Corn Lily


©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
December 2004
Updated August 2005, Nov. 2007; April 2009, Jan 2010, Feb 2010, May 2010, Oct 2010, Jun 2011, June 2016 (references)



     Veratrum is a genus of rhizomatous-bulbous herbs closely allied to the lily family (Liliaceae).  Species of Veratrum are commonly known as false hellebore, skunk cabbage, corn cabbage, cow cabbage, and corn lily.  The most appropriate is corn lily, because plants often grow in dense patches much like corn fields, and the terminal inflorescences superficially resemble corn tassels.  Moreover, skunk cabbage is a common name for plants of the Arum family, Lysichiton americanus Hultén & H. St. John, a western North American species, or Symplocarpus foetidus (L.) Salisb. ex Nutt., an eastern North American species, both unrelated to corn lily.  Hellebore—from the genus Helleborus, classified in the unrelated butter-cup family (Ranunculaceae)—is native to the Euro-Mediterranean where it may not be as familiar to persons living elsewhere.  The common name, false hellebore, which is based on superficial resemblance of some hellebore species to corn lily—in the greenish to whitish flowers—would obviously be less familiar. 

     The genus Veratrum is conservatively estimated to have ~27 species distributed in the temperate to boreal regions of the northern hemisphere (Liao et al. 2007).   The center of diversity lies in eastern China where there are 13 species, eight of which are endemic.  A related genus Melanthium with four species in eastern North America was recently included in Veratrum  (Zomlefer 1997; Zomlefer et al. 2000, 2003; Liao et al. 2007).


      Many of the species may be further divided.  For example, within the Section Veratrum—recognized by the green V-shaped nectar glands (Zomlefer et al. 2003)—are corn lily plants from south-central Wyoming to Colorado-New Mexico state line that have been distinguished as V. tenuipetalum A. Heller by the slightly smaller (narrow elliptic) tepals (   Other  differences are evident in the green V-shaped nectar glands near the base of the tepals  appearing wider in V.  californicum (e.g., above from the Klamath Mountains of California) than in V. tenuipetalum (e.g., from Colorado-New Mexico State line); however, plants in the Sierra Nevada of California also can have narrower V-shaped glands.  The type for V. californicum is from Nevada County, CA.  The taxonomic significance of floral characters in size, shape and position of nectar glands on the tepals is difficult to evaluate due to their small size and subtle variation in size, especially as tepals develop and wither. 

     Veratrum tenuipetalum may also include plants from New Mexico to Arizona and northern Mexico that collectively can be distinguished by the shorter flexuous inflorescence branches near the base of the primary flowering stem, often branching to near apex (Spjut 2010).  In contrast, V. californicum usually differs by the longer and more stiff inflorescence branches near the base of the main flowering stem that ends in a relatively short unbranched tail. This narrower interpretation of V. californicum is found in Utah and southwestern Wyoming to western Montana, across parts of Idaho and Nevada to Washington, Oregon, and California (Spjut 2010).  Additionally, rhizomes of Veratrum tenuipetalum are deeper in the soil and more entangled with roots of other plants; thus, more difficult to remove in which the clonal plants do not grow as close together as in var. californicum.  Nevertheless, these differences all appear relatively minor that would seem to justify varietal status rather than species status.

     Veratrum californicum var. caudatum (A. Heller) C. L. Hitchcock (long tailed corn lily) differs by the main flowering stem ending in a longer (unbranched terminally 1/3–1/2) tail (McNeal & Shaw 2002), and by other features such as longer flower bracts, paler green nectar glands on tepals, and rhizomes developing not as deep in soil.  It is found in more swampy environments in which three broad ecological complex types may be recognized: (1) a northern Rocky Mountain ecotype associated with grand fir and/or western red cedar that occurs at higher elevations than var. californicum in northern Idaho as shown above, but also found at various elevations in the Cascade Ranges of Washington and Oregon—up to 6,600 ft, (2) a  Coast Range element occurring in forest prairies and oak woodlands from the Washington-Puget Trough to the Willamette Valley in Oregon, and also in the Coast Ranges from Washington to northern California, and (3) a coastal maritime ecotype found in tidal marshes—where it may even become “partially submersed” at high tide as reported by one collector—at scattered locations from Washington to northern California.   Unlike var. californicum, var. caudatum may stand-up to heat and withstand frost more than var. californicum.  For example, if seasonal precipitation is below normal, var. californicum may fall to the ground before fruit matures (Spjut documented field observations, 2007); its distribution seems limited to the montane environments where temperatures are relatively cool so it can reproduce by seed (Spjut 2010). These differences support the species status of var. caudatum; however, the geographical distribution of var. californicum overlaps more with var. caudatum than with V. tenuipetalum  (Consortium of Pacific Northwest Herbaria 2010; McNeal & Shaw 2002), and intermediates are common in Idaho. 

     Not all white corn lily plants flower each year, often not much more than 5% (Taylor 1956b).  In drier years fewer plants flower (Spjut-documented comm. 2007, 2008, 2009, noting frequency of flowering in southern Idaho, Utah, Oregon and Colorado).  However, flowering in corn lily seemed above normal in 2009 in Idaho, Oregon and California; 73 of 75 sites studied had at least one individual in flower and >80% of the individuals at more than half of the sites were observed to have flowered (as determined from a review of photographs), possibly because of preceding years of drought.  Sites with less than 20% flowering  were evident for 4 of 7 sites in the Sawtooth National Forest in Idaho and all of five sites studied in Nevada—where desert vegetation in 2009 had received above average precipitation and had remained green until early July.  Taylor (1956b) reported that "frequency of blooming varies greatly by species and by habitat,” noting that “areas in the Sierras densely populated with literally millions of plants of V. californicum were cruised without seeing a single bloom,” and that 10 years may pass without flowering as he judged from the “frequency of crown doubling” for V. viride var. eschscholtzii (A. Gray) Breitung.  In a survey conducted by Spjut in 2010, sites observed in 2009 with high flowering frequency showed very little or no flowering.  Yet nearby sites in some places, such as along the western shore of Lake Cascade showed >90% of the plants flowering, where last year apparently they did not flower.

     Veratrum californicum also appears to intergrade with V. viride Aiton, a species distributed in the boreal region of North America (Taylor 1956b) as shown in photos below.  Veratrum  viride may be subdivided into two varieties (or subspecies or species), one found in the Pacific NW from Alaska to northern California and northern Idaho, Montana and Wyoming, and the other in the eastern North America from Quebec to northern Georgia (McNeal & Shaw 2002).  It is usually identified by the smaller narrower tepals, green to yellowish green in color, which is also in reference to its epithet (viride), in contrast to white flowers of V. californicum, and by the drooping flower branches and leaves (western North American plants).  Both V. californicum and V. viride can occur near each other such as along the western shore of Cascade Lake in Idaho, where V. viride is seen in wetter more shaded environments such as in forest understory or openings, particularly under conifers near forest margins (Spjut 2010, unpubl.).  Generally, where the species are sympatric, V. viride occurs at higher elevations than V. californicum (Spjut 2010, unpubl.).  Populations of corn lily in the white-fir and red fir forest meadows of the Klamath Region of California and southern Oregon show a range of intermediates in flower color and inflorescence flaccidness.  A mixed species population is suggested for the occurrence in Carter Meadow along the Cecilville-Callahan Road near Russian Peak, and in western Nevada near Reno.  Also, plants of V. californicum found in the ponderosa pine woodlands ranging from western Idaho to just east of the Cascades in Oregon often have flexuous to recurved inflorescence branches similar to V. viride.  This variation may be the result of historical introgression.

     Veratrum californicum (white corn lily) var. californicum is often found in open permanently to seasonally wet herb-grass-rush (and/or sedge) meadows in montane forest regions, especially in open valley headlands where snow may linger and where snow runoff leads to streams with riparian communities of aspen (Populus tremuloides), alder (Alnus spp.) and willow (Salix spp.).  The meadows are usually surrounded by various conifer forest types that may include ponderosa pine, subalpine fir, white fir, red fir, Englemann spruce, Douglas fir, lodgepole pine, incense cedar, Jeffrey pine, and/or sugar pine.  Aspen is commonly present near the conifer forest margins, and corn lily is often in the aspen understory extending to the meadow borders.  Occasionally, corn lily occurs within the understory of lodgepole pine, white fir, or snowbush  (Ceanothus velutinus) communities that appear to be the result of a gradual succession of vegetation from meadow to forest communities (see also Murray 2003).  In drier grassland and sagebrush communities, or occasionally antelope bush communities, corn lily often forms dense patches in open areas of topographical depressions where springs come to the surface, and along the edges of riparian aspen-sagebrush or western birch-sagebrush vegetation in valleys.  Corn lily is also a frequent understory component of ponderosa pine forests where it can be consistently present as single scattered individuals among herbs and other bushes as commonly seen in the Boise National Forest of Idaho, or corn lily may appear in more dense stands in ponderosa pine savannas in areas from western Idaho west to the Cascade Range of Oregon and south to Mount Lassen, California.  The elevation varies, generally not found below 8,500 ft in Utah, to most often occurring from 4,000-6,500 ft on the wetter slopes in California and Oregon, at higher elevations along the eastern Sierra Nevada Range to the southern California mountains (Consortium of California Herbaria 2010) and Nevada, and below 5,000 ft in Idaho.  Details on associated herbs and grasses are given in Sawyer et al. (2008) for California in which they recognize the “Veratrun californicum Herbaceous Alliance” that includes several associations such as with Bistorta bistortoides, with Juncus nevadensis, or with Senecio triangularis, while other associations are evident from specimen data in the Consortium of California Herbaria.

