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 Veratntm vir.ide Art. Can. J. Plant Sci. 67:7'l'7-'786.

of

Canadian weeds.

This paper prorrides a summary of biological data on Veratum viride Ait., false

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hellebore. It is a herbaceous perennial, native to wet habitats in North America. Subspeciesviride ocr:urs primarily in eastern Canada and subspecies eschschohzii (A. Gray) Love and Lrive in the west. Both subspecies have the chromosome number of n :16,2n = 32.Ingested material of false hellebore is poisonous to humans and livestock.

Key words: False hellebore, Veratrum viride Att., weed biology [Biologie des ntauvaises herbes du Canada. Veratrum viride Ait.l

Titre 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-espdce eschscholtzii (A, Gray) Love et Love dans I'Ouest. Les deux formes portant le nombre chromosomien n 16,2n 32. Le v6rdtre vert est toxique ir I'ingestion pour I'homme

et pour les aninraux de ferme.

Mots cl6s: v6rdtre vert (varaire verl) Verotrum viride Ait., biologie des mauvaises herbes

Stem erect, up to 2 m tall, pubescent on Veratrum 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 and et ar. 1e80). Liliaceae, rily family, Liliac6es.

1. Name

2. Description and l\ccount of

::ilj}il,]il;i1i:1"'":li?#*.""j"T,il:."dt'.i Variation white tomentose beneath. Inflorescence a

false

The taxonomic status of eastern and western panicle up to 7 dm long, diffuse to densely populations of lalse hellebore has been the branched, lower branches usually drooping,

subject

of

some clisagreement among upper branches spreading or erect, panicle

branches becoming mostly erect during fruiting, pedicels to 5 mm long. Perianth yellowgreen to deep green with six tepals 6-10 mm

hellebore in its natil'e habitat, growing it under cultivation and examining numerous herbarium collections, we agree with those who consider )Veratrum viride to be one species with trryo geographic subspecies, utbsp. viride arnd sulDsp. eschscholtzii (A. Gray) Lrive & llcive. Veratrum virtide - long-lived perennial reproducing b5r seeds and crown buds (Fig. l). Rootr;tocks short, thick, erect rhizomes with lvhorls of thick fleshy roots.
can. J. prant Sci. 67: 77,7-786(Jury

taxonomists. However, after studying

long, oblong-lanceolate to oblong-elliptic,

narrowing to a short claw, at least the lower edges of tepals are pilose, the outer sides generally woolly. Stamens usually 6-9 mm long, shorter than the tepals, ovary usually glabrous. Capsule glabrous, 2-3 cm long.
Seeds 8-10 mm long, narrow, surrounded

by a thin papery wing; 20-40 seeds per capsule. The chromosome count of

re87)
777

;":,ffi'K'-rJ"1lnl."*il"'ff1"i":ilS:

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CANADIAN JOURNAL OF PLANT SCIENCE

in British

Columbia, Washington

State,

Quebec and New Brunswick. Voucher specimens for these counts have been deposited in the Agriculture Canada Herbarium in Ottawa (DAO acronyms after Holmgren et al.

(1981)).

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 103.241.206.34 on 12/02/13 For personal use only.

eastern North America and subsp. eschscholtzii (fig. lA) in the west. They can usually be recognized by the fcllowing
characters. Subsp. viride - leaves sparsely pubescent below, at least along the veins; lower panicle branches usually drooping with the middle and upper branches usually erect; stamens generally 6-7 mm long. Subsp. eschscholtzii - leaves usually white tomentose below; lower and middle panicle

Two subspecies of Veratrum viride occur in North America: subsp. viride (Fig. 18) in

poisoning in mature cattle and sheep are rare whereas young animals sometimes eat false hellebore with fatal results. Large quantities of false hellebore must be eaten by animals for serious poisoning to occur. Chickens have been poisoned after ingestion of seeds or young shoots. Symptoms of poisoning include salivation, muscular tremors, low temperature and death resulting from heart paralysis (Fyles 1920; Gress 1935; Reynard and Norton 1942;

Kingsbury 1964). Sheep in western rangelands forage leaves of false hellebore after frost with apparent immunity (Fyles l92O; Dayton 1960). A recent study

branches drooping, and upper branches

usually spreading or erect; stamens generally

8-9 mm long. The differences between eastern subsp. viride and western subsp. eschscholtzii are
largely quantitative. Western plants tend to
have more open panicles with drooping lower branches, and tomentose pubescence on the underside of the leaves. However, herbarium

