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486 COLCHICACEAE/UVULARIA<br />

free, pale yellow, promptly deciduous; stamens 6, ca. 1/3–1/2 <strong>the</strong> length <strong>of</strong> <strong>the</strong> perianth; filaments<br />

shorter than an<strong>the</strong>rs; ovary superior, 3-locular; style 1; stigmas 3; fruit a 3-angled dehiscent<br />

capsule; seeds 1–3 per locule.<br />

AA genus <strong>of</strong> 5 species (Nordenstam 1998; Utech & Kawano 2002) endemic <strong>to</strong> e North<br />

America. They are rhizoma<strong>to</strong>us spring-flowering herbs with arching stems, 2-ranked leaves,<br />

and pendulous flowers. The <strong>plants</strong> vegetatively superficially resemble Polygonatum,<br />

SOLOMON’S-SEAL (Convallariaceae, order Asparagales), but can be easily distinguished by <strong>the</strong><br />

free perianth parts (vs. perianth parts fused nearly <strong>to</strong> <strong>the</strong> tips in Polygonatum). Some are cultivated<br />

as ornamentals. According <strong>to</strong> Yatskievych (1999), “Young shoots <strong>of</strong> Uvularia species have<br />

been cooked and eaten like asparagus, and <strong>the</strong> starchy roots<strong>to</strong>cks <strong>of</strong> some species have also<br />

been boiled and eaten.” The roots were also used as a salve for wounds and sores (Bush 2001).<br />

Elaiosomes/arils (= external food bodies) are present on <strong>the</strong> seeds, and <strong>the</strong> seeds <strong>of</strong> at l<strong>east</strong> some<br />

species are dispersed by ants (Handel et al. 1981). The common name, BELLWORT, is probably derived<br />

from <strong>the</strong> bell-shaped nodding flowers and <strong>the</strong> Anglo-Saxon, wyrt, a plant or herb. (Latin:<br />

uvula, <strong>the</strong> s<strong>of</strong>t ana<strong>to</strong>mical structure hanging in <strong>the</strong> human throat, diminutive <strong>of</strong> uva, grape,<br />

apparently in reference <strong>to</strong> <strong>the</strong> pendulous flowers; <strong>the</strong> plant was once thought <strong>to</strong> be a remedy for<br />

diseases <strong>of</strong> <strong>the</strong> uvula—Shosteck 1974)<br />

REFERENCES: Anderson & Whitaker 1934; Dietz 1952; Wilbur 1963; Kawano & Iltis 1964;<br />

Whigham 1974; Seibert & Savidge 1991; Reveal 1992; Hawashi et al. 1998; Kudoh et al. 1999;<br />

Bush 2001; Wijesinghe & Whigham 2001; Utech & Kawano 2002; Singhurst et al. 2003a.<br />

1. Leaves perfoliate (= with leaf base completely surrounding <strong>the</strong> stem); flower borne on a flower<br />

stalk (resembling a side branch) bearing a single perfoliate leaf-like bract; capsules obtusely 3angled,<br />

� truncate (= cut <strong>of</strong>f squarely) at tip, with 2 attenuate beaks on each <strong>of</strong> <strong>the</strong> 3 lobes _____ U. perfoliata<br />

1. Leaves sessile; flower borne on a bractless flower stalk from axil <strong>of</strong> a leaf along main stem; capsules<br />

sharply 3-angled, ellipsoid and � pointed at tip, not beaked _____________________________ U. sessilifolia<br />

Uvularia perfoliata L., (perfoliate, with <strong>the</strong> leaf surrounding or embracing <strong>the</strong> stem), PERFOLIATE<br />

BELLWORT, MEALY BELLWORT, LARGE BELLWORT. Plant with slender s<strong>to</strong>lons in addition <strong>to</strong> short rhizome;<br />

stems 15–40(–60) cm tall at flowering time, terete; leaves elliptic <strong>to</strong> ovate-lanceolate, 3.5–<br />

12.5 cm long, 1.5–5(–6) cm wide, perfoliate, <strong>the</strong> margins smooth; flower 1 per leafy perfoliate<br />

bract (commonly 1 but sometimes up <strong>to</strong> several flowers per stem); perianth (10–)17–30(–37)<br />

mm long, granular-papillose on inner surface; an<strong>the</strong>rs 6–10 mm long; connectives <strong>of</strong> filaments<br />

exserted more than 0.8 mm beyond an<strong>the</strong>rs; ovary sessile; capsule 7–13(–15) mm long, truncate<br />

apically; seeds arillate. Moist woods, hardwood bot<strong>to</strong>ms and lower slopes, <strong>of</strong>ten in ravines or in<br />

or near hillside seeps, acidic sandy soils or calcareous substrates; Sabine (BRIT), San Augustine<br />

(TAMU), Shelby (TOES 1993), and Nacogdoches (Singhurst et al. 2003a) cos. in <strong>the</strong> Pineywoods;<br />

first documented for TX by Nixon et al. (1987); se Canada and widespread in <strong>the</strong> e U.S. w <strong>to</strong> OH<br />

and TX. Spring. A life his<strong>to</strong>ry study <strong>of</strong> this species in North Carolina (Whigham 1974) found<br />

that <strong>the</strong> primary pollina<strong>to</strong>r was a flightless staphylinid beetle, ants dispersed <strong>the</strong> seeds, asexual<br />

reproduction was much more important than sexual reproduction, <strong>the</strong> species was largely selfincompatible,<br />

and growth and reproduction occurred during <strong>the</strong> “very short period <strong>of</strong> time between<br />

<strong>the</strong> onset <strong>of</strong> favorable growing conditions and closure <strong>of</strong> <strong>the</strong> overhead canopy.” Seibert<br />

and Savidge (1991) found that <strong>the</strong> principle pollina<strong>to</strong>rs <strong>of</strong> a similar species, U. grandiflora Sm.,<br />

were queen bumblebees; <strong>the</strong>y questioned Whigham’s (1974) finding concerning pollination because<br />

“<strong>the</strong> two Uvularia species are morphologically and phenologically very similar.” The lifehis<strong>to</strong>ry<br />

strategy <strong>of</strong> this species has been described as a “waiting strategy,” with predominantly<br />

vegetative reproduction under suboptimal closed canopy conditions and both vegetative and<br />

sexual reproduction under optimal conditions in canopy gaps (Kudoh et al. 1999). (TOES 1993:<br />

V) � m/306<br />

Uvularia sessilifolia L., (sessile-leaved), SESSILE-LEAF BELLWORT, SMALL BELLWORT, WILD-OATS,<br />

STRAW-LILY. Plant with long slender rhizomes; stems 15–40(–45) cm tall at flowering time,

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