Canellaceae

The Canellaceae are a family of flowering plants in the order Canellales.<ref name="judd2008">Walter S. Judd, Christopher S. Campbell, Elizabeth A. Kellogg, Peter F. Stevens, and Michael J. Donoghue. 2008. Plant Systematics: A Phylogenetic Approach, Third Edition. Sinauer Associates: Sunderland, MA, USA. Template:ISBN</ref> The order includes only one other family, the Winteraceae.<ref name="apweb">Template:Citation.</ref> Canellaceae is native to the Afrotropical and Neotropical realms. They are small to medium trees, rarely shrubs, evergreen and aromatic.<ref name="heywood2007">Vernon H. Heywood (with David J. Mabberley). 2007. "Canellaceae" page 84. In: Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. Flowering Plant Families of the World. Firefly Books: Ontario, Canada. (2007). Template:ISBN.</ref> The flowers and fruit are often red.

Several species of Canellaceae are important in herbal medicine or as a substitute for cinnamon, which is obtained from genus Cinnamomum in family Lauraceae. Canella winterana is the only species known in cultivation.<ref name="rhs">Anthony Huxley, Mark Griffiths, and Margot Levy (1992). The New Royal Horticultural Society Dictionary of Gardening. The Macmillan Press, Limited: London. The Stockton Press: New York. Template:ISBN (set).</ref>

The family is divided into five genera,<ref name="kubitzki1993">Klaus Kubitzki. 1993. "Canellaceae". pages 200-203. In: Klaus Kubitzki (editor); Jens G. Rohwer, and Volker Bittrich (volume editors). The Families and Genera of Vascular Plants volume II. Springer-Verlag: Berlin; Heidelberg, Germany / New York, US. Template:ISBN (Berlin) Template:ISBN (New York)</ref> but studies of DNA sequences have indicated one of these genera should be split.<ref name="salazar2008">Jackeline Salazar and Kevin Nixon. 2008. "New Discoveries in the Canellaceae in the Antilles: How Phylogeny Can Support Taxonomy". Botanical Review 74(1):103-111. {{#invoke:doi|main}}</ref> These genera together comprise about 25 species. In the Greater Antilles, many of these species are rare and restricted to small ranges. As of 2008, five of the species were newly recognized and not yet named.<ref name="salazar2008"/>Template:Update inline

Description

Some common properties include:<ref name="kubitzki1993"/><ref name="salazar2008"/><ref name="takhtajan2009">Armen L. Takhtajan (Takhtadzhian). Flowering Plants second edition (2009), pages xxxvi & 31. Springer Science+Business Media. Template:ISBN. Template:ISBN. {{#invoke:doi|main}} (see External links below)</ref>

