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746 MELANTHIACEAE/ZIGADENUS<br />

addressed <strong>the</strong>se issues, showing that Melanthium is derived from within Veratrum, and supporting<br />

<strong>the</strong> monophyly <strong>of</strong> Veratrum in <strong>the</strong> broad sense. Therefore, treatment <strong>of</strong> Melanthium<br />

within Veratrum now seems most appropriate. � Veratrum (FALSE or WHITE HELLEBORE) species<br />

have long been important medicinally and as poisonous <strong>plants</strong>. They have been used medicinally<br />

since ancient times in Eurasian folk medicine, by various Native American tribes, and<br />

from <strong>the</strong> early 1800s until recently by medical doc<strong>to</strong>rs for <strong>the</strong> control <strong>of</strong> hypertension; <strong>the</strong>y<br />

have also been used in sorcery, mystical rites, as crow poisons, and as insecticides (Kupchan et<br />

al. 1961; Zomlefer 1997a; Burrows & Tyrl 2001; Alm 2002). The hypotensive, tera<strong>to</strong>genic, and/or<br />

neuro<strong>to</strong>xic properties are <strong>the</strong> result <strong>of</strong> Veratrum alkaloids (e.g., veratradine, veratramine), a<br />

physiologically powerful group <strong>of</strong> very complex molecules. Approximately 100 <strong>of</strong> <strong>the</strong>se compounds<br />

have been isolated, even though relatively few species have been thoroughly analyzed<br />

(Zomlefer et al. 2003). (Latin: vere, true, and aater, black, in reference <strong>to</strong> <strong>the</strong> black rhizomes<br />

found in some species—McNeal & Shaw 2002)<br />

REFERENCES: Zimmerman 1958; Bodkin 1978, 1998; Zomlefer 1997a; Tamura 1998d; Zomlefer et<br />

al. 2001; Bodkin & Utech 2002; McNeal & Shaw 2002; Zomlefer et al. 2003.<br />

Veratrum virginicum (L.) W.T. Ai<strong>to</strong>n, (<strong>of</strong> Virginia), BUNCHFLOWER. Perennial, short rhizoma<strong>to</strong>us<br />

herb lacking <strong>the</strong> odor <strong>of</strong> onion or garlic; stem base slightly bulbose, with brown fibers derived<br />

from old leaf bases; stem � erect, 0.6–1.5(–2) m tall, downy or scurfy above; leaves mostly basal,<br />

those on <strong>the</strong> stem reduced and mostly bract-like; basal leaves 20–50(–80) cm long, 4–32 mm<br />

wide, broadly linear; inflorescence a terminal panicle; pedicels 4–22 mm long, subtended by<br />

small bracts, <strong>the</strong>se densely floccose on margins and lower surfaces; flowers perfect or sometimes<br />

those near tip <strong>of</strong> inflorescence functionally staminate, numerous; tepals 6, 5–13 mm long,<br />

fused at <strong>the</strong> base, clawed, with a pair <strong>of</strong> yellow <strong>to</strong> greenish brown, nectariferous glands at base<br />

<strong>of</strong> � ovate blade portion, whitish <strong>to</strong> pale yellow, turning greenish yellow or reddish purple; claw<br />

< 1/2 as long as blade; stamens 6, attached near or above middle <strong>of</strong> each claw; styles 3; ovary<br />

superior; fruit a 3-lobed capsule 10–18 mm long; seeds flat; 2n = 16 (Bodkin & Utech 2002).<br />

Bogs, swales, savannahs, meadows, forest margins; Angelina, Jasper (ASTC), Anderson, Jefferson,<br />

and Milam (Turner et al. 2003) cos. in <strong>the</strong> Pineywoods and Post Oak Savannah; widespread<br />

in e U.S. w <strong>to</strong> KS, OK, and TX. May–Jul. [Melanthium dispersum Small, Melanthium virginicum<br />

L.] This species contains a number <strong>of</strong> poisonous alkaloids (Yatskievych 1999). � m/306<br />

ZIGADENUS Michx.<br />

DEATH-CAMAS, POISON-SEGO, STAR-LILY, ZYGADENE<br />

A A monotypic genus <strong>of</strong> <strong>the</strong> se U.S. represented only by Z. glaberrimus. Recent chromosomal,<br />

molecular, and morphological studies (Zomlefer et al. 2001; Zomlefer & Judd 2002) have necessitated<br />

significant changes in generic circumscription <strong>of</strong> <strong>the</strong> tribe Melanthieae (which includes all<br />

East TX members <strong>of</strong> <strong>the</strong> family). In <strong>the</strong> past, Zigadenus has been variously circumscribed—<br />

Zomlefer (1997a) provisionally recognized 17 species, included <strong>the</strong> monotypic segregate<br />

Amianthium, and noted that <strong>the</strong> genus represents a “heterogeneous, non-monophyletic grouping”<br />

needing study. Tamura (1998b) circumscribed <strong>the</strong> genus more narrowly (ca. 10 species) and<br />

excluded Amianthium. Schwartz (2002) had a more inclusive view, recognizing 18–22 species. In<br />

all <strong>of</strong> <strong>the</strong>se conceptions, <strong>the</strong> genus was a heterogeneous assemblage <strong>of</strong> only distantly related species.<br />

This is a clear example <strong>of</strong> new research requiring scientific names <strong>to</strong> be changed (despite<br />

some nomenclatural confusion). Extensive recent research (Zomlefer et al. 2001) has shown that<br />

Z. glaberrimus is an isolated taxon on a separate evolutionary branch from <strong>the</strong> rest <strong>of</strong> <strong>the</strong> tribe—it<br />

thus has <strong>to</strong> be recognized as a separate genus. Because Z. glaberrimus is <strong>the</strong> type species <strong>of</strong><br />

Zigadenus, different generic names have <strong>to</strong> be used for <strong>the</strong> o<strong>the</strong>r groups <strong>of</strong> species previously recognized<br />

in <strong>the</strong> polyphyletic Zigadenus (Zomlefer & Judd 2002). For example, Z. densus and Z.<br />

nuttallii are now treated in Stenanthium and Toxicoscordion respectively. Previously sometimes<br />

spelled Zygadenus, an orthographic variant that does not have nomenclatural priority. (Greek:<br />

zygos, a yoke, and aden, a gland, referring <strong>to</strong> <strong>the</strong> glands on <strong>the</strong> tepals sometimes being in pairs)

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