Burseraceae
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The Burseraceae are a moderate-sized
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of 17-19
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
and about 540
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of woody
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s. The actual numbers given in taxonomic sources differ according to taxonomic revision at the time of writing. The Burseraceae are also known as the torchwood family, the
frankincense Frankincense, also known as olibanum (), is an Aroma compound, aromatic resin used in incense and perfumes, obtained from trees of the genus ''Boswellia'' in the family (biology), family Burseraceae. The word is from Old French ('high-quality in ...
and
myrrh Myrrh (; from an unidentified ancient Semitic language, see '' § Etymology'') is a gum-resin extracted from a few small, thorny tree species of the '' Commiphora'' genus, belonging to the Burseraceae family. Myrrh resin has been used ...
family, or simply the incense tree family. The family includes both
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s and
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s; its species are native to tropical regions of
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,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
,
Australasia Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...
, and the
Americas The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.''Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sing ...
. Because of taxonomic revision, as the family size (in terms of genera and species) differs according to the time period of study; so, too, does the family's higher-
level Level or levels may refer to: Engineering *Level (optical instrument), a device used to measure true horizontal or relative heights * Spirit level or bubble level, an instrument designed to indicate whether a surface is horizontal or vertical *C ...
relationships, including order. Burseraceae is a genetically-supported
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group since APG III and is frequently cited within the Sapindales. It is recognized as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the Anacardiaceae. The Burseraceae are characterized by the generally non- allergenic resin they produce in virtually all plant tissue and their distinctive smooth, yet flaking, aromatic bark.Judd, W.S., Campbell, C.S., Kellogg, E.A., Stevens, P.F., and M.J. Donoghue. 2008. Plant Systematics: A Phylogenetic Approach 3rd ed. Sinauer Associates, Inc., Sunderland, Massachusetts, USA.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 8, June 2007 nd more or less continuously updated since http://www.mobot.org/MOBOT/research/APweb/ The origins of the family can be traced to the
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
(about 65 Mya) when ''Beiselia mexicana'' first diverged in
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
.Weeks, A., Daly, D.C. and B.B. Simpson. 2005. The phylogenetic history and biogeography of the frankincense and myrrh family (Burseraceae) based on nuclear and chloroplast sequence data. Molecular Phylogenetics and Evolution, 35: 85–101. The subsequent divergences in the family lineage and migration of species in the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(53 Mya) out of North America have led to the current distribution of the species being primarily associated with the tropics. Though the family likely originated in North America, the greatest genetic diversity presently is found in the Southern Hemisphere. Tabonuco ('' Dacryodes excelsa'') and gumbo limbo ('' Bursera simaruba'') represent the economic, ethnobotanical, and ecological significance of the Burseraceae in the Western Hemisphere, while frankincense (''
Boswellia sacra ''Boswellia sacra'', also known as ''Boswellia carteri'' and others, and commonly called the frankincense tree or the olibanum tree, is a tree in the genus '' Boswellia,'' in the Burseraceae family, from which frankincense, a resinous dried sap ...
'') and myrrh ('' Commiphora myrrha'') represent the same in the Eastern Hemisphere.


Key characteristics

The Burseraceae trees or
shrubs A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
are characterized by
resins A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Comm ...
(having triterpenoids and ethereal oils)Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia University Press, New York, New York, USA. that are present within the plant tissue from the vertical resin canals and ducts in the bark to the leaf veins.Harley, M.M., Song, U. and H.I. Banks. 2005. Pollen morphology and systematics of Burseraceae. Grana, 44: 282–299.Heywood, V.H. 1993. Flowering Plants of the World. Oxford University Press, New York, New York, USA.Mabberley, D.J. 1997. The Plant Book: A portable dictionary of the vascular plants. Cambridge University Press, Cambridge, UK. In fact, the
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
of the Burseraceae is the smooth yet peeling or flaking aromatic bark. The clear, nonallergenic resins may smell like almonds, but at least the most well known resins, frankincense and myrrh, have a distinct odor. The leaves are generally alternate, spiral, and odd-pinnately compound with opposite, frequently long-petiolulate, entire to serrate, pinnately veined leaflets whose symmetry is distinctive in some genera. However, some members are known to have trifoliate or unifoliate leaves. The leaf and leaflet stalks and axis may be brown and scurfy, while the leaf base is swollen and may be concave adaxially. The family members tend to be without
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...
s. The determinate, axillary
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
s carry small, radial, unisexual flowers. The plants tend to be
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
. The flowers may have four or five faintly connate but imbricate
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s with an equal number of distinct, imbricate
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s. Also, the stamens, that may contain nectar discs, have distinct glabrous filaments that occur in one or two whorls and in numbers equaling or twice the number of petals; the tricolporate
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
is contained within two
locule A locule (: locules) or loculus (; : loculi) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usually refers to a chamber within an ovary ...
s of the
anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s that open longitudinally along slits. The
gynoecium Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl (botany), whorl of a flower; it consists ...
contains 3–5 connate
carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...
s, one
style Style, or styles may refer to: Film and television * ''Style'' (2001 film), a Hindi film starring Sharman Joshi, Riya Sen, Sahil Khan and Shilpi Mudgal * ''Style'' (2002 film), a Tamil drama film * ''Style'' (2004 film), a Burmese film * '' ...
, and one stigma that is head-like to lobed. Each locule of the superior ovary has two
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the sporangium, megasporangium), ...
s with axile placentation that are anatropous to campylotropous. The one- to five-pitted
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
is a
drupe In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
that opens at maturity. The endosperm is usually lacking in the embryo.