• These trees, rarely shrubs, are evergreen and glabrous.
• The stems have nodes with three (rarely two) leaf gaps and three leaf traces. The xylem has narrow rays. The bark is aromatic, with prominent and unusual appearing lenticels.
• The leaves have a peppery taste, are alternate, spiral, or distichous in arrangement, simple, entire, coriaceous, petiolate, pinnately nerved, without stipules, with translucent (pellucid) glands. The parenchyma is without palisade layer in Pleodendron and Canella. The stomata are paracytic in American genera, and anomocytic in the Old World.
• The inflorescences are terminal or axillary, in a panicle (Canella) or a raceme; otherwise, the flowers are solitary (by reduction) and axillary.
• The flowers are actinomorphic, hypogynous, and usually trimerous. The receptacles are barely excavated, and the hypogynous disc is absent.
• The three (rarely 2) sepals are thick, coriaceous, and imbricate.
• The petals number (4-)5-12, in 1-2 (-4) unlike whorls or spirally arranged, slender, imbricate in bud, usually free (connate at the base in Canella and halfway to the apex in Cinnamosma).
• The androecium is monadelphous, adnate to the ovary. Stamens number 6-12, apparently derived from the fusion of two whorls in Warburgia and Canella. Anthers are extrorse and bithecal, with two sporangia per theca, attached to the outside of the staminal tube, and sessile; dehiscence is by a longitudinal slit, connective not projecting beyond thecae or only slightly so.
• The gynoecium is syncarpous. The ovary has two to six carpels, unilocular and superior. The style is short and thick; the stigma is apical and capitate, with two to six lobes. Placentation is parietal. Ovules number from two to many in one or two rows on each of the two to six placentas; they are hemianatropous to campylotropous, bitegmic, and crassinucellate.
• The fruit is a berry with a persistent calyx, with two or more seeds. Cinnamosma macrocarpa, in the Madagascan genus Cinnamosma, has the largest fruit in the family, sometimes reaching Template:Convert by Template:Convert.
• Seeds have exotestae (the outer layer of the testa) only; the tegmen (the inner layer of the testa) is collapsed. The seed coat has oily idioblasts; the endosperm is abundant and oily (ruminate in Cinnamosma). The embryo is small and straight to slightly curved, with two cotyledons.
• Pollen occurs in monads, and is delicate and monosulcate (usually with 10% of the grain trichotomosulcate); apertures are distal, exine, generally tectate, and granular, intectate, and reticulate in Cinnamosma; grains are small and hardly ornamented in Cinnamodendron and Warburgia, largest and most highly decorated in Canella and Pleodendron. The pollen is generally similar to that of the Myristicaceae, which had at one time caused some systematists to believe the two families were closely related.
• The chromosome number 2n is 22, 26, or 28.<ref name="ehrendorfer2000">Friedrich Ehrendorfer and Maria Lambrou. 2000. "Chromosomes of Takhtajania, other Winteraceae, and Canellaceae: phylogenetic implications". Annals of the Missouri Botanical Garden 87(3):407-413.</ref>

Synapomorphies for Canellaceae include monadelphous stamens, parietal placentation, and campylotropous ovules.<ref name="salazar2008"/>

Other notable traits include the conspicuous lenticels, the aromatic bark, the peppery taste of the leaves, the three (rarely two) fleshy sepals, and the berry with reniform seeds.<ref name="salazar2008"/>

Some sources indicate Cinnamodendron has 20-40 stamens, contrary to the sources that are regarded here as reliable. The very large stamen numbers (20 to 40), are probably counts of thecae or microsporangia.

Ecology

Canellaceae has species in both xeric and wet forests.

In Canella winterana, the flowers are protogynous. The berries are usually red, and probably eaten by birds, which contribute to seed dispersal (ornithochory). The trees are attacked by larvae of different insects, including dipterans.

Phytochemistry

File:Canella winterana Guadeloupe2.JPG

Fruit of Canella winterana, at Pointe des Châteaux, Guadeloupe, October 2008

File:Canella winteriana Guadeloupe.JPG

Flowers of C. winterana, at Pointe des Châteaux, Guadeloupe, October 2008

Monoterpenes are common, as are drimane-type sesquiterpenes, including cinnafragrins, cinnamodial, and capsicodendrin. These three sesquiterpenes are shared with only the Winteraceae in angiosperms. Canellaceae also have alkaloids of the aporphine type, such as N-(cinnamoyl)-tryptamine, lignans of the aryl-tetralin type, cinnamaldehydes, and allylphenols. Crystals of calcium oxalate are in the leaf mesophyll. Most species are cyanogenetic. Protocyanidins, flavonols, saponins, sapogenins, and ellagic acid are absent.Template:Citation needed

Uses

The saro, or green sandalwood, (also known locally as mandravasarotra), Cinnamosma fragrans, is native to Madagascar and is exported from there to India to be burned in ceremonies. It is not related to the true sandalwoods, which are in the family Santalaceae.Template:Citation needed

Most species of Canellaceae produce bark that is similar in odor and flavor to cinnamon, but they are closer related to the family Piperaceae including black pepper (Piper nigrum) than to true cinnamons, which are in the family Lauraceae (still within Magnoliids).

The white cinnamon, Canella winterana, a native of Florida and the Antilles, is used as a condiment, with tonic properties.Template:Citation needed

Commercial production of "white cinnamon" from C. winterana has ceased,<ref name="zanoni2004">Thomas A. Zanoni. 2004. "Canellaceae". page 81. In: Nathan Smith, Scott A. Mori, Andrew Henderson, Dennis Wm. Stevenson, and Scott W. Heald (editors). Flowering Plants of the Neotropics. Princeton University Press and The New York Botanical Garden. Template:ISBN.</ref> but small-scale, local production continues. The Canellaceae have long had local use as aromatic plants and as herbal medicines.

