Bot. Rev. (2008) 74:103–111
DOI 10.1007/s12229-008-9002-z
New Discoveries in the Canellaceae in the Antilles:
How Phylogeny can Support Taxonomy
Jackeline Salazar1,2,3 & Kevin Nixon2
1
L.H. Bailey Hortorium, Department of Plant Biology, Cornell University, Ithaca, NY 14853, USA
Departamento de Biología, Universidad Autónoma de Santo Domingo, Zona Universitaria,
Santo Domingo, Dominican Republic
3
Author for Correspondence; e-mail: jackeline_salazar@hotmail.com
2
Published online: 22 April 2008
# The New York Botanical Garden 2008
Abstract Six genera have been described in the family Canellaceae, four of them
from the Neotropics and the other two from Africa and Madagascar. The
Caribbean genera are Canella, Pleodendron and Cinnamodendron. Canella is a
monotypic genus widespread in the region, and Pleodendron is present in the
Greater Antilles and Costa Rica. Cinnamodendron occurs in the Greater Antilles
(Cuba, Hispaniola, and Jamaica) as well as in South America. A recent phylogenetic
analysis of the family shows that Cinnamodendron is not monophyletic because the
South American species and the Antillean species are recovered in two different
clades. The Antillean species formed a clade sister to Pleodendron. The
synapomorphies of the Antillean species of Cinnamodendron are tetramerous
flowers with eight petals, eight stamens, four carpels, and four placentae. Based
on the results from the phylogenetic analysis major taxonomy changes are expected
for the family.
Resumen Seis géneros han sido descritos en la familia Canellaceae, cuatro de
estos para el neotrópico y los otros dos para Africa y Madagascar. En las
Antillas se encuentran los géneros Canella, Pleodendron y Cinamodendron.
Canella es un género monotípico de amplia distribución en la región del caribe y
pleodendron se encuentra presente sólo en las antillas mayores y Costa Rica.
Cinnamodendron ha sido dado tanto para las Antillas mayores como para America
del Sur. Un análisis filogenético previo de la familia indica que el género
Cinnamodendron no es monofilético. Existe una separación de las especies
Sudamericanas y Antillanas en clados diferentes. Las especies de las Antillas
forman un clado que es hermano de Pleodendron. Los carácteres sinapomórficos de
las especies antillanas de Cinnamodendron son: flores tetrámeras con ocho petalos,
ocho estambres, cuatro carpelos y cuatro placentas. Basados en los resultados de la
filogenia del grupo, se anticipan cambios taxonómicos para la familia.
Keywords Antilles . Canellaceae . Canella . Cinnamodendron . Pleodendron .
Phylogeny
NO9002; No of Pages
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J. Salazar, K. Nixon
Taxonomic History
The Canellaceae are a small tropical plant family of aromatic trees, rarely treelets and
shrubs, with a disjunct occurrence between tropical and subtropical America (the
Caribbean Region, Costa Rica and South America) and Africa (Madagascar and Eastern
Africa). Six genera and ca. 21 species have been traditionally recognized in this family
(Cronquist, 1981; Takhtajan, 1997; Zanoni, 2004; Hammel & Zamora, 2005): Canella
P. Browne (monotypic widespread Caribbean taxon), Capsicodendron Hoehne (a
monotypic endemic taxon from Brazil), Cinnamodendron Endlicher (Greater Antilles
and South America), Pleodendron Tieghem (Greater Antilles and Costa Rica),
Cinnamosma Baillon (Madagascar) and Warburgia Engler (from Eastern and South
Africa). The highest diversity of this family occurs in the Neotropics with four genera
and approximately 13 species (Endlicher, 1842; Miers, 1858; van Tieghem, 1899;
Urban, 1922, 1928; Uittien, 1926; Sleumer, 1936; Occhioni, 1943, 1947, 1948, 1949;
Steyermark, 1952; Zanoni, 2004, Hammel & Zamora, 2005). The number of taxa has
increased since new species have been discovered recently in Brazil and the Greater
Antilles (Salazar, 2006). The Greater Antilles is a center of diversity and endemicity for
the family and has three genera (Canella, Pleodendron and Cinnamodendron) and
seven species (Miers, 1858; van Tieghem, 1899; Urban, 1922, 1928; Fawcett &
Rendle, 1926; Sleumer, 1936; León & Alain, 1953; Adams, 1972; Liogier, 1983, 1994;
Bornstein, 1989; Liogier & Martorel, 2000; Zanoni, 2004).