(Campbell et al. 1985) showed that false hellebore contains the steroidal alkaloidjervine in its roots and rhizomes. Jervine is a teratogen and causes deformities in the fetuses of sheep, goats, cattle, chickens and mice. Many cases of human poisoning have resulted from misuse of medicinal preparations or other accidents. Underhill (1959) reported the poisoning of three boys by false hellebore in British Columbia. The boys ate blanched stems mistaking it for skunk cabbage (Lysichitum americanum Hult. & St. John). Symptoms included nausea. vomiting. cramps

specimens examined showed a wide range of pubescence in plants from both sides of the continent. Some collections from the Yukon and Northwest Territories had leaf vestiture indistinguishable from eastern plants. In ad-

and blurred vision. After 12

branches

dition, some western plants had lower of the inflorescence mostly erect

symptoms disappeared. Anonymous (19'72) reported cases of false hellebore poisoning ol two people in Nelson and four people in Kootenay, British Columbia. Boudreault (1979) wrote that 1 - 3 g ofcrushed dry root
is enough to

15

h,

the

kill a man. Boivin (1948) relates

whereas some eastern plants had some lower branches drooping. Youngken (1953) reported that western plants growing from Alaska

the experience of ingesting false hellebore and

suffering weakness, dizziness, nausea, fevet and chills for about 12 h.

(b;)

to

northern Washington State were more

Beneficial

pubescent than plants growing farther south.

3. Economic Importance
(a) Detrimental

ally weedy in fields and pastures in eastern Canada, especially in the eastern townships
ofQuebec and along the St. John River valley of New Brunswick. It also grows occasionally on rangelands in British Columbia. Its repu-

False hellebore is occasion-

Indians in North America in medicine and rituals. Josselyn (1672) reported that young Indians would elect their chiefs by choosing
the person whose stomach could withstand the poisoning action of false hellebore root the

False hellebore was used by

tation as a poisonous plant makes it

undesirable

in

these habitats. Cases of

longest. Taylor (1940) described false hellebore use by Cherokee Indians on skin abrasions, probably for the strong counter-irritant effect. Turner (1975) reported that coastal Indian groups of British Columbia, although

MULLIGAN AND

MUNRO

VERA1RUM V|RIDE ATT

7'79

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l. A-D. Veratum viride. A. Flowering specimen of V. viride subsp. eschscftolrzll growing in situ 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 grown
Fig.
in greenhouse, showing a few drooping panicle branches. C. First year seedling (dght side) and dormant overwintering root bud after one season's growth. D. Habitat of false hellebore in the Eastern Townships,
Quebec.

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CANADIAN JOURNAL OF PLANT SCIENCE

recognizing false hellebore as one of the most

violently poisonous local plants, used it cautiously as a medicine. Long (1981) and
Johnston (1982) noted that false hellebore was

4. Geographical Distribution False hellebore, native to North America,

occurs in both the eastern and western parts of the continent. In eastern Canada, false helebore is found in eastern and central New Brunswick and in southeastern Quebec, the Eastern Townships, along the St. Lawrence River Basin and occasionally, on the Gasp6 Penninsula (Fig. 2). Lcive et al. 1958) and Cayouette (1972) reported false hellebore from northern Saguenay Co., around Ellen Lake, and Lavoie (1984) reported collections from Rividre Moisie north of Sept-Iles. In the United States, false hellebore is mostly an Appalachian element, ranging south to North
Carolina and Tennessee, eastward through the

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of several plants used by Indians to commit suicide. The Blackfoot Indians took powdered root for headaches and for itching. Kalm (1979) described how corn seed was soaked in a decoction of false hellebore root to protect seeds from birds after planting. Birds would suffer from vertigo, thus frightening other birds away. Juice from boiled roots was combed through childrens' hair to
one
effectively kill lice. Edwards (1980) discusses folklore and use of false hellebore by Bella Coola Indians in British Columbia. This