     The natural occurrence of white corn lily (Veratrum californicum) appears to be small clumps of clonal plants consisting of 3 to 4 shoots; the clumps scattered at various  intervals, often 100 m or more, along margins of wet  meadows or riparian vegetation.  Where meadows have a long history of grazing by domesticated animals, corn lily appears invasive to the extent that the open meadows or stream banks are dominated by corn lily (Potter 1998; Cosgriff et al. 2004; Murray 2007; Sawyer et al. 2008; Spjut 2010;  USDA Forest Service FEIS Environmental Impact Statement, Jarbidge District (NV), partial report online).   A clear example of corn lily's ability to increase in density and range as a result of grazing is shown in the following photos of a private land meadow within the Fremont National Forest of Oregon. 



     Comparison of satellite (Google Earth, 29 June 2006) with ground photos taken by Richard Spjut (Aug 2008) of a private parcel area within the Fremont National Forest in southeastern Oregon containing white corn lily (Veratrum californicum var. californicum).  The upper left photo was taken looking southwest along the fence, and the upper right photo shows a more direct west view. From Google Earth, corn lily appears as the paler green patches along the fence as well as inside the fence.  As cattle evidently wander along the perimeter of the fence in search of a path to the greener pasture inside, much of the vegetation—except corn lily—gets destroyed.  Where sagebrush and corn lily come into contact, the disturbed ecotone seems to favor corn lily more than sagebrush; consequently, corn lily advances into the sagebrush community—until perhaps moisture becomes a limiting factor.  Corn lily also appears to increase in density in the trampled area.  Normally, rhizomes of corn lily produce a new layer of tissue near the bulbous base each year as the older basal portions disintegrate (Taylor 1956b); however, from experience in collecting in areas where stands of corn lily are known to have been heavily grazed, it was not uncommon to find several rhizomes connected together; thus, grazing may induce branching and growth of rhizomes, which are underground stems; the activity of grazing perhaps analogous to one pruning bushes.

     The degree to which corn lily can spread over time has not been clearly documented.  In Lower Carter Meadow of the Klamath Mountains of California, photos taken of corn lily five years apart (2005, 2010) show very little difference in density and size of population.  A comparison of images taken in the mid 1960's for Morris Meadows along the Stuart Fork in the Trinity Alps Wilderness with internet images reportedly taken in 2009 appear to show some increase in V. californicum during the past 50 years (meadows grazed mainly by back packer's horses en route to lakes).  Thus, one may wonder just how old are some of the 5–10 acre high density stands of corn lily in the Warner Mountains, western Idaho, and northern Utah?  Additionally, the high density of V. californicum var. californicum occasionally seen as a major understory component of lodgepole forests and white fir forests in southeastern Oregon and in the Warner Mountains of California, among snow bush (Ceanothus velutinus) communities in Idaho and in quaking aspen communities in many western States appear to be the result of a long term natural succession of meadows to forest vegetation as already mentioned above; Murray (2003) showed examples of changes that have occurred in some meadows in the Klamath Mountains over a period of 50+ years.

     Ecologists in the Manti-la Sal National Forest (Utah) and in Ogden have been concerned about the long term trend in the spread of corn lily in the open subalpine meadows—where increase in the population size and density of corn lily is strongly suspected to be the result of a long history of sheep grazing—to the extent that the US Forest Service has tried to control its spread by chemical spraying and mechanical tilling, but with little success (Anderson & Thompson 1993; Cosgriff et al. 2004; Spjut-Thompson comm. Oct 2004). Fortunately, there is a chemical in the plant (cyclopamine) that is currently being utilized for the possibility of development of a new anticancer drug; see  Thus, by allowing harvests of corn lily plants for cancer research, the spread of corn lily might be controlled as well as help restore meadows back to their natural appearance.

     The World Botanical Associates (WBA) has been conducting geographical surveys of Veratrum californicum throughout most of the species range in the western United States since Sep 2004.  This terminated in 2012 as a result of a cyclopamine derivative (IPI-926) failing to show clinical benefit for treating pancreatic cancer (Spjut 2016); however, cyclopamine is continues to be investigated for treating colorectal cancer (Qualtrough et al. 2015).  Voucher specimens were prepared for more than 150 locations from where samples were collected for chemical screening of antitumor active compounds, cyclopamine and cycloposine (Spjut 2010, unpubl.).   Some vouchers have been sent to the Botanical Research Institute of Texas (BRIT) and to the University of Georgia (GA); most have been retained by the WBA.  As noted below samples were also collected of Veratrum viride in Alaska by the WBA for the National Cancer Institute in July 2003. 

     Additionally, in 2004, we obtained a permit that allowed us to gather ~250 kg of root-rhizome from within a national forest in northern California.  Upon visiting the same site in 2008, corn lily was observed to be as abundant as it appeared in 2004.  In collecting root-rhizomes, plants were left untouched (as required by the permit) so as not to completely eradicate it from any one spot.  These observations suggest that corn lily can quickly return.  In this case it was not cattle but direct human disturbance that may have helped the plant to spread.  Taylor (1956b) has suggested “that Veratrum plants are inherently capable of prolific annual increase by multiple crown formation” as evidenced from spraying plants with colchicine.

     The genus Veratrum has been reported to contain more than 100 alkaloids, among which is cyclopamine, known from V. californicum since the late 1960's  to cause birth defects in livestock such as Cyclops.  This poisoning occurs when plants are ingested by a pregnant animal at day 14 gestation.  The discovery was the result of observations on sheep developing Cyclops when grazing around areas where corn lily plants occurred in the Muldoon Canyon of Idaho (Panter et al. 2010). Scientists at the USDA Agricultural Research Service, Plant Poisonous Laboratory in Logan, Utah were asked to investigate the cause of the birth defects (Keeler 1968; James 1999; Heretsch et al. 2010; Panter et al. 2010).  This resulted in the isolation of the active (poisonous) compound, cyclopamine.  Later, oncologists at John Hopkins, who were looking for chemicals to target a specific embryonic developmental pathway (Sonic hedgehog signaling pathway) related to a genetic (protein deficiency) disorder, “Gorlin’s syndrome,” which can lead to medulloblastoma, had recalled studies on the inhibitory effects of cyclopamine. This further led to their finding of cyclopamine suppression of specific cultured brain tumor cells from mice, and from human medulloblastoma (Berman et al. 2002).  We at the WBA had also recognized the importance of this research on Veratrum when collecting samples of plants for the National Cancer Institute in Alaska during the summer of 2003 (DeVelice 2003).  Subsequently, the Infinity Pharmaceuticals Inc. has developed a derivative of cyclopamine as a potential new anticancer drug, currently scheduled for Phase II clinical studies for treating various cancers associated with the Sonic Hedgehog pathway (Mann 2010).

     General samples of Veratrum californicum (s.l.) had also been collected in the 1960's for the National Cancer Institute's antitumor screening program, and had showed preliminary activity in May 1964 from a root-bulb sample collected by Robert Perdue Jr. from Colorado (Perdue 4623, NA) and confirmed activity in Walker 256 in Aug 1968 from an aqueous extract of a stem-leaf sample collected by Arthur Barclay in Aug 1962 from Panther Meadow, Mt. Shasta, California (Barclay 1396, NA).  Recollections by Edward Terrell in Aug 1968 from Mt. Shasta (Terrell 4196, NA) and by Richard Spjut in July 1970 from Gumboot Lake, ~16 mi SW of Mt Shasta (Spjut 444, HSC, NA, accessioned by the USDA ARS as M-3418, PR-20369), failed to reconfirm as reported by Morris Kupchan, while a notation in the procurement record for P-388 activity is questionable and that the smaller recollection by Terrell 4196 was sufficient.  Morris Kupchan, who had isolated antitumor active compounds from many plants, had earlier done considerable work on the chemistry of Veratrum (e.g., Kupchan 1961; Kupchan et al. 1961).  Steroidal alkaloids had not been regarded to have potential in cancer chemotherapy based on similar antitumor active compounds isolated by Kupchan from  Solanum spp., while one might also suspect tannin activity since the original activity was from an aqueous extract.  A similar report was also made by Monroe Wall for the recollection obtained by Barclay from Colorado in July 1967, but this report did not get listed in the CPAM (1977) because preliminary activity did not confirm, while the author for this web page has not seen the original screening report.   Additionally, aqueous and alcohol extracts prepared from fresh samples of rhizomes and aerial parts collected by the University of Arizona from Apache County AZ  in Aug 1961 were inactive.   However, an aqueous-ethanol extract from a root [-rhizome] sample of V. nigrum L. collected in Mar 1974 from Taiwan by the National Defense Medical Center showed confirmed activity in KB (CPAM 1977). 

     It may be noted that rhizome samples of Veratrum can vary ecologically and geographically in alkaloid content (Spjut 2010, unpubl., 2016).  Cyclopamine was found in plants within a relatively narrow range of the species and partially correlated with vegetation type, probably due to mycorrhizal flora in which mycorrhizae associations have been reported for V. viride (Cazares et al. 2005). Thus, the original samples and recollections made for the NCI may not have contained cyclopamine, and subsequent samples that might have been collected by Perdue from Utah during the 1960's may not have been screened, because the NCI guidelines precluded screening active plants (Spjut 1985, 2010).   Cyclopamine has also been reported in the genus Fritillaria (Li et al. 2006) and has been produced in cultured cells of Veratrum calfiornicum (US Patent, Ritala-nurmi et al. 2009).

