giving up their oxygen to other substances in the cell. When injury occurs the reduction is checked more rapidly than the oxidation, with the result that the pigment accumulates. It is also possible that injury causes the admission of oxygen to the cells. In order to test the effect of the indophenol reaction on leaves of Monotropa they were torn in two and placed in a mixture of equal parts of aqueous I percent paraphenylene diamine and saturated aqueous alpha naphthol. “It was found that the result depends some- what on the condition of the reagent. In the most favorable cases the cells which were torn open became pale purple in color almost at once, showing that the reagent readily penetrated them. Usually the cell contents (cytoplasm, nucleus and vacuole) became at first uniformly tinged with purple. After a while the nuclei would usually assume a deeper purple than the remainder of the cell contents. The cells lying a little further from the torn surface, which were injured but not actually torn open, showed at first a pale yellowish color which in some cases became deeper with time and in other cases gave way to a purplish tint. In most of these cells the nuclei gradually became deeper in color than the other cell contents. Later the cytoplasm became in some cases so deep in color as to obscure the nuclei. Cells lying still further from the torn surface changed very slowly (many remaining unchanged after some hours) so that it was evident that the reagent penetrated from the torn surface and not through the outer cell walls (which are normally in contact with the ain. In. most cases the general result, after a few minutes, was a deep purple band along the torn edge: inside the purple band was a yellow- ish one of irregular outline, followed by nearly colorless intact cells further away from the torn edge. In I percent aqueous paraphenylene diamine the results were similar but the purple color was replaced by a dirty brownish-red (with more or less purplish tinge). It should be pointed out that these results are most striking with 4 More rapid penetration from cut or torn surfaces is commonly observed in the entrance of reagents into leaves, petals, etc. OSTERHOUT: THE NUCLEUS AS A CENTER OF OXIDATION 347 reagents which have stood long enough to take up oxygen, in conse- quence of which the paraphenylene diamine becomes reddish in color, while the alpha naphthol assumes a dirty grayish-purple. When the reagents are freshly made up the action is very slow unless hydrogen peroxide be added. When a mixture is made up of equal parts of each of the reagents previously mentioned and 0.3 percent hydrogen peroxide the results are similar to those just de- scribed. But if stronger hydrogen peroxide be used a greater amount of purple coloration is observed in the cells. When 3 percent hydrogen peroxide is used (in place of 0.3 percent) the following changes may be observed. A pronounced purple color appears at once in the torn cells: this spreads rapidly to the adjacent cells, which are still intact, and may extend through several rows of intact cells. In these intact cells the first appearance of change is the formation of purple granules of indophenol in the vacuole. The vacu- ole becomes filled with these granules which show active Brownian movement. Occasionally some of them come in contact with the nucleus (or the film of cytoplasm which covers the nucleus) and stick fast to it. At this time nucleus and cytoplasm are usually free from granules or coloration. The purple color grows more intense until the details of cell structure become obscured. The general conclusion is that while the indophenol reaction indi- cates that the nucleus is the center of oxidation it does not give as definite information on this point as the formation of natural pigments within the cell as the result of the oxidation of substances normally present. SUMMARY Injury produces in the leaf-cells of the Indian Pipe (Monotropa uniflora), a darkening which is due to oxidation. The oxidation is much more rapid in the nucleus than in the cytoplasm and the facts indicate that this is also the case with the oxidation of the uninjured cell. PHYSIOLOGICAL SPECIALIZATION OF PARASITIC FUNGI GEORGE M. REED University of Missouri One of the important developments in plant pathology in recent years has been the discovery of races of well-defined morphological species of parasitic fungi which are restricted to particular hosts. These’ specialized races can be distinguished from each other only by their ability to grow on some host plants and not on others. It is now well established that species of parasitic fungi, identical in their structural features as found on a more or less wide range of plants, may consist of numerous races or strains which differ in their capacity to infect the various hosts. Apparently Schroeter (136), as early as 1879, was the first to call attention to this phenomenon in connection with certain rusts on Carex. It is, however, to Eriksson (34) that we are indebted for a realization of the importance and significance of the host specialization of fungous parasites and for the impetus to the numerous investiga- tions devoted to this phase of plant pathology. Eriksson’s demon- stration of races of Puccinia graminis, P. glumarum, P. dispersa and P. coronata, distinguishable from each other only on the basis of the hosts that they are able to successfully attack, is the real starting point for a general recognition of the phenomenon of host specialization of fungous parasites. Various terms have been introduced to apply to these races or strains which show no anatomical differences, but are distinguishable only by their physiological behavior in the choice of hosts. Schroeter (138), in 1893, suggested the term sister species (Species sorores). Klebahn (76), in 1892, described them as biologische Spezies. Rostrup (121, 122), in 1894, suggested the term diologiske Arter, and, in 1896, proposed another term biologische Rassen. In 1894 Hitchcock and Carleton (63) proposed the term physiological species. Eriksson (34), in 1894, introduced the generally used expression Specialisierte Formen or formae speciales. Magnus (96), in 1894, employed the term Gewohnheitsrassen or adapted races. Marchal (97) applied the term races spéciali ées, while Ward (174), Salmon (123) and others have used the expression biologic forms or biological forms. 348 > REED: SPECIALIZATION OF PARASITIC FUNGI 349 Whatever term has been applied, the underlying conception has been the same, namely, that these races, strains, forms, etc., of distinct morphological species of fungous parasites differ, not in discernible structural features, but in their physiological behavior, as indicated by their ability to infect some hosts and not others. They differ in their ability to establish the parasitic relation with particular hosts and thus secure the necessary food for their normal development. The phenomenon is distinctly physiological and is doubtless quite com- parable to the well-known behavior of saprophytic fungi on different chemical substrata. Various saprophytes, structually similar, vary in their ability to utilize different chemicals as sources of food, de- pendent on their capacity to secrete the necessary enzymes. While the strains of parasites may differ essentially in their ability to secure food from a particular host, we must keep in mind the possibility of a more complicated series of relations in which toxin and antitoxin production are involved. Many investigators of the phenomenon of host specialization have made a large number of species on the basis of the results of their inoculation tests. This is especially the case in the rusts where Klebahn, Eriksson, Schneider, Fischer and others have raised many forms to specific rank, although no distinct structural differences can be observed. It may be noted that the races of Puccinia dispersa, P. sessilis, P. Ribesit-Caricis, P. extensicola, Coleosporium Campanulae, Melampsora populina, M. Tremulae and others, referred to below, are regarded as good species by some students. Fischer (45), in connection with the rusts, accepts as species the following: 1. All rusts which are structurally distinct. 2. All rusts which have a different life-cycle; for example, forms which are distinguished by the presence or absence of certain spore-forms. 3. All forms which differ in their choice of hosts, in so far as the hosts belong to different genera. In heteroecious rusts species are recognized when the hosts of one generation, aecidial or uredo and teleuto, belong to two different genera. Fischer unites under one species as formae speciales or specialized races all rusts which differ only physiologically and whose hosts are species of a single genus. Whether a particular rust is a physiological species or a specialized race is thus determined by the range of its hosts. It is doubtless true that many rusts, and other parasitic fungi as well, which can be distinguished only by the hosts upon which they grow, are just as distinct forms as others which are characterized by 24 350 BROOKLYN BOTANIC GARDEN MEMOIRS minute structural differences. The poplar rusts of the group Melamp- sora Tremulae, with their uredo and teleuto stages on Populus alba and P. tremula and their aecidial stage on such widely separated hosts as Larix decidua, Pinus silvestris, Mercurialis perennis and Chelidonium major, certainly differ from each other in a fundamental way. The difference between these is doubtless as significant as some of the minute structural differences which distinguish other species. It is, however, certainly important to recognize the fact that these rusts, mildews, etc., referred to as specialized races, physiological species, etc., can be distinguished only by cultural tests. This can most easily be done by grouping them together on the basis of struc- tural similarity. Klebahn (92) has done this recently in his scheme for illustrating the relationships of the willow and poplar rusts, and also in the case of the Ribes-Carex rusts. Arthur (10) adopts the same plan in combining the Compositae-Carex rusts under Puccinia ex- tensicola Plowr. In a similar way Tranzschel (155) combines the various Centaurea-Carex rusts under the name Puccinia Centaureae- Caricts. At the present time, a large number of the parasitic fungi have been investigated from the standpoint of specialization to particular hosts and the phenomenon has been found to be of wide occurrence. The present paper is an attempt to bring together the results of numerous investigations bearing on this point. THE RustTs—UREDINEAE Puccinia graminis Pers. Extensive studies have been made in both Europe and the United States on the specialization of the black stem rust of the cereals and other grasses. Eriksson (34, 37, 38, 41) in Sweden, Jaczewski (68) in Russia, Carleton (25, 26), Freeman and Johnson (57), Arthur (2, 5, 6, 7, 8, 10, 11), and Stakman and co- workers (143-149) in the United States, have reported the results of their cultural experiments with this rust. Experiments have been reported in which inoculation tests with both uredospores from the various grass hosts and aecidiospores from the barberry have been used. The general results of these experiments may best be sum- marized as follows: +e S51 SPECIALIZATION OF PARASITIC FUNGI REED e MNjDpiuiDAKd UNnpYiDd a4DBINA UNapAozy | g UNpYyDg adD3INA UNapdoTT | eg dd4DB INA UNAPAO TT ¢ Unaunjuods unapso fT eg Unpisnd UWnapdéojzy zg SNIDUD] SNIOTT g' DUNO DINISAT g AOYDII DINISAT ge SNISNQOA SNUATT g SISUIPDUDI SNWUWATT] VDIDAaUlo]s SYKJIDCT x SISUIPDUDI S1SOABDULDID. zg WNAOJ297 SNUMOAT g SUDBANG SNMOAT e'SNJIAdI SNULOAT e SIMAOLADINAA DIUUDULY IIT zg SNYDII UWNALIYIDUIYAL rijjand unyjunxoyjupy zsusuajpad snangado] y 7 Snpppnauas snandaqo] y ¢ DAafIUoJO]S SYsod3 Y [DIDADXA SYSOAS YF g' DQID SUSOAB YF ge WNIDISII UOAKGOAD 7 ponwf puray 1 DaAyns DUaY "UNIDIVGSQNS UNIJASVA J, ‘sisuaradsuout Uos0ga10q ‘mndagsD WNIIY *DIDISIAI DIAIIION MNUIANUL UNAPAO LT *S4]JOW SNIIOTT ‘yapur “ds DINISay “DIDSNIQO DIUOID “DIDAIULOIS SUKJID “SNIDIPI SNWMOAT "SNYDIA WNAIYIDUIY AA “DIUDUIAD DILYGOUU YF *staqsadqp snangadoyy "DAQDIS S1JSOAB YF “SUpldajs DUAL ‘sisuajpad Duaay "1ddYOOY DUILY ‘pnypf Duaay ‘DAYDS DUAL ‘uoTzL]NIOUT [eIoyIzAe Aq pozoayut A[IY SIS SISOP] ¢ ‘uolzENIOUT YeIoylyAe Aq pazoayur A]IAVIY SYSOP] z *91N}LU UT PUNO} SPM JOVI JY} YOIYM UO S}SOF] , “DUIO DINISAT "SISUIRAD SNULOAT ‘DIpaUu VEU “SNYDIA WNAIYIDUBY Ad ‘sasuaypad snangedo] y *suaasaqnd Duany “pays DUAR TysMozov f—vissny aDUaaY AVY AAZVIVIOIdS ‘sapromodqg DIdjnA MNgXYPOYyIsSyp WNIASIA T, ‘mndadsp Unaty q “SUSUALADUDI STADIDY J ‘mnsn {fa uNnYyryy “DIAND DIYIADUDT “DaInjas 1149190 Y “DAOYINUA] DINIS IT “SAPLOANIIS DINISAT “SNANKUL DINISAT “‘pypAauUiops SYKJIDC "SISUAJIAPDUL SNUMOAT “SKYIDISKYIDAQ SNULOAT "SISUIAAD SNULOAT “DUUIXDUL DET “SN1]DIA WNABYIDUBY Ad ‘sasuaqpad snAnzaqoyy “SUtdas DUAR "10049 DUALLY “pans DUAR UOSSYLIG]—UdpaMg BROOKLYN BOTANIC GARDEN MEMOIRS sey punyisnd wnapso fy punyngnl wnapdo fT 2 SNIWUIBAIA SNUK IT pSNISNGOL SNUA TT PSIsUapdUnd SNUK TT 7g WNAOJIA SNUMOLT eo SuDdand snuiodg eDAYDS DUIBY ‘2dD3 INQ UNapso fT pUindsauay UOAK 018 Y “MNULANUM WNApPAO LT puyus Uostdods “mnDgnl Wnapso FT eg MNILAAGIS UOAKGOAD YY : : “MNSOWOI UNapso FT psuadad Uodk gods V “SNILAIQIS SHUT “MNIPAWMdaquUr UOAKG OAD “pIDAIULOJS SUKJIN "SNUADUIAAD SNUA CALA d s MUTT eUniDILAQUir UOAK Oda “SNUYDIAS SNULOAT “snulpIas snuoLg z WNIDSUOIAa UOAKG OAS [7 “SIUMLABUL SNULOAT. ‘suadad Uodk ods y “MUNIDISIAI UOAKG OAD *suadad Uodk ods ‘MmNAOpAasap UOAK OAD I V f4aSap V punuiuns Uuodtdoda Y ‘MNUIMDI UOAK Odd Y ‘MNUWMDI UOAKG Odd Y ya1Dada9 aypIasy ’ ‘ajpadaI a1DIAS ‘ajpadaI ayvIas spray AVY GAZVIVIOAdS 'a]Dad499 aqpIay easuaipad WnNaly J z SISUIUDUDI SIADIDY J pPDAagyfvnndg DIADINIWUD I eUnyuanwmal WnYOT e aUuadad WN1]0'T A ECCT S pDIDIS1AI 1149190 Y e DINIDE X1AjSK ET u0ya]IV—S9I"IS powuy TysMozov [—vissnyy UOSSYLI]—UdpaMs JesloWolg pur ureuryRe{G—soajejC pou panuyuojy—avuaapy AVY GAZIIVIOAdS a 353 SPECIALIZATION OF PARASITIC FUNGI REED a eet iit SIS Sst SSS zg Dyund snuodg z Snaapapdoy SNULOAT ¢'DAUDS DUAR e'sisuajpag snanzaqo] ¥ puinsaua] UAL G Odd Y punjnridgs uodt gods y puyuus uoskdoss y e'suadad uodk ods FY eUnipaudagqur Uodkdods y 7, unjosUuoja UOLK Odd Y UNIDISIAI UOAKG Odd Y PUnNULUDI UOAKGOdB pAdDB INA WHNIYL T, PUNPIBAN] WNIYLA T, PDUyads wnIgyed J, punauojod WnIyIA T, pUNIIOIOUOUL WNIWYLA T, PUNANP WNIYL T, pMnII0IIp WNIYLA T, PUNJIDG MOI WNIYVU T, ¢ QADS INA WNIYU ysnapunjg&sa snjoqosods ronjod xvas FT zunppuunskd wnpyynd aspajna wnapso py z Mnpiy(pd a4Dd]Na UNapdo TT padpapna Unapdo fT zunaunjuods mnapdojzy] JasIowWarg puv uRUTyLIG—so}eJg powUy) “DIDISVAI 1149190 ‘adDB NA WNap¢o fT “MNULANU UNapso FT “munyoqnl unapao zy “DayUnsis DINISAT “SNIIUIGAIA SHUT “SISUAPDUDI SNUWADT “‘pypdaUioys SUXJIDCT "MNAIUNAL UOAKG OAD YF “mMospany aid UOAKG OAS Y ‘AAD INA WNILLA T, “MUNSO[IA WNIYLA “MNPIBAN] WNIYCA T, “mnaUojod WnII7;Ld T, “WUNIIOIOUOUL UNIVYLA T, “WMNANP WNIYLA T, “MNJIDGWUWOI WNIUYVA T, ‘auuadad UN1]0'T “adDBINA UWNAPAO FT ‘DaJUDSIB DINISAT “MNUIUDI UOAKGOd3 Y *‘suadad UOdK 04d *AdADB INA WNIYVA T, ‘9]Dad99 a]DI9S *adDd INA WNAapso FT “DAYS DUALY ‘QADBING WNIYLA T, WA. ANY AAZIVINAIS uoyopIe—saIeIs powuy il TysMozov f—vissny UOSSyLIG]—UIPIMG panuyuojy—sipoag ANY AAZIIVIONdS — ——— BROOKLYN BOTANIC GARDEN MEMOIRS 354 pSuadad Uosk dois Fy e MNIpPIMAgqUL UOAKG OAS YF ZWNIDSUOoJa UOAKG OAD eg UNAOJAasap UOALG OAD YY Z WN4IDISUI UOAKGOA3 Y padD0 NA WNIYIA T, zg WNIIOIOUOM WNINYLA J. py UNANp WNIT, g WMNIIOIUIP WNINY4VA J, pUnjavd mod WNIYIT, *paqoayur Ap}YS1[s A[UO a1aM Si9yIO ayIyM apqiydadsns 931Inb a19M sorIyJaIVA IWOS , yanguos 194t4 J, AOVY AAZIIVIOIS g ‘1'9/Dada9 aqpIay poynjod xr4jsk FT zunjopiunskd wnpYypvd a4D3]NA WNapso_T , Wnpiypd adD5jNA UNapsOTT z (UDIMISSkqgD) 2d4Dd1NA UNapso LT pa4Dd]Na Unapso fT z Unaunjuods unapso FT pungyisnd wnap4.o HT punqnqnl wnapso fT punsojidsana uUnapso fT PSNIVUIGAU SNUATT PSNISNQOL SNUK ITT puUunorDUL snuck) pPSISUapDUDI SNUKI rstyapjskyonag snucpay 3 WNAOJIA] SNWMOLT jastowarg pur ULWYLIS—saqwIS powuy uo}apIVJ—sa}2IS pou TysMozov [—vIssn yy UOSSYOII—UdIpaMs panuyuoj—toyis [ ANY AAZVIVIOAIS wy 35 SPECIALIZATION OF PARASITIC FUNGI REED ¢'9]Dada9 aqpIaS 2 DIDISIAI 1149190 ¢ AdDBINA UNaps«o LT zSNIDUD] SNIIO FT 2 DIDAIULOTS SYkjIDG ¢ SISUIPDUDI SUSOABDULDIDID zg UWNAOIIA] SNULOAT e DAYDS DUILY 2 Sisuajpad snanzaqoly zSnDINIIUaS SnAnIado] F {Ddafiuojojs SYsods gDUIUDI SYSOAG TDQID SUSOdd VY "StADSINA DQID SY4soss Y “DQID SYSOAB YY *pdaftuopojs SYSOdB V7 "DULUDI SYSOAB YW SYsods YY AOVY AAZITVIONdIS *psojigsavd DALY *DSOjtgSaDI DALY “DIVUIJOG DALY aDdAL py AOVYL GAZVIVIOAS ¢'a]Dada9 aqpIay 7 adDB/N2 UWNaps~o fT pungpgnl wnapso fT pMUunompUL snUK TT PSNIND]S SNUA TT PSNVsuapuod snuc]T pSISUuapDUnI SNUAIT zg WNA0JIA] SNWMOLT undauay UosK gods Y ruynus uost goss Psloulsrg pue ueUTYe}IG—sd9}ej}C pou uoJaIpPIVJ—so}eIS poywuy TysMozoe [—vIssny UOSSYLI7]—UdPIMS panUuyUojy—YIDG MOI 191 ANY AAZITWIOIdS 356 29109409 ayvIay e aUuadad WNIYOT gf MNILIDIY MNYOT pDIDISUdI 1149190 ¢0dDBINA WNApAO LT ge uningnl wnapsopzyT zg SNIDUD] SNIIOLT psisuayvag DINISAT p40UD]9 DINISAT eg SNIUIBAIL SNUG PDIpAdaMioys SsyKjID. eg WNAOIIA] SNULOAT DADS DUaLy e DNID{ DUaLy ge SNYDIA WNAIYIDUIYAA YF zg sisuaypad snangado] yy z Snypynoiwuas snandaqo] yy pasuapad Unayy q *aqpadad aqDIAS “pssadquod DOT *LOUDIA DINISA “pypdawmops SikqIDCT “SNYDIA WNLAAYIDUIYAA YF “pans DUAL ‘asuaypad WNATY J *aqDada9 a1DIAS “40UDIA DINISAT "DAUDS DUAL ‘asuaypad Wnaly J sasuaypag-19]Yq AVY AAZVIVIOAdS “‘DRYDS DUAL “SNYDIA UNABYIDUAYAA PF LdaYyJDUIYAd YF AVY, AAZITVIOAMS | ‘yuan voids vaad y | apdad YW AVY AAZIIVIOAdS | "S098 Ida SUSOABDUDIDI BROOKLYN BOTANIC GARDEN MEMOIRS SUSOABDULDIDI AVL AAZIIVIOAdS ‘DULOAIS DOT “sisuaqDagd DOG “pssadqdusod DOT ‘sisuaypad DOT “DSSadd uo) DOT ‘DISIDI DOT J9SloWolq pur ULUTYLIS—SI}CYS pou) uosuyo (—seqv1s poy | TySMozov [—vissnyy UOSSYIIG]—udpaMs apog AOVY AAZVIVIOdS REED: SPECIALIZATION OF PARASITIC FUNGI 357 According to the various workers the specialized race Avenae occurs on a wide range of hosts. Eriksson (41) lists it on twenty species belonging to fourteen genera; Jaczewski (68) lists it on seven species belonging to six genera; Carleton (25) records it on nineteen species belonging to fifteen genera; Stakman and Piemeisel (149) record it on thirty-three species belonging to twenty-one genera. The specialized race Tritict is reported by Carleton on seventeen species belonging to seven genera; Stakman and Piemeisel list it on thirty- three species belonging to nine genera; Jaczewski lists it on six species belonging to five genera; Eriksson records it on the four cereals and states that this race seems less sharply fixed in its host relations than any of the others. Stakman and Piemeisel report another race, Tritict compacti, from the Palouse wheat region of western United States. This race is reported on twenty species belonging to six genera. The specialized race Secalis also occurs on a number of hosts, Eriksson reporting it on eleven species belonging to five genera, Jaczewski reporting it on six species belonging to four genera and Stakman and Piemeisel recording it on twenty-three species belonging to nine genera. According to Eriksson, Jaczewski and Carleton the specialized race Agrostis occurs only on species of Agrostis, while Stakman and Piemeisel report it on thirteen species of grasses be- longing to ten genera. The specialized race Arrhenatheri occurs on both Avena sativa and Arrhenatherum elatius. The races Airae, Poae, Calamagrostis and A perae are limited to species of a single genus. Eriksson and Henning (43) separated the timothy rust from the black-stem rust and made it a distinct species, Puccinia Phlei-pratensis. The separation was based largely on the fact that they were unable, with one doubtful exception, to infect the barberry with teleutospores from timothy. Other workers have made similar attempts but with- out success. Stakman and Piemeisel consider it best to regard timothy rust as a race of Puccinia graminis. Eriksson (34) reports successful inoculations with uredospores from timothy on Avena sativa, Festuca elatior and Secale cereale, negative results being obtained with Triticum vulgare and Hordeum vulgare. Johnson (69) transferred timothy rust, using uredospores, to Avena sativa, Arrhenatherum elatius, Dactylis glomerata, Festuca elatior, Poa compressa and Secale cereale. Inoculations on thirteen other grasses, including Triticum vulgare and Hordeum vulgare, gave negative results. Stakman and Piemeisel (149), however, were able to infect eighteen species of grasses belonging to thirteen genera, these including Avena sativa, Hordeum vulgare and Secale cereale, but not Triticum vulgare. They regard this race as being closely related to the race Avenae. It may also be noted that Eriksson lists barley (Hordeum vulgare) 358 BROOKLYN BOTANIC GARDEN MEMOIRS and rye (Secale cereale) as hosts for the same specialized race Secalis. Carleton, on the other hand, places wheat and barley as common hosts for the race Tritici, although he gives no information regarding the relation of the rye rust to the other forms. Other variations in the hosts for the different races, as reported by these three workers, also occur. Eriksson lists Dactylis glomerata as a host for the race Avenae; Jaczewski lists it for the race Secalis; and Carleton records it as a host for both races Avenae and Tritici. Agropyron repens is a host for the two races Secalis and Tritici according to Jaczewski. Carleton records the race Avenae on Hordeum murinum, while Eriksson places this host, along with the other species of Hordeum, as a host for the race Secalis. It is important to note that many grasses are listed by Stakman and Piemeisel as common hosts for several races. They suggest that the six races which they experimented with may be divided into two groups on the basis of their parasitism. The races Tritici, Tritici compactt and Secalis form one group; these vary in their capacity for infecting certain hosts but all three vigorously infect Agropyron cristatum, A. elongatum, A. smithii, Bromus tectorum, Hordeum jubatum, H. vulgare, Elymus canadensis and Hystrix patula. The other three . races, Agrostis, Avenae and Phleipratensis, also vary in their infecting capacity but all vigorously attack Alopecurus geniculatus, A. pratensis, Dactylis glomerata, Holcus lanatus and Koeleria cristata. Three hosts, Bromus tectorum, Hordeum vulgare and Secale cereale, are infected by all six races. Stakman and Piemeisel do not regard the barley rust as a distinct race. Barley is a very favorable host for races Secalis, Tritici, and Tritict compacti, as well as being susceptible to the other three races. In the field barley seems especially to harbor the race T7itict. Freeman and Johnson (57) have confined their work almost ex- clusively to the cereal hosts of Puccinia graminis. They conclude that their experiments indicate the existence of four specialized races: Tritict on wheat, Hordei on barley, Secalis on rye, and Avenae on oats. The rust on wheat can be transferred to barley and rye, but not to oats; the rust on barley can be transferred to the other three cereals, rye, oats and wheat; the rust on rye can be transferred to barley, but not to wheat nor oats; the rust on oats can be transferred to barley. These workers also report that Hordeum jubatum can be infected with the rust of both wheat and barley; Agropyron repens with therust of wheat; and Dactylis glomerata with the rust of oats. So far as the cereals are concerned it appears that the different grains may be hosts for more than one specialized race of the black-stem rust. Stak- man and Piemeisel, however, do not regard Hordei as distinct from Tritict. REED: SPECIALIZATION OF PARASITIC FUNGI 359 Stakman’s (143) results with the cereal rusts, in general, confirm those of Freeman and Johnson. He found that uredospores from barley infected rye, barley and wheat, but not oats; uredospores from oats infected wheat, rye and oats, but not barley; .uredospores from rye infected rye, but not wheat nor barley; uredospores from wheat infected barley, rye and wheat, but not oats. Uredospores from Agropyron repens gave the following results: on wheat, on oats, on barley and on rye. Stakman (144) briefly mentions some other results with this rust. He states that uredospores from Agropyron repens, A. tenerum, A. caninum, A. smithii and Hordeum jubatum readily infect barley and rye, very slightly wheat and practically fail to infect oats. Uredo- spores from Dactylis glomerata and Poa nemoralis infect oats, but not the other cereals. Practically no success was attained in trying to infect any of the cereals with uredospores from Agrostis alba, A. stolonifera, Anthoxanthum odoratum, Calamagrostis canadensis, Poa pratensis and P. compressa. Bolley and Pritchard (20) state that uredospores from barley (Hordeum jubatum) and Avena fatua were able to infect wheat and uredospores from wheat produced infection on barley and Hordeum jubatum. Pritchard (110), in North Dakota, suggests that distinct races occur on wheat and barley, although he gives no experimental evi- dence. Another race occurs on rye, oats, Avena fatua, Agropyron repens, A. tenerum and Hordeum jubatum, as indicated by inoculations with uredospores from the gramineous hosts and aecidiospores from the barberry. Gassner (58) has reported brief results on this rust in South America. He was able to infect wheat with uredospores taken from barley, oats, rye, Lolium temulentum and Dactylis glomerata. Barley was also infected by uredospores from wheat. As noted before, the infecting capacity of aecidiospores from bar- berry has been tested, as well as that of the uredospores from various gramineous hosts. Eriksson (41) has infected the barberry with teleutospores from more than fifty different grasses. In some cases the aecidiospores thus experimentally produced were used to inoculate various grasses. In general, the aecidiospores from the barberry are restricted in their ability to infect grasses in the same way as uredo- spores from the grasses which were used as a source for the teleuto- spores for inoculating the barberry. It should be noted, however, that aecidiospores from the barberry, produced by inoculation with teleutospores from Bromus madritensis, Briza maxima, Festuca myurus and Phalaris canariensis, recorded as hosts for the specialized race 360 BROOKLYN BOTANIC GARDEN MEMOIRS Avenae, infected Secale cereale as well as Avena sativa. Eriksson, however, gives no information regarding the infecting capacity of uredospores from the four grasses mentioned. Jaczewski (68) made fairly complete tests with aecidiospores from the barberry, obtained by inoculation with teleutospores from various gramineous hosts. He records exact correspondence between the results obtained with the aecidiospores and uredospores from the grasses used as a source for the teleutosporic infections of the barberry. Pritchard (110) found that aecidiospores from barberry in the open infected Avena sativa, A. fatua, Agropyron tenerum, A. repens, Hordeum jubatum and Secale cereale, but not Hordeum vulgare nor Triticum vul- gare. They thus correspond to the infecting capacity of uredospores from the gramineous hosts. Stakman’s (143) results are in harmony with those mentioned. In one trial, the barberry was infected with teleutospores from wheat, although standing in the open. The aecidiospores, when inoculated onto various plants, infected the following: wheat, barley, oats, rye and Triticum monococcum. In another test, aecidiospores from the barberry, produced by inoculation with teleutospores from Agropyron repens, infected wheat, barley, and rye, but not oats, while a similar series, starting with teleutospores from wheat, infected wheat, barley, rye, but not oats. These results are in close correspondence to those previously mentioned for uredospores. Arthur (7, 8, 10) has infected the barberry with teleutospores from Agrostis alba; the aecidiospores produced infected Hordeum vulgare and Triticum vulgare but not Avena sativa. Teleutospores from Elymus canadensis also infected the barberry but the aecidiospores developed failed to infect Secale cereale and Triticum vulgare. Other results of Arthur are the following: aecidiospores from barberry, arising from inoculation with teleutospores from Agropyron tenerum, infected Avena sativa and aecidiospores, arising from inoculation with teleutospores from Sitanion longifolium, infected Triticum vulgare. Freeman and Johnson (57) studied the variations in the size and shape of the uredospores of the different races. While the uredospores of the same race varied considerably in these points, yet they found well-defined differences in the uredospores of the various races. Stakman and Piemeisel have made similar studies and confirm the conclusions of Freeman and Johnson. Puccinia coronata Corda.. Next to Puccinia graminis the crown rust of grasses has been the most extensively investigated rust from the standpoint of its heteroecism and its restriction to hosts. De- Bary (17) first established the fact that the aecidial stage of a crown rust on grasses occurred on Rhamnus. Plowright (108) seems to have REED: SPECIALIZATION OF PARASITIC FUNGI 361 been the first to suggest that the aecidial stage of the rust on Rhamnus cathartica and Rh. Frangula belonged to two different groups of gramineous hosts. This supposition was later confirmed by Eriksson (34, 37, 42), Klebahn (77-81, 90) and Miihlethaler (101, 102). These same workers have further discovered the existence of races of the rust on the groups of grasses which have their aecidial hosts on different species of Rhamnus. The specialization of Puccinia coronata Corda has been found to be as follows: I, SPECIALIZED RACES WITH THEIR AECIDIAL STAGE ON Rhamnus catharlica, Rh. utilis, Rh. dahurica, Rh. saxatilis AND Rh. Imeretina (Puccinia coronifera Kleb.) 1. Avenae on Avena sativa and A. brevis. 2. Alopecurt on Alopecurus pratensis, A. arundinaceus and, to some extent, on Avena sativa. 3. Festucae on Festuca arundinacea, F. elatior, F. gigantea, F. varia and F. alpina. 4. Lolit on Lolium remotum var. aristatum, L. perenne, L. rigidum, L. temulentum, L. ttalicum, Festuca elatior and Holcus lanatus. 5. Glyceriae on Glyceria aquatica. 6. Agropyri on Agropyron repens. 7. Epigaet on Calamagrostis epigeios and to some extent on Avena sativa. 8. Holct on Hoicus lanatus and Lolium perenne. g. Bromi on Bromus erectus, B. erectus var. condensatus, B. inermis, B. secalinus, B. stertlis, B. tectorum, B. commutatus and probably B. asper. 10. Arrhenatherit on Arrhenatherum elatius. II. SPECIALIZED RACES WITH THEIR AECIDIAL STAGE ON Rhamnus Frangula, Rh. Purshiana, Rh. Alaternus, Rh. californica AND Rh. Imeretina (Puccinia coronata (Corda) Kleb.) 1. Calamagrostis on Calamagrostis arundinacea, C. lanceolata, C. phragmitoides, C. calybea and Phalaris arundinacea. . Phalaridis on Phalaris arundinacea, Calamagrostis arundinacea and C. lanceolata. . Agrostis on Agrostis vulgaris and A. stolonifera. . Holci on Hoicus lanatus and H. mollis. . Agropyri on Agropyron repens. nb wn III. Race WITH ITs AECIDIAL STAGE ON Rhamnus alpina, Rh. pumila, Rh. Imeretina AND Rh. Purshiana (Puccinia alpinae-coronata Mihlethaler) Calamagrostis on Calamagrostis varia and C. tenella. IV. RACE WITH ITs AECIDIAL STAGE ON Rhamnus dahurica (Puccinia himalensis (Barclay) Diet.) Brachypodii on Brachypodium silvaticum, V. RACE WHOSE AECIDIAL STAGE IS UNKNOWN Melicae on Melica nutans. 362 BROOKLYN BOTANIC GARDEN MEMOIRS It will be noted that the specialized races of groups I, II, and III have Rhamnus Imeretina as a common aecidial host. Rhamnus Purshiana also occurs as an aecidial host for races of groups IJ and III. It also appears that specialized races on Holcus and Agropyron occur in both groups I and IJ. In connection with the gramineous hosts certain specialized races also overlap. For example, Avena sativa is a host for specialized races Avenae, Alopecuri and Epigaet (Group 1); Festuca elatior is a host for specialized races Lolit and Festucae (Group 1); Calamagrostis arundinacea, C. lanceolata and Phalaris arundinacea are hosts for specialized races Calamagrostis and Phalaridis (Group II). Carleton (26), in this country, has tested the host relations of the crown rust on oats. He finds that the rust on Avena sativa can be transferred to Avena sativa patula, A. sativa orientalis, A. satwa nuda, A. fatua, A. pratensis, Alopecurus alpestris, Phleum pratense, Ph. asperum, Dactylis glomerata, Aira caespitosa, Holcus mollis, Eatonta sp. indet., Koeleria cristata, Anthoxanthum odoratum, Festuca sp. indet., Phalaris arundinacea, Polypogon monspeliensis, Trisetum subspicatum, Brizopyron siculum and Poa annua. Carleton also transferred the rust from Phalaris caroliniana to Avena sativa and Dactylis glomerata. The rust on Arrhenatherum elatius was also transferred to Avena sativa. Aecidiospores from Rhamnus lanceolata readily infected Phalaris caroliniana and Avena sativa. Arthur (6, 11, 14) reports the successful infection of Avena sativa with aecidiospores from Rhamnus lanceolata, Rh. caroliniana and Rh. cathartica. He also succeeded in infecting Rhamnus alnifolia with teleutospores from Calamagrostis canadensis. ‘Teleutospores from Holcus lanatus and Scolochloa festucacea failed to infect Rhamnus cathartica. Treboux (159, 160) reports the results of experiments with the crown rust carried out in southern Russia which are quite at variance with those obtained in Europe. He finds that aecidiospores from Rhamnus cathartica obtained from a common source infected fifty- one species of grasses belonging to the genera Alopecurus, Agropyron, Agrostis, Aira, Arrhenatherum, Avena, Brachypodium, Briza, Bromus, Calamagrostis, Dactylis, Eatonia, Festuca, Glyceria, Hierchloa, Holcus, Hordeum, Koeleria, Lolium, Melica, Phalaris, Poa, Phleum, Poly- pogon, Sclerochloa, Secale, Sesleria and Triticum. In these experi- ments with Puccinia coronifera Kleb., Treboux obtained positive results with three hosts of P. coronata (Corda) Kleb.—A grostis stolontf- era, Calamagrostis arundinacea and Phalaris arundinacea. Treboux has also used the aecidiospores from Rhamnus Frangula to successfully infect nine species of grasses, among them Avena sativa, a host belong- ing to P. coronifera Kleb. Treboux’s tests with uredospores show the REED: SPECIALIZATION OF PARASITIC FUNGI 363 same lack of specialization to particular hosts as Carleton reports for the crown rust in the United States. Puccinia glumarum (Schm.) Eriks. and Henn. Eriksson (34) is the only investigator to report on the host relations of this rust. He claims the existence of five specialized races: Tritict on Triticum vulgare. Secalis on Secale cereale and Triticum vulgare. Elymi on Elymus arenarius. Agropyri on Agropyron repens. Hordei on Hordeum vulgare. Puccinia dispersa Eriks. and Henn. Ericksson (34, 40) first separated the hosts of this rust into five distinct groups each sup- porting a distinct specialized race. Later each race was raised to specific rank, based on the host relations and life history. 1. Secalis on Secale cereale. This race has its aecidial stage on different species of Anchusa. 2. Agropyrti on Agropyron repens (Puccinia agropyrina Eriks.). Aecidial host unknown. 3. Bromi on Bromus species (Puccinia bromina Eriks.). According to Miiller (99), this race has its aecidial stage on Pulmonaria mon- tana and Symphytum officinalis. 4. Tritict on Triticum vulgare (Puccinia triticina Eriks.). Aecidial host unknown. 5. Holci on Holcus lanatus and H. mollis (Puccinia holcina Eriks.). Aecidial host unknown. 6. Triseti on Trisetum flavescens (Puccinia Triseti Eriks.). Aecidial host unknown. Miiller, Ward, and Freeman have studied the rust of the bromes. Miiller (103) found that aecidiospores from Pulmonaria montana infected Bromus arvensis, B. brachystachys, B. erectus, B. mollis and B. secalinus. Aecidiospores from Symphytum officinalis also infected these species of Bromus and, in addition, B. brizaeformis. Miiller obtained the following results with uredospores: (1) uredospores from Bromus erectus infected B. arvensis, B. brachystachys, B. erectus, B. macrostachys and B. mollis; (2) uredospores from B. arvensis infected B. arvensis, B. brachystachys, B. inermis and B. mollis; (3) uredospores from B. mollis infected B. brachystachys, B. macrostachys and B. mollis. Ward (171, 172) has carried out a large series of experiments with this rust. In his inoculation tests he used species of Bromus belonging to each of the five recognized subdivisions of the genus. Uredospores from eleven different species, belonging to three different sections of the genus, were used. Ward found marked differences in the sus- ceptibility of the bromes to the uredospores from different hosts. In kW bY & 364 BROOKLYN BOTANIC GARDEN MEMOIRS general, he concludes that species of the same section of the genus as the one serving as a source of uredospores were more fully infected than the species of other sections. The evidence for Ward’s con- clusion is not very striking except in the case of the two hosts Bromus mollis and B. sterilis. Some of Ward’s data may be indicated as follows: Source of Uredospores Host Inoculated | Bromus| Bromus \Wespaostes| Barorace| arven- | brizae-| Prourus secalt- |arduen-| Bromus | sts | forms mollis nus | nensis| Sterilis Serrafalcus: | | Bromus arvensis.............:: | 12/135| 10/9 33/97 6/8 | Bromus brachystachys.......... 13/14 | 14/14 TAT | Bromus brizaeformis...........) 7/17] 7/15| 14/25 | 3/5 | BL OMUUSHKG. CUSCUM Bieta ie nchtee | 2T/27 a1 | 14/59 Bromus macrostachys........... | 14/15 | 16/17 5/19 5/5 | BrOMUS MOLtOVINIS 22,2 see oe - 2/6 1/25 BROMUSINOLIS See eee Tao 8/15 | 21/26| 119/154 | 3/8 1/3] er Bromus pendulinus.. 2. .50.0:- 8/6 | 43/40) 30/50 | | 17/65 (BLOMUS. SCCOLUNUS:. 3.0 act aes oe 14/14 |14/15| 31/61 |16/16| 8/8 IBY OIMUSHUCSTULIUES eemeere exe icles exe 2/Ae | | 1/4 Libertia: | Bromus arduennensis........... | | 13/14 | | 8/7 Stenobromus: | (BLOMUS EUSSONUA ee wie ee eee oe | 10/26 6/53 | 37/60 BEGMUS MOAN SES ates e)toisiee « | | 11/13 1/13 | 43/68 BRONUVUSHINGXUNTUUS eae ieiereeieier | 1/74 | 2/82 BLOMUS SLCTUTS akties einen | 4/148 | | 126/146 Freeman (56) has made a further study of the brome rust, using uredospores from Bromus mollis and B. sterilis. He states that twenty- two different species of Bromus remained free from infection following inoculation with rust from both hosts. Eleven species were infected with uredospores from B. mollis but not with uredospores from B. sterilis. Only one host, B. sterilis, was infected by uredospores from B. sterilis, and not by uredospores from B. mollis. Five species were infected with spores from both grasses. Puccinia Stipina Tranzschel. Under this name, Tranzschel (156) groups the North American rust Puccinia Stipae Arth., with uredo and teleuto on Stipa spartea and aecidial stage on Aster ericoides, A. multiflorus, A. Novae-angliae and Solidago canadensis, and the Euro- pean rust, Puccinia Stipae Bubak, with uredo and teleuto stages on Stipa capillata and aecidial stage on Thymus and Salvia. Klebahn (87, 91) finds evidence for two specialized races in the European rust: 5 The denominator indicates the number of leaves inoculated and the numerator the number infected. REED: SPECIALIZATION OF PARASITIC FUNGI 365 1. Thymi-Stipae WKleb.; aecidial stage on Thymus serpyphyllum, T. angustifolius and T. vulgaris. 2. Salviae-Stipae Kleb.; aecidial stage on Salvia silvestris and S. pratensis. Puccinia sessilis Schneid. This rust, which has its uredo and teleuto stages on Phalaris arundinacea and its aecidial stage on various plants of the Liliaceae, Amaryllidaceae, Araceae and Orchidaceae, has been broken up into a number of distinct species, based largely upon the choice of the aecidial host. Klebahn (77-83, 86, 87, 89-91) has carried out the cultural tests with the rust and recognizes the following relations: 1. Puccinia Smilacearum-Digraphidis (Sopp.) Kleb. a. Smilacearum-Digraphidis typica Kleb.; aecidial stage on Con- vallaria majalis, Maianthemum bifolium, Polygonatum multiflorum, P. officinale, P. verticillatum and Paris quadri- folia. b. Convallariae-Digraphidis (Sopp.) Kleb.; aecidial stage on Convallaria majalis. c. Paridi-Digraphidis (Plowr.) Kleb.; aecidial stage on Paris quadrifolia. . Puccinia Allii-Phalaridis Kleb.; aecidial stage on Allium ursinum. 3. Puccinia Orchidearum-Phalaridis Kleb.; aecidial stage on Gymna- denia conopea, Listera ovata, Orchis maculata, O. latifolia, Platanthera bifolia and P. Chlorantha. 4. Puccinia Ari-Phalaridis (Plowr.) Kleb.; aecidial stage on Arum maculatum. 5. Puccinia Schmidtiana Diet.; aecidial stage on Leucojum aestivum and L. vernum. It may be especially noted that, according to Klebahn, Puccinia Smilacearum-Digraphidis includes a race which occurs on a number of hosts and two races which occur on only one host. The hosts for the second two races are also hosts for the first race. It is further to be noted that the species recognized are also to be distinguished on the basis of host relations. Puccinia Caricis (Schum.) Rebent. Klebahn (91) indicates the following specialization in the Urtica-Carex rust, the races being indi- cated by the choice of uredo and teleuto host: 1. Urticae-acutae on Carex acuta, C. Goodenoughti, to a less extent C. stricta. 2. Urticae-hirtae on Carex hirta. 3. Urticae-acutiformis on Carex acutiformis, C. Kochiana, to a less extent on C. pseudocyperus. 4. Urticae-vesicariae on Carex vesicaria. 25 No 366 BROOKLYN BOTANIC GARDEN MEMOIRS Tranzschel (155) suggests the possibility of two other races, one on Carex pallescens and a second on Carex vaginata. Puccinia Centaureae-Caricis Tranz. Tranzschel (155) suggests that the various Centaurea-Carex rusts may best be grouped under the above name. ‘The following rusts are thus included: 1. Puccinia Caricis-montanae Ed. Fischer. Bandi (15) claims to have found evidence for the occurrence of specialization in this heteroecious rust which forms its uredospores and teleuto- spores on Carex montana and its aecidiospores on species of Centaurea. He mentions two specialized races based on the choice of aecidial host: a. On Centaurea scabiosa. b. On Centaurea montana. . Puccinia tenuistipes Rostrup; aecidial stage on Centaurea Jacea; uredo and teleuto stages on Carex muricata. 3. Puccinia arenarticola Plowr.; aecidial stage on Centaurea nigra; uredo and teleuto stages on Carex arenaria. 4. Puccinia Jaceae-leporinae Tranz.; aecidial stage on Centaurea Jacea; uredo and teleuto stages on Carex leporina. 5. Puccinia Jacea-capillaris Tranz.; aecidial stage on Centaurea Jacea; uredo and teleuto stages on Carex capillaris. 6. An unnamed rust with aecidial stage on Centaurea orientalis; uredo and teleuto stages on Carex gynobasis. Puccinia extensicola Plowr. This name is applied to a group of rusts which have their aecidial stage on Compositae and their uredo and teleuto stages on species of Carex. Arthur (2, 3, 5, 6, 8, 9, 10, 11, 13) has listed a number of forms under distinct names, but later suggests that they are merely specialized races of Puccinia extensicola. The following rusts are regarded as belonging here: 1. Puccinia Caricis-Erigerontis Arth.; aecidial stage on Erigeron annuus, E. canadensis and E. philadelphicus; uredo and teleuto stages on Carex festucacea. 2. Puccinia Caricis-Asteris Arth.; aecidial stage’on Aster acuminatus, A. adscendens, A. cordifolius, A. paniculatus and Solidago gramintfolia; uredo and teleuto stages on Carex festiva, C. foenea C. retrorsa, C. rosea, C. scoparia and C. trisperma. 3. Puccinia Caricis-Solidaginis Arth.; aecidial stage on Solidago caesia, S. canadensis, S. graminifolia, S. rigida, S. serotina and S. ulmifolia; uredo and teleuto stages on Carex Jamesiit, C. scoparia, C. sparganoides and C. stipata. It is suggested that a distinct race may occur on Carex scoparia and Solidago graminifolia. Puccinia silvatica Schroet. This rust is reported as having its NO REED: SPECIALIZATION OF PARASITIC FUNGI 367 uredo and teleuto stages on various species of Carex, while the aecidial stage occurs on Taraxacum officinalis, Crepis biennis, Lappa officinalis and three species of Senecio. Schroeter (137) proved that the aecidium on Taraxacum was connected with the rust on Carex brizoides and C. praecox. Later he connected the aecidium on Senecio nemorensis with the rust on Carex brizoides. Dietel (27) connected the aecidium on Lappa with the Carex rust and Juel (72) and Bubak (24)sestablished the connection between the aecidium on Crepis and the Puccinia on Carex. Wagner (167, 169), however, claims that a particular collec- tion of teleutospores from Carex would not infect Taraxacum, Lappa and Senecio but only one of these aecidial hosts. Some collections of teleutospores infect one aecidial host while other collections infect a still different aecidial host. There is an indication, then, of a special- ization to particular hosts in this rust. Puccinia Ribesii-Caricis Kleb. This rust has its aecidial stage on species of Ribes and its uredo and teleuto stages on various species of Carex. Arthur (2, 4, 5, 6,7, 8, 11, 13, 14), in this country, has carried out inoculation tests with the Ribes-Carex rust for a number of years. He finds that the aecidial hosts include Ribes aureum, R. Cynosbati, R. gracile, R. prostratum, R. rotundifolium and R. uva-crispa; other species not infected are R. floridum, R. oxyacanthoides and R. rubrum. Teleutospores were used from Carex arctata, C. crinita, C. debilis, C. gracillima, C. intumescens, C. pallescens, C. pubescens, C. squarrosa, C. tenuis, and C. tetanica. Fraser (55) reports the successful infection of Ribes oxyacanthoides with teleutospores from Carex arctata and Carex crinita. There appears to be no indication of the existence of specialized races. In Europe, Klebahn (87, 91) has extensively studied the Rzbes- Carex rust and finds evidence for the existence of five specialized races. The specialization occurs very largely in the choice of the uredo and teleuto host, as all the races pass over to practically the same species of Ribes as aecidial hosts. Klebahn distinguishes the following on the basis of cultural tests: 1. Puccinia Pringsheimiana Kleb.; uredo and teleuto tages on Carex acuta, C. caespitosa, C. Goodenoughu, and C. stricta; aecidial stage on Ribes alpinum, R. aureum, R. Grossularia, R. rubrum and R. sanguineum. 2. Puccinia Ribesii-Pseudocyperi Kleb.; uredo and teleuto stages on Carex pseudocyperus; aecidial stage on Ribes alpinum, R. aureum, R. Grossularia, R. nigrum, R. rubrum and R. san- guineum. 3. Puccinia Ribis nigri-Paniculatae Kleb.; uredo and teleuto stages on ' Carex paniculata and C. paradoxa; aecidial stage on R. alpinum, R. aureum, R. nigrum, R. rubrum and R. sanguineum. 368 BROOKLYN BOTANIC GARDEN MEMOIRS 4. Puccinia Magnusti Kleb.; uredo and teleuto stages on Carex acuti- formis and C. riparia; aecidial stage on Ribes alpinum, R. aureum, R. nigrum and R. sanguineum. 5. Puccinia Ribis nigri-acutae Kleb.; uredo and teleuto stages on Carex acuta and C. stricta; aecidial stage on Ribes alpinum, R. aureum, R. nigrum and R. sanguineum. Puccinia’Bistortae (Str.) DC. This rust has its uredo and teleuto stages on species of Polygonum and its aecidial stage on various umbellifers. It is broken up into distinct races based upon the choice of the aecidial host. By some, these races are regarded as true species. According to Klebahn (91) they are as follows: 1. Puccinia Angelicae-Bistortae WKleb. with the aecidial stage on Angelica silvestris and Carum carvt. 2. Puccinia Conopodii-Bistortae Kleb. with the aecidial stage on Conopodium denudatum. Puccinia mammillata Schroet. This rust, also with its uredo and teleuto stages on Polygonum, as a result of the work of Bubak (21) and Semadeni (140), is separable into two races: 1. Puccinia Angelicae-mammillata WKleb. with the aecidial stage on Angelica silvestris (Aecidium Bubakianum Juel). 