"Putsk" by the Indians and considered strong medicine. Roots were

species was called

New England States to Maryland, and westward to Ohio (Griggs 1938; Pease l9&;Bean

boiled in water for a bath to get rid of scabies, aches, pains and to remove human smell for hunting. Pieces of false hellebore root were also used as luck charms. Medical extracts of false hellebore were used to help cure many disorders (Baker 1859; Millspaugh 1887; Jefferis and Nichols 1894). Osol and Farrar (1955) recorded that false hellebore contains many alkaloids which have proven useful in the treatment of hypertension disorders in humans. The alkaloids are esters of highly hydroxylated parent alkanolamine bases and include cevine, germine. pro-

et al. 1966; Johnson 1970). In western North America, false hellebore

occurs in south central Alaska and along the panhandle, eastward into southern and central Yukon, and in the Mackenzie district of the Northwest Territories. Its occurrence is occasional along coastal British Columbia, becoming abundant on the Queen Charlotte and Vancouver Islands. It is found through the interior to the Rocky Mountains and foothills

of British Columbia

and Alberta (Fig. 2; Hulten 1968; Porsild and Cody 1980; Moss 1983). In the western United States, false

toverine. cevadine. veratridine

and protoveratrine (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-

hellebore is found in the Olympic and Cascade Mountains of Washington State, south through Oregon to northern California. It also

grows in the Rocky Mountains of Montana

and Idaho (Munz and Keck 1959; Peck 196l).

rations caused a decrease in systolic and diastolic pressure, decreased heart rate and increased peripheral blood flow (Kauntze and Trounce l95l). The modern formulation, veriloid, was removed from general use for
hypotensive therapy about 1961 because of side effects (Hightower 1979).

5. Habitat
(a) Climatic requirements - False hellebore growth is probably most adversely affected by a lack of moisture. Its disjointed distribution reflects high moisture requirements. It can tolerate sustained winter temperatures to

the noxious weeds act of Manitoba (1917).

However,

(c) Legislation

False hellebore is listed in

-40'C in alpine areas of eastern and western Canada. False hellebore inhabits boggy areas

it

does not occur

in or

near

in eastern North America offering winter protection from snow cover. In western
mountainous areas, false hellebore grows to 2500 m on moist slopes or in wet meadows.

Manitoba. It is not listed in other provincial or federal seed or weed lesislation.

MULLIGAN AND MUNRO

VERATRUM VIRIDE

AI'I

'781

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-t)

Fig. 2. Distribution map of false hellebore in Canada and adjacent United States and locations of plants used for chromosome counts by G. A. Mulligan and D. B. Munro (from herbarium specimens in CAN,

: -\,\

lt

O HERBARIUIVI SPECIMEN O CHROMOSOME COUNT

-/-L

DAO. MT).

It grows in

areas where the annual rainfall is above 60 cm per annum in eastern Canada but

with pH 5.4 in Quebec. Taylor (1956) grew

false hellebore in pots of sandy loam to which

in northwestern Canada it will grow where

annual rainfall is as low as 20 cm per annum (Anonymous 1974).

nitrogen, phosphorus and potassium were added in various combinations. Additions of

phosphorus and potassium did not affect growth. Plant growth was suppressed by additions ofnitrogen, especially on fertile sandy
loam.

(b\ Substratum
a

- False hellebore srows on variety of substrates. lrom organiCto sandy

North

loam soils. Taylor (1953) found that false

hellebore flourished in very wet, well-drained

soil of schist origin in western

America. It also was found on fine volcanic ash in the Cascade Mountains. Soils were low

in nitrogen and phosphorus and had moderate potassium. The pH ranged from 4.5 to 5.9 on Alaskan sites and was 6.6 on Mt. Baker. Munro (unpubl. data) tested soil pH for l8

collection sites

of

false hellebore

in

the

(c) Communities in which species occurs False hellebore generally inhabits moist sites along watercourses, around lakes, marsh edges, alpine meadows, and on seepage areas in western mountains. It also inhabits uncultivated fields and pastureland in eastern North America (Fig. 1D). Pastures in the Eastern Townships of Quebec are often close-cropped by cattle except for the tall stalks of false

provinces of New Brunswick and Quebec. The pH ranged from 4.5 to 6.9 on soils varying from sandy loam to organic humous. Lavoie (1984) found false hellebore on soils

hellebore. Common associated plants

Thalictrum dioicum

are

L., Fragaria virginiana

Duchesne, Rubus pubescens Raf. and Alnus rugosa (Du Roi) Spreng. (Munro, pers. obs.).

't82

CANADIAN JOURNAL OF PLANT SCIENCE

In the White Mountains of New Hampshire false hellebore is frequent in springy places above the treeline (Pease 1964). It is found where drainage is poor and trees are often drowned by meltwater in small bogs. As-

in the Eastern Townships during 1950 were still present 31 yr later (Munro, pers. obs.).