The bark of the red cinnamon or false Winter's bark, Cinnamodendron corticosum, is used as a substitute for Winter's bark (Drimys winteri, a member of Winteraceae) in Chile and Argentina, where it is called canelo, a name that is also applied to cinnamon. In Africa, several species of Warburgia have medicinal uses. The barks of Warburgia salutaris and Warburgia ugandensis are used to treat fevers, colds, and malaria.Template:Citation needed Other species are used for timber or in the production of resins used as glue.Template:Citation needed

Fossils

Fossil leaves of Canella are known from the Pliocene of Bahia (Brazil).Template:Citation needed Pollen of Pleodendron is known from the Oligocene of Puerto Rico.Template:Citation needed

Systematic position

Depending on the classification system and the characters considered, Canellaceae has been placed close to Annonaceae, Myristicaceae or Winteraceae.<ref name="heywood2007"/> In his last book, Armen Takhtajan defined the order Canellales as consisting of Canellaceae and Winteraceae.<ref name="takhtajan2009"/> This circumscription is followed in the APG IV system, in which the order Canellales is sister to another small order, the Piperales.<ref>Template:Cite journal</ref> These two orders combined with another two sister-orders Laurales and Magnoliales form together the clade Magnoliids.<ref>Template:Citation.</ref>

Included taxa

Theoretical introduction to Taxonomy

In this article, the genus Capsicodendron is maintained in synonymy with Cinnamodendron, although preliminary molecular phylogenetic studies separate Capsicodendron from Cinnamodendron and place Capsicodendron closer to Cinnamosma and Warburgia than to Cinnamodendron. This placement is not corroborated by morphology. The currently recognized genera in Canellaeae can be distinguished as follows:<ref name="salazar2008"/>

• Petals fused into a tube to the middle of their length

Cinnamosma Baill., 1867, Madagascar

• Petals free or slightly connate at the base

* Petals 5, slightly connate at the base, inflorescence a terminal panicle

Canella P. Browne, 1757, Florida, Antilles, northern South America

* Petals 6-12, free, flowers solitary, terminal or axillary, or in axillary inflorescences

* Petals 12, in 3-4 whorls, stamens 12, carpels 6

Pleodendron Tiegh., 1899, Greater Antilles, Costa Rica

* Petals 6-10, in two whorls, stamens 6-10, carpels 2-5(-6)

* Petals 6-10, stamens 6-10, carpels 2-4(-6), leaves elliptic to obovate, ripe fruit up to 2 cm in length

Cinnamodendron Endl., 1840 (including Capsicodendron Hoehne, 1933), Greater Antilles to southern Brazil

* Petals 10, stamens 10, carpels 5, leaves oblanceolate-spatulate to elongate, ripe fruit 3-6 cm long

Warburgia Engl., 1895, eastern and southern Africa

History

Canella winterana was an important medicinal plant of the natives of the American tropics, and it was soon adopted as such by the Europeans, as well. Dr. Diego Álvarez Chanca accompanied Christopher Columbus on his second voyage, after which he wrote of a cinnamon (canela in Spanish) which was unlike any of the species of cinnamon used in Europe.<ref name="dalby2001">Andrew Dalby. 2001. "Christopher Columbus, Gonzalo Pizarro, and the Search for Cinnamon". Gastronomica: The Journal of Food and Culture 1(2):40-49. (See external links below).</ref> He had probably reported the use of C. winterana.<ref name="salazar2008"/>

In 1737, in his Hortus Cliffortianus, Linnaeus combined Canella with Drimys, a genus now in Winteraceae, and Cinnamomum, now in Lauraceae, to form a taxon which he called Winterania.<ref name="linnaeus1737">Carolus Linnaeus. 1737. Hortus Cliffortianus:488. Lubrecht and Cramer. Amsterdam, Netherlands. (See External links below).</ref> In 1753, in the first edition of Species Plantarum, Linnaeus divided Winterania into four species.<ref name="linnaeus1753">Carolus Linnaeus (Carl von Linné). 1753. Species Plantarum, 1st edition, vol. 1, page 371. Holmiae: Impensis Laurentii Salvii (Lars Salvius). (A facsimile with an introduction by William T. Stearn was published by the Ray Society in 1957).</ref> Three of these are now in Cinnamomum, and the fourth, which he called Laurus winterana, consisted of what are now Canella winterana and Drimys winteri. These four species were included in a broadly defined Laurus.