     As a final note,  a number of plant derived anticancer agents such as taxol, camptothecin and others, that are currently being used in cancer chemotherapy, come from poisonous plants.  For general info on relationships between antitumor plants, medicinal plants and poisonous plants, see Spjut, R. W.  2005.  Relationships Between Plant Folklore and Antitumor Activity: An Historical Review.  Sida 21(4): 2205–2241.


Veratrum californicum

East slopes of Cascade Range, WA
Swauk Meadow, SPJ-16722, July 2010

Veratrum californicum

Just east of Pullman, WA, SPJ-16723, July 2010.  Some plants appearing to have been sprayed by chemical as shown on right.

Veratrum californicum var. caudatum

Near Grangeville, ID, SPJ-16740, July 2010

Veratrum californicum var. californicum
Independence Mts, NV, SPJ-16750, July 2010.  Note old flowering stalks from last year still standing.

Veratrum californicum var. californicum
Warner Mts, CA, just south of the Dismal Swamp, SPJ-16756, July 2010.  Occurring extensively in the understory of lodgepole pine forest, probably the result of lodgepole pine invading the meadow over a long period of time, rather than corn lily invading the forest.

Veratrum californicum var. californicum
Warner Mts, CA, SPJ-16765, July 2010.  A large stand with only a few plants in flower.

Veratrum californicum var. californicum
Trinity Alps Wilderness, CA, July 1973. Growing abundantly along edge of Granite Lake, cattle observed grazing
in the area


Veratrum californicum
var. californicum

Morris Meadows
Stewart Fork Trail,
Trinity Alps Wilderness,
Trinity Co., CA, July 1969


Veratrum californicum
Russian Wilderness,
Taylor Lake,
~6,500 ft elev., CA, July 2005.


Veratrum x californicum
Carter Meadows, Siskiyou Co., Salmon Mts., 5,800 ft., Klamath National Forest, CA
Oct 2004. The dense patches among alder thickets is probably related to past grazing in the meadow (Cosgriff et al. 2004; Murray 2007). Appearing intermediate to V. viride.

Veratrum x californicum
Putative hybrid with V. viride in a mixed
species population shown below
Klamath National Forest,
Carter Meadow, elev. 5,800 ft,
Salmon Mts., CA
July 2005.

Veratrum x viride
Aug 2010, the V. x californicum did not flower this year.

Veratrum californicum
Lower Carter Meadows.
 4800 ft elev., Siskiyou Mts.,
Klamath National Forest, CA, July 2005.
One additional photo on right, Aug 2010. Population shows little change in growth during past 5 years.
None of the plants flowered in 2010.

Veratrum californicum
Mt Ashland, OR, Spjut 16441, Aug 2008, intermediate to V. viride

Veratrum californicum
Marble Mountains Wilderness, Deep Lake, ~6,000 ft elev., CA, July 2005.  A USDA Forest person who was checking campsites for fire prevention in the area  reported that cattle graze the meadows during summer months

Veratrum californicum
Marble Mountains Wilderness,
Meadow below Boulder Peak,
CA, ~7,000 ft elev., July 2005.


Veratrum californicum
var. californicum

Haypress Meadows
Marble Mts. Wilderness,
Siskiyou Co., CA, July 2004


Veratrum californicum var. californicum

Toiyabe Range, Kingston Creek, NV, SPJ-16477, July 2009

Veratrum californicum var. californicum

Toiyabe Range, Birch Creek, NV,
SPJ-16478, July 2009

Veratrum californicum var. californicum

Humboldt-Toiyabe National Forest, CA, subalpine meadow ~ 9,600 ft, June 2005

Veratrum californicum var. californicum
Manti-La Sal National Forest, Utah,  alpine meadow
~ 10,000 ft, June 2005

Manti-La Sal National Forest, Utah,
10,200 ft, SPJ-15961; August 2005

Veratrum californicum var. californicum

Wasatch Cache National Forest, UT, Albion Basin, Little Cottonwood Canyon,
elev. ~ 9,500 ft.  SPJ- 15962, August 2005

Veratrum californicum var. californicum

Boise National Forest, ID
Corn lilies growing singly in wooded understory of pine woodland, instead of in wet meadows, 4,000 ft.  SPJ-15963
August 2005


Veratrum californicum var. californicum

Payette National Forest, ID
Corn lilies growing singly in wooded understory of pine woodland, instead of in wet meadows, 4,000 ft.  SPJ-15964
August 2005


Veratrum californicum var. californicum

Boise National Forest, ID
Corn lilies growing singly in seral montane chaparral, primarily among snowbush, Ceanothus velutinus, 6,100 ft.  SPJ-15966, August 2005

Veratrum californicum var. californicum

Boise National Forest, ID
Corn lilies growing singly and in clumps, openings of steep slopes in spruce-fir forest, and singly among shrubs in riparian vegetation, nectary pale green, 6,100 ft.  SPJ-15967, August 2005

Veratrum californicum var. californicum
Sawtooth National Forest, Idaho,
wet ravine ~ 7,200 ft elev.
June 2005


Veratrum californicum var. californicum

West shore of Lake Cascade, ID,
SPJ-16745, July 2010.
High density stand with >90% flowering.  Population probably developed from vegetative  reproduction by rhizomes.


Veratrum californicum var. californicum
Nevada Co., Tahoe National Forest, California
Possibly type locality
  4,550 ft, August 2006


Veratrum californicum var. caudatum
Vicinity of Oregon State University campus, Nov. 2004.  Localized patch of ~200 plants in
swampy meadow.  Close-up of infructescence showing septicidal capsules, some still containing seeds.




Veratrum insolitum
Klamath National Forest, CA,
Elev. ~4,000 ft, July 2005

Veratrum insolitum
near Bald Hills, CA,
July 2005


Veratrum tenuipetalum
Columbine, CO, near WY state line, Aug 2008


Veratrum tenuipetalum
San Juan NF, CO, between Durango and Colorado Springs, SPJ-16392,, July 2008

Veratrum tenuipetalum
Navajo Peak, CO/NM state line, July 2008

Veratrum tenuipetalum
Medicine Bow, NF, WY, Aug 2008

Veratrum viride
Kenai Peninsula, AK
Spjut & Marin 15396, July 2003


Veratrum viride
West shore of Cascade Lake, ID
Spjut 16511, Aug 2009


Veratrum viride x V. californicum
and V. californicum
West shore of Cascade Lake, ID
Spjut 16510, Aug 2009

Veratrum viride
Mount Hood, OR
Spjut 16523, Aug 2009


Veratrum sp.
Northern CA,  Aug 2009
Note: narrow leaves, zigzag inflorescence branches

Veratrum sp.
Western NV,  Aug 2009
Tepals cuneate, not overlapping, nectar glands irregularly developed, inflorescence branches dark green as in V. viride


Veratrum californicum
Sierra Nevada, Greenhorn Mts., CA
Aug 2009, SPJ-16558


Selected References:

American Cancer Society. Chemotherapy and other drugs for pancreatic cancer.. Accessed June 2016.

Anderson, V. J. and R. M. Thompson. 1993. Chemical and mechanical control of false Hellebore (Veratrum californicum) in an alpine community. U.S. Dept. of Agriculture, Forest Service, Intermountain Research Station, Research paper INT; 469.. Ogden, UT (324 25th St., Ogden 84401) . 6 p.

Anonymous.  1967. Veratrum alkaloids in the therapy of myasthenia gravis.  Can. Med. Assoc. J. 96(23): 1534–1535.

Bastin, 1895.  E. S. Structure of Veratrum viride.  Amer. J. Pharm. 67 (4): 1–8. Botanical Medicine Monographs and Sundry.  The Southwest School of Botanical Medicine  This is about the morphological structure of the root-rhizome.

Berman, D. M., S. S., Karhadkar,  A. R. Hallahan,  J. l. Pritchard, C. G. Eberhart, D. N. Watkins, J. K. Chen,  M. K. Cooper, J. Taipale, J. M. Olson, and P.A. Beachy.  2002.  “Medulloblastoma growth inhibition by Hedgehog Pathway Blockade.  Science 297:  1559–1561. “Constitutive Hedgehog (Hh) pathway activity is associated with initiation of neoplasia, but its role in the continued growth of established tumors is unclear. Here, we investigate the therapeutic efficacy of the Hh pathway antagonist cyclopamine in preclinical models of medulloblastoma, the most common malignant brain tumor in children. Cyclopamine treatment of murine medulloblastoma cells blocked proliferation in vitro and induced changes in gene expression consistent with initiation of neuronal differentiation and loss of neuronal stem cell-like character. This compound also caused regression of murine tumor allografts in vivo and induced rapid death of cells from freshly resected human medulloblastomas, but not from other brain tumors, thus establishing a specific role for Hh pathway activity in medulloblastoma growth.”

Binns W., L. F. James, R. F. Keeler and L. D. Balls.  1968.  Effects of teratogenic agents in range plants.  Cancer Res. 28(11): 2323–3236.