7. Growth and DeveloPment

(a) Morphology False hellebore is one of from the soil in the plants emerge to first the spring. These early shoots, from overwintering rhizomes, are thick and robust and grow rapidly. They provide false hellebore with an

(Michx.) Nutt., Cornus canadensis L., Kalmia polifulia Wang., and Vaccinium
Can. J. Plant Sci. Downloaded from pubs.aic.ca by 103.241.206.34 on 12/02/13 For personal use only.
orycoccus L. (Griggs 1938). Franklin and Dryness (1973) describe false hellebore as an important component of subalpine meadow communities of the Cascade Mountains in western-Washington State. The Valeriana sitchensis Bong-Veratrum viride (Sitka valerian, false hellebore) ecological zone is a lush community with associated herbaceous species which generally stand I m or more in height. Associated plants are Lupinus latfolius Agardh, Carex spectabilis Dewey, Castilleja parviflora Bongard, Erythronium

sociated plants are Calamagrostis canadensis

early advantage for successful competition with other vegetation.

nial and overwinters as underground rhizomes. Taylor (1956) reported that rhizomes

(b) Perennation

False hellebore is peren-

have pronounced winter dormancy and require cold treatment to break it. False hellebore has an upright rhizome with whorls of fleshy roots and is surmounted by a large crown bud. A small second bud, adjacent to

Polygonum bistortoides Pursh, Mitella

breweri Gray, Epilobium angustfolium L., and Heracleum lanatum Mx. This community
occurs on steep, well-watered slopes of varying exposure that tend to have deep soils and a long growing season. Avalanches frequently

spp., Anemone occidentalis

Watson,

the main bud,

will initiate the subsequent

occur on these slopes. The community

survives because the major constituent plant species survive winter as underground parts
protected from snow creep and avalanching. The Abies amabilis Dotglas-Veratrum viride (Pacific silver fir, false hellebore) communi-

year's growing shoot. Meristematic activities occur at the junction of the crown bud and rhizome. A new whorl of roots is added to the top of the root system each growing season. Some rhizomes have two concentric circles of root primordia suggesting periodic initiation. False hellebore crowns examined in Alaska were estimated to be over a century old. New growth of transplanted rhizomes was usually restricted to top growth. Plants
established best when they were transplanted with their profuse network of lateral roots

occurs in the Cascade Mountains of southern Washington and northern Oregon States. This ecological zone is characterized by a lush herbaceous understory on deep soils subjected to temporary water seepage.

ty

intact.

(c) Physiological data

data are available.

No physiological
commences

6. History
False hellebore, native to North America, has

now essentially the same Canadian range as that described by Macoun (1886). However, more recent collecting has extended the known range of this species in a few remote areas. Abundant cleared land has no doubt decreased the concentration ofthis species in
agricultural areas due to the lowering of water tables. However, colonies of false hellebore visited by Mulligan and Calder (pers. obs.)

during the first half of June in eastern Canada and mature seeds are produced within a month of flowering. Plants in western Canada begin flowering in the first half of July and seeds mature about a month later. Plants at the northern limits of distribution may not flower until early August. Seeds are shed gradually, with the capsules dehiscing and the seeds being dispersed by wind currents.

(d) Phenology - Flowering

(e) Mycorrhiza - Thete are no reports of mycorrhizal associations with false hellebore.

MULLIGAN AND MUNRO

VEMTRUM VIRIDE

ATT

'783

8. Reproduction

as the snow melted. Young seedlings were

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 103.241.206.34 on 12/02/13 For personal use only.

- Taylor (1956) found that false hellebore usually did not flower until 7 -lO yr after germination. Subsequent flowering occurred in occasional years. Flowering might or might not occur when the crown duplicated. False hellebore has protandrous flowers with exposed nectar secreted by the thickened bases of the tepals. This suggests that false hellebore is cross-pollinated, possibly by insects. However, we have not made field observations and could find no information on its floral biology in the literature.
(b) Seed production and dispersal Taylor (1956) noted that false hellebore produced seed intermittently; heavy seed-crop years were generally followed by five or more years of low seed production. The authors believe that wind dispersal of the 20 - 40 seeds per capsule is aided by the papery wing surround-

(a) Floral biology

back early

sensitive to direct sunlight; the tops dying in the growing season. Plants developed only one true leaf the frrst year.