In 1756, Patrick Browne applied the name Canella to the species now known as Canella winterana.<ref name="ipnicanella">Canella. In: International Plant Names Index.</ref> He did not add a specific epithet to create a binomial.<ref name="browne1756">Patrick Browne. 1756. The Civil and Natural History of Jamaica:275. T.Osborne & J. Shipton: London, UK.</ref> The generic name is derived from canela, the Spanish word for cinnamon, but the Spanish word is derived from the Latin canna, meaning "a reed", or from the related Greek kanna, which refers to a piece of rolled bark.<ref name="quattrocchiI">Umberto Quattrocchi. 2000. CRC World Dictionary of Plant Names volume I. CRC Press: Boca Raton; New York; Washington, DC, USA. / London, UK. Template:ISBN (vol. I). (see External links below).</ref>

The genus Canella was not adopted by Linnaeus, who resurrected Winterania in the second edition of Species Plantarum in 1762.<ref name="linnaeus1762">Carolus Linnaeus (Carl von Linné). 1762. Species Plantarum, 2nd edition, vol. 1, page 636. Holmiae: Impensis Laurentii Salvii (Lars Salvius).</ref> He assigned to Winterania a single species, Winterania canella, which was equivalent to the species he had previously called Laurus winterana.

In 1784, Johan Andreas Murray divided Winterania into two monospecific genera, the constituent species of which were Canella alba and Wintera aromatica.<ref name="murray1784">Johan Andreas Murray. 1784. pages 443 and 507. In: Caroli a Linné eqvitis Systema vegetabilivm : secvndvm classes ordines genera species cvm characteribvs et differentiis, 14th edition. Johann Christian Dieterich: Gottingen, Germany.</ref> The name Canella alba was validated by Murray in 1784,<ref name="ipnicanella"/> but it had long been in use. Linnaeus attributed the name to Samuel Dale, who used it in his Pharmacologia,<ref name="linnaeus1737"/> the first edition of which was published in 1693.<ref name="boulger_undated">George Simonds Boulger. date?. "Samuel Dale", entry 385. In: Dictionary of National Biography, 1885-1900, Volume 13.</ref> Patrick Browne mentions its use by Mark Catesby.<ref name="browne1756"/> Canella alba was renamed as Canella winterana by Joseph Gaertner in 1788 in his classic work De Fructibus et Seminibus Plantarum (The Fruits and Seeds of Plants).<ref name="gaertner1788">Joseph Gaertner. 1788. pages 373 and 374. In: De Fructibus et Seminibus Plantarum. Sumtibus Auctoris, Typis Academiae Carolinae. Stuttgart, Germany. (A facsimile edition was published by Nabu Press in 2010. Template:ISBN).</ref> The name change was required by the rules of botanical nomenclature. Wintera aromatica is now known as Drimys winteri and is in the family Winteraceae.

The family Canellaceae was established by Carl von Martius in 1832 and was defined as consisting of only the genus Canella.<ref name="reveal2008onward">James L. Reveal. 2008 onward. A checklist of suprageneric names for extant vascular plants. At: Home Page of James L. Reveal and C. Rose Broome. (See External links below).</ref><ref name="martius1832">Template:Cite book</ref> Stephan Endlicher divided Canella in 1840, creating the new genus Cinnamodendron. Cinnamosma was erected in 1867, Warburgia in 1895, and Pleodendron in 1899. Capsicodendron was erected in 1933. Some authors accept Capsicodendron and assign to it two species, Capsicodendron pimenteira and Capsicodendron dinisii.<ref name="zanoni2004"/> Other authors subsume Capsicodendron into Cinnamodendron and C. pimenteira into C. dinisii.<ref name="salazar2008"/>

Molecular phylogenetic studies of DNA sequences have shown Cinnamodendron, as traditionally circumscribed, is polyphyletic, consisting of two distinct groups.<ref name="salazar2008"/> These groups are morphologically different and their ranges do not overlap.