2. Puccinia Mei-mammillata Semadeni with the aecidial stage on Meum mutellina. Puccinia Polygoni-amphibii Pers. Several workers claim the existence of at least two rusts on the various species of Polygonum on the basis of minor morphological characteristics. Puccinia Poly- goni-amphibii Pers. is recorded on Polygonum amphibium and P. Polygoni-Convolvuli DC. on Polygonum convolvulus. P. and H. Sydow (151) in their monograph of the rusts, however, claim that the differences are not sufficient to distinguish the species and consequently list the various Polygonums as hosts for the one rust, Puccinia Poly- goni-amphibu. Tranzschel (152, 153) first demonstrated the heteroecism of this Polygonum rust, connecting the uredo and teleuto stages on Polygonum amphibium with the aecidial stage on Geranium palustre and G. pratense. He further found that the uredo and teleuto stages on Polygonum convolvulus was connected with the aecidial stage on Geranium pusillum. A number of other workers have confirmed the connection between the uredo and teleuto stages on Polygonum and the aecidial stage on Geranium. Jacob (66, 67) has carried out the most extensive series of cultural experiments with the Polygonum rust, using teleutospores from Poly- gonum amphibium, P. convolvulus and P. dumetorum. The aecidio- spores produced experimentally on the various species of Geranium, REED: SPECIALIZATION OF PARASITIC FUNGI 369 and also uredospores produced by aecidiosporic inoculations were also used. As the result of the experiments, Jacob concludes that the rusts on Polygonum amphibium and on P. convolvulus and P. dume- torum are distinct. The relationship of these two forms may be indicated as follows: 1. Puccinia Polygoni-amphibit Pers.; aecidial stage on Geranium albanum, G. columbinum, G. dissectum, G. lucidum, G. molle, G. nodosum, G. pratense, G. pusillum, G. pyrenaicum, G. rivulare, G. rotundifolium and G. sanguineum; uredo and teleuto stages on Polygonum amphibium. 2. Puccinia Polygont Alb. et Schw. (P. Polygoni-Convolvuli DC.); aecidial stage on Geranium columbinum, G. dissectum, G. molle, G. pusillum, and G. rotundifolium; uredo and teleuto stages on Polygonum convolvulus and P. dumetorum. It may be noted that the same species of Geranium occur as hosts for both races, although the host list for Puccinia Polygoni-amphibii includes more species than the other. Puccinia absinthii DC. Klebahn (91) suggests the probability of specialized races in this rust although no cultural experiments have been carried out. He reports, however, minute differences in the spore measurements of the different forms. The following races are indicated: 1. Absinthit on Artemisia absinthium. 2. Artemisiae on Artemisia vulgaris. 3. Abrotani on Artemisia abrotanum. : Puccinia bullata (Pers.) Winter. This rust, according to Semadeni (135, 136), has a race specialized to Silaus pratensis and one to Thys- selinum palustre. Puccinia carduorum Jacky. Probst (112) recognizes three special- ized races in this rust. I. Crispi on Carduus crispus and C. personata. 2. Deflorati on Carduus defloratus. 3. A third race probably occurs on Carduus nutans. Puccinia Centaurea Mart. Jacky (64) and Hasler (62) have both studied the specialization of this rust. Jacky suggested the occurrence of two specialized races: (a) Jaceae on Centaurea Jacea and (b) Ner- vosae on C. nervosa. Hasler separates out three rusts, making them species and finds additional host specialization in one of these. His arrangement is as follows: 1. Puccinia Centaureae-vallesiacae Hasler on Centaurea vallesiaca, C. alba, C. rhenana and, to a less extent, on C. cyanus. 2. Puccinia Jaceae Otth on Centaurea Jacea, C. rhenana and, to a less extent, on Centaurea austriaca, C. Jacea var. longifolia, C. phrygia and C. transalpina. 370 BROOKLYN BOTANIC GARDEN MEMOIRS 3. Puccinia Centaureae (Mart.) Hasler. a. Scabiosae on Centaurea scabiosa. b. Nigrae on Centaurea nigra. c. Nervosae on Centaurea nervosa. d. Transalpinae on Centaurea transalpina, C. alba, C. austriaca, C. Jacea var. longifolia, C. nervosa, C. nigrescens and C. bhrygia. Puccinia chaerophylli Purt. Semadeni (139, 140) claims that a race of this rust occurs on Anthriscus silvestris and another on Chaero- phyllum aureum. Puccinia Epilobii-tetragoni (DC.) Winter. Dietel (29) was unable to infect Epilobium hirsutum with aecidiospores from E. tetragonum. This is the only indication of specialization in this rust. Puccinia Geranii-silvatici Karst. This rust has been reported on a few species of Geranium in widely separated localities. It occurs commonly on Geranium silvaticum in Europe. Its restriction to widely separated regions has led to the suggestion that the rust consists of geographically specialized races. Jacob (66, 67) has shown by cul- tural experiments that there is no specialization in Europe, for both Geranium silvaticum and G. rotundifolium are readily infected by teleutospores from the former. Puccinia Helianthi Schw. Arthur, Kellerman, and Jacky have made inoculation tests using teleutospores of this rust. Jacky (65), using teleutospores from Helianthus annuus, infected’H. annuus, H. cucumertfolius, and H. californicus, but failed to infect H. maximiliant, H. multiflorus, H. rigidus, H. scaberrimus and H. tuberosus. Kellerman (74, 75) obtained negative results with teleutospores from H. annuus on eighteen species of Helianthus. His results with teleutospores from H. ambiguus were negative on twelve species and also negative with teleutospores from H. decapetalus on eight species. He found, however, that teleutospores from H. mollis infected H. annuus and H. mollis but not fourteen other species; teleutospores from H. grosse-serratus infected H. annuus, H. decapetalus, H. giganteus, FI. grosse-serratus, H. Kellermanit, H. orygalus and H. tracheifolius, but not H. maximiliani nor H. mollis; teleutospores from H. tuberosus infected only H. annuus. Arthur (3, 4, 5, 6) secured the following results: (1) Teleutospores from Helianthus mollis infected H. annuus, H. hirsutus, H. mollis, H. occidentalis, H. strumosus and H. tomentosus, but not H. grosse- serratus, H. Kellermani, H. laetiflorus, H. longifolius, H. orygalus and HT, tuberosus; (2) teleutospores from H. grosse-serratus infected H. annuus, H. grosse-serratus, H. maximiliant and H. tomentosus but not FH. decapetalus, H. hirsutus, H. laetiflorus, H. mollis, H. occidentalis, REED: SPECIALIZATION OF PARASITIC FUNGI SHA H. orygalus, H. strumosus nor H. tuberosus; (3) teleutospores from H. laetiflorus infected H. annuus, H. divaricatus, H. Kellermani, H. laetiflorus, H. mollis, H. occidentalis and H. tomentosus, but not H. grosse-serratus, H. hirsutus, H. orygalus, H. strumosus, nor H. tuberosus. About the only conclusion that one can draw is that H. annuus and H. tomentosus are readily infected with teleutospores from a variety of hosts. The evidence for distinct specialized races is not very clear. Tranzschel (158) states that he was able to infect Helianthus annuus with teleutospores from Xanthium strumarium; the aecidiospores produced on Helianthus annuus infected both HH. annuus and Xan- thium strumarium. Puccinia Hieracii (Schum.) Mart. Jacky (64) first investigated this rust and found some evidence for specialization, distinguishing one race on Hieracium villosum and suggesting the possibility of several others. Probst (113), more recently, has studied the rust and has come to the conclusion that there are two subspecies, each with several specialized races. He arranges the forms as follows: 1. Puccinia Piloselloidarum Probst. . Hoppeani on Hieracium hoppeanum. . Peleteriani on Mieracium peleterianum. Pilosellae (a) on Hieracium pilosella. Pilosellae (b) on Hieracium pilosella. Velutint on Mieracium pilosella velutinum. Auriculae on Mieracium auricula and H. peleterianum. Floretini on Hieracium florentinum var. obscurum. Ziziant on Hieracium bauhini, H. florentinum var. alethes, H. pratense and H. zizianum. 2. Puccinia Hieracit. a. Silvatict on Hieracium glaucum, H. humile, H. ochroleucum, H. pictum, H. silvaticum and H. trebevicianum. b. Silvatici pleiotricht on Hieracium silvaticum var. pleiotrichum. c. Stuvatict gentili on Hieracium folcanum and H. intybaceum. d. Schmidt on Hieracium Schmidtii, H. humile and H. ochro- leucum. e. Cinerascentis on Hieracium cinerascens and H. ochroleucum. Puccinia Leontodontis Jacky. Probst (112) states that three specialized races occur in this rust: 1. Hispidi on Leontodon hispidus. 2. Autumnalis on Leontodon autumnalis. 3. Pyrenaict on Leontodon pyrenaicus. Puccinia petroselini (DC.) Lindr. Semadeni (139, 140) dis- tinguishes two specialized races of this rust, one on Aethusa cynapium and the other on Petroselinum sativum. Tene Ro v8 52 BROOKLYN BOTANIC GARDEN MEMOIRS Puccinia Pulsatillae Kalchbr. Bubak (22) distinguishes the fol- lowing races of this rust, basing the separation on the distribution and character of the sorus, not on cultural experiments: 1. Concortica on Pulsatilla alpina and P. sulphurea. 2. Atragenicola on Atragene alpina. 3. Genuina on Anemone silvestris and Pulsatilla patens. . Pulsatillarum on Pulsatilla pratensis and P. vulgaris. Puccinia Ribis DC. Eriksson (39) distinguishes a specialized race Rubri, for he found that teleutospores from Ribes rubrum would infect this species but not R. nigrum nor R. Grossularia. Uromyces alchimillae (Pers.) Lév. Fischer (53) has carried out a few experiments with this rust which tend to show the existence of a host specialization. The results reported, however, do not indicate any close correspondence between the plants infected with a particular collection of uredospores and the systematic grouping of the host plants within the genus Alchimilla. Fischer found that uredospores from hosts belonging to the section Vulgares infected plants belonging to sections Pubescentes and Splendentes. Uromyces caryophyllinus (Schrank) Winter. Fischer (49-51) gives experimental evidence for the occurrence of specialized races in the carnation rust. The rust is heteroecious, the aecidial stage oc- curring on Euphorbia Gerardiana and the uredo and teleuto stages on various Caryophyllaceae. Fischer finds that aecidiospores from Euphorbia Gerardiana in one locality are able to infect only Tunica prolifera, while aecidiospores collected on the same host in another region are able to infect only Saponaria ocymoides. In still other localities, however, a race of rust is found that is able to infect both Tunica and Saponaria. Uromyces Dactylidis Otth and U.Poae Rabh. These rusts, probably indistinguishable by any well-defined structural character- istics, are further alike in having their aecidial stage on species of Ranunculus. The former develops its uredo and teleuto stages on Dactylis glomerata, while the latter has the corresponding stages on several species of Poa, Agrostis alba also being listed asa host. A large number of workers have contributed to our knowledge concerning the heteroecism of these rusts. Krieg (93, 94) and Klebahn (89, 91) have worked with Uromyces Dactylidis from the standpoint of specialization and the following races are indicated: 1. Aecidial stage on Ranunculus bulbosus and R. repens. 2. Aecidial stage on Ranunculus lanuginosus, to a slight extent on R. bulbosus (Uromyces lanuginosi-dactylidis Kleb.). 3. Aecidial stage on Ranunculus aconitifolius, R. alpestris, R. glacialis, and R. platanifolius (Uromyces platanifolii-dactylidis Krieg.). aS REED: SPECIALIZATION OF PARASITIC FUNGI 33 4. Aecidial stage on Ranunculus silvaticus (Uromyces silvatict-dactylidis Krieg.). 5. Aecidial stage on Ranunculus acer and R. polyanthemos. Juel (73) distinguishes several forms in Uromyces Poae: 1. Ficariae-nemoralis on Ranunculus Ficaria and Poa nemoralis. . Ficariae-trivialis on Ranunculus Ficaria and Poa nemoralis and P. palustris. Ficariae-pratensis on Ranunculus Ficaria and Poa pratensis. 4. Repentis-nemoralis on Ranunculus repens and R. bulbosus and Poa nemoralts. 5. Repentis-trivialis on Ranunculus repens and Poa trivialis and P. annua. 6. Auricomi-pratensis on Ranunculus auricomus and Poa pratensis and P. nemoralis. . Cassubici-pratensis on Ranunculus cassubicus and Poa pratensis. . Repentis-pratensis on Ranunculus repens and Poa pratensis. . Bullati-bulbosae on Ranunculus bullatus and Poa bulbosa. Uromyces Fabae (Pers.) de Bary. Jordi (70, 71) has distinguished the following specialized races on this autoecious rust which is recorded as occurring on species of Lathyrus, Lens, Pisum, and Vicia: 1. On Lathyrus vernus and probably on Pisum sativum. 2. On Vicia Faba and Pisum sativum. 3. On Vicia cracca, V. hirsuta and Pisum sativum. It should be noted that Piswm sativum is listed as a host for all three specialized races. Uromyces Geranii (DC.) Otth and Wartm., and U. Kabatianus Bubak. The first mentioned species is reported as occurring on a large number of different species of Geranium, while the second men- tioned has been reported only on Geranium pyrenaicum. Bubak (23) has called attention to minor morphological differences between the two rusts. Bock (19) carried out tests with uredospores of Uromyces geranit from Geranium silvaticum and was able to infect sixteen species while thirteen gave negative results. Jacob (66, 67) also conducted experi- ments with the rust, using teleutospores from Geranium silvaticum and aecidiospores produced from the teleutosporic inoculation. A con- siderable number of species of Geranium were successfully infected. A few gave negative results. Jacob also used teleutospores of Uro- myces Kabatianus from Geranium pyrenaicum and found that this rust could also be transferred to a considerable number of other species of Geranium. The host range of the two rusts is very similar and the differences between the two depend upon minor structural differences and not on host relations. iS) 2 XO Co SI 374 BROOKLYN BOTANIC GARDEN MEMOIRS Uromyces Pisi (Pers.) Winter. This rust is heteroecious forming its aecidial stage on various species of Euphorbia and its uredo and teleuto stages on species of Lathyrus and Pisum. The aecidial my- celium is perennial in the Euphorbia host. It may also be noted that aecidial stages on Euphorbia, especially E. cyparissias, have been connected with a number of different species of Uromyces on legumes. The aecidia belonging to these different rusts are quite indistinguish- able by structural features. Jordi (70, 71) suggests that specialized races of U. Pisi occur on Lathyrus pratensis and Vicia cracca; the latter race is by some recognized as a species—Uromyces Fischeri- Eduard. Uromyces proeminens (DC.) Lév. P. and H. Sydow (151) list twenty-eight species of Uromyces on the species of Euphorbia, basing their separations largely on the studies of Tranzschel (157) in this group of rusts. Nine species are recorded as autoecious and have all four spore-forms present, the others either being short-cycled or the life history incompletely known. The structural differences between many of these species are very slight and the characters used as a basis for separation are, in many cases, quite variable. In fact Arthur (12) has grouped several of the full-cycled forms under the one species Nigredo proeminens, suggesting the occurrence of specialized races. Arthur (1, 2, 3) has obtained the following results with inoculation experiments: aecidiospores from Euphorbia nutans infected E. nutans but not E. maculata, E. marginata nor E. humistrata; aecidiospores from E. humistrata infected E. humistrata and E. nutans but not E. maculata; uredospores from EF. dentata infected E. dentata but not E. humistrata, E. nutans nor E. marginata; uredospores from E. nutans infected E. nutans but not FE. maculata. Arthur (12) suggests that a race is restricted to the section Poin- settia, a second race to the section Dichrophyllum, a third race to the prostrate species, and the fourth to the more upright species of the section Chamaesyce of the genus Euphorbia. Uromyces Scirpi (Cast.) Burr. ‘This rust has its uredo and teleuto stages on Scirpus maritimus and its aecidial stage on Glaux maritima, HMippuris vulgaris, Berula angustifolia, Daucus carota, Oenanthes aquatica, O. crocata, Pastinaca sativa and Sium latifolium in Europe. The similar rust in this country forms its uredo and teleuto stages on Scirpus americanus, S. campestris, S. fluviatilis and S. robustus, and its aecidial stage on Cicuta bulbifera, C. maculata, Glaux maritima, Oenanthes Californica and Sium cicutaefolium. A good deal of experi- mentation has been carried on by Klebahn and others to determine the host relations of the rust and various races have been separated out as distinct species. Klebahn (89, 91) distinguishes the following: REED: SPECIALIZATION OF PARASITIC FUNGI 315 1. Uromyces Pastinacae-scirpi Kleb.; aecidial stage on Pastinaca sativa. 2. Uromyces Berulae-scirpi Kleb.; aecidial stage on Berula angusti- folia. Dietel (28) was able to infect with teleutospores from a single host Sium latifolium and Hippurus vulgaris. Gymnosporangium tremelloides Hartig. There is but little indi- cation in the literature of the existence of specialized races in Gymno- sporangium. Most of the species of this genus are quite restricted in host range, often occurring on only a few species of a single genus of host. Klebahn (89) gives some evidence for a host specialization of Gymnosporangium tremelloides, the teleutospores of which are produced on Juniperus communis. On the basis of cultural tests he suggests the following: 1. Gymnosporangium Ariae-tremelloides WKleb. Aecidial stage on Sorbus aria and S. torminalts. 2. Gymnosporangium Mali-tremelloides Kleb. Aecidial stage on Pyrus malus. Ochrospora Ariae (Fuckel.) Syd. Fischer (45) stated that he was unable to infect Aruncus silvestris with uredospores of Ochrospora ariae from Sorbus aucuparia. Tranzschel (152, 153), Fischer (45) and Klebahn (89) showed that Sorbus aria, S. scandica and Pyrus malus were all hosts of the same race, but Aruncus silvestris could not be infected with spores from the same source. Later, however, Fischer (48) was able to infect Sorbus aucuparia and Aruncus silvestris with the same aecidiosporic material from Anemone nemorosa. Pucciniastrum Abieti-Chamaenerii Kleb. and P. Epilobii (Pers.) Otth. These two rusts, according to Klebahn (83, 87), are dis- tinguished almost solely on the basis of their life cycle. The former is heteroecious with its aecidial stage on Abies balsamea and A. pectinata and its uredo and teleuto stages on Epilobium angustifolium, E. Dodonacum and E. latifolium. The latter has its uredo and teleuto stages on a number of other species of Epilobium and its aecidial host is as yet not known. Phragmidium disciflorum (Tode.) James. Bandi (15) suggests that a host specialization occurs in this rust. He reports one race on Rosa cinnamomea, R. pimpinellifolia and R. rubrifolia and a second race on R. canina and R. centifolia. Coleosporium Campanulae (Pers.) Lév. This rust has its aecidial stage on Pinus montana, P. rigida and P. sylvestris, and its uredo and teleuto stages on a considerable number of different species and general of the Campanulaceae. Wagner (170) and Klebahn (87, 91) have made extensive studies with reference to the specialization of the rust on different hosts. Klebahn makes the following races: 376 BROOKLYN BOTANIC GARDEN MEMOIRS 1. Campanulae-rapunculoides Kleb. on Campanula rapunculoides and, in favorable cultures, on C. bononiensis, C. glomerata, C. glomerata var. dahurica, C. lamtifolia, C. latifolia, C. nobilis, Phyteuma orbiculare and Ph. spicatum; not on Campanula rotundifolia, nor C. trachelium. . . Campanulae-trachelii Kleb. on Campanula trachelium and, in favor- able cultures, on C. bononiensis, C. glomerata, C. glomerata var. dahurica, C. latifolia var. macrantha, C. nobilis, C. rapunculoides and Wahlenbergia hederacea; not on Campanula pusilla, C. rotundifolia nor C. turbinata. 3. Campanulae-rotundifoliae Kleb. on Campanula rotundifolia and, in favorable cultures, on C. bononiensis, C. glomerata var. dahurica, C. pusilla, C. turbinata, Phyteuma orbiculare, Ph. spicatum and Wahlenbergia hederacea; not on Campanula rapunculoides nor C. trachelium. Wagner indicated the following additional races: 4. Campanulae-Phyteumatis Wagner on Phyteuma spicatum. Campanulae-macranthae Wagner on Campanula macrantha (C. latifolia var. macrantha). 6. Campanulae-patulae Wagner on Campanula rotundifolia and C. patula. Kklebahn, however, does not believe that these can be distinguished from the first three. It may also be noted that a number of plants occur as hosts for more than one race. Coleosporium Senecionis Fr. Wagner (168) first gave evidence of host specialization in this rust. Fischer (54) has also investigated the rust and the following races are indicated: I. Senecionis silvatict on Senecio silvaticus, S. viscosus and S. vulgaris (Senecionis | of Wagner). . Senecionis Fuchsii on Senecio Fuchsii and S. nemorensis (Senecionis II of Wagner). 3. Senecionis subalpini on Senecio subalpinus (Coleosporium subalpinum Wagner). 4. Senecionis doronici on Senecio doronicum. Melampsora Euphorbiae (Schub.) Cast. This rust, completing its life cycle on species of Euphorbia, has been studied by W. Miller (105, 106) who claims the existence of the following races: 1. Euphorbiae-cyparissiae W. Miiller on Euphorbia cyparissias. 2. Euphorbiae-exiguae W. Miiller on Euphorbia exigua. 3. Euphorbiae-pepli W. Miiller on Euphorbia peplus. Melampsora Euphorbiae dulcis Otth. This autoecious rust on Euphorbia is very similar to Melampsora Euphorbiae. As a result of the experimental work of W. Miiller (105, 106) and Klebahn (91) two races are indicated: bo on iS) REED: SPECIALIZATION OF PARASITIC FUNGI ohh 1. Euphorbiae-dulcis s. str. on Euphorbia dulcis. 2. Euphoriiae-strictae W. Miiller on Euphorbia stricta and E. platy- phyllos. Melampsora populina Lév. and Melampsora Tremulae Tul. A number of different species of Melampsora are recorded as having their uredo and teleuto stages on the various kinds of poplars. These rusts, however, constitute a group of closely related forms and differ but little in their structural characteristics. The main differences appear to be in the choice of hosts, especially in the aecidial stage. The relationships between the poplar rusts may best be indicated by grouping them under the above names. Melampsora populina is distinguished from M. Tremulae by the fact that the teleutospores are subcuticular while in the case of MW. Tremulae they are subepidermal. The uredo and teleuto hosts are also different species of Populus and serve further as a means of distinguishing between the two. Klebahn (87, 91) has made a special study of these rusts. Melampsora populina includes two rusts which can be distinguished only by the choice of the aecidial host. These are: 1. Melampsora Allii-populina Kleb.; aecidial stage on Allium cepa and A. ursinum; uredo and teleuto stages on Populus balsamif- era and P. nigra. 2. Melampsora Larici-populina Kleb.; aecidial stage on Larix decidua; uredo and teleuto stages on Populus balsamifera and P. nigra. Melampsora Tremulae includes four or five rusts whose uredo and teleuto stages occur on Populus alba and P. tremula, rarely on other species; the aecidial stage is found on widely separated host plants. The following belong in this group: 1. Melampsora Larici-Tremulae Kleb.; aecidial stage on Larix decidua. 2. Melampsora pinitorqua Rostr.; aecidial stage on Pinus silvestris. 3. Melampsora Rostrupii Wagner; aecidial stage on Mercurialis perennis. 4. Melampsora Magnusiana Wagner; aecidial stage on Chelidonium major. 5. Melampsora Klebahni Bubak; aecidial stage on Corydalis cava, C. digitata, C. fabacea, C. laxa and C. solida. This may not be _ distinct from the preceding one. Melampsoras of Salix. The Melampsoras on different species of Salix constitute a complex group of interrelated rusts. The structural differences between the large number of commonly recognized species are comparatively insignificant. In order to segregate the different species it is necessary to rely, to a large extent, upon differences in the choice of host. The willow rusts also afford many parallels among the poplar rusts and in some cases it is not possible to distinguish the rusts on these two genera except by the choice of host. 378 BROOKLYN BOTANIC GARDEN MEMOIRS Klebahn (87, 91) and Schneider (133-135) have been the principal investigators of the willow rusts from the standpoint of host special- ization. While many forms have been segregated out, Klebahn (92) has recently indicated the value of grouping these on the basis of structural features. The relationship of some of these different willow rusts, based upon structural characteristics and physiological special- ization, may be indicated. Melampsora Larici epitea Kleb. The aecidial stage occurs on Larix decidua and the uredo and teleuto stages on various species of Salix. Klebahn (87, 91) and Schneider (133-135) have carried out a number of inoculation tests and have distinguished the following races: a. Larici-epitea typica Kleb. on Salix aurita, S. cinerea, S. caprea, S. hippophaéfolia and S. viminalis. b. Larici-daphnoides Kleb. on Salix daphnoides. c. Larici-retusae Ed. Fischer on Salix retusa and S. herbacea. d. Larici-nigricantis Schneider on Salix nigricans, S. glabra and S. hegetschwetlert. e. Larici-purpureae Schneider on Sali purpurea. f. Larici-reticulatae Schneider on Salix reticulata and S. hastata. Klebahn states that minute structural differences can be observed Ine@."O; and: Melampsora Ribesii-purpureae Kleb. The aecidial stage occurs on species of Ribes and the uredo and teleuto stages on various species of Salix. Specialized races occur, according to Klebahn (87, 91) and Schneider (133-135), as follows: a. Ribesii-purpureae Kleb. on Salix purpurea and S. purpurea X S. viminalis; to a less extent on S. daphnoides. b. Ribesti-auritae Kleb. on Salix aurita and possibly on S. Caprea and S. cinerea. c. Ribesii-grandifoliae Schneider on Salix grandifolia and S. aurita, and possibly on S. arbuscula. Klebahn places (a) and (6) under a separate species Melampsora Ribesii-epitea Kleb. Melampsora Evonymi-Capraearum Kleb. The aecidial stage appears on species of Evonymus and the uredo and teleuto stages on various species of Salix. Schneider (133-135) distinguishes two spe- cialized races: a. Evonymi-capraearum typica Schneider on Salix aurita, S. Caprea, and S. cinerea. b. Evonymi-incanae Schneider on Salix incana and S. caprea, not on S. aurita nor S. cinerea. Melampsora Larici-pentandrae Kleb. and M. Salicis-albae Kleb. REED: SPECIALIZATION OF PARASITIC FUNGI 379 According to Klebahn (84, 87, 91), Melampsora Larici-pentandrae Kleb., with its aecidial stage on Larix decidua and uredo and teleuto stages on Salix fragilis and S. pentandra, is hardly distinguishable from Melampsora Salicis-albae Kleb., with its aecidial stage on species of Allium and its uredo and teleuto stages on Salix alba, except on the basis of host relations. Melampsora Allii-fragilis Kleb. and M. Galanthi-fragilis Kleb. According to Klebahn (84, 85, 87, 91), a similar relation exists between Melampsora Allu-fragilis Kleb., with its aecidial stage on species of Allium and uredo and teleuto stages on Salix fragilis and S. pentandra, and Melampsora Galanthi-fragilis Kleb., with its aecidial stage on Galanthus nivalis and uredo and teleuto stages on the same two species of Salix. Melampsorella Caryophyllacearum (DC.) Schroet. Bubak (24), Klebahn (87), and Fischer (45) have given some evidence for special- ization in this rust. Fischer, using the same collection of aecidio- spores, infected Stellaria media, S. graminea, Arenaria serphyllifolia. Malachium aquaticum and Cerastium sp. indet., while Cerastiwm arvense, Moehringia trinervig and M. mucosa were not infected. Kle- bahn obtained somewhat similar results. Melampsoridium betulinum (Tul.) Kleb. Klebahn (87, 91) sug- gests the occurrence of at least two specialized races of this rust which has its aecidial stage on Larix decidua and its uredo and teleuto stages on species of Betula. He found that aecidiospores from Larix, pro- duced by inoculation with teleutospores from Betula pubescens, in- fected B. pubescens and B. nana very abundantly and B. verrucosa sparingly; aecidiospores from Larix, produced by inoculation with teleutospores from Betula verrucosa, infected B. verrucosa abundantly, B. nana sparingly, and B. pubescens not at all. Accordingly the fol- lowing races are indicated: 1. Betulae-verrucosae Kleb. on Betula verrucosa and B. nana. 2. Betulae-pubescentis Kleb. on Betula pubescens, B. nana and, to a slight extent, on B. verrucosa. The above review lists the occurrence of host specialization in more than fifty different rusts. The nature of the specialization of these rusts may be indicated by arranging them in the following groups: 1. Heteroecious rusts in which the aecidial host (or hosts) is common to several specialized races which occur on the uredo and teleuto hosts: Coleosporium Campanulae (Pers.) Lév., C. Senecionis Fr., Melampsora Evonymi-Caprearum Kleb., M. Larici-epitea Kleb., M. Ribesii-purpureae Kleb., Melampsorella Caryo- phyllacearum Schroet., Melampsoridium betulinum (Tul.) Kleb., 380 BROOKLYN BOTANIC GARDEN MEMOIRS Ochropsora Ariae (Fuck.) Syd., Puccinia graminis Pers., P. Caricis (Schum.) Rebent., P. Polygoni-amphibit Pers., Uro- myces caryophyllinus (Schrank) Winter, and Uromyces Pisi (Pers.) Winter. Certain other rusts might also be looked for in this group as Puccinia coronata (Corda) Kleb., P. loliit Niels. (P. coronifera Kleb.) and P. bromina Eriks. 2. Heteroecious rusts in which the uredo and teleuto host (or hosts) is common to several races which occur on the aecidial hosts: Gymnosporangium tremelloides Hartig, Melampsora populina Lév., M. Tremulae Tul., Puccinia Bistortae (Str.) DC., P. mammillata Schroet., P. sessilis Schneid., P. silvatica Schroet., Uromyces Dactylidis Otth, U. Scirpi (Cast.) Burr. 3. Heteroecious rusts in which the specialized races are recognized by their selection of both aecidial and uredo and teleuto hosts: Puccinia Centaureae-Caricis Tranz., P. coronata Corda, P. dispersa Eriks. and Henn., P. extensicola Plowr., P. glumarum (Schm.) Eriks. and Henn., P. Ribesii-Caricis Kleb., P. Stipina Tranz., Uromyces Poae Rebent. 4. Autoecious rusts and those whose life history is incompletely known. The following belong in this group: Melampsora Euphorbiae (Schub.) Cast., M. Euphorbiae-dulcis Otth, Phrag- midium disciflorum (Tode.) James, Puccinia Absinth DC., P. bullata (Pers.) Winter., P. Carduorum Jacky, P. Centaureae Mart., P. Chaerophylli Purt., P. Epilobii-tetragoni (DC.) Winter., P. Heliantht Schw., P. Hieracit (Schum.) Mart., P. Leontodontis Jacky, P. Petroselini (DC.) Lindr., P. Pulsatillae Kalchbr., P. Ribis DC., Uromyces Fabae (Pers.) de Bary, U. proeminens (DC.) Lév. POWDERY MILDEWS—ERYSIPHACEAE The first work carried on to determine whether a host special- ization occurs among the species of the powdery mildews was that of Neger (107), the results of which were published in 1902. Since then a number of workers have contributed to the evidence for special- ized races in this well-defined group of parasites. In fact, at the pres- ent time, one or more species of five of the six genera of the Ery- siphaceae have been tested. In most cases, however, the data are very meager and it is not possible to draw any definite conclusions. In a few cases, notably for Erysiphe graminis and E. cichoracearum, the facts are better established. Erysiphe cichoracearum DC. The host relations of this species was first reported upon by Neger (107) who obtained the following ijl REED: SPECIALIZATION OF PARASITIC FUNGI 381 results: (1) conidia from Artemisia vulgaris, Lactuca muralis and Lithospermum arvense infected plants of the same species but did not infect each other nor some other plants tested; (2) conidia from Eieracium murorum infected H. murorum and Leontodon taraxacum, the latter very slightly; (3) conidia from Senecio vulgaris infected S. vulgaris and Lactuca muralis; (4) conidia from Lappa major, Plantago major, Pulmonaria officinalis and Verbascum thapsiforme failed to infect any host inoculated. Salmon (128) reports the successful infection of Plantago major and P. media, using conidia from the former; negative results were obtained with Plantago lanceolata, Eupatorium cannabinum and Galium A parine. The writer (115, 116) has carried on a very extensive series of experiments with this mildew, particularly with the cucurbit hosts. In the main, the mildew as found on the Hubbard squash (Cucurbita maxima) was used for inoculating the various plants. It was found that this mildew readily infected Cucurbita maxima (seven varieties), C. moschata (three varieties), C. pepo (seventeen varieties), C. foeti- dissima, Cucumis dipsaceus, C. melo (nine varieties), C. sativus (eight varieties), Cyclanthera explodens, Echinocystis lobata, Lagenaria vulgaris (six varieties), Momordica charantia and Sicyos angulatus. Partial infection of the following was obtained: Cuitrullus vulgaris (seven varieties), Cucumis anguria (two varieties), Ecballium elaterium, Melothria scabra and Momordica balsamina. Coccinea cordifolia, Luffa acutangula and L. Aegyptiaca proved to be entirely resistant. To a slight extent it was possible to transfer the cucurbit mildew to the sunflower (Helianthus annuus) and plantain (Plantago rugellii). Efforts to transfer it to Aster cordifolius, A. laevis and Solidago caesia failed. It was also found that a race of mildew occurred on Aster cordifolius, A. laevis and A. sagittifolius, not passing over to Cucurbita maxima nor Solidago caesia. Another race occurs on Solidago caesia, infecting this species but not passing over to asters or cucurbits. Erysiphe graminis DC. Marchal (97) was the first to demonstrate host specialization in this powdery mildew. As a result of his tests he concluded that the following races may be distinguished, although he gives us no details of his evidence: 1. Tritict upon Triticum vulgare, T. Spelta, T. polonicum and T. turgidum; not on T. durum, T. monococcum nor T. dicoccum. 2. Hordei upon Hordeum hexastichon, H. vulgare, H. trifurcatum, H. nudum, H. jubatum and H. murinum; not on H. maritimum, H. secalinum nor H. bulbosum. 3. Secalis upon Secale cereale and S. anatolicum. 4. Avenae upon Avena sativa, A. fatua, A. orientalis and Arrhenatherum elatius. 26 382 BROOKLYN BOTANIC GARDEN MEMOIRS . Poae upon Poa annua, P. trwialis, P. pratensis, P. caesia, P. mutalensis, P. nemoralis and P. serotina. . Agropyrt upon Agropyron. 7. Bromi upon various species of Bromus. Salmon (123, 128) has infected Avena nuda, A. brevis and A. sativa with conidia from A. nuda; conidia from A. sterilis infected A. pratense and A. sativa; and conidia from A. sativa infected A. sativa, A. brevis, A. nuda, A. orientalis, A. sterilis and A. strigosa. Attempts to infect twelve other grasses, belonging to different genera, with the oat mildew failed. The writer (120) has carried out an extensive series of experiments with the powdery mildew on Avena sativa. Infection occurred on the following: A. barbata, A. brevis, A. fatua, A. fatua var. glabrata, A. ludoviciana, A. nuda, A. nuda var. chinensis, A. nuda var. elegantissima A. planiculmis, A. pratensis, A. pubescens, A. purpurea, A. sativa (sixteen varieties), A. sativa orientalis (six varieties), A. sterilis, A. strigosa and A. sulcata. In most cases, in a large number of trials, one hundred percent of infection was secured. The oat mildew also infected Arrhenatherum avenaceum. Negative results were obtained with Avena bromoides and A. sempervirens, as well as grasses belonging to other genera. This race, then, extends over a wide range of species and varieties of Avena, but, with the exception of Arrhenatherum avenaceum, is restricted to this genus. Salmon (123, 132) has reported only a few results with the powdery mildew of wheat. He successfully infected Triticum vulgare and T. Spelta with conidia from the former. He also states that young seedlings of Hordeum silvaticum could be infected with the same mildew. Vavilov (164) has tested out, under field conditions, seven hundred and fifty-five ‘‘pure-lines’’ belonging to the different species and varieties of Triticum. He used pure lines belonging to thirty varieties of Triticum vulgare, seven of T. compactum, ten of T. turgidum, nine of T. Spelta, fifteen of T. durum, three of T. polonicum, five of T. dicoccum and four of T. monococcum. In general the pure lines be- longing to the different varieties of 7. vulgare, T. compactum and T. Spelta are extremely susceptible to the mildew, while the pure lines of the varieties of 7. durum, T. turgidum, T. polonicum and T. monococcum proved, in the main, to be quite resistant. The pure lines of some varieties of T. dicoccum proved to be highly susceptible, while the pure lines of other varieties were markedly resistant. Dif- ferences in the susceptibility of the pure lines were noted and certain varieties, notably 7. vulgare var. fuliginosum and T. dicoccum var. picnurum stood out as distinctly immune. However, the pure lines On ON REED: SPECIALIZATION OF PARASITIC FUNGI 383 belonging to the various varieties of all the species of Triticum, with the exception of certain ones, as those just noted, proved quite sus- ceptible when they were tested under greenhouse conditions. Vavilov suggests that the greenhouse conditions are much more favorable to the development of the powdery mildew and thus more or less immune races may be successfully attacked by the fungus. The writer (117, 118, 120) has carried on a very extensive series of tests with the powdery mildew of the wheat. One hundred and sixty-one varieties belonging to the eight recognized types or species of Triticum have been tested under greenhouse conditions, many tests having been made with nearly all of these. The results are summarized in the following table: VARIETIES OF SPECIES OR TYPES OF Triticum IN RELATION TO THE WHEAT MILDEW ] 5 ame Le | 100% infec- o-90% 50-89% | 10-49% -9% Species of type Varieties | tion infection | infection | infection | infection COMmPpachinn Are = | 2a I 2 | I | 0 . | ID ICOCCUM ms eae 24 8 3 6 g) | 4 DI OE EOTENS esters Bose ote 45 36 O 6 2 I Monococcum......... ye || fo) I 2 I I POLGHICUIT A eee TOMN | 9 I oO O fo) SyPA TIES £e Oe one ee II 8 I | 2 Ce) (o) DUGOUT ee ee a 3 2 2 o. (| (0) Vulgare?® . 52,78 35 5 6 3 3 ——— hota serrate aeciek ies 161 TOT 5 14 27 10 9 It is thus seen that the great majority of the varieties are highly susceptible to the wheat mildew. It may be specially noted that T. dicoccoides Kcke. the wild wheat of Palestine, proved to be quite susceptible. The distinctly resistant varieties included T. dicoccum var. Khapli, Russian emmer and Spring emmer and T. vulgare var. caesium and pyrothrix. Various other species of Triticum have also been tested, T. bicorne Forsk and T. caudatum Gren. & Godr. proving to be highly susceptible and T. triaristatum Gren. & Godr. and T. triuncinale Rasp. proving to be resistant. The writer (119, 120) has also found that the powdery mildew of §Summary includes Triticum Cienfuegos Lagh., T. dicoccoides Kcke., T. Frey- cenetitt Host., T. Meyert, T. Tumonia Schrad. and T. durum Desf. var. plenissimum. * Summary includes T. abyssinicum Steud. and T. dicoccum Schrank var. campy- lodon. 8 Summary includes T. Thaoudar Reut. and T. dicoccum Schrank var. cladurum. 9Summary includes T. durum Desf. var. africanum and Triticum species indet. var. Tibeian. 384 BROOKLYN BOTANIC GARDEN MEMOIRS wheat readily passes over to certain species of Aegilops which some systematists regard as a subgenus of Triticum. Aegilops aristata, Aeg. Aucheri, Aeg. cylindrica, Aeg. ligustica, Aeg. speltoides, Aeg. squarrosa, Aeg. triaristata and Aeg. ventricosa proved very susceptible, while Aeg. ovata and Aeg. triticoides gave negative results. It is evident that the powdery mildew of wheat can successfully attack a very wide range of varieties belonging to the various species of Triticum. It cannot, however, pass over onto Avena sativa, Hor- deum vulgare, Secale cereale and other grasses tested. The powdery mildew of rye has been tested out by the writer (110); Secale cereale, S. anatolicum and S. montanum were readily infected, S. dalmaticum proving resistant. This mildew will not pass over onto any other cereal, nor such grasses as Bromus mollis, Dactylis glomerata, Festuca elatior, Glyceria fluitans, Hordeum jubatum, Lolium perenne, Phleum pratense and various species of Poa. Treboux (161) also states that conidia from Secale cereale infected S. cereale but not Hordeum vulgare nor Triticum vulgare. Salmon (128) reports meager data for the quack grass mildew; conidia from Agropyron repens infected A. caninum and A. tenerum, but not A. repens, A. acutum nor A. glaucum. Salmon (128) reports the successful infection of Poa annua, P. nemoralis and P. pratensis with the mildew on the latter. The writer (110) has successfully used the mildew on P. pratensis to infect P. compressa, P. nemoralis, P. pratensis and P. trivialis. A number of other grasses tested by these workers have given negative results. The powdery mildew of barley has been tested by both Salmon (125) and the writer (117). According to the former the mildew from Hordeum vulgare is readily transferred to H. decipiens, H. distichon, H. hexastichon, H. intermedium, H. vulgare and H. zeocriton, while it is not transferable to H. jubatum, H. murinum, H. secalinum nor H. sil- vaticum; in addition H. bulbosum, H. maritimum generally gave nega- tive results, infection occurring in only a few cases. The writer has obtained positive results, using the mildew from H. vulgare, on H. distichon, H. nudum, H. Steudelii X trifurcatum, H. tetrastichon, H. trifurcatum, H. vulgare and H. Zeocriton. Young seedlings of H. nodosum also proved susceptible, although older plants gave negative results. Negative results were obtained with H. bulbosum (except that a few tufts of conidia were produced on one plant), H. maritumum and HT. jubatum. Efforts to transfer this mildew to various grasses, including the other cereals, failed. Salmon (128) reports the successful infection of Dactylis glomerata with conidia from the same host; negative results were obtained with this mildew on Agropyron repens, Lolium temulentum and the cereals. REED: SPECIALIZATION OF PARASITIC FUNGI 385 Salmon (123, 126, 127) has made extensive tests with the mildew on the brome grasses using conidia from a number of different species. Some of Salmon’s more important results are indicated in the following table: Plants Used as Hosts Source of Conidia 25 ae 25 <8 22 = § =:2 aN = = Seo RS SS | Se Re oe ke le x38 ss S8 S8 S& S$ | ss] 8 = S38 S38 Sk ANTS as aS AN Seiad FBGA ae as aS as ae} ae: | iS Qe | ia Q 2 5 ie Sis a re Sr ra ie (lege 1. Bromus interruptus..... 3/3" |48/48|10/10| 0/42| 0/29 16/16 | 0/20) 0/25 | 11/26! 2. Bromus hordeaceus var. | | | | DIOOKESCENS 2 2 a. ae | 33/33|34/34 38/47| 0/28, 27/27 | 0/13| 9/22" | 10/16 3. Bromus commutatus... . | 0/7.| 0/9 |21/21| 0/7 | 4/8! | o/7 | 6/7 6/7 4. Bromus arvensis....... | 0/9 8/8 | | 5. Bromus tectorum....... | | | 8/8 | 7/8 From his data, Salmon concluded that four, or possibly even five, specialized races exist within this genus. In looking over his data, however, there seems to be very little, if any, difference between some of them. The races on B. interruptus and B. hordeaceus differ only in their capacity for infecting B. commutatus, the mildew on B. horde- aceus infecting this host, while that on B. interruptus does not. The race on B. commutatus differs from that on B. hordeaceus in not in- fecting B. mollis and B. interruptus. The race on B. arvensis infects this same species but not B. mollis. Finally the race on B. tectorum differs from that on B. hordeaceus in being able to infect B. sterilis. It is evident that these races are not distinctly marked off from one another. But little work has been done using ascospores from various hosts. Marchal (98) mentions the following results: (1) ascospores from Hordeum vulgare infected H. vulgare, H. distichon, H. trifurcatum and H. Zeocriton, but not Avena sativa, Secale cereale nor Triticum vulgare; (2) ascospores from Secale cereale infected S. cereale, but not Hordeum vulgare nor Triticum vulgare; (3) ascospores from Triticum vulgare infected 7. vulgare but not Agropyron caninum, Avena sativa, Hordeum vulgare nor Secale cereale. Salmon (124, 132) found that ascospores from Hordeum vulgare infected H. vulgare, H. trifurcatum and H. Zeocriton, but not H. bulbosum, H. jubatum, H. maritimum, H. secali- num, Avena sativa, Secale cereale nor Triticum vulgare; ascospores from Bromus commutatus infected B. commutatus and B. hordeaceus, but not 10 The denominator indicates the number of leaves inoculated, the numerator indicates the number infected. 1! Subinfection. 386 BROOKLYN BOTANIC GARDEN MEMOIRS B. racemosus. These results correspond exactly with the infecting capacity of conidia from the same hosts. The evidence is quite conclusive for the existence of highly special- ized races in the grass mildew. For the most part these races are definitely restricted to the species of a single genus of host plants. Erysiphe Galeopsidis DC. Neger (107) tested the infecting capacity of the mildew on Galeopsis tetrahit and found that this host was infected, while negative results were obtained on Calamintha acinos, Glechoma hederacea and Stachys recta. Salmon (128) obtained positive results with the mildew from Ballota nigra on this same host but failed to infect Salvia verticillata and Leonurus cardiaca. Erysiphe Polygoni DC. Neger (107) used the mildew from the following hosts: Galium silvaticum, Heracleum spondylium, Hypericum perforatum, Ranunculus repens and Trifolium incarnatum. Positive results were obtained when the mildew was sown on plants of the same host from which it was obtained and negative results on all other hosts tested. In one case he noted a slight infection of Galium silvaticum with conidia from Ranunculus repens but this was probably a foreign infection. Salmon (123) successfully infected Piswm arvense with conidia from P. sativum. Other legumes gave negative results. Conidia from Trifolium pratense infected this host but gave negative results on seven other species of this genus as well as on species of other genera tested. Microsphaera Astragali (DC.) Trev. The only results recorded for species of this genus are those of Neger (107). He infected Astra- galus glycyphyllus and A. cicer with conidia from the former. Three other hosts gave negative results. Uncinula aceris (DC.) Sacc. and U. salicis (DC.) Winter. Neger (107) used conidia of the former species from Acer pseudoplatanus to successfully infect A. pseudoplatanus and A. campestre. Conidia of the second species from Salix purpurea infected S. purpurea and S. capreda. Phyllactinia corylea (Pers.) Karst. Neger (107) reports one test with conidia from Corylus avellana, these failing to infect the same host. Voglino (166), using conidia from Corylus, infected Corylus but not Carpinus, and conidia from Carpinus infected Carpinus but not Corylus. He further found that ascospores from Carpinus infected Carpinus but not Fagus, while ascospores from Fagus infected Fagus but not Carpinus. Sphaerotheca Humuli (DC.) Burr. Salmon (128) used conidia of this mildew from Potentilla reptans to infect P. reptans; no infection occurred on Agrimonia Eupatoria, Alchemilla arvensis, A. vulgaris, Fragaria (cult. sp.) Poterium officinale nor Spiraea ulmaria. REED: ‘SPECIALIZATION: OF PARASITIC FUNGI 387 Steiner (150) found that the mildew on Alchemilla was confined to the species of this genus. He also claimed to be able to distinguish specialized races within this genus of host plants. He found that conidia from Alchemilla pastoralis and A. flexicaulis were alike in infecting power except that conidia from the former host would not infect A. Alpigena and only slightly A. pubescens, while conidia from A. flexicaulis infected A. Alpigena slightly and A. pubescens not at all. Conidia from A. impexa would not infect A. Alpina vera nor A. nitida, while conidia from A. pastoralis at least partially infected these hosts. Steiner also found that conidia from the Vulgares section of the host genus would not produce full infection on species of the Alpinae sec- tion, although conidia from species of the latter section vigorously infected species of the former. Steiner noted marked differences in the relation of the species of Alchemilla to the mildew, dividing them into the following groups: (1) Susceptible species, as A. impexa; (2) immune species, as A. conjuncta; (3) species susceptible to the mildew from some hosts, while immune to that from others, as A. micans. Sphaerotheca Humuli (DC.) Burr. var. fuliginea (Schlecht) Sal- mon. Salmon (128) has made a few tests with conidia of this mildew from Plantago lanceolata and Taraxacum officinale. Conidia from the former infected the same host but not Plantago major nor Taraxacum officinale; conidia from Taraxacum officinale infected T. officinale but not Fragaria (cult. sp.), Plantago media nor P. lanceolata. Oidium on Euonymus japonicus., Salmon (131) reports the fol- lowing results with this mildew whose identity was not fully deter- mined; conidia from Euonymus japonicus infected E. japonicus var. aureus, var. albomarginatus, var. ovatus aureus, var. microphyllus, var. President Gunter, E. radicans var. microphyllus and var. Silver Gem, but not E. nanus, E. americanus var. angustifolius, E. chinensis, E. europaeus, E. radicans var. carrieret, Celastrus scandens, C. articulatus, C. orixa nor Prunus laurocerasus var. latifolia. ADDITIONAL FUNGI Physiological specialization has also been investigated in a few other groups of fungi by a number of different workers, but, outside of the rusts and powdery mildews, no extensive studies have been made. Synchytrium taraxaci de B. and Wor. In the Chytridiaceae, Liidi (95) has tested the infecting capacity of swarmspores of Syn- chytrium taraxact from Taraxacum officinale. He tried to infect nine- teen species of Compositae which do not belong to the subdivision Cichoraceae, but with negative results in every case. He also used 388 BROOKLYN BOTANIC GARDEN MEMOIRS twenty-one species which belong to genera of this subgroup, but was able to infect only four species of the genus Taraxacum: T. officinale, T. ceratophorum, T. palustre and T. erythrospermum. Three other species of this genus tested remained free from the fungus. In many of his experiments, Liidi kept control plants of T. officinale and these were readily infected by the swarmspores. Albugo candida (Pers.) Roussel. Eberhardt (31, 32) has made inoculation tests with this parasite. His results are as follows: 1. Conidia from Capsella Bursa-pastoris infected C. Bursa-pastoris, Arabis alpina, Iberis amara and Lepidium sativum. Conidia from Capsella Heegeri infected C. Bursa-pastoris and Lepidium sativum. 