(d) Vegetative reproduction - False hellebore occasionally produces forked rhizomes with independent crowns. When a flowering stalk is produced one or several suppressed buds develop to produce independent crowns.

Experimental trials were carried out on severed crowns to test vegetative reproduc-

tion. It was found that a low percentage of

plants reproduced by this method (Taylor
19s6).

9. Hybrids
There is no evidence of any natural hybridi-

zation involving this species. 10. Population Dynamics

ing the

seed.

(c) Viability of seeds and germination Taylor (1956) reported that false hellebore
seeds have pronounced

Mulligan and Calder (pers. obs.) surveyed false hellebore populations in the Eastern Townships of Quebec during 1950. False hellebore occasionally grew in extensive

populations

dormancy. He found

that stratification at 3 - 5"C for 4 - 4.5 mo resulted in prompt germination. Stratifi cation for shorter periods resulted in slow or low

herbaceous plants. In one 6-m x 6-m quadrat in Brome County, 214 plants were present.

to the exclusion of

other

germination. Higher stratification temperatures resulted in poorer germination. Germination in all tests ranged from22 to 85% . The seeds sprouted at about 5"C. Constant sub-

Taylor (1956) studied seedling germination and found that stand densities of 100 - 150 false hellebore seedlings per 0.3 m2 were not limiting to growth of the seedlings.

11. Response to Herbicides and Other

Chemicals Information is not available on control of false hellebore by herbicides. However, control of the closely related western false hellebore

freezing temperatures were ineffective in

breaking seed dormancy. Seeds subjected to alternating temperatures between 5"C and
subfreezing every 2 wk germinated in several months. Stratihed seed subjected to weak light germinated little better than seed exposed only to the dark. Seeds sealed in containers containing perlite and held at 5'C for 2 yr and

(Veratrum califurnicum Durand) had been

studied in Idaho (Williams and Kreps 1970; Williams and Cronin 1981). Dense plots of western false hellebore were treated in early July with amine salts of 2,4-D at 2.2 kg ha just as the last leaf had expanded. A followup spray applied a year later removed all remaining plants. Control of western false hellebore in these plots after 10 yr was 100%.

4 mo at 2oC, germinated after exposure to light. These seeds had a 46%

then held viability. Unsealed
seeds held at 5'C declined in viability by the end of I yr with only a small number germinating after 2 yr. Seeds germinated best when covered with 2 mm of sand,

12. Response to Other Human

vermiculite or shredded sphagnum. Seeds sown outdoors in plots and held over the
winter produced seedlings (Fig. lC) as soon

Manipulations
False hellebore roots can be grubbed out in

spring when the soil is wet. Where land is

784

CANADIAN JOURNAL OF PLANT SCIENCE

badly overrun, cultivation and drainage will reduce the number ofplants. Seeding can be prevented by early cutting. Mountainous lands can be treated as pasture if mature animals are allowed to graze after plants are no longer young and succulent (Fyles 1920).

Canada/Noms populaires

et

scientifiques des

plantes 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 and other 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 plantes
medicinales du Quebec. Marcel Broquet Inc., La Prairie, Qu6bec. 205 pp. Campbell, M. A., Brown, K. S., Hassell, J. R.,

Can. J. Plant Sci. Downloaded from pubs.aic.ca by 103.241.206.34 on 12/02/13 For personal use only.

13. Response to Parasites (a') Insects and other nondomestic organisms - No records were found of insects on false hellebore. A collection of false hellebore from Shefford County, Quebec was
observed to have aphids on the inflorescence.

(b) Microorganisms and viruses - Conners (1967) listed the following fungi on false hellebore in Canada: Phyllosticta melanoplaca

Thum. (Coelomycetes)

in B.C.;

Cerco-

sporella veratri Pk. (Hyphomycetes) in B.C; Puccinia veratri Duby (Uredinales) in B.C.

Horigan,

E. A. and

Keeler,

R. F.

1985.

Fr. (Basid. Imperf.),Ilelotium spp., Patinella

and Que.; Sclerotium durum

Inhibition of limb chondrogenesis by a Veratrum alkaloid; temporal specificity in vivo and in vitro.

aloysii-sabaudiae Sacc. (Disco. Inoperc.) were found on Alaskan material of false hellebore. The United States Department of Agriculture (1960) lists a number of fungi on

Dev. Biol. lll 464-470 Cayouette, R. 1972. Etudes sur

Saguenay.

la flore de

VI.

Les Liliac6es et leur distribution.

1967. An annotated index of plant

Nat. Can. 99:457-468.

Conners,
diseases

l.L.

false hellebore in the United

States:

in Canada and fungi recorded on plants

Colletrotrichum liliacearum (Schw.) Ferr. in N.J.; Cylindrosporum veratrinum Sacc. & Wint. in N.Y.; Cercosporella terminalis Pk. in N.Y.; Phyllochora melanoplaca (Desm.) Sacc. in N.Y.

Canada and Greenland. Agriculture Canada, Ottawa, Ont. Publ. 1251. 381 pp. Dayton, W. A. 1960. Notes on western range

in Alaska,

forbes, USDA For. Ser. Washington. Agric'

Handb. 16l:254 pp.

(c) Higher plant parasites - There are no records of higher plant parasites on false
hellebore.

T. 1980. Bella Coola Indian and European medicines. Beaver (Winter) 1980: 5- I I . Franklin, J. F. and Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. USDA For.
Edwards, G.
Ser. Gen. Tech. Rep. PNW-8. 417 PP. Fried, J., White, H. L. and Wintersteiner, O. 1950. The hypotensive principles of Veratrum viride. I. Am. Chem. Soc.72: 4621-4630' Fyles, F. 1920. Principle poisonous plants of

ACKNOWLEDGMENTS
We would like to thank the following people: Cal Sime, Warden, Yoho National Park and W. J. Cody, of BRC, for supplying additional live plants; Drs. A. Stahevitch and S. L Warwick of BRC, for providing helpful suggestions for the manuscript; W. Wojtas of BRC, for preparing chromosome
slides.

Bull. 329 (2nd series).

Canada. Agriculture Canada Experimental Farms 120 PP.

Gress,

E. M. 1935. Poisonous plants

of

Pennsylvania. Penn. Dep. Agric. Gen. Bull.531.

5l

pp.

Griggs, R. F. 1938. Mt. Washington Observatory Alex, J. F., Cayouette, R. and Mulligan' G. A. 1980. Common and botanical names of weeds in
News Bull. 4: 5. Hightower, C.E. 1979. Plants that kill and cure.

MULLIGAN AND MUNRO

VERATRUM VIRIDE AIT.

785

Vet. Hum. Toxicol. 2l:360-362 Hitchcock, C. L., Cronquist, A., Ownbey, M. and Thompson, J, W. 1969. Vascular plants of the Pacific Northwest. Part I. University of
Washington Press, Seattle, Washington, 914 pp.

Holmgren, P. K., Keuken, W. and Schofield, E. K. 1981. Index herbariorum. Part I. The herbaria of the world. 7th ed. Dr. W. Junk B. V.
Publishers. The Hague/Boston. 452 pp. Hulten, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press, Stanford,

of Veratrum eschscholtzii Gray. L The glycosides. J. Am. Chem. Soc. 75: 2133-2135. Klohs, M. W., Draper, M. D., Keller, F., Malesh, W. and Petracek, F. J. 1953b. The isolation of desacetylneo-protoveratrine from
Veratrum viride Ait. J. Am. Chem.Soc. 75: 3595-3596. Klohs, M. W., Draper, M., Keller, F., Koster,

S., Malesh, W. and Petracek, F. J, 1954. Alkaloids of Veratrum eschschohzii Gray II. The
ester alkaloids. J.

Am. Chem. Soc. 76: 1152-1153.

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Calif.

1008 pp.

Jacobs,

W. A. and Craig, L. C. 19,M.

The

Kupchan, S. M. and Gruenfeld, N. 1959. The hypotensive principles of cryptenamine, a

Veratrum viride alkalord preparation. J. Am. Pharm. Assoc. Sci. Ed. 48: 727-730. Lavoie, G. f984. Contribution d la connaissance de la flore vasculaire et invascualaire de la
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This article has been cited by: 1. Liliaceae Juss 751-807. [CrossRef] 2. Leo J Schep, David M Schmierer, John S Fountain. 2006. Veratrum Poisoning. Toxicological Reviews 25:2, 73-78. [CrossRef] 3. David Kleijn, Thomas Steinger. 2002. Contrasting effects of grazing and hay cutting on the spatial and genetic population structure of Veratrum album, an unpalatable, long-lived, clonal plant species. Journal of Ecology 90:2, 360-370. [CrossRef] Can. J. Plant Sci. Downloaded from pubs.aic.ca by 103.241.206.34 on 12/02/13 For personal use only.