Lyonieae are clearly supported as the sister group to a large clade comprising Vaccinieae, Andromedeae, and Gaultherieae. The Lyonieae are diagnosed by bands of fibers in the phloem (char. #7; Fig. 27), a lignified leaf epidermis (char. #21; Fig. 22), and S-shaped filaments (char. #53, which have been lost in some species of Pieris, and have evolved in parallel in some Styphelioideae and Ericoideae). The phylogenetic relationships within this clade are discussed in detail in Kron & Judd (1997) and Kron et al. (1999b). The monophyly of the Gaultherieae + Andromedeae + Vaccinieae clade is not apparent morphologically, although the clade has a bootstrap value of 98% in the combined analysis (see Fig. 7). Most of the members of this clade, however, possess paracytic stomata (char. #28), although anomocytic stomata are present in Andromeda and Zenobia (Fig. 28).
Lyonieae Kron & Judd, Amer. J. Bot. 86: 1298. 1999. – Type genus: Lyonia Nutt.

Shrubs, rarely lianas, evergreen or deciduous, with, non-ericoid entire, serrulate, or serrate leaves, convolute or less commonly revolute in bud. Secondary phloem with bands of fibers; leaf epidermal cells ± lignified. Indumentum of stalked glandular and/or elongate and eglandular hairs, or ferrugineous, peltate scales. Inflorescence usually axillary, rarely terminal, racemose, paniculate, or fasciculate; bracts present, bracteoles paired, variable in position; calyx usually articulated with pedicel. Flowers usually 5-merous, but rarely 4- to 8-merous, actinomorphic. Calyx lobes usually small, persistent; corolla sympetalous, usually urceolate to tubular, the lobes small; stamens usually 10, the filaments usually S-shaped, unicellular-pubescent, roughened, or smooth, sometimes with paired spurs on the anther-filament junction or upper portion of filament; anthers slightly to clearly roughened, with a white deposit of disintegration tissue dorsally, dehiscing by terminal pores. Ovary usually 5-locular, with axile placentation, superior; style impressed; stigma ± truncate. Fruit a loculicidal capsule; seeds usually with elongate testa cells; embryo with 2 cotyledons.
Full Detailed Description of Vaccinioideae
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