3. Conidia and oospores from Lepidium sativum infected L. sativum and Capsella Bursa-pastorts. 4. Conidia from Arabis alpina infected A. alpina, A. Halleri, A. hirsuta, A. turrita, Capsella Bursa-pastoris, Cardamine pratensis, Iberis amara, Lepidium sativum, Senebiera coronopus, but not Brassica napus, B. nigra, B. oleracea, Raphanus sativus nor Sinapis arvensis. 5. Conidia from Brassica Rapa infected B. Rapa, B. nigra, B. oleracea (var. botrytis, capitata, congyloides), Diplotaxis tenuifolia and Sinapis arvensis, but not Capsella Bursa-pastoris, Iberis amara nor Lepidium sativum. On the basis of these experiments, Eberhardt concludes that there are two specialized races of the parasite: 1. On Arabis-Capsella-Lepidium. 2. On Brassica-Diplotaxis-Sinapis. Melhus (99) has also tested the infecting capacity of Albugo candida on the radish (Raphanus. sativus). He was able to infect this host, twenty-two varieties being equally susceptible, also Raphanus caudatus, Brassica alba (white mustard) and Brassica oleracea (cabbage, fifteen varieties). In the case of the latter plants infection was less certain than for the radish. The following plants gave negative results: Brassica rapa (turnip, ten varieties), B. nigra (black mustard), B. campestris (rutabaga, three varieties), Capsella Bursa-pastoris (shep- herd’s purse), Lepidium sativum (garden cress), L. virginicum (wild pepper grass), Sisymbrium officinale, S. altissimum (hedge mustard), Iberis umbellata (candytuft), Nasturtium officinale (water cress) and Cheiranthus Cheiri (wall flower). Peronospora parasitica (Pers.) de Bary. Gaumann (59) has carried out a few experiments with this fungus and finds a high degree of host specialization. The fungus that occurs on Capsella cannot infect other Cruciferae. The same is true of the race on Brassica. tN REED: SPECIALIZATION OF PARASITIC FUNGI 389 In some cases the parasite seems to be restricted to a single species, for the fungus on Sisymbrium officinale does not infect S. sophia. On the other hand the same race occurs on Brassica oleracea and B. rapa. Taphrina aurea (Pers.) Fr. Giesenhagen (60), as a result of his work on the Exoasceae, suggests that Taphrina aurea, which infects three species of Populus, is becoming specialized into races, each of which is adapted to a single species of Populus. Claviceps purpurea (Fr.) Tul. Stager (142) has found five special- ized races in the ergot of rye, Claviceps purpurea. One race occurs on rye and also on seventeen other species of grasses; a second race occurs only on Glyceria fluitans; a third is confined to species of Lolium; a fourth to Poa annua; while the fifth is found on Brachypodium sil- vaticum and Milium effusum. Both conidia and ascospores, where tested, are limited in the same fashion. Stager did not find any such specialization in Claviceps mucro- cephala. This ergot is reported on only three grasses. Plowrightia morbosa (Schw.) Sacc. Gilbert (61) reports that Plowrightia morbosa (Schw.) Sacc. is specialized on the choke cherry (Prunus virginiana) and wild plum (Prunus americana). Ascospores, conidia, and pycnospores from the former host were inoculated into the wild plum without giving any evidence of infection. On the other hand, ascospores and conidia under like conditions, readily infected the choke cherry, giving rise to normal knots. A study of the distri- bution of the fungus on the two hosts lends confirmatory evidence as to the specialization, for in one locality the disease may be prevalent on one host while absent from the other. Rhytisma acerinum (Pers.) Fr. Miiller (104) has made a study of this parasite on various maples and concludes that it consists of several specialized races. One race Platanoides is found on Acer platanoides infecting only slightly Acer campestris and A. pseudo- platanus; a second race Campestris occurs on Acer campestris, to a slight extent on A. platanoides, but does not occur on A. pseudoplata- nus; a third race, which is given specific rank as Rhytisma pseudopla- tani, occurs only on Acer pseudoplatanus. Tubeuf (162) inoculated Acer pseudoplatanus, A. platanoides, A. campestris and A. negundo with ascospores from the first named host, infection occurring only on this one maple. Colletotrichum lindemuthianum (Sacc. and Magn.) Briosi and Cavara. Barrus (16) has studied the relation of bean varieties to the common anthracnose, Colletotrichum lindemuthianum. He tested the susceptibility of one hundred sixty-one varieties to a culture of this organism and found that, while most of the varieties were susceptible in varying degrees, a few seemed to be immune. When, however, 390 BROOKLYN BOTANIC GARDEN MEMOIRS other cultures of the organism were used to inoculate the varieties a different arrangement of susceptibility became evident. Varieties quite immune to the one strain were severely attacked by another strain. All the varieties proved quite susceptible to at least one of the strains used. This indicates the existence of distinct races of this parasite with fairly definite host limitations. Edgerton and Moreland (33) have made similar studies with cul- tures of this same fungus. Their results also indicate. differences in the infecting capacity of strains of the fungus isolated from different varieties of beans. Some beans, like the snap beans, appear to be quite susceptible to a number of different strains. Other varieties, while quite susceptible to certain strains, are resistant to strains from a different source. Edgerton and Moreland have also studied cultures of Glomerella gossypii (South.) Edg., the cotton anthracnose fungus. They do not find evidence for the existence of specialized strains in this fungus, for all the cultures isolated from different sources were able to infect a large number of cotton varieties. GENERAL DISCUSSION—BRIDGING Hosts The above review of the investigations with reference to host specialization of parasitic fungi indicates that the phenomenon is of general occurrence. The work done is particularly extensive in con- nection with the rusts and the powdery mildews, but sufficient has been accomplished in other groups to make clear the presence of specialized races. It is highly probable that the same phenomenon is of wide occur- rence among other groups of plant parasites. A large number of so-called species of the Imperfect Fungi, as Cercospora, Phyllosticta, and Septoria, may really be only specialized races of a relatively small number of forms distinct on structural grounds. In several genera of the Imperfect Fungi, as those mentioned, a very large number of species have been recorded; in fact many of them are, mainly at present at least, identified by the host upon which they grow. Cultural ex- periments may result in grouping many of these together, at the same time making clear the physiological host relations. We are not, however, to assume that host specialization is of universal occurrence. There are several cases on record where the fungus shows no evidence of the specialization of its hosts. A striking case of this sort is that of Puccinia subnitens Diet. Arthur (5) has been able to infect ten hosts, belonging to the families Chenopodiaceae, Cruciferae, and Capparidaceae, with teleutospores from Divstichlis spicata. Bethel (18), using teleutospores from the same grass, recently a REED: SPECIALIZATION OF PARASITIC FUNGI Shed has succeeded in producing aecidia on twenty-two species, belonging to fifteen genera, distributed among six different families, as follows: 1. Polygonaceae: Polygonum aviculare, P. erectum and P. ramosissi- mum. 2. Chenopodiaceae: Salsola pestifer, Chenopodium album, C. glaucum, C. lanceolatum, C. pagonum, Monolepis nuttaliiana and Kochia scoparia. Amaranthaceae: Amaranthus retroflexus and A. blitoides. Nyctaginaceae: Abronia fragrans. Cruciferae: Capsella Bursa-pastoris, Lepidium densiflorum, L. medium, Erysimum asperum, Sophia pinnata, Roripa palustris, Thaspi arvense and Sisymbrium altissimum. 6. Capparidaceae: Cleome serrulata. It is also suggested that species of Papaveraceae may be aecidial hosts for the same rust. Bock (19) has carried out rather extensive cultural tests with Puccinia Gentianae (Str.) Link. He reports no evidence for host specialization, finding that a large number of species of Gentiana were readily infected with rust from two different species. Another illustration of the same condition is reported by Camilla Popta (109) who has been able to infect a number of umbellifers with the same race of Protomyces macrosporus. The following plants were infected with the fungus from Aegopodium podograria: Cicuta virosa, Seseli montanum, Libanotis vulgaris, Palimba chalrau, Bubon gemmif- erum, Pachypleurum alpinum, Bunium virescens, Ferula_ thyrstflora, Trinia vulgaris and Athamanta cretensis. If now we compare the degree of specialization found in the different parasitic fungi we find the greatest divergences. Ina general way the specialized races may be grouped on the basis of their host range as follows: I. Specialized races restricted to certain species of a genus of hosts. Here belong the specialized races of Erysiphe graminis within the genera Bromus and Hordeum; of Puccinia dispersa within the genus Bromus; the specialized races of Phragmidium disciflorum, Puccinia Centaureae, P. Epilobii-tetragoni, P. Helianthi, P. Hieracii, P. Pulsatillae, P. Ribis, P. Ribesti-Caricis, P. Caricis-montanae, Uromyces Poae, U. proeminens and Melampsora Euphorbiae. 2. Specialized races restricted to a particular genus of host plants. A number of specialized races of this sort are known. Among the Erysiphaceae we find the specialized races of Erysiphe graminis on Agropyron, Dactylis, Poa and Secale and of Erysiphe cichoracearum on Aster and Solidago. Among the rusts we find the races of Puccinia graminis on Aira, Poa, Calamagrostis and Apera; of P. coronata on on —& Ww 392 BROOKLYN BOTANIC GARDEN MEMOIRS Glyceria, Agropyron and Bromus; most of the races of P. glumarum and P. dispersa. 3. Specialized races occurring on two or more genera but belonging to the same family. As examples, we may mention the specialized races Avenae, Secalis and Tritict of Puccinia graminis: Loli, Calama- grostis and Phalaridis of P. coronata; Secalis of P. glumarum; Orchide- arum-phalaridis of P. sessilis; the races of P. Ribesii-Caricis, P. extensicola, P. silvatica, P. Bistortae, P. mammullata, Albugo candida and Claviceps purpurea. 4. Races occurring on hosts belonging to different families as Puccinia subnitens, Uromyces Scirpi and Erystphe cichoracearum. When we compare the specialization of a parasite in relation to a particular host we also find the greatest differences. For example Puccinia gramints avenae occurs, according to Eriksson (41), on twenty species of grasses belonging to fourteen genera, Carleton (25) recording it on nineteen species belonging to fifteen genera, Jaczewski (68) on seven species belonging to six genera and Stakman and Piemeisel (149) on thirty-three species belonging to twenty-one genera. All agree that this rust occurs on a wide range of more or less unrelated hosts. On the other hand, Puccinia coronata avenae is restricted to species of Avena and possibly Arrhenatherum. The powdery mildew of oats, Erysiphe gramints avenae, is also sharply restricted to Avena, infecting to some extent Arrhenatherum. The crown rust and powdery mildew are similar in infecting a large number of species and varieties of Avena. A similar condition is found in the case of the parasites occurring on Secale cereale: Claviceps purpurea secalis occurs on eighteen species belonging to eleven genera; Puccinia graminis secalis, according to Eriksson (41) on eleven species belonging to five genera and according to Stakman and Piemeisel (149) on twenty-three species belonging to nine genera; Puccinia glumarum secalis on Secale cereale and Triticum vulgare; Erysiphe graminis secalis on two species of Secale; and Puc- cima dispersa secalis on Secale cereale. Many of the specialized races, while in the main restricted to certain hosts, yet are able to infect to a greater of less extent a number of other plants. Generally these races are distinguished by their ability to fully infect certain hosts while their development on others is weak and limited. Very good illustrations of such races are found among several of the rusts. The various races of Coleosporium Campanulae are not sharply limited to definite hosts, but the hosts of one race may also be attacked by other races. The same is true of the races of Puccinia graminis, P. coronata, P. Ribesti-Caricis, Melamp- sora Larici-epitea and Uromyces Poae. REED: SPECIALIZATION OF PARASITIC FUNGI 393 It has repeatedly been suggested by many investigators that special- ized races of parasitic fungi may extend their normal host range by passing through certain so-called “bridging hosts.” There are many cases known where a particular host plant can be infected by two or more races of a parasite. Such hosts may enable a specialized race to infect a wider range of plants. Ward (172) apparently was the first to emphasize this possibility as a result of his studies of the behavior of Puccinia dispersa on various bromes. Ward supposed that Bromus arduennensis, which is readily infected with uredospores of Puccinia dispersa from Bromus mollis of the section Serrafalcus, as well as by uredospores from B. arduen- nensts of the section Libertia, served as a means for the rust on bromes of the section Serrafalcus to pass over on to bromes of the section Libertia. The following data indicate Ward’s results as bearing on this point. . Uredospores Uredospores from Bromus from Bromus Section Libertia: arduennensis mollis MST TIESUSEITE ALCTUIFETIS EUS ace oes. Se ASG NG It fein 0.5 SNS os Be 8/7” 13/14 Bromus arduennensts var. villosus. 2.0.0.0 ce ee 10/10 1/14 Section Serrafalcus: , SP ONUUSHINLOULUSP OA Mey Sti Ay es end Bie i a esl Oe wee 1/8 119/154 BS TONUUSESCEQLINLUS Sol sa Seis aan ee Hh es 8/8 31/61 Stenobromus: LER OURDSLDETE 0 Sea OE ee ee 0/6 1/74 ES VONUUSESICNIIUSt A ASI Ay: poets hc sae ene s Mee nik Sane 0/8 4/148 An examination of the above data leads one to conclude that the rust on the two hosts, Bromus arduennensis and B. mollis, are practi- cally identical in their capacity for infecting other bromes. The rust on Bromus arduennensis does not have any wider host range than the rust on B. mollis. Both grasses appear rather to be hosts for the same strain of rust. Ward’s evidence that Bromus Krausei and B. pendulinus may serve as bridging hosts is perhaps stronger. The following data indicate the relation of these bromes to the rust on Bromus sterilis and B. mollis. Uredospores Uredospores from Bromus from Bromus Serrafalcus: sterilis mollis IEF AOL OCIS OH TIS GT ste Cees a Oe a ee ae 14/29 27/27 IBTOMUS PENAUIINUS.. oye Sc oe eee Vdd ea oe TFBS 50/50 ESUONLISHTLOLLEL ORNL S ewe a a See a a ee ee 1/25 2/26 SNOT SHIIVOLLES eae OTE Rosey SRS Ee So ce AE oi elem he 1/137 119/154 BT OMUSLUCSTILUSTS ME A, SAA ono his Sal ties Jeo einer 1/4 3/4. Stenobromus: ESP OMLUESESUCEULES A APM At hots. gente athe ke Bee eva Loe 126/146 4/148 ROLES IO Do, BORE SC OT cc Oe Or 37/60 6/53 2 The denominator of the fraction indicates the number of leaves inoculated and the numerator the number infected. 13 The denominator of the fraction indicates the number of leaves inoculated and the numerator the number infected. 394 BROOKLYN BOTANIC GARDEN MEMOIRS From this data it is evident that Bromus Krausei and B. pendulinus are susceptible to the rust on both B. mollis and B. sterilis. From the standpoint of bridging hosts, however, what we need to know is the infecting capacity of uredospores from B. Krausei and B. pendulinus, produced by inoculation with uredospores from B. mollis and B. sterilis. It may well be that B. Krauset and B. pendulinus are merely hosts for the rust on both B. mollis and B. sterilis. At least the evidence is not complete for proving that they are bridging hosts. Freeman and Johnson (57) conclude that barley is a bridging host enabling the specialized races of Puccinia graminis on wheat and rye to infect oats. They find that the rust on wheat will not infect oats, but will infect barley; the same is true of the rust on rye. When, however, the rust on barley, produced by inoculation from either wheat or rye is sown on oats, infection occurs to a very slight extent. The data upon which this conclusion is based are as follows: (1) uredo- spores from wheat to barley (26/31),4 to barley (28/42), to barley (16/16), to oats (2/54); (2) uredospores from rye to barley (23/31), to oats (1/22). As noted before, the barely rust is able to infect all four cereals, but rye and oats less completely than wheat and barley. The indications are that wheat rust and rye rust, as a consequence of growing on barley for one or more generations, are able to infect oats. It is evident, however, that the data obtained are rather meager and very much more extensive series of inoculations should be carried out. Evans (44) has crossed a wheat (Bob’s Rust Proof) resistant to rust (Puccinia graminis) with another wheat (Wol Koren) which is highly susceptible. Evans found that the hybrid, although of more vigorous growth than either parent, was much more severely attacked by rust than the susceptible parent. It was also noted that in pot cultures in the greenhouse the hybrid produced an abundance of teleutospores, which rarely occurred on either parent under the same conditions. Evans next tried to determine the infecting capacities of the rust after it had developed on the hybrid. He found that the rust from the hybrid infected the susceptible parent much more severely than the rust originally found on it. Not only that but the rust on the hybrid readily attacked the resistant parent. Accordingly it is suggested that hybrid plants may play an important part in the trans- mission of parasites from susceptible to resistant varieties by increasing the virulence of the parasite. Stakman and Piemeisel (149) record many grasses as hosts for more than one race of Puccinia graminis. In fact Bromus tectorum, 4 The denominator of the fraction indicates the number of leaves inoculated and the numerator the number infected. REED: SPECIALIZATION OF PARASITIC FUNGI 395 Hordeum vulgare, and Secale cereale are infected by all six races that they worked with. They insist, however, that these races are all distinct and that bridging hosts are not present. The grasses which harbor more than one race of rust are, of course, important in the spread of these races, even though they do not enable them to increase their usual host range. Johnson (69) reports that certain grasses enable the timothy rust to extend its normal range. He found that this rust would not directly infect Hordeum vulgare nor Triticum vulgare. However, when the timothy rust was transferred to Avena sativa, the uredospores produced on this host infected Hordeum vulgare. Further uredospores pro- duced on Festuca elatior by inoculation from timothy infected both Hordeum vulgare and Triticum vulgare. It was also found that uredo- spores from Dactylis glomerata, produced by inoculation from timothy, infected Triticum vulgare. Stakman and Jensen (145), however, find no evidence for bridging hosts in the timothy rust.. Neither Avena sativa nor Dactylis glomerata increased the host range. They also report that Hordeum vulgare is a host for the timothy rust. Stakman and Piemeisel have further extended the host range of this rust and emphasized its relation to the race Avenae. Arthur (5) suggests that Helianthus annuus may be a bridging host for various races of the sunflower rust, Puccinia Helianthi, special- ized to a narrow range of species of Helianthus. H. annuus seems to be readily infected by means of teleutospores from other sunflower hosts. Arthur (3, 4) and Kellerman (74, 75) report successful infec- tions with teleutospores from H. mollis and H. grosse-serratus; Arthur (5) further reports successful infection with teleutospores from H. laetiflorus and Kellerman (75) with teleutospores from H. tuberosus. The teleutospores from these hosts vary in their ability to infect other Helianthus species and, according to results reported, are not able to ‘infect each other except that, according to Arthur, infection of H. mollis occurred when teleutospores from H. laetiflorus were used. Neither Arthur nor Kellerman have reported positive tests with the sunflower rust found in nature on H. annuus or produced on it experi- mentally by using spores from other species. Jacky (65) in Europe reports a few tests with teleutospores from H. annuus; these were able to infect only three out of eight species tested. As yet no one has clearly shown that the rust on H. annuus has a wider range of hosts than the rust on H. mollis, H. grosse-serratus, etc. In fact, the evidence is much stronger that H. annuus is a very susceptible host to the various races of rust occurring on other species of Helianthus, if such races really exist, than that H. annuus is a bridging host. 396 BROOKLYN BOTANIC GARDEN MEMOIRS In various heteroecious rusts it has been suggested that the aecidial host may act as a bridge for races occurring on the uredo and teleuto hosts to pass over on to other species normally beyond their range of infection. In the case of Pucvinia graminis the aecidial host, Berberis vulgaris, is common to all the races specialized on different kinds of grasses. In the aecidial stage the difference between the races on the gramineous hosts might disappear and the aecidiospores produced on the barberry might have a much wider range of infection. Several investigators have published data bearing on this point. Eriksson (41) has infected the barberry using teleutospores from more than fifty different grasses. In some of these cases the aecidiospores produced were used to inoculate various gramineous hosts. Some of Eriksson’s results may be indicated in the following summary: Aecidiospores Sown on ew [ole eis wsxlSulegiky S| S| 218s Teleutospores from SE Sx eS Sis Sis 8 Rs x8 \8 avs gs s Sle sis s SS (SES SISSISS SOIESIS SIS EIS S/S SISSIES 8| alate Ibs etl vom i Skat 8] s] aS NE | A pe a (a | AVENAE | | AVECNOSSCLLUD nee an 2/2 1/I 0/2/6/2 0/2 IBTIZOU NOX ae a I/I o/1\1/I o/I Bromus arvensis .......... 1/1 |o/I | o/I Bromus brachystachys..... L/e \O/a io/1/0/1 O/I Bromus madritensts....... 1/1 \1/1 }0/1/1/1 0/1 Dactylis glomerata ........ 1/T | Festuca MYUTUS. 0... ose: 1/I o/1)1/1 o/I Festuca tenutflora......... ie «| 0/1 0/1 o/I IOClena@ SCLDGE, 32045 rey o/I 0/1 o/1\0/1 Milium effusum .......... 2/2 o/I 0/1 o/I Phalaris canariensis.......| 1/1 G/1|1/1 o/I Phleum asperum.......... 1/1 0/1/0/1 o/I Vulpia bromoides......... 1/1 |0/1\0/1 o/I SECALIS Agropyron repens......... 0/1 1/I 1/1\1/1| 0/1 Bromus secalimus:...:.... o/I 1/1 Elymus sibiricus.......... \1/I Hordeum vulgare.......... 0/2 2/2\1/1 1/2 MS ECOLCNGETCOLCH Ee te 0/2 2/2\2/2 0/2 AIRAE ATG UOLVICU Seni 1/1 POAE IPOGACOESIA Te ere o/1|1/1 WOUIGONUDT ESS ay. ee AOL o/1\1/I 1/1\0/1\0/1 TRITICI | Triticum vulgare.......... 1/3 |2/3\1/2 6/7 In general, the aecidiospores from the barberry are restricted in ’ The denominator of the fraction indicates the number of tests and the numer- ator the number that were successful. REED: SPECIALIZATION OF PARASITIC FUNGI 397 their capacity for infection in the same way as uredospores from the same gramineous hosts used as a source of the teleutospores for in- fecting the barberry. The only marked variation from this is in the case of aecidiospores from the barberry produced by inoculation with teleutospores from Bromus madritensis, Briza maxima, Festuca myurus, and Phalaris canariensis, all hosts for the race Avenae, which infected not only Avena sativa but also Secale cereale, a host of race Secalis of the rust. Jaczewski (68) found close correspondence in the infecting capacity of aecidiospores from the barberry arising from teleutospore inocu- lations from definite plants and that of the uredospores from the same gramineous hosts. In fact his establishment of the nine specialized races of the black stem rust in Russia is based as much on aecidiospore inoculations as on uredospore inoculations. Stakman (143) also found no essential differences in the infecting capacity of uredospores from wheat and Agropyron repens and that of aecidiospores from the barberry arising as a result of inoculation with teleutospores from these same hosts. Pritchard (110) also found a correspondence between the infecting capacity of uredospores and aecidiospores. Bolley and Pritchard (20) assert that aecidiospores from a single barberry hedge have been used to infect wheat, oats, barley, Hordeum jubatum, Agropyron tenerum and A. repens. ‘The origin of the infec- tion of the barberry was not known but probably was due to teleuto- spores from Hordeum jubatum. In 1905 aecidiospores from barberry readily infected barley and Hordeum jubatum, less readily wheat, and oats hardly at all. Arthur (10), however, has come to the conclusion that the “bar- berry acts as a bridging host between each and every other gramineous host.’’ The evidence that he gives in support of his statement may be indicated: (1) aecidiospores from barberry, produced by inocula- tion with teleutospores from Agrostis alba, infected wheat and barley, but not oats; (2) aecidiospores from barberry produced by inocula- tion with teleutospores from Agropyron tenerum, infected oats; (3) aecidiospores from the barberry, produced by inoculation with teleuto- spores from Sitanion longifolium, infected wheat; (4) aecidiospores from the barberry, produced by inoculation with teleutospores from Elymus canadensis, failed to infect wheat and rye. No one has recorded any data on the infecting capacity of uredospores developed on Sitanion longifolium; accordingly we have no information whether this rust is a distinct race or whether S. longifolium is merely a host for specialized race Tritici. Further, Agropyron tenerum is a host for specialized race Avenae; hence the results recorded are just what one 27 398 BROOKLYN BOTANIC GARDEN MEMOIRS would expect. Agrostis alba, however, seems to harbor a distinct race of rust and so the result recorded is not in line. Taking the results of aecidiospore inoculations as a whole, there seems to be no good reason for assuming that aecidiospores from the barberry, produced by teleutospores from a known grass, have any greater range of hosts than uredospores from the same grass. It appears that the racial strains of the black stem rust are not so sharply fixed in their host restrictions in either the uredo or aecidial stage. Further, the nature of the specialization is different in Europe from what it isin America. It is not surprising, then, that these races are able to grow on other hosts. There is, however, no clear indication that the barberry acts in any way as a bridging host and that it enables the races on different grasses to increase their range. The possibility of the aecidial host serving as a means for extending the host range of specialized races is quite apparent in Puccinia coronata Corda. Miihlethaler (102) records Rhamnus Imeretina as an aecidial host for specialized races of three of the main subgroups of the crown rust: Puccinia coronata (Corda )Kleb., P. coronifera Kleb. and P. alpinae coronata Mihlethaler. Rhamnus Purshiana is likewise an aecidial host for the races on two subgroups—P. coronata (Corda) Kleb. and P. alpinae coronata Miihlethaler. There is, however, no evidence at hand to indicate that, as a matter of fact, these species do, in any way, act as bridging hosts. There are many other cases where a particular species of plant is a host for two or more specialized races of a parasite and it might be possible for these to enable the different races to extend their host range. A few cases of this sort may be mentioned. According to Jaczewski (68) Agropyron repens and A. caninum are hosts for the races Tritici and Secalis of Puccinia graminis and this might serve to enable one race to pass over on to the hosts of the other. There is, however, no experimental proof in support of the suggestion. Stak- man and Piemeisel (149) record a number of hosts as common to several or all of the six races they studied. However, no bridging occurs, each race being distinct. According to Miihlethaler (102), Festuca elatior is a host for races Festucae and Loli of Puccinia coronata. Phalaris arundinacea is the only uredo and teleuto host for the special- ized races of Puccinia sessilis with their aecidial stages on Liliaceae, Orchidaceae, Amaryllidaceae and Araceae. Several species of Ribes are common aecidial hosts for the specialized races of Puccinia Ribesu- Caricis. Similar conditions are found among a large number of other forms—Uromyces Dactylidis, U. Poae, U. Fabae, U. Scirpi, Coleo- sporium Campanulae, Melampsora Larici-epitea, M. populina, M. Tremulae, etc. REED: SPECIALIZATION OF PARASITIC FUNGI i ie The occurrence of bridging hosts has been suggested in other groups of parasites as well as in the rusts. Salmon (126, 127), in connection with his work on the powdery mildew of the bromes, has suggested the possibility that Bromus hordeaceus may act as a bridging host for the mildew on Bromus racemosus and B. commutatus. He found that the mildew on B. racemosus failed to infect B. commutatus (0/12),'° while it readily infected B. hordeaceus (34/34). Furthermore conidia from B. commutatus failed to infect B. racemosus (0/36), while infecting B. hordeaceus (40/49). Salmon supposes that B. hordeaceus may act as a bridge for the mildew on B. racemosus to pass over to B. commutatus and also the reverse. In one case, Salmon infected B. hordeaceus with conidia from B. racemosus. The conidia produced on the former were then used to infect B. commutatus. Salmon, how- ever, did not test the infecting capacity of the conidia thus produced on B. commutatus. Steiner (150), in his work with the mildew (Sphaerotheca Humuli (DC.) Burrill) of Alchemilla, reports the occurrence of bridging hosts. He states that conidia from Alchemilla connivens and A. pubescens readily infected A. pastoralis and A. impexa but gave negative results when sown on A. micans. On the other hand, A. micans is readily infected with conidia from A. pastoralis and A. impexa. Accordingly Steiner supposes that A. pastoralis and A. impexa may carry the mil- dew over to A. micans from A. connivens and A. pubescens. Some evidence is also given to indicate that A. impexa is a bridging host between A. nitida and A. fallax. Steiner’s conclusions, however, are based on only a few tests. In the Erysiphaceae the question has been raised whether the ascospores and conidia from a particular plant possess the same in- fecting capacities. Marchal (98), Salmon (124, 132), and Voglino (166) have, in a few cases, used ascospores for inoculation tests and they report that the capacity of ascospores for infection is identical with that of conidia from the same host. It is, however, somewhat surprising that the evidence that various races of parasites may increase their range by means of bridging hosts is so vety meager, if such really occurs. In those cases where the suggestion of bridging has been most emphasized one is not impressed with the data supplied. In fact, in all such cases the races of the parasite are not sharply limited in their host range. They may infect some hosts more readily and more vigorously than others, but the virulence of the parasite does not seem to be increased or decreased by developing on congenial or uncongenial hosts. 16 The denominator of the fraction indicates the number of leaves inoculated and the numerator the number infected. 400 BROOKLYN BOTANIC GARDEN MEMOIRS The notion of bridging hosts, of course, implies that the fungus undergoes a physiological change in consequence of its new habitat and thus becomes able to attack other hosts. The change is certainly closely associated with variation in virulence so well known in the case of pathogenic bacteria. It is quite likely that fungous parasites vary in virulence or can be made to do so by suitable experimental methods. At present, however, we have no good evidence that this has been done in any particular case. The results of Evans (44), referred to above, point in this direction. In this case, however, the data are not at all extensive. The facts might be explained by an increase in susceptibility of both resistant and susceptible parent in response to a change in the environment; or external factors may have favored a more successful invasion on the part of the rust parasite. Passing reference may be made to the work of Salmon (129, 130), Ward (173), Stakman (143) and Spinks (141) which clearly indicates that a plant may be rendered more or less susceptible to fungous invasion by means of certain agencies. The work of these investi- gators shows that mineral starvation, excess of nutrients, mechanical injuries, anaesthetics, etc., modify the relations of a plant to fungous invasion. It has been pointed out by Eriksson (35), Ward (172, 174), Vavilov (164, 165) and others that a specialized race tends to occur on more or less closely related hosts. There are, however, great differences among the specialized races in this respect. As pointed out above, the host range of these races may be narrow or wide. Within a single species of parasite we may have a race occurring on many hosts belonging to different genera and another race restricted to a single genus or even certain species of a genus. Puccinia graminis, as well as other fungi, includes races of such wide differences in host range. Attempts have been made to utilize the infective capacity of a parasite to determine the genetic relationship of hosts. Eriksson (35) applied this test in determining the possible relation of a rye-wheat hybrid to the two parents. Ward (171, 172) reports a fairly close correspondence between the hosts of the more or less well-defined races of Puccinia dispersa bromi and the grouping of the bromes on other grounds. Vavilov (154, 155) has used Puccinia dispersa tritict and Erysiphe gramints tritici as a test in determining the relationship of types and varieties of Triticum. He also used Puccinia graminis avenae and P. coronata avenae as a similar test in connection with species and varieties of Avena. It is interesting to note that the parasites on wheat gave practically the same results and these are both quite narrowly specialized races. The rusts on oats, however, did not give corresponding results, Puccinia graminis avenae infecting a wider REED: SPECIALIZATION OF PARASITIC FUNGI 401 range of varieties than P. coronata avenae. ‘The former, as noted above, occurs on a number of species of grasses belonging to different genera, whereas the latter is closely restricted to the genera Avena and Ar- rhenatherum. It has been suggested that there is a connection between stability of host species and the occurrence of specialized races. Edgerton and Moreland (33) suppose that the explanation of the difference between Bean Anthracnose and Cotton Anthracnose is due to the fact that bean varieties are quite distinct and well marked, with little or no crossing and, accordingly, no intergrading forms. On the other hand, cotton varieties readily cross and thus a series of intergrading forms occurs onto which the anthracnose fungus may spread. This explana- tion, however, cannot have any very general explanation. The evi- dence indicates that the willows hybridize quite readily and yet a number of specialized races of willow rusts are recorded. Magnus (96) suggested that the existence of these specialized races may indicate an adaptation on the part of the parasite to live on particular hosts. He makes a distinction between adaptive races and biologic forms. The former term is applied to strains or races of a parasite which tend to infect certain hosts more readily than others. On the other hand, the biologic form is sharply restricted to its hosts. Dietel (30) points out that a parasite may first have attacked a wide range of hosts, gradually becoming broken up into races adapted to certain hosts and finally limited to them. It is possible that the rela- tive abundance of the hosts may have been a factor in this process. The presence or absence of hosts in a given locality may also have played a part. The question has often been raised as to whether these specialized races of fungi are constant or fixed. Montemartini (100) has recently raised the question again as a result of his studies with certain para- sites. He inclines to believe that these races are not fixed and definite and so capable of being carried from one region to another but rather that they are local adaptive forms, perhaps dependent on the vari- ability or distribution of the host plants. He suggests that they are not permanent but temporary, owing their origin to the various nutri- tive conditions afforded by different hosts. I have already called attention to the fact that some specialized races are distinguished by their ability to infect some hosts more fully than others. The races of Puccinia Ribesu-Caricis, P. Hieracu, P. graminis, Uromyces Dactylidis and Coleosporium Campanulae afford illustrations of this condition. The infection or non-infection of certain hosts is dependent to a great extent upon particularly favorable experimental conditions. It is well known that various factors do 402 BROOKLYN BOTANIC GARDEN MEMOIRS influence in a striking way the capacity of a parasite for infecting hosts. The extent of the stability of specialized races must vary greatly in different cases. Some of the forms above considered are doubtless as fixed and constant as many parasites which show structural differ- ences. Mention may be made of the races of Melampsora Tremulae as illustrating a good case of physiological species. In studying the races of Erysiphe graminis one also gets a strong impression of their constancy and definiteness and they seem as real as though separable by structural features. In other cases, however, this is not true. One is not impressed with the definiteness of races in Puccinia Helianthi, P. Hieraciu and others. These are not characterized by well-defined host limitations. Perhaps one is justified in distinguishing different types of specializa- tion as physiological species, races, strains, etc. But little data are available for comparing the specialization of the same fungus in widely separated localities. The specialization of Puccinia graminits is apparently the same in Sweden and Russia but it has taken a quite different course in North America. Treboux (159, 160) finds a different condition in Puccinia coronata in southern Russia from what Eriksson (37, 42) finds in Sweden and Miihlethaler (102) finds in Switzerland. Carleton’s (26) results in the United States also diverge widely from those of Eriksson. Arthur (11, 13) does not find evidence for specialization in Puccinia Ribesit-Caricis in this country, while Klebahn (87, 91) reports several fairly well- defined races of this rust in Europe. On the other hand, there seems to be no essential difference in the specialization of Erysiphe graminis in Europe and North America. The same sharp host limitation seems to occur in both countries. Further data are necessary before we are able to determine the relation between the specialization of parasites in different regions. In a few cases the races are also characterized by minor structural differences. Freeman and Johnson (57) and later Stakman and Piemeisel (149) have noted variations in the size and shape of the uredospores which distinguish the races of Puccinia graminis. Kle- bahn (87, 91) has noted similar differences in the spores of the various races of Puccinia Absinthi, Melampsora Larici-epitea and others. Fischer (46, 47), as well as others, has suggested that physiological specialization is a starting point for the origination of forms distinct on structural grounds. The suggestion is plausible, for it is possible to arrange a series of forms ranging from races differing in host rela- tions through all stages to others showing constant structural differ- ences. REED: SPECIALIZATION OF PARASITIC FUNGI 403 The general occurrence of specialized races of parasitic fungi makes their study particularly important. It is especially desirable to know the exact host relations of the different races as well as to deter- mine whether the races are stable and constant or whether they are capable of a change of virulence and consequently able to extend their host range. on! ~ I N iy 20. 21. 22. 23. 24. 25. 26. CA CR Cone ONC ee LITERATURE CITED . Arthur, J. C. Cultures of Uredineae in 1899. Bot. Gaz. 29: 268-276. 1900. —— Cultures of Uredineae in 1900 and Igor. Journ. Mycol. 8: 51-56. 1902. — Cultures of Uredineae in 1902. Bot. Gaz. 35: 10-23. 1903. —— Cultures of Uredineae in 1903. Journ. Mycol. 10: 8-21. 1904. — Cultures of Uredineae in 1904 Journ. Mycol. 11: 50-67. 1905. —— Cultures of Uredineae in 1905. Journ. Mycol. 12: 11-27. 1906. — Cultures of Uredineae in 1906. Journ. Mycol. 13: 189-205. 1907. — Cultures of Uredineae in 1907. Journ. Mycol. 14: 7-26. 1908. — Cultures of Uredineae in 1908. Mycologia 1: 225-256. 1909. — Cultures of Uredineae in 1909. Mycologia 2: 213-240. IgI0. . — Cultures of Uredineae in 1910. Mycologia 4: 7-33. 1912. — Uredinales. North American Flora 7 (Part 3): 161-268. 1912. — Cultures of Uredineae in I912, 1913 and 1914. Mycologia, 7: 61-89. IQI5. Cultures of Uredineae in 1915. Mycologia 8: 125-141. 1916. . Bandi, W. Beitrage zur Biologie der Uredineen (Phragmidium subcorticium, Puccinia Caricis montanae). Hedwigia 42: 118-152. 1903. . Barrus, M. F. Variation of Varieties of Beans in their Susceptibility to An- thracnose. Phytopathology I: 190-195. I9QII. . de Bary, A. Neue Untersuchungen iiber Uredineen. Zweite Mitteilung. Monatsberichten der Akademie der Wissenschaften zu Berlin (1866): 205- 215. . Bethel, E. Puccinia subnitens and its Aecial Hosts. Phytopathology 7: 92- 94. I917. . Bock, R. Beitrage zur Biologie der Uredineen. Centralbl. f. Bakt. Abt. II, 20: 564-592. 1908. Bolley, H. L. and Pritchard, F. J. Rust Problems. Facts, Observations and Theories; Possible Means of Control. N. Dak. Agr. Exp. Stat. Bull. 68: 607-676. 1906. Bubak, Fr. Uber einige Umbelliferen bewohnende Puccinien. 1. Sitzungsber. K6nigl. bbhmischen Gesell. d. Wissenschaften in Prag (1900): Nr. 26: 1-8. Ueber die Puccinien vom Typus der Puccinia Anemones—virginianae Schweinitz. Sitzungsber. d. Kénigl. bdhmischen Gesell. der Wissenschaften in Prag, Math. naturwissen. Klasse. No. 2: 1-11. Ig0l!. Einige neue oder kritische Uromyces-Arten. Sitzungsber. d. Konigl. bohmischen Gesell. der Wissenschaften in Prag (1902): I-23. Infektionsversuche mit einigen Uredineen. II. Bericht (1903). Cen- tralbl. f. Bakt. Abt. II, 12: 411-426. 1904. Carleton, M. A. Cereal Rusts of the United States. U.S. Dept. Agr. Div. Veg. Physiol. and Path. Bull. 16: 74 pp. 1899. Investigations of Rusts. Bur. Pl. Ind. Bull. 63: 32 pp. 1904. 404 49. 50. BROOKLYN BOTANIC GARDEN MEMOIRS . Dietel, P. Uber die Aecidien von Melampsora Euphorbiae dulcis Otth und Puccinia silvatica Schroet. Oesterreichische bot. Zeitschr. 39: 256-259. 1889. Uber den Generationswechsel von Uromyces lineolatus (Desm.) Schroet. Hedwigia 29: 149-152. 1890. Uber Rostpilze mit wiederholter Aecidienbildung. Flora, 81: 394-404. 1895. . — Waren die Rostpilze in friihern Zeiten plurivor? Bot. Centralbl. 79: 81-85, 113-117. 1899. . Eberhardt, A. Zur Biologie von Cystopus candidus. Centralbl. f. Bakt. Abt. II, 10: 655-656. 1904. Contribution a l'étude de Cystopus candidus. Centralbl. f. Bakt. Abt. II. 12: 235-249; 426-439; 614-631; 714-727. 1904. . Edgerton, C. W. and Moreland, C. C. Experiments on Varietal Resistance to the Bean and Cotton Anthracnose Diseases. La. Agr. Exp. Sta. Bull. 155. 24 pp. I9g16. . Eriksson, J. Uber die Specialisirung des Parasitismus bei den Getreiderost- pilzen. Ber. d. d. bot. Gesell. 12: 292-331. 1894. Ein parasitischer Pils als Index der inneren natur eines Pflanzenbastards. Bot. Notiser (1895): 251-253. ; Neue Untersuchungen iiber die Specialisirung, Verbeitung und Herkunft des Schwarzrostes (Puccinia graminis Pers.). Jahr. f. wissen. Bot. 29: 499- 524. 1896. Neue Beobachtungen iiber die Natur und das Vorkommen des Kronen- rostes. Centralbl. f. Bakt. Abt. II, 3: 291-308. 1897. A General Review of the Principal Results of Swedish Research into Grain Rust. Bot. Gaz. 25: 26-38. 1898. Etude sur le Puccinia Ribis DC. des Groseillers rouges. Rev. générale de Botanique 10: 497-506. 1808. Nouvelles études sur la rouille brune des céreales. Ann. Sci. Nat. (Ser. 8) g: 241-288. 1899. Uber die Specizialisierung des Getreideschwarzrostes in Schweden und in anderen Landem. Centralbl. f. Bakt. Abt. II, 9: 590-607, 654-658. 1902. Neue Studien iiber die Spezialisierung der grasbewohnenden Kronen- rostarten. Arkiv. for Botanik, 8 (No. 3). 26 pp. 1908. . Eriksson, J. and Henning, E. Die Getreideroste, ihre Geschichte und Natur, sowie Massregeln gegen dieselben. 1896. . Evans, J.B. P. South African Cereal Rusts, with Observations on the Problem of Breeding Rust-resistant Wheats. Journ. Agr. Sci. 4: 95-104. IQII. . Fischer, Ed. Die Uredineen der Schweiz. Beitrage zur Kryptogamenflora der Schweiz. 2 (Heft II). 1904. Die biologischen Arten der parasitischen Pilze und die Entstehung neuer Formen im Pflanzenreich. Atti della Societa elvetica di Scienze naturali adunata in Locarno. 86 Sessione: 49-62. 1904. Der Speciesbegriff bei den parasitischen Pilzen. Verhandl. der Schweiz- er. Naturforschenden Gesellsch. 88 Jahresversammlung. pp. 300-308. 1905. Beitrage zur Entwicklungsgeschichte der Uredineen. Centralbl. f. Bakt. Abt. II. 28: 139-143. 1910. Uber die Specialisation des Uromyces caryophyllinus (Schrank) Winter. (Vorlaufige Mitteilung.) Mycol. Centralbl. 1: 1, 2. 1912. Beitrage zur Biologie der Uredineen. 3. Die Specialisation des Uromyces caryophyllinus (Schrank) Winter. Mycol. Centralbl. 1: 307-313. I912. 53: REED: SPECIALIZATION OF PARASITIC FUNGI 405 Beitrage zur Biologie der Uredineen. 4. Weitere Versuche tiber die Specilisation des Uromyces caryophyllinus (Schrank) Winter. Mycol. Centralbl. 3: 145-149. I913. Beitrage zur Biologie der Uredineen. 5. Puccinia Pulsatillae Kalchbr. (Syn. Puccinia de Baryana Thiim) und Theoretisches iiber die Specialisation. Mycol. Centralbl. 3: 214-220. 1913. Mykologische Beitrage. 1. Die Wirtswahl bei der Alchimillenbewohn- enden Uromyces. Mitteilungen d. Naturforschenden Gesellschaft in Bern (1915): 214-234. I916. . —— Mykologische Beitrage. 11. Mitteilungen d. Naturforschenden Gesell- schaft in Bern (1916). 1917. . Fraser, W.P. Cultures of Heteroecious Rusts. Mycologia 4: 175-193. I912. . Freeman, E. M. Experiments on the Brown Rust of Bromes (Puccinia dis- persa). Ann. Bot. 16: 487-494. 1902. . Freeman, E. M. and Johnson, E.C. The Rusts of Grains in the United States. Bur. Pl. Ind. Bull. 216. 87 pp. IgI1. . Gassner, Gustav. Die Getreideroste und ihr Auftreten im subtropischen dstlichen Siidamerika. Centralbl. f. Bakt., Abt.'II, 44: 305-381. 1915. . Gaumann, E. Centralbl. f. Bakt. Abt. II, 45: 575-577. 1916. . Giesenhagen, K. Die Entwickelungsreihen der parasitischen Exoasceen. Flora 81: 267-361. 1895. . Gilbert, E. M. Biologic Forms of Black Knot. Phytopathology 3: 246, 247. 1913. . Hasler, A. Beitrage zur Kenntnis der Crepis- und Centaurea-Puccinien vom Typus der Puccinia Hieracii. Centralbl. f. Bakt. Abt. II, 21: 510-511. 1908. . Hitchcock, A. S. and Carleton, M. A. Second Report on Rusts of Grain. Kans. Agr. Exp. Sta. Bull. 46. 9 pp. 1894. . Jacky, E. Die Compositen-bewohnenden Puccinien vom Typus der Puccinia Hieracii und der Spezialisierung. Zeitschr. f. Pflanzenkr. 