Cazares, E., J. M. Trappe, & A. Jumpponen. 2005. Mycorrhiza-plant colonization patterns on a subalpine glacier forefront as a model system of primary succession. Mycorrhiza 15: 405–416. Lyman glacier in the North Cascades Mountains of Washington has a subalpine forefront characterized by a well-developed terminal moraine, inconspicuous successional moraines, fluting, and outwash. These deposits were depleted of symbiotic fungi when first exposed but colonized by them over time after exposure. Four major groups of plant species in this system are (1) mycorrhiza independent or facultative mycotrophic, (2) dependent on arbuscular mycorrhizae (AM) (3) dependent on ericoid mycorrhiza (ERM) or ectomycorrhizae (EM), and (4) colonized by dark -septate (DS) endophytes. We hypothesized that availability of mycorrhizal propagules was related to the success of mycorrhiza-dependent plants in colonizing new substrates in naturally evolved ecosystems. To test this hypothesis roots samples of 66 plant species were examined for mycorrhizal colonization. The plants were sampled from communities at increasing distances from the glacier terminus to compare the newest communities with successively older ones. Long established, secondary successional dry meadow communities adjacent to the glacier forefront, and nearby high alpine communities were sampled for comparison. DS were common on most plant species on the forefront. Nonmycorrhizal plants predominated in the earlier successional sites, whereas the proportion of mycorrhizal plants generally increased with age of community. AM were present, mostly at low levels, and nearly absent in two sites of the forefront. ERM were present in all species of Ericaceae sampled, and EM in all species of Pinaceae and Salicaceae. Roots of plants in the long established meadow and heath communities the forefront and the high alpine community all had one or another of the colonization types, with DS and AM predominating.

Cholakova M., V. Christov, N. Kostova, R. Todorova, E. Georgieva and E. Nikolova.  2005.  Biological activity of Veratrum alkaloids.  Exp. Pathol. Parasitol. 8(2): 16–19.  “The biological activity and the chemical classification of alkaloids isolated from Veratrum plants are discussed.”  The article mentions that cyclopamine has been isolated from several species.

Consortium of California Herbaria CAS-DS · CDA · CHSC · DAV · HSC · IRVC · OBI · PGM · RSA-POM · SBBG · SD · SDSU · SJSU · UC-JEPS · UCR · UCSB · UCSC. 270 records for Veratrum californicum, 12 records for V. viride. Feb 2010.

Consortium of Pacific Northwest Herbaria. Managed by the University of Washington Herbarium, Burke Museum of Natural History and Culture, Box 355325 University of Washington, Seattle, WA 98195.  129 records for Veratrum californicum, 215 records for V. viride. Feb 2010.

Contois, M., J. Cahill, N. Chavez, C. Cacace, G.I.  Wechsler, M. Pauli and E. Kosman.  2004.  Estimating Corn Lily Abundance in Bear Trap Meadow, July 2004.

Cosgriff, R., V. J. Anderson, and S. Monsen. 2004.  Restoration of communities dominated by false hellebore.  J. Range Mgmt. 57:365–370.  “False hellebore (Veratrum californicum Durand) is a native component of high-elevation, meadow-riparian areas of the mountain West that has increased due to historic heavy grazing. In 1991, a study was established in dense stands of false hellebore to evaluate mechanical and chemical control methods to reduce false hellebore and increase the abundance of the other native herbaceous species in these tall-forb communities. Four control methods consisting of the herbicide glyphosate (N-phenophonomethylglycine), mow, mowing in 2 consecutive years (remow), and tillage were used in 1991-1992. Each method was evaluated based on (1) reduction of false hellebore stem densities; (2) response of residual understory species; and (3) effectiveness of seeding a perennial grass and forb mixture to sustain initial treatment control. Stem density of false hellebore and nested frequency data for all species were collected in 1991, 1992, 1995, and 1999. The glyphosate treatment was effective in reducing false hellebore stem density which allowed for recovery of the remnant tall-forb community. The till treatment, while effectively reducing false hellebore stem density, also eliminated the other species in the community, leaving it open to invasive weeds. The mow and remow treatments did not reduce false hellebore stem density, but did allow for recovery of other components of the tall-forb community. Seeding following control treatments had no effect on false hellebore stem densities due to poor establishment. The mechanical treatments were generally more cumbersome in application and limited to gentle topography and well-drained sites without surface rocks. The application of herbicides is much easier and is adaptable to all types of terrain. The use of the herbicide glyphosate gave the best balance of false hellebore control and recovery of the tall-forb community.”

CPAM (Confirmed Plant and Animal Materials).  1977. T–Z. (p. 582).

Cseri J., M. Dankó, L. Kovács, G. Szücs and E. Varga. 1980. Analysis of the sensitizing effect of veratrum alkaloids to potassium on frog muscle.  Acta Physiol. Acad. Sci. Hung. 56(3): 289–301. “The sensitizing effect of veratrum alkaloids to potassium is not specific. Reducing the concentration of chloride in Ringer's solution, or treating the muscle with nicotine in a concentration close to threshold after pretreatment with subliminal concentration of cevadine result in a marked mechanical response of the muscle. However, cevadine does not alter the sensitivity of the muscle to caffeine. On the basis of these observations it has been suggested that veratrum alkaloids sensitize the muscle membrane essentially to depolarizing processes. 2. Cevadine, 0.01 mM, fails to depolarize the muscle membrane but increases the depolarizing effect of 10 mM potassium. The depolarizing effect of a reduction of the concentration of chloride from 120 mM to 30 mM is also increased in cevadine pretreated muscle. Cevadine pretreatment increases the depolarizing effect of nicotine, too. 3. The above sensitizing effects are unanimously Na-dependent. Accordingly there is no mechanical response and increased depolarization in muscles equilibrated in sodium-free (choline) Ringer's solution before the cevadine treatment. 4. On the basis of the present data it is suggested that the membrane, when sensitized by veratrum alkaloids, can be triggered by different depolarizing processes and the depolarization increases as the result of increased Na permeability. The increased depolarization at the threshold level becomes sufficient for the automatic regenerative processes of the action potential to develop which activate the contractile elements. However, the mechanical response is a prolonged contraction rather than a contracture, its long period being the result of a very slow repolarization caused by the well-known inhibitory effect of veratrum alkaloids on Na inactivation.”

DeVelice, R. L. 2003. In search of a cure. Sour Dough Notes: 23, Editor's Note.

Fluck H. and H. R. Hegi.  1956. [Studies on the alkaloids of the surface organs of Veratrum album L. I. Qualitative studies on the alkaloids of the leaves.] Pharm. Acta Helv. 31(9): 428–447. German.

Fluck H. and H. R. Hegi.  1960. [Studies on the alkaloids in the aerial organs of Veratrum album L. Part 3. Fluctuations in the alkaloid content of the aerial organs, especially the low-lying leaves, of Veratrum album L.] Pharm. Acta Helv. 35: 1–12.

Fuska J., A. Fusková, A. Vassová and Z. Votický.  1981. New substances with cytotoxic and antitumor effects. IV. In vitro effect of some veratrum alkaloids and their derivatives on leukemia P388 cells. Neoplasma 28(6): 709–714. “Some Veratrum alkaloids and their derivatives exhibited an in vitro cytotoxic effect on leukemia P388 cells, depending on the structure of the skeleton of the molecule, particularly on the type of the heterocycle attached to C-20. Veracintine and 20-(2-methyl-1-pyrrolin-5-yl)-4-pregnen-3-one, which proved to be the most effective, inhibited incorporation of uridine, and to a lesser extent that of L-valine into P388 cells fractions. After a brief reaction (15 min), these substances became irreversibly bound in the P388 cells and stopped their further in vitro proliferation. The cytotoxic effect of veracintine became enhanced by sublethal doses of tubercidine (phase of maximum lethality of G1).

Heretsch, P., L. Tzagkaroulaki, and Athanassios Giannis. 2010. Cyclopamine and Hedgehog Signaling: Chemistry, Biology, Medical Perspectives. Angew. Chem. Int. Ed. 49: 2–12 (online preprint). "When Odysseus left the devastated city of Troy after ten years of siege he could not foresee the perils he still had to face. The encounter with the cyclops, a giant with only one eye placed in the middle of its forehead, was doubtlessly one of the creepiest and most dangerous of his adventures. In the end, Odysseus could only escape with the help of a sheep. Whether Homers cyclops was inspired by the observation of terribly malformed neonates remains speculative. However, when sheep herders in Idaho in the middle of the 20th century faced an increasing number of cyclops-like sheep in their herds, a unique  cascade of chemical, biological, and medicinal discoveries was initiated. This Minireview tells this story and shows its impact on modern biomedical research."

Incardona, J. P. W. Gaffield,  R. P.  Kapur and H. Roelink. 1998. The teratogenic Veratrum alkaloid cyclopamine inhibits sonic hedgehog signal transduction. Development 125: 3553–3562.  “The steroidal alkaloid cyclopamine produces cyclopia and holoprosencephaly when administered to gastrulation-stage amniote embryos. Cyclopamine-induced malformations in chick embryos are associated with interruption of Sonic hedgehog (Shh)-mediated dorsoventral patterning of the neural tube and somites. Cell types normally induced in the ventral neural tube by Shh are either absent or appear aberrantly at the ventral midline after cyclopamine treatment, while dorsal cell types normally repressed by Shh appear ventrally. Somites in cyclopamine-treated embryos show Pax7 expression throughout, indicating failure of sclerotome induction. Cyclopamine at concentrations of 20-100 nM blocks the response of neural plate explants to recombinant Shh-N in a dose-dependent manner. Similar concentrations have no effect on the post-translational modification of Shh by cholesterol in transfected COS-1 cells. Comparison of the effects of cyclopamine to those of the holoprosencephaly-inducing cholesterol synthesis inhibitor AY-9944 shows that cyclopamine does not induce malformations by interfering with cholesterol metabolism. Although AY-9944 does not interrupt Shh signaling in ovo, it blocks the response to Shh-N in explants cultured without an exogenous cholesterol source. As predicted by current models of the regulation of cholesterol metabolism, the response to Shh-N in AY-9944-treated explants is restored by providing exogenous cholesterol. However, exogenous cholesterol does not restore Shh signaling in cyclopamine-treated explants. These findings suggest that cyclopamine-induced teratogenesis is due to a more direct antagonism of Shh signal transduction. ”

Infinity Pharmaceuticals,  Inc (IPI). 2009. Infinity Announces Hedgehog Pathway Preclinical Data in Ovarian Cancer Data Demonstrate Significant Inhibition of Tumor Growth in Primary Ovarian Cancer.  Globe NewsWire.

