Plants usually woody, or herbs, sometimes lacking chlorophyll. Leaves spiral, sometimes decussate, margin often toothed. Inflorescence racemose; bracteoles paired, basal. Flowers (4 or)5-merous. Calyx imbricate. Corolla connate, imbricate. Stamens 10, sometimes with spurs or awns, dehiscing by pores, pollen in tetrads, rarely single. Ovary superior or inferior, placentation axile, rarely parietal, often many ovules per locule. Style ± as long as corolla, slender. Fruit a capsule or berry, rarely a drupe; calyx persistent. Herbs, subshrubs, shrubs, or trees, (rarely vines), perennial, deciduous or evergreen, usually autotrophic, sometimes mycotrophic (subfam. Monotropoideae), usually chlorophyllous and autotrophic, sometimes achlorophyllous and heterotrophic (subfam. Monotropoideae), aromatic compounds (e.g., methyl salicylate) sometimes present (Gaultheria). Stems (absent in some Monotropoideae) erect or decumbent to prostrate, glabrous or hairy, (aerial stems sometimes produced from suckers, rhizomes, or corms), pith solid (hollow, with diaphragms in Agarista). Leaves (reduced or absent in some Monotropoideae), usually cauline, sometimes in basal rosettes (subfam. Monotropoideae), usually alternate or pseudoverticillate, sometimes opposite or, rarely, whorled, simple; stipules absent; petiole present or absent; blade plane or acicular, often coriaceous, margins entire or toothed, plane or revolute. Inflorescences terminal or axillary racemes, umbels, corymbs, panicles, fascicles, spikes, or solitary flowers. Flowers usually bisexual, rarely unisexual (subfam. Ericoideae), radially symmetric (sometimes slightly bilaterally symmetric in subfam. Monotropoideae and subfam. Ericoideae); perianth and androecium hypogynous (epigynous in some Vaccinioideae); hypanthium absent; sepals absent or (2-)4-5(-7), distinct or connate basally; petals (2-)4-5(-8), rarely absent or highly reduced, connate or distinct, not sticky (covered with sticky exudate in Bejaria), corolla absent or rotate to crateriform, campanulate, cylindric, globose, or urceolate (salverform in Epigaea); intrastaminal nectary disc present or absent; stamens (2-)5-8(-10) [14, 16, 20]; filaments distinct; anthers inverted during development, often with awns, dehiscent by pores or short slits (at apparent apex) or slits (lateral); pistils 1, 4-5-carpellate; ovary superior (inferior in some Vaccinioideae), incompletely (2-)5-10-locular (1-locular in some Monotropoideae), often furrowed or lobed externally; placentation axile or parietal; ovules anatropous, unitegmic, tenuinucellate; styles 1, straight or declinate (curved in Elliottia), hollow; stigmas 1, capitate or peltate to funnelform, usually 5-lobed. Fruits capsular and dehiscent (loculicidal, septifragal, or septicidal), or drupaceous (axis fibrous or soft in some Monotropoideae) or baccate (rarely each surrounded by accrescent or fleshy calyx in Gaultheria) and indehiscent. Seeds 1-10(-1000+), tan to yellowish brown or brown, ellipsoid, ovoid or spheroidal, or fusiform to flattened, or oblong (sometimes 3-sided); testa thin (bony in subfam. Arbutoideae and subfam. Vaccinioideae); embryo usually straight, fusiform, rarely minute and undifferentiated; endosperm abundant, cellular, fleshy. Fls regular or somewhat irregular, hypogynous to sometimes epigynous, mostly perfect and 4–5-merous; cor gamopetalous or much less often polypetalous, the lobes convolute or imbricate; stamens free from the cor or attached to its very base, as many or more often twice as many as the pet, rarely intermediate in number; anthers becoming inverted during ontogeny, so that the morphological base is apical, opening by pores at the apparent apex, or by more or less elongate slits extending downward from the apex, or rarely opening longitudinally their whole length, the apparent tip often drawn out into a pair