Ericaceae

 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’. However, the leaves are flat and fairly large in the genus Enkianthus, a group of shrubs found in eastern Asia that is placed by phylogenetic analysis as the sister group of other Ericaceae (Kron et al. 2002).

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)
    |--Enkianthus [Enkiantheae, Enkianthoideae]KJ02
    |    |--E. campanulatusKJ02
    |    |--E. deflexusSN88
    |    `--E. perulatus [incl. E. japonicus]LO98
    `--+--MonotropoideaeKJ02
       `--+--ArbutoideaeKJ02
          `--+--+--Harrimanella [Harrimanelloideae]KJ02
             |  |    `--H. hypnoidesKJ02
             |  `--+--StyphelioideaeKJ02
             |     `--VaccinioideaeKJ02
             `--+--Cassiope [Cassiopeae, Cassiopoideae]KJ02
                |    |--C. fastigiataO88
                |    |--C. mertensiana [incl. C. mertensiana ssp. californica, C. mertensiana ssp. ciliolata]H93
                |    |--C. selaginoidesO88
                |    `--C. tetragonaKJ02
                `--Ericoideae [Ledeae, Ledoideae, Rhododendroideae, Rhodoroideae]KJ02
                     |  i. s.: LedumH93
                     |           |--L. glandulosum (see below for synonymy)H93
                     |           |--L. groenlandicumB-PW00
                     |           `--L. palustreRKD02
                     |--EriceaeKJ02
                     |--PhyllodoceaeKJ02
                     |--RhodoreaeKJ02
                     |--BejarieaeKJ02
                     |    |--Ledothamnus guyanensisKJ02
                     |    |--Bryanthus gmeliniKJ02
                     |    `--BejariaKJ02
                     |         |--B. racemosaKJ02
                     |         `--B. resinosaKJ02
                     `--Empetreae [Coremateae, Empetraceae, Empetroideae]KJ02
                          |--Corema conradiKJ02
                          |--Ceratiola ericoidesKJ02
                          `--EmpetrumKJ02
                               |--E. atropurpureumKJ02
                               |--E. hermaphroditumKJ02 [=E. nigrum ssp. hermaphroditumH93]
                               |--E. nigrumKJ02
                               `--E. rubrumKJ02
Ericaceae incertae sedis:
  SciadopityitesG02
  Pernettyopsis megabracteataH03
  Oxycoccus palustrisD03
  Bruckenthalia spiculifoliaJK80
  Vaccinioides Kvaček & Walther 1990HL08
    `--V. echitiformis (Rüffle, Müller-Stoll & Litke) Kvaček, Walther & Wilde in Kvaček & Walther 1990 (see below for synonymy)HL08
  Xylococcus bicolorH93
  Kalmiophyllum marcodurense Kräusel & Weyland 1959CBH93
  ErecipitesCBH93
  Philippia johnstoniiPB27
  Waldemaria Klotzsch 1862KC01
  EriciphyllumP92
  OrphanidesitesP92
  Corallobotrys acuminataDD73

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

Ledum glandulosum [incl. L. glandulosum var. australe, L. glandulosum var. californicum, L. glandulosum var. columbianum, Rhododendron neoglandulosum, L. glandulosum ssp. olivaceum]H93

Vaccinioides echitiformis (Rüffle, Müller-Stoll & Litke) Kvaček, Walther & Wilde in Kvaček & Walther 1990 [=Apocynophyllum echitiforme Rüffle, Müller-Stoll & Litke 1976]HL08

*Type species of generic name indicated

References

[B-PW00] Behan-Pelletier, V., & D. E. Walter. 2000. Biodiversity of oribatid mites (Acari: Oribatida) in tree canopies and litter. In: Coleman, D. C., & P. F. Hendrix (eds) Invertebrates as Webmasters in Ecosystems pp. 187–202. CABI Publishing: Wallingford (UK).

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[DD73] Deb, D. B., & R. M. Dutta. 1973. Contribution to the flora of Tirap Frontier Division. Journal of the Bombay Natural History Society 69 (3): 547–573.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8. Botany 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.

[HL08] Henniger, M., & R. Leder. 2008. Eozäne Makroflora der Querfurter Mulde. Mauritiana 20 (2): 229–251.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[JK80] John, J., & K.-P. Kolbe. 1980. The systematic position of the “Theales” from the viewpoint of serology. Biochemical Systematics and Ecology 8: 241–248.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[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: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[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: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 341–379.

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