9: 193-224, 263- 295, 330-346. 1899. . —— Beitrag zur Kenntnis der Rostpilze. Centralbl. f. Bakt. Abt. II, 9: 746- 804, 841-844. 1902. . Jacob, Gina. Zur Biologie Geranium bewohnender Uredineen. (Vorlaufige Mitteilung.) Mycol. Centralbl. 3: 158, 159. I913. . —— Zur Biologie Geranium bewohnender Uredineen. Centralbl. f. Bakt. Abt. II, 44: 617-658. I915. . Jaczewski, A. von. Studien iiber das Verhalten des Schwarzrostes des Ge- treides in Russland. Zeitsch. f. Pflanzenkr. 20: 321-359. I9gI0. . Johnson, E. C. Timothy Rust in the United States. Bur. Pl. Ind. Bull. 224. 20 pp. IQgII. . Jordi, E. Culturversuche mit Papilionaceen-bewohnenden Rostpilzen. (Vor- laufige Mitteilung.) Centralbl. f. Bakt. Abt. II, 10: 777-779. 1903. Beitrage zur Kenntniss der Papilionaceen-bewohnenden Uromyces- arten. Centralbl. f. Bakt. Abt. II, 11: 763-795. 1904. . Juel, H. O. Mykologische Beitrage V. Ofversigt af k. Vetenskaps-Akad. Forhandlinger, 53: 213-224. 1896. Ein Beitrag zur Kenntnis des Uromyces Poae Rabenh. Svensk Botanisk Tidskrift 2: 169-174. 1908. . Kellerman, W. A. Uredineous Infection Experiments in 1903. Journ. Mycol. Q: 225-238. 1903. 99. 100. BROOKLYN BOTANIC GARDEN MEMOIRS . —— Uredineous Infection Experiments in 1904. Journ. Mycol. 11: 26-33. 1905. . Klebahn, H. Kulturversuche mit heteroecishen Uredineen. Zeitschr. f. Pflanzenkr. 2: 258-275, 332-343. 1892. . —— Kulturversuche mit heteroecishen Uredineen. IJ. Bericht (1893). Zeitschr. f. Pflanzenkr. 4: 7-13, 85-90, 129-139. 1894. . — Kulturversuche mit heteroecishen Rostpilzen. III. Bericht (1894). Zeitschr. f. Pflanzenkr. 5: 13-18, 69-79, 149-156. 1895. . —— Kulturversuche mit heteroecishen Rostpilzen. IV. Bericht (1895). Zeitschr. f. Pflanzenkr. 5: 257-268, 327-333. 1895. . — Kulturversuche mit heteroecischen Rostpilzen. V. Bericht (1896). Zeitschr. f. Pflanzenkr. 6: 257-270, 324-338. 1896. . — Kulturversuche mit heteroecischen Rostpilzen. VI. Bericht (1897). Zeitschr. f. Pflanzenkr. 7: 325-345, 8: 11-30. 1897-08. Kulturversuche mit heteroecischen Rostpilzen. VII. Bericht (1898). Zeitschr. f. Pflanzenkr. 9: 14-26, 88-99, 137-160. 1899. . — Kulturversuche mit Rostpilzen. VIII. Bericht (1899). Jahr. f. wissen. Bot. 34: 347-404. 1900. . — Kulturversuche mit Rostpilzen. IX. Bericht (1900). Jahr. f. wissen. Bot. 35: 660-710. Igotl. . —— Kulturversuche mit Rostpilzen. X. Bericht (1901). Zeitschr. f. Pilanzenkr 1207-44) 132-151. 1902! Kulturversuche mit Rostpilzen. XI. Bericht (1902). Jahrbuch d. Hamburgischen wissenschaftlichen Anstalten 20: I-56. 1903. . —— Drie wirtswechselnden Rostpilze. 1904. Kulturversuche mit Rostpilzen. XII. Bericht (1903 und 1904). Zeitschr. f. Pflanzenkr. 15: 65-108. 1905. — Kulturversuche mit Rostpilzen. XIII. Bericht (1905 und _ 1906). Zeitschr. f. Pflanzenkr. 17: 129-157. 1907. Kulturversuche mit Rostpilzen. XIV. Bericht (1907-1911). Zeit- schr. f. Pflanzenkr. 22: 321-350. 1912. . — Uredineen. Kryptogamenflora der Mark Brandenburg 5a: 69-904. 1gi4. . —— Aufgaben und Ergebnisse biologischer Pilzforschung. Vortrage aus dem Gesamtgebiet der Botanik, Heft I. 41 pp. 1914. . Krieg, W. Experimentelle Untersuchungen iiber Ranunculus-Arten bewohn- ende Uromyces. Centralbl. f. Bakt. Abt. II, 19: 697-714, 771-788. 1907. Neue Infektionsversuche mit Uromyces Dactylidis. Centralbl. f. Bakt. Abt. II, 25: 430-436. 1909. . Liidi, R. Beitrage zur Kenntniss der Chytridiaceen. Hedwigia 40: I-44. 1901. . Magnus, P. Die systematische Unterscheidung nachst verwandter para- sitischer Pilze auf Grund ihres verschieden biologischen Verhaltens. Hed- Wigia 33: 362-366. 1894. . Marchal, E. De la specialisation du parasitisme chez l’Erysiphe graminis. Compt. Rend. 135: 210-212. 1902. De la specialisation du parasitisme chez l’Erysiphe graminis. Compt. Rend. 136: 1280, 1281. 1903. Melhus, I. E. Experiments on Spore Germination and Infection in Certain Species of Oomycetes. Wis. Agr. Exp. Sta. Res. Bull. 15: 25-91. Ig9I1I. Montemartini, L. Sopra la specializzazione dei funghi parassiti, con particolare riguardo alla specializzazione delle ruggini dei cereali. Rivista di Patalogia Vegetale, year 8: 33-44, 145-158. 1916. Rr Iol. 102. 103. 104. 105. 106. 107. 108. 109. 110. 1111 iG Dp. Tse II4. E15; REED: SPECIALIZATION OF PARASITIC FUNGI 407 Miihlethaler, F. Infektionsversuche mit Kronenrosten. Centralbl. f. Bakt. Abt. II, 26: 58. i910. —— Infektionsversuche mit Rhamnus befallenden Kronenrosten. Centralbl. f. Bakt. Abt. II, 30: 386-419. Ig11. Miiller, F. Beitrage zur Kenntniss der Grasroste. Beihefte zum Bot. Cen- tralbl. 10: 181-212. Igor. Miiller, K. Zur Biologie der Schwarzfleckenkrankheit der Ahornbaume, hervorgerufen durch den Pilz Rhytisma acerinum. Centralbl. f. Bakt. Abt. II, 36: 67-98. 1912. Miiller, W. Versuche mit Uredineen auf Euphorbien und Hypericum. Cen- tralbl. f. Bakt. Abt. II, 17: 210, 211. 1906. — Zur Kenntnis der Euphorbia-bewohnenden Melampsoren. Centralbl. f. Bakt. Abt. II, 19: 441-460, 544-563. 1907. Neger, F. W. Beitrage zur Biologie der Erysipheen. Flora 90: 221-272. 1902. Plowright, C. B. A Monograph of the British Uredineae and Ustilagineae. 1889. Popta, Miss C. M. L. Beitrage zur Kenntnis der Hemiasci. Flora 86: 1-46. 1899. Pritchard, F. J. A Preliminary Report on the Yearly Origin and Dissemina- tion of Puccinia graminis. Bot. Gaz, 52: 169-192. IQII. Probst, R. Versuche mit Kompositen-bewohnenden Puccinien. (Vorlaufige Mitteilung.) Centralbl. f. Bakt. Abt. II, 19: 543, 544. 1907. — Infektionsversuche mit Kompositen-bewohnenden Puccinien. Annales Mycol. 6: 289-300. 1908. Die Spezialisation der Puccinia Hieracii. Centralbl. f. Bakt. Abt. II, 22: 676-720. 1909. Reed, George M. Infection Experiments with Erysiphe graminis DC. Trans. Wis. Acad. Sciences, Arts and Letters 15: 135-162. 1906. Infection Experiments with the Mildew on Cucurbits, Erysiphe cichor- 1907. Infection Experiments with Erysiphe cichoracearum DC. Bull. Univ. Wisc. (Science Series) 3: 337-416. 1908. The Mildews of the Cereals. Bull. Torrey Bot. Club 36: 353-398. 1909. Infection Experiments with the Powdery Mildew of Wheat. Phyto- pathology 2: 81-87. 1912. The Powdery Mildews—Erysiphaceae. Trans. Amer. Micro. Soc. 32: 219-258. I913. The Powdery Mildews of Avena and Triticum. Mo. Agr. Exp. Sta. Res. Bull. 23. 19 pp. 1916. . Rostrup, E. Mykologiske Meddelelser, IV. Botanisk Tidsskrift 18: 36-47. 1894. Biologiske Arter og Racer. Botanisk Tidsskrift 20: 116-125. 1896. . Salmon, E. S. On Specialization of Parasitism in the Erysiphaceae. Beihefte zum Bot. Centralbl. 14: 261-315. 1903. Infection Powers of Ascospores in Erysiphaceae. Journ. of Botany 41: 159-165, 204-212. 1903. Cultural Experiments with the Barley Mildew, Erysiphe graminis DC. Annales Mycol. 2: 70-99. 1904. 408 126. 127. 128. 129. 130. 13. 132) 133. 134. 135. 136. a7 138. 140. BROOKLYN BOTANIC GARDEN MEMOIRS On Erysiphe graminis DC. and its Adaptive Parasitism within the Genus Bromus. Annales Mycol. 2: 255-267, 307-343. 1904. Recent Researches on the Specialization of Parasitism in the Ery- siphaceae. The New Phytologist 3: 55-60. 1904. On Specialization of Parasitism in the Erysiphaceae, II. The New Phytologist 3: I109-I2I. 1904. — Cultural Experiments with “ Biologic Forms” of the Erysiphaceae. Phil. Trans. Royal Society of London B. 197: 107-122. 1904. Further Cultural Experiments with ‘“‘ Biologic Forms” of the Ery- siphaceae. Ann. Bot. 19: 125-148. 1905. Cultural Experiments with an Oidium on Euonymus japonicus Linn. Annales Mycol. 3: I-15. 1905. — On Specialization of Parasitism in the Erysiphaceae, III. Annales Mycol. 3: 172-184. 1905. Schneider, O. Versuche mit schweizerischen Weidenmelampsoren. Centralbl. f. Bakt. Abt. II, 13: 222-224. 1904. Weitere Versuche mit schweizerischen Weidenmelampsoren. Cen- tralbl i. BaktAbt, Lis) 232-234.) 1005; Experimentelle Untersuchungen iiber schweizerische Weidenrostpilze. Centralbl. f. Bakt. Abt. II, 16: 74-93, 159-176, 192. 1906. Schroeter, J. Entwicklungsgeschichte einiger Rostpilze II, III. Cohn’s Beitrage zur Biologie der Pflanzen 3: 51-93. 1879. Die Pilze Schlesien Uredinei. Kryptogamen-Flora vy. Schlesien. 3: 291-381. 1889. Zur Entwicklungsgeschichte der Uredineen. Jahrsber. d. Schles. Ges. f. vaterl. Cult. (Bot. Sect.) 71: 31, 32. 1893. ’ . Semadeni, O. Kulturversuche mit Umbelliferen bewohnenden Rostpilzen. Centralbl. f. Bakt. Abt. II, 10: 522-524. 1903. Beitrage zur Kenntnis der Umbelliferen bewohnenden Puccinien. Cen- tralbl. f. Bakt. Abt. II, 13: 73-81, 214-221, 338-352, 439-448, 527-543. 1904. . Spinks, G. T. Factors Affecting Susceptibility to Disease in Plants. Journ. Agr. Science 5: 231-247. I913. . Stager, R. Infektionsversuche mit Gramineen-bewohnenden Clavicepsarten. Bot. Zeitung 61: II1I-158. 1903. . Stakman, E. C. A Study in Cereal Rusts, Physiological Races. Minn. Agr. Exp. Sta. Bull. 138, 56 pp. 1914. . —— A Preliminary Report on the Relation of Grass Rusts to the Cereal Rust Problem. Phytopathology 4: 411. 1914. 5. Stakman, E. C. and Jensen, Louise. Infection Experiments with Timothy Rust. Journ. Agr. Res. 5: 211-216. 1915. . Stakman, E. C. and Piemeisel, F. J. Infection of Timothy by Puccinia gram- inis. Journ. Agr. Res. 6: 813-816. 1916. Biologic Forms of Puccinia graminis on Wild Grasses and Cereals. Phytopathology 6: 99, 100. 1916. . — A New Strain of Puccinia graminis. Phytopathology 7: 73. 1917. . — Biologic Forms of Puccinia graminis on Cereals and Grasses. Journ. Agr. Res. 10: 429-502. 1917. . Steiner, J. A. Die Specialization der Alchemillenbewohnenden Sphaerotheca Humuli (DC.) Burr. Centralbl. f. Bakt. Abt. II, 21: 677-726. 1908. . Sydow, P. and H. Monographia Uredinearum. 3 Volumes. 1904-1915. 2. Tranzschel, W. Versuche mit heteroecischen Rostpilzen (Vorlaufige Mit- teilung). Centralbl. f. Bakt. Abt. IJ, 11: 106. 1903. E71. 172. REED: SPECIALIZATION OF PARASITIC FUNGI 409 Neue Falle von Heterécie bei den Uredineen. Trav. du Mus. Bot. de l’Acad. Impér. d. Sci. St. Pétersbourg 2: 14-30. 1904. Beitrage zur Biologie der Uredineen. Bericht iiber die im Jahre 1904 ausgefiihrten Kulturversuche. Trav. du Mus. Bot. de l’Acad. Impér. d. Sci. St. Pétersbourg 2: 64-80. 1904. Beitrage zur Biologie der Uredineen. III. Trav. du Mus. Bot. d. l’Acad. Impér. d. Sci. St. Pétersbourg 7: I-19. 1909. Ueber einige Aecidien mit gelbbrauner Sporenmembran. Trav. du Mus. Bot. d. l’Acad. Impér. d. Sci. St. Pétersbourg 7: I1I-116. I9gI0. Die auf der Gattung Euphorbia auftretenden autécischen Uromyces- Arten. Annales Mycol. 8: 1-35. IgIo. Culturversuche mit Uredineen in den Jahren 1911-1913. (Vorlaufige Mitteilung.) Mycol. Centralbl..4: 70, 71. 1914. . Treboux, O. Infektionsversuche mit parasitischen Pilzen, III. Annales Mycol. 10: 557-563. 1912. . —— Infektionsversuche mit parasitischen Pilzen, IV. Annales Mycol. 12: 480-483. I914. . —— Uberwinterung vermittels Mycels bei einigen parasitischen Pilzen. Mycol. Centralbl. 5: 120-126. I9g14. . Tubeuf, F. K. von. Infektionsversuche mit Uredineen der Weisstanne. Centralbl. f. Bakt. Abt. II, 9: 241. 1902. Rassenbildung bei Ahorn-Rhytisma. Naturwiss. Zeitschr. Forst.- und Landwirtschr. 11: 21-24. 1913. . Vavilov, N. I. Beitrage zur Frage iiber die verschiedene Widerstandsfahigkeit der Getreide gegen parasitische Pilze. Arbeiten d. Versuchsstation fiir Pflanzenziichtung am Moskauer landwirtsch. Institut, 1 Folge, I-110. 1913. Immunity to Fungous Diseases as a Physiological Test in Genetics and Systematics, Exemplified in Cereals. Jour. of Genetics, 4: 49-65. 1914. . Voglino, P. Contribuzione alla studio della Phyllactinia corylea. Nuovo Giornale Botanico Italiano 12: 313-327. 1905. . Wagner, G. Culturversuche mit Puccinia silvatica Schroeter auf Carex brizoides L. Hedwigia 34: 228-231. 1895. Beitrage zur Kenntnis der Coleosporien und der Blasenroste den Kiefern (Pinus silvestris L. und P. montana Mill.). Zeitschr. f. Pflanzenkr. 6: 9-13. 1896. Beitrage zur Kenntniss der Puccinia silvatica Schroeter und der Puccinia sessilis Schneider. Ber. d. d. Bot. Gesell. 14: 212-215. 1896. Beitrage zur Kenntniss der Coleosporien und der Blasenroste der Kiefern (Pinus silvestris L. and Pinus montana Mill.) III. Zeitschr. f. Pflanzenkr. 8: 257-262. 1898. Ward, H. M. On the Relations between Host and Parasite in the Bromes and their Brown Rust, Puccinia dispersa Erikss. Ann. Bot. 16: 233-315. 1902. Further Observations on the Brown Rust of the Bromes, Puccinia dispersa (Erikss.) and its Adaptive Parasitism. Annales Mycol. 1: 132-151. 1903. Experiments on the Effect of Mineral Starvation on the Parasitism of the Uredine Fungus Puccinia dispersa on Species of Bromus. Proc. Roy. Soc. (London) 71: 138-151. 1902. Recent Researches on the Parasitism of Fungi. Ann. Bot. 19: 1-54. 1905. RELATION OF MARL PONDS AND PEAT BOGS W. W. ROWLEE Cornell University The filled-in lakes and ponds of western New York are of two distinct types, the bogs often called peat or cranberry bogs, and the marl ponds. ‘These ponds are alike in that both occupy depressions in the terrain and both are filled with water from springs at their bottom or near their shores. They also resemble each other in that both are subject to filling in with material produced by organic life in and around them. They differ from each other however in the character of the water they contain, in the flora which inhabits the water and the adjacent shore and the method by which they are filled in. The glaciation of the country left a terrain with potholes and other depressions particularly favorable to peat-bog formation. The material with which peat bogs are filled consists mainly of sphagnum and heath-like plants always much disintegrated and accumulated principally around the shores. Peat bogs have long been a subject of interest. Mitchill in 1798 studied them and set forth their general structural characteristics in a paragraph of his report as commissioner for the Agricultural Society of New York, as follows: “As the peat is formed, layer over layer, in the course of successive vegetations, it can be easily explained how trunks of trees, fossil wood, and bodies and bones of animals came to be buried so deep below the present surface; because at the same time when the trees fell, and animals died, in the places where they are now found, they were upon the top, and, by the perpetual growth of the plants around, they have in many places, become covered to a great depth.” He was particularly impressed with the bones of extinct species of animals found in bogs in Orange County and other parts of eastern New York. Dachnowski in 1912 gives a very comprehensive dis- cussion and classification of the distribution of species in the several areas on the surface of a bog. The flora now found on the peat bogs corresponds to the flora of colder climates. The flora of the marl ponds corresponds more closely to that of the seashore in the same or more southerly latitudes. Marl ponds are filled in not only near the shores but in all parts of the pond where the water is not too deep. The water in the marl ponds is 410 ROWLEE: RELATION OF MARL PONDS AND PEAT BOGS 411 decidedly hard, that is, it is impregnated with lime while the water of peat bogs is soft, that is, it is not alkaline in its reaction. In a series of careful studies Davis has shown that marl is com- posed mainly of the remains of the alga Chara. Chava thrives in hard water and its cell walls are impregnated with calcium carbonate. In many marl ponds a complete transition from the living Chara to characteristic marl can be seen. Since Chara grows submerged and the principal bog plants grow emerged it is evident why there is filling at the shore in the one and in all parts of the pond in the other. In sounding many peat bogs in western New York, the writer was somewhat surprised to find many of them underlayed with marl. Fic. 1. Marl bog, northeast side of Lowry’s Pond, West Junius, Seneca Co., N. Y. Species of sedges the dominant vegetation form. The assumption had been that the alkaline or nonalkaline character of the water originally filling the depressions determined whether bog vegetation or marl pond vegetation would develop in it. Is it possible that a pond might be alkaline during one stage of its existence and then become non-alkaline in a later stage? An affirmative conclusion seems inevitable. It is Dachnowski’s view that there are changes in the vertical or historical succession in the bogs, for he says: “While working on the ecology of ravines near Ann Arbor, Michigan, I became convinced that the reactions of plants on their habitat were equally as great and profound, in some cases, as the influence of edaphic and climatic 412 BROOKLYN BOTANIC GARDEN MEMOIRS factors. In various places the decomposed remains of an earlier vegetation led to mechanical and chemical changes in the soil, the extent of which was more effective toward breaking up the flora into a heterogeneous formation, accompanied by a frequent replacement of one dominating group by another.”’ There are some bogs without evidence of marl at the bottom. Davis reports all the many peat bogs examined by him in Maine as resting on sand, clay or rock bottom, none on marl. Most of the peat bogs in the Adirondack region of New York have no marl at the bottom. These presumably were from the first supplied by springs of non-alkaline water. Some of the peat bogs of central and western Fic. 2. Peat bog near McLean, Tompkins Co., N. Y. Chamaedaphne, Andro- meda, Ledum and other heaths together with Sphagnum form the dominant vegeta- tion. 4 New York have great masses of marl under them. Such a one is near Peterboro in Madison County and another on Gorham Creek in On- tario County. Here it is apparent that some agency changed the composition of the water to such an extent that oxylophytes found conditions congenial. The most extensive marl ponds in the region are in the vicinity of the limestone belt of central New York. They occur at West Junius in Seneca County and southwest of Rochester, especially at Bergen in Genessee County. Isolated and much smaller marl ponds ROWLEE: RELATION OF MARL PONDS AND PEAT BOGS 413 occur at Tully and near Cortland. The limestone outcrop in these regions accounts for a continuous supply of water impregnated with lime. The peat bogs with the greatest thickness of marl under them occur not far from this same limestone belt, while the peat bogs with little or no marl are usually farthest from the limestone outcrop. It does not seem at first thought as though Chara could be the agency causing the radical change in the history of the vegetation of the pond and the accumulations in these depressions. In so far as vegetation is concerned calcium carbonate is relatively insoluble. Fic. 3. Transition bog near Cortland, N. Y. A thick bed of marl is over- laid with about four feet of fibrous peat. Marl was excavated from the hole in the foreground. Sedges are prominent in the vegetation now covering the surface of the bog. Not so, however, is the lime in spring water. The water with available (more or less) free lime is what Chara takes in and in its life processes converts into calcium carbonates secreted in its walls. That an im- mense amount of lime is converted is shown by the bulk of marl in the ponds. Where the amount of the lime in the original soil was not large, rain water constantly tended to wash it out and in the course of time the lime content of the water would be decreased. The ponds were artesian pools fed by these springs and as the character of the water changed there was, if our theory is correct, a corresponding 28 414 BROOKLYN BOTANIC GARDEN MEMOIRS change in the flora. There would be then two phases in the life of our bogs. rst. The marl ponds in which lime-loving plants predominated and especially Chara and which are filled with and often surrounded by beds of marl or “bog lime.”’ 2nd. Peat bogs which from the bottom up are composed of non- alkine peat which in all their history have been inhabited by oxylo- phytes. Between these two phases gradations of all degrees occur in the peat bogs of western New York. The succession might, of course, be in the other direction if the calciferous phase should become pre- dominant over a previous carboniferous one. But such a case is unknown in western New York. PATHOLOGICAL PROBLEMS IN THE DISTRIBUTION OF PERISHABLE PLANT PRODUCTS' Cy. L.- SHEAR Bureau of Plant Industry, U. S. Department of Agriculture INTRODUCTION Under our present social and economic conditions, public interest is being aroused and directed to questions and problems which have been largely overlooked or neglected in less strenuous times. The conservation of our natural resources, especially our food products, greatly to our discredit as a nation, has, until very recently, been too largely neglected. Max O’Rell is said to have made the statement some years ago that Europe could be fed on what America wastes. This statement is probably somewhat exaggerated, but unfortunately has too many facts to support it. At present, these questions, so far as food products are concerned, are of vital importance, not merely figuratively but literally. We wish to call attention here to loss and waste occurring in connection with the distribution of fruits and vegetables. A large proportion of the fruit and truck crops grown never reach the consumer. Part of this loss occurs on the farm and in the orchard, and part in transit and distribution. Adams* in his recent work on marketing perishable farm products, asserts that at least 25 percent of the perishables which arrive at the wholesale markets is hauled to the dump pile because it is unfit for human consumption. This statement we fear is not based upon sufficient data to be accepted. It is, however, the opinion of one writer on the subject. In 1910, according to Dr. Pennington,’ the New York Board of Health condemned and destroyed over twelve million pounds of fruit and over seven million pounds of vegetables. This presumably does not represent the total loss, as a considerable amount probably escaped detection. It is easy to see that such great destruction of food 1 Published by permission of the Secretary of Agriculture. 2 Adams, Arthur B. Marketing perishable farm products. Studies in history, economics, and public law, Columbia University. 72°: 25. New York. 1916. 3 Pennington, Mary E. Proper handling of foodstuffs from farm to market. In Report of the Mayor’s Market Commission of New York City, p. 257, New York, 1913. 415 416 BROOKLYN BOTANIC GARDEN MEMOIRS products must be a serious drain on our food supply and must add materially to the cost of living. No adequate estimate can, however, be made of the enormous economic loss represented in such cases. Think of the time, labor, and investment involved in the planting, cultivating, harvesting, and hauling of these products, and of the freight or express, refrigeration and delivery charges paid! This is one of the most expensive forms of economic loss imaginable. CAUSES OF LOSSES The important problems which confront investigators, producers, carriers, and consumers, are the causes and means of prevention of these enormous losses. The producers and transportation companies have heretofore been too much inclined to look upon their share of these losses as among the natural hazards of their business. The carriers, however, are now realizing more fully than ever before, the great reduction in their income due to the payment of claims from shippers for loss resulting from decay and spoilage of products in transit. According to the report of the American Association of Refrigeration,* the total amount of claims paid by 180 railroads in 1914 for loss of perishable freight was $4,977,383.09; of this amount over one half, or $2,687,393.36 was for fruit and vegetables. This, of course, does not represent all the railroads of the country nor all the losses on the roads represented. In order to devise means of reducing or preventing this enormous destruction of food products, it is, of course, first necessary to deter- mine the causes and their relations and importance. The deteriora- tion of fruits and vegetables in transit is due chiefly to the action of parasitic or saprophytic fungi. Natural ripening processes and changes in the cell contents caused by the accumulation of respiration products or smothering, may also render the articles unfit for food. These changes are usually hastened by high temperature and lack of ventila- tion. Each kind of fruit has, of course, its own natural keeping qualities. Some kinds, like strawberries, raspberries, blackberries, and figs, soon become spoiled under optimum conditions, while others, like apples, may be kept in good condition for relatively long periods. The structure and composition of the ordinary perishable plant products and their relations to the keeping and carrying qualities of such products are fairly well known and no discussion of them will be attempted here. There are many other factors, however, involved in determining the keeping and carrying qualities of fruits and vegetables, such as ‘ Bulletin No. 2. Issued by Commission on Railway and Steamship Refrigera- tion of the American Association of Refrigeration, p. 82, June, 1916. SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 417 soil and climatic conditions under which they are grown, methods of cultivation and fertilization, nature of the variety, condition as to maturity at time of harvesting, methods and care in harvesting, grading, packing, and handling previous to shipment, and methods of loading, stowing, and bracing in the cars. Any or all these factors may be and frequently are involved in the final decay due to parasitic or saprophytic fungi occurring in the field or in transit. Hence it is of the utmost importance to obtain as complete knowledge as possible of the various organisms which attack the particular product, their life histories, the time, mode, and conditions of infection and development and also their relations to methods of handling and their temperature, moisture, and host relations. These problems are primarily pathological. Growers and shippers long ago discovered that storing fruits and vegetables at low temperature prolongs their keeping. This observa- tion finally led to the development of commercial cold storage and refrigeration methods and practices. These methods and _ practices have developed thus far largely along empirical lines. It happens that growth in most of the organisms which destroy perishable plant products is inhibited at from 33° to 36° F. Therefore, if fruit or vegetables, though infected with fungi, are placed under such tempera- ture conditions before development of these organisms is too far advanced, growth of the fungi will be temporarily suspended. In some cases, therefore, refrigeration may simply delay the destruction of the product and shift or render uncertain the responsibility for its loss which may occur before it reaches the consumer. It will appear evident, therefore, that in order to devise methods of preventing or avoiding such losses, all the factors involved in any particular case must be accurately determined as well as their relations and relative importance. Because it is known that certain fungi destroy certain fruits and vegetables and that these fungi occur in the orchard or on the farm, it has been inferred by some that the presence of such organisms on decayed products at destination is sufficient evidence that the responsibility for the loss rests with the grower. This may be true in the case of some particular product affected with some particular disease when shipped without refrigera- tion. In the case of refrigerated products, however, our experience and that of others has shown that in order to determine the real cause or causes and the responsibility for loss in any specific case, the whole history of picking, packing, handling and treatment of the product must be known, or at least its history from the field to destination. This has been very strikingly brought out in the investigations of 418 BROOKLYN BOTANIC GARDEN MEMOIRS citrus fruits and also in recent investigations of raspberry, strawberry, and cranberry losses. The practice of refrigeration of fruits and vegetables in transit is for two purposes, viz.: to retard the natural ripening processes which continue after the crop is harvested and to prevent the development of destructive fungi which are assumed to be present and are likely to develop unless a constant low temperature is maintained. In many cases it is practically impossible to eliminate the organisms which cause decay and all the handling in such cases must be with the presumption of their presence and the possibility of their rapid develop- ment under favorable conditions. Each product and each fungus has its own peculiarities and reac- tions under various conditions and treatment. This may be illustrated by citing a few specific cases. STEM—END Rot AND ANTHRACNOSE OF WATERMELON Meier® has given an account of a decay of watermelons in the field and in transit, caused by a species of Diplodia. It has been found that this organism is a wound parasite, and infection takes place through the stem end of a melon after it has been cut from the vine. A practical method of preventing this infection by the appli- cation of a fungicide before shipment has been found to be the simplest means of preventing decay from this cause. The anthracnose of watermelons can also be largely controlled by proper field treatment. LEAK OF POTATOES This trouble, which is most prevalent on the Pacific coast, has been found by Hawkins® to be due chiefly to Pythium debaryanum. It has been shown that infection occurs in the field and through wounds only; hence, the most practical means of prevention is to avoid as much as possible injury in digging and handling and to sort out all wounded potatoes before shipping. This is a case in which trans- portation methods and facilities are not the controlling factor in deter- mining the condition of the product upon its arrival in the market, but are of minor importance. Potatoes, watermelons, and similar products which are not usually shipped under refrigeration can fortunately be more or less satis- factorily insured against loss by proper treatment previous to ship- ment. The temperature, ventilation and handling of such products ® Meier, F. C. Watermelon stem-end rot. Journal of Agricultural Research 6: 149-152. Ap. 24, 1916. ® Hawkins, Lon A. The disease of potatoes known as “‘Leak.’’ In Jour. Agri. Res, 6: 627-640. 1 fig. pl. XV. 10916. SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 419 en route, however, have considerable influence upon their condition at destination, even when all practical field treatments and precautions have been taken. Citrus FRUITS In the case of citrus fruits, it has been found that where the chief cause of decay is Penicillium, one of the important factors in its control is to avoid, as far as possible, all injury to the fruit in picking and packing, as the fungus enters only through wounds. It is also necessary to ship this fruit under proper refrigeration in order to insure its arrival in good condition in distant markets. While proper care in picking, packing, and handling are of primary importance in determining the keeping qualities of these fruits, proper refrigeration and prompt delivery are also essential to prevent loss from this and other organisms. CRANBERRY Rots Cranberries under proper conditions of cultivation and handling possess excellent shipping and keeping qualities and as they are mostly distributed during cool weather do not require refrigeration. Spray- ing to prevent fungous diseases in the field, careful picking and hand- ling with temporary storage in cool ventilated houses and packing in proper packages will ordinarily insure their reaching market in good condition with the usual means of transportation. Most of the losses occur before shipment and much loss of fruit held for late shipment is due to the natural ripening processes of the fruit, the action of the respiration products and smothering. Proper venti- lation would prevent the latter. RASPBERRY ROTS Ramsey’ reports the results of studies of shipments of raspberries from the Pacific coast under various conditions. The decay was attributed to Botrytis and Penicillium. He found that care in handling and prompt cooling were among the most important factors in suc- cessful shipment, but that maintaining a uniform low temperature in transit was also essential. STRAWBERRY LEAK In the cases of strawberries which have been investigated by Stevens and Wilcox,® of the Bureau of Plant Industry, for the past 7 Ramsey, H. J. Factors governing the successful shipment of red raspberries from the Puyallup Valley. U.S. D. A. Bul. 274: 1915. 8 Stevens, N. E., and Wilcox, R. B. Rhizopus rot of strawberries in transit. USS DeAC Bulle samue4—7. 19L7- 420 BROOKLYN BOTANIC GARDEN MEMOIRS two years, it has been found that “leak,” a decay caused by Rhizopus, which is the most rapid-growing destructive organism attacking this fruit, can be controlled by proper methods of picking, handling, and shipping. Rhizopus spores seem to be practically omnipresent and it is impossible to eliminate them. All handling of strawberries must, therefore, be based upon the assumption of their presence. The fungus, however, cannot gain entrance through the uninjured tissues and does not develop seriously at a temperature below 45-50° F. Southern-grown strawberries having fair natural shipping qualities, carefully picked and handled, and not subjected to too high tempera- tures before shipment, can with proper refrigeration and transportation be delivered in northern markets in good condition. If a carload of strawberries shows much “‘leak’”’ at destination, it may be due to delay or rough handling of cars in transit, and faulty refrigeration; or it may be due to improper treatment by the grower or shipper. Only a full knowledge of all the facts in any particular case can deter- mine the exact cause or causes and responsibility. The presence of the fungus on the fruit at destination is not sufficient to throw the blame on the grower or shipper. Brown Rot oF PEACHES The brown-rot fungus, Sclerotinia cinerea, which is one of the most serious causes of decay of peaches, is very common and widely distrib- uted and is found in practically all peach orchards in humid regions. It is doubtful whether a shipment of peaches grown in such a region could be found which did not contain spores of this organism. Not- withstanding the general presence of this fungus on peaches, if the fruit is picked at the proper stage of development, and properly handled, packed and refrigerated in transit, such fruit may, and usually does, reach distant markets and the consumer in good condi- tion. The fact that a carload of peaches arrives at destination in a decayed condition and the brown-rot fungus is present, does not necessarily indicate that the grower is to blame for the loss. Mr. J. A. Ruddick,’ Canadian Dairy and Cold Storage Commis- sioner, states that Canadian peaches from the Niagara district are successfully shipped from Canada to Liverpool and London, the time in transit to London being twelve days, and also that in 1910 twenty-three thousand cases were shipped from Cape Town, South Africa, to London, arriving in good condition. Seventeen days was the minimum time in transit. Other shipments from the same place ® Ruddick, J. A. Cold storage for apples and other fruit. Evidence of Mr. J. A. Ruddick before the Select Standing Committee on Agriculture and Colonization, IQIO-II, pp. 106-109. Ottawa, I9QII. SHEAR: DISTRIBUTION OF PERISHABLE PLANT PRODUCTS 421 were made to Canada by way of New York, also arriving in good con- dition and selling at one shilling each. This indicates some of the possibilities of shipping such a perishable fruit as the peach when properly handled and treated, even though “inherently liable to deterioration and decay.” Stevens,’ citing the fact that peaches become infected with the brown-rot fungus only in the orchard or before shipment, says: ‘‘We may be sure that if infected at destination, shipment was also infected at the starting point. It appears clear to the writer that in both these cases (Sclerotinia libertiana on lettuce being the other case mentioned) the responsibility rests with the shipper just as much as it would if a consignment of horses infected with glanders but not yet showing the disease was placed upon the cars.”’ We have been unable to discover facts or data to support the statement that in such fruit responsibility for losses in transit rests entirely with the shipper. Investigations have shown that the brown- rot fungus makes little or no growth at a temperature of 32—35° F. (o-2° C.),!! and that if the fruit is kept at this temperature brown rot does not develop. Of course, all practical field treatment to reduce infection should be practiced. As a result of the above quoted and similar statements some railroad representatives have taken the position that the presence in a shipment of spoiled fruit or vegetables at destination of destructive fungi known to originate in the field, is sufficient to justify the con- clusion that the carrier is free from any responsibility for the loss. Such a general conclusion as this is not in accord with the facts and is fraught with great possibilities of injustice. Every effort should be made to correct this mistake. In the past the transportation companies have undoubtedly paid many unjust claims. Now there seems to be danger of the pendulum swinging to the other extreme, resulting in the rejection of just claims. It is possible for transportation companies to prevent any just claims for losses due to destructive fungi by furnishing proper cars and refrigeration service and delivering the products on schedule time. With the recent improvement of refrigerator cars a sufficiently uniform low temperature throughout the load can be maintained to avoid the trouble which so frequently occurs of having variations in temperature, of 20 degrees or more, between the top and bottom of the car, as reported by Ramsey.” Under such conditions decay 10 Stevens, F. L. Some problems of plant pathology in reference to transporta- tion. Phytopathology 5: 108. Ap. 1915. 11 Brooks, Chas. and Cooley, J. S. Temperature relations of apple-rot fungi. Jour. Agr. Res. 8: p. 163, Jan. 1917. PA RAISeY A) ble ies aiGs 422 BROOKLYN BOTANIC GARDEN MEMOIRS and loss in some products are sure to occur, no matter how great care has been given them before shipment. The grower and the shipper, however, are subject to many hazards and conditions, some of which are beyond their control. Chief among these are climatic factors. There are many things, however, which can be done by the producer to prevent or reduce losses of this kind. All practical means of prevention should be utilized and as soon as the cause of the trouble in any specific case is determined, every reasonable and practicable effort should be made by the pro- ducer or shipper to remove such cause. On the other hand, the carrier should modify and improve his equipment and methods when neces- sary to insure the delivery of perishable products in a sound condition. The various cases described above show something of the variety and complexity of the problems involved and the need of thorough investigations to discover the causes and remedies in each case. The numerous factors involved in the case of any two products or diseases are frequently not the same and when they happen to be the same are not of equal importance. Most of them are primarily pathological or have very direct pathological bearings. Where fungi are concerned, as in most cases, full knowledge must be obtained of their host relations, time, mode and conditions of infection, tempera- ture and moisture relations and the effect on their development of various methods of treatment of the fruit or vegetable during its production, harvesting, packing, handling and transportation. The present agencies interested in and at present studying these problems are the pathologists, pomologists, horticulturists, refrigera- tion engineers, the specialists in markets and marketing, the railway freight claims associations, and the commercial inspection services. The most complete cooperation of all these agencies is necessary 1n order to solve these important problems most quickly and thus reduce as far as possible this great economic loss. EXPLANATION OF PLATES IX-XI 1 PLatE IX. Two flasks of Missionary strawberries kept two days at ordinary room temperature. a. Containing fruit in natural condition free from wounds. b. Containing same quantity of fruit inoculated with Rhizopus. PLateE X. Three wounded strawberries above; three sound berries below. All were sown with spores of Rhizopus. Photographed after two days at ordinary room temperature. The three wounded berries entirely destroyed, the three others sound. PLATE XI. Pile of spoiled cranberries discarded in sorting and screening. Loss chiefly due to fungous disease and to the effect of ripening processes and their pro- ducts or smothering. BROOKLYN BOTANIC GARDEN MEMOIRS. , VOLUME I, PLATE IX SHEAR: STRAWBERRY LEAK BROOKLYN BOTANIC GARDEN MEwoirs. VOLUME I, PLATE X. SHEAR: STRAWBERRY LEAK “cs. ; ian Pa ‘ 7 a =A #1 teed Cars VoLUME I, PLATE XI. BROOKLYN BOTANIC GARDEN MEMOIRS. #) ute sh y bah seh i ite Fai 4 Pe ya tition, Here iy . CRANBERRY ROTS SHEAR: TUBERS WITHIN TUBERS OF SOLANUM TUBEROSUM F. C. STEWART New York State Agricultural Experiment Station, Geneva At the New York Agricultural Experiment Station, in 1915, several bushels of seed potatoes not needed for the spring planting were left over summer in a cellar. The potatoes were of the variety Sir Walter Raleigh. They were stored in slatted crates which were piled one above another three crates deep in a single row along the cellar wall. The cellar was cool, moderately damp and dimly lighted. Its floor and walls were of cement. Fic. 1. A new tuber protruding from a slit in the side of an old seed tuber of Solanum tuberosum. Nat. size. No attention was given the potatoes until the latter part of Sep- tember. It was then observed that instead of producing sprouts in the usual manner they had formed large numbers of new tubers. Some of the new tubers were in sessile clusters of several small tubers 423 424 BROOKLYN BOTANIC GARDEN MEMOIRS each while others were borne singly on sprouts one to three centi- meters long and were of considerable size. The formation of new tubers directly from old ones in this manner is so common as to attract little attention. The unusual features of the present case were: (1) The large size of the new tubers. Many of them had a weight of 25-30 grams, several of 50-60 grams, and one weighed 67 grams. (2) The formation of new tubers within old Fic. 2. A new tuber protruding from a slit at the bud end of an old seed tuber. Nat. size. (Compare Fig. 3.) ones. Fifteen of the old tubers had large new tubers protruding from slits in their sides (Figs. 1-2). In one instance a new tuber weighing about 28 grams was wholly included within the parent tuber. This, like most of the protruding new tubers, was considerably flattened by the pressure to which it had been subjected during its growth. This phenomenon of large new tubers within old ones was a sight calculated to excite wonder in the beholder. To the mycologist it suggested the bursting of the volva in the egg stage of the phalloids. eal STEWART: TUBERS OF SOLANUM TUBEROSUM 42 To one familiar with Gager’s interesting paper! on ingrowing sprouts of potato tubers it appeared probable that tubers forming on ingrowing sprouts had enlarged until the pressure produced became sufficient to rupture the tissues of the parent tuber. Upon dissection of the tubers this was found to be true. Fic. 3. The specimen shown in Fig. 2 with one side cut away to show the origin of the ingrowing sprout which bears the new tuber. A slight change in pose has brought to view a second new tuber. Nat. size. The old tuber shown in Fig. 2 was carefully dissected to determine the origin of the sprout bearing the new tuber which was emerging from a slit at the bud end. It was found to have started from the ‘“eye’’ on the right side of the old tuber where the two external, sessile new tubers are seen. Fig. 3 shows the same tuber, in a slightly different position, with the tissue cut away so as to expose the new tuber and the sprout bearing it. In accomplishing this one of the 1Gager, C. Stuart. Ingrowing sprouts of Solanum tuberosum, Bot. Gaz. 54: 515-524. I912. 426 BROOKLYN BOTANIC GARDEN MEMOIRS small external tubers was removed. The changed pose in Fig. 3 brings to view a second new tuber not shown in Fig. 2. The larger of the new tubers was borne on a very short branch three centimeters from the point of origin of the sprout; and the smaller one on a similar branch about a centimeter beyond. The course of the ingrowing sprout was perpendicular to the surface of the parent tuber at the point of origin. Apparently, the direction of growth had been inward from the beginning. Close observation of this and some other specimens revealed nothing to indicate that the sprouts had started externally and turned inward. Whether it was the tip of the sprout or the expanding tubers which first broke through the cortex cannot be determined in this case; but in other specimens (among them the one shown in Fig. 1) tt was clear that the new tuber had been responsible for the rupture of the cortex. The ingrowing sprouts exhibited the lenticel-like openings observed by Gager. These signify nothing except that the sprouts were formed in a humid atmosphere. The fibrous roots observed by Gager were lacking and the sprouts were but slightly branched. The strange behavior of these tubers cannot be ascribed to low vitality. This is shown by the fact that a large number of tubers from the same lot were planted and a good stand of vigorous plants obtained. In the main, these observations agree with those made by Gager and add nothing to them except to show that tubers of considerable size may form on ingrowing sprouts and produce a striking freak of nature. The internal tubers observed by Gager were small ones. THE DUPLICATION OF A LEAF-LOBE FACTOR IN THE SHEPHERD’S-PURSE! GEORGE HARRISON SHULL Princeton University In two previous papers (Shull, 1911, 1914) I have demonstrated the existence of two independent Mendelian factors in the shepherd’s- purse (Bursa bursa-pastoris), each. of which produces the triangular form of capsule. In the latter paper I discussed at some length some of the criteria and the significance of such ‘‘duplicate’’ factors. I gave also a practically complete list of the relevant literature which had appeared before 1914, and called attention to certain miscon- ceptions which had found expression in a number of the papers cited. It is not necessary, therefore, in presenting a new case of dupli- cation of factors in this species, to repeat at any length the discussion in this earlier paper. It is important however to direct attention to the discussions there presented, since several papers along similar lines, or on closely related matters, which have appeared more recently, do not include a reference to my paper, even when from the terminology used it is evident that the authors have had it before them. Several writers are now making the desired distinction between “duplicate”’ and “‘plural’’ factors, and it is to be hoped that in the future, in the interest of precision and accuracy, all those who discuss size-inheritance and related phenomena, will abandon the expression ‘“‘multiple”’ factors because of its erroneous implications. It has been shown (Shull, 1909, 1910, 1911) that the form of leaf in shepherd’s-purse is controlled by certain Mendelian genes which have been designated Aa and Bb, the presence of A resulting in an elongation of the primary lobes of the leaf, while the B gene divides the leaf to the midrib, and brings to light certain characteristic second- ary lobing. The several possible combinations of these genes give the four rosette types: AB = heteris, aB = rhomboidea, Ab = tenuts, and ab = simplex. These four forms are illustrated in Figs. 1-4. I have now studied the progenies of a considerable number of wild Bursas from places as diverse as Chile, Hawaii, Japan, China, 1 Contribution from the Station for Experimental Evolution, of the Carnegie Institution of Washington, and from the Genetical Laboratory of Princeton Uni- versity. 427 428 BROOKLYN BOTANIC GARDEN MEMOIRS (171) (172) (183) (184) Fic. 1.—heteris. Fic. 2.--rhomboidea. (301) (302) (303) (304) FIG. 3.—tenuis. Fic. 4.