Shrubs, trees, or lianas, often epiphytic, evergreen or deciduous, with alternate, non-ericoid leaves with margin entire to serrulate or serrate, convolute or less commonly revolute in bud. Leaf epidermal cells lignified or not. Indumentum of elongate and unbranched, or peltate, and glandular to non-glandular hairs, and occasionally papillae. Inflorescence terminal or axillary, racemose, paniculate, or fasciculate, or of solitary flowers; bracts present, bracteoles usually paired, variable in position; flower articulated or continuous with pedicel. Flowers usually 4- or 5-merous, actinomorphic, pendulous. Calyx lobes usually 4 or 5, variable in size, usually persistent, sometimes fleshy; corolla usually sympetalous, urceolate to cylindrical or campanulate, deciduous, with the usually 4 or 5 lobes small to less commonly large; stamens usually 5—10, usually included, the filaments straight to S-shaped, distinct or connate, unicellular-pubescent or glabrous (smooth or roughened), rarely papillose; anthers tetrasporangiate, inverting early in development, ± roughened, with or without a pair of dorsal appendages (spurs), or these occurring instead on the distal portion of the filament, or with 2 or 4 projections (awns) arising from the dorso-apical portion of the anther, with or without disintegration tissue, lacking a fibrous endothecium, dehiscing by terminal pores, and these often associated with tubules. Pollen in tetrads, lacking viscin threads. Ovary usually 4- or 5-locular, or 10-locular due to secondary division of each locule, with axile placentation, superior to inferior; style usually impressed; stigma truncate. Fruit a loculicidal capsule, berry, or drupe; seeds with variable testa cells; embryo with 2 cotyledons.

The Vaccinieae are easily recognized because they possess a distinctive combination of synapomorphic features, i.e., anther tubules (char. #62, also found in parallel in a few Gaultherieae), an inferior ovary (char. #72; Fig. 29), and indehiscent, brightly colored fruits
(chars. #78, 82), i.e., usually berries, but drupes in Gaylussacia. It is worth noting that fleshy, colorful fruits (either drupes or berries) have also evolved in some Styphelioideae (i.e., Styphelieae), some Gaultherieae, Empetreae, and Arbutoideae (Fig. 30), likely in connection
with a shift from wind to bird dispersal. Generic relationships within Vaccinieae are very problematic (Kron et al., 1999a; Luteyn, 1996) and are currently being investigated by Kron and Luteyn.

genus: Vaccinium L.

Thibaudieae Benth. & Hook.f., Gen. Pl. 2: 564. 1876.
Vacciniiae Klotzsch, Monatsber. Königl. Preuss. Akad. Wiss.
Berlin 1857: 4. 1857 (as Vaccinieae).

Shrubs, small trees, or lianas, often epiphytic, evergreen or deciduous, with, non-ericoid entire or serrate leaves, convolute in bud. Leaf epidermal cells not lignified. Indumentum of short- to long-stalked glandular or non-glandular hairs. Inflorescence usually axillary, rarely terminal, paniculate, racemose or fasciculate, or solitary axillary flowers; bracts present, bracteoles usually paired, variable in position; calyx articulated with the pedicel or not. Flowers usually 4- or 5-merous, rarely 3 or 6-7-merous, actinomorphic. Calyx lobes small to large, persistent, the calyx winged or not; corolla almost always sympetalous, usually urceolate or tubular, sometimes winged, the lobes small to large; stamens 5—10(-14), the filaments ± straight, sometimes connate, unicellular pubescent, roughened, or smooth, sometimes spurred (with either one spur or, more commonly, a pair of spurs) on the anther-filament junction; anthers slightly to clearly roughened, usually lacking disintegration tissue, usually with tubules, dehiscing by pores or slit-like pores at the end of the tubules. Ovary (3-)4—5(-7)-locular, or 10-locular due to secondary division of each locule, with axile placentation, inferior; style impressed; stigma ± truncate. Fruit fleshy, a berry or 10-pitted drupe; seeds with short to elongate testa cells; embryo with 2 cotyledons.

    Traditionally this tribe was broadly circumscribed, including genera
here placed in Lyonieae, Gaultherieae, and Oxydendreae (Stevens, 1971).
As traditionally delimited the group clearly is paraphyletic (Fig. 7; 
see also discussion in Stevens, 1995; Kron et al., 1999b). In this 
classification Andromedeae is restricted to only Andromeda and Zenobia 
and may be characterized by a lack of multicellular hairs on the leaves
(disregarding the reduced non-glandular hairs associated with the 
marginal teeth of Zenobia; see discussion in Kron et al., 1999b). The
leaves of Andromeda are entire, and thus lack these tooth-associated 
hairs (or any other multicellular hairs). Despite the lack of 
distinctive morphological synapomorphies, the monophyly of this small
clade is strongly supported in the molecular (Figs. 4,5,6) and combined
analyses (Fig. 7).

     Andromedeae Klotzsch, Pfl. Abbild. Beschr.: 1838. – Type genus: 
ndromeda L.Andromedinae D. Don, New Philos. J. Edinburgh 17: 157. 1834 
 (as Andromedeae).

     Shrubs, evergreen, with, non-ericoid, entire or serrulate leaves, 
convolute or revolute in bud. Secondary phloem with bands of fibers in 
Zenobia, these lacking in Andromeda; leaf epidermal cells ± lignified 
in Andromeda, but unlignified in Zenobia. Indumentum of multicellular 
hairs lacking except for ± globose, eglandular hairs associated with 
serrations of Zenobia. Inflorescence axillary or terminal, 
± fasciculate; bracts present, bracteoles paired, basal; calyx 
articulated with pedicel. Flowers 5-merous, actinomorphic. Calyx 
lobes small, persistent; corolla sympetalous, urceolate or campanulate,
the lobes small; stamens 10, the filaments ± straight, unicellular-
pubescent or roughened, with 2 or 4 awns arising from the dorso-apical
portion of the anther; anthers slightly to clearly roughened, with a 
white deposit of disintegration tissue dorsally in Zenobia, but this 
lacking in Andromeda, dehiscing by terminal pores. Ovary 5-locular, 
with axile placentation, superior; style impressed; stigma ± truncate.
Fruit a loculicidal capsule; seeds with elongate or ± isodiametric 
testa cells; embryo with 2 cotyledons.

Although the Gaultherieae clade is present in all of the equally parsimonious trees discovered in the combined analysis (Fig. 7), its monophyly is not (as yet) well supported. It is probably most closely related to the Andromedeae, and it is likely significant that 4-appendaged anthers (char. #61; Fig. 31) and a base chromosome number of eleven (char. #89) characterize Gaultherieae and also occur in Zenobia. A strongly supported subclade (comprising Diplycosia, Tepuia, Gaultheria, and Pernettya) is diagnosed by the presence of methyl salicylate (char. #88). This fragrant compound, which gives the plants a distinct wintergreen odor, has been lost in several species, especially within Gaultheria. Generic relationships are problematic within this subclade, and phylogenetic relationships within this group are currently being investigated (Powell & Kron, 2001).