Taxonomy

Some discrepancy exists in the literature about the size of the Burseraceae. Records say that the family has 17 to 18 genera and 500 to 540 to 726 species. Other authors cite different numbers: 16–20 genera and 600 species; 20 genera and 500–600 species;Lawrence, G.H.M. 1951. Taxonomy of Vascular Plants. The Macmillan Company, New York, New York, USA. According to a pollen studies and molecular data, the family is split into three
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
s: the Protieae, Bursereae, and Canarieae. The Protieae are composed of ''Protium'' (147 species and largest in this tribe), ''Crepidospermum, Garuga'', and ''Tetragastris''. The Bursereae, which are further split into
subtribe Subtribe is a taxonomic category ranking which is below the rank of tribe and above genus. The standard suffix for a subtribe is -ina (in animals) or -inae (in plants). The first use of this word dates back to the late 19th century. An example of ...
s Boswelliinae and Burserinae, contain ''Commiphora'' (nearly 200 species and largest in the family), ''Aucoumea, Beiselia, Boswellia, Bursera,'' and ''Triomma''. Finally, the Canarieae are composed of ''Canarium'' (75 species and largest in this tribe), ''Dacryodes, Haplolobus, Pseudodacryodes, Rosselia, Santiria, Scutinanthe'', and ''Trattinnickia''. The morphology of the fruit, which is a drupe, helps to distinguish between the three tribes. Though the groupings have slightly changed since the 1990s, the Protieae are described as having a two- to five-parted drupe with either ‘free or adhering parts’ which are ‘not fused in the endocarp’; The Bursereae are described as having a drupe with parts that are fused in the endocarp, but an exocarp with dehiscing valves; and the Canarieae as simply having a drupe with parts that are fused in the endocarp. This is a list of the 19 genera of the Burseraceae with placement in three tribes (and subtribes where applicable):


Subfamilies and genera

;Bursereae *'' Aucoumea'' *'' Beiselia'' *''
Boswellia ''Boswellia'' is a genus of trees in the order (biology), order Sapindales, known for its fragrant resin. The biblical incense frankincense is an extract from the resin of the tree ''Boswellia sacra'', and is now produced also from ''Boswellia fre ...
'' (frankincense) *'' Bursera'' (Palo Santo) *'' Commiphora'' (myrrh) *'' Garuga'' *'' Triomma'' ;Canarieae *'' Ambilobea'' *'' Canarium'' *'' Dacryodes'' *'' Haplolobus'' *'' Pseudodacryodes'' *'' Rosselia'' *'' Santiria'' *'' Scutinanthe'' *'' Trattinnickia'' ;Protieae *'' Protium'' (including '' Tetragastris'') (copal) ;Unplaced *†'' Barghoornia'' (
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
Klondike Mountain Formation) *'' Pachylobus''