Canellaceae species from the West Indies, specially the widespread Canella
winterana Gaertn., are traditionally used for medicinal, fishing, aphrodisiac,
ritualistic, and aromatic purposes (Salazar, 2006). Plant parts are mainly boiled for
drinking and bathing, and more rarely consumed in alcohol. The bark is also grilled
when used as a condiment.
Beverages prepared with these plants are consumed to treat fever with
Canella winterana (Roig, 1991), stomach problems with Canella winterana and
Cinnamodendron ekmanii Sleumer from the Dominican Republic (Salazar, 2006),
pain with Canella winterana (Ayensu, 1981), sexual diseases such as syphilis with
Canella winterana (Ayensu, 1981), and as a post-partum beverage to “clean” the
uterus (‘matriz”) in the Dominican Republic with Canella winterana and
Cinnamodendron sp (Salazar, 2006). Some species of Canellaceae are also used
to catch octopus, Cinnamodendron ekmanii (Salazar, 2006), and fish, Canella
winterana (Liogier, 2000). Their use as an aphrodisiac is a custom mainly in
the West Indies, where Canella winterana (Liogier, 2000), Cinnamodendron
angustifolium Sleumer (Liogier, 2000) and C. sp from Haiti (Salazar, 2006),
C. corticosum Miers. from Jamaica (Salazar, 2006), and C. ekmanii from the
Dominican Republic (Peguero et al., 1995) have been reported as having this use.
Teas are consumed as aromatic beverages, for e.g. Canella winterana (Duss,
1972), Cinnamodendron ekmanii and C. sp (Peguero et al., 1995; Salazar &
Peguero, 1994), Furthermore, the bark of C. corticosum Miers in Jamaica is also
used as a condiment (Hutchinson, 1964; Salazar, 2006)
Most of the Antillean species are regional or local endemics, very rare, and most
of them can be considered as threatened (Salazar, 2006). According to the IUCN
Red List 2007 three species are catalogued as threatened: Cinnamodendron
New discoveries in the Canellaceae in the Antilles
105
corticosum from Jamaica listed as vulnerable (Bellingham, 1998), C. cubense
considered as endangered (Areces-Mallea, 1998), and Pleodendron macranthum
Tiegh. from Puerto Rico listed as critically endangered (World Conservation
Monitoring Centre, 1998). The others Cinnamodendron species from The West
Indies are not listed, but status of their populations is unknown, for e.g. the Haitian
Cinnamodendron angustifolium was collected only once from the type locality
(Salazar, 2006).
The family Canellaceae was published in 1832 by Martius with Canella as
described by Browne in 1756 as the sole genus. The taxonomy of the family has
been confused and controversial since the type genus, Canella, was published.
Members of this family were classified within the Winteraceae mainly because the
bark of Canella and of Drimys J. R. Forster & G. Forster (Winteraceae) have similar
therapeutic effects, and both have been known in folk medicine as “Cortex
winteranus.” However, the most important reason for the taxonomic confusion
between these two genera is that both were merged in a single taxon called
Winterania by Linnaeus (1737), Laurus winterana (Linnaeus, 1754), and Winterania
canella (Linnaeus, 1762–1763). Later Murray (1784) treated them as two different
taxa, Canella alba Murray for the Caribbean species and Wintera aromatica Murray
for the species that is today classified as Winteraceae.
Until relatively recently, the Canellaceae have been assigned either to the
Parietales (Warburg, 1895; Gilg, 1925; Lawrence, 1951), Annonales (Dahlgren,
1983), or to the Magnoliales (Bonnet, 1876; Takhtajan, 1981; Cronquist, 1988).
However, recent molecular and morphological analyses suggest that the family
belongs to the magnoliid clade sensu APG II (2003), which includes several lineages
that traditionally formed the sub-class Magnoliidae sensu Cronquist (1981). Within
this group, the family has been assigned to the order Canellales (APG II, 2003;
Cronquist, 1957) or Winterales (Doyle & Endress, 2000), and as sister taxon to
Winteraceae (Doyle & Endress, 2000; Soltis et al., 2000; Judd et al., 2002; APG II,
2003; Bremer et al., 2003; Soltis & Soltis, 2004; Endress, 2004).
