Neotropical Winteraceae

Jon L.R. Every

Royal Botanic Gardens, Kew, UK. 

Description

Evergreen trees or shrubs, glabrous, commonly with suckers, fresh bark and leaves aromatic with a burning, pepper-like taste. Leaves alternate, spiral, commonly irregularly crowded, exstipulate, margins entire, simple, often coriaceous, glaucous-white below with wax plugging stomata, often finely pellucid-dotted, sometimes very strongly curved adaxially (supervolute), venation pinnate, brochidodromous, tertiaries reticulate (NB! there is considerable polymorphism and variation in both foliage and floral characters). Inflorescence terminal, intercalary or axillary, cymose, in umbellate clusters or solitary. Flowers bisexual, hypogynous, bracts or bracteoles caducous; sepals 2-4(-6), connate in bud, forming a calyptra, rupturing early at anthesis, sometimes glandular; petals (0-)2-many, in 1-3 whorls, outer members sometimes, nearly always free, imbricate; stamens spirally arranged, 3-many, filaments stout, fleshy, anthers basifixed, opening by longitudinal slits; ovary superior, (1-)5-12-many, seemingly apocarpic but can become syncarpous, unilocular; style minute to absent, stigma bilobed (not decurrent), one per carpel. Fruit a cluster of follicles or berries, indehiscent. Seed few to numerous, smooth to blistered, testa dark grey to black, hard, brittle, endosperm copious.

Notes on delimitation

  • Together with the Canellaceae in the order Canellales (APG 2, 2003).
  • The Canellales are found in the Magnoliids and are most likely sister to the Piperales (APG 2, 2003).
  • Molecular and floral ontogeny studies places Drimys J.R. Forst. & G. Forst. as sister to Tasmannia R.Br. ex DC. and not in the same genus split into two sections (Doust, 2004). Tasmannia is an Old World genus of c. 40 spp. with unisexual flowers.

Distribution in the Neotropics

  • Winteraceae generally grow in moist montane tropical environments.
  • From Veracruz in central-eastern Mexico, throughout the Neotropics to the southernmost point of South America.
  • D. granatensis Mutis ex L.f. is found in cloud forests throughout the Neotropics at 3,000-3,500 m alt.
  • D. roraimensis (A.C. Sm.) Ehrend & Gottsb. is found in the Venezuelan montane and cloud forests, dwarf forests, slopes of tepuis, along streams and in swamps, at 1,800-2,500 m alt.
  • D. granandensis L. occurs from southern Mexico through to Peru and Chile.
  • D. winteri J.R. Forst. & G. Forst. occurs in Argentina, Chile and Peru.

Distinguishing characters (always present)

  • Alternate, simple leaves arranged spirally and somewhat irregularly clustered.
  • Leaf blade with secondary veins that do not extend to the leaf margin but link up with the neighbouring one.
  • Underside of leaf waxy and seemingly white/blue in colour.
  • During or after flowering the twig elongates monopodially from the apical bud of the inflorescence.
  • Flowers bisexual with the sepals forming a cap-like structure which covers the stamens and carpels in bud.
  • Stamens spirally arranged.
  • Superior ovary.
  • Fruits indehiscent.

Other important characters

  • Fresh material has a distinct smell with a burning, pepper-like taste.
  • Transparent gland dots are regularly visible on the leaf blades.
  • Partial inflorescence of 2-21, sweetly scented predominantly white flowers.
  • Bracts and bracteoles are rapidly deciduous.
  • Gynoecium can be apocarpic or fused in the middle and becoming syncarpous.
  • Fruit in clusters of follicles or berries.

Key differences from similar families

Canellaceae have:

  • 3 sepals.
  • 5-12 petals.
  • 6-numerous stamens fused into a tube.
  • 2-6 connate carpels.

Number of genera

  • One: Drimys J.R.Forst. & G.Forst.

Status

  • Drimys is endemic to the Americas.

General notes

  • Due to the basal position of Winteraceae in the angiosperm phylogenetic tree the flower morphology is variable and not as fixed as more recently evolved families.
  • There is considerable polymorphism and variation in both foliage and floral characters. This instability can lead to difficulty in determining the species of the specimen.

Important literature

APG 2. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Botanical Journal of the Linnean Society. 141. pp. 399-436.

Doust, A. N. & Drinnan, A. N. 2004. Floral development and molecular phylogeny support the generic status of Tasmannia (Winteraceae) Amer. J. Bot. 91 (3): 321-331.

Frame, D. 2004. Winteraceae. In: Smith, N., Mori, S. A., Henderson, A., Stevenson, D. W. and Heald, S. V. (eds). Flowering Plants of the Neotropics. pp.398-9. The New York Botanical Garden, Princeton University Press, Princeton.

Heywood, V.H., R.K. Brummitt, A. Culham & O. Seberg (eds). 2007. Flowering plant families of the world, pp.336-7. Royal Botanic Gardens, Kew.

Judd, W. S., Campbell, C. S., Kellogg, E. A., Stevens, P. F., Donoghue, M. J. 2008. Plant Systematics: a phylogenetic approach 3rd ed. pp.244-5. Sinauer Associates, Massachusetts.

Maas, P. J. M. & Westra, L. Y. Th. 2005. Neotropical Plant Families. 3rd ed. pp. 116-7. A.R.G. Gantner Verlag K.G., Ruggell.

Miller, J. S. 2005. In: Steyermark, J. A., Berry, P. E., Yatskievych, K. & Holst, B. K. (eds). Flora of the Venezuelan Guayana. Volume. 9 Rutaceae -Zygophyllaceae. pp 524-5. Missouri Botanical Garden, St. Louis.

Pennington, T.D. 2004. Illustrated guide to the trees of Peru, p.162. David Hunt, Milbourne Port, Sherborne.

Stevens, P. F. 2008. Angiosperm Phylogeny Website. Version 9 onwards. http://www.mobot.org/MOBOT/research/APweb/.

Vink, W. 1993. Winteraceae. In: Kubitzki, K. Rohwer, J. G. & Bittrich, V. (eds), Families and Genera of Vascular Plants. Volume 2. Flowering plants. Dicotyledons. pp.630-638. Springer-Verlag Berlin.

Watson, L. & Dallwitz, M. J. (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000.

How to cite

Every, J.L.R. (2009). Neotropical Winteraceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Winteraceae.htm.