Order

According to the literature, the Burseraceae have not been lumped with other families nor split up into several others. However, they have "jumped" orders several times. For example, in the early 19th century, the family seems to have been placed in the Burserales, with the Anacardiaceae and Podoaceae. In the mid-19th century and early 20th century, the family was placed in the Geraniales. Then, by the mid- and late-20th century, the family was moved to the Rutales. Finally, in the late 20th century, the family was (and today still is) located within the Sapindales. Families that are consistently found in the same order as the Burseraceae (except when in the Burserales) include the
Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...
, Meliaceae, and Simaroubaceae. Only in recent studies were the Burseraceae and the Anacardiaceae seen as sister groups. The Sapindales are contained within the malvids of the rosid
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
within the eudicotyledons. The order contains nine to 15 families,Gadek, P.A., E.S. Fernando, C.J. Quinn, S.B. Hoot, T. Terrazas, MC. Sheahan, and M.W. Chase. 1996. Sapindales: Molecular delimitation and infraordinal groups. Am. J. Bot. 83: 802–811. 460 genera, and from 5,400 to 5,670 to 5,800 species. The currently recognized families include Aceraceae, Anacardiaceae, Burseraceae, Hippocastanaceae, Julianaceae, Meliaceae, Rutaceae, Sapindaceae, and Simaroubaceae. The Sapindales are a clade supported by DNA-based analyses on ''rbc''L, ''atp''B, and 18S sequences. Within the Sapindales are two clades that contain gum and resin: the Rutaceae-Meliaceae-Simaroubaceae clade and the Burseraceae-Anacardiaceae clade. The Burseraceae are thus not the only family in the Sapindales with this characteristic. The synapomorphies of the clade include pinnately compound alternately- or spirally-arranged leaves that may be palmately compound, trifoliate, or unifoliate, and small four- or five-merous flowers having a characteristic nectar disk and imbricate petals and sepals. Some of these characteristics also occur in the Rosales. However, the Sapindales and Rutales may actually form a complex, since many families "jump" between them. Indeed, ''rbc''L sequence studies seem to indicate that a sapindalean/rutalean complex exists and may better represent the relationships of the families than the separate orders would. A study based on the
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
-encoded
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
''rbc''L reconstructed cladograms that include families within both the Sapindales and Rutales. One such
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
indicated that the Sapindales are robust and that the Burseraceae (and Anacardiaceae) are within a single clade of their own. This grouping seems to make sense, as both the Burseraceae and Anacardiaceae have secretory canals in the phloem and resin canals in the leaves, and are unique in the Sapindales for having biflavones in the leaf tissue. However, the two families have several distinguishing characteristics. The resin of the Burseraceae is nonallergenic and two ovules per carpel occur, whereas the resin of the Anacardiaceae can be allergenic or poisonous and one ovule per carpel is found. The Burseraceae-Anacardiaceae clade is sister to a robust cluster of three other families, the Sapindaceae-Aceraceae-Hippocastanaceae clade. The Rutaceae-Meliaceae-Simaroubaceae clade is sister to the Burseraceae-Anacardiaceae and Sapindaceae-Aceraceae-Hippocastanaceae clade. The ''rbc''L technique is supported and considered acceptable until such time as other analytical methods become better developed.


Biogeography

The Burseraceae are distributed throughout the world and primarily in the tropics, especially
Malesia Malesia is a biogeographical region straddling the Equator and the boundaries of the Indomalayan and Australasian realms. It is a phytogeographical floristic region in the Paleotropical kingdom. It was first recognized as a distinct region ...
, Africa, and Central and South America. The three tribes can be linked to a specific region of the world, although this is not obligatory. For example, members of the tribe Protieae are generally found in South America, those of the Bursereae are found in Africa and
Mesoamerica Mesoamerica is a historical region and cultural area that begins in the southern part of North America and extends to the Pacific coast of Central America, thus comprising the lands of central and southern Mexico, all of Belize, Guatemala, El S ...
, while members of the Canarieae are found in Malesia. However, each tribe has a representative genus present in all the tropical regions: ''Dacryodes'' (Canarieae), ''Protium'' (Protieae), and ''Commiphora'' (Bursereae). The Burseraceae are found in a variety of habitats, including hot, dry
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
and
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
, as well as in coastal
mangrove A mangrove is a shrub or tree that grows mainly in coastal saline water, saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen a ...
forest and
rain forest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
habitats. One study found that the family originated in North America during the
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
(about 65 Mya), when the earliest fossils of the Sapindales are found. During the Early to Middle Eocene (about 53 Mya), family members dispersed to eastern
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
(i.e. Europe and Asia) via the Boreotropical Land Bridge and the continents in the Southern Hemisphere, which is now the area of the greatest generic diversity of this family. More specifically, the earliest diverging genus was ''Beiselia'' (of the Bursereae subtribe Boswelliinae) in either North America, Mexico, or the
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
in the Paleocene. Similar results from other studies find that ''Beiselia mexicana'', a native of Mexico, is basal to the remaining Burseraceae. These results may indicate that the family originated in Mexico. The next divergence was in the Early Eocene when the Burserinae (i.e. ''Commiphora'') diverged and emigrated from North America into Africa,
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
, and India. ''Commiphora'' dispersed throughout Africa during the Middle Eocene (about 44 Mya) and from Africa to Madagascar during the Oligocene (about 30 Mya) via the Mozambique Channel Land Bridge; the spread to India was more recent (about 5 Mya). The Canarieae and Boswelliinae (subtribes of Bursereae) dispersed from western
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
and spread eastward during the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
;
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
of Canarium, for example, from the
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
date to the Late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
(23 Mya). Finally, the Protieae originated in North America like the rest of the family, then migrated to Africa and Asia through the Tethys seaway in the Late
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(about 37 Mya), but then made its way back to South America via long-distance dispersal. By the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
(about 23Mya), all three Burseraceae tribes were extant and dispersed throughout the Northern Hemisphere. The mechanism of seed dispersal via animal link vectors ( endozoochoric dispersal) may explain how most Burseraceae were able to expand their range so efficiently across the globe. ''Beiselia, Boswellia'', and ''Triomma'' have dry fruits better suited for wind dispersal, but most Burseraceae have fleshy, edible fruit that is eaten by many animal dispersers. The seeds may provide a high reward in fat (24–73%) and protein (2.7–25.9%) if digested, but many animals eat just the fleshy part of the fruit and either discard the endocarp right away or excrete it some time later. Some known Burseraceae fruit consumers include hornbills (''Buceros bicornis, Ceratogyma atrata, C. cylindricus, Penelopides panini''), oilbirds (''Steatnoris caripensis''), fruit pigeons, warblers, vireos, orioles, flycatchers, tanagers, woodpeckers, loeries, primates (''Cercopithecus'' spp., ''Lophocebus albigena''), lemurs (''Varecia variegate'' subsp. ''variegate''), and sun bears (''Helarctos malayanus''). The fruits may also have been water dispersed.