of slender (sometimes connate) tubules (awns), and the body sometimes with 2 or 4 evident, slender appendages (spurs); pollen generally in tetrahedral tetrads; ovary superior to sometimes inferior, mostly 4–5-locular (8- or 10-locellar in Gaylussacia and some Vaccinium); placentation axile; ovules usually several or numerous in each locule (1 per locule in Arctostaphylos, 1 per locellus in Gaylussacia); fr capsular, often with a persistent axis, or sometimes a berry or drupe; seeds (5) ± numerous, sometimes winged, with copious endosperm and a straight (often very short) dicotyledonous embryo; seed-coat a single layer of cells; shrubs, small trees, or lianas, seldom almost herbaceous, with simple, exstipulate lvs (these often small and firm, with revolute margins, and then said to be ericoid, but also often quite ordinary) and small (often urceolate) to large fls typically in bracteate racemes or corymbs, but sometimes solitary and terminal or axillary. Genera 1–7 belong to the tribe Rhododendreae, 8–18 to the Andromedeae, 19 to the Arbuteae, 20 and 21 to the Ericeae, and 22 and 23 to the Vaccinieae. 125/3500. Calyx persistent; sepals free or united Flowers hermaphrodite, actinomorphic or slightly zygomorphic Shrubs or undershrubs, less frequently trees; leaves alternate or whorled, rarely opposite, simple, usually evergreen; stipules absent Fruit a capsule, berry, or drupe Ovary superior, several-celled, with numerous ovules on axile or rarely basal placentas, rarely 1-ovulate; style simple Stamens usually double the number of the corolla lobes or, if the same number, alternate with them, hypogynous; filaments and anthers usually free; anthers 2-celled, opening by pores or pore-like slits Corolla hypogynous, gamopetalous, inserted below a fleshy disk; lobes contorted or imbricate Seeds with fleshy endosperm and straight embryo, sometimes winged SELECTED REFERENCES Anderberg, A. A. 1993. Cladistic interrelationships and major clades of Ericaceae. Pl. Syst. Evol. 184: 207-231. Cipollini, M. L. and E. W. Stiles. 1992. Antifungal activity of ripe ericaceous fruits: Phenolic acid interactions and palatability for dispersers. Biochem. Syst. & Ecol. 20: 501-514. Harborne, J. B. and C. A. Williams. 1973. A chemotaxonomic survey of flavonoids and simple phenols in leaves of the Ericaceae. Bot. J. Linn. Soc. 66: 37-54. Judd, W. S. and K. A. Kron. 1993. Circumscription of Ericaceae (Ericales) as determined by preliminary cladistic analyses based on morphological, anatomical, and embryological features. Brittonia 45: 99-114. Kron, K. A. et al. 2002. Phylogenetic classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. (Lancaster) 68: 335-423. Luteyn, J. L. et al. 1996. Ericaceae of the southeastern United States. Castanea 61: 101-144. Small, J. K. 1914. Ericaceae. In: N. L. Britton et al., eds. 1905+. North American Flora.... 47+ vols. New York. Vol. 29, pp. 33-102. Stevens, P. F. 1969. Taxonomic Studies in the Ericaceae. Ph.D. thesis. University of Edinburgh. Stevens, P. F. 1970. Calluna, Cassiope and Harrimanella: A taxonomic and evolutionary problem. New Phytol. 69: 1131-1148. Stevens, P. F. 1971. A classification of the Ericaceae: Subfamilies and tribes. Bot. J. Linn. Soc. 64: 1-53. Stevens, P. F. et al. 2004. Ericaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 7+ vols. Berlin, etc. Vol. 6, pp. 145-194. Wood, C. E. Jr. 1961. The genera of Ericaceae in the southeastern United States. J. Arnold Arbor. 