—simplex. Fics. 1-4. Climax leaves from eight rosettes representing the four phenotypes in pedigree No. 15406. Heteris and rhomboidea possess one or more B factors; tenuis and simplex lack them. SHULL: DUPLICATION .OF A ‘LEAF-LOBE FACTOR 429 Australia, Tasmania, India, Ceylon, South Africa, the Sahara, and from widely distributed points in Europe and North America, and find that the forms everywhere fall into one or more of these four rosette types. This does not mean, however, that with respect to leaf-form there are only four biotypes of this species in existence, for nearly every lot of material from a new locality presents minor details of lobing which lead to their easy recognition as new and distinct biotypes. In all of the earlier crosses between types respectively dominant and recessive for either of the above-mentioned character-pairs, there appeared in the F,2 close approximations to the monohybrid ratio, 3:1, or undoubted modifications of that ratio,—the modifications being due, in most cases at least, to the facts (a) that the A factor has been in some combinations not completely dominant, and ()) that both A and B require for their manifestation a certain minimum oppor- tunity in the way of favorable environment, including cultural treat- ment. These results having been well established while the situation in regard to the capsules called for further extensive investigation, a number of my cultures which concurrently involved the rosette char- acters and the capsule characters, have been grown under conditions not ideal for the development of the leaf lobes, though adequate for the determination of capsule form. For this reason my records with respect to the leaf types in certain families are of such incompleteness as to make the recorded ratios of no particular value. In nearly all cases, however, a small portion of each pedigree has been given suf- ficiently good treatment that the composition of the several families with respect to the rosettes could be inferred with small probability of error. The discovery to be detailed below, that in certain races there are two independent Mendelian factors which affect the leaf-form in identical ways, each dividing the leaf to the midrib and bringing out the secondary lobing which is seen unmodified in rhomboidea and modi- fied by the action of the A factor in the case of heteris, has revived my interest in the inheritance of the rosette characters and investigations are now in progress which I hope will give in time a full insight into the composition of the rosettes with respect to the major factors affect- ing the leaf lobes. The duplication of the gene which produces the triangular capsule has been found almost universally distributed geographically, as will be shown in detail in a later report, but the duplication of the leaf- lobe factor, B, appears to be relatively much less frequent. Before presenting the evidence of dimery in respect to the B lobes of the leaves, it will be advantageous to have before us the 29 430 BROOKLYN BOTANIC GARDEN MEMOIRS cases of monomery with respect to this character, as indicated by the occurrence of monohybrid ratios in the F2 from crosses between plants respectively having and lacking the characteristic “B’’-lobing. TABLE I Composition of Fz Families which Show Monohybrid Ratios, from Crosses Involving the Presence of the B Factor in Wild Biotypes of Shepherd’s Purse Origin New Carlisle, Ohio Chicago, Illinois Cardiff, Wales Groningen, Holland Pedigree Numbers Pr Fi Fe 040) 054 056 06130 (15361 14359 |15362 15363 040 | 0514 0693 07203 tay eee | 15367 14361 15368 15369 15324 14348 15325 115326 15327 15328 14350 | 2306 /15331 14352 |15332 15333 14353 15334 15340 14355 15341 15342 13219 |14523 heteris AB sim- plex ab 16 72 SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 431 TABLE I (continued) Pedigree Numbers | 3 | rhom- ; | ome, i * a heter idea | tenuis Ratio One Serger er ae eee | ees 13218 15373 ——= || aitels, — Tae 22550508 | X |14365 |15374 | = 48 — 8 |6.00:1 | 1376 15375 = 85 = NOM 5 -3ite oL 13222 BE a | | ‘ x 14368 | 5382 | 3) 240m, he OL® 2,501.1 1376 [15383 | 184 | 68.°)) 34) | 138° 13:50) > 1 Berlin, Germany oa | a 1322 | | 5225 a 15385 | 2 9 | 1 | 2 |3.67:1 ZED 15387 | Tn 5 6 5 13226 15394 =| LS = 355.) 3:o6tne X 14373 15395 =) | wea oe My 7I | 3.3021 1376 15396 | —| 246 | — | 80 | 3.08:1 050 DX 0688 06212) 336 71 109 20) 3.133 059 eS. 0689 06196 103 BA | a7 | TAL | 2260.2 1 056 06197 | 1177 | 315 | 344 | 79 |3-29:1 11498 8 8 2 6 E Landau, Germany x pone mere eeay p | 3 ale et : 11425 12348 13189 20 Sel On 183-29 sk 11500 13190 — 13 — Bhs aieer X 112349 |13191 =) Ls) = 2° |19:00 92/17 11429 13192 = t aELO — AZ) | 2:Aqe 1150 | a eee 13200 fi 25 | oy 14 ee I 11425 13201 59 | PG ish i ea Vicenza, Italy — 140 |15586| 43 5 9 | — |5.03:1 13239 1542C al SO = Gi Osa eon Peking, China xX 114385 |15421 ikea ake — |} 13 |2.54:1 | 1376 15422 = 25 = A O:25071 All of the cases of monomeric B lobes which I have thus far demon- strated in wild plants are shown in Table 1. 2 These families may possibly have a duplication of the B factor, but the evidence for such duplication is much less adequate than is the evidence for the occurrence’ of a 3: I ratio in one or more F2 families from these same crosses. It should be observed however, that both 15:1 and 3: 1 ratios might occur in different F, families from the same cross, since the original wild plant used in the cross might have been homozygous for one B factor and heterozygous for the other. 432 BROOKLYN BOTANIC GARDEN MEMOIRS Several of these ratios deviate widely from the 3:1 ratio, but in each such case the inclusion in the monomeric group has been based on some special consideration. In some cases one or more pedigrees from the same parentage gave a convincing approximation to the 3 : I ratio when grown under favorable conditions; in other cases small samples of the families have been grown under good conditions, and the inclusion of the particular pedigrees in one or another of the tables has been based upon the constitution of these small well-grown samples regardless of the indecisive ratios displayed by the family as a whole, when grown under conditions which tended to suppress the dominant leaf characters here under discussion. In still other cases a number of families belonging to later generations have been grown and have given full confirmation of the classification of the wild biotype from which the pedigree in question originated. Because of the pre- liminary character of the present report, it is not considered neces- sary to present in greater detail, the evidences in support of the conclusion that the families included in this table have a single B factor. It need only be stated that families whose records are inde- cisive for the particular point at issue, have been included for the sake of completeness, and to avoid the immorality of arbitrarily _ selecting for presentation those cases which are deemed to support convincingly the author’s hypothesis. Seeds of a specimen of shepherd’s-purse, received March 31, 1g11, from Tucson, Arizona, through the kindness of Dr. D. T. MacDougal have yielded a pedigree line which has given me much difficulty in the classification of the rosettes, even under the most favorable environment I could provide, owing to the fact that in this particular strain there is so strong a tendency to precocious develop- ment of the stems that the leaf characters even in the climax leaves, are frequently of the relatively undifferentiated juvenile type. It was just in this difficult material that, during several years, two facts impressed me with the probability that there were present in this strain two independent factors corresponding with the B factor of the above notation. These facts were (a) the occurrence of a rela- tively small number of tenuis (Ab) plants in two F»2 families (11413, 11414) derived from a cross between the Tucson biotype and a speci- men of B. bp. tenuis from the eastern United States. No exact count of the fenuis rosettes in these F,. families was made, but it was noted that these tenuis rosettes were sufficiently distinct from the rest of the family, that their number could probably have been deter- mined with small degree of error. Only desultory attention was being given at that time, however, to rosette characters, owing to seemingly insurmountable difficulties of classification in this biotype, and to my SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 433 interest in certain peculiarities of inflorescence and capsule characters, which will be discussed elsewhere. (b) The second fact which sug- gested the probable occurrence of duplication of the B factor in this strain was the frequent preponderance of rhomboidea-like plants in the pure-bred families. A large portion of the plants which, because of precocious development, did not reach the full typical adult con- dition, showed, nevertheless, highly developed rhomboidea characters, even when no trace of the elongated A lobes was discernible. Other plants in the same pure-bred families showed conspicuous elongation of the primary lobes, thus making certain that the A factor was present. If two B factors were present, namely B and B’, associated with only one A factor, the greater ease with which the A character, as compared with the B character, was suppressed by unfavorable conditions, would be readily comprehensible. With this clue to a possible interpretation of the rosette characters in the Tucson strain, especial note was taken of the F.2 families from crosses between the same Tucson biotype and B. Heegeri simplex, but in every case the number of individuals which fruited with rel- atively undifferentiated “juvenile’’ condition of the rosettes was so great that the F. ratios gave no fully convincing proof of the correct- ness of the hypothesis. The results of such crosses are given in Table 2. TABLE 2 The Composition of the Fz Progenies from a Cross between Bursa bursa-pastoris heteris from Tucson, Arizona, and B. Heegert simplex, grown at the Station for Experimental Evolution Fecigrec Dydmbers heteris rhomboidea | tenuzs | simplex | Ratio Pi F Pp | AB ab Ab | ab B:6 11505 13202 | 87 44 16 | 4 6:55 y-0 x 12353 13203 | 189 66 18 I2 S50 211 11425 | 13204 | 7 59 13 9 copay een 13205 18 13h 7 (6) V ALASE= I Seon oie 13206 45 2a 4 Bs jh 72h eur Ahi 354 13207 35 18 6 4 5.30: 1 aoe 13208 142 122 | 33 14 5-62 : I Wl Gtal se eeqa lho rouge const | 592 | = 364 | 97 51 | 6.46 : I Fiepected....s\.....- +. Veet ema be cape! We aan # i her 17 15.00: 1 The results in the several families were quite consistent, but the deviation from the expected 15 : 1 ratio were in all cases very con- siderable. If it is taken into account, however, that the demon- strated difficulty in genetical studies with this Tucson strain arises 434 BROOKLYN BOTANIC GARDEN MEMOIRS from the suppression of dominant characters it will be obvious that these ratios are modifications from a higher ratio (e. g., 15 : 1) rather than from the 3 : I ratio to which the empirical results in fact more closely approximate. A new attempt to test the constitution of the rosette in the shep- herd’s-purse of Tucson, was made with seeds received from Dr. Forrest Shreve on November 29, 1914. The same care was given to the culture of the families involved in this new experiment as was exercised in those recorded in Table 2. The better greenhouse facili- ties available at Princeton as compared with those at Cold Spring Harbor, where the previous cultures had been grown, made it possible to secure a closer approximation to the expected ratios, as seen in. Table 3, the deviation being the same as before 7 direction but less in amount. TABLE 3 The Composition of the F, Progenies from a Cross between Bursa bursa-pastoris heteris from Tucson, Arizona, and B. Heegert simplex grown at Princeton University BSE umber heteris \Phomibeddea | tenuis simplex | Ratio P; | F, F AB | aB | Ab a B:b ee 14387 15423 143 76 ee ee 11-53 25 1542 I 2 I | “75001 cur: Site ale 5424 Whe ae 7 ‘ if NOtAlSmrtsredeece canter te aeaie etek 157 83 13 9 LO:Ol0 BPO. apch 3s eine | 85 OF) -aleeei2 4 15.00: Fortunately for genetical studies on the rosette characters of shepherd’s-purse, such extensive suppression of characters occurs in nearly all of the other biotypes which have been under observation, only as a result of distinctly unfavorable environmental conditions. The study of a large number of these biotypes from other regions, in crosses with B. Heegert simplex, has brought to light several other cases of probable duplication of the B factor, as shown in Table 4. The cultures in Table 4 also show for the most part distinctly defective ratios, due certainly in the main to the fact that they were being grown in an extensive study of the capsule determiners, and that in consequence suitable conditions were not provided in many cases for full development of the rosette characters. Here and there a ratio closely agrees with the expected Fy, ratio, 15 : 1, the best fits being seen in certain families grown from seeds from Bremen, Germany. By chance these families from Bremen grew under more favorable conditions than many of the other cultures listed in this table and this fact doubtless explains in part why they show a closer approxima- SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 435 TABLE 4 The Composition of F, Progenies from Crosses of Bursa Heegeri simplex and Members of Wild Biotypes of B. bursa-pastoris which Probably Possessed Dimeric B lobes Pedigree Numbers ae heteris rhom- tenuts sintplex Ratio Paes Pirate F AB Aeidee Ab a Bib F 1334 | | 15337 | 112 2y7, 20 2 6.427214 Sronwgen; x 1142545) 15338 58 20 12 4 4.88 : 13 Holland | 13179 | 15339 189 109 14 EQ) sel OAysaT | 13220 15379| = 286 | — 35 SSL 7a x 14367 | 15380-}+-— — 56 — ea Ocoee t 1376 15381 42 8 4 I 10;00:2)1 | 13222 | x 14368 15384 216 86 II 7 163753 0 1376 | | | | 13225 | | 15391 60 13 16 20 2103)" 1° Berlin Pb xe MLAB 7 2n fF r5302 53 16 15 7 2 LAane | 13179 | 15393 208 38 8 10 13.67 2: 5 527) 15307 | 170 26 15 2° NER SQ cr x 14374 15398 72 25 2 I 72g | 1376 o 32-33 : 13229 | 15401 | 116 30 18 7 5.84 : 13 X | 14377 | 15402 | 162 5 2 2 AU 75 30 13179 | 15403 243 74 4 2 52.83 : 1 | 13237 | 15404 | 218 69 14 9 12.48 : I X | 14378 | 15405 189 do Wi eS 5 14.56: 1 1376 15406 | 210 Wa 15 5 14335 51 | 15407 — 151 — 17 | 8.88:1 | 15408 = 103 = 12 | 8.58:1 13232.\| 15409 — 106 — EY .|\e9i6321 Bremen x 14379 | 15410 — 282 — 18 || r5:67020 1376 | TSAco | se 167 = 12 13/02) 351 | 15412 — 262 _— 6r | az0cr 15413 a TOSS see 23e {| (So: O5e sn 13237 | 15415 -— 202 | — 27 | AeASh x 14383 15416 — 298 — 267 ites Grea 1376 15417 257 = 19 | 13.53 :1 1337 | Wales x 14357 | 15355 — 85 _ 6 ae 1338 iu | 3 These families passed the winter in the open field, and suffered considerable injury. This doubtless accounts tor their very great deviation from the 15: 1 ratio. Small samples from each of these tamilies retained in the greenhouse vie!ded ratios much nearer to 15:1. Only for this reason are these families included here 436 BROOKLYN BOTANIC GARDEN MEMOIRS tion to the expected ratios. In families No. 15405 and 15406, which were given the most careful and detailed study, there was little evi- dence of the suppression of dominant characters, except that in a few specimens it was a little difficult to be quite sure whether the A lobe was present or not and it is not improbable therefore that a few heteris plants have been erroneously included in the rhomboidea group, but this does not affect the ratios relative to the presence or absence of the B factor. On the whole, the three families derived from F, plants of pedigree 14378, showed the least marked tendency to the suppression of the dominant lobing, and these families show a close approximation to the expected ratio 45 AB :15aB:3 Ab:1ab. The close agreement with this ratio in these families, indicates not only the duplication of the B factor but also the independence of the two B factors from the A factor. While 15:1 ratios in the F, give evidence of duplication, it is highly important to carry the analysis forward at least into the F3 generation in order to secure more convincing proof that the B factor was really duplicated in the dominant parent of the original cross. Until now the only families beyond the Fz which have been grown from material in which the B factor is duplicated, have been derivatives from the earlier Tucson cultures and, as before, these families were grown primarily for the study of the capsules, and only incidental attention was given to the rosettes. The ratios in these families are also defective, therefore, but they give, nevertheless, strong support to the hypothesis that the B factor is duplicated in the Tucson plants. These F; families are brought together in Table 5. This table has been arranged into the three groups which are expected in the Fs; of a cross involving duplication of determiners. In the first section are the families which bred true to the B lobing; in the second section are those which split in the ratio 15 : I, and in the third section are those which split into 3:1. The results may be summarized as follows: 19 F; families contained neither tenuis nor simplex individuals, seeming to indicate that the I9 parents of these had at least one of the B factors homozygous; 3 families showed ratios which may be assumed to represent the 15 : 1 class, showing that the 3 parents of these had both B and B’ present in the hetero- as probably having a duplication of the B lobe. As stated in footnote to Table 1, 15: 1 and 3:1 ratios might both occur in the F: families grown from plants in a single F, family, if the wild form used in the cross happened to be heterozygous for one or the other of the B factors. Thus BBB'b’ X bbb’b’ = BbB’'d’, yielding 15 : 1, and Bbb’b’ yielding 3 : I. This situation appears to have been realized in two cases, involving F, families 14357 from Wales and 14368 from Berlin. These two crosses are included in both Tables 1 and 4. SHULL: DUPLICATION OF A LEAF-LOBE FACTOR TABLE 5 437 Composition of F; Families from Fz, Parents Having B Lobes in Crosses between B. Heegeri simpiex and a Biotype of B. bursa-pastoris heteris from Tucson, Arizona Pedigree Numbers heteris rhomboidea tenuts stmplex | Ratio F F AB aB Ab | ab B:b 2 3 14476 ee ae me -- Lo 14477 49 155 maa ris 1:0 13202 14479 I4 9 mae = 1:0 14482 22 155 = = I-20 14483 30 98 — == r0) 14487 30 3 = Fi 1:0 14488 zante | 68 = = 1:0 14489 6 21 — = LC. 13203 14490 — 80 — a 50/51 14495 a | 30 = = 1:0 14496 = 245 — — E20 14499 2 G/ — — 120 | 14503 2 74 = ae 1:0 14506 = 115 — — iste, poo | 14508 = 60 = = jee, | 14509 28 57 = — 3410) | 14511 2 44 = = LO eae | "14513 56 18 — == 1:0 14514 = 13 — —_ 1-20 13202 | 14480 ; 93 216 16 i 3 x 16.225505 13203 14492 II 12 I — 23-00) -85 13204 AG 5 76 42 5 a 4-752 14478 58 9 19 3 3:05 00 13202 14484 = 125 = 82 1.52701 14485 36 I 18 = ZOO 14500 59 104 12 27 AslOrsck 7 |\eskaso4 — 66 = Ag 2.44:1 a 14505 3 60 = | 24 2362231 14510 gI 61 36 | — AOE zygous state, 7. e., BbB’b’; while 7 families gave ratios which may be appropriately referred to the 3 : I group, indicating that the parents of these had but one of the B factors and this one heterozygous, e. g., Bbb'b’ or bbB’b’. According to theoretical expectation for the number of families given in Table 5 these three groups should occur in the ratio 4 The occurrence ot one simplex in this family is of unknown significance. It may represent an unusually defective 15:1 ratio, but the deviation is so much greater than in any other family that other possibilities seem more likely to be true. From all my experience with these pedigrees, it seems probable that this “simplex” individual is merely a rhomboidea modified by the suppression of the B lobing. There remain also the possibility of ‘“‘loss mutation”’ and of a technical error. 438 BROOKLYN BOTANIC GARDEN MEMOIRS 14 :8:8 instead of 19: 3:7. A nearer agreement might well have been expected even with this small number of families. The dis- crepancy is probably due in part to the small size of some of the fami- lies. Thus among the families which were of necessity included in the first section of the table there are six families, the largest of which had no more than 33 individuals and if these families belonged properly to the second group of the table, only 1 or 2 individuals of the recessive type were to have been expected. That some of these small families which contained no recessives, had them nevertheless potentially present may be taken as a certainty. Only 5 such families need be shifted from the first section of the table to the second section, to bring about as perfect agreement with the expected ratio as is the- oretically possible among 29 F3 families. A more positive demonstration of the duplication of the B factor may be expected from the F; families derived from the Bremen bio- types in which the suppression of the dominant rosette characters is less extensive. Further experiments involving these biotypes are in progress. DISCUSSION The discovery of a second case of duplication of determiners in Bursa bursa-pastoris owes its chief interest to the facts pointed out in one of my former papers (Shull, 1914), (a) that cases of actual duplication of genes appears to be rare, and (0) that there is some likelihood that the duplication of factors may involve ‘“‘a series of special genotypic phenomena.” It will be recalled that only in the red pericarp of wheat, yellow endosperm of maize, the occurrence of a ligule in oats, and of triangular capsules in shepherd’s-purse, was the demonstration of dimery con- sidered adequate. Several new instances have been brought to light more recently, and at least one of these must be admitted as fully demonstrated (Ikeno, 1917). This relates to the quantity and dis- tribution of chlorophyll in Plantago major, in which two seemingly identical factors, G and H, determine independently the normal full green pigmentation of the plant. Either of these two factors may be entirely wanting, without modifying in any way the self-green color, but when both are absent, the plants are conspicuously variegated with white or pale yellowish blotches and stripes, the total quantity of chlorophyll being considerably reduced. Hallqvist (1916) has found the pinnatifid leaves of Brassica Napus apparently produced independently by two factors, the recessive undivided type reappearing in the F; in only one plant in 16. As the lobed plants showed considerable variation the results in F3 will be necessary to confirm the author’s conclusion in this case. SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 439 Of cases in which two or more factors do not produce visibly identical but only more or less similar results as in the black glume color in oats (Nilsson-Ehle, 1908, 1909), there are many more in- stances. These do not represent instances of duplication at all, of course, though they may be expected to grade into cases which would be indistinguishable from duplication. Several of the eye-color and body-color factors of Drosophila appear to be of this nature, and some real duplication may also be present in this group. Some of these Drosophila characters should have been included in my former paper, but they had not been to my knowledge cited as examples of ‘‘multiple’’ factors. They have since been so treated, and with obvious propriety, by Morgan, Sturtevant, Muller and Bridges (1915). The characters specifically mentioned by these authors are (a) pink eye-color which is determined independently by factors associated respectively, one with the sex (or X) chromosome, and the other with the ‘“third”’ chromosome; and (b) dark body-color, which is independently pro- duced by two genes which have been designated “black”’ and ‘‘ebony,”’ one in the “second”? and the other in the ‘‘third’’ chromosome. Black and ebony are not identical but merely so similar that their separation is not practicable when associated in the same family. Howard and Howard (1912, 1915) have shown that velvet chaff of wheat is independently produced by two factors, Z and S, but here also the factors are clearly not duplicates of each other, for S produces short hairs and L long silky hairs, while plants containing both factors have a mixture of both types of hairs on the glumes. The same authors have found the long awns of “‘bearded”’ wheat to result from the com- bined action of two factors B and 7, each of which produces short awns in the absence of the other, but 7 produces shorter awns than B and the T awns are most conspicuous in the distal part of the spike while the B awns are more evenly distributed on the spike. In this case the action of both B and T is cumulative, the fully awned form appearing only when both B and T are homozygous, 7. e., BBTT. An exceedingly interesting case of duplication, should it stand the test of further analysis, is reported by Gates (1915) in a cross between Oenothera rubricalyx and Oe. grandiflora; for, starting with a heterozygous type supposedly monomeric with respect to the char- acteristic red pigmentation of the rubricalyx bud, he secured in the F,. two 15 : I ratios and two 3 : I ratios, in addition to one 4 : I and four 5:1 ratios. In the F; he records 4 families with a 2:1 ratio, one 3:1, two 4:1, four 15:1, and six pure rubricalyx (4. e€., 1 : 0), besides three families in which the pigmentation of all individuals was intermediate. Gates interprets the several 15:1 ratios as evidence that the R factor has become duplicated, but owing to the 440 BROOKLYN BOTANIC GARDEN MEMOIRS notorious inharmonies between the inheritance ratios in the Oenotheras and the expectation based on the usual Mendelian methods of segre- gation and recombination, one may well suspend judgment regarding this case as an instance of duplication, until it has been shown by further analysis of one of these 15:1 ratios, that the rubricalyx individuals will yield three kinds of families, characterized respec- tively by the ratios I :0, 15 : 1, and 3: 1, and that these three kinds of families are produced in approximately the ratio 7: 4:4. Unless this should be the result of the further study, the 15’: I ratio noted in several of the Fy and F; families must have been brought about by some combination of circumstances, other than the typical Mendelian distribution of two duplicate factors for the rubricalyx pigmentation. Gates discusses at some length two of the several methods by which one may reasonably suppose duplication of factors to come about. He seems to imply (Gates, 1915, p. 204) that my discussion of this subject does not adequately cover the several possibilities. He then proceeds to present two of the same possibilities as if they were original propositions of his own. These several possibilities are (a) the occurrence of independent mutations affecting in the same or closely similar manner non-homologous chromosomes; (b) the mating of non-homologous chromosomes; and (c) the transposition of parts of chromosomes by what I have called a ‘“‘sort of longitudinal crossing- over”’ (Shull, 1914, p. 139). Only the first two of these propositions are considered by Gates and he agrees with me that both of the proc- esses (a) and (b) have probably actually resulted in the duplication of factors. He thinks that repeated mutations were responsible for the duplication of red pericarp color in Nilsson-Ehle’s wheats, and that mismating of chromosomes will explain the duplication which he believes to have taken place in his Oenothera rubricalyx crosses. Upon unpublished evidence Bridges (1917, p. 454) refers to two cases of duplication in Drosophila which seem to result from essentially the longitudinal rearrangement of genotypic materials that I had in mind when suggesting the possibility of ‘longitudinal crossing-over,”’ though the details of the process as understood by him are somewhat different. He states that a section from the mid-region of one X chromosome appears to have been removed from its accustomed place or locus in that chromosome, and to have become attached to the end of the other X chromosome, its mate. The full account of this case will be awaited with interest. Accepting the validity of these several methods of duplication, one may well ask in each specific case whether circumstances make possible a judgment as to which method was probably responsible for the duplication in question. I have assumed that the complexity SHULL: DUPLICATION OF A LEAF-LOBE' FACTOR 441 of the structure of the triangular capsule of Bursa bursa-pastoris as compared with the Heegeri type of capsule, is strong evidence against the duplication of the factor for this complex character through inde- pendently repeated mutations affecting different chromosome pairs (Shull, 1914, p. 141). The character under consideration in the present paper, namely the B lobing of the rosette leaves, appears to justify the same observation. The production of leaves divided at frequent intervals by sinuses reaching to the midrib, and bearing characteristic secondary lobes and sinuses, involves the control of the number and direction of cell divisions through very long and com- plexly branched cell lineages, and it is scarcely conceivable that such specific control of these long cell lineages should be exactly duplicated by independent mutations affecting different chromosomes. It appears to me much more logical to assume that such a rearrange- ment of the genotype has taken place that the two B determiners which are allelomorphic to each other in the homozygous monomeric strains, assumed new positions, whereby they became associated with chromosomes belonging to different pairs, and thus capable of inclu- sion in the same germ cell. As these two factors, B and B’, are apparently entirely independent of each other, it may be taken for granted that they are associated with different chromosome pairs. They could become thus asso- ciated by either of the two methods, (b) or (c), but in the absence of known linkage relations, there is nothing to indicate which of these two methods has been the more probably responsible for the dupli- cation of the B factor,—whether a rearrangement of whole chromo- somes or the rearrangement of parts of chromosomes through a so- called “longitudinal crossing-over.” All these suggestions as to the origin of duplicate determiners assume the duplication to be a derivative condition; but it may also be in some cases the primitive condition from which monomeric and recessive strains may have arisen as a result of repeated “loss’’ muta- tions, as stated in my previous paper (1914, p. 137). Studies of the geographical distribution of the duplicated factors may throw some light upon the relative age of the monomeric and polymeric types, for if wild biotypes almost universally possess the duplicated factors, it may be assumed that this condition is either primitive or at least relatively old, while a much restricted and more or less definitely circumscribed range may be accepted as a criterion of relatively recent origin from the monomeric condition. In regard to the B leaf-lobe factor it will be noted by reference to the tables, that plants showing the duplication (Table 4) have been found at Tucson, Arizona, at Cardiff, Wales, at Groningen, Holland, 442 BROOKLYN BOTANIC GARDEN MEMOIRS at Bremen and Berlin, Germany, and perhaps at Peking, China, while monomeric B lobes (Table 1) have been demonstrated in strains from all these places except Tucson, Arizona. Besides these places in which the two types have been found associated together, the mono- meric condition has been found at Chicago, Illinois, at New Carlisle, Ohio, at Landau, Germany, and probably at Vicenza, Italy, in which places duplicated factors for this character have not yet been dis- covered. Excepting only Landau, Germany, these localities in which duplication of the B factor has not yet been found, have been repre- sented in my cultures by only one wild B-lobed plant from each locality. It may be merely a matter of chance that the first plant from each of these localities had but one of the B factors. It should also be noted that from the only region in which monomeric B lobes have not been found, namely at Tucson, Arizona, only two wild plants have yet been tested, a number quite too small to give any confidence in the suggested inference that no biotypes with monomeric B lobes occur at that place. It is obviously necessary to make the study of geographical distribution of these B factors much more extensive before safe conclusions may be drawn as to the primitive or derivative condition of the B lobe with respect to duplication. This is a work in which many students might lend assistance by crossing together the several wild biotypes from their own localities. SUMMARY The leaf lobes of shepherd’s-purse are controlled by Mendelian factors A, producing elongated sharp lobes, and B which divides the leaf to the midrib and brings to light certain characteristic secondary lobing. The action of these factors is easily suppressed or obscured by unfavorable environmental conditions, and the inheritance ratios are usually more or less defective on this account. In previous papers both of these characters have been reported to be monomeric, 7. @., each was found to be controlled by a single factor. It is shown in the present paper that two factors, B and B’, exist in certain strains and that these two factors produce the same char- acteristic lobing of the leaves, but are inherited independently of each other and of the factor A. The biotypes having the B factor duplicated appear to be less widely distributed than those which are monomeric with respect to the B lobes. More extensive data are needed on this point, but if the present indications are confirmed, the relatively less frequency of the dimeric condition is taken to mean that the duplication of this factor has taken place at a relatively recent date. SHULL: DUPLICATION OF A LEAF-LOBE FACTOR 443 The morphological complexity of the character produced by the B and B’ factors is believed to indicate that duplication has come about through a physical rearrangement in the genotype rather than by a repeated mutation affecting in like manner chromosomes belong- ing to distinct pairs. My acknowledgments and thanks are due to the following corre- spondents, to whom I am indebted for seeds of the wild biotypes which were used in the present study: D. T. MacDougal, F. Shreve, J. M. Shull, C. A. Shull, A. H. Trow, Tine Tammes, G. Bitter, E. Baur, L. Gross, G. Molon, and T. Z. Chang. i LITERATURE CITED Bridges, C. B. Deficiency. Genetics 2: 445-465. 1917. Gates, R. R. On Successive Duplicate Mutations. Biol. Bull. 29: 204-220. 1915. Hallqvist, Carl Ein neuer Fall von Dimerie bei Brassica Napus. Bot. Not. 1: 39-42. I916. Howard, A., and Howard, Gabrielle L. C. On the Inheritance of Some Characters in Wheat I. Mem. Dept. Agr. India 5: No. 1. 1912. — On the Inheritance of Some Characters in Wheat II. Mem. Dept. Agr. India 7: 273-285. I9I5. Ikeno, S. Variegation in Plantago. Genetics 2: 390-416. 1917. Morgan, T. H., Sturtevant, A. H., Muller, H. J.,and Bridges, C.B. The Mechanism of Mendelian Heredity. Pp. xiii + 262. New York: Henry Holt & Co. IQI5. Nilsson-Ehle, H. Einige Ergebnisse von Kreuzungen bei Hafer und Weizen. Botaniska Notiser, pp. 257-294. 1908. — Kreuzungsuntersuchungen an Hafer und Weizen. Lunds Universitets Arsskrift, INSEE Atdi2 Bde s-oNr: 2. pp: 122. VlQoo: Shull, G. H. Results of Crossing Bursa bursa-pastoris and Bursa Heegeri. Proc. 7th Internat. Cong. Zoél. 1907, Boston 1912. ‘‘ Advance reprint ”’ in 1910, 6 pp. 1907. — Bursa bursa-pastoris and Bursa Heegeri: Biotypes and Hybrids. Carnegie Institution of Washington. Publ. No. 112, pp. 57. 1909. — Defective Inheritance-ratios in Bursa hybrids. Verh. d. naturf. Verein Briinn 49: 157-168. I9gII. —— Duplicate genes for capsule-form in Bursa bursa-pastoris. Zeitschr. f. ind. Abstamm.- u. Vererb. 12: 97-149. I914. ISOLATION AND SPECIFIC CHANGE EDMUND W. SINNOTT Connecticut Agricultural College Those regions of the earth which are so isolated biologically that the dispersal of plants or animals between them and other areas is difficult or impossible are characterized, as is well known, by large numbers of species and genera which are peculiar to them or are ‘endemic.’ In general, the more definitely isolated the region the higher is its proportion of local forms. Why isolation should be associated so universally with the presence of these endemic types is a problem which has excited speculation. It is evident that once a local race is established, isolation will operate effectively to maintain it, both by preventing its dispersal abroad and by excluding invaders which might supplant it. The difficult problem has been to account | for the actual origin of the endemic types themselves in the first place. In an attempt to throw light on this problem a study has been made of the floras of a number of islands which are isolated to a greater or less degree from adjacent land masses and have developed a large body of local species and genera of vascular plants.' Certain elements of the endemic flora in all these islands are doubt- less not of local origin but are ‘‘relicts,’’ remnants of types once much more widely spread, which owe their preservation to freedom from the keener competition of the mainland. These types do not con- cern our problem. As to just how abundant they are we do not know, but those forms which stand well apart and have no near relatives in the islands or elsewhere are probably to be looked upon as relicts. From the close similarity of most of the endemic species and genera with others on near-by islands or on the adjacent mainland, however, and from their frequent occurrence as groups of related forms, it is evident that the bulk of the endemic element in these floras is actually of local origin. Several hypotheses have been put forward to account for the origin of these endemic forms. Some investigators have pointed to natural selection as the primary factor, believing that new types are produced by this agency to fit exactly the peculiar conditions in each region, rather than a wide environmental range. Others, less con- 1 These islands are New Zealand, Ceylon, Hawaii, the Galapagos, Juan Fernan- dez, St. Helena, Sokotra and Mauritius. 444 SINNOTT: ISOLATION AND SPECIFIC CHANGE 445 vinced of the efficacy of selection, believe that each region has its own characteristic environmental complex, different from that of all others, which modifies directly the germ plasm of the animal and plant types living under it and stamps upon them their local distinc- tions. Both of these views regard the environment as the most important factor in specific change and look upon isolation as the agency which, through providing a comparatively simple and constant environment, allows a much closer adjustment to it by the plant and animal population than is possible on wider areas, and hence leads to the production of large numbers of local species. Still another view considers that most, if not all, of these endemic and peculiar forms would have developed anyway under the progressive evolution of their type, and owe their local character not to a dependence, direct or indirect, upon a specific environment, but merely to the fact that they have been unable to’ become dispersed abroad. An analysis of the insular floras under investigation presents certain facts which have a bearing on the problem. It makes evident, in the first place, that endemism is by no means uniformly characteristic of all the elements in the flora but that it occurs very much more fre- quently in certain of the great groups of vascular plants than in others. The vascular cryptogams, for example, which comprise an important part of the vegetation of these islands, include but few species or genera which are confined to any one island or island group. The glumaceous monocotyledons—Gramineae, Cyperaceae and Juncaceae —which are also abundant, are represented infrequently among the endemic forms, though they are somewhat commoner there than are the vascular cryptogams. It is in the petaloideous monocotyledons and the dicotyledons that the great bulk of the endemics occurs throughout all of these insular floras. Not only hosts of the species but almost all of the local genera belong to these groups. Certain families, like the Orchidaceae and the Compositae, often contain almost nothing but endemic species. How great is this disparity in the extent to which endemism occurs is evident from the following table, which is an average of the eight island groups investigated. | Species Genera | Endemic Non-endemic Endemic Non-endemic rs ey Tse Saas Waseular Cry ptogams...4........- | 23.2% 76.87% 1.9% 98.1% Glumaceous Monocotyledons.......| 31.4% 68.6% 2.2% 97.8% Petaloideous Monocotyledons...... 59.0% 41.07% 9.7% 91.3% DicotyledoOns'.,>. he hi cite epee ects 61.7% 38.3 % 11.4% 88.6% __ What bearing have these facts on our problem of the origin of local types? They offer little support, in the first place, to the theory 30 446 BROOKLYN BOTANIC GARDEN MEMOIRS that natural selection has presided over the appearance of these new forms, for groups which have developed few or no endemic species are apparently as successful elements of the vegetation as are those in which such species have been abundantly produced. In fact, Willis? has gathered evidence from the flora of Ceylon which seems to show that the non-endemic species are more successful, as a whole, than the endemic ones, a fact which militates strongly against the theory of selection. Of course we are confronted here, also, with one of the major difficulties urged against natural selection, namely that it can never create but can only eliminate. Nor do our figures support the theory that local forms owe their origin to the direct action of the environment, for such a theory can- not well explain the abundance of endemic species in some groups and their rarity in others. It may be argued that the vascular crypto- gams and glumaceous monocotyledons are more primitive and slow- changing types than the petaliferous groups, and are thus able longer to resist the pressure of the environment and to maintain their original characters. We have little evidence, however, that this is actually the case. Ferns under cultivation seem to be very plastic, and our knowl- edge of the genetics of the Gramineae, at least, does not indicate that they are a particularly rigid group. Both of these views look to the environment as the factor, either direct or indirect, which is chiefly responsible for the origin of new forms, and both are open to the objection (among others) that although the whole flora is subject to the same environment, these new forms develop only in certain groups. Our third alternative largely dis- regards the environment. It looks upon the actual production of new types as due to factors within the organism rather than in its surroundings, and considers that the locally developed species and genera in the floras under discussion would have appeared in these regions whether isolation existed or not. Isolation is thus regarded merely as the agency which keeps these new forms local and endemic by preventing their dispersal beyond the place of their birth. Of course such a theory allows for the play of selection in weeding out all new forms which were distinctly unsuited to the environment under which they appeared. But is not this view also open to the objection which we have offered to the others, that it cannot account for the rarity of endemism in certain groups and its extreme commonness in others? A study of the methods of reproduction in plants belonging to these two cate- gories suggests an answer to this question. Vascular cryptogams in the great majority of cases have bisexual gametophytes and are 2 Willis, J. C., The evolution of species in Ceylon, with reference to the dying out of species. Annals of Botany 30: I-23. I9g16. SINNOTT: ISOLATION AND SPECIFIC CHANGE 447 doubtless almost invariably self-fertilized. In the glumaceous mono- cotyledons, although crossing is certainly not uncommon, it will probably be agreed that self-fertilization is also the general rule. In these two groups we have noted that local species and genera are very rare. In the petaloideous monocotyledons and in the great majority of dicotyledons, on the other hand, the flowers are attractive ‘to insects and cross-fertilization preponderates. As far as our knowl- edge goes, there are few petaliferous species which are not at least occasionally cross-pollinated. These facts are significant when we remember that it is among such forms that local types are produced in great abundance. In short, our analyses of these insular floras sup- ports strongly the theory recently emphasized by Lotsy and others that hybridization is the most important factor in the production of new forms; self-fertilized types changing slowly, cross-fertilized ones, rapidly. The unequal development of endemism in certain floral elements, therefore, which neither the theory of selection or that of the direct effect of the environment can well explain, is thus under- standable as the result of differences in methods of reproduction, and is quite consistent with the theory that the production of new forms is due primarily to internal causes. The evidence brought forward by our study of isolated insular floras therefore seems to indicate that isolation is not a factor in evolution but simply in distribution; that it cannot create an endemic species but can merely keep a species endemic. SUMMARY 1. Isolated regions are characterized by possessing large numbers of local, or endemic, species and genera. 2. In the insular floras investigated, endemism is not equally distributed among all plant groups, the local species and genera being rare among vascular cryptogams and glumaceous monocotyledons but very common among petaloideous monocotyledons and dicotyledons. 3. This fact seems to indicate that the environment, whether acting directly or by means of natural selection, has not been the determining factor in the development of endemic forms. 4. Those groups which are poor in endemics are predominantly self-fertilized, those which are rich in endemics, predominantly cross- fertilized; suggesting that hybridization has been the most potent factor in the development of these new forms. 5. Isolation is therefore regarded not as the factor which, by pro- viding a simple, limited and peculiar environment, is responsible for the origin of locally developed species and genera; but merely as the factor which, by prohibiting dispersal, maintains the endemic char- acter of local types which owe their origin to other causes. THE RELATIONS OF CROWN-GALL TO OTHER OVERGROWTHS IN PLANTS ERWIN F. SMITH Bureau of Plant Industry, U. S. Department of Agriculture In the time assigned the most I can hope to do is to give the barest outline of the suggested relationships. Some of these are well determined; others are only suspected and are mentioned here as hopeful lines of research rather than as definite conclusions. Indeed, I am quite willing to admit that our work on crown gall has opened up more problems than it has settled, but, one way or another, all of this present uncertainty will make for progress and an eventual better understanding of the whole mechanism of overgrowth. My own belief is that all overgrowths are correlated phenomena, are the response of the organism to essentially similar (but not necessarily identical) stimuli, the visible difference in response when brought about by parasites being due to number and location of the parasites, age and kind of tissues invaded, and volume, direction, and velocity of the stimulus exerted. In other words, in every case, I think the stimulus is primarily a physical stimulus due to changed osmotic pressures rather than a direct chemical stimulus. Overgrowths, therefore, do not always involve the presence of a parasite although as observed in nature parasites are probably responsible for most of them. I. Factors governing type of overgrowth in crown gall. A. The host reaction depends on the type of tissue infected. (1) Vascular vs. parenchymatic. For example, depend- ing on the tissue in which it originates the vessels in a tumor may be numerous or few, the paren- chyma abundant or scanty. (2) Nexus of cells stimulated, 7. e., unipotent, multi- potent, or totipotent cells. Thus, according to the tissues infected by the crown gall schizomycete, we have it causing either organoid galls or histioid galls. (3) Rate of growth. The rate of growth depends on the readily available supply of food and water, on the age of the tissues when infected and on the species 448 SMITH: CROWN GALL AND OTHER OVERGROWTHS 449 attacked—some species are not subject to this disease; old tissues respond slowly. (4) Individual differences. ‘There are, I believe, indi- vidual differences in susceptibility as well as species differences. B. The host reaction depends also on the activities of the parasite (Bact. tumefaciens) which are variable. (1) Loss of virulence on culture media. The cause of virulence is not known. The effect of long continued growth on culture media is to reduce the virulence of the organism and finally to destroy it altogether. (a) “Old” vs. “resistant” cultures of Paris daisy organism. Our first isolation from Paris daisy was extremely virulent in the begin- ning but lost all power to produce galls in about three years. Another isolation which we called ‘Resistant Daisy” is now slowly losing virulence at the end of three and one half years. (b) New vs. old cultures of poplar isolation. An isolation from a poplar tumor was extremely virulent for some time (several years) but has now lost all power to produce tumors. Along with this loss has come a progressive thickening and toughening of the pellicle on bouillon. This was true also of the daisy isolation which lost its virulence. Apparently this loss of virulence is not correlated with loss of power to produce formic acid for, according to the chemist, the non-virulent poplar organism still pro- duces that substance. Loss of power to infect must be related, however, to loss of some chemical or physical property once possessed, and surely we ought to be able to discover the exact nature of this loss. Some strains of Bact. tumefaciens lose virulence much sooner than others. One of our strains (from hop) is still virulent after nine years on culture media." (2) Not every isolation is a distinct strain. I speak of “strains’’ only when I know that cultural 1 Ten years, as this now goes through the press. 