Economic significance and ethnobotanic uses

Several representative species within the Burseraceae typify the economic and ethnobotanic significance of the family. First, ''Dacryodes excelsa'' of the Canarieae is an important old-growth species found in the Caribbean. Second, ''Bursera simaruba'' of the Burserinae is a fast-growing ornamental that is one of a few representatives of the primarily tropical family in the United States. Finally, the namesakes of the family ''Boswellia carterii'' (frankincense) and ''Commiphora abyssinica'' (myrrh) are important economically and medicinally in several parts of the world. Though this is a small subset of the large number of potentially important species, these four members exemplify the wide use and importance of the Burseraceae. The latter three are frequently cited in the literature for their renowned importance. Commonly known as tabonuco (or gommier, also candlewood), '' Dacryodes excelsa'' is a large, dominant tree found in
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
and other parts of the Caribbean. The seeds of the tree are a source of food for birds. Like all members, the tree releases sap from the bark when wounded. The clear sap oozes from the tree and hardens to a white, aromatic waxy
resin A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...
that can be used to make candles and incense. Before the arrival of the Spaniards to Puerto Rico, the native Taínos used the resin to make torches. The wood itself is useful for constructing housing, furniture, boxes, small boats, and a variety of other wood-based products; the utility of the wood is comparable to that of mahogany and birch. In addition, species like '' Canarium littorale, Dacryodes costata, Santiria laevigata,'' and Santiria tomentosa from Malaysia, as well as '' Aucoumea klaineana'' and '' Canarium schweinfurthii'' from Africa, also produce valuable wood for construction projects and carpentry. Several species in the genus ''Canarium'' are used as edible fruits (for example, '' Canarium album'', the Chinese olive) and nuts (pili nuts, ''
Canarium ovatum ''Canarium'' is a genus of about 120 species of tropical and subtropical trees, in the family Burseraceae. They grow naturally across tropical Africa, south and southeast Asia, Indochina, Malesia, Australia and western Pacific Islands; in ...
''). Species in the genus ''Bursera'', especially the so-called elephant tree, grow primarily in Mexico, where their secretions are a raw material in making
varnish Varnish is a clear Transparency (optics), transparent hard protective coating or film. It is not to be confused with wood stain. It usually has a yellowish shade due to the manufacturing process and materials used, but it may also be pigmente ...
. The
Maya Maya may refer to: Ethnic groups * Maya peoples, of southern Mexico and northern Central America ** Maya civilization, the historical civilization of the Maya peoples ** Mayan languages, the languages of the Maya peoples * Maya (East Africa), a p ...
also used a ''Bursera'' sp. to make
incense Incense is an aromatic biotic material that releases fragrant smoke when burnt. The term is used for either the material or the aroma. Incense is used for aesthetic reasons, religious worship, aromatherapy, meditation, and ceremonial reasons. It ...
. However, the ''Bursera'' may also be considered an ornamental genus and a common representative of the family in the United States, especially in Florida (''B. simaruba'') and the Southwest (''B. odorata, B. microphylla''). Naked Indian (also known as gumbo limbo), or ''Bursera simaruba'', in particular, is found in
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
, Mexico, the Caribbean,
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
, and
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. The tree is also named the ‘tourist tree’ for its very distinctive flaking red bark; apparently, the tree occurs in tropical areas where many white tourists go on vacation.Plant Creations, Inc. Updated: August 28, 2007. Bursera simaruba. http://www.plantcreations.com/bursera_simaruba.htm The resin from this tree can be used to make varnish and turpentine. In addition, the resin may also be used similarly to tiger balm (containing ''Cinnamomum camphora'' of the