42: 10-80. Plants usually woody, or herbs, sometimes lacking chlorophyll. Leaves spiral, sometimes decussate, margin often toothed. Inflorescence racemose; bracteoles paired, basal. Flowers (4 or)5-merous. Calyx imbricate. Corolla connate, imbricate. Stamens 10, sometimes with spurs or awns, dehiscing by pores, pollen in tetrads, rarely single. Ovary superior or inferior, placentation axile, rarely parietal, often many ovules per locule. Style ± as long as corolla, slender. Fruit a capsule or berry, rarely a drupe; calyx persistent. Calyx persistent; sepals free or united Flowers hermaphrodite, actinomorphic or slightly zygomorphic Shrubs or undershrubs, less frequently trees; leaves alternate or whorled, rarely opposite, simple, usually evergreen; stipules absent Fruit a capsule, berry, or drupe Ovary superior, several-celled, with numerous ovules on axile or rarely basal placentas, rarely 1-ovulate; style simple Stamens usually double the number of the corolla lobes or, if the same number, alternate with them, hypogynous; filaments and anthers usually free; anthers 2-celled, opening by pores or pore-like slits Corolla hypogynous, gamopetalous, inserted below a fleshy disk; lobes contorted or imbricate Seeds with fleshy endosperm and straight embryo, sometimes winged Flowers hermaphrodite, actinomorphic or slightly zygomorphic Shrubs or undershrubs, less frequently trees; leaves alternate or whorled, rarely opposite, simple, usually evergreen; stipules absent Fruit a capsule, berry, or drupe Ovary superior, several-celled, with numerous ovules on axile or rarely basal placentas, rarely 1-ovulate; style simple Stamens usually double the number of the corolla lobes or, if the same number, alternate with them, hypogynous; filaments and anthers usually free; anthers 2-celled, opening by pores or pore-like slits Corolla hypogynous, gamopetalous, inserted below a fleshy disk; lobes contorted or imbricate Seeds with fleshy endosperm and straight embryo, sometimes winged Fls regular or somewhat irregular, hypogynous to sometimes epigynous, mostly perfect and 4–5-merous; cor gamopetalous or much less often polypetalous, the lobes convolute or imbricate; stamens free from the cor or attached to its very base, as many or more often twice as many as the pet, rarely intermediate in number; anthers becoming inverted during ontogeny, so that the morphological base is apical, opening by pores at the apparent apex, or by more or less elongate slits extending downward from the apex, or rarely opening longitudinally their whole length, the apparent tip often drawn out into a pair of slender (sometimes connate) tubules (awns), and the body sometimes with 2 or 4 evident, slender appendages (spurs); pollen generally in tetrahedral tetrads; ovary superior to sometimes inferior, mostly 4–5-locular (8- or 10-locellar in Gaylussacia and some Vaccinium); placentation axile; ovules usually several or numerous in each locule (1 per locule in Arctostaphylos, 1 per locellus in Gaylussacia); fr capsular, often with a persistent axis, or sometimes a berry or drupe; seeds (5) ± numerous, sometimes winged, with copious endosperm and a straight (often very short) dicotyledonous embryo; seed-coat a single layer of cells; shrubs, small trees, or lianas, seldom almost herbaceous, with simple, exstipulate lvs (these often small and firm, with revolute margins, and then said to be ericoid, but also often quite ordinary) and small (often urceolate) to large fls typically in bracteate racemes or corymbs, but sometimes solitary and terminal or axillary. Genera 1–7 belong to the tribe Rhododendreae, 8–18 to the Andromedeae, 19 to the Arbuteae, 20 and 21 to the Ericeae, and 22 and 23 to the Vaccinieae. 125/3500. SELECTED REFERENCES Anderberg, A. A. 1993. Cladistic interrelationships and major clades of Ericaceae. Pl. Syst. Evol. 184: 207-231. Cipollini, M. L. and E. W. Stiles. 1992. Antifungal activity of ripe ericaceous fruits: Phenolic acid interactions and palatability for dispersers. Biochem. Syst. & Ecol. 20: 501-514. Harborne, J. B. and C. A. Williams. 