veratrum viride

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THE BIOLIDGY OF CANADIAN WEEDS. TT.Veratrum viride Ait.GERALD A. MULLIGAN ANd DEREK B. MUNRO

Biosystematics trl.esearch Centre, Ottawa, Ontario, Canada KIA 0C6. Received 6 Oct.1986, accepted 22 Jan. 1987.

MuLLrcAr\, G. A. AND MuNRo, D. B. 1987. The Biology of Canadian weeds.Veratntm vir.ide Art. Can. J. Plant Sci. 67:7'l'7-'786.This paper prorrides a summary of biological data on Veratum viride Ait., falsehellebore. It is a herbaceous perennial, native to wet habitats in North America. Sub-speciesviride ocr:urs primarily in eastern Canada and subspecies eschschohzii (A. Gray)Love and Lrive in the west. Both subspecies have the chromosome numberof n :16,2n = 32.Ingested material of false hellebore is poisonous to humans andlivestock.

Key words: False hellebore, Veratrum viride Att., weed biology[Biologie des ntauvaises herbes du Canada. Veratrum viride Ait.lTitre abrdg6: Veratrum viride Ait.Cette morrographie r6unit les donn6es biologiques concernant Veratrum viride Ait.,v6rAtre vert. C'r:st une herbac6e p6renne des endroits humides en Am6rique du Nord.La sous-espbce viride se rencontre principalement dans l'est du Canada et la sous-espdceeschscholtzii (A, Gray) Love et Love dans I'Ouest. Les deux formes portant le nombrechromosomien n : 16,2n : 32. Le v6rdtre vert est toxique ir I'ingestion pour I'hommeet pour les aninraux de ferme.Mots cl6s: v6rdtre vert (varaire verl) Verotrum viride Ait., biologie des mauvaises herbes

1. Name Stem erect, up to 2 m tall, pubescent onVeratrum viridet Ait. (V. eschscholtzii A. middle and upper part, hollow. Leaves many,Gray) - false hellebore, varaire vert (Alex oval to oblong-elliptic, up to 35 cm long andet ar. 1e80). Liliaceae, rily family, Liliac6es. ::ilj}il,]il;i1i:1 ' :li?#*. j T,il:. dt'.i2. Description and l\ccount of Variation white tomentose beneath. Inflorescence aThe taxonomic status of eastern and western panicle up to 7 dm long, diffuse to denselypopulations of lalse hellebore has been the branched, lower branches usually drooping,subject of some clisagreement among upper branches spreading or erect, panicletaxonomists. However, after studying false branches becoming mostly erect during fruit-hellebore in its natil'e habitat, growing it ing, pedicels to 5 mm long. Perianth yellow-under cultivation and examining numerous green to deep green with six tepals 6-10 mmherbarium collections, we agree with those long, oblong-lanceolate to oblong-elliptic,who consider )Veratrum viride to be one narrowing to a short claw, at least the lowerspecies with trryo geographic subspecies, edges of tepals are pilose, the outer sidesutbsp. viride arnd sulDsp. eschscholtzii (A. generally woolly. Stamens usually 6-9 mmGray) Lrive & llcive. long, shorter than the tepals, ovary usuallyVeratrum virtide - long-lived perennial glabrous. Capsule glabrous, 2-3 cm long.reproducing b5r seeds and crown buds Seeds 8-10 mm long, narrow, surrounded(Fig. l). Rootr;tocks short, thick, erect by a thin papery wing; 20-40 seeds perrhizomes with lvhorls of thick fleshy roots. capsule. The chromosome count ofcan. J. prant Sci. 67: 77,7-786(Jury re87) ; :,ffi'K'-rJ 1lnl. *il 'ff1 i :ilS:

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'7'78 CANADIAN JOURNAL OF PLANT SCIENCEin British Columbia, Washington State,Quebec and New Brunswick. Voucher speci-mens for these counts have been deposited inthe Agriculture Canada Herbarium in Ottawa(DAO acronyms after Holmgren et al.(1981)).