Lauraceae Lauraceae, or the laurels, is a plant Family (biology), family that includes the bay laurel, true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genus (biology), genera worldwide. They are dicotyled ...
) to relieve sprains and muscle aches. The leaves are used to brew a tea to relieve inflammation. The bark serves as an antidote to skin irritation caused by ''Metopium toxiferum'' (also known as poisonwood, Florida poison tree, and hog gum) of the Anacardiaceae. The gumbo limbo grows quickly and can be used to make a living fence especially out of cut limbs that are placed straight into the ground or for restoration projects as a pioneer species. The tree is highly tolerant of high-intensity wind such as hurricane-force winds, so is planted in areas where hurricanes occur frequently, such as Florida and the Caribbean. The seeds of this species are also a source of food for birds. Frankincense, or olibanum, ('' Boswellia carterii'') and myrrh ('' Commiphora abyssinica'') have long been valued for the aromatic resins they produce. These resins are extracted via tapping, or cutting of the bark to make it release sap. The liquid sap hardens and is gathered, sold as is or further processed and mixed with spices, seeds, and roots to make various forms of incense.Dharmananda, S. Created: May 2003. Myrrh and Frankincense. http://www.itmonline.org/arts/myrrh.htm Both species are native to parts of Northeast Africa (
Somalia Somalia, officially the Federal Republic of Somalia, is the easternmost country in continental Africa. The country is located in the Horn of Africa and is bordered by Ethiopia to the west, Djibouti to the northwest, Kenya to the southwest, th ...
, frankincense; Somalia and
Ethiopia Ethiopia, officially the Federal Democratic Republic of Ethiopia, is a landlocked country located in the Horn of Africa region of East Africa. It shares borders with Eritrea to the north, Djibouti to the northeast, Somalia to the east, Ken ...
, myrrh) and
Arabia The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
(
Oman Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
and
Yemen Yemen, officially the Republic of Yemen, is a country in West Asia. Located in South Arabia, southern Arabia, it borders Saudi Arabia to Saudi Arabia–Yemen border, the north, Oman to Oman–Yemen border, the northeast, the south-eastern part ...
, frankincense), but their distribution and use have been extended beyond these regions to India and China. The best frankincense is grown in Oman and the incense is widely used in worship in India. The ancient Egyptians prized frankincense for the resin they used to make the characteristic dark eyeliner and myrrh as an embalming agent for deceased pharaohs. At that time, myrrh was worth more than gold. In modern times resins from these trees are used in Chinese herbal medicine and Indian Ayurvedic medicine to treat several ailments. Pills containing small doses of frankincense and other ingredients are valued in
oriental medicine Traditional Asian medicine is a collective term for several types of traditional medicine practiced in Asia. These include the medical traditions of: * East Asia ** Traditional Chinese medicine, China *** Traditional Tibetan medicine, Tibet ** Kam ...
for promoting blood flow and the movement of the '' qi'' (‘life force’ or ‘spiritual energy’). and myrrh is similarly claimed to promote blood flow, stimulate the stomach and digestion, and to be useful in treating diabetes, menopause, uterine tumors, amenorrhoea, and
dysmenorrhea Dysmenorrhea, also known as period pain, painful periods or menstrual cramps, is pain during menstruation. Its usual onset occurs around the time that menstruation begins. Symptoms typically last less than three days. The pain is usually in th ...
. Both frankincense (containing triterpene acids) and myrrh are used to relieve pain and inflammation as in arthritis and asthma.Hanus, L.O., Rezanka, T., Dembitsky, V.M. and A. Moussaieff. 2005. Myrrh- Commiphora Chemistry. Biomed. Papers, 149(1): 3–28.


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External links

{{Authority control Sapindales families