One of these groups is related to the African genera Cinnamosma and Warburgia, and might be paraphyletic over them. It consists of eight species, one of which was named in 2005.<ref name="hammel2005">Barry E. Hammel and Nelson A. Zamora. 2005. "Pleodendron costaricense (Canellaceae), a new species for Costa Rica". Lankesteriana 5(3):211-218.</ref> Two other species in this group have not been formally named and described in the scientific literature.<ref name="salazar2008"/> This group is restricted to South America. Since it includes the type species, Cinnamodendron axillare, it will retain the name Cinnamodendron.

The other group of Cinnamodendron species is most closely related to Pleodendron and is restricted to the Greater Antilles. It consists of six species, two of which remain unnamed.<ref name="salazar2008"/> The name Antillodendron has been proposed for this group, but this name is considered by some to be invalid because it was not effectively published.<ref name="tropicos_antillodendron">Antillodendron In: Tropicos At: Missouri Botanical Garden. (See External links below).</ref>

External links

Pictures

• Flowers of Cinnamosma madagascariensis
• Flowers of Canella winteranaTemplate:Dead link
• Warburgia salutaris in fruit
• Flowers of Cinnamodendron ekmanii

Words

• Neotropical Canellaceae Template:Ifsubst style="color:green">At: Family Index Template:Ifsubst style="color:green">At: Neotropikey Template:Ifsubst style="color:green">At: Projects and Programmes Template:Ifsubst style="color:green">At: Tropical America Project Template:Ifsubst style="color:green">At: Kew in depth Template:Ifsubst style="color:green">At: Scientific Research and Data Template:Ifsubst style="color:green">At: Kew Gardens
• List of Genera in Canellaceae Template:Ifsubst style="color:green">At: Dicotyledons Template:Ifsubst style="color:green">At: List Genera within a Family Template:Ifsubst style="color:green">At: Vascular Plant Families and Genera Template:Webarchive Template:Ifsubst style="color:green">At: About the Checklist Template:Ifsubst style="color:green">At: World Checklist of Selected Plant Families Template:Ifsubst style="color:green">At: Data Sources Template:Webarchive Template:Ifsubst style="color:green">At: ePIC Template:Webarchive Template:Ifsubst style="color:green">At: Scientific Databases Template:Ifsubst style="color:green">At: Kew Gardens
• List of Genera in Canellaceae Template:Ifsubst style="color:green">At: Canellaceae Template:Ifsubst style="color:green">At: List of families Template:Ifsubst style="color:green">At: Families and Genera in GRIN Template:Ifsubst style="color:green">At: Queries Template:Ifsubst style="color:green">At: GRIN taxonomy for plants
• Canella winterana Template:Ifsubst style="color:green">At: Canella Template:Ifsubst style="color:green">At: Canellaceae Template:Ifsubst style="color:green">At: Canellales Template:Ifsubst style="color:green">At: Magnoliids Template:Ifsubst style="color:green">At: Magnoliophyta (flowering plants) ... Template:Ifsubst style="color:violet">In: ··· Embryophyta Template:Ifsubst style="color:green">At: Streptophytina Template:Ifsubst style="color:green">At: Streptophyta Template:Ifsubst style="color:green">At: Viridiplantae Template:Ifsubst style="color:green">At: Eukaryota Template:Ifsubst style="color:green">At: Taxonomy Template:Ifsubst style="color:green">At: UniProt
• Canellaceae Template:Ifsubst style="color:green">In: Volume 3 Template:Ifsubst style="color:green">At: Family List Template:Ifsubst style="color:green">At: FNA (Flora of North America) Template:Ifsubst style="color:green">At: eFloras
• Canellaceae Template:Ifsubst style="color:green">In: Leslie Watson and Michael J. Dallwitz (1992 onwards), The families of flowering plants
• NCBI Taxonomy Browser: Cannellaceae