1973. A chemotaxonomic survey of flavonoids and simple phenols in leaves of the Ericaceae. Bot. J. Linn. Soc. 66: 37-54. Judd, W. S. and K. A. Kron. 1993. Circumscription of Ericaceae (Ericales) as determined by preliminary cladistic analyses based on morphological, anatomical, and embryological features. Brittonia 45: 99-114. Kron, K. A. et al. 2002. Phylogenetic classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. (Lancaster) 68: 335-423. Luteyn, J. L. et al. 1996. Ericaceae of the southeastern United States. Castanea 61: 101-144. Small, J. K. 1914. Ericaceae. In: N. L. Britton et al., eds. 1905+. North American Flora.... 47+ vols. New York. Vol. 29, pp. 33-102. Stevens, P. F. 1969. Taxonomic Studies in the Ericaceae. Ph.D. thesis. University of Edinburgh. Stevens, P. F. 1970. Calluna, Cassiope and Harrimanella: A taxonomic and evolutionary problem. New Phytol. 69: 1131-1148. Stevens, P. F. 1971. A classification of the Ericaceae: Subfamilies and tribes. Bot. J. Linn. Soc. 64: 1-53. Stevens, P. F. et al. 2004. Ericaceae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 7+ vols. Berlin, etc. Vol. 6, pp. 145-194. Wood, C. E. Jr. 1961. The genera of Ericaceae in the southeastern United States. J. Arnold Arbor. 42: 10-80. Herbs, subshrubs, shrubs, or trees, (rarely vines), perennial, deciduous or evergreen, usually autotrophic, sometimes mycotrophic (subfam. Monotropoideae), usually chlorophyllous and autotrophic, sometimes achlorophyllous and heterotrophic (subfam. Monotropoideae), aromatic compounds (e.g., methyl salicylate) sometimes present (Gaultheria). Stems (absent in some Monotropoideae) erect or decumbent to prostrate, glabrous or hairy, (aerial stems sometimes produced from suckers, rhizomes, or corms), pith solid (hollow, with diaphragms in Agarista). Leaves (reduced or absent in some Monotropoideae), usually cauline, sometimes in basal rosettes (subfam. Monotropoideae), usually alternate or pseudoverticillate, sometimes opposite or, rarely, whorled, simple; stipules absent; petiole present or absent; blade plane or acicular, often coriaceous, margins entire or toothed, plane or revolute. Inflorescences terminal or axillary racemes, umbels, corymbs, panicles, fascicles, spikes, or solitary flowers. Flowers usually bisexual, rarely unisexual (subfam. Ericoideae), radially symmetric (sometimes slightly bilaterally symmetric in subfam. Monotropoideae and subfam. Ericoideae); perianth and androecium hypogynous (epigynous in some Vaccinioideae); hypanthium absent; sepals absent or (2-)4-5(-7), distinct or connate basally; petals (2-)4-5(-8), rarely absent or highly reduced, connate or distinct, not sticky (covered with sticky exudate in Bejaria), corolla absent or rotate to crateriform, campanulate, cylindric, globose, or urceolate (salverform in Epigaea); intrastaminal nectary disc present or absent; stamens (2-)5-8(-10) [14, 16, 20]; filaments distinct; anthers inverted during development, often with awns, dehiscent by pores or short slits (at apparent apex) or slits (lateral); pistils 1, 4-5-carpellate; ovary superior (inferior in some Vaccinioideae), incompletely (2-)5-10-locular (1-locular in some Monotropoideae), often furrowed or lobed externally; placentation axile or parietal; ovules anatropous, unitegmic, tenuinucellate; styles 1, straight or declinate (curved in Elliottia), hollow; stigmas 1, capitate or peltate to funnelform, usually 5-lobed. Fruits capsular and dehiscent (loculicidal, septifragal, or septicidal), or drupaceous (axis fibrous or soft in some Monotropoideae) or baccate (rarely each surrounded by accrescent or fleshy calyx in Gaultheria) and indehiscent. Seeds 1-10(-1000+), tan to yellowish brown or brown, ellipsoid, ovoid or spheroidal, or fusiform to flattened, or oblong (sometimes 3-sided); testa thin (bony in subfam. Arbutoideae and subfam. Vaccinioideae); embryo usually straight, fusiform, rarely minute and undifferentiated; endosperm abundant, cellular, fleshy.General Information
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