James, L. F. 1999. Teratological research of the USDA-ARS Poisonous Plant Research Laboratory.  J. Nat. Toxins 8: 63–80. “Research on teratogenic plants started at the USDA-Agricultural research Service Poisonous Plant Research Laboratory in the mid 1950's when Dr. Wayne Binns, Director of the laboratory, was asked to investigate the cause of a cyclopain facial/skeletal birth defect in lambs.  Dr. Lynn F. James joined the staff shortly after.  These two people worked as a team wherein most planning was done jointly with Binns supervising most of the laboratory work and James the field studies.  It was determined that when pregnant ewes grazed Veratrum californicum on day 14 of gestation a significant number of lambs had the cyclopic defect.  Skeletal and cleft palate birth defects in calves was associated with pregnant cows grazing certain lupine species during 40–70 days of gestation.  Shortly thereafter research work was initiated on locoweed which caused abortions, wasting, right heart failure, skeletal birth defects, and fetal right heart failure.  Dr. Richard F. Keeler, a chemist who joined the staff in the early 1960's isolated and characterized the teratogens in V. californicum as the steroidal alkaloids cyclopamine, jervine, and cycloposine. He also described the terotogen in lupines as the quinolizidine alkaloid angyrine and the piperidine alkaloid ammodendrine.  Drs. Russell Molyneux and James identified the toxin in locoweed as the indolizidine alkaloid swainsonine.” “In 1974 the editor of Nutrition Today (Vols. 9 and 4) wrote 'The idea that birth defects occurring in humans may be in some way related to diet is not widely held...' Dr. Lynn James pointed out in this issue that such defects in animals can be produced with absolute predictability and regularity by foods ordinarily beneficial to livestock.  Management strategies have been developed to prevent or minimize the economic impact of the cyclopian lamb and the crooked calf condition on livestock producers and well on the way to doing the same with locoweed.  It is of interest to note that livestock researching on Veratrum, lupines and locoweed and toxins therefrom are now significant research tools for specific human health problems.

Keeler, R. F. 1968. Teratogenic compounds of Veratrum californicum (Durand). IV. First isolation of veratramine and alkaloid Q and a reliable method for isolation of cyclopamine. Phytochemistry 7: 303–306.

Keeler, R. F. 1969.  Toxic and teratogenic alkaloids of western range plants. J. Agric. Food Chem. 17: 473–482.

Keeler, R. F., and W. Binns. 1971. Teratogenic compounds of Veratrum californicum as a function of plant part, stage, and site of growth. Phytochemistry, 10: 1765–1769. “The level of the teratogen cyclopamine in Veratrum californicum varied considerably among plants from various collection sites. The variation was not correlated with differences in soil type, pH, soil nutrients, elevation, drainage, or sunlight. However, marked variation in both total alkaloid and percentage cyclopamine occurred as a function of stage of growth of the plant. The levels of both were highest in early growing season in the leaves, in midgrowing season in the stems and in late growing season in the root/rhizome system.”

Keeler, R. F. 1978. Cyclopamine and related steroidal alkaloid teratogens: their occurrence, structural relationship, and biologic effects.  Lipids. 13(10): 708–715. “A spontaneous congenital deformity is produced in lambs whose dams consume Veratrum californicum on the 14th day of gestation. The deformity is generally expressed as cyclopia, cebocephaly, anophthalmia, or microphthalmia. This teratogenic effect is produced by certain steroidal alkaloid teratogens from the plant - most notably the compound cyclopamine. Cyclopamine is a C-nor-D-homo steroid with fused furanopiperidine rings E and F at right angles to the plane of the steroid because of spiro attachment at C-17 of the steroid. Among veratrum alkaloids, only those with an intact furan ring E were teratogenic in sheep, whereas those in which the peperidine ring is not rigidly positioned at right angles to the steroid were not. Many ruminants and laboratory animals are susceptible to the teratogen. It has wide species and tissue specificity and appears to have a direct effect on the embryo, not as a consequence of metabolic alteration of its structure nor as an indirect effect through a maternal influence. Other plant sources, notably potatoes, tomatoes, and eggplant contain related spirosolane steroidal alkaloids. Among naturally occurring spirosolanes, solasodine is teratogenic in hamsters, but neither tomatidine not diosgenin, the non-nitrogen containing analog of solasodine, is teratogenic. Results of these and other studies suggest that a basic nitrogen positioned alpha with respect to the steroidal plane and at appropriate distance beyond the D ring confers the teratogenicity on the molecule. Potato sprouts with high alkaloid content are teratogenic in hamsters, but tubers and peels are not.

Keeler, R. F. and W. Binns   2005. Teratogenic compounds of Veratrum californicum (Durand). V. Comparison of cyclopian effects of steroidal alkaloids from the plant and structurally related compounds from other sources Teratology 1: 5–10. Cyclopamine, and its glycoside alkaloid X, along with jervine and veratrosine, induced cyclopian malformations in offspring born to ewes ingesting these compounds on the fourteenth day of gestation. Other steroidal alkaloids with somewhat similar structures and various other steroidal compounds including certain hormones and steroidal sapogenins did not induce the malformation.”

 Klohs,M. W.,  M. D. Draper, F. Keller, S. Koster, W. Malesh and F. J. Petracek.  1953. The Alkaloids of Veratrum fimbriatum Gray. J. Am. Chem. Soc.; 75(20):  4925 - 4927. "A chemical investigation of Veratrum fimbriatum Gray has yielded two new hypotensively active germine esters, germanitrine (C39H6901lN), and germinitrine (C39H57011N), as well as neogermitrine, jervine and pseudojervine. In addition, the alkaloidal ester, veratroylzygadenine, previously isolated from Zygadenus venenosus Wats was found to be present in this species of Veratrum. On hydrolysis, the triester germanitrine affords germine, acetic acid, tiglic acid and (I)-a-methylbutyric acid. The hydrolysis of the triester germinitrine, yielded germine, acetic acid, tiglic acid and angelic acid. On methanolysis, germanitrine readily loses a labile acetyl group yielding germanidine."

Krayer O., S. M. Kupchan, C. V. Deliwla and B. H. Rogers. 1953. Studies on Veratrum alkaloids. XVIII. Chemical and pharmacological relations between Zygadenus and Veratrum alkaloids. Naunyn Schmiedebergs Arch. Exp. Pathol. Pharmakol. 219(5): 371–385.

Krupp H., L. Lendle and K. Stapenhorst.  1952.  [The use of nicotin and veratrum alkaloids as insecticides.]. Arzneimittelforschung 2(6): 258-62.

Kupchan S. M., J. C. Grivas, C. I. Ayres, L. J. Pandya and L. C. Weaver. 1961.  Veratrum alkaloids. XLVI. Structure-activity relationships in a series of analogs of the protoveratrines.  J. Pharm. Sci. 50: 396–403.

Kupchan, S. M.  1961. Hypotensive Veratrum ester alkaloids. J. Pharm. Sci. 50: 273–287.

Kupchan, S. M. and R. H. Nehnsler.  1961. Veratrum alkaloids.  XLIV. Structure-activity relationships in a series of synthetic hypotensive esters of protoverine.  J. Med. Pharm. Chem. ;3: 129–155.

Kutney J. P., J. Cable, W. A. Gladstone, H. W. Hanssen, E. J. Torupka and W. D. Warnock.  1968. The total synthesis of Veratrum alkaloids. I. Verarine. J. Am. Chem. Soc. 90 (19): 5332–5334.

Li, H-l., J. Tang, R-h. Liu, M. Lin, B. Wang, Y-f. Lv, H-q. Huang, C. Zhang, and W-d. Zhang. 2007. Characterization and identification of steroidal alkaloids in the Chinese herb Veratrum nigrum L. by high-performance liquid chromatography/electrospray ionization with multi-stage mass spectrometry.  Rapid Communications in Mass Spectrometry 21(6): 869–879. Electrospray ionization multi-stage mass spectrometry (ESI-MSn) was performed to study the fragmentation behaviour of seventeen steroidal alkaloids (4 protoverine-type alkaloids, 10 germine-type alkaloids and 3 zygadenin-type alkaloids) from the Chinese herb Veratrum nigrum L. The MSn spectra of the [M+H]+ ions for steroidal alkaloids provided a wealth of structural information on the substituted groups. In positive ion mode, the three types of alkaloids showed very different characteristic ions: m/z 436 or 418 for protoverine-type alkaloids; m/z 438, 420 or 402 for germine-type alkaloids; m/z 440 or 422 for zygadenin-type alkaloids. These fragments were used to deduce their mass spectral fragmentation mechanisms. Furthermore, the primary compounds in methanolic extracts of the herb of Veratrum nigrum L. were investigated by using liquid chromatography (LC)/ESI-MSn. As a result, 21 steroidal alkaloids (5 protoverine-type alkaloids, 14 germine-type alkaloids and 2 zygadenin-type alkaloids) were selectively identified from 27 determined peaks. Eleven compounds were unambiguously identified by comparing with standard compounds and ten compounds were tentatively identified or deduced according to their MSn data. Two of these compounds (xingangermine and deacetyl xinganveratrine) were found to be novel steroidal alkaloids. In addition, the chemical structures of two pairs of steroidal alkaloid isomers were deduced by comparing their fragment ions. Given the important structural information of known and unknown steroidal alkaloids in crude herbal extracts, this study is useful for identifying these types of steroidal alkaloids in crude materials rapidly and selectively.”