Two subspecies of Veratrum viride occurin North America: subsp. viride (Fig. 18) ineastern North America and subsp.eschscholtzii (fig. lA) in the west. They canusually be recognized by the fcllowingcharacters. Subsp. viride - leaves sparselypubescent below, at least along the veins;lower panicle branches usually drooping withthe middle and upper branches usually erect;stamens generally 6-7 mm long. Subsp.eschscholtzii - leaves usually white tomen-tose below; lower and middle paniclebranches drooping, and upper branchesusually spreading or erect; stamens generally8-9 mm long.The differences between eastern subsp.viride and western subsp. eschscholtzii arelargely quantitative. Western plants tend tohave more open panicles with drooping lowerbranches, and tomentose pubescence on theunderside of the leaves. However, herbariumspecimens examined showed a wide range ofpubescence in plants from both sides of thecontinent. Some collections from the Yukonand Northwest Territories had leaf vestitureindistinguishable from eastern plants. In ad-dition, some western plants had lowerbranches of the inflorescence mostly erectwhereas some eastern plants had some lowerbranches drooping. Youngken (1953) report-ed that western plants growing from Alaskato northern Washington State were morepubescent than plants growing farther south.3. Economic Importance(a) Detrimental - False hellebore is occasion-ally weedy in fields and pastures in easternCanada, especially in the eastern townshipsofQuebec and along the St. John River valleyof New Brunswick. It also grows occasionallyon rangelands in British Columbia. Its repu-tation as a poisonous plant makes itundesirable in these habitats. Cases of

poisoning in mature cattle and sheep are rarewhereas young animals sometimes eat falsehellebore with fatal results. Large quantitiesof false hellebore must be eaten by animalsfor serious poisoning to occur. Chickens havebeen poisoned after ingestion of seeds oryoung shoots. Symptoms of poisoning includesalivation, muscular tremors, low temperatureand death resulting from heart paralysis (Fyles1920; Gress 1935; Reynard and Norton 1942;Kingsbury 1964). Sheep in westernrangelands forage leaves of false helleboreafter frost with apparent immunity (Fylesl92O; Dayton 1960). A recent study(Campbell et al. 1985) showed that false helle-bore contains the steroidal alkaloidjervine inits roots and rhizomes. Jervine is a teratogenand causes deformities in the fetuses of sheep,goats, cattle, chickens and mice.Many cases of human poisoning haveresulted from misuse of medicinal prepara-tions or other accidents. Underhill (1959)reported the poisoning of three boys by falsehellebore in British Columbia. The boys ateblanched stems mistaking it for skunk cabbage(Lysichitum americanum Hult. & St. John).Symptoms included nausea. vomiting. crampsand blurred vision. After 12 - 15 h, thesymptoms disappeared. Anonymous (19'72)reported cases of false hellebore poisoning oltwo people in Nelson and four people inKootenay, British Columbia. Boudreault(1979) wrote that 1 - 3 g ofcrushed dry rootis enough to kill a man. Boivin (1948) relatesthe experience of ingesting false hellebore andsuffering weakness, dizziness, nausea, fevetand chills for about 12 h.(b;) Beneficial - False hellebore was used byIndians in North America in medicine andrituals. Josselyn (1672) reported that youngIndians would elect their chiefs by choosingthe person whose stomach could withstand thepoisoning action of false hellebore root thelongest. Taylor (1940) described false helle-bore use by Cherokee Indians on skin abra-sions, probably for the strong counter-irritanteffect. Turner (1975) reported that coastalIndian groups of British Columbia, although

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MULLIGAN AND MUNRO VERA1RUM V|RIDE ATT

Fig. l. A-D. Veratum viride. A. Flowering specimen of V. viride subsp. eschscftolrzll growing insitu on Vancouver Island, British Columbia, showing many drooping lower panicle branches. B.Flowering specimen of V. viide subsp. vlrlde collected in the Eastern Townships of Quebec and grownin greenhouse, showing a few drooping panicle branches. C. First year seedling (dght side) and dormantoverwintering root bud after one season's growth. D. Habitat of false hellebore in the Eastern Townships,Quebec.