450 BROOKLYN BOTANIC GARDEN MEMOIRS and other differences exist; otherwise, I speak only of “isolations.” There are cer- tainly two of these crown gall strains, and probably many. (3) Feeble and virulent strains exist in nature, 1. e., there 1s variation in virulence of colonies from the same source—hop, carnation, rose, sugar beet, etc. The author believes that the crown gall bacteria not only lose virulence on cul- ture media but also in the tissue of the gall. Examples are: (a) of three colonies selected as typical from plates poured from a hop tumor in 1910 only one proved infectious; (b) of six colonies plated from a witch broom on carnation, all of which looked alike and typical for crown gall, only one was found to be able to cause tumors when inoculated; (c) of seven colonies selected as typical from a plate poured from a rose gall only three proved infectious, and of these two were actively infectious, while the other was only feebly so; (d) of five colonies selected as typical from a plate poured from a tumor on Pelargonium none proved infectious, al- though in advance we felt quite sure of all of these colonies; (e) of thirty colonies selected from plates poured from natural tumors on sugar beets only five were infec- tious and all feebly so (Bul. 213, pp. 192-194 and Plate XXXVI). The first and natural supposition when a culture has lost virulence is that some in- truder has displaced the right organism; and when only certain colonies on a plate are in- fectious, that the others are intruders how- ever much they may resemble the right organism. I cannot say that we have abso- lutely excluded this hypothesis, to which I held strongly in the beginning, but we are gradually coming to believe that it does not explain all the facts. II. Some other types of plant galls. (1) Nonparasitic intumescences. These can be produced in several ways: by overwatering and exposure to SMITH: CROWN GALL AND OTHER OVERGROWTHS 451 excessively moist air; by exposure to very dilute vapors of ammonia or of formaldehyd; by applica- tion of a great variety of soluble substances not the products of parasites, e. g., copper salts; by painting the surface with vaseline and other insolu- ble substances which interfere with the gas exchange; by freezing lightly (Harvey), etc. (2) Non-cancerous bacterial tumors—olive knot (due to Bact. savastanot), beet tumor (due to Bact. beticola). In these the bacteria are abundant and easily seen occupying the vascular bundles and the spaces between cells. Bacterial cavities are produced and the overgrowths are only superficially like crown galls. The secondary tumors are not developed from tumor strands. When the bacteria are intra- cellular the cells are destroyed. 3) Nematode galls. In galls due to Heterodera radicicola giant cells, 7. e., several to many nucleate cells, are com- mon. Parasitic nematodes which do not produce galls. Here the anal excretions are voided outside of the plant (Cobb). Occurrence of several-nucleate cells in crown gall. (4) Various fungous galls. Parenchymatic vs. woody; _perish- able vs. persistent; witch brooms (see newer work on crown gall). Parenchyma strands (Dodge). (5) Insect galls which show features resembling crown galls. (a) Galls bearing leaves; galls bearing roots; galls bearing modified trichomes. We have now suc- ceeded in producing on internodes by _ bacterial inoculation crown galls bearing roots, leafy shoots, flower buds, and modified trichomes. (b) Galls with cortex carrying bright colors—purple, red, yellow. Crown galls produced on pale green balsam stems show a red color in their cortex provided the plants bear colored flowers, but not if they bear white flowers. The production of red pigment in the cortex has been seen also in axillary (teratoid) crown galls developed on red-flowered Pelargoniums. Etiolation. Excess of chlorophyll. (c) Galls opening by opercula—strange but not more so than twin-leaf fruits opening in a similar manner, or than a double vascular cylinder in Ricinus with death of intermediate pith and separation into two cylinders. (Jour. Ag. Res. Jan. 29, ’17, pl. 37). BROOKLYN BOTANIC GARDEN MEMOIRS (d) Galls with very definite and distinct strata of gall tissue—parenchymatic, vascular and _ protective, e. g., cynipid galls. Kiister’s prosoplasmatic galls. Many insect galls differ from crown galls in that (1) the parasites are few or reduced to a single organism, and (2) are extracellular, whereas in crown gall the parasites are more numerous and are intra- cellular. Many differences in structure, even of the more complex galls, can be explained, I think, by these two differences, especially if we assume (3) that the kind of reaction depends on the volume, direction, and velocity of the stimulus, its constant or intermittent flow, and on location, distance, and mobility or immobility of the source of the stimulus. As in various insect galls so in crown galls, there is a tendency toward the production of more primitive tissues and of various anomalously formed organs. III. Crown galls are formed by extrusion of chemical substances. I have recently produced galls with diluted crown-gall products and this, it seems to me, suggests a new method of attacking gall problems in general, especially those in which the gall para- sites can be cultivated pure in sufficient quantity for chemical analysis, e. g., various fungi. Striking results have been ob- tained but many tests are yet to be made with the crown-gall substances in various dilutions, mixed and separate on a variety of tissues of responsive ages. Various types of cell growths have been produced by the action of ammonia, acetic acid, formic acid, aldehyd, etc. (all products of Bacterizim tumefaciens, the crown-gall organism) in less than killing doses, that is, various degrees of hypertrophy and hyperplasia of cells and mixtures of the two have been observed. Sometimes there is great stretching of cells as in certain fungous and insect galls. Giant cells in the animal pathologist’s sense of that word, namely, cells containing several to many nuclei, such as occur in the common nematode galls, are to be searched for in all sorts of plant galls and to be produced, if possible, experi- mentally, 7. e., with gall-forming substances. In due time we shall be able, I believe, to get these multinucleate cells at will. Probably they are weakened cells. “Two very important things to be determined are whether the size of the cell depends on the volume or rate of movement of the stimulus or on the kind of stimulus, and whether mixed stimuli applied in varying proportions change the manner of cell reaction. SMITH: CROWN GALL AND OTHER OVERGROWTHS 453 IV. Other effects of parasitic excretions. I believe also from what I have seen and have obtained to some extent by experiment that thyloses, fasciations, distortions of tissues, and various duplications, simplifications and inverse tissue differentiations are caused by the excretions of feeble parasites although in nature probably all are not so caused. Finally, I would suggest that the application of chemical substances in various dilutions to growing plants or plant organs, such substances In particular as are known or suspected to be produced by living organisms, or are present in soils as a result of decompositions, may prove to be a hopeful way of attacking certain unsolved and difficult problems in plant pathology, e. g., the aetiology of the mosaic diseases, the cause of various growth limitations, etc. LITERATURE Those who wish details on crown gall are referred to the following papers: For the older work: Science, N. S., April 26, 1907; Phyto- pathology, Vol. I, No. 1, Feb., 1911; Science, N.S., February 2, 1912; 1° Congrés International de Pathologie Comparée, Tome II, Paris, 1912; 17th International Congress of Medicine, London, 1913, Sec- tion III, General Pathology; and Bulletins 213 and 255, Bureau of Plant Industry, to be had from the Superintendent of Documents, Government Printing Office, price 40 and 50 cents, respectiyely. For the newer work so far as published: The Journal of Agri- cultural Research, April 24, 1916; The Journal of Cancer Research, April, 1916; Science, N.S., June 23, 1916; The Journal of Agricultural Research, January 29, 1917; Bulletin of the Johns Hopkins Hospital, Sept., 1917; and Proceedings of the American Philosophical Society, Vol. 56, 1917. Separates of most of these papers may be had from the writer. CONTACT STIMULATION GEORGE E. STONE Amherst, Massachusetts The experiments presented here have been carried on since 1904, at which time the writer observed some rather remarkable stimulated growth responses induced in sunflowers when surrounded by wire netting. At the time these observations were made we were carrying on investigations relative to the effect of varying atmospheric electrical potentials on plant growth, and for this purpose we made use of sun- flowers established in large earthen pots or wooden boxes located at different elevations in the open air. In some of these experiments the plants were surrounded with wire netting and in contact with the same but not with the soil, while in others (normals) no wire netting was used. In some instances the wire netting was not only in contact with the plants, but with the soil in which the plants were growing; the soil being grounded by the use of copper plates in the bottom of the boxes and by insulated wire, which led to the earth. In other instances the plants were in contact with wire netting and the soil, but were not grounded. The problem under consideration at that time, however, more particularly concerned itself with the influences of atmospheric electricity on plant growth, for which purpose organisms of various kinds, including bacteria, were exposed to elevations varying from thirty to sixty feet. Some of the earlier experimenters have maintained that when plants were grown in the free atmosphere surrounded with wires, they failed to develop, and would eventually die in consequence of being deprived of the bene- ficial effects supposed to be derived from atmospheric electricity. In passing we may state that we have never observed any remarkable mortality among plants in consequence of their being surrounded with wires even when the experiments were performed at more or less high elevation above the ground, and under ideal conditions for determining the effects of atmospheric electricity on vegetation. Moreover the growing of plants in conservatories where the electrical conditions of the atmosphere are quite different from those out-of-doors demon- strates the fallacy of this idea. On the other hand, we found that plants were greatly stimulated by wire enclosures, especially when they came in contact with the plants, and also the same stimulation was noted when plants were grown thickly together and the leaves 454 STONE: CONTACT STIMULATION 455 touched those of other plants. An electrically charged atmosphere, however, exerts a marked stimulation on plants and it is possible to modify the function of organisms located at more or less high eleva- tions by the use of metal coverings. The old idea that milk sours more rapidly during thunder storms, and that plant growth is greater following electrical storms has in reality a fundamental basis. The discovery of contact stimulation led us to modify our methods of studying the effect of electrical potential on plants since we found that when the plants were not in contact with one another, or with the surrounding wire mesh quite different results were obtained. The observations and results obtained by contact of plants with one another and with wires, etc., was so significant that we undertook the investigation of this phase of the problem at that time, and have devoted considerable attention to it since. Some of our earlier experi- ments were conducted out-of-doors and parallel experiments were carried on in a conservatory. Repeated tests of the air in our con- servatory with a water-drip apparatus and electrometer have in- variably shown that under ordinary weather conditions there exists no atmospheric electricity in conservatories, the glass apparently acting as a screen. The nature of the stimulation due to contact is probably in no way associated with atmospheric electrical phenomena, or at any rate, the growth responses do not appear to be identical with those resulting from ordinary electrical stimulation. The response to contact is induced by the use of various materials, such as wire, twine, wood and metal stakes, excelsior, sphagnum moss, soil particles or even by the plants being in contact with one another. The same reactions are produced whether the different contact materials used are suspended and only touch the leaves of the plant, or whether they touch both the leaves and soil in which the plants are growing. While there is no evidence to show that these reactions are associated with any changes in the electrical tension of the at- mosphere, surrounding the plants, they may, however, be connected with electrical phenomenon. The reactions resulting from contact stimulations are not unlikely quite primitive and universal to plants and probably common to the lower forms of life in general. Probably all organs will prove to be sensitive to contact but from our observa- tion the leaves appear to be especially so. The nature of the reactions appear to be fundamentally similar to those of touch, from which it would seem the more highly differentiated reactions of tendrils and wound responses, etc., originated. 456 BROOKLYN BOTANIC GARDEN MEMOIRS Fic. 1. Showing growth of sunflowers surrounded by wire netting. Note difference in size of plants which have penetrated through the wire-mesh enclosure, a feature of common occurrence. METHODS In the study of the effects of contact of various materials with plants we employed several methods and the experiments were carried on under different conditions. With the exception of some of the earlier experiments which were made out-of-doors, most of them were STONE: CONTACT STIMULATION 457 conducted in a conservatory where the heat, light and soil conditions were uniform and normal. Some of the experiments were carried on in direct sunlight; others in darkness, while others again were con- ducted under bell glasses where it was possible to maintain uniform atmospheric moisture and transpiration conditions. Contacts with wire were made with a two-inch-mesh galvanized iron wire netting, and in some cases a one-inch-mesh wire was employed. These were made up into cylinders 4 to 6 feet high and varied in diameter from 8 to 26 inches, according to the size and number of plants employed. These wire cylinders were placed around the plants. In some in- stances the plants were grown between trellises made of wire netting placed from 6-10 inches apart, in which case the tips or free end of the leaves of the plants came in contact with the wire on two sides. Wooden stakes (dowels) and iron posts driven into the soil were also employed as contact material, the dowels being used in some instances to support loose twine which surrounded the plants and in other cases they were used alone in various combinations. Fishnet made of twine with a mesh of about two inches was employed in a similar manner to that of the wire cylinders, and in some cases the plants were more or less loosely tied up with twine. In the study of the effects of contact on the stems and roots of seedlings, excelsior was employed either loosely in long strands, or cut up more or less in fine shreds as the nature of the experiment required. In the root, contact experi- ments were made with soil particles which ranged from 16 mm.—o.I mm. in size, the various grades of material being obtained by sifting through sieves and bolting cloth. The particles were repeatedly washed and sterilized and submerged in jars of water, the latter being previously boiled to exclude air, inasmuch as the presence of air would greatly modify the growth of the seedlings (1). EFFECTS OF CONTACT OF PLANTS WITH ONE ANOTHER When plants are grown close together, as is the case of some crops, they often come in contact with one another and even in nature con- tact stimulation plays an important réle, particularly where certain organisms grow so close to one another that they touch. The growth of some coniferous trees is often such that they are much congested, and in nurseries where many thousands of coniferous seedlings and other nursery plants are grown close together a contact stimulation may sometimes occur. We have, however, no authentic data based upon experiments regarding the behavior of coniferous and other seedlings grown under nursery conditions. Neither have we attempted to differentiate growth which may be due to contact in thick stands of forest growth from those originating from other causes, but some 458 BROOKLYN BOTANIC GARDEN MEMOIRS gardeners and foresters have intimated that they have observed indi- cations of an accelerated growth in height as a result of coniferous seedlings coming in contact with one another. The stimulation effect of contact, however, can be observed in the growth of crops and the Fic. 2. Showing growth of tomatoes im situ in contact and not in contact with one another. method of close growing of certain economic plants has its advantages. The configuration of plants, however, is greatly modified by close growing, as may be observed in the handling of single-stemmed chrysanthemums by florists, the growth of corn and various other TABLE I Showing Growth of Tomatoes (Lycopersicum esculentum Mill.) in Contact and Not tn Contact with One Another! Average Height and Diameter of Plants (Centimeters) | Percentage Gain by Contact Height | Diameter ies. * Height | Diameter Nonialess tannic satiate 19.41 .46 | Contact) pescni cise ere ee 47 | 60% | 295 plants. The stimulative effect of contact of one plant with another is shown in Fig. 2. The tomatoes in this case were grown in a well- lighted conservatory in soil similar in all respects and the water supplied was such that each plant obtained similar amounts. (Cf. Table 1.) 1 36 plants used. STONE: CONTACT STIMULATION 459 RESPONSE OF PLANTS TO CONTACT STIMULATION WITH WIRES AND TWINE The investigations relating to the effects of contact with wires, etc., are given in Tables 2-9. All of these experiments were made in a well-lighted conservatory, the plants being grown in a good uniform grade of loam either in solid beds or in benches. In this series I-6 plants were enclosed by wires, or dowels, and twine, and in some cases only fish netting was employed. The normal plants were in some instances grown free from contact with one another and in others not. In the case of only a single plant being surrounded by wire netting contact would occur only with the wire, whereas when two or more Fic. 3. Showing growth of castor beans in contact and not in contact with wire netting. Plants removed with as little disturbance as possible from original position for photographic purpose. plants were grown tolerably close together they would eventually be in contact with each other as well as with the wire netting, etc. Hence contact stimulation would result not only from the use of wires, etc., but from the contact of plants with one another, or in other words the so-called normal plants were not in all instances free from contact, inasmuch as when they were grown in groups they would eventually touch one another and growth would be influenced. We therefore have two series of experiments, namely: (a) those in which the 460 BROOKLYN BOTANIC GARDEN MEMOIRS normal plants were perfectly free from contact with one another and (b) those in which the so-called normal plants were more or less in contact with one another. Both of these normal series were compared with those in direct contact with wires, etc. The plants in this series of experiments all show a gain in height by contact with different material. The effects of contact with wire netting (2-in. mesh) as compared with no contact whatsoever in the normal plants is given Fic. 4. Showing growth of tomato plants in contact and not in contact with wire. in Tables 2-4. In the former experiment (Table 2) 12 plants were utilized, all of which were grown separately, 6 being surrounded by cylinders of wire netting and 6 grown free from any contact whatso- ever. The plants in contact with wire netting showed a gain in height of 31 percent compared with those of the normals and the same percentage gain is given in Table 3, in which case the contact plants STONE: CONTACT STIMULATION 461 TABLE 2 Showing Growth of Sunflowers (Helianthus annuus L.) in Contact with Wire Netting. Average of One Experiment with Twelve Plants* Percentage Gain Average Height in Height by (Centimeters) Contact INormal tes is.5.0 85 ke oe ee 131.9 Gontact wites..7. 2) s. oe 173.0 31% TABLE 3 Showing Growth of Sunflower (Helianthus annuus L.) in Rows between Wire Netting. Average of Two Experiments with 54 Plants® Average Dimension and Weight in Centimeters and Grams ; | Internodes | , | Percentage Gain in | Height Length | Weight | Height by Contact a EA ee A | | eee (ne A, | ENO a letter reeves 5. oiovee cus | 114.1 6.9 307.9 Contact wire 4522s o eee. | 151.2 8.9 302.9 31% TABLE 4 Showing Growth of Castor Bean (Ricinus communis L.) in Contact with Wire Netting! Average Dimension and Weight in Centimeters and Grams cod Diameter Entecodes) | Weight Percent Gain in Stem Nantes | Length | Height by Contact SS SS ee ee INormalignvent-- 73 1.8 9.2 | 7.8 | 457 Contact wire...| 99 1.5 Solel t= wee 3 lee oA 35% were grown between parallel rows of wire netting, the netting being of sufficient distance apart to come in contact with the leaves of the plant. Neither the normal plants nor those in contact with the wire touched each other. The experiments with the castor beans shown in Table 4 were identical with those given in Table 2. This showed a gain of 35 percent in the contact plants over the normal. The re- maining 5 tables (5-9) show the effects of surrounding plants with wire netting, dowels and twine, and fish netting. Since in this series, from 2-6 plants are grown close together, they also show to a certain extent the effects of contact with one another. Both the normal and contact plants were in pairs in Tables 5, 6 and 8a while 7 and 8b were arranged in threes and those in Table 9 contained 6 plants. In Table 7a, a wire netting was suspended overhead by twine and did not come in contact with the soil. From the data given in these tables it is hardly possible to draw exact deductions as to the relative 2 Normal plants separated from one another. 3 Normal plants separated from one another. 4 12 single plants used, 6 in contact and 6 not in contact. 31 462 BROOKLYN BOTANIC GARDEN MEMOIRS value of the various methods employed in inducing responses to con- tact stimulation. Deductions, however, based upon a large series of experiments not included here justify us in stating that surrounding plants loosely and irregularly with twine does not produce the same degree of stimulation or response, as by the more thorough contact derived from the use of other material such as wire netting with a uniform mesh, or, in other words, plants react more pronouncedly to a larger contact surface than to a relatively smaller one, although there probably exists a definite size of mesh which would produce the best result, and this would undoubtedly vary with different species. There appears, however, to exist some difference in the degree of stimulation arising from the same size mesh, as shown by the behavior of some species when the contact is applied to the leaves. The leaves, for example, of the sunflower and corn do not respond so freely as those of tomatoes and the castor bean presumably because the leaf apices are the most sensitive as in the case of tendrils. The latter species, possessing different type leaves, would appear to act differently on this account. TABLE 5 Showing Growth of Castor Bean (Ricinus communis L.) in Contact with Wire Netting’ Average Dimension and Weight in Centimeters and Grams Internodes | | Percentage Gain i | | } | ie ia l Per ain in | Height | Diameter Stem | Wake | on | Weight | Height by Contact | Normail:.5...- =.) 32-3 rae <= ly eto |e ih gee Contact wire...| 55.1 1.18 IS rES Ie 426 200 70% The data derived from these experiments are not sufficient to allow of deductions being drawn which would be of any value in determining the relative value of the various-sized meshes in stimulating growth. In some cases where galvanized iron netting with a one-inch mesh was employed, the stimulation appeared to be equally as great as with the two-inch mesh. Neither is it possible by these tests to determine accurately the difference in the degree of stimulation which resulted from the use of wires and that from the contact of the plants them- selves. In all cases where single plants were employed they were removed far enough away from one another so as not to touch. The stimulated growth, therefore, was due entirely to the material which surrounded them. On the other hand when plants were grown in such a manner as to touch one another there existed two sources of contact. In the experiment shown in Table 9, six sunflower plants 572 plants used. Plants in pairs. STONE: CONTACT STIMULATION 463 were grown close together (about 8 inches apart) and the two series of normal plants which consisted of twelve or six in each set were in contact with one another throughout their period of development. Hence, the percentage gain in growth in height is due to the additional contact of the plants to the various materials which surrounded them. The largest increase in growth is shown in Table 5 but this data was obtained only by the use of 12 plants and probably is exceptional. When closely grown plants are surrounded with wire netting, etc., and especially large-leafed plants such as the common sunflower the leaves do not have an opportunity to assume their normal shape and in this way there occurs a tendency to shade the stems. The dis- placement of the leaves by the wires thus shading the plants or the slight shadow cast by the wire enclosures apparently has little effect on the growth in length of the internodes, inasmuch as the same reactions can be obtained by growing the plants in darkness. The data concerning the results given in the tables (2-9) follow. TABLE 6 Showing Growth of Castor Bean (Ricinus communis L.) and Corn in Contact with Twine® Average Dimension, etc., in Centimeters Experiments No. of Plants Height Faterupdee Percentage Gain in J elg j - as Used Masabes | ane Height by Contact Goa Normals... 2......: 24 81.7 t46. £58 Contact twine.... 93-6 130 sO: 12%. 0 2F Corn (Zea Mays L.) a | | (Dye Normalln., 5 <0. fi: 36 [226 10.0 12.4 Contact twine... . | 145.6 10.1 14.0 9% TABLE 7 Showing Growth of Sunflower (Helianthus annuus L.). (a) Average of Twelve Plants in Contact with Wire Netting, (b) Average of Twelve Plants in Contact with Stakes (Dowels) and Twine’ Percentage Gain Average Height in Height by Experiments (Centimeters) Contact (Gy NGENTAE 2 os ci ieg oir cas 137 Contactewites 42728 35.- 164 19% (>) BNocma lcs bye se. faa 132 Contact stakes and twine.178 34% 6 Plants in pairs. 24 plants in (a), 36 in (0). 7 Plants in triplets. 464 BROOKLYN BOTANIC GARDEN MEMOIRS TABLE 8 Showing Growth of Sunflower (Helianthus annuus L.) in Contact with Wire Netting® | ; : 3 : : Average Dimensions and Weight in Centimeters and Grams Experiment ae Internodes | Pp Gai Height! oe ay a eat Wekee in Hagen | Number Length by Contact (a) Normal....... 108.80 1.35 TAc2e. Nip MOLOws Ne Aes | Contact wire..| 138.91 | 1.33 14,8. | 10:4) |p 267-9 27% (b) Normal......., 81.5 | 0.80 17 Ala, O06 __Contact wire. .| 109.2 | o:84 ||) 242504) 4:5 cpl O256 33% TABLE 9 Showing Growth of Sunflower (Helianthus annuus L.) Surrounded by Wire and Twine. 60 Plants Used in Each Test. Experiment Made in Greenhouse in Benches® | Average Dimension and Weight in Centimeters and Grams | | ri Percent Gain in Height | Diameter | Internode | Weight | Moisture Height by Length Percent Gontace Normal 5 oes | 132.5 1.59 8:02) 22133 87.6 Contact wire......| 151.0 1.46 9.01 | 216.0 89.6 15% Contact twine=....- 155.0 1.43 | 871 || 210.0) 7) 90:4 16% Contact fish net...) 159.0 1.40 | 10.50 l“x60!0" =z9r'8=| 20% The average diameter of the stems of the normal was 1.2 cm. and for those in contact 1.24 cm. The average number of internodes for the normal was 13 and for the contact plants 13.6; while the average length of internodes for the normal was 6.8 cm. and that for the contact plants 8.9 cm. or a gain of 30 percent. The average weight for the normal plants was 243 grams and for the contact 216 grams. The diameter, number and length of internodes was slightly greater in the contact than in the normal. The average weight was 12 percent greater in the normal than in the contact plants. The moisture contents of the plants were greater in the contact plants than in the normal ones. The most important difference between the normal and contact plants is in the length of internodes. RESPONSE OF PLANTS TO CONTACT WITH EXCELSIOR All of the experiments with excelsior were made with seedlings and were carried on in darkness. These were made in large flower pots containing either soil or sawdust. The excelsior was packed loosely over the, soil or sawdust as the case might be at the time the seeds 8 12 plants used in each experiment. (qa) Plants in pairs; (0) in triplets. 9 Plants in sixes. STONE: CONTACT STIMULATION 465 were planted, to a height of about Io inches and often less, depending upon the nature of the plant in use. The excelsior was cut in lengths varying from 1 to 10 cm. or according to the nature of the plant employed. Since these experiments were all made in darkness and the plants were in all cases covered with receptacles, such factors as light and transpiration were controlled, and under these conditions the seedlings were dependent largely on the reserved material con- tained in the seeds. There was, however, no contact of the plants with each other. Moreover in this series, the same contact material was used throughout, namely excelsior, and any specific reaction which might arise from the use of different materials in contact with the TABLE I0 Showing Growth of Horse Beans (Vicia Faba L.) in Contact with Excelsior® Average Height and Weight in Centimeters and Grams Experiments | # Porriiage Gan by No. of Plants F : ees Height Weight * Height Weight (a a Normalee 9: see souk 149 14.03 0.96 Contact excelsior....... 18.20 1.28 29% 33% (py miNormaless i: Se alk. 374 15.10 0.74 Contact excelsior....... s 18.50 1.05 18% 41% Mone IN Ontiiall crs. shay eee [Gree 1.37 | , I22 | Contact excelsior....... | 20.41 bf Om 77/7, 9% Average mormal: ..5cj550 0-6 15.48 1.02 | OIPACE Jo the he ie | 19.03 F631.” | 23%. 3. 5870 TABLE II Showing Growth of Hypocotyls of Lupines (Lupinus albus L.) in Contact with Excelsior. Average of Two Experiments with 128 Plants Average Height of Hypocotyls (Centimeters) Percentage Gain Hypocotyls by Contact IN ODI testi hi teie tw ache ee 19.28 Contact excelsior.......... 21.65 12% plants would not be present in these. There would of course occur variations in the stimulus imparted to the different species owing to difference in the relative degree of contact of the plants with the excelsior—a feature which would be determined by the fineness, compactness and amount of the excelsior employed, as well as by the 10 Experiments in which light was excluded. 11 Experiments in which light was excluded. 466 BROOKLYN BOTANIC GARDEN MEMOIRS surface area of the organs in touch with the same. Inasmuch as the plants used were seedlings and a larger number were employed than in the preceding series, the results are likely to be more uniform. On the other hand it should be noted that with the use of seedlings grown under good heat conditions (in which case they would develop rapidly) the duration of stimulus would be much less prolonged, and the ultimate effect of contact on the configuration of the plants would be less pronounced. The most marked stimulating effect of contact would occur in general in those experiments which were the most prolonged, namely with the larger seedlings. With the excep- tion of corn all of the measurements given are either of the hypocotyls or stems. The results of these experiments are given in Tables 10-15, all of which show a stimulated growth due to contact with excelsior. The average weight in all instances where determined was greater in the contact than in the normal. In one water determination with lupine, there was 4 percent more moisture in the contact-stimulated plants than in the normal. In the experiments with corn (Table 14) measurements of leaves were taken as well as the cotyledons although it was our original intention to include in our measurements the cotyledons only, since when the leaves break through the cotyledons the growth of these organs is greatly retarded. The data giving the measurements of leaves in the corn, however, have a limited value as the growth of the leaves displayed more or less erratic behavior. In one experiment the leaves protruded above the excelsior and conse- quently they were not in contact. In another instance the leaves showed a very decidedly accelerated growth in the normal plants which was caused by a brief and accidental exposure to light. By taking proper precautions in further experiments this did not occur again. Some of the more sensitive cotyledons of the Gramineae are characterized by a marked growth correlation following mutilation or decapitation of the cotyledons in etiolated seedlings, and similar reactions occur to etiolated seedlings which are exposed to light. The function apparently of the cotyledons is to protect the true leaves in protruding upwards through the soil and as soon as they are exposed to light their growth ceases rather abruptly. Correlated with the retarded growth of the cotyledons is a greatly accelerated growth of the leaves which may amount to over 100 percent increase in two or three hours. The exposure of the cotyledons to light even for a brief period is sufficient to check their growth and greatly ac- celerate the development of the leaves and this is what happened on one occasion. The normal plants would be the most affected by any such exposure as they were not covered with excelsior, while the contact plants STONE: CONTACT STIMULATION 467 TABLE 12 Showing Growth of Peas (Pisum sativum L.) in Contact with Excelsior. Average of Two Experiments with Eighty-two Plants'* Average Height of Plants (Centimeters) Percentage Gain Stems by Contact Normal ooo ..2. ey ee 13.07 Contact excelsior.......... 16.66 27% TABLE 13 Showing Growth of Cucumbers (Hypocotyls) (Cucumis sativus L.) in Contact with Ex- celstor'8 Average Height of Hypocoty] (Centimeters) Percentage Gain Hypocotyls by Contact INOEmale ats oe ee oe 15.04 ContactiexcelSions 4 Hy.5-2 16.30 8.3% were to a more or less extent protected from light by the excelsior. One of the experiments with lupines which was carried on at the same time behaved in a similar manner to that of corn. In one case 31 out of 36 of the normal plants developed leaves averaging 7 cm. in length while none developed in any of the excelsior contact plants—a feature due to the same cause, namely, to a brief and accidental exposure of the plants to light. TABLE 14 Showing Growth of Cotyledons and Leaves of Corn (Zea Mays L.) in Contact with Ex- celsior. Average of Two Experiments with 72 Plants'4 Average Length of Cotyledons and Leaves (Centimeters) | Percentage of Gain by Contact Cotyledons Leaves ze Cotyledons Leaves IN Oniiia lire, oaeee- ene oye ci 8.41 17275 Contact excelsion= .. 2/205... : = 1007 - SF inSiag et 19% | 4% TABLE 15 Showing Growth of Turnip (Hypocotyls) (Brassica Rapa L.) in Contact with Excelsior. Average of Two Experiments with 194 Plants Average Height of Hypocotyls (Centimeters) Percentage Gain Hypocotyls by Contact INOnmaliey shes t cerca 8.57 Gontact/excelsion.-= 2-1. 2414 9.66 11% 12 Experiments in which light was excluded. 13 Experiments in which light was excluded. No. of plants used 26. 14 Experiments in which light was excluded. 15 Experiments in which light was excluded. 468 BROOKLYN BOTANIC GARDEN MEMOIRS RESPONSE OF Roots TO CONTACT STIMULATION Excelsior In the experiments so far enumerated no account has been taken as to the effects which various substances coming in contact with stems may have upon the growth of roots and other organs. It is quite natural to expect that if one organ is affected by a stimulus other organs will be, inasmuch as the organism as a whole responds to stimuli of quite insignificant character. Incidentally, we observed in our early experiments with seedlings grown in sawdust that the roots in some cases respond when the serial portions of the plant were in contact with excelsior. In 1912 experiments were carried on for the purpose of comparing the growth of roots in soil under different conditions. These experiments consisted in the growing of roots in boxes with glass sides. In one series the roots were grown in holes or channels along the edge of the glass, and in the other series no holes were provided. In both series there would be contact, but where the roots followed the channels, there was less contact of the roots with the soil particles than where they had to force themselves through the soil. These experiments being limited, however, did not furnish data of any particular value, although from the more or less meager data obtained, they seemed to indicate that the presence of grooves or channels in the soil produces less stimulating effect on the growth of roots than when more thorough contact exists. Most of our investigations relating to the effects of contact on roots were made with excelsior, and in some cases sphagnum moss and cocoa fibers were used. In many experiments we have also utilized various- sized particles of gravel, sand, and silt suspended in water. The excelsior which we employed did not produce any bad effect on the growth of roots when submerged in water. On the other hand it appeared to clear up the water in some instances, inasmuch as it was apparent that it absorbed certain accumulated products which sometimes occurred, and which were more or less detrimental to the growth of the roots. The water containing the excelsior cultures was clearer and possessed a more agreeable odor than those cultures where it was not used. As the seeds were suspended over water on a fine-mesh cotton cloth netting, these would occasionally become too moist and in this way the water became more or less turbid, owing to the presence of various extract substances derived from the same. The presence of excelsior in the water had a tendency to prevent any abnormality in the growth of the roots which might follow from the presence of foreign substances, and in this respect it acted like sand and charcoal in removing certain impurities such as copper sulphate. STONE: CONTACT STIMULATION 469 TABLE 16 Showing Growth of Roots of Lupines (Lupinus luteus L.) in Water Containing Excel- stor and Sphagnum Moss Average Length of Primary Roots (Centimeters) Treatment Roots IN GEitial os tno coe Ae 7.3 Spharnummosse eee ae ee 8.8 Excelsioncoarse.4-6 see 10.4 For the purpose of obtaining more uniform conditions for root growth, we subsequently adopted larger containers provided with loose partitions, in which case all the plants were subject to like con- ditions. The excelsior experiments were made in either glass or earthen jars with tap water which had been previously boiled to exclude the air. The excelsior, which was of the ordinary commercial form, was in long strands of more or less irregular shape and in cross section was about I mm. in diameter. We employed three different grades, namely, the coarse, loose, curly form as it is obtained com- mercially, and the same cut from I cm. to 4 cm. long. The uncut grades were packed loosely in the jars containing the water, while the finer grades were much more compact and greatly increased the contact surface to which the roots were subjected. In all cases the excelsior was boiled before using it to exclude air, inasmuch as the presence of air would be capable of modifying growth. The seeds employed were of a good quality and were separated by sieves and carefully selected before planting, which insured a uniform size and TABLE I7 Showing Growth of Stems and Roots of Peas (Pisum sativum L.) in Water Containing Excelsior. Average of Four Experiments Average Length of Stems and Primary Roots (Centimeters) Treatment Stems Roots INvaynaierl ie 2 See Se rete ware ile 4.5 Excelsiot coarser rin ean LOr 7.0 xcelsion fine sae ete ae er Fier 8.8 Secondary roots predominated in the fine excelsior, many in the coarse excelsior, practically absent in the normal. corresponding germinating capacity. The experiments were con- ducted in a dark place, although in this series not in absolute darkness. The amount of light, however, which prevailed was insufficient for photosynthesis or for phototropic curvatures. Neither the stems nor roots were in contact with one another, hence, any stimulating growth 16 5 plants used in each test. 17 One hundred and sixty plants used. 470 BROOKLYN BOTANIC GARDEN MEMOIRS which followed was due to the excelsior alone. All of the plants were in the seedling stage and the duration of the experiments in no case exceeded fifteen days. The investigations relating to contact stimu- lation of roots were limited to the use of a few species which show different types of reaction. The results of these experiments are shown in Tables 16-18, although several other species were employed which are not included here. In all cases, the ‘“‘excelsior coarse’”’ implied the loose commercial form which came in contact with roots occasionally, while ‘‘excelsior fine’”’ was in these particular experiments, cut up into lengths 4 cm. long and furnished considerable contact. When such plants as lupines and peas which possess strong and fairly good-sized primary roots were in contact with excelsior the reaction was characterized mainly by an accelerated growth of the primary root system, together with considerable secondary root development whereas in the case of delicate roots, such as mustard (Table 18 ), the growth of the primary roots is less and the secondary root development is greatly accelerated. Practically little or no difference existed in the growth of hypocotyls and stems in any case, although this feature is not always constant. In both the lupines and peas (Tables 16 and 17) there occurred a considerable accelerated growth of the primary roots. In the latter case (Table 17), which represents an average of four experiments, there was a decided increase in the number and length of secondary roots from the normal to the fine excelsior. In one experiment where the secondary roots were counted, their average number was as follows: INOnitial ee eek seer Ta ee Ce eee ons Deeg 17 Coarse excel SiO ts Pie ee oe ee ESE: Cuciae “roenenaeac 32 RUNeLeXCelSIOl a eee ee cicre ota 40 This feature was more pronounced in the mustard, however, where the primary roots in contact with the excelsior were much less de- TABLE 18 Showing Growth of Hypocotyls and Roots of Mustard (Brassica alba Botss.) in Water Containing Excelsior'® Average Length of Hypocotyls and Primary Roots (Centimeters) Treatment Hypocotyls Roots INorinial. ss ais. Sons ea 12a Tad Excelsionicoarsen ian 13.3 6.8 Excelsior iinereenn an eeeeeace 11.9 8.9 veloped (Table 18). In this case there were no secondary roots on the normal plants; many in the coarse excelsior and very numerous 18 20 plants used in each test. STONE: CONTACT STIMULATION 471 and well developed on those in contact with fine excelsior. The number of secondary roots or those in contact with the fine excelsior averaged 18 to a plant and the ratio of the total length of the entire root system of the normal and fine excelsior was I to 6 in favor of the latter. Measurements made of the total surface area of a single typical root from one of the normals and one of the fine excelsior showed that the total surface area of the latter was over three times that of the normals. These experiments show, at least in young seedlings, that roots respond to contact and that the response is confined very largely to these organs, although more than one type of growth correlation may occur. They indicate also that different species will respond to contact in a different manner. In other words, secondary root development is stimulated more in some species than in others by contact, and this excessive development of the secondary root system is correlated with a lesser development of the primary root system. Soil Particles Since roots are sensitive to contact to various materials it would naturally be supposed that the nature of the soil constituents or particles would exert an influence upon growth and configuration of plants, and particularly upon the root itself. As contact is effected by the surface area involved, variation in the size and shape of the soil particles would be expected to produce different results. Conse- quently, a series of experiments were carried on, but not completed, with the idea of determining what effect, if any, soil particles have on the growth of roots, and how the various-sized particles effect development. For this purpose we had at our disposal several care- fully prepared grades of gravel, sand, silt and clay which had been sifted through sieves and bolting cloth. The size of the particles ranged from 16mm. to0.I mm.and in some cases to.05 mm. The ex- periments were conducted in glass jars filled with water previously boiled to exclude air. Each jar was completely filled with some particular grade of material which had previously been thoroughly washed with water and sterilized. We thus had a medium in which the particles of gravel, sand, etc., were surrounded by water, and as far as possible free from air. A fine-mesh cotton netting was placed over the jars on which rested the seeds, and as germination took place the radicles penetrated downward between the submerged soil particles. All experiments made with soil particles in water were carried on in darkness in a moist chamber where transpiration was limited and the temperature condition alike. The plants were in fact under identical conditions throughout, except as regards the substratum. 