Ericaceae Durande

Pedraza, P. (2009). Neotropical Ericaceae.

Morphology

Description

Terrestrial or epiphytic shrubs, subshrubs, perennial herbs, or fleshy achlorophyllous mycotrophs, sometimes lianoid, rarely trees, sometimes rhizomatous or stoloniferous, commonly producing tannins, frequently with hypocotyl maturing as lignotuber up to 1 m diam.;  indumentum of uni- to multicellular hairs or scales, these sometimes glandular . Stems terete or subterete, sometimes conspicuously winged . Leaves alternate , rarely opposite, verticillate , whorled , or lacking and then replaced by bract -like scales, simple , usually petiolate , exstipulate , newly unfolding leaves often a conspicuous red colour;  lamina coriaceous to membranous , evergreen to deciduous , the margin entire or sometimes serrulate - crenate , the venation pinnate or plinerved; leaf scars usually with a single vascular bundle scar, nodes usually with one trace and one gap. Inflorescence axillary or rarely terminal , sometimes long-pendent, racemose, paniculate, fasciculate , or flowers solitary, the parts often viscid or secretory in bud ; individual flowers pedicellate or rarely sessile in axils of caducous to deciduous or persistent floral bracts, these sometimes conspicuously glandular ;  pedicel usually bibracteolate; bracteoles deciduous or persistent , small or large. Flowers mostly hermaphrodite , but rarely functionally unisexual (more rarely plants dioecious ), actinomorphic or slightly zygomorphic , mostly (3-)5(-7)- merous , typically obdiplostemonous, hypogynous or epigynous and with a typically biseriate perianth , typically without floral odors, rarely with extrafloral nectaries;  calyx aestivation valvate , imbricate , or reduplicate, continuous or articulate with  pedicel , synsepalous, rarely fleshy and accrescent to  fruit ,  calyx tube terete or angled to winged ;  corolla membranous to thick-carnose, polypetalous or more commonly sympetalous, cylindric, campanulate or urceolate, terete or angled to winged opposite lobes, sometimes basally gibbous;  stamen (5-)8-10(-14), in 2 whorls, usually twice as many as petals or rarely just as many, equal with each other or alternately unequal, borne on edge of an obscure to prominent nectariferous disc ; filaments equal or unequal, usually straight or rarely S-shaped (geniculate), sometimes also basally papillose , distinct or connate , shorter or longer than  anther ; anther inverting during development, 2-celled, equal or unequal, often distally with 2 distinct or connate tubules or terminal awns, sometimes provided with abaxial spurs; white, powdery, disintegration tissue present or lacking; thecae smooth to coarsely granular , with or without a basal appendage ; tubules when present conical and rigid or cylindric and flexible, of equal or ca. 1/2 the diameter of thecae, longer to shorter than thecae; dehiscence normally introrse, but rarely extrorse or latrorse, by longitudinal slits or more typically by apical to subapical pores;  pollen grains in tetrahedral tetrads or rarely single, sometimes with viscin threads;  pistil single;  ovary superior or inferior, 4-5(-10)-carpellate, usually with as many locules as carpels or with twice as many locules as carpels or rarely loculate in lower portion and 1-locular above; placentation axile , rarely intruded parietal ; ovules numerous per locule or rarely solitary, anatropous to campylotropous with a single integumentary layer;  style single, fluted , hollow;  stigma simple but occasionally weakly lobed . Fruit a loculicidal or septicidal capsule , berry , or drupe , with a usually persistent , rarely with accrescent and fleshy calyx ; seeds small, ca. 1-1.5 mm long, usually numerous (1 per locule in Gaylussacia Kunth), winged or tailed (only in Bejaria Mutis ex L.), sometimes enclosed in a mucilaginous sheath , testa thin with elongated or isodiametric cells,  endosperm fleshy , embryo straight, usually white or sometimes green.

Distribution

Distribution in the Neotropics

• Found throughout the Neotropics in montane regions, mostly (but not exclusively) in moist and cool environments.
• The highest number of species is concentrated in the northern Andes above 1,000 m.

Diagnostic

Distinguishing characters (always present)

• In the Neotropics Ericaceae have evolved different life-forms and may occupy many different habitats.
• The species of this family are sometimes quite plastic and many shift between terrestrial and epiphytichabit. However, in the Neotropics Ericaceae grows continuously throughout the year in a seasonal habitats, producing flushes of intensely red-pigmented new leaves . This makes the family very easy to spot in the field.
• The leaves of many Ericaceae are characteristically coriaceous, sometimes "ericoid", and without stipules.
• The great majority of the species have flowers with fused petals; the corollas vary in size, shape, and colors (white, pink, red, yellow, green), but some speciose genera have flowers consistently brightly colored or with contrasting colour patterns (see examples in Cavendishia Lindl.).
• The stamens of Ericaceae are very characteristic because of the presence of different staminal appendages (spurs, awns and tubules), either on the dorsal surface of the anther/filament or at the functional apex of the anther.