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780 CANADIAN JOURNAL OF PLANT SCIENCErecognizing false hellebore as one of the mostviolently poisonous local plants, used itcautiously as a medicine. Long (1981) andJohnston (1982) noted that false hellebore wasone of several plants used by Indians tocommit suicide. The Blackfoot Indians tookpowdered root for headaches and for itching.Kalm (1979) described how corn seed wassoaked in a decoction of false hellebore rootto protect seeds from birds after planting.Birds would suffer from vertigo, thus fright-ening other birds away. Juice from boiledroots was combed through childrens' hair toeffectively kill lice. Edwards (1980) discussesfolklore and use of false hellebore by BellaCoola Indians in British Columbia. Thisspecies was called Putsk by the Indians andconsidered strong medicine. Roots wereboiled in water for a bath to get rid of scabies,aches, pains and to remove human smell forhunting. Pieces of false hellebore root werealso used as luck charms.Medical extracts of false hellebore wereused to help cure many disorders (Baker1859; Millspaugh 1887; Jefferis and Nichols1894). Osol and Farrar (1955) recorded thatfalse hellebore contains many alkaloids whichhave proven useful in the treatment of hyper-tension disorders in humans. The alkaloids areesters of highly hydroxylated parent alkanola-mine bases and include cevine, germine. pro-toverine. cevadine. veratridine andprotoveratrine (Jacobs and Craig 1944,1945;Fried et af. 1950; Klohs et al. 1952a,b,1953a,b, 1954; Myers et al. 1952, 1955;Kupchan and Gruehfeld 1959). Active prepa-rations caused a decrease in systolic anddiastolic pressure, decreased heart rate andincreased peripheral blood flow (Kauntze andTrounce l95l). The modern formulation,veriloid, was removed from general use forhypotensive therapy about 1961 because ofside effects (Hightower 1979).(c) Legislation - False hellebore is listed inthe noxious weeds act of Manitoba (1917).However, it does not occur in or nearManitoba. It is not listed in other provincialor federal seed or weed lesislation.

4. Geographical DistributionFalse hellebore, native to North America,occurs in both the eastern and western partsof the continent. In eastern Canada, false hele-bore is found in eastern and central NewBrunswick and in southeastern Quebec, theEastern Townships, along the St. LawrenceRiver Basin and occasionally, on the Gasp6Penninsula (Fig. 2). Lcive et al. 1958) andCayouette (1972) reported false helleborefrom northern Saguenay Co., around EllenLake, and Lavoie (1984) reported collectionsfrom Rividre Moisie north of Sept-Iles. In theUnited States, false hellebore is mostly anAppalachian element, ranging south to NorthCarolina and Tennessee, eastward through theNew England States to Maryland, and west-ward to Ohio (Griggs 1938; Pease l9&;Beanet al. 1966; Johnson 1970).In western North America, false helleboreoccurs in south central Alaska and along thepanhandle, eastward into southern and cen-tral Yukon, and in the Mackenzie district ofthe Northwest Territories. Its occurrence isoccasional along coastal British Columbia, be-coming abundant on the Queen Charlotte andVancouver Islands. It is found through theinterior to the Rocky Mountains and foothillsof British Columbia and Alberta (Fig. 2;Hulten 1968; Porsild and Cody 1980; Moss1983). In the western United States, falsehellebore is found in the Olympic and CascadeMountains of Washington State, souththrough Oregon to northern California. It alsogrows in the Rocky Mountains of Montanaand Idaho (Munz and Keck 1959; Peck 196l).5. Habitat(a) Climatic requirements - False helleboregrowth is probably most adversely affectedby a lack of moisture. Its disjointed distribu-tion reflects high moisture requirements. Itcan tolerate sustained winter temperatures to-40'C in alpine areas of eastern and westernCanada. False hellebore inhabits boggy areasin eastern North America offering winterprotection from snow cover. In westernmountainous areas, false hellebore grows to2500 m on moist slopes or in wet meadows.

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MULLIGAN AND MUNRO - VERATRUM VIRIDE AI'I '781

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: -\,\ ltf IO HERBARIUIVI SPECIMENO CHROMOSOME COUNT -/-LFig. 2. Distribution map of false hellebore in Canada and adjacent United States and locations of plantsused for chromosome counts by G. A. Mulligan and D. B. Munro (from herbarium specimens in CAN,DAO. MT).It grows in areas where the annual rainfall isabove 60 cm per annum in eastern Canada butin northwestern Canada it will grow whereannual rainfall is as low as 20 cm per annum(Anonymous 1974).(b\ Substratum - False hellebore srows ona variety of substrates. lrom organiCto sandyloam soils. Taylor (1953) found that falsehellebore flourished in very wet, well-drainedsoil of schist origin in western NorthAmerica. It also was found on fine volcanicash in the Cascade Mountains. Soils were lowin nitrogen and phosphorus and had moder-ate potassium. The pH ranged from 4.5 to 5.9on Alaskan sites and was 6.6 on Mt. Baker.Munro (unpubl. data) tested soil pH for l8collection sites of false hellebore in theprovinces of New Brunswick and Quebec.The pH ranged from 4.5 to 6.9 on soils vary-ing from sandy loam to organic humous.Lavoie (1984) found false hellebore on soils