472 BROOKLYN BOTANIC GARDEN MEMOIRS The normal or check series were run in water alone. The results given by these experiments, which were limited in numbers, are similar to those obtained by the use of excelsior in contact with roots in water, namely, the various grades of gravel, sand, and silt gave rise to different reactions on the part of the organism which resulted in a stimulated growth correlation, as exemplified in the development of the primary organs and secondary root system. The results ob- tained by the use of mustard, peas and soy beans, etc., in the experi- ments so far carried on, indicate generally that there exists little difference in the height of the hypocotyls or stems of the normals, and those in contact with the different-sized particles of soil constit- uents. The growth in length of the primary root is lessened and the secondary root system is greatly increased as we approach the finer grades of contact material. In some species with relatively large roots the coarser particles, namely 16-8 mm., appear occasionally to stimulate primary organs, but this reaction is variable, and associated with growth correlations as manifested in the more or less increased development of the secondary root system. The reaction of roots to different-sized particles will undoubtedly be found quite variable. Species with relatively large and strong primary roots such as the horse bean, peas, etc., react differently from seedlings with a delicate root system, such as the mustard and turnip. The roots of the latter species, even when grown in contact with excelsior or soil particles, produce a marked secondary root system characterized by little or TABLE 19 Showing Growth of Hypocotyls and Roots of Mustard (Brassica alba Boiss.) in Contact with Different-sized Particles of Sand and Gravel in Water'® Average Length of Hypocotyls and Primary Roots (Centimeters) Size of Particles (Millimeters) Hypocotyls Roots INormal eee eat Soo 8.7 1247 TO—Sieees Peas eo ee oes eioe 8.2 12.6 SoA ret goer et Aes Rade ite) 8.7 Baoan Rar rcesuaie ence see ema ener 11.5 7.6 2 TARAS See SAR eRe AG EN 11.9 4-4 DONS ties Sele ane Cee eee 11.6 4-3 OF5=O0 25a eer cee ele TL:2 4.2 OP25 SO 81t ere ten aeereteat forte 1257, 3.4 no primary root development. The same tendency exists in species characterized by larger roots to increase their secondary root system. On the other hand, the primary root system is greatly accelerated by contact with excelsior in species with large roots. (Cf. Tables 16-17 and 18.) The experiments with mustard given in Table 19 show 19 75 plants used in each test. STONE: CONTACT STIMULATION 473 much difference in the growth of hypocotyls and roots in the different grades. This feature is associated with the greatly accelerated development of the secondary root system, and extending from the coarser grades to the finer ones. Other than the production of secondary roots in the plants in the 16-8 mm. grades there was little difference between the growth of the latter and those of the normal. There were no secondary roots in the normal grades or water cul- ture plants in this case, although they were fairly well established in the 16-8 mm. grade, from which grade the increase in numbers and total length of the secondary roots were quite noticeable. The average development of the primary and secondary organs as well as the surface area of the same was greater in the contact plants than in the normals. Soil particles and excelsior submerged in water have a similar stimulating effect on mustard as will be seen by comparing Tables 18 and Ig. THEORETICAL CONSIDERATIONS While the general tendency of plants and plant organs is to avoid contact with one another, the histological units or cells which com- posed the individual are in contact with one another, and the same holds true to a certain extent with different organs when in the em- bryonic or bud stage. It is the exception rather than the rule to find the various members or organs of different plants, or even those of the same plant such as roots, branches, leaves, etc., in contact with one another, or in other words it appears to be a universal law in nature that the various organs of plants occupy space by themselves. Uniformity and regularity in the arrangement of cells and organs is more common to the lower than to higher organisms, since in the higher organisms this feature is sacrificed to some extent by biological necessity and adaptation. Primarily the arrangements of organs in plants or angles of divergence are determined by laws which are common to gravitational and electro-magnetic phenomena, and the arrangement of the various organs of plants appears to be determined by the action of these forces upon their ultimate structural units, molecules, micellae, atoms, electrons, or whatever they may be. The angles which various organs assume in plants closely resemble those which are illustrated in the formation of certain types of crystals, and the behavior of iron filings under the influence of a magnet. Plants are susceptible to all of the common environmental influences which surround them, but the modus operandi of these various external agencies on protoplasm is little known and especially concerning the mechanism and nature of conductivity of impulses. The reaction to contact results from a mechanical impulse, inasmuch as when the 474 BROOKLYN BOTANIC GARDEN MEMOIRS external cells of the organism come in contact with solid particles a reaction follows. To affirm, however, that the stimulus is mechanical in nature does not explain anything, because we know nothing of the nature of the so-called mechanical impulse. The bombardment of organisms with electric waves (negative electrotropic response) may be of the nature of a mechanical impulse and such may hold true for other forms of radiant energy. Phototropic, thermotropic, electrotropic and contact stimuli may not materially differ from one another in the nature of their action on organisms, that is, in the sense of acting ina mechanical manner on plants. The stimulation resulting from such contact is apparently transmitted to the living zones which induce definite adjustments or coordinations of the vital processes, the nature of the response being determined by the nature of the stimulus involved and the organs stimulated. The external cells of a leaf, for example, coming in contact with an object would probably react to the stimulus through the cuticle and exterior cell walls. The reaction of plants to contact is probably one of the most primitive forms of responses and quite universal in the vegetable kingdom.” The nature of the response to contact resembles mostly that of touch or at least a primitive and rudimentary form of this sense. There are several types of contact stimulation that have long been recognized such as occur in the response of tendrils, tentacles, stamens, etc., when brought in contact with different substances. These reactions are associated with different types of irritability. Most of the known reactions to contact such as are illustrated by tendrils, etc., are closely associated with biological adaptations, and as such they have been subject to considerable modification. It is not at all improbable, however, that these various types of reactions are modifications and differentiations of a more simple and_ universal form of contact response. The various forms of response movements which are associated with irritability have been classified under the so-called tropistic, nastic, and taxic reactions. The first type of response is characterized by the movement of organs toward a definite position in response to a stimulus, while nastic reactions are those which are independent of the direction of the stimulus. Taxic reac- tions are those shown by motile organisms. There are several different types or manifestations of irritable phenomenona such as_ photo- 20 Some experiments made with mucor in Petri dishes containing beef extract and fine particles of quartz sand showed contact responses. Both series of Petri dishes contained the same amount of beef extract, but in one the fine sand was added for the purpose of obtaining contact with the mycelium. The sporangia were black and mature in the Petri dishes without sand, while in the dishes containing the fine sand, the aerial hyphae were much more elongated and the sporangia light in color and less mature. STONE: CONTACT STIMULATION 475 tropism, geotropism, etc., which are characterized by both positive and negative reactions, but for our purpose it is necessary to consider only a few which are somewhat similar in character, and which have not unlikely originated from a simple fundamental type. (a) Contact Stimulation Probably a universal type and one of the most fundamental forms of response in plants, which is characterized by a stimulated growth of various organs due to contact of one organ with another, or with various substances. The extent of the reaction is determined by the degree of contact, and probably influenced by the nature of the contact substance. (b) Haptotropism Illustrated by various tendrils reactions; tentacles of Drosera. (c) Haptonastie Illustrated by certain types of contact of some tendrils and tentacles of Drosera.” (d) Seismonastie Illustrated by movement of stamens of Centaurea, Berberis, and Mimosa leaves. (e) Thigmotaxis Reaction of motile organisms (Chlamydomonas) resulting from contact of cilia with different substances. (f) Wound Stimuli and Reactions Wound stimuli in general caused by mutilation, presence of organ- isms, and direct contact of plant members with one another induce various reactions, the nature of which depends upon the host plant, and nature of the cause responsible for the same. The general response to wound stimulation is manifested by the production of abnormal growth due to active cell division, and in some cases enlarge- ment of the cells. These reactions are characterized by local effects and the reactions are often disproportionate to the stimulus. In some types of reactions there would seem to be involved more than one class of phenomena. For example, the haptotropistic reac- tions of tendrils, or at least the subsequent effects involved in the formation of the spiral, etc., by the stimulus, and which is manifested 21 Stark (3) has observed that many plants and some roots react to contact stimulation similar to tendrils when rubbed with a stick or cork. 476 BROOKLYN BOTANIC GARDEN MEMOIRS by the production of mechanical tissue would seem to be similar to that induced by other irritable phenomenona. In the formation of the spirals in tendrils there is considerable stress to be overcome and the constantly increasing weight of the various members of the plant, such as is associated with the development of foliage, fruit, etc., together with the effects of rain, swaying movements due to wind, is most admirably met by the production of mechanical tissues. These additional strains are taken care of by the same type of response as that induced by the stretching of plants with weights. Since the leaves of plants appear to be quite sensitive to contact it is readily conceivable how the more specialized haptotropistic responses to contact exempli- fied by tendrils could readily be evolved from simple contact irritability. There are, however, a large class including various types of reaction phenomena which do not fall under tropistic, nastic and taxic responses. These are the so-called wound reactions—traumatotaxis (reaction of cell nucleus). Traumatotropisms, illustrated by decapitated root, and positive galvanic currents on roots, are apparently direct responses to wound stimuli, as probably are the reactions first observed by Darwin resulting from the attachment of different substances to the root tip. In wound reactions, however, contact is involved to a greater or less extent, and more or less injury and abnormal conditions are associated with this class of phenomena. Most of the responses following wounds are local in their effect, although the organism as a whole may be affected even from relative insignificant mechanical injuries, as shown by the modification in the developments and functions of the several organs in mutilated plants. Representative types of this class are seen in the various accelerated growths produced by insects, fungi, bacteria, mechanical injuries, etc., and generally the reaction continues long after the primary or excitory cause has disappeared. Local accelerated growths, however, do not always follow as a result of the intrusion of pathogenic organism inasmuch as the nature of the response is determined to a large extent by the character of the tissue affected. For example, eel-worm infestation of roots may give rise to galls, whereas on stems such a response may not necessarily follow and even on roots these reactions differ. In many so-called wound reactions the degree of response is disproportionate to the stimulus responsible for the same. This is illustrated by feeble lightning discharges on trees, in which case the stimulus (lightning causing burning) lasts but a few thousandths of a second. The reaction, however, to such insignificant injury—often hardly perceptible and characterized by the destruction of a few cambium cells, may manifest itself for years in an accelerated growth of the annular rings adjacent to the injury. The flow of tissue in STONE: CONTACT STIMULATION 477 sucker growth around stumps, the enclosure of twigs, grass, etc., by mushrooms, and the imbedding of tree guards illustrate stimulated growth responses not generally associated with wounding. When more or less long stubs remain following the destruction of limbs on trees, they act as a stimulus, and large masses of tissue ultimately accumulate around the base of the same. If, however, the stub is removed close to the tree, healing follows, and when the callus unites growth becomes normal. Again the feathery growths, consisting of small twigs which are common on the trunks of elm trees, give rise to the same type of reactions, namely, the formation of excrescence or tuberous growths at their point of junction to the tree. The presence of blocks of wood on trees to prevent the contact of guy wires with the bark often stimulates the flow of tissue, and while there is appar- ently no injury associated with this connection there exists a more or less severe tension or pressure on the bark. The tendency of tissue to grow over even loosely attached tree guards and wire attach- ments, even when it would require but little force to dislocate them, is universal. Signboards placed on trees, although often insecurely attached, are sometimes entirely obliterated by a greatly stimulated callous growth, and tree guards loosely attached to trees often become imbedded in the tissues, when it would require but little exertion on the part of the underlying tissue to dispose of the same. It may be assumed, however, that the presence of nails driven into the wood for the purpose of holding the sign in place would act as a stimulus, but the same reaction follows if the nails are not present and the sign merely held securely by means of wires. Moreover, the growth response here conforms to the sign as a whole rather than to any par- ticular point of attachment by the use of nails, etc. Similar response occurs in the growing of roots around large masses of gravel in the soil or when limbs or roots come in contact with one another, in which case natural grafts may occur. The tissue, however, exterior to that which is in contact is often destroyed in natural grafts, etc., which would have the same effect as wounds; although even here the growth stimulation is much greater than that resulting from mere wounding. In this type of response there are two factors involved, namely con- tact and growth acceleration. The reaction, however, is local and does not differ materially whether injuries occur or not, as is shown by growth stimulations arising from other causes, such as frost cracks, etc. There are other similar types of reactions apparently differing somewhat in the nature of the response to contact from those enumer- ated, namely, those arising from restricted growth. Contact in such cases occurs, but injury may be absent, although abnormal growth may result. This type seems to be characterized in their manner of re- sponding by a greatly increased osmotic tension of the cells. 32 478 BROOKLYN BOTANIC GARDEN MEMOIRS When roots are growing between large boulders and are restricted in their development, the flow of tissue is such that the root may become enormously flattened. The reaction in such cases is similar to that produced by a wound, although evidence of wounding may be entirely absent. Such roots, however, often lift enormous weights which would indicate that considerable osmotic tension exists in the cells. When rapidly developing organs, such as a squash, for example, is placed in a harness and subject to weight, it will assume a much distorted shape which illustrates stimulated growth phenomena. Again, when ferns and even delicate mushrooms push through con- crete they show little or no evidence of wounding, although the reac- tions in such cases are characterized by a large increase in the osmotic tension of the cell, sometimes equaling 50 atmospheres (4). The experiments of Pfeffer (5), in enclosing roots in plaster casts, thus restricting growth and greatly increasing the osmotic tension of the cells, are typical of this class of responses. To what extent, if any, cell fusion of sexual elements, development of attachment organs in fungi and algae, and outgrowths in spirogyra filaments when in con- tact with certain crystals, are related to contact, is problematical since in some instances chemotropic phenomena would have to be carefully differentiated from any other which might prevail in interpreting such phenomena. Also to what extent contact stimulation may influence cell enlargement and cell division resulting from the intrusion of foreign elements would be merely a matter of speculation at the present time. There are, however, numerous instances of cell enlarge- ment associated with crystals (raphides), pathogenic and non-patho- genic organisms, which resemble contact stimulation, although the recent important contributions on this subject by Dr. E. F. Smith (6) would indicate, in some cases at least, that cell responses to pathogenic organisms are associated with chemical or physical phenomena. Some of the responses of plants associated with pathological phenomena are not characterized by cell stimulation, but with color reactions due apparently to excretions from organisms. Intense color reactions are also associated with marked cell proliferation caused by chemical substances absorbed by plants from the soil, as shown by the reactions of Platanus Orientalis to the toxic properties contained in illuminating gas. These reactions, however, as are similar ones in poplars and wil- lows, which develop large masses of parenchyma under the bark and cause rupture of the same, are associated with two factors, namely, the direct effect of the toxic substances on the cambium inducing rapid cell division, and decrease in the tissue tensions of the cortex following the collapse of the same by poisons. Various chemical substances (banding substances) applied to trees produce local growth STONE: CONTACT STIMULATION 479 acceleration, but these apparently affect the outer tissues, and to a certain extent the underlying vital layer. Dr. H. von Schrenk (7) and Dr. E. F. Smith (6) have shown that ammonia compounds cause intumescences in cauliflower. The absorption of the chemical sub- stances by the roots in some cases produces a rupturing of the epidermal tissue which is followed by cork formation, and in cases of malnutri- tion excrescences are often formed on the fruit. Finally, contact stimulation has an important bearing on experimental work, especially with that done with plant food, fertilizers or soils, inasmuch as stakes and wires are often employed as support. Any differences existing in the contact of the plants, either by the use of supports or that occa- sioned by the proximity of plants to one another, thus causing contact, would be effective in modifying the results. The relative height and development of two plants which would affect the contact surface would also be important factors in experiments of this nature. Indeed in this respect we have noticed on more than one occasion experiments where differences existing in the degree of contact would account for all the variations in the growth of the plants that were supposed to be due to other causes rather than to the particular treatment which they received. LITERATURE CITED 1. Hall, A. D., Brenchley, W. E., and Underwood, S. M. The Soil Solution and the Mineral Constitutions of the Soil. The Jour. of Agr. Sci. 6: 278-301. Sept. I9I4. 2. True, R. H., and Oglevee, C.S. The Effects of the Presence of Insoluble Sub- stances on the Toxic Action of Poisons. Bot. Gaz. 39: I-21. 1905. 3. Stark, Peter. Ber. d. d. Bot. Ges. 33°: 389. , 4. Stone, George E. The Power of Growth Exhibited by Ferns. Bull. Torrey Bot. Club 36: 221-225. May 1909. 5. Pfeffer, W. Druck und Arbeitsleistung durch Wachsendepflanzen, p. 93. 6. Smith, E. F. Mechanism of Tumor Growth in Crowngall. Jour. of Agr. Res. 8: Jan. 1917. 7. Schrenk, H. von. Intumescences Found as a Result of Chemical Stimulation. Ann. Rept. Mo. Bot. Gard. 16: 125-148. 1905. DUPLICATION AND COHESION IN THE MAIN AXIS IN CICHORIUM INTYBUS AVeD. SLOuUr New York Botanical Garden The terms duplication and cohesion may be used to designate a very special type of fasciation which the writer has observed in the variety of chicory cultivated under the name ‘‘red-leaved Treviso.” The most distinctive characteristic of this type of fasciation is the duplex nature of the main axis. From a single root a double stem arises with the two parts, as a rule, strongly united. The duplex nature is indicated by a pair of grooves which extend in the direction of the long axis and round out the two stem-elements. This is quite well shown in number 5 of the accompanying plate XII. This type of fasciation differs from the banded and cone types most usually seen in that here two stem-elements of equal size and rank are clearly in evidence throughout a segment of unequal diameters, giving a symmetry that is bilateral. Also the fasciation is confined to the middle and lower portions of the stem and decreases upwards, the main axis often becoming at its apex quite symmetri- cally simplex. I have been unable to find in the literature any reference definitely mentioning fasciation of this particular type. It appears not to have been noted and described even in this rather well-known variety of chicory. In the degree and the extent of the duplication there is much variation. The most extreme condition of duplication is seen in the complete separation of the two stem-elements with each perfectly formed and without lesions, as is shown in no. 7 of Plate XII. In many cases the two elements are indicated only by grooves which extend from close to the base of the plant to a height of about three feet. The length of the grooves and the corresponding segment of evident duplication may, however, be reduced to mere traces, as is shown in no. 3 at a or in no. 4 from a to 8, as indicated. In a few plants of this strain there is no evidence of duplication and the phyllotaxy is of a single and regular spiral only. Each stem-element has its own phyllotaxy in so far as this can be expressed. In the most pronounced duplex condition there are clearly two separate spirals in evidence. The phyllotaxy of one is not a’ 480 STOUT: DUPLICATION AND COHESION 481 continuation of the other and more branches and leaves are produced upon the two than upon a single stem. Branches and leaves occa- sionally develop, however, from the very center of the grooves and the bases of leaves thus placed often extend across a side of both stem- elements. Such leaves and branches are always simple as far as I have observed. In the uppermost portions of the stem the phyllo- taxy is often completely regular, but immediately above the seg- ment of noticeable duplication the phyllotaxy is irregular and this irregularity may extend to the apex or terminate in the banded type of fasciation. Torsion is frequently seen. The two elements may twist about one another in spirals with the two parts equal in development and the spirals in the same direction. In other cases the two stem-ele- ments exhibit incompatible growth either as to direction of torsion or rate of growth, or both, and mechanical lesions result. These tear apart the two elements or often cause one to break, thus producing much irregular and twisted development of the main axis. Various conditions which thus arise are shown in Plate XII. Usually the lateral branches above such lesions are poorly developed (no. 9), the more vigorous growth being seen in the lower laterals. In this strain of red-leaved Treviso, irregularities of development are seen in the growth of the cotyledons. For most seedlings two quite normal and separate cotyledons are produced as shown in nos. I, 2, and 4of Text Fig. 1. In numerous cases even of sister plants the two cotyledons are fused; only the basal portions of the stems may be fused as in nos. 5, 6 and 15, or the fusion may extend upward toward the apex as shown in nos. 7, 8 and g. In some cases what appears to be only one cotyledon is in evidence, as shown in nos. 10, II, 12 and 16; either the two have become completely fused or the growing point of one failed to develop. The very complete series of stages of fusion leading to a single cotyledon suggests that the initial growing points of the two cotyledons may completely fuse or perhaps are never differentiated. There are also numerous cases of crumpled and irregularly developed cotyledons either in one or both of a pair or in a single one, as may be seen in HOS. 3,53; 14, £5 and 17. In only two cases have I observed any tendency to an increase of cotyledons; in each of these seedlings one of the cotyledons was some- what bisected, as shown in nos. 19 and 20. The duplication seen in the main axis is not accompanied by duplication or increase in the normal number of cotyledons. In a very few instances there is no development of the plumule. This may occur in seedlings having two normal unfused cotyledons as 482 BROOKLYN BOTANIC GARDEN MEMOIRS shown in no. 4, as well as in seedlings having only one cotyledon, as shown in nos. 16and17. Such plants usually die soon. In some cases, however, adventitious buds arise, but as far as observed these make only a feeble growth. The drawings 16) and 17) show such feeble adventitious growth of seedlings nearly three months old, at which time many sister plants had rosettes measuring ten inches in diameter. af nA es a a Fic. 1. Young seedlings of the variety red-leaved Treviso. Scale about one- half natural size. 1 and 2. Two normal separated cotyledons with plumule. 3. Two cotyledons, both somewhat crumpled. 4. Two cotyledons; no plumule. 5. Cohesion at base of cotyledons only. 6. Cohesion at base of cotyledons; one crumpled. 7, 8,9. Decided cohesion. 10, II, 12. Only one cotyledon present. 13. Two cotyledons; separation involved some lesion. 14. Much crumpled and poorly developed cotyledons. 15. Some cohesion at base. 16a. Single cotyledon; no plumule. 6. Same seedling two months later with adventitious bud. 17a. Single much crumpled cotyledon with seed coat attached at apex; no plumule. 6. Same seedling two months later. 18. Much fusion of cotyledons. 19, 20. Decided lobing of one cotyledon. The growth interrelations of reduction or fusion of cotyledons with duplication in the main axis are not clear. In the former there is a fusion or reduction in the number of organs normally formed; in the latter there is a tendency to the production of two main stems STOUT: DUPLICATION AND COHESION 483 instead of one, accompanied by their cohesion or incomplete separation. In both the factor of cohesion is present, but in the cotyledons it leads to reduction in the number normal to the species, while in the stem it tends to reduce the duplication to the single stem-element that is normal to the species. The occurrence of the sort of duplication described above has now been observed in four generations. Thirty-five plants of the 1913 crop of the variety red-leaved Treviso were grown from commercial seed. In 14 plants the main axes were decidedly duplex, the length of the segments showing duplication ranging from a few inches to about three feet. The stems of the other 21 plants showed no sign of duplication. All plants of this generation which were tested were found self-sterile from physiological incompatibility, so it was not possible to obtain self-fertilized seed. Numerous crosses were at- tempted between plants, but only one cross attempted was compatible. In 1914, 12 plants were grown as progeny of the cross between two plants which were quite alike in exhibiting duplication of rather inter- mediate development. In all 12, duplication developed. The vari- ability in degree of duplication was marked and ranged from very slight indications to very decided cases of torsion and lesion. One plant developed too late in the season to be tested for self-fertility, but the other 11 were all self-sterile. However, some compatible crosses were made from which 43 plants were grown in I9QI5. Of the 1915 crop, 39 of the 43 plants exhibited duplication to some degree. In one plant the two elements were completely separate from the very base upward (no. 7). In several plants torsions and lesions were strongly developed. Four plants were apparently not in the least fasciated and in each of these the phyllotaxy was regular and single. In the 1916 crop, 150 plants were grown. Two of these were froma self-fertile plant (the only one of the 1915 crop that was found to be self- compatible in any degree). The 148 other plants of this generation were from six different crosses involving eight different parent plants, all - of which, however, exhibited duplication in some degree. This genera- tion was descended from three generations of parentage that exhibited duplication. There was duplication to some extent at least in 144 of these plants. Six appeared to have a single stem-element with regular phyllotaxy; these six plants were distributed among the offspring of three different crosses. One plant of the 1916 crop exhibited a noticeable fasciation of the ribbon type in the upper branches in addition to duplication and cohesion in the main axis. The tips of the main branch and of various laterals were broadly flattened and the branching was reduced so that 484 ~ BROOKLYN BOTANIC GARDEN MEMOIRS flowers and short spur branches were clustered at the extreme ends, giving a peculiar rosette-like appearance. The two types of fasciation were quite distinct on the plant, both as to general appearance and location. From the performance of these pedigreed cultures, it seems clear that the character of duplication and cohesion persists in successive generations of this variety of chicory to such an extent as to appear strongly heritable. It is not completely so, for a few normal single- stemmed plants do occur quite irregularly in various generations and lines of descent. The heredity of the character of duplication has also been tested by crossing plants of the 1915 crop showing typical fasciation with a plant of wild stock (plant A) which had a short, slender, main stem. No tendency to duplication has been seen in plants of wild stock which have been grown or in plants of various generations derived by crossing the wild plant A with plants of the cultivated variety known as Barbe de Capucin. This cross here in question involved, therefore, on one side parents with duplication, and on the other a plant of a stock free from duplication. : Thus far 81 plants of an F; generation have been grown; nineteen of these had the wild plant A as a seed parent. Of these 81 plants only three possessed the grooves which are seen in most typical cases of duplication of the main axis. In only one of these were the grooves pronounced (see no. 10); in the other two there were only slight indi- cations of grooves (see no. 11). In 78 plants there was not the slightest indication by grooves of any duplication. However, in tracing the phyllotaxy from base upward, irregularities were seen in 48 plants. Two leaves or branches were often opposite or the direction of the spiral would appear to shift from left to right or vice versa. In 30 plants the spiral of the phyllotaxy proceeded very regularly from base to tip in a way that indicated a normal single stem-element (see no. 12). Of the 19 plants having the wild plant for a seed parent, 7 were quite normal and in 12 there was a mixed or irregular phyllotaxy. Judged by performance in the F, generation, the character of duplication is only incompletely and partially dominant. An inter- mediate type is frequent in which the only suggestion of a duplex con- dition of the main axis is seen in an irregular phyllotaxy. I cannot at the present time contribute any information regarding the sources, Causes, or nature of the stimulus operating in duplication nor any definite facts regarding the attending anatomical development. When this paper was presented it was suggested by Dr. Erwin F. Smith that possibly infection by some organism, bacterial or otherwise, was necessary to the development of duplication as here observed. If this on "IX aLvid ‘| 3WN10A AUOSIHD NI NOILVIOSVY + LOOLS “SUIOWSIN NaGYVv OINVLOG NATMOOY@ STOUT: DUPLICATION AND COHESION 485 should be the case it is evident that while the susceptibility is very decidedly limited to the variety it can be transmitted directly or indirectly though incompletely to large numbers of the offspring of a hybrid generation. SUMMARY An unusual type of fasciation occurs in the variety of chicory known as red-leaved Treviso. It consists of a very decided duplication in the main axis of the stem, giving two stem-elements with, however, a decided cohesion of the two. In this variety of chicory there are also various irregularities in the development of cotyledons and plumule. All degrees of fusion between the two cotyledons are in evidence; in some seedlings only one cotyledon is present. Occasionally no plu- mule develops. The character of duplication and cohesion of the main axis is strongly but not completely heritable. There is wide variation in the degree of duplication and a few plants with a normal unduplicated main axis occur. In an F, hybrid generation of crosses between plants with dupli- cation and plants of wild stock which exhibit no tendency to such fasciation the character of duplication is incompletely dominant both as to degree of expression and number of plants affected. An inter- mediate type is strongly in evidence in which the only indication of duplication is seen in a mixed and irregular phyllotaxy. EXPLANATION OF PLATE XII Nos. I-9 inclusive are of stems of plants of the variety red-leaved Treviso; no. 7 is of the 1915 crop, all others are of the 1916 crop. No. 1. Stem showing no duplication. Phyllotaxy regular. No. 2. No duplication but stem thicker than in no. 1. Phyllotaxy regular. No. 3. Stem of small plant. Short segment of duplication at a. No. 4. Duplication from a to b; phyllotaxy above a is irregular with tendency for branches to be paired. No. 5. Very decided duplication from a to 6. Lesion separates the two stem- elements in lower center. Stem-elements parallel below but twisted above. No. 6. Decided duplication with parts much twisted. No. 7. The two stem-elements separate from the base. No lesions. No. 8. Longitudinal lesions strongly developed. One stem-element much contorted. No. 9. Much torsion with transverse lesion of one stem-element. Nos. 10, 11 and 12 are F, hybrids of red-leaved Treviso X unfasciated plant of a wild variety. No. 10. Duplication evident in lower two thirds of stem. No. 11. Duplication indicated by a very slight but broad groove near base. Phyllotaxy mixed and irregular especially near base. No. 12. No duplication. Phyllotaxy regular. A QUANTITATIVE STUDY OF RAUNKIAER’S GROWTH- FORMS AS ILLUSTRATED BY THE 400 COM- MONEST SPECIES OF LONG ISLAND, N. Y. NORMAN TAYLOR Brooklyn Botanic Garden The value of sorting species of plants into different categories, based on their growth-forms, has been pointed out so often that there scarcely seems further need of going over the subject again. The weakness of such a sorting and the percentages based on it, due to the fact that species, not individual plants, are considered, is ob- vious. Such percentages as have been published show not so much an actual response to climatic factors, as they do the multiplicity of forms that may have been developed. For most regions that is all that can be done, as anything like a plant census of a given region is usually impossible. Yet upon such a census, or some approximation to it, there could be based percentages of different growth-forms that reflect more accurately than any species percentage the actual climatic response of vegetation to climate. The importance of getting, if possible, some growth-form per- centages that should be quantitative rather than those based on species only resulted in a study of the flora of Long Island, N. Y., with this in view. The island is roughly 120 miles long and 12-16 miles wide and, excluding ferns and their allies, has about 1120 species of native plants. It is diversified as to vegetation, as there are good- sized areas of ‘“‘scrub,” mostly oak and Ericaceae, considerable de- ” ciduous forest, some extensive ‘‘pine-barrens,’’ salt marshes, a small prairie, and the downs at Montauk and Shinnecock. In a general study of the flora and vegetation of the island, distri- bution maps for each of the native species were made and have been posted up for several years. Such maps indicate actual collections represented by specimens in herbaria, field notes by the writer, all published records of species and descriptions of different vegetative areas by nearly all who have written about Long Island for the last 250 years. From Daniel Denton’s History of New York, through the period’ when numerous Quaker journals were issued, down to the modern observations of professional botanists, these records have been accumulated. The opportunity, therefore, of getting something 486 TAYLOR: RAUNKIAER'S GROWTH-FORMS 487 like a comprehensive view of the flora and vegetation of the island is excellent. Within the last few months it has been possible to separate the distribution maps of the species into two groups. The one which interests us just now is the smaller, consisting of the 400 commonest species. These make up the great bulk of the vegetation of the island, the other group of about 719 species being scattered and nothing like so common. When these four hundred species are sorted into the different growth-form categories of Raunkiaer and their percentages reckoned, we find them grouped as follows:! Numbers of Percentages of the Growth Form Species Commonest Species INGE ely tare h tennis 6 1.50 IMS aie is oleae I2 3.00 IVD Gap ete Abaca PA ES cosas 2's 34 8.50 IS Riese ek ete eeiatace Ra 17 4.2 OL roe nae 29 7.25 LE Ron cee ene Pace ee Meee ee 120 30.00 Cis et ents Ri a ee rae 84 21.00 loll okey ee on Relea ae 27 6.75 LDS OR a re or en ty ea 57 14.25 Stemssucculents),,....-2...5.. 8 2.00 ATASILES He Ae bile bid Sa obs 6 1.50 The amount of deviation from the normal spectrum of Raunkiaer or from the growth-form percentages of the total flora of Long Island, or from the percentages of the local flora area? should show an inter- esting relation. The figures are as follows: |Mc|ms|mc| N|cu| H | Gc | HH | T | | Wocalifloray.2 im st ON n ses +52 4.03 17-18 |3.51 15.29 | 33-29 | 20.23 | 11.74 13.00 89 |4.37 |6.34 |2.77 |5. 89 | 33. 15 | 20.10 | 10.90 | 13.94 400 commonest Long Island | | SRECICS eso ohsincs biel 1.50 3.00 8. 50 4.25 7. 25 | 30. 00 | 21.00| 6.75 | 14.25 Normal spectrum.......... 6 | 17 | 20/9 | 27 | 5) I 13 It will be seen from this table that the percentage of large and medium-sized trees, the herbs that root near the surface, and the annuals are somewhere near what the normal spectrum would lead one to expect. In fact the growth-form percentages of these 400 commonest Long Island species are in remarkable agreement with the percentages of the total Long Island flora and of that whole region 1 Abbreviations for the different growth-forms are the same as those in general use. See Journ. Ecol. 1: 16-26. 1913 and Am. Journ. Bot. 2: 23-31. 1915. 2See Mem. N. Y. Bot. Gard. 5: 1-683. 1915 and Am. Journ. Bot. 2: 23-31. 1915. Stem succulents and parasites are omitted from now on as being too small to signify. 488 BROOKLYN BOTANIC GARDEN MEMOIRS near New York which is here called the local flora. To all those who have wondered how much an actual plant census of any region would derange the Raunkiaer scheme, these figures will come as a surprise. It has been supposed by some that wherever there was a serious dis- parity between the growth-form percentages of a region and the normal spectrum, the fact that species, not individuals, were being considered was obscuring the truth.® The fact that there is such a remarkable agreement between the percentages based on species and those based on frequency, and that both these sets of figures disagree radically from the normal spectrum, tends to increase the doubt as to the validity of the spectrum as laid down by Raunkiaer. In an earlier paper on the growth-forms of New York and vicinity, it was pointed out that “‘for no region in the world has there been published such a large percentage of these plants with bulbs, rhizomes, corms, and other subterranean methods of winter protection.’’ Considering that 20.23 percent of geophytes in the local flora area should have occasioned this remark and that for the 400 commonest Long Island species, the figure is 21 percent, the case for the normal spectrum which calls for only 3 percent of these geophytes seems decidedly weak. When it is remembered that in the normal spectrum our ordinary deep and shallow-rooted herbs call for only 30 percent, the aquatics I percent, and chamaephytes 9 percent, we have a total of only 40 percent for all herbaceous plants on the most favorable assumption. Actually many of the chamaephytes are low woody plants, so that the normal spectrum allows only about 35 percent for herbaceous plants of all kinds, excluding annuals, or 48 percent including them. The percentage for the same groups in the local flora area is about 79 per- cent, for all Long Island 83 percent, and for the 400 commonest species it is 78 percent. There can be here no question of the wrong assign- ment into the Raunkiaer growth from categories, for by lumping the chamaephytes (about half of which may be woody), hemicryptophytes, geophytes, aquatics, and annuals, we separate at once the woody from the herbaceous species. Does this difference of 30 percent in the herbaceous element of the vegetation of Long Island from that of the normal spectrum really mean that the region is so far off normal or that the normal spectrum itself is in need of further study? It has been shown that there is a rather definite relation between the percentages of herbaceous and woody elements in temperate and tropical floras, but unfortunately the figures as published deal only with dicotyledons.t. For our purposes, however, they show the per- s'Am: Journ. Bot. 2330: “1915; 4Sinnott, E. W., and Bailey, I. W. The origin and dispersal of herbaceous angiosperms. Ann. Bot. 28: 566-567. I914. TAYLOR: RAUNKIAER’S GROWTH-FORMS 489 centages of herbaceous plants based on the calculations of Sinnott and Bailey. Their table is as follows: ‘TEMPERATE REGIONS ay see a geese «| eae Northeastern United States (Gray)................ 2,280 1,748 1G | Northern United States (Britton and Brown)....... 2,662 2,089 78 Southeastern United States (Small)............... 4,608 Baie 72 Southern United States (Chapman)............... 2,266 1,666 74 Rocky Mountains (Coulter)...2.....2.2...00000% 2,206 1,910 87 Kos Angeles (Abrams)............... ee eS Oe 802 62 78 Blonidapweys, (Small) cj 005 oocc.c eed re ea Sue ae ead 9 od ater 415 225 54 Great Britain (Hooker)..................0.00000% 927 821 89 France (Cusin and Ansberque)................... | 3,924 | 3,492 89 CGenuany, (Mnglen)ie. 222. vcd) so isee + pane ane: LLL 947 85 Switzerland (Schinz and Keller).................. 1,899 | 1,726 gI Russian Empire (Ledebour)...................... 14,704 12,588 86 INotwayA(DIVEE) Ein tam acti cicircteig cs one etn, sesie ases 857 741 86 Keeland? (Stefansson) ss) 4! 22 ees eae: 221 || 200 90 Euilesmereland= (Simmons): . 1252 2..02 «cnt. . nse se. 76 7 93 Haroes:(Warming etial.):. - . 26 ic eem ene nee ns ne | 164 | 150 gl SySS UOT Gl oy W22)) kee ca) ee ee A Oe ee 4,481 | 3,554 79 Creten(Raulin)ie eee eet een. re Oe, 6 1,461 1,161 79 Sicilya Glornabene) ie i io oie oo atic he 2 Gee es 8 Ses 1,697 1,295 7 Sistas (OST) ep teerpece yay ee tes thac, Penne ene Ad cxtoeie 2 a ae ena 2,949 DAG, 84 Flora Orientalis (Boissier)...............0....06.. 9,771 8,110 83 apank (Matsumura): 68 oe oe aie es ee odes RONG 1,861 57 Braziledvineller)), Ase vel ass. {hb ds Goede oes 15,981 | 4,092 26 Ditto Amazon: Valleysonly. 6 cc.u oa cee ee be whee 2,209 265 12 British West Indies (Grisebach).................. | 2,249 675 30 Tropical Africa (Oliver and Thiselton-Dyer)........| 8,577 | 3,560 42 Biritisnelndias(ELOOKEN) F- ais Fa iY tate ee 4 Fc? te re ue +. © = 7 oe de cian 2K ; NIL TRELEASE: THE ANCIENT OAKS OF AMERICA 501 folia. Berry, Prof. Paper U. S. Geol. Surv. 98L. pl. 11. 7. Q. Lehmanni. Hollick, Md. Geol. Surv. Miocene. pl. 483.—PLIOCENE: 8. Q. lambertensis. Berry, Prof. Paper U. S. Geol. Surv. 98L. pl. 11. PLATE XXII. BicoORNES.—EOCENE: I. Q. negundoides. Lesquereux, Rep. U.S. Geol. Surv. 7. pl. 21. 2. Q. bicornis. Ward, Bull. U. S. Geol. Surv. 37. pl. 9.— MyYRICAEFOLIAE.—CRETACEOUS: 3. Q. praeundulata. Hoilick, Monog. U. S. Geol. Surv. 35. pl. 31 (as Q. stnuata Newb., a preoccupied name). 4. Q. antiqua. Hollick, Monog. U. S. Geol. Surv. 35. pl. 13.—MIOCENE: 5. Q. Ramaleyt. Cockerell, Bull. Torr. Bot. Cl. 33: 309.—SpPuRIC-ILICES.—CRETACEOuS: 6. Q. Haydeni. Lesquereux, Rep. U. S. Geol. Surv. 7. pl. 19. 7. Q. spuric-ilex. Knowlton, Monog. U. S. Geol. Surv. 17. pl. 48.—MIOcENE: 8. Q. Applegatei. Knowlton, Rep. U. S. Geol. Surv. 20°. pl. r. 9. Q. distincta. Lesquereux, Mem. Mus Comp. Zool. 6%. pl. 2 (as Q. voyanda). THE ABSORPTION OF CALCIUM SALTS BY SQUASH SEEDLINGS* RODNEY H. TRUE AND R.. B. HARVEY Bureau of Plant Industry, U. S. Department of Agriculture In the course of a former investigation! carried on with the white lupine (Lupinus albus L.) as a test plant, it was found that the rate of absorption of electrolytes by seedlings from solutions of the usual mineral nutrients was influenced to a great degree by the chemical character of the substances offered. In simple solutions it appeared that the usual salts of potassium were not absorbed producing rather an outgoing current of ions from the seedlings into the solutions; in magnesium solutions a small absorption took place in the more dilute solutions, while in calcium solutions absorption was much greater and took place in the most dilute solutions. It appeared from the behavior of the lupine in the solutions of calcium sulphate and calcium nitrate that in each case the quantity absorbed is limited, even though the supply may contain a large excess, and the effect of the anion appears to be subordinate to that of the Ca ion in determining the quantity absorbed. It was found by preliminary experiments by the present authors that this condition of things does not apply to all kinds of plants. The common garden squash, sweet corn and soy bean were found to behave quite differently with respect to the anion employed with the Ca ion. In this paper the records of a series of experiments are presented showing the absorption from a series of solutions of three of the commoner inorganic salts of calcium by the seedlings of Cucur- bita Pepo L. of the horticultural variety known as Early Prolific Marrow. Before considering the evidence on which the conclusions here advanced are based a word concerning the method is in order. Selected seedlings obtained from seeds germinated in chopped sphag- num were grown in carefully prepared solutions contained in prac- tically insoluble glass beakers. The concentration of ions of each solution was noted daily by taking conductivity readings by means of an accurate wheatstone bridge. The temperature throughout was * Published by permission of the Secretary of Agriculture. 1 True, R. H., & Bartlett, H. H. Am. Journ. Bot. 2: 278. 1915. 