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TRILLIUM/TRILLIACEAE<br />

1131<br />

Schilling 2002; Judd et al. 2002). Species <strong>of</strong> all <strong>the</strong> genera are cultivated (Zomlefer 1996). While<br />

<strong>the</strong> traditional view was that <strong>the</strong>re were 2 genera (Paris, Tr illium) in <strong>the</strong> family, more recently,<br />

different authors have recognized ei<strong>the</strong>r 4 genera (e.g., Tamura 1998d) or only a single genus<br />

(Judd. et al. 2002). However, <strong>the</strong> most recent study, based on both molecular and morphological<br />

data, suggests that 6 genera may be warranted (Farmer & Schilling 2002). The genera have<br />

been variously treated in terms <strong>of</strong> family affiliation—since 1846 members <strong>of</strong> <strong>the</strong> family have<br />

been placed in five different families (Farmer & Schilling 2002). Some authorities have put<br />

<strong>the</strong>m in a very broad and clearly polyphyletic (but practical) Liliaceae (e.g., Correll & Johns<strong>to</strong>n<br />

1970; Cronquist 1988). However, based on phylogenetic analyses, we are following Dahlgren et<br />

al. (1985), Zomlefer (1996), Tamura (1998d), Judd et al. (1999, 2002), and Farmer and Schilling<br />

(2002) in recognizing a narrowly defined Trilliaceae; as such, <strong>the</strong> family is monophyletic (Ka<strong>to</strong><br />

et al. 1995b). The family is related <strong>to</strong> <strong>the</strong> Melanthiaceae (in <strong>the</strong> narrow sense as treated here),<br />

and additional research is needed <strong>to</strong> clarify that relationship (Zomlefer 1997a). Indeed, some recent<br />

molecular and morphological analyses (e.g., Chase et al. 2000; Fuse & Tamura 2000;<br />

Rudall et al. 2000b; Zomlefer et al. 2001) have suggested that Trillium and related genera be included<br />

in Melanthiaceae, a course followed by <strong>the</strong> Angiosperm Phylogeny Group (APG II 2003).<br />

While clearly related <strong>to</strong> that family, such a treatment <strong>of</strong> Trillium and allies is, however, problematic<br />

in that it groups <strong>to</strong>ge<strong>the</strong>r <strong>plants</strong> <strong>of</strong> radically different morphologies (1 showy versus<br />

many small flowers, 3 bracts in a whorl versus basal leaves, berry fruit versus capsules, etc.). In<br />

fact, “<strong>the</strong>re are no morphological synapomorphies that unite <strong>the</strong>se two groups as separate from<br />

o<strong>the</strong>r Liliaceous <strong>plants</strong>” (Farmer & Schilling 2002). As a result, we are taking what we consider<br />

<strong>to</strong> be an intermediate, practical approach in recognizing <strong>the</strong> Trilliaceae as a separate family. For<br />

a detailed discussion <strong>of</strong> <strong>the</strong> groups formerly treated as Liliaceae in <strong>the</strong> broad sense, see <strong>the</strong> family<br />

synopsis <strong>of</strong> <strong>the</strong> Liliaceae (here treated in a restricted sense) on page 726. Some members <strong>of</strong><br />

<strong>the</strong> Trilliaceae have been used medicinally (e.g., species <strong>of</strong> Paris, Trillium), and Paris quadrifolia<br />

L., HERBA PARIS, native <strong>to</strong> Europe and n Asia, is a well known medicinal and poisonous plant still<br />

used in homeopathy “for neurological disorders, gout, rheumatism, and o<strong>the</strong>r inflamma<strong>to</strong>ry<br />

maladies” (Zomlefer 1996); � its fruits are considered very poisonous (Zomlefer 1996). (subclass<br />

Liliidae—Cronquist; order Liliales—APG II)<br />

FAMILY RECOGNITION IN THE FIELD: <strong>plants</strong> with leaves (actually leaf-like bracts) in a single whorl<br />

<strong>of</strong> 3 at summit <strong>of</strong> <strong>the</strong> stem, flower 1 per plant, 3 showy petals, and fruit fleshy and berry-like.<br />

REFERENCES: Dahlgren et al. 1985; Ka<strong>to</strong> et al. 1995b; Zomlefer 1996; Tamura 1998d; Fuse &<br />

Tamura 2000; Rudall et al. 2000b; Farmer & Schilling 2002.<br />

TRILLIUM L. WAKE-ROBIN, BIRTHROOT,<br />

TRILLIUM, TOAD-SHADE, SQUAW-ROOT, CARRION-FLOWER<br />

Perennial rhizoma<strong>to</strong>us herbs, lacking <strong>the</strong> odor <strong>of</strong> onion or garlic; stems unbranched, erect or<br />

nearly so, glabrous or slightly scabrous; leaf-like bracts (<strong>of</strong>ten referred <strong>to</strong> as leaves since <strong>the</strong>y are<br />

<strong>the</strong> primary pho<strong>to</strong>syn<strong>the</strong>tic structures <strong>of</strong> <strong>the</strong> plant) usually 3, in a distinctive whorl at summit<br />

<strong>of</strong> stem, glabrous, with net venation; flowers perfect, solitary, sessile or pedicelled, at summit <strong>of</strong><br />

stem with leaf-like bracts; perianth parts 6, in two series, free, <strong>the</strong> outer series (sepals) green or<br />

with some purplish pigmentation, persistent; inner perianth segments (petals) white <strong>to</strong> pink,<br />

purple, greenish, or yellow, not persistent; stamens 6; an<strong>the</strong>rs linear, ca. as long as or longer than<br />

filaments; ovary superior; fruits fleshy, berry-like.<br />

AA genus <strong>of</strong> 43 species (Case 2002) with some authorities recognizing up <strong>to</strong> ca.48–50 species<br />

(Zomlefer 1996; Tamura 1998d; Kazempour Osaloo et al. 1999). They are rhizoma<strong>to</strong>us herbs with<br />

an apical whorl <strong>of</strong> three leaf-like bracts and a single flower and are disjunctly distributed in w<br />

North America, e North America, and <strong>the</strong> Himalayas <strong>to</strong> e Asia, with <strong>the</strong> center <strong>of</strong> diversity in<br />

<strong>the</strong> Appalachian mountain region (Zomlefer 1996; Tamura 1998d). The genus is <strong>of</strong>ten considered<br />

an Arc<strong>to</strong>-Tertiary element (Tamura 1998d), and according <strong>to</strong> Kazempour Osaloo et al.

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