Useful tips for generic identification

A key to the genera of Neotropical Ericaceae is available at http://www.nybg.org/bsci/res/lut2/familydescription.html#keys

General Description

Number of genera

46 genera (for more details see http://www.nybg.org/bsci/res/lut2/main.html):

• Agarista D.Don
• Anthopteropsis A.C.Sm.
• Anthopterus Hook.
• Arbutus L.
• Arctostaphylos Adans.
• Bejaria
• Cavendishia
• Ceratostema Juss.
• Chimaphila Pursh
• Comarostaphylis Zucc.
• Demosthenesia A.C.Sm.
• Didonica Luteyn & Wilbur
• Diogenesia Sleumer
• Disterigma ( Klotzsch ) Nied.
• Gaultheria Kalm ex L.
• Gaylussacia
• Gonocalyx Planch. & Linden
• Kalmia L.
• Lateropora A.C.Sm.
• Ledothamnus Meisn.
• Lyonia Nutt.
• Macleania Hook.
• Monotropa L.
• Mycerinus A.C.Sm.
• Notopora Hook.f.
• Oreanthes Benth.
• Orthaea Klotzsch
• Orthilia Raf.
• Pellegrinia Sleumer
• Pernettya Gaudich.
• Pieris D.Don
• Plutarchia A.C.Sm.
• Polyclita A.C.Sm.
• Psammisia Klotzsch
• Pterospora Nutt.
• Pyrola L.
• Rhododendron L.
• Rusbya Britton
• Satyria Klotzsch
• Semiramisia Klotzsch
• Siphonandra Klotzsch
• Sphyrospermum Poepp. & Endl.
• Tepuia Camp
• Themistoclesia Klotzsch
• Thibaudia Ruiz & Pav.
• Utleya Wilbur & Luteyn
• Vaccinium L.

Status

• Over 800 species native to the Neotropics and ca. 94% of them are endemic.
• Few native species of Cavendishia are locally cultivated in the northern Andes.
• Rhododendronsimsii Planch., endemic to China, occurs as a cultivar and rarely as an escape in the Neotropics.

Literature

Important literature

(a complete bibliography can be found at http://www.nybg.org/bsci/res/lut2/main.html)

Kron, K. A., W. S. Judd, P. F. Stevens, D. M. Crayn, A. A. Anderberg, P. A. Gadek, C. J. Quinn & J. L. Luteyn. 2002a. Phylogenetic classification of Ericaceae: molecular and morphological evidence. The Botanical Review 68(3): 335-423.

Kron, K. A., E. A. Powell & J. L. Luteyn. 2002. Phylogenetic relationships within the blueberry tribe(Vaccinieae, Ericaceae) based on sequence data from matK and nuclear ribosomal ITS regions, with comments on the placement of Satyria. American Journal of Botany 89(2): 327-336.

Luteyn, J. L. 2002. Diversity, adaptation, and endemism in neotropical Ericaceae: Biogeographical patterns in the Vaccinieae. The Botanical Review 68(1): 55-87.

Luteyn, J. L. & P. Pedraza-Peñalosa. 2006. Neotropical Blueberries: The Plant Family Ericaceae. http://www.nybg.org/bsci/res/lut2. [continuously updated].

Luteyn, J. L. & P. Pedraza-Peñalosa. 2007. Rapid color guide to the Ericaceae of the National Park Machu Picchu. The Field Museum at http://fm2.fieldmuseum.org/plantguides/.

Luteyn, J. L. & P. Pedraza-Peñalosa. 2007. Rapid color guide to the Ericaceae of Bolivia. The Field Museum at http://fm2.fieldmuseum.org/plantguides/.

Luteyn, J. L. & R. L. Wilbur. 2005. Flora Costaricensis. Fieldiana, Botany, new series 45: 1-107.

Stevens, P. F., J. L. Luteyn, E. G. H. Oliver, T. L. Bell, E. A. Brown, R. K. Crowden, A. S. George, G. J. Jordan, P. Ladd, K. Lemson, C. B. McLean, Y. Menadue, J. S. Pate, H. M. Stace & C. M. Weiller. 2004. Ericaceae. Pp. 145-194. In: Kubitzki, K., (ed.) The families and genera of vascular plants VI. Flowering Plants.Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales. Springer-Verlag, Berlin.

Ericaceae, R. Ross. Flora Zambesiaca 7:1. 1983

Morphology General Habit

Small trees, shrubs or sub–shrubs, never herbs

Morphology Leaves

Leaves simple, alternate, opposite or whorled, exstipulate, usually evergreen

Morphology Reproductive morphology Flowers

Flowers bisexual, actinomorphic or slightly zygomorphic, in the axils of the leaves, in terminal umbels, or on simple racemes which may be terminal or axillary or both

Morphology Reproductive morphology Flowers Calyx

Calyx and corolla normally 4–merous or 5–merous, occasionally 3–merous Sepals free or fused, normally equal but in a few genera one sepal larger than the others

Morphology Reproductive morphology Flowers Corolla

Corolla normally gamopetalous, 3–5–lobed, lobes contorted or imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens normally twice as many as the petals, occasionally equal in number to them and alternating with them, or, more rarely, intermediate in number, insertedd on a hypogynous or epigynous disc; filaments normally free, occasionally adnate to the corolla at their base; anthers innate or versatile, opening by terminal or lateral pores, often with appendages

Morphology Reproductive morphology Flowers Gynoecium

Carpels normally equal in number to the petals, occasionally one less, or all but one aborted, united to form a normally 3–5–locular, occasionally 1–locular, ovary, which is normally superior but is inferior in some genera; seeds numerous in most genera, 1 or 2 per loculus in a few, placentation axile or basal

Morphology Reproductive morphology Fruits

Fruit a capsule, usually loculicidal, a drupe or a berry

Ericaceae, R. Ross. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Shrubs or undershrubs, less frequently trees; leaves alternate or whorled, rarely opposite, simple, usually evergreen; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic or slightly zygomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx persistent; sepals free or united

Morphology Reproductive morphology Flowers Corolla

Corolla hypogynous, gamopetalous, inserted below a fleshy disk; lobes contorted or imbricate

Morphology Reproductive morphology Flowers Androecium

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

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, several-celled, with numerous ovules on axile or rarely basal placentas, rarely 1-ovulate; style simple

Morphology Reproductive morphology Fruits

Fruit a capsule, berry, or drupe

Morphology Reproductive morphology Seeds

Seeds with fleshy endosperm and straight embryo, sometimes winged