Members of the Canellaceae form a monophyletic group, which share several
synapomorphies such as monadelphous stamens, parietal placentation, and
campylotropous ovules (Salazar, 2006). In addition, plants of this family can be
easily identified by their lenticels on the trunk and branches, aromatic bark and
leaves with peppery taste, simple and alternate-distichous leaves, flowers with fleshy
perianth, three sepals (rarely two) and berry fruit containing reniform seeds (Salazar,
2006).
The first taxon described in this family was the Caribbean species Canella
winterana Gaertn. Although this name was published by Browne in, 1756, the
species was well known prior to its taxonomic description because of its medicinal
use. The bark of Canella was introduced into Europe between the late 16th and early
17th centuries, and it was known as “Canelle blanche” (Bonnet, 1876). However, it
is likely that by then this plant was already known to Europeans because Dr. Diego
Alvarez Chanca, who accompanied Columbus on his second voyage (Dalby, 2001;
Gerbi, 1978), reported the presence of a cinnamon (=“canela” in Spanish), which
was different from the one that was known in Europe. It is believed that this account
refers to Canella winterana (Gerbi, 1978).
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J. Salazar, K. Nixon
The Endemic Species of Cinnamodendron and Pleodedron
Cinnamodendron was published by Endlicher (1840). The first published species of
this genus, the Brazilizan C. axillare Endl. ex Walp., was originally described as
Canella axillaris (Nees & von Martius, 1824). Two additional Brazilian species have
been described: Cinnamodendron dinisii Scwacke (Schwacke, 1898) and C.
sampaioanum Occhioni (Occhioni, 1948). The other South American taxa are:
C. tenuifolium Uittien based on material collected in Surinam in 1921 (Uittien,
1926), and C. venezuelense, a Venezuelan endemic published by Steyermark (1952).
Four species of Cinnamodendron are recognized in the Caribbean Islands, and they
are all confined to the Greater Antilles: Cinnamodendron. angustifolium Sleumer from
Hispaniola, C. corticosum Miers from Jamaica, C. cubense Urb. from Cuba, and
C. ekmanii Sleumer from Hispaniola (Miers, 1858; Urban, 1922; Sleumer, 1936).
The first species of this genus described for the region was the Jamaican
C. corticosum (Miers, 1858). Also, the same taxon was described as C. rubrum by
Grisebach (1864). Bonnet (1876) suggested that the Jamaican species, described by
Sloane (1686–1692, 1725) as Arbor baccifera laurifolia, aromatica, Fructu viridi
calyculato, racemoso refers to C. corticosum, but the illustration and description of
the plant, including characters such as terminal inflorescences and fruit with
gelatinous pulp, clearly suggest that this species correspondeds to Canella winterana
(Salazar, 2006).
Cinnamodendron macranthum Baill. was described as an endemic species from
Puerto Rico (Baillon, 1882). Later Pleodendron was erected based on C. macranthum,
and the combination P. macranthum Tiegh. was made (van Tieghem, 1899). This
genus has been reported for two islands in the Greater Antilles: Hispaniola (P. ekmanii
(Urban, 1928; Liogier, 1983)) and Puerto Rico (P. macranthum (van Tieghem, 1899;
Liogier, 1994; Liogier & Martorel, 2000)). Pleodendron was considered as a genus
endemic to the Greater Antilles until an additional species, P. costaricense N. Zamora,
Hammel & R. Aguilar, was described recently from Costa Rica (Hammel &
Zamora, 2005).
Recent Discoveries
Four of the Greater Antilles endemics have been traditionally placed in
Cinnamodendron: C. angustifolium, C. corticosum, C. cubense, and C. ekmanii).
However, a recent molecular cladistic analysis of the Canellaceae based on ITS,
matK, trnL–trnF, trnDGUC–trnTGGU and rbcL DNA regions from the nuclear and
chloroplast genome places these taxa in a distinct clade that does not include the five
South American species of this genus, C. axillare, C. dinisii, C. sampoianum, C.
tenuifolium, and C. venezuelense (Salazar, 2006) (Fig. 1). Indeed, the clade with the
species from the Greater Antilles is sister to a group composed exclusively by taxa
belonging to Pleodendron (Fig. 1). The same study found that the South American
species of Cinnamodendron and the Brazilian genus Capsicodendron form a
monophyletic group that is part of a clade containing the Old World genera,
Warburgia and Cinnamosma (Salazar, 2006) (Fig. 1).