502 TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 503 maintained at 18° C. by automatic control so accurate that the range of variation was seldom above four tenths of a degree Centigrade during the course of an experiment running a fortnight. The con- tainers remained in darkness except during the short time required for the determination of the conductivity which took place in rather faint diffused light. Since it is obviously unsafe to draw conclusions from a comparison of ohms, results were always calculated to concen- trations expressed as gram-normals of the salt in question dissolved in a million liters of water (grm. norm. X 107-*). The water was obtained by twice distilling Potomac River water from glass with electric heat in a laboratory from which gas was excluded. Each experiment was usually continued until signs of deterioration began to appear in the seedlings. CaLcium NITRATE Several experiments were carried out with squash seedlings in calcium nitrate solutions. Since they were in close agreement but one is presented here, that running from May first to May fifteenth, last, inclusive. The distilled water used in making up the solutions had an initial conductivity equal to that of a solution containing 11.7 grm. norm. X 10-6 Ca(NOs3)o. Nine cultures each containing 5 seed- lings and 500 cc. of solution were set up in a series ranging in con- centration from 18.2 to 867.0 grm. norm. X I10-®. Daily observations were made until signs of exhaustion began to appear. Since in the cultures containing less than 50 grm. norm. the behavior of the seed- lings varied so little in the different members of the series only a part of the record is shown here in order not to confuse the table with several nearly coinciding curves. In the curve representing the record of the culture in distilled water a dashed line is employed (Fig. 1). It will be observed that in both distilled water and in cultures containing calcium nitrate up to a concentration of 100 grm. norm. < 10-* the solutions gain in concentration for two or three days, a course which in the distilled water is followed by a very slight ab- sorption until near the close of the experimental period. At no time, however, were the plants able to regain any considerable proportion of the electrolytes lost to the medium during the first few days. With the dilute solutions of the salt (under 100 grm. norm.) this period of leach gradually passes over into one of active absorption as a result of which these solutions are reduced to a lower ion content than the distilled water. As the initial salt content of the solution is increased to approxi- mately 500 grm. norm. the slight leach seen in the weaker solutions 504 BROOKLYN BOTANIC GARDEN MEMOIRS fails to appear, immediate though slow absorption being the rule. This intake gradually speeds up through the succeeding days, the absorption being roughly related to the original quantity of salt in the solution. This active absorption continues to the end of the experiment at which time the appearance of the seedlings and the WO &50 700 650 ro Ss x.108 iJ | S NORMAL CONCENTRATION Fic. 1. For explanation see text. diminishing absorption indicate approaching exhaustion. At this final stage of the experiment the plants have reduced the ion content of solutions of the salt to an approximately like minimum, which lies between 35 and 50 grm. norm. X 10~® Ca(NOs3)e. This concentration lies at about 25 grm. norm. X 10~° below the concentration reached TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 505 by the distilled water, and seems to represent a fairly well-defined irreducible minimum. This point is marked by a lower ion content than the similar minimum seen in the distilled water. It seems clear that the substances giving to distilled water its conductivity are not absorbed to as dilute a minimum concentration as is calcium nitrate. It will be noted that only in the two more concentrated members of the series are there any absorbable electrolytes remaining at the end of the experiment, and therefore only in these cultures is the total capacity of these plants to absorb this salt measured. In these cases the total absorbed salt equals 565 and 713 grm. norm. X 107° re- spectively. An inspection of the graphs shows that in nearly all solutions of the salt there occurs a time at which the plants reduce the conductivity of the solution to a minimal concentration, a point that may be assumed for present purposes to represent that of maximum absorption. In some cases experiments have been closed before this point has been reached in a number of members of the series (usually the more con- centrated ones), owing to the well-marked exhaustion of the plants in a number of the cultures. In such a case the absorption maximum may not have been reached, although probably in most cases it has been approached. It is interesting to compare the original concentration of the differ- ent solutions with the corresponding concentration at the time of maximum absorption. By this means one is able to ascertain how much of the salt is absorbed or how much net loss the plants have suffered calculated at the time of greatest absorptive efficiency. Carrying out the calculation referred to for the cultures included in this experiment the results seen in the following table (1) are obtained. A glance shows that about 37.0 grm. norm. Ca(NQOs3). are required by five squash seedlings growing in 500 cc. of solution to enable them to protect such ions as are mobilized from their reserves against the leaching action of the distilled water. This stated in terms of the quantity of Ca( NOs). per plant would be about 0.00028 grm. absorbed during a period of about two weeks. As the salt content is increased, absorption increases in approxi- mately the same magnitude leaving an unabsorbable residue of approximately 34 grm. norm. X 10-® Ca(NOs3)o. This minimum is here calculated as Ca(NO3)2, but it should be clearly understood that the substances actually composing this residue may be, and probably in considerable part are, of quite other composition. Indeed, the substances indicated as Ca(NOs)2 at all stages of the experiment, especially the later ones, doubtless consist in part of other materials, largely coming by exosmosis from the interior of the test plants. The 35 506 BROOKLYN BOTANIC GARDEN MEMOIRS TABLE I Net Absorption by Squash Seedlings from Solutions of Ca(NOs3)2 Original Concent. of Minimum Concent. of Maximum Net Absorp- Solution Solution as tion Calc. as Ca(NOs)2 & 1076 Ca(NOs)2 10-6 Ca(NOs)2 1076 L2Orgkma NOL. ee eae SOLOs STM NOLIN etc eee — 38.0 grm. norm. NOc2 we Siete ners AIBYAO). fh! (peadtacircheheanr > Ppigey, “ Pi Sete, 5 At eae es Bylo). ae eR AAA STA: = 19.5) i Boye ee SiGe aelottrara sete 3250 be Sta hades caer bees afer Anas “ ASO. ial tee yaa eRe Ry eg ae Ree. O00 es " MIS EG/ e Tad Wa Eee) Ae 29:0) Bote hares T6ol) ay te 45.5 “ Panto heey ae 26:3 lee FRA ORY ‘ 62200 os AE Ma eee haG BVA oy Ov Raat hs 230A * Oley sl eee eR Oe BAe Oma eM ola 62535 “ 18236) re AMER aie Roars oe 2916) ntl eee eee RR 153 Omnis yy 351.8 : MALL Tae se SORE 20 alee SIP Ea tRnido ie B12 7a i Sich es) Me Soins ees 47.8 DPM Stn clateeae ATO ae ‘ 6205 DL eter tie 125.0) 0 TRAE ore 567.6 “ my 86720) + ep cit oA ey 152.0) Nit ttinet dene: TAS te i concentration indicated in any case merely states the net result of ion interchange in a given culture stated in terms of such a solution of the given salt as would have a like electrical conductivity. As the quantity of salt is still further augmented, the unabsorbed residue increases, indicating a clear surplus of this constituent. It is interesting to note, however, that the unabsorbed residue does not increase at a rate parallel with the quantity offered; the absorption increasing also though at a lagging rate. The maximum quantity of Ca(NO3)2 absorbed by five plants out of a concentration of 867 grm. norm. X 10-° is about 714.1 grm. norm. This expressed in weight of salt absorbed per seedling would be about 0.0055 grm. This may be taken to represent approximately the maximum quantity of Ca(NOs)e absorbed by a squash seedling living in darkness in a temperature of 18° C. during the time required by the seedling to exhaust its available reserves. CALCIUM SULPHATE A somewhat similar experiment was made in which calcium sulphate was used, in the hope that we might get some light on the comparative effect due to the sulphate and the nitrate anions. A series of 14 cul- tures was set up using squash seedlings as before covering a range of concentration between distilled water and 830 gram normals of the salt in a million liters. The experiment ran from July 12 to 23, 1916. A graphic record? of the course of the concentration changes in the several cultures is shown in the accompanying diagram (Fig. 2): 2 In order not to crowd the figure only one curve representing an original CaSO, concentration less than 50 grm. norm. is given. TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 507 A glance at the curves shows that in some respects the course of absorption is markedly different from that seen in the case of Ca( NOg)o. Alike in the distilled water control and in the solutions originally containing 11.08 and 15.6 grm. norm. respectively, the solutions gain ions and not even at the time of greatest absorption are the plants able to reduce the ion concentration to that seen at the beginning of the experiment. These quantities of the salt in question are insufficient to prevent the solutions from acting practically like distilled water HO, 550 00 + 900 850 850 G0 B00 750 V [A a NORMAL CONCENTRATION X 106 BS Y [| | LIAS 58 — g OLAS 12 13) G14 a da ae DAYS Fic. 2. For explanation see text. by withdrawing a preponderating quantity of ions from the seedlings. When the concentration of CaSO, reaches about 30 grm. norm. in a million liters, an equilibrium of some sort seems to be established between the plant and the medium with the result that neither ab- sorption nor leach is marked. A similar equilibrium point appears at approximately the same concentration in the case of Ca(NOs)2 solutions. It is important to note that the concentration of the calcium sulphate solutions at the time of maximum absorption in 508 BROOKLYN BOTANIC GARDEN MEMOIRS these most dilute solutions is higher than is the case of those originally having a greater salt content, seeming to indicate that at this extreme dilution the plants are not only forced to yield ions to the solutions but are unable by reabsorbing them to reduce the concentration to a point lying much below the equivalent of 50 grm. norm. in a million liters. As the original concentration of CaSO, is increased to 31.3 and 52.1 grm. norm. respectively absorption increases to a point showing net gains by the plants. Leaching of ions by the plants, if it takes place, is more than met here by their greater absorptive activity. This activity when greatest reduces the residual ion content markedly below that seen in the case of the originally more dilute solutions. It seems that with the addition of even slightly larger quantities of CaSOu, the absorptive function becomes more active and is able more nearly to exhaust the quantity of ions offered. As the quantity of CaSO, is increased to concentrations rising from 101.9 to 824.4 grm. norm. in a million liters the plants reduce the ion content of all solutions but absorption even at its greatest leaves a residue which increases as the quantity offered increases. Absorption, while in general increasing as the quantity of ions present increases, lags far behind the quantities offered. It thus comes about that the curve representing the residual ion content approximately parallels that representing the original ion content of the solutions. From these data it would seem that when squash seedlings are grown in darkness at 18° C. in solutions of CaSO, alone, they are unable to absorb as many ions as they lose when the solution contains less than about 30 grm. norm. of the salt in a million liters. When the solutions contain somewhat more than this quantity of this mixture of ions the plants can reduce the more dilute members of the series to an ion content of about the concentration seen at this equilibrium point. When the supply of ions is far in excess of absolute requirements, the seedlings absorb greater quantities than in weaker solutions but the quantity of residual ions left in the solution increases in nearly the same proportion as the quantity offered. In the following table (Table 2) are shown (1) the original con- centration of each solution, indicating the quantity of ions offered in each case; (2) the concentration of each solution at the time of maximum absorption, or the residual ion content of each solution and (3) the quantity of ions absorbed from each solution at the time of maximum absorption. TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 509 TABLE 2 Net Absorption by Squash Seedlings from Solutions of Calcium Sulphate Original Concentration Minimum Concentration Maximum Absorption as Grm. Norm. as Grm. Norm. as Grm. Norm. CaSOs X10-6 CaSO; X 10-6 CaSO; X 10-8 ARON ere eres oo oes Se AGiGn eet eee ew — 38.7 BIC OSL ents con eas an se Cop [as BE ie ne on ae ee — 409.32 Rij Oa frst ste tae le eons See BOLO sar eaisee ee eens ee a Aad. GING i ie ge eee en eae 217 (AS Yies PREY ete Fey hy tio be = 334 Banco serve ot agar cts & Ses BT eT ep ee 14.4 ONCO Bre top cy iaetialanc meine seus 7 OS Or Pas eck eae ee aaa 25-9 11/10 ER Ae eo pe AU GLO en pet eee 57-4 EQ Hedin a: oiekest ic st Ak a keewe TAG 2M 2. ee 51.3 ATE) Se ots ihn csi node, Secu Move sol BERR seys Ma ceva ons 57 ANT APs oA Rt eds Ae Seed ct Ba5sS tit othe eae Oe 78.8 GoGy eek as een a aes Eases fe AAS pikes Bleach Mae 87.3 OSAP Am TS ee tk. he et Sees 5287 yee ee 125-7) FAT Cee Oe er ene, NE GES 0) eee oes 112.0 SALA aioe He cht oy ee TORO nea tise. cet eee 119.8 CALCIUM CHLORIDE In the foregoing experiments we have had to do with salts in which both cation and anion are required for the normal growth of higher plants, and we find squash seedlings behaving toward them in sharply contrasting ways. From solutions containing G and NO3 i ions root absorption is very active. When Ce ions are accompanied by SO, ions, however, absorption is relatively small. In view of these facts it is a matter of no small interest to test the behavior of plants in solutions in which the Ca ion is accompanied by the Cl ion. A series of solutions was made up containing a graded series of concentrations running up to 582.4 grm. norm. in a million liters. The experiment ran from Aug. 9 to Aug. 26, 1916. A graphic record of the concentration changes observed is shown in the accompanying group of curves (Fig. 3). It is perhaps hardly necessary to do more than to point out certain of the more important features to be observed. In general a very striking similarity to the corresponding set of curves obtained from Ca(NO3)2 solutions may be noted. The solutions hav- ing an original concentration of 32.5 and less show a loss of electrolytes for the first five or six days after which time absorption begins and continues until near the end of the experiment, but in none is the loss during the early days fully regained, although in that originally containing 32.5 grm. norm. the net loss is practically negligible. At this concentration we find again evidence of a critical concentration of some sort below which the plants can not absorb and above which 510 BROOKLYN BOTANIC GARDEN MEMOIRS they are able to do so. This required minimum quantity is probably of considerable physiological significance. At higher initial con- centrations, from 53.5 to 384.0 grm. norm. loss of ions during the early days of the experiment becomes less marked as the concentration increases. Absorption usually begins more promptly and proceeds with greater speed as the quantity of ions present increases until in all cases but one the concentration of ions remaining in the solution is reduced __[SauAsH CaCl, 18°C. 700 700 650 550 50 De) Ss 00 450 S ~ 400 Ss 3 0 S = 300 & 50 S = lf 200 1/50 PERE Bae ae aes Bee, CO OE ZA Se aaa) Fic. 3. For explanation see text. to the unabsorbable minimum at an average concentration of approxi- mately 27 grm. norm. X 107®. Only when the original concentration of 582.4 grm. norm. X 107° is reached do we find the quantity of ions offered greater than can be reduced by the seedlings to the unab- sorbable minimum. These relations are seen in the numerical data given in Table 3, in which thé concentration relations are shown as they exist at the time of maximum absorption. Here again the similarity to the situation seen in solutions of Ca(NQOs)z is striking. If we are justified in concluding that the depressing action of the SO, anion is responsible for the great reduction of root absorption TRUE & HARVEY: ABSORPTION OF CALCIUM SALTS 511 TABLE 3 Net Absorption by Squash Seedlings in Solutions of Calcium Chloride Original Concentration of Minimum Concentration of Maximum Net Absorp- Solution Grm. Norm. Solution as Grm. Norm. tion as Grm. Norm. CaCh X10 CaCle X 10-6 CaCle X 10-8 ABQ EO LIM MOLIN e r5 ale) os 255.1 - 36:5 Stil) NOMS... — 34.3 grm. norm. ae ae ~~ se “a “cc aa 9.2 eae 25.1 ae ier ee — 15.9 LS a7hy 6 to tg geht eee Oe Een a 26.5 1 nae ey On mem ae “ee “a ac ae Bonptere bl celts, Sak xs 33. CRs ieee Pan ea. hee = Tet ae “ee se ae “c Seb) a ee eee A he Pah PENT + 10.8 oe oe ~ ae ae aa 75-3 eh Re axceeeeae ZO et ete Mere ees 54.8 TOA? SYM as ik eee eee LOu7) | os ites tod ate 96.8 “ : 191.8 . . Sy eae 19.9 men zn SopeN ote T7100 oe: FLY SPE ee eee 32.3——— Eee S57 mae 5o240 hgh s ae k PAT piacere tlc Sey 441.4 “ seen in the solutions of CaSO; when compared with that seen in solu- tions of Ca(NOs)e of a similar range of concentration, we are also justified in concluding that the influence on process of absorption due to the Cl anion is as favorable as that exerted by the NO; anion. DISCUSSION OF RESULTS It seems clear that for both the squash and white lupine in the seedling stage the calcium ion favors the absorption of ions. The lupine while finding this ion necessary does not absorb it in as large quantities as does the squash. The maximum requirement per cul- ture of five seedlings of the lupine under the conditions of these experi- ments is not over 175 grm. norm. X 10-® Ca(NOs3)2 and about 125 grm. norm. X 10-® CaSQ,.’ The squash on the other hand reduces an original concentration of 518 grm. norm. Ca(NOs3)2 X 107* to 47.8 grm. norm., thus finding but a very small surplus present, and may absorb more than 700 grm. norm. when a concentration of 867 grm. norm. X 10~° is offered. It is more quickly satisfied in the case of CaSOu,, this plant absorbing a maximum of about 120 grm. norm. only from solutions increasing in concentration up to about 825 grm. norm. X 10-°. It takes in there- fore from an excess supply about as much CaSQ, as does the lupine. This comparison gives additional evidence of the well-known indifference of the white lupine toward calcium in several combinations. From the evidence at hand it appears that in the presence of the Ca ion, the effect exerted by the NO; and SO, anions on absorption by the lupines is not markedly different. With the squash the anion effect comes strikingly to the front. The Ca ion accompanied by the 3 Calculated from True, R. H., and Bartlett, H. H. Am. Journ. Bot. 2: 262 and 265. I915. 512 BROOKLYN BOTANIC GARDEN MEMOIRS NO; ion is from four to five times as favorable for absorption under the conditions of these experiments as the Ca ion accompanied by the SO, anion. This seems to indicate a striking and specific difference in the influence of these anions on the absorptive activities of the squash. When the Ca ion is accompanied by the Cl ion, absorption is influenced very much as in the case of Ca acting with NO; ions. The favorable effect of NO; and Cl ions is contrasted with the action of the SO, ion. The strong influence exerted by the specific characteristics of the different species of plants is seen in the contrasting behavior of the lupine and the squash in the presence of Ca accompanied by the NO; ion. It should be borne in mind that the probable physiological inter- action of a given pair of ions is perhaps such as to make it unsafe to speak strictly of the specific action of any single ion irrespective of that of its companion ion or ions. It is obvious from what has been here shown that any theory of cell permeability which may be framed to account for the income and outgo of the living plant with respect to electrolytes must reckon with the striking differences that exist in the behavior of plants toward even such fundamental factors as the required mineral nutrient ions. INHERITANCE STUDIES ON CASTOR BEANS ORLAND E. WHITE Brooklyn Botanic Garden Ricinus, though a monotypic genus involving only a single widely recognized species (R. communis), possesses a multitudinous number of forms, which from time to time have been temporarily ranked as species. These forms breed true to many of the numerous characters which distinguish them, as shown by data obtained from growing several generations of fifty or more types in the experimental breeding plots of the Brooklyn Botanic Garden. Numerous crosses between even the most extreme types have given perfectly fertile F; and F, generation hybrids. Hybridization studies to determine the manner of inheritance of a dozen or more of these characters have been followed through the F,, F2, and, in some cases, the F3 generations. Several thousand plants were involved in these studies. MATERIALS AND METHODS Seeds of the various types were secured through Farquhar & Co. of Boston, P. Henderson & Co. of New York City, and from various botanic gardens. Many of these types are known in seedsmen’s catalogues as varieties or sub-species, and these, much to my surprise (since the castor-oil plant is monoecious and wind-pollinated), bred true immediately to many of their more prominent characteristics, such as stem color, seed color and color pattern, and height. Further observations on plants of different varieties grown close together demonstrated that very little cross-fertilization took place (certainly not more than five percent), even when conditions were most favor- able. This rather unexpected tendency to self-fertilization in a monoecious plant adapted apparently to wind-pollination is largely due to the slightly earlier maturity of the male flowers and to the comparative isolation of the flowers of each plant through the preven- tion of air currents by the large leaf surfaces. As the stigmatic surfaces of the female flowers become exposed and mature, the pollen from flow- ers on the same plant has already fallen or falls upon them in small clouds, thus insuring, to a large extent, self-fertilization. Difficulty is experienced under Long Island climatic conditions in making bagged inflorescences on outdoor cultures, set a normal amount 513 514 BROOKLYN BOTANIC GARDEN MEMOIRS of seed. Because of this, the F; generation plants, in some cases, were grown in isolated cultures, instead of being bagged. In this way large quantities of seed were obtained for growing F. populations. In most cases, enough seed from bagged F; plants was obtained to check up the F» results from the unbagged seed. A large number of F; families were grown from seed of unbagged Fy, plants. The per- centage of cross-fertilization among the Fy» individuals appeared to be small. EXPERIMENTAL WORK Stem Color Stem color in castor beans can be roughly classified into five cate- gories (see Plates X XIII and XXIV)—bright green, green with reddish blush on sunny side, carmine or rose red, mahogany red, and purple (dark red). The development of each type of red coloration depends, to some extent, on sunlight, particularly the red blush class. In shade, plants of this class have green stems. These red color types are similar chemically, so far as solubility tests with their pigments are concerned, since all the red pigments are soluble in water containing a trace of chloroform, but remain insoluble in alcohol, xylol, or in pure chloroform. The red pigment is a sap color, occurring in the epidermal palisade cells in the leaf and stem (see Plate X XIII), also in parenchymatous areas of the stem, especially in the pith in some varieties. The different shades are apparently due (as observed microscopically) to different concentrations of the pigment in the cells and to a difference in the amount of pigmented cell area. From evidence thus far obtained, the writer is inclined to regard the hereditary differences between the red types as due to the presence and absence of several color dilution factors, each of which modifies the expression of a red pigment producing factor common to all, except, of course, the green-stemmed class. Still further modi- fications in coloration appear to be due to the presence and absence of a pattern restriction factor, in the absence of which the stems and leaves are mahogany red, or purplish red if ‘‘bloom” is present. In crosses, red blush X other red blush varieties always gave only red blush F, and F, offspring. Red blush varieties X green- stemmed varieties and the reciprocal always gave all red blush in Fy and approximately 3 red blush: 1 green in Fy». ACE allysOb tained esse eee tie eine 113 red blush: 43 green Mheoreticallyvexpected... 0-4 0540-5 1G FG flail des 200 In F3, seed from unbagged Fy. green-stemmed plants generally gave all green plants, while unguarded seed from Fy, red blush plants either gave all red blush or both red blush and green-stem families. WHITE: INHERITANCE STUDIES ON CASTOR BEANS 515 Red blush X mahogany and reciprocal gives rose or carmine- stemmed plants in Fy, and approximately 1 red blush: 2 rose: 1 mahogany in Fe, the actual figures being 47 red blush: 144 rose: 47 mahogany, theoretically expected 59.5 red blush: 119 rose: 59.5 mahogany. No Fs; generation of this cross has been grown, but from the above ratio the plants with rose-colored stems are expected to produce all three Fy types again, while the other two types are ex- pected to breed true. As previously stated, there are forms with rose-colored stems that breed true. These when crossed with red blush varieties give all rose-stemmed F, plants, indistinguishable from their rose-stemmed parent. In Fs, these produced 429 rose: 145 red blush, the the- oretically expected proportions being 430.5 rose: 143.5 red blush. Associated with the types of stem coloring are pigmented areas in other parts of the plant. The mahogany-red-stemmed plants have mahogany-red leaves and fruits. The rose- and red-blush-stemmed types have green leaves with red or reddish-green midribs. The dark purplish-red- (mahogany bloom) stemmed plants have dark purplish- red leaves and fruits. Plate XXIII is a much enlarged micro-photo- graph in natural colors showing the pattern and distribution of the pigmented areas in the leaf of the mahogany type. The pattern resembles one of the mottling patterns in castor-bean seed coats, and possibly may be due to the same cause. It occurs only in the forms with dark red or purplish-red leaves and stems, and may be regarded as resulting from the absence of the pattern restriction factor previ- ously mentioned. Bloom Bloom, similar in appearance to that on grapes, in castor beans consists of a scale-like waxy substance, which, under the microscope, resembles a piece of cracked frozen ground. It is easily rubbed off. This covers the whole plant and is especially noticeable on the stems and fruit capsules. In some varieties, it is produced more freely than in others. Many forms are known which breed true to its absence. When it occurs on plants with mahogany stems, a dark purple or purplish-black effect is produced (see Fig. D, Plate XXIV). Crosses of bloom X no-bloom give either complete or partial domi- nance of bloom in F;. In Fs, approximately 3 with bloom: I no- bloom were obtained (actual numbers being 1,108 bloom: 377 no- bloom, the theoretically expected being 1,113 bloom: 371 no-bloom). In F3, seeds from unguarded F, plants without bloom produced only plants without bloom. Seeds from unguarded F», plants with bloom either bred true in F; or gave 3 bloom: 1 no-bloom families. In one 516 BROOKLYN BOTANIC GARDEN MEMOIRS cross of bloom X no-bloom, the F, plants had bloom, but were only lightly covered as compared with their ‘“‘bloom”’ parent. In Fs, the plants approximated a ratio of 3 bloom: 1 no-bloom as usual, though many of those with bloom were lightly covered as in F,. Dehiscent and Indehiscent Seed Pods The seed pods or seed capsules of Ricinus, in most varieties, are dehiscent, the seeds being thrown out of the mature ripe capsule with great force. These are known as “poppers” in regions where the plant is cultivated commercially. A few varieties have inde- hiscent capsules, the seed being retained within the pod for several months. This characteristic considerably reduces the cost of harvest- ing the beans, as only two or three collections a season are necessary, whereas ‘“‘popper”’ varieties must be harvested twice a week or more, to prevent loss. The castor-bean plant matures its seed over a very long season. The “non-popping”’ variety with which the writer experimented has thin-walled, comparatively brittle capsules, while all the popping varieties have capsules with thick, leathery walls. The dehiscent characteristic is probably due to cells similar to those found in the fern sporangium, which rapidly lose their moisture content when the capsule matures and contract, thus breaking apart the cap- sule and expelling the seed. Crosses of ‘popper’? with the ‘‘non-popper”’ variety gave all ‘“poppers”’ in F; and approximately 9 “popper’’: 7 ‘“‘non-popper”’ in F2 (the actual figures being 343 pop.: 259 n.-pop., the theoretically expected being 338.4 pop.: 263 n.-pop.). In F3, seed from unguarded F» “‘popper’’ segregates gave either all “poppers,” or “poppers’’ and “‘non-poppers.”’ Seed from unbagged ‘“‘non-popper’”’ F2 segregates produced in most cases only ‘“‘non-poppers.’’ The “popper’’ and ‘‘non-popper”’ vari- eties involved in the crosses were very distinct, but the F2 populations were somewhat difficult to classify, as many of the “‘non-poppers”’ would, under very favorable conditions, slightly pop. These were usually thin-walled, brittle capsules, showing, perhaps, that the nature of the capsular tissue (thick, leathery or thin and brittle) modified the “popping” or “non-popping”’ characteristics. On the assumption that two pairs of characters are concerned in this cross, each of which is primarily determined by the presence and absence of a single factor, the results may be interpreted by regarding the “popper”’ character as due to the presence of both the factor for popping (A), and the factor for thick, leathery capsules (B). In the presence of A and the absence of B, the capsules would have thin, brittle tissues, but pop slightly, although not sufficiently to class them as “‘poppers.’’ In the ‘ WHITE: INHERITANCE STUDIES ON CASTOR BEANS 517 presence of B and absence of A, the capsules would be thick and leathery, but non-popping. When both A and B are absent, the capsules would be thin, brittle, and non-popping. On this provisional hypothesis, “‘non-poppers”’ of the aB class crossed with those of the Ab class would give all AB or “‘poppers” in Fy anda 9 : 7 ratio in Fo. The two types used in the above-recorded crosses would be represented by the formulae AABB = “poppers,” aabb = “non-poppers.”’ Crosses of these would give a 9 : 7 F» ratio, such as that actually obtained. Seed-coat Colors Seed-coat colors in castor beans are white, brownish yellow, various shades of red, gray, brown, and black. With one exception, all forms, so far as the writer knows, have seed coats in which the ground color is modified by one of several mottling patterns, although the mottling patterns are inherited, as in garden beans, independently of the ground color. The nearest approach to a self color in the writer’s collection is a black-seeded variety having in some cases very few mottling marks and in others none at all. Efforts have been made to discover a self-colored white-seeded variety, but so far with no success. In crosses, chocolate brown is dominant over black. red, white and gray. No Fs data are available from any crosses excepting those of red X brownish gray and its reciprocal. The F, is brown on a gray background. In Fs, segregates of various degrees of redness appear as a minority. By counting all those Fy. segregates with a red cast, an approximation to a ratio of 3 brownish gray : 1 reddish gray is obtained. The actual results are 172 non-red : 40 reddish gray or red, the theoretically expected results being 159 non-reds : 53 red gray. No reds as brilliant as the grand parental type appeared, showing that more than a single pair of factors is involved. In F3, seed from unbagged F», light red segregates gave all light reds in the majority of cases. Seed of the same kind from medium red F, segregates also bred true. Seed from unbagged brown F>2 segregates gave browns of various shades in some cases in F3, while in others, browns, reds, and brown grays were produced. Reds as brilliant as the red ancestor were secured from red F2 segregates. At least three types of seed-coat color mottling can be distinguished definitely in castor-bean seeds. One is coarse-veined, one is fine- veined, dotted and splotched, while the third is characterized by a very few large splotches. When the seed-coat is black, the mottling 518 BROOKLYN BOTANIC GARDEN MEMOIRS is obscured. As in the case of the stem colors, the different types of mottling appear to involve the presence and absence of several pairs of restriction factors. Only two of these patterns have been studied in detail. These are the coarse- and fine-veined types. Crosses between coarse and fine always give in F; all fine, indistinguishable from the ‘‘fine’’ pattern parent. In Fo, approximately 3 fines : 1 coarse are obtained, the actual figures being 163 fine : 49 coarse (theoretical expectation 159 fine :53 coarse). Unbagged Fy. segre- gates having coarse mottled seed generally breed true in F3, the few cases where plants with fine mottled seeds have appeared being un- doubtedly due to foreign pollen contamination. Unbagged F». segre- gates with fine mottled seeds either bred true in F3 or gave both fine and coarse-mottled progeny. Coarse-mottled x the large splotched type gives a dominance of the former in F;. No F2 progeny have been grown. INTERPRETATION The inheritance of five of the sets of characters described in pre- ceding pages—green and red blush stems, red blush and mahogany stems, red blush and rose red stems, bloom and no-bloom, fine and coarse seed pattern—so far as the F,; and F» data are concerned, are most simply interpreted as due to the presence and absence of a single genetic factor in each case, making in all five genetic factors. The inheritance of dehiscent and indehiscent capsules is assumed to involve primarily two pairs of factors. No evidence of close linkage was found between any of these seven pairs of factors, although the data were taken with this end in view. SEED SHAPE AND DIMENSION Castor-bean seeds differ as to shape in being oval or orbicular (about as long as broad). In crosses between varieties breeding true to the two types, the F, plants are all oval seed, while in Fo», orbicular seeds are present in considerably over one fourth of the progeny sug- gesting a 9:7 ratio. Varieties of Ricinus vary remarkably in their seed dimensions and weight. Some of the commercial varieties have seeds less than a centimeter long, which run about 4,550 to a pound of 450 gm., while the seeds of some of the large Zanzibar ornamental types are over 2.5 cm. long and run only 450 to a pound. Between these are numerous forms breeding true to almost every gradation in size and weight. A large number of crosses between these types have been made, the F, plants showing various degrees of intermediacy. All the WHITE: INHERITANCE STUDIES ON CASTOR BEANS 519 different F, plants of each were practically uniform as to seed size (see Plates XXV, XXVI, and XXVII). Most of the large-seeded forms and some of the very small-seeded forms (with indeterminate growth period) require a long season to produce mature seeds, so that even when Fy» populations were started in the greenhouse in pots several months before planting out, only about two thirds of the segregates matured seed. One year, attempts to overcome this difficulty by growing the plants in 10 cm. pots for a year were unsuccessful. The difference in seed maturity between the outdoor and these pot-grown plants was very slight. Because of these difficulties, several of the F2 populations shown in Plate X XVII repre- sent only part of the segregates—the small- and intermediate-seeded classes. In F: populations from small X large seed or the reciprocal, small-seeded types similar to the small-seeded grandparent and even smaller were obtained in every case, while in some of the crosses involving nearly complete F.2 populations (Plates XXV, XXVI), the large-seeded type was also obtained. In all crosses, as expected from studies of size characters in maize, poultry, and other plants and animals, numerous intermediates were present, so that a complete F2 population represented a gradating series ranging from those similar to or smaller than the small-seeded parent to those similar to the parent with large seeds. Seeds from unbagged F2 small-seeded and large-seeded segregates have given similar F3 progeny, showing the extremes to breed true. Various F, intermediates have also bred true in F3, while other inter- mediates have given the whole Fy, series again. Still others have shown very much less variation. OTHER CHARACTERS Numerous characters, other than those described in preceding pages, have been studied from the standpoint of heredity, but not in sufficient detail, to admit of interpretation. Crosses between low- growing (dwarfs), early seed-maturing types with determinate growth, and tall, late-maturing types with indeterminate growth gave inter- mediates in F,, which in F, gave all three types, though accurate classification so far has been impracticable. Crinkled, much notched leaved types crossed with ordinary leaved types gave either dominance of the ordinary type or intermediates in F;. Some types have a loose, few-seeded fruiting spike, while others have a dense compact spike with a larger number of pods. Crosses between them give either intermediates or dominance of the loose spike. In F2, both types reappear, together with many intermediates. 520 BROOKLYN BOTANIC GARDEN MEMOIRS HETEROZYGOSIS F, hybrids between several of the forms produced, as in many maize crosses, a much larger amount of seed than either parent, the environment being practically the same for all. F, hybrids between still other forms, however, failed to show this increased productivity. This is also true of maize F, hybrids. This increased productiveness in F, should be, as in the case of maize and tomatoes, of great commercial value, since crossing castor- bean varieties, where no particularly accurate results are desirable, is very simple and could be done rapidly. Plants of the two types to be crossed could be grown separately and one lot used entirely as a pollenizer. A large quantity of pollen from the same spike matures at the same time. Hence, these spikes could be cut off when nearly mature and laid on paper sheets till the pollen was shed—a matter of a day or two. The pollen could then be collected in a powder gun or similar device and shot over the newly matured pistils each morning. The male flowers on the plants used as_ seed-bearers, for the most part, can be easily rubbed off without injuring the flower spike. The amount of selfed seed by this method would be very small, most of the mature seed being crossed. As ordinarily planted commercially, one bushel of beans running 1,500 beans to the pound will plant anywhere from 6 to 40 acres, depending on the distance apart and the number of beans planted per hill (1-3 beans). Some of the commercial varieties run as high as 3,000 beans or more per pound. Castor beans produce from 10 to 40 bushels per acre, depending on the variety, soil, climate and length of frostless season. With these facts in view, it seems unnecessary to urge the commercial importance of using F; generation hybrid seed produced by the method mentioned above. Experiments to determine which varieties crossed together would give the greatest yields in a given locality should be made in regions where the beans are grown commercially, since, as previously stated, there is great variation in varieties as regards this characteristic. This characteristic of increased productivity in F, progeny of certain varieties, combined with such characters as ‘“‘non- popping”’ and high oil content, should help toward putting castor-bean growing on a better commercial basis in this country. No experiments regarding increasing the oil content through “‘selection,’”’ so far as the writer knows, have been made. Varieties with seed yielding 30 to 45 percent oil are said to be already common commercially. Experi- ments with, and chemical analysis of the innumerable varieties would possibly give us strains with a much larger oil content. BROOKLYN BOTANIC GARDEN MEMOIRS y, “7 PLAT xx I WHITE [NHERITAN( 5 BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME I, PLATE XXIV. WHITE: INHERITANCE STUDIES ON CASTOR BEANS BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME |, PLATE XXV aN Ve eS HH YN MG 8 te “e YOY a WHITE: INHERITANCE STUDIES ON CASTOR BEANS BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME I, PLATE XXVI WuiITE: INHERITANCE STUDIES ON CASTOR BEANS BROOKLYN BOTANIC GARDEN MEMOIRS. VOLUME |, PLATE XXVII WHITE; INHERITANCE STUDIES ON CASTOR BEANS | 3 ‘we aoe VOLUME |, PLATE XXVIII. MOIRS. a BROOKLYN BOTANIC GARDEN ME HERITANCE STUDIES ON {CASTOR BEANS N HITE: I Ww WHITE: INHERITANCE STUDIES ON CASTOR BEANS 521 EXPLANATION OF PLATES XXIII-XXVIII PLATE XXIII is a microphotograph of a highly magnified portion of leaf epidermis of the mahogany type showing the red pigment pattern. PLATE XXIV illustrates the various stem colors. Fig. A is red blush and green bloom; Fig. B is green; Fig. C is red blush and green without bloom; Fig. D is mahogany red or dark red-purple with bloom; Fig. E is mahogany or dark red without bloom; Fig. F is rose or carmine red. PLATE XXV illustrates inheritance of seed dimension. A. The ovule or maternal parent is represented by 30 seeds, each seed representing a random selected sample from a single plant, showing the degree of variation in seed dimension within the variety. B. 30 seeds selected at random from a single plant of the maternal parent variety, showing the degree of variation in seed dimension on the same plant. C. 6 seeds representing 6 plants of the paternal parent. D. F; progeny. Each F, seed is a random selected sample from a single plant, illustrating the uniformity in seed dimension of the F; progeny. EE. Three F, families are shown, each separated by a white line. Each F2 segregate represented by a single random selected seed. PLATE XXVI illustrates inheritance of seed dimension and coarse and fine pattern. A. Maternal parent. B. Seed similar to paternal parent in size, seed coat pattern and in seed shape. C. F; progeny. Note the uniform appearance both as to size and pattern in the F; progeny. D. F2 progeny. Only one F2 family is represented. The coarse and fine pattern segregates are arranged in a separate series. PLaTE XXVII. A series of crosses illustrating inheritance of seed dimension, pattern and seed shape. The different crosses are separated by a white line. Each seed was selected at random and represents a single plant. Only part of the F, progeny of these families are shown, as the large-seeded segregates did not mature. In all these crosses excepting cross no. 4 (from the top), the maternal parent is placed on the left, the paternal on the right, and the F; generation between them in one row, while the row or rows below represent the Fz progeny. In cross no. 4, the group of 7 beans on the left is all that matured of an F2 progeny, the F2 parent and two grandparents of which are in the same row and arranged as mentioned above, the ovule parent on the left, etc. In some of the F2 populations, attempts to classify the beans as to shape have been made. PLATE XXVIII. A. Two leaf types. Varieties with the crinkled, notched type being much more ornamental than the other. B. Loose and compact forms of fruit- ing spikes. The compact form associated with certain other characteristics in a variety means greater productiveness. I IUD TA mM | Hi 5185 99 er heer eer eae Toaphere rene yeyty en a cenaette poabdan oer reer rte Lippe yet vy aor yt ot erry Sahat ehped saidieat ys eth Pe hpsbat > § Teena is rls rarest) inate ee carer i divmetctedee + Joe ey Peg +4 ry tyag bye Gohsaet-onid “ AsliserteG: ih hs rPheAig | bAdbaP thy Ne? eva eceinen urn > er" put sab Pah ney Nh Yh en niannretiil cf pap bole hy elnino} e182 1 by ete a ke mere Barer santo pebelic het iste te Tee habearaer ime forute NOE ee eetied ae * Ve a 4" f : , agiraopistrntines teat seoati ta aac peramniconget rh Aro bess mae nat cs Sopyvhyromanney adr pry ort a ates rte itod Me " mek ne aiat WF fe ahh © ed Mite atan gn Botettrern talents te ; 4 eae : Nie ee PAP ead nye eee acataed > \ ae + H ry ces(anis fo tieet Sulpeune Ary “el ptliy joaddittombe tr patien vrs tonne tao) eipititanicneet side tape ricaeeaes vehi vy) thai sane fat a east tats Sate it eect it tr tsberets } + te Sapaeleane ere: soe fa to aa Hite bal A sft) 9) " ty Mf itethsete oe i} pyory He by r viii eet ment of) y Ree ig By mat FS Ah a pes apace Geb nbenay 14 Rete er j (Fay Crag oi et diseith ahs oy eth BIEN Bi Mh - te vibe of cole ee Wettig) Saroente Ae HodeF Re ORNLAK beet ds pret oo of SE A) Ho gi a ge rrr. ne ee ay Sarge tes CAMP Be Mb PAL fr Megane ters rat v Di acta sie vet eegntie yeertet shane frites angen uit pert iSitattbetrigis ” Se oe bbe i ¥ ain i ein} aaa if pla autinigehgernts uit Ne usecht eu NV Peete ead Mabry: frie lar nd ta ey tar rr 1 He J rar f at 7 rt + : bg pede ay “0? Oo" ath di lugproru tats ear otyiniyt ee h= tie aime My PREM i rag } hake ta rg RET bem cae Bt bay AP a Wy oseeewenel rts 6 tes Mish bab eth ita HG dpe bb \Tohapt ah ae w ree ot Rite vee Uva aK Mae ata faeetieh Neperey taht fie peill PF ont vevutone ray gaan a Wa ty rug bein isvinanra citi Por ACRE? Rep yhae gies of tit ca ont bd sy Wrgiy -aopet eaeleh@ oe i pelbQoy PemdiET Wie | BiAT Hr peed Ten iune 4 Fle tiee 4 bie Meo bent ted heh hat ans 4 ry oheoe h2 iste chracdetrntehreenetiy: Wish) pepsin (enone ites ty 0 NH peeve siebed tiene Oe f TTA ALA dae + holy eure Teentert ta! eraredie Lope reat shai aed eb $eu neh ae oo Lerky eile ken cale Sebieceasceyers? ocelh baer ir a pehats praet e phe ie We dhe qateredtilenri itt On oe hoc g Bets Lire eased ast etirbumreperlt den ie habe yf tate ne Rodi eets parpedenagauhues #8 (M4 epee ee iether bet st hind nes f \, a) poeyaieat) tor Aree bye ae eiririnhh igen pris or arabe til seine re wegen keane en yved ye sabe ae pepe pele = peteltbed +d) of ti Fisvpeaeniteh , Siaaeue raping vdede butiegobiatets tas ihe " tit tee Biter see at ried e bY pet ey one eqrere eons pase tlt natant rok bpso-t\ 0 iehehenngeasestttintt ra shes d>)¥8 ponoy rue son + tarde rere actet b rd He UE aad it pied yeh ried weyrnetd bwied iY fevbtaserovartt4 hve 6 sibel y ety eet feat ais rie greene pees area 4 draibeten bent t na) fy wre chobewetntciy greneen * fupnepene he eed (ri ie had bas shah} +! if crete ete Pred pone Reet My Pe himnre Urfete fegseswmints yet meornhardetsunan py obi ger Mh i weiter 4t medi toe Lott grrone arent apis ser ie valde ibeesalts pecey, hae bE e/ ERA Rwe tear heey ee hd PO ara terse tes eens we é Sickie phena ded f ‘ heard hy oer hal bla Ppl naoe dn oe fr orvedem Ww eneviged et) be OUFHTTR ARE i Semen + Mere ate tieod sare ages re raat Mrrasetdeetempesrechege ania tty des ia asi See ee concer bairestecd caste 4 enene crt: ; Se Soret al serhewet (ride vitveepcrew nett at tte ; ie. Neat prilled toned sce mabenay ae but Hhederw ada a 1 “ ereryri Hears anavinettitte purlesositrant peje ot rl eee deiet thie een Cie iphebetrer teat inne aah byegeieeyd see) att Patnuere at due leh thet ere pire roe naie oe 7 rn eveweconyer aueeed eet in” ¢ Hy varied syareeaecney trait ear ptnee sec tree it an ttt Tad aaa borer tag teeed ety fete aatiet sett EET Lie cenivr een elt ttae Met ; Arid torah r" vente Reyes sate bbedr hii} pad sp rope it + WER HR Te AEH ONES CeBEHCH OOTY ant Hebru secur, aperassetsnecpres coment ainanitneahinatccs ty Snts tte henaeen gang attract q Wtegren ier el * ve ne (rp hett a aicgehent ; ee ath a oth ak it + we whe erte ti meet pede bt ii fos suc set arrepeled rheneuserer ted tegecren ty trl he te eireondewts Sie is baeeaites * ard oe ow bits hy Serrated $i yin) ort ripe are) bape diese we vet p(t re re y " facitiens vshaqeianiere! 9s wpa Pheer ee ey Oriented iber april OF reret aeeamtaanas ina sirwce at . hey) sais here eben eeeny earavini't raisers erent ee nego elit be “ ae Bish eis rereit iach eat tie bes ih) oeetalanesarhol Eee ecianrt aii dante f wren (ett vara att te i“ re e Spree arene ympat eet eartt ere bod pris aa eatidnesd Pitter: piu ei 4 ‘ een put batts , pene ny yatatesutiegeis etre (eae or Ort erel Cae wigehe eh Hey “+ tet ne itiyeee beneag sre? qecraate Wane de | oP be ‘ f rahe el Ser raseree Stor earaer etre & Ar ehrense Tah letasntt saree ie ibe ate 4 farihvee ety ae mentors Biers rir Leet ogeet tre est atthe aed ce nn te Sie : eet esecgnee Roce f venir y AR werwit f weer naire iis ttt ities ae eet ae (uh Or goose yah eyed iis 4} Qee Soe becnedheesty Gh, + eee nt i eile janes . | Wannatiets Ud ede be eae wernt Pest Tare tivents tne toon haa