Li H-J., Y. Jiang and P. Li. 2006. Chemistry, bioactivity and geographical diversity of steroidal alkaloids from the Liliaceae family.  Nat. Prod. Rep. 23: 735–752.

Liang G. Y. and N. J. Sun. 1984. [Chemical studies on active principles of Veratrum stenophyllum. III. Studies on the structure of beta 1-chaconine and the partial structures of stenophylline C and stenophylline D]. Yao Xue Xue Bao. 19(6): 431-436.

Liang G. Y. and N. J. Sun. 1984. [Chemical studies on active principles of Veratrum stenophyllum. II. Studies on the structure of a new blood pressure lowering agents--stenophylline A]. Yao Xue Xue Bao. 19(3): 190-194.

Liang G. Y.  1984. [Studies on the alkaloids of Veratrum genus]. Yao Xue Xue Bao. 19(4): 309-20.

Liao W.-J., Y.-M. Yuan and D.-Y. Zhang.  2007. Biogeography and evolution of flower color in Veratrum (Melanthiaceae) through inference of a phylogeny based on multiple DNA markers  Plant Systematics and Evolution 267:177–190. “Veratrum (Melanthiaceae) comprises ca. 27 species with highly variable morphology. This study aims to construct the molecular phylogeny of this genus to infer its floral evolution and historical biogeography, which have not been examined in detail before. Maximum parsimony, maximum likelihood, and Bayesian analyses were performed on the separate and combined ITS, trnL-F, and atpB-rbcL sequences to reconstruct the phylogenetic tree of the genus. All Veratrum taxa formed a monophyletic group, within which two distinct clades were distinguished: species with white-to-green perianth formed one highly supported clade, and the species with black-purple perianth constituted another highly supported clade. Phylogenetic inference on flower color evolution suggested that white-to-green perianth was a plesiomorphic state and black-purple perianth was apomorphic for Veratrum. When species distribution areas were traced as a multi-state character, parsimonious optimization inferred that Veratrum possibly originated in East Asia. Our study confirmed previous phylogenetic and taxonomic suggestions on this genus and provided a typical example of plant radiation across the Northern Hemisphere.”

Ma R., A. Ritala, K. M. Oksman-Caldentey and H. Rischer. 2006. Development of in vitro techniques for the important medicinal plant Veratrum californicum. Planta Med. 72(12): 1142–1148. “Veratrum californicum (Liliaceae) is an important monocotyledonous medicinal plant which is the only source of the anticancer compound cyclopamine. An in vitro culture system for somatic embryogenesis and green plant regeneration of Veratrum californicum was developed. Embryogenic calli were induced from mature embryos on induction medium. Five basal media supplemented with different growth regulators were evaluated for embryogenic callus induction, modified MS medium with 4 mg/L picloram showing the best result for embryogenic callus production. Fine suspension cell lines were established by employing friable embryogenic calli as starting material and AA medium and L2 medium as culture media. The suspension cell lines cultured in AA medium with 4 mg/L NAA appeared to be fresh yellow and fast growing. The suspension cells were cryopreserved successfully and recovered at a high rate. Green plants were regenerated from embryogenic calli maintained on solid medium with 73 % regeneration ability (green plants/100 calli) in 27-month-old culture. The in vitro plantlets contained the steroid alkaloids cyclopamine and veratramine. This in vitro system will form the basis for metabolic engineering of Veratrum cells in the context of biotechnological production of pharmaceutically important secondary metabolites. DMSO:dimethyl sulfoxide fw:fresh weight NAA:naphthaleneacetic acid 2,4-D:2,4-dichlorophenoxyacetic acid picloram:4-amino-3,5,6-trichloro-2-pyridinecarboxylic acid dicamba:3,6-dichloro-2-methoxybenzoic acid.

Maj J. [Investigations on Veratrum lobelianum Bernh.; Veratrum extracts.] 1955. Acta Pol. Pharm. 11(Suppl): 108-110. Polish.

Mann, D. 2010. IPI-926: Potential of natural products for drug development: Investigations into Veratrum californicum (cow cabbage) as a source. Society for Range Management: Symposium—Medicinal Uses of Veratrum. Annual Meeting with Weed Science Society of America, Denver, CO, Feb 7–11 (30 minute oral presentation, no abstract).

McNeal, Jr., D. W. and A. D. Shaw. 2002. Veratrum. In Flora North America 26, Liliaceae: 72–76.

Meilman E. Clinical studies on veratrum alkaloids. III. 1953. The effect of protoveratrine on renal function in man. J. Clin. Invest. 32(1): 80–89.

Meilman E. and O. Krayer.  1950. Clinical studies on veratrum alkaloids; the action of protoveratrine and veratridine in hypertension. Circulation. Feb;1(2):204-13.

Monsen, S. 2010. Wildland harvesting - site identification, lifting processes, and restoration measures. Society for Range Management: Symposium—Medicinal Uses of Veratrum. Annual Meeting with Weed Science Society of America, Denver, CO, Feb 7–11 (30 minute oral presentation, no abstract).

Murray, M. P.  1997. High elevations meadows and grazing.  Internatl. J. Wilderness 3(4):24–27.Abstract: “High elevation grazing of cattle and sheep is a legal activity in wilderness areas administered by the U.S. Forest Service (USFS) and Bureau of Land Management (BLM) and occurs in about one-third of the U.S. Wilderness System (USWS). General effects of grazing on species composition and soil properties are described based on reported findings for three extensive types of high elevation meadows-grass, herbaceous, and moist sedge. The challenge for wilderness managers is to keep grazing within limits that protect the naturalness of meadow ecosystems. In general, where excessive grazing occurs, shifts in species composition from preferred livestock forage to less desirable, nonpalatable, and exotic species is observed. Soils of each meadow type respond differently to grazing pressure, and careful management should reflect these differences. Suggestions are offered for careful control of livestock distribution, timing, and stock numbers in order to protect naturalness of high elevation wilderness meadows.”

Murray, M. P.  2003. Tree encroachment on Klamath Mountains meadows. Fremontia 31:  13–18.

National Institutes of Health, National Cancer Institute. 1966. Screening data from the Cancer Chemotherapy National Service Center Screening Laboratories.  Plants collected and extracted by the University of Arizona. P. 116 in regard to test results for fresh samples of root and flower-fruit from Veratrum californicum collected in Apache County, AZ, Aug 1961; aqueous and alcohol extracts tested in CA, KB, LE, and SA.

Neuss, N., A new alkaloid from Amianthium muscaetoxicum Gray. Journal of the American Chemical Society, 1953. 75: p. 2772-2773.

Oatis Jr, J. E , P. Brunsfeld , J. W Rushing, P. D. Moeller, D. W. Bearden, T. N Gallien6  and G. Cooper IV.  2008   Isolation, purification, and full NMR assignments of cyclopamine from Veratrum californicum.  Chem. Centr. J. , 2:12doi:10.1186/1752-153X-2-12.  “The Hedgehog signaling pathway is essential for embryogenesis and for tissue homeostasis in the adult. However, it may induce malignancies in a number of tissues when constitutively activated, and it may also have a role in other forms of normal and maladaptive growth. Cyclopamine, a naturally occurring steroidal alkaloid, specifically inhibits the Hedgehog pathway by binding directly to Smoothened, an important Hedgehog response element. To use cyclopamine as a tool to explore and/or inhibit the Hedgehog pathway in vivo, a substantial quantity is required, and as a practical matter cyclopamine has been effectively unavailable for usage in animals larger than mice.  In this paper, we report a rapid and efficient isolation and purification of large quantities of cyclopamine from the roots and rhizomes of Veratrum californicum Dur. (the Corn Lily or Western false hellebore). We also provide unambiguous assignments of the carbon and proton resonances by using the multinuclear spectra and the spin coupling networks. This method could meet a very real need within diverse scientific communities by allowing cyclopamine to become more readily available.

Olney H. O.  1968. Growth substances from Veratrum tenuipetalum.  Plant Physiol 43(3):293-302. “Young leaves and buds of Veratrum tenuipetalum yielded non-indolic growth accelerators and inhibitors in the acidic ether fraction. The titer of accelerators decreased while the inhibitors increased as leaves matured. This was also true when comparing extracts of immature and fruiting inflorescences. Indole 3-acetic acid was at no time detected in leaf, bud, and inflorescence extracts but indole 3-acetyl-aspartic acid was readily recovered from all of these.The alkaline ether fraction of leaf base meristems, buds, roots, and rhizomes was rich in crystalline and amorphous alkaloids and phenolic acids. Reducing the quantity of major substances through crystallization, precipitation, and filtration permitted thin film chromatographic separation of the minor alkaloids and phenolic acids in the presence of the major ones. The unchromatographed mixture, and also certain of the purified major and minor alkaloids studied, strongly inhibited growth in germinating oat and winter rye seeds. In addition, profound changes in morphology and cytology of the seminal roots resulted. DNA disappeared partially to completely from affected tissue in 2 day germinated seeds.

Panter, K., K. Welch, D. Gardner and S. Lee. 2010. False Hellebore (Veratrum californicum): Historical perspectives and management implications for livestock and wildlife. Society for Range Management: Symposium—Medicinal Uses of Veratrum. Annual Meeting with Weed Science Society of America, Denver, CO, Feb 7–11 (30 minute oral presentation, no abstract).