New discoveries in the Canellaceae in the Antilles
107
Fig. 1 Strict consensus of 42 most parsimonious trees from the combined analysis of ITS, matK, trnL–
trnF, trnD–trnT and rbcL and 49 informative morphological characters by 29 taxa: 22 Canellaceae and
seven outgroups. Bootstrap support values above branches (Salazar, 2006). Lineages of Cinnamodendron
from the Antilles are indicated with solid circles and lineages of Cinnamodendron from South America are
indicated with open circles
The sister relationship between Pleodendron and the Antillean species of
Cinnamodendron is also well supported by cladistic analyses of morphological
characters and of a combined DNA and morphology data sets (Salazar, 2006).
Morphological synapomorphies supporting these two groups are the presence of
many-seeded fruits with fibrous pulp. The two groups of this clade are easily
distinguished because Pleodendron possess hexamerous flowers (12 petals, 12
stamens, 6 carpels, and 6 placentae). In contrast, the Antillean Cinnamodendron
species have tetramerous flowers (8 petals, 8 stamens, 4 carpels, and 4 placentae).
108
J. Salazar, K. Nixon
The South American species of Cinnamodendron do not form a monophyletic
group because the Brazilian genus Capsicodendron is nested within this group. The
synapomorphies of the Cinnamodendron-Capsicodendron clade are a stipe in the
fruit and the presence of a connective projection above the anther (Salazar, 2006).
It is clear that the Greater Antillean and South American species traditionally
classified within Cinnamodendron belong to two distinct phylogenetic clades, which
are distantly related (Kubitzki, 1993; Zanoni, 2004). These two groups present very
well defined morphological synapomorphies that support the separation of the
Cinnamodendron species into two distinct genera, as it has been suggested by
Salazar (2006). Cinnamodendron taxa from South America are characterized by
having flowers with 6–10 petals, generally 10; 8–10 stamens, commonly 10;
bicarpellate or tricarpellate, few ovules (4–6); stipite, a few seeded fruit (4 or less)
with a gelatinous pulp; big seeds (up to 12 mm long). In contrast, the Antillean clade
of “Cinnamodendron” is characterized by having tetramerous flowers (8 petals,
8 stamens, 4 carpels; ovary with 4 placentae), many ovules (more than 14); not
stipitate, many seeded fruit (more than 8) with a non-gelatinous pulp; small seeds
(up to 3 mm long) (Fig. 2).
In summary, the phylogenetic analysis of Canellaceae in combination with
morphological characters has clarified the taxonomy of the genus Cinnamodendron,
which has been traditionally reported for South America as well as the Greater
Antilles. The Antillean species of Cinnamodendron must be placed in a genus
different from the Cinnamodendron species from South America, because the type
for the genus Cinnamodendron is C. axillare from Brazil (Salazar, 2006). A formal
description of this new genus will be published in a separate study.
Fig. 2 Species of Cinnamodendron reported in the Greater Antilles. Cinnamodendron corticosum (a),
Cinnamodendron cubense (b), Cinnamodendron ekmanii (c, d)
New discoveries in the Canellaceae in the Antilles
109
Acknowledgements We thank the staff at the following herbaria for facilitating specimens: K, S,U, MO,
NY, HUH, UPR, CICY, FLAS, BM, US, MU, MEXU, G, IJ, SJ, UPRRP, HAC, HAJB, RB, HB, SP, MBM,
USP, UFSC, UPCB, HBR, FLOR, UEC, BM, BH, and INB. Alberto Veloz, Brígido Peguero, e Idelfonso de
los Angeles from the Botanical Garden of Santo Domingo provided assistance in the field. George Proctor
and Keron Campbell (Museum of Natural History of Jamaica) facilitated the work in Jamaica. Hilaire
Vilmond made possible the work in Haiti. Eugenio Santiago-Valentin, Marcos Caraballo, and Tomas Carlos
supported the work in Puerto Rico. Rosalina Berazain from Jardín Botánico de La Habana made possible the
work in Cuba.
We appreciate the comments and suggestions done by Dennis Stevenson, Eugenio Santiago-Valentin,
and Javier Francisco-Ortega.
Funding for Jackeline Salazar was provided by Cornell University (Harold E. Moore Jr. Endowment
Fund, The Mario Einaudi Center for International Studies: LASP/Tinker Graduate Student Field Research
Grants, Department of Plant Biology), Botanical Society of America, American Society of Plant
Taxonomists, International Association for Plant Taxonomy, The Linnean Society of London, and the
Botanical Garden of Santo Domingo.
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