Vaccinioideae

 Shining fetterbush Lyonia lucida, photographed by Will Cook.

Belongs within: Ericaceae.
Contains: Vaccinieae, Gaultherieae.

The Vaccinioideae are a clade within the Ericaceae strongly supported by molecular analysis, though the only synapomorphies of the clade—the presence of disintegration tissue on the back of the anthers, and a base chromosome number of 12—are homoplasious. Members include the Lyonieae, a group of shrubs (rarely lianas) with more or less lignified leaf epidermal cells and usually S-shaped stamen filaments (Kron et al. 2002).

Characters (from Kron et al. 2002): Shrubs, trees, or lianas, often epiphytic. Leaves evergreen or deciduous, alternate, non-ericoid, 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, filaments straight to S-shaped, distinct or connate, unicellular-pubescent or glabrous (smooth or roughened), rarely papillose; anthers tetrasporangiate, inverting early in development, more or less 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 dorso-apical portion of 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.

<==Vaccinioideae
    |--Oxydendrum [Oxydendreae]KJ02
    |    `--O. arboreumKJ02
    `--+--+--VaccinieaeKJ02
       |  `--+--GaultherieaeKJ02
       |     `--Andromedeae [Andromedinae]KJ02
       |          |--Zenobia pulverulentaKJ02
       |          `--AndromedaKJ02
       |               |--A. glaucophyllaKJ02
       |               |    |--A. g. f. glaucophyllaKJ02
       |               |    `--A. g. f. nanaKJ02
       |               `--A. polifoliaKJ02
       `--LyonieaeKJ02
            |--Craibiodendron yunnanenseKJ02
            |--+--AgaristaKJ02
            |  |    |--A. buxifoliaKJ02
            |  |    |--A. populifoliaKJ02
            |  |    `--A. salicifoliaKJ02
            |  `--PierisKJ02
            |       |--P. floribundaKJ02
            |       |--P. formosaKJ02
            |       |--P. japonicaKJ02
            |       |--P. nanaKJ02
            |       `--P. phillyraeifoliaKJ02
            `--Lyonia Nuttall 1818 (nom. cons.)KJ02, L95
                 |--L. buchiiJ87
                 |--L. ferrugineaKJ02
                 |--L. ligustrinaKJ02
                 |--L. lucidaKJ02
                 |--L. macrocalyx (Anthony) Airy Shaw 1937 [=Pieris macrocalyx Anthony 1927]P06
                 |--L. microcarpaJ87
                 |--L. nezikiBBO02
                 |--L. ovalifoliaP06
                 |--L. rubiginosa (Pers.) Don 1834 (see below for synonymy)L95
                 |    |--L. r. var. rubiginosaJ87
                 |    |--L. r. var. costataJ87
                 |    `--L. r. var. stahlii (Urb.) Judd 1981 [=L. stahlii Urb. 1908, Xolisma stahlii (Urb.) Small 1914]L95
                 |--L. sikkimensisP06
                 |--L. truncataJ87
                 |    |--L. t. var. truncataL95
                 |    `--L. t. var. proctorii Judd 1990L95
                 `--L. villosaP06

Lyonia rubiginosa (Pers.) Don 1834 [=Andromeda rubiginosa Pers. 1805, Xolisma rubiginosa (Pers.) Small 1914]L95

*Type species of generic name indicated

References

[BBO02] Begerow, D., R. Bauer & F. Oberwinkler. 2002. The Exobasidiales: an evolutionary hypothesis. Mycological Progress 1 (2): 187–199.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum—Biological Sciences 32 (1): 1–136.

[KJ02] Kron, K. A., W. S. Judd, P. F. Stevens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn & J. L. Luteyn. 2002. Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68: 335–423.

[L95] Liogier, H. A. 1995. Descriptive Flora of Puerto Rico and Adjacent Islands: Spermatophyta-Dicotyledoneae vol. 4. Melastomataceae to Lentibulariaceae. Editorial de la Universidad de Puerto Rico.

[P06] Panda, S. 2006. Lyonia macrocalyx (J. Anthony) Airy Shaw (Ericaceae)—a new record for India from Arunachal Pradesh and Nagaland. Bulletin of the Botanical Survey of India 48: 207–210.

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