Potter. D. A. 1998.  Forested communities of the Upper Montane in the Central and Southern Sierra Nevada. General Technical Report, PSW-GTR-169.

Qualtrough, D., P. Rees, B. Speight, A. C. Williams and C. Paraskeva. 2015.  The hedgehog inhibitor cyclopamine reduces -Catenin-Tcf transcriptional activity, induces E-Cadherin expression, and reduces invasion in colorectal cancer cells. Cancers 2015, 7, 1885-1899; doi:10.3390/cancers7030867. "Colorectal cancer is a major global health problem resulting in over 600,000 deaths world-wide every year with the majority of these due to metastatic disease. Wnt signalling, and more specifically -catenin-related transcription, has been shown to drive both tumorigenesis and the metastatic process in colorectal neoplasia, yet its complex interactions with other key signalling pathways, such as hedgehog, remain to be elucidated. We have previously shown that the Hedgehog (HH) signalling pathway is active in cells from colorectal tumours, and that inhibition of the pathway with cyclopamine induces apoptosis. We now show that cyclopamine treatment reduces -catenin related transcription in colorectal cancer cell lines, and that this effect can be reversed by addition of Sonic Hedgehog protein. We also show that cyclopamine concomitantly induces expression of the tumour suppressor and prognostic indicator E-cadherin. Consistent with a role for HH in regulating the invasive potential we show that cyclopamine reduces the expression of transcription factors (Slug, Snail and Twist) associated with the epithelial-mesenchymal transition and reduces the invasiveness of colorectal cancer cells in vitro. Taken together, these data show that pharmacological inhibition of the hedgehog pathway has therapeutic potential in the treatment of colorectal cancer."

Ritala-nurmi, Anneli (Helsinki, FI) Rischer, Heiko (Espoo, FI) Oksman-caldentey, Kirsi-marja (Helsinki, FI) and Rui, Ma (Jilin, CN). 2009.   Plant cell lines established from the medicinal plant Veratrum californicum. US Patent Application 20090305338, Filed 9/26/2007, published 12/10/2009.

Ruskin, A. and J. A., Rider. Veratrum viride, bio-assayed, in treatment of essential hypertension. 1950. Tex. State J. Med. 46(2): 80-84.

Sawyer, J. O., T. Keeler-Wolf & J. M. Evens. 2008. A manual of California vegetation 2nd ed. CNPS, Sacramento, CA.

Schaffner, U. D. Kleijn, V. Brown and H. Müller-Schärer. 2001. Veratrum album in montane grasslands: a model system for implementing biological control in land management practices for high biodiversity habitats. BiocontrolNews and Information 2001 22 (1): 19N – 28N.

Schep, L. J.; D. M. Schmierer and J. S. Fountain.  2006. Veratrum Poisoning.  Toxicological Reviews. 25(2): 73-78.  "Several species of the Veratrum genus are associated with toxicity in humans and animals. The principal toxins are steroid alkaloids; some have a modified steroid template, whereas others differ in their esterified acid moieties. These alkaloids act by increasing the permeability of the sodium channels of nerve cells, causing them to fire continuously. Increased stimulation, associated with the vagal nerve results in a reflex that causes the triad of responses known as the Bezold-Jarisch reflex: hypotension, bradycardia and apnoea. Clinically, various Veratrum extracts were marketed for clinical use as antihypertensive drugs, but because of their narrow therapeutic index were withdrawn from the market. Following the ingestion of Veratrum alkaloids, expected signs and symptoms include vomiting and abdominal pain, followed by cardiovascular effects such as bradycardia, hypotension and cardiac conduction abnormalities and death. Similar symptoms arise in other mammalian species ingesting these alkaloids; teratogenic effects may occur to the fetuses of animals that have grazed on Veratrum californicum. Treatment consists of supportive care, with an emphasis on haemodynamic stability with fluid replacement, atropine and vasopressors. The onset of symptoms occurs between 30 minutes and 4 hours, and the duration of the illness can range from 1 to 10 days; however, with prompt supportive care, patients typically make a full recovery within 24 hours.”

Shakirov, R., V. V. Kul'kova and I. Nakhatov. 1994. Alkaloids of Veratrum lobelianum, verdinine and 3,15-DI-O-(2-methylbutyroyl)germine.  Khimiya Prirodnykh Soedinenii 1: 100–104 [Chemistry of Natural Compounds 31(1): 79–85, 1995].  “The results are given of an investigation of the alkaloid composition of the epigeal part of Veratrum lobelianum Bernh. Veralomidine, rubijervine, germinaline, and the new bases verdinine (1) and 3,15-di-O-(2-methylbutyroyl)germine (2) were isolated. The structures of (1) and (2) have been established on the basis of their physicochemical properties and transformations. This is the first time that veralomidine has been isolated from a plant.

Society for Range Management: Symposium—Medicinal Uses of Veratrum. Annual Meeting with Weed Science Society of America, Denver, CO, Feb 7–11. “Veratrum californicum has been recognized as a poisonous plant that can be teratogenic in pregnant female sheep · The teratogenic agent has been identified as the alkaloid cyclopamine · IPI-926, a derivative of cyclopamine, is in clinical trials for cancer. Wildland harvesting practices and techniques are being developed · Site restoration measures are under investigation to generate a sustainable resource · The environmental impact is a significant consideration in harvesting.  Chair: Steve Monsen.”

Spjut, R. W. ["Rich", "President's Message"] 2016. The Search for the White Corn Lily. Mimulus Memo, June: 5–7 [California Native Plant Society, Kern Chapter].

Spjut, R. W. 2010. Potential harvest sites of Veratrum californicum in relation to its taxonomy, geographical distribution, and presence of cyclopamine and cycloposine. Society for Range Management: Symposium—Medicinal Uses of Veratrum. Annual Meeting with Weed Science Society of America, Denver, CO, Feb 7–11 (30 minute oral presentation, no abstract). Richard Spjut has been conducting geographical surveys of corn lily for the Infinity Pharmaceutical since Sep 2004, in Utah, Colorado, New Mexico, Arizona, California, Nevada, Oregon, Idaho, and Wyoming.

Spjut, R. W.  1985.  Limitation of a Random Screen: Search for New Anticancer Drugs in Higher Plants.  Economic Botany 39(3): 266–288.

Tang J., H-L. Li, Y-H. Shen, H-Z. Jin, S-K. Yan, R-H. Li, and W-D. Zhang.  2007. Four New Germine Esters from Veratrum dahuricum. Helvetica Chimica Acta 90(4): 769–775.  “Four new alkaloids, compounds 1-4, based on the germine (=4,9-epoxycevane-3,4,7,14,15,16,20-heptol; 5) framework, were isolated from the rhizomes of V. dahuricum, together with germine proper. The X-ray crystal structure of germine (5) was solved, and all compounds were characterized by circular dichroism, 1D- and 2D-NMR (1H,1H-COSY, DEPT, HSQC, HMBC), as well as HR-MS analyses.”

Tang J., H-L. Li, Y-H. Shen, H-Z. Jin, S-K. Yan, R-H. Li, and W-D. Zhang.  2007. Antitumor activity of extracts and compounds from the rhizomes of Veratrum dahuricum. Phytotherapy Res. 22: 1093–1096.  “The antitumor activity of six extracts (ethanol extract, petroleum ether fraction, CHCl3 fraction, ethyl acetate fraction, n-butanol fraction and total alkaloids) from the rhizomes of Veratrum dahuricum, and six compounds (veratramine (1), jervine (2), germine (3), veramitaline (4), veratrosine (5) and cyclopamine (6)) from the ethanol extract were investigated in vitro. The 12 samples exhibited cytotoxic activity against human tumor cell lines A549, PANC-1, SW1990 and NCI-H249. Among these samples, CHCl3 fraction, the total alkaloids, compounds 1 and 6 showed higher inhibitory activity, compound 3 selectively exhibited significant cytotoxicity to SW1990 and NCI-H249.”

Taylor, C. A. 1956a.  Alkaloid yields of Veratrum fimbriatum as influenced by site, season and other factors.  Econ. Bot. 10: 166–173. "Alkaloid content of Veratrum fimbriatum is much greater during rapid growth in early spring than the rest of the year.  Top growth at the expense of stored food reserves did not deplete alkaloids,.  Individual plants were in two categories regarding high or low percentage alkaloid producers."

Taylor, C. A. 1956b.  The culture of false hellebore.  Econ. Bot. 10: 155–165.  “Veratrum plants and seeds have winter formancy which is relieved by cold.  The germination behavior, storage life of seeds, requirements for nursery culture and possibilities of vegetation propagation are described in this article—also the habits of several species in the wild.”

USDA ARS Record of Plant Procurement for Veratrum californicum from California

USDA ARS Record of Plant Procurement for Veratrum californicum from Colorado

USDA Forest Service. Jarbidge Ranger District Rangeland Management Project. FEIS Environmental Impact Statement (Humboldt-Toiyabe National Forest). Incomplete. Not dated,  but includes references published in 2008.

Wilkins R. W., J. R. Stanton and E. D. Freis. 1949. Essential hypertension; therapeutic trial of veriloid, a new extract of Veratrum viride. Proc. Soc. Exp. Biol. Med. 72(2): 302–304.

Wolters B. 1970. [Antimicrobial activity of Veratrum alkaloids]. Planta Med. 19(2): 189–196.

Yasuhiro, T., T. Kukuchi, W. Zhao, J. Chen and Y. Guo. 1998. (+)-Verussurine, a New Steroidal Alkaloid from the Roots and Rhizomes of Veratrum nigrum var. ussuriense and Structure Revision of (+)-Verabenzoamine1. J. Nat. Prod. 61 (11): 1397–1399.  “Two minor steroidal alkaloids, 1 and 2, have been isolated from the roots and rhizomes of Veratrum nigrum var. ussuriense. Their structures have been determined by the use of spectral data as 7-O-acetyl-15-O-(2-methylbutyroyl)-3-O-veratroylgermine (1) and 15-O-(2-methylbutyroyl)-3-O-veratroylgermine (2). By spectral data comparison with verabenzoamine, the structure of the latter compound has been revised from the previously reported 7-O-acetyl-15-O-(2-methylbutyroyl)-3-O-veratroylgermine (1) to 15-O-(2-methylbutyroyl)-3-O-veratroylgermine (2). Accordingly, alkaloid 1 [7-O-acetyl-15-O-(2-methylbutyroyl)-3-O-veratroylgermine] must be new, and it was given the trivial name verussurine.”

Yagi, A. and T. Kawasaki. 1962. [Alkaloids of Japanese Veratrum genus plants. V. Alkaloids of Veratrum stamineum.] J. Pharmacol. 82: 210–213.

Youngken H. W. 1953. Studies on Veratrum. II. Veratrum eschscholtzii A. Gray; observations on seed germination and early growth of seedlings of Veratrum species. J. Am. Pharm. Assoc. Am. Pharm. Assoc. (Baltim.) 42(1): 39-45.

Youngken H. W. 1952. A pharmacognostical study of roots of different species of Veratrum. J. Am. Pharm. Assoc. 41(7): 356-361.

Zhao W., Y. Guo, S. Wang, T. Shao, Y. Tezuka and T. Kikuchi.  1998. [Chemical research on stilbenes from Veratrum macckii Reg.]. Zhongguo Zhong Yao Za Zhi. 23(10): 619–620, 640.“OBJECTIVE: To study the components in rhizome of Veratrum macckii. METHODS: Column chromatography and preparative thin layer chromatography with silica gel were employed for the isolation and purification of constituents. The structures were elucidated by IR, MS and 1H-NMR analysis. RESULTS: Two compounds were obtained and elucidated as resveratrol and 2,3',4,5'-tetrahydroxystilbene. CONCLUSION: The two compounds were separated from V. macckii for the first time.”

Zhao, W., W. Hao, Y. Tezuka,  T. Kikiuchi,  J. Chen and Y. Guo. 1991. Studies on the Constituents of Veratrum Plants. II. Constituents of Veratrum nigrum L. var. ussuriense. (1). Structure and 1H-and 13C-Nuclear Magnetic Resonance Spectra of a New Alkaloid, Verussurinine, and Related Alkaloids. Chem. & Pharm. Bull. (Pharm. Soc. Japan): 39(3): 549–554. “Alkaloidal constituents of the roots and rhizoma of Veratrum nigrum L. var. ussuriense (Liliaceae), which are used as a source of the Chinese crude drug "Li-lu, " were examined and a new alkaloid named verussurinine and six known alkaloids have been isolated. The structure of verussurinine was determined to be 16-O-(2-methylbutyroyl)germine (1) by means of spectroscopic methods, and six other alkaloids were identified as germidine (2), germerine (3), 15-O-(2-methylbutyroyl)germine (4), verazine (5), jervine (6), and neogermbudine (7), Complete assignments of the proton and carbon-13 nuclear magnetic resonance (^1H- and ^<13>C-NMR) signals of these alkaloids are also presented.

Zhou C-X., J-Y. Liu, W-C. Yeb, C-H Liu and R-X. Tan. 2006. Neoverataline A and B, two antifungal alkaloids with a novel carbon skeleton from Veratrum taliense. Tetrahedron 59(30): 5743–5747. “Bioassay-guided fractionation of the ethanol extract of the roots and rhizomes of Veratrum taliense yielded two new and thirteen known steroidal alkaloids. The structures of the two new compounds, neoverataline A and B, were established by extensive spectroscopic analyses to be 3,4-secocevane-4,9-olid-14,15,16,20-tetra-ol-3-oic acid and 3,4-secocevane-4,9-olid-7,14,15,16,20-penta-ol-3-oic acid, respectively, and are a novel carbon skelton. All of the fifteen alkaloids were subjected to in vitro antifungal assays, which showed that the verazine- (veramitaline, stenophylline B, stenophylline B-3-O-β-Image-glucopyranoside, veramiline-3-O-β-Image-glucopyranoside) and jerveratrum-type (jervine, jervine-3-O-β-Image-glucopyranoside) alkaloids exhibited strong antifungal activities against the phytopathogenic fungus Phytophthora capisis with MICs of 160, 120, 160, 80, 80 and 120 μg·L−1, respectively. Furthermore, the verazine-type alkaloids stenophylline B, stenophylline B 3-O-β-Image-glucopyranoside and veramiline 3-O-β-Image-glucopyranoside were shown to also inhibit the growth of another fungal phytopathogen Rhizoctonia cerealis with MICs of 160, 120 and 120 μg mL−1. The MICs of triadimefon (an antifungal agrochemical used herein as a positive control) against P. capisis and R. cerealis were 120 and 80 μg mL−1, respectively. A preliminary structure–activity relationship regarding these alkaloids has been formulated. ”

Zhou C. X., J. Tanaka, C. H. Cheng, T. Higa and R. X. Tan.  1999.  Steroidal Alkaloids and Stilbenoids from Veratrum taliense.  Planta Med. 65(5): 480–482. “Phytochemical investigation of roots and rhizomes of Veratrum taliense yielded a new and six known steroidal alkaloids as well as a new and one reported stilbene derivative. By a combination of spectral methods (IR, MS, (1)H- and (13)C-NMR, COSY, HMQC, HMBC, and NOESY), the structure of the new alkaloid was established as 15-angeloylgermine while the known ones were identified as 15-(2-methylbutyroyl)germine, jervine, 3-veratroylzygadenine, germine, veramiline 3- O-(beta- D-glucopyranoside and stenophylline B-3- O-beta- D-glucopyranoside. The new stilbenoid, named veraphenol, was determined to be 2-(3',5'-dihydroxyphenyl)-6-hydroxybenzofuran, and the known one was shown to be resveratrol. The IN VITRO enzyme assay indicated that 3-veratroylzygadenine and resveratrol are inhibitors of xanthine oxidase. The enzyme inhibitory action of resveratrol, the most active compound found so far in V. TALIENSE, is dose-dependent with the IC (50) value at 30 microM (the IC (50) value of allopurinolused as a positive control in the study is 10 microM).

Zimmerman J. H. 1958. A monograph of Veratrum. Ph.D. dissertation, University of Wisconsin, Madison, Wisconsin, USA.

Zomlefer, W. B, W. S. Judd,  W. M. Whitten and N. H. Williams. 2006. A synopsis of Melanthiaceae (Liliales) with focus on character evolution in tribe Melanthieae. Aliso 22: 566–578.

Zomlefer, W. B., W. M. Whitten, N. H. Williams and  and W. S. Judd.  2003. An Overview of Veratrum s.l. (Liliales: Melanthiaceae) and an Infrageneric Phylogeny Based on ITS Sequence Data. Syst. Bot. 28: 250–269. “A synopsis of Veratrum, including commentary on species and character evolution within the genus, is presented. The circumscription and relationships of infrageneric taxa are evaluated using parsimony analyses of ITS (nuclear ribosomal) DNA sequence data of 26 representative taxa. Proposed new infrageneric circumscriptions, strongly supported by tree statistics and topologies, are correlated with potential morphological synapomorphies at the proper level of universality. Based on our analyses, Veratrum is circumscribed broadly (including Melanthium) and divided into two sections and two subsections (most with novel circumscription). This modified infrageneric classification involves reassignment of Veratrum subgenus Pseudoanticlea as subsection Pseudoanticlea. Although interspecific relationships are not highly resolved, the molecular data provide strong support for placing several species previously of unknown affinities and also validate several generalizations concerning character evolution within Veratrum.”

 Zomlefer, W. B., N. H. Williams, M. Whitten and W. S. Judd.  2000.  Generic circumscription and relationships in the tribe Melanthieae (Liliales, Melanthiaceae), with emphasis on Zigadenus: evidence from ITS and trnL-F sequence data. Am. J. Bot. 88: 1657–1699.  “The circumscription and relationships of genera within the tribe Melanthieae (29 representative taxa) were evaluated using parsimony analyses of ITS (nuclear ribosomal) and trnL-F (plastid) DNA sequence data, alone and in combination. Proposed new generic circumscriptions, strongly supported by the tree statistics and topologies in all analyses, are correlated with potential morphological synapomorphies at the proper level of universality. Based on the molecular cladograms, Stenanthium is biphyletic, and the traditional Zigadenus s.l. (sensu lato) is polyphyletic. Amianthium and Schoenocaulon are distinct entities; the Veratrum complex is conservatively treated as one large monophyletic genus (including Melanthium). Although some generic relationships are not highly resolved, the analyses provide strong support for Zigadenus glaberrimus as sister to the rest of the tribe, and Amianthium muscitoxicum as closely related to Veratrum s.l. As a result of these analyses, seven genera (some with novel circumscription) are recognized within the tribe Melanthieae: Amianthium, Anticlea, Schoenocaulon, Stenanthium, Toxicoscordion, Veratrum, and Zigadenus.”

Zomlefer W. B. and K. D. Perkins 1999. Phylogeny of the Melanthiaceae. Available at: and

Zomlefer W. B. 1997a The genera of Melanthiaceae in the southeastern United States. Harvard Papers in Botany 2: 133-177.