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The Burseraceae are a moderate-sized
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of 17-19
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and about 540
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s. The actual numbers differ according to the time period in which a given source is written describing this family. The Burseraceae are also known as the torchwood family, the
frankincense Frankincense (also known as olibanum) is an aromatic resin used in incense and perfumes, obtained from trees of the genus ''Boswellia'' in the family Burseraceae. The word is from Old French ('high-quality incense'). There are several species o ...
and
myrrh Myrrh (; from Semitic, but see '' § Etymology'') is a gum-resin extracted from a number of small, thorny tree species of the genus ''Commiphora''. Myrrh resin has been used throughout history as a perfume, incense and medicine. Myrrh mi ...
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, including only woody plants with secondary growth, plants that are ...
s and
shrub A shrub (often also called a 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 ...
s, and is native to tropical regions of
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
.
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologica ...
, and the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
. Just as the family size (in terms of genera and species) differs according to the time period of the study, so, too, does its placement in the higher ranks. Nevertheless, the family is a genetically supported
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group currently and frequently cited within the
Sapindales Sapindales is an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem. The APG III system of 2009 includes it ...
and is recognized as a sister group to the
Anacardiaceae The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of the Anacardiaceae bear fruits that are drupes and in some cases produce ...
. The Burseraceae are characterized by the nonallergenic resin they produce in virtually all plant tissue and the 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 The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...
. 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), E ...
(about 65 Mya) when ''Beiselia mexicana'' first diverged in
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
.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 the species in the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
(53 Mya) from North America have led to the current distributions of the species that are primarily associated with the tropics. Though the family likely originated in North America, the greatest genetic diversity presently is in the Southern Hemisphere.
Tabonuco ''Dacryodes excelsa'' is a tree native to Puerto Rico with a habitat that extends into the Lesser Antilles in the Caribbean region. Its English vernacular names include gommier and candlewood. Its Spanish common name is ''tabonuco''. According ...
(''
Dacryodes excelsa ''Dacryodes excelsa'' is a tree native to Puerto Rico with a habitat that extends into the Lesser Antilles in the Caribbean region. Its English vernacular names include gommier and candlewood. Its Spanish Spanish might refer to: * Items from o ...
'') and
gumbo limbo ''Bursera simaruba'', commonly known as gumbo-limbo, copperwood, chaca, West Indian birch, naked Indian, and turpentine tree, is a tree species in the family Burseraceae, native to the Neotropics, from South Florida to Mexico and the Caribbean ...
(''
Bursera simaruba ''Bursera simaruba'', commonly known as gumbo-limbo, copperwood, chaca, West Indian birch, naked Indian, and turpentine tree, is a tree species in the family Burseraceae, native to the Neotropics, from South Florida to Mexico and the Caribbean ...
'') represent the economic, ethnobotanical, and ecological significance of the Burseraceae in the Western Hemisphere, while frankincense (''
Boswellia sacra ''Boswellia sacra'' (commonly known as frankincense or olibanum-tree) is a tree in the Burseraceae family. It is the primary tree in the genus ''Boswellia'' from which frankincense, a resinous dried sap, is harvested. It is native to the Arabian ...
'') and myrrh (''
Commiphora myrrha The genus of the myrrhs, ''Commiphora'', is the most species-rich genus of flowering plants in the frankincense and myrrh family, Burseraceae. The genus contains approximately 190 species of shrubs and trees, which are distributed throughout the ...
'') represent the same in the Eastern Hemisphere.


Key characteristics

The Burseraceae
trees 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, including only woody plants with secondary growth, plants that are u ...
or
shrubs A shrub (often also called a 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 ...
are characterized by
resins In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
(having
triterpenoids Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all produce triterpenes, including squale ...
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 character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
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 an odor that is distinct from almonds, smelling like incense. 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). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s. The determinate, axillary
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s carry small, radial, unisexual flowers. The plants tend to be
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
. The flowers may have four or five faintly connate but
imbricate Aestivation or estivation is the positional arrangement of the parts of a flower within a flower bud before it has opened. Aestivation is also sometimes referred to as praefoliation or prefoliation, but these terms may also mean vernation: the ar ...
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 The term ''sepalum'' was coined b ...
s with an equal number of distinct, imbricate
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s. Also, the stamens, that may contain
nectar disc Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualis ...
s, 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 seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
is contained within two
locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) 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'' usu ...
s of the
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s that open longitudinally along slits. The
gynoecium Gynoecium (; ) 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) ''pistils'' ...
contains 3–5 connate
carpel Gynoecium (; ) 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) ''pistils'' ...
s, one
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
, 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 megasporangium), and the fe ...
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 that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
is a
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or '' pyrena'') of hardened endocarp with a seed (''kernel'') ...
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 usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
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 subtribes 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 Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Agg ...
, but an
exocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggr ...
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 The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...
(and subtribes where applicable):