with pH 5.4 in Quebec. Taylor (1956) grewfalse hellebore in pots of sandy loam to whichnitrogen, phosphorus and potassium wereadded in various combinations. Additions ofphosphorus and potassium did not affectgrowth. Plant growth was suppressed by ad-ditions ofnitrogen, especially on fertile sandyloam.(c) Communities in which species occurs -False hellebore generally inhabits moist sitesalong watercourses, around lakes, marshedges, alpine meadows, and on seepage areasin western mountains. It also inhabits uncul-tivated fields and pastureland in eastern NorthAmerica (Fig. 1D). Pastures in the EasternTownships of Quebec are often close-croppedby cattle except for the tall stalks of falsehellebore. Common associated plants areThalictrum dioicum L., Fragaria virginianaDuchesne, Rubus pubescens Raf. and Alnusrugosa (Du Roi) Spreng. (Munro, pers. obs.).

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't82 CANADIAN JOURNAL OF PLANT SCIENCEIn the White Mountains of New Hampshirefalse hellebore is frequent in springy placesabove the treeline (Pease 1964). It is foundwhere drainage is poor and trees are oftendrowned by meltwater in small bogs. As-sociated plants are Calamagrostis canadensis(Michx.) Nutt., Cornus canadensis L.,Kalmia polifulia Wang., and Vacciniumorycoccus L. (Griggs 1938).Franklin and Dryness (1973) describe falsehellebore as an important component of subal-pine meadow communities of the CascadeMountains in western-Washington State. TheValeriana sitchensis Bong-Veratrum viride(Sitka valerian, false hellebore) ecologicalzone is a lush community with associated her-baceous species which generally stand I m ormore in height. Associated plants are Lupinuslatfolius Agardh, Carex spectabilis Dewey,Castilleja parviflora Bongard, Erythroniumspp., Anemone occidentalis Watson,Polygonum bistortoides Pursh, Mitellabreweri Gray, Epilobium angustfolium L.,and Heracleum lanatum Mx. This communityoccurs on steep, well-watered slopes of vary-ing exposure that tend to have deep soils anda long growing season. Avalanches frequentlyoccur on these slopes. The communitysurvives because the major constituent plantspecies survive winter as underground partsprotected from snow creep and avalanching.The Abies amabilis Dotglas-Veratrum viride(Pacific silver fir, false hellebore) communi-ty occurs in the Cascade Mountains ofsouthern Washington and northern OregonStates. This ecological zone is characterizedby a lush herbaceous understory on deep soilssubjected to temporary water seepage.6. HistoryFalse hellebore, native to North America, hasnow essentially the same Canadian range asthat described by Macoun (1886). However,more recent collecting has extended theknown range of this species in a few remoteareas. Abundant cleared land has no doubtdecreased the concentration ofthis species inagricultural areas due to the lowering of watertables. However, colonies of false helleborevisited by Mulligan and Calder (pers. obs.)

in the Eastern Townships during 1950 werestill present 31 yr later (Munro, pers. obs.).7. Growth and DeveloPment(a) Morphology - False hellebore is one ofthe first plants to emerge from the soil in thespring. These early shoots, from overwinter-ing rhizomes, are thick and robust and growrapidly. They provide false hellebore with anearly advantage for successful competitionwith other vegetation.(b) Perennation - False hellebore is peren-nial and overwinters as underground rhi-zomes. Taylor (1956) reported that rhizomeshave pronounced winter dormancy and re-quire cold treatment to break it. False helle-bore has an upright rhizome with whorls offleshy roots and is surmounted by a largecrown bud. A small second bud, adjacent tothe main bud, will initiate the subsequentyear's growing shoot. Meristematic activitiesoccur at the junction of the crown bud andrhizome. A new whorl of roots is added tothe top of the root system each growing sea-son. Some rhizomes have two concentriccircles of root primordia suggesting periodicinitiation. False hellebore crowns examinedin Alaska were estimated to be over a centuryold. New growth of transplanted rhizomeswas usually restricted to top growth. Plantsestablished best when they were transplantedwith their profuse network of lateral rootsintact.(c) Physiological data - No physiologicaldata are available.(d) Phenology - Flowering commencesduring the first half of June in eastern Canadaand mature seeds are produced within a monthof flowering. Plants in western Canada beginflowering in the first half of July and seedsmature about a month later. Plants at thenorthern limits of distribution may not floweruntil early August. Seeds are shed gradually,with the capsules dehiscing and the seedsbeing dispersed by wind currents.(e) Mycorrhiza - Thete are no reports ofmycorrhizal associations with false hellebore.

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MULLIGAN AND MUNRO - VEMTRUM VIRIDE ATT '7838. Reproduction(a) Floral biology - Taylor (1956) found thatfalse hellebore usually did not flower until7 -lO yr after germination. Subsequent flower-ing occurred in occasional years. Floweringmight or might not occur when the crownduplicated. False hellebore has protandrousflowers with exposed nectar secreted by thethickened bases of the tepals. This suggeststhat false hellebore is cross-pollinated, pos-sibly by insects. However, we have not madefield observations and could find no informa-tion on its floral biology in the literature.(b) Seed production and dispersal - Taylor(1956) noted that false hellebore producedseed intermittently; heavy seed-crop yearswere generally followed by five or more yearsof low seed production. The authors believethat wind dispersal of the 20 - 40 seeds percapsule is aided by the papery wing surround-ing the seed.(c) Viability of seeds and germination -Taylor (1956) reported that false helleboreseeds have pronounced dormancy. He foundthat stratification at 3 - 5 C for 4 - 4.5 moresulted in prompt germination. Stratifi cationfor shorter periods resulted in slow or lowgermination. Higher stratification tempera-tures resulted in poorer germination. Germi-nation in all tests ranged from22 to 85% . Theseeds sprouted at about 5 C. Constant sub-freezing temperatures were ineffective inbreaking seed dormancy. Seeds subjected toalternating temperatures between 5 C andsubfreezing every 2 wk germinated in severalmonths. Stratihed seed subjected to weak lightgerminated little better than seed exposed onlyto the dark. Seeds sealed in containers con-taining perlite and held at 5'C for 2 yr andthen held 4 mo at 2oC, germinated afterexposure to light. These seeds had a 46%viability. Unsealed seeds held at 5'C declinedin viability by the end of I yr with only a smallnumber germinating after 2 yr. Seeds germi-nated best when covered with 2 mm of sand,vermiculite or shredded sphagnum. Seedssown outdoors in plots and held over thewinter produced seedlings (Fig. lC) as soon

as the snow melted. Young seedlings weresensitive to direct sunlight; the tops dyingback early in the growing season. Plantsdeveloped only one true leaf the frrst year.(d) Vegetative reproduction - False helleboreoccasionally produces forked rhizomes withindependent crowns. When a flowering stalkis produced one or several suppressed budsdevelop to produce independent crowns.Experimental trials were carried out onsevered crowns to test vegetative reproduc-tion. It was found that a low percentage ofplants reproduced by this method (Taylor19s6).9. HybridsThere is no evidence of any natural hybridi-zation involving this species.10. Population DynamicsMulligan and Calder (pers. obs.) surveyedfalse hellebore populations in the EasternTownships of Quebec during 1950. Falsehellebore occasionally grew in extensivepopulations to the exclusion of otherherbaceous plants. In one 6-m x 6-m quad-rat in Brome County, 214 plants were present.Taylor (1956) studied seedling germinationand found that stand densities of 100 - 150false hellebore seedlings per 0.3 m2 were notlimiting to growth of the seedlings.11. Response to Herbicides and OtherChemicalsInformation is not available on control of falsehellebore by herbicides. However, control ofthe closely related western false hellebore(Veratrum califurnicum Durand) had beenstudied in Idaho (Williams and Kreps 1970;Williams and Cronin 1981). Dense plots ofwestern false hellebore were treated in earlyJuly with amine salts of 2,4-D at 2.2 kg hajust as the last leaf had expanded. A follow-up spray applied a year later removed allremaining plants. Control of western falsehellebore in these plots after 10 yr was 100%.12. Response to Other HumanManipulationsFalse hellebore roots can be grubbed out inspring when the soil is wet. Where land is