• Distribution Map Template:Ifsubst style="color:green">And Genus List Template:Ifsubst style="color:green">At: Canellaceae Template:Ifsubst style="color:green">At: Canellales Template:Ifsubst style="color:green">At: Trees Template:Ifsubst style="color:green">At: APweb Template:Ifsubst style="color:green">At: botanical databases Template:Ifsubst style="color:green">At: About Science & Conservation Template:Ifsubst style="color:green">At: Missouri Botanical Garden
• Canellaceae in BoDD – Botanical Dermatology Database
• page xxxvii Template:Ifsubst style="color:green">In: Flowering Plants (Takhtajan)
• CRC World Dictionary of Plant Names: A-C At: Botany & Plant Science At: Life Science At: CRC Press
• The search for cinnamon Template:Ifsubst style="color:green">In: Gastronomica
• page 488 Template:Ifsubst style="color:green">In: Hortus Cliffortianus Template:Ifsubst style="color:green">At: View Record for title 2450 Template:Ifsubst style="color:green">At: Titles/H Template:Ifsubst style="color:green">At: Titles Template:Ifsubst style="color:green">At: Biodiversity Heritage Library
• page 371 Template:Ifsubst style="color:green">In: Species Plantarum, 1st ed. (1753) At: View Record of title 25 At: Titles by Carl von Linné (1707-1778) Template:Ifsubst style="color:green">At: Authors / L Template:Ifsubst style="color:green">At: Authors Template:Ifsubst style="color:green">At: Biodiversity Heritage Library
• Canella Template:Ifsubst style="color:green">At: Plant Names Template:Ifsubst style="color:green">At: IPNI
• Canella Template:Ifsubst style="color:green">In: The Civil and Natural History of Jamaica Template:Ifsubst style="color:green">At: Patrick Browne Template:Ifsubst style="color:green">At: Authors Template:Ifsubst style="color:green">At: Botanicus
• Samuel Dale Template:Ifsubst style="color:green">In: Dictionary of National Biography, 1885-1900 Template:Ifsubst style="color:green">At: Wikisource
• page 636 Template:Ifsubst style="color:green">In: Species Plantarum 2nd edition, volume 1 Template:Ifsubst style="color:green">At: View Record for title 26 Template:Ifsubst style="color:green">At: Titles by Carl von Linné (1707-1778) Template:Ifsubst style="color:green">At: Authors / L Template:Ifsubst style="color:green">At: Authors Template:Ifsubst style="color:green">At: Biodiversity Heritage Library
• Canella on page 443 Template:Ifsubst style="color:green">And Wintera on page 507 Template:Ifsubst style="color:green">In: Systema Vegetabilium (1784) Template:Ifsubst style="color:green">At: View Record Template:Ifsubst style="color:green">At: Titles by Johann Andreas Murray Template:Ifsubst style="color:green">Or Titles by Johan Anders Murray Template:Ifsubst style="color:green">At: Authors/M Template:Ifsubst style="color:green">At: Authors Template:Ifsubst style="color:green">At: Biodiversity Heritage Library
• page 373 Template:Ifsubst style="color:green">And page 374 Template:Ifsubst style="color:green">In: De Fructibus et Seminibus Plantarum Template:Ifsubst style="color:green">At: Joseph Gaertner Template:Ifsubst style="color:green">At: Author List Template:Ifsubst style="color:green">At: Botanicus Digital Library
• Families CA-CL Template:Ifsubst style="color:green">In: Indices Nominum Supragenericorum Plantarum Vascularium Template:Ifsubst style="color:green">At: Home Page of James L. Reveal and C. Rose Broome
• page 168 Template:Ifsubst style="color:green">Of Nova genera et species plantarum, vol. 3 Template:Ifsubst style="color:green">At: View Record for title 12 Template:Ifsubst style="color:green">At: Titles by Karl Friedrich Philipp von Martius Template:Ifsubst style="color:green">At: Authors Template:Ifsubst style="color:green">At: Biodiversity Heritage Library
• Antillodendron (search exact) Template:Ifsubst style="color:green">At: Name Search Template:Ifsubst style="color:green">At: Tropicos Template:Ifsubst style="color:green">At: About Science and Conservation Template:Ifsubst style="color:green">At: Missouri Botanical Garden