Crowberry Empetrum nigrum, photographed by Ekaterina Smirnova.

Belongs within: Ericales.
Contains: Monotropoideae, Arbutoideae, Styphelioideae, Vaccinioideae, Ericeae, Phyllodoceae, Rhodoreae.

The Ericaceae is a cosmopolitan family of mostly cooler-climate plants (including tropical montane). Many species have small, needle-like leaves, referred to as 'ericoid'.

Characters (from Kron et al. 2002): Evergreen or deciduous shrubs, lianas, trees or herbs (these sometimes lacking chlorophyll), sometimes epiphytic. Leaves alternate, decussate, or whorled, sometimes distally clustered on shoots, entire to serrate, and often with margins strongly revolute and leaves needle-like; stipules absent; leaf epidermal cells lignified or not, hypodermal cells often present; stomata usually anomocytic or paracytic; vascular bundles frequently associated with fibre sheath; nodes usually unilacunar. Buds protected by two to numerous scales or naked. Indumentum a mixture of unicellular and multicellular hairs, or sometimes only unicellular-pubescent, the multicellular hairs extremely various, glandular or non-glandular, unbranched and short to elongate, variously branched, or peltate. Inflorescences terminal or axillary, usually indeterminate and often racemose, with variously developed bracts and usually a pair of bracteoles, these sometimes recaulescent, replacing calyx lobes; flowers pendulous to erectly held, articulated with pedicel or not. Flowers showy to inconspicuous, perfect to imperfect, usually tetra- or pentamerous, actinomorphic to zygomorphic. Sepals (1-)4-5(-9), usually connate, persistent or deciduous, occasionally colourful and fleshy; petals (3-)4-5(-9), usually connate and typically campanulate, cylindrical, or urceolate, occasionally rotate, deciduous or pesistent. Stamens (2-)5-10(-16), free from corolla or adnate, sometimes connate, included or exserted, filaments straight to variously curved, unicellular-pubescent or glabrous; anthers tetrasporangiate or bisporangiate, inverting (late, or more commonly early) in development, with two or four apparently terminal appendages (awns, spurs) or these sometimes on filaments, opening by pores or short to long slits, sometimes through narrowed tubules, endothecium usually absent. Pollen usually tricolpate to tricolporate, in tetrahedral tetrads, these sometimes reduced, or occasionally in monads, sometimes associated with viscin threads. Nectary present or absent. Ovary (2-)4-5(-14) carpellate, superior to inferior, with usually axile to intruded parietal placentations, rarely apical or basal; ovules one to many per carpel, anatropous to nearly campylotropous, unitegmic, tenuinucellate, embryo sac usually of Polygonum type; style short to long, hollow, sometimes expanded apically; stigma truncate to capitate or slightly to strongly lobed, sometimes cup-shaped, funnel-shaped, or flabellate to pinnatifid. Fruit loculicidal to septicidal capsules, berries, drupes (with one or several pits), or occasionally dry and indehiscent; seeds small, testa usually single layered, with cells isodiametric to strongly elongated, sometimes winged or tailed, sometimes mucilaginous on wetting; embryo fusiform or spatulate, white or less commonly green, with two cotyledons, to extremely reduced and undifferentiated; endosperm cellular, well developed, with haustoria at both ends; germination epigeal.

<==Ericaceae (see below for synonymy)
    |  i. s.: Sciadopityites G02
    |         Pernettyopsis megabracteata H03
    |         Oxycoccus palustris D03
    |--Enkianthus [Enkiantheae, Enkianthoideae] KJ02
    |    |--E. campanulatus KJ02
    |    |--E. deflexus SN88
    |    `--E. perulatus [incl. E. japonicus] LO98
    `--+--Monotropoideae KJ02
       `--+--Arbutoideae KJ02
          `--+--+--Harrimanella [Harrimanelloideae] KJ02
             |  |    `--H. hypnoides KJ02
             |  `--+--Styphelioideae KJ02
             |     `--Vaccinioideae KJ02
             `--+--Cassiope [Cassiopeae, Cassiopoideae] KJ02
                |    |--C. fastigiata O88
                |    |--C. mertensiana KJ02
                |    |--C. selaginoides O88
                |    `--C. tetragona KJ02
                `--Ericoideae [Ledeae, Ledoideae, Rhododendroideae, Rhodoroideae] KJ02
                     |  i. s.: Ledum palustre RKD02
                     |--Ericeae KJ02
                     |--Phyllodoceae KJ02
                     |--Rhodoreae KJ02
                     |--Bejarieae KJ02
                     |    |--Ledothamnus guyanensis KJ02
                     |    |--Bryanthus gmelini KJ02
                     |    `--Bejaria KJ02
                     |         |--B. racemosa KJ02
                     |         `--B. resinosa KJ02
                     `--Empetreae [Coremateae, Empetraceae, Empetroideae] KJ02
                          |--Corema conradi KJ02
                          |--Ceratiola ericoides KJ02
                          `--Empetrum KJ02
                               |--E. atropurpureum KJ02
                               |--E. hermaphroditum KJ02
                               |--E. nigrum KJ02
                               `--E. rubrum KJ02

Ericaceae [Andromedaceae, Arbutaceae, Arctostaphylaceae, Arctostaphyleae, Azaleaceae, Azaleoideae, Diplarchaceae, Ericineae, Hypopithydes, Hypopityaceae, Kalmiaceae, Kalmiae, Ledaceae, Menziesiaceae, Oxycoccaceae, Prionotaceae, Rhododendra, Rhododendraceae, Rhodoraceae, Salaxidaceae, Stypheliaceae, Vaccinia, Vacciniaceae]

*Type species of generic name indicated


[D03] Dus̩n, P. 1903. The vegetation of western Patagonia. In Reports of the Princeton University Expeditions to Patagonia, 1896-1899, vol. 8 РBotany (W. B. Scott, ed.) pp. 1-34. The University: Princeton (New Jersey).

[G02] Gomez, B. 2002. A new species of Mirovia (Coniferales, Miroviaceae) from the Lower Cretaceous of the Iberian Ranges (Spain). Cretaceous Research 23: 761-773.

[H03] Heads, M. 2003. Ericaceae in Malesia: Vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311-449.

[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.

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263-276.

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: An introductory note. In The Himalayan Plants vol. 1 (H. Ohba & S. B. Malla, eds) The University Museum, University of Tokyo, Bulletin 31: 19-46.

[RKD02] Relys, V., S. Koponen & D. Dapkus. 2002. Annual differences and species turnover in peat bog spider communities. Journal of Arachnology 30 (2): 416-424.

[SN88] Suzuki, M., & S. Noshiro. 1988. Wood structure of Himalayan plants. In The Himalayan Plants vol. 1 (H. Ohba & S. B. Malla, eds) The University Museum, University of Tokyo, Bulletin 31: 341-379.

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