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784 CANADIAN JOURNAL OF PLANT SCIENCEbadly overrun, cultivation and drainage willreduce the number ofplants. Seeding can beprevented by early cutting. Mountainous landscan be treated as pasture if mature animalsare allowed to graze after plants are no longeryoung and succulent (Fyles 1920).13. Response to Parasites(a') Insects and other nondomesticorganisms - No records were found of in-sects on false hellebore. A collection of falsehellebore from Shefford County, Quebec wasobserved to have aphids on the inflorescence.(b) Microorganisms and viruses - Conners(1967) listed the following fungi on false hel-lebore in Canada: Phyllosticta melanoplacaThum. (Coelomycetes) in B.C.; Cerco-sporella veratri Pk. (Hyphomycetes) in B.C;Puccinia veratri Duby (Uredinales) in B.C.and Que.; Sclerotium durum Fr. (Basid.Imperf.),Ilelotium spp., Patinellaaloysii-sabaudiae Sacc. (Disco. Inoperc.)were found on Alaskan material of falsehellebore. The United States Department ofAgriculture (1960) lists a number of fungi onfalse hellebore in the United States:Colletrotrichum liliacearum (Schw.) Ferr. inN.J.; Cylindrosporum veratrinum Sacc. &Wint. in N.Y.; Cercosporella terminalis Pk.in N.Y.; Phyllochora melanoplaca (Desm.)Sacc. in N.Y.(c) Higher plant parasites - There are norecords of higher plant parasites on falsehellebore.

ACKNOWLEDGMENTSWe would like to thank the following people: CalSime, Warden, Yoho National Park and W. J.Cody, of BRC, for supplying additional live plants;Drs. A. Stahevitch and S. L Warwick of BRC, forproviding helpful suggestions for the manuscript;W. Wojtas of BRC, for preparing chromosomeslides.Alex, J. F., Cayouette, R. and Mulligan' G. A.1980. Common and botanical names of weeds in

Canada/Noms populaires et scientifiques desplantes nuisibles du Canada. Agriculture Canada,Ottawa, Ont. Publ. 1397 (rev.) 132 pp.Anonymous. 1972. Beware - poisonous plant.The Daily News, Nelson, British Columbia 7l(No. 99): 2.Anonymous. 1974.The national atlas ofCanada.4th rev. ed. MacMillan Co., Toronto, Ont.266 pp.Baker, P. D. 1859. Veratrum viride in chorea andother convulsive diseases. South. Med. Surg. 15 : 4.Bean, R. C., Richards, C. D. and Hyland, F.1966. Checklist of the vascular plants of Maine(rev.). Josselyn Bot. Soc. Orono, Maine. BuIl. 8:71 pp.Boivin, B. 1948. Centurie de plantes Canadiennes.Nat. Can. 75: 202-227.Boudreault, M. 1979. Guide pratique des plantesmedicinales du Quebec. Marcel Broquet Inc., LaPrairie, Qu6bec. 205 pp.Campbell, M. A., Brown, K. S., Hassell, J. R.,Horigan, E. A. and Keeler, R. F. 1985.Inhibition of limb chondrogenesis by a Veratrumalkaloid; temporal specificity in vivo and in vitro.Dev. Biol. lll 464-470Cayouette, R. 1972. Etudes sur la flore deSaguenay. VI. Les Liliac6es et leur distribution.Nat. Can. 99:457-468.Conners, l.L. 1967. An annotated index of plantdiseases in Canada and fungi recorded on plantsin Alaska, Canada and Greenland. AgricultureCanada, Ottawa, Ont. Publ. 1251. 381 pp.Dayton, W. A. 1960. Notes on western rangeforbes, USDA For. Ser. Washington. Agric'Handb. 16l:254 pp.Edwards, G. T. 1980. Bella Coola Indian andEuropean medicines. Beaver (Winter) 1980: 5- I I .Franklin, J. F. and Dyrness, C. T. 1973. Naturalvegetation of Oregon and Washington. USDA For.Ser. Gen. Tech. Rep. PNW-8. 417 PP.Fried, J., White, H. L. and Wintersteiner, O.1950. The hypotensive principles of Veratrumviride. I. Am. Chem. Soc.72: 4621-4630'Fyles, F. 1920. Principle poisonous plants ofCanada. Agriculture Canada Experimental FarmsBull. 329 (2nd series). 120 PP.Gress, E. M. 1935. Poisonous plants ofPennsylvania. Penn. Dep. Agric. Gen. Bull.531.5l pp.Griggs, R. F. 1938. Mt. Washington ObservatoryNews Bull. 4: 5.Hightower, C.E. 1979. Plants that kill and cure.

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