Subfamilies and genera

;Bursereae *'' Aucoumea'' *'' Beiselia'' *''
Boswellia ''Boswellia'' is a genus of trees in the 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 '' B. frereana''. Frankincens ...
'' (frankincense) *''
Bursera ''Bursera'' is a genus with about 100 described species of flowering shrubs and trees varying in size up to high. It is the type genus for Burseraceae. The trees are native (often for many species endemic) to the Americas, from the southern Uni ...
'' *''
Commiphora The genus of the myrrhs, ''Commiphora'', is the most species-rich genus of flowering plants in the frankincense and myrrh family, Burseraceae. The genus contains approximately 190 species of shrubs and trees, which are distributed throughout th ...
'' (myrrh) *'' Garuga'' *'' Triomma'' ;Canarieae *'' Ambilobea'' *''
Canarium ''Canarium'' is a genus of about 100 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; inclu ...
'' *''
Dacryodes ''Dacryodes'' is a genus of about 60 species of trees in the family Burseraceae. The generic name is from the Greek ' meaning "tear(drop)", referring to how resin droplets form on the bark surface. Description ''Dacryodes'' species grow as shrub ...
'' *'' Haplolobus'' *'' Pseudodacryodes'' *''
Rosselia ''Rosselia'' is a genus of plants in the family Burseraceae. The sole species is ''Rosselia bracteata'', endemic to Papua New Guinea. It is a tree and grows primarily in the wet tropical biome(s). Both the genus and the species were circumscribe ...
'' *'' Santiria'' *'' Scutinanthe'' *'' Trattinnickia'' ;Protieae *'' Protium'' (including ''
Tetragastris ''Tetragastris'' is a genus of plants in family Burseraceae The Burseraceae are a moderate-sized family of 17-19 genera and about 540 species of flowering plants. The actual numbers differ according to the time period in which a given sourc ...
'') (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 The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Wash ...
)


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 The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of the Anacardiaceae bear fruits that are drupes and in some cases produce ...
and
Podoaceae Anacardioideae is a subfamily of plants in the family Anacardiaceae The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of t ...
. 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, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
,
Meliaceae Meliaceae, the mahogany family, is a flowering plant family of mostly trees and shrubs (and a few herbaceous plants, mangroves) in the order Sapindales. They are characterised by alternate, usually pinnate leaves without stipules, and by syncar ...
, and
Simaroubaceae The Simaroubaceae are a small, mostly tropical, family in the order Sapindales. In recent decades, it has been subject to much taxonomic debate, with several small families being split off. A molecular phylogeny of the family was published in 200 ...
. Only in recent studies were the Burseraceae and the
Anacardiaceae The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of the Anacardiaceae bear fruits that are drupes and in some cases produce ...
seen as sister groups. The
Sapindales Sapindales is an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem. The APG III system of 2009 includes it ...
are contained within the
malvids The rosids are members of a large clade (monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classificatio ...
of the
rosid The rosids are members of a large clade ( monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classifica ...
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
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. Therefore, the Burseraceae are not the only family with this characteristic. The
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
of the Sapindales include pinnately compound, alternate and spiral 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 Rosales () is an order of flowering plants. Peter F. Stevens (2001 onwards). "Rosales". At: Trees At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below) It is sister to a clade consisting of Faga ...
. 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
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
-encoded
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
''rbc''L reconstructed cladograms that include families within both the Sapindales and Rutales. One such
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
indicated that the Sapindales are robust and that the Burseraceae (and Anacardiaceae) are within a single clade. 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 other methods become better developed for the analysis.


Biogeography

The Burseraceae are distributed throughout the world and primarily in the tropics, especially
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
, 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 in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
, while members of the Canarieae are found in Malaysia. 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 barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
and
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the Canopy (forest), canopy does not close. The open canopy allows sufficient light to rea ...
, as well as in coastal
mangrove A mangrove is a shrub or tree that grows in coastal saline water, saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evoluti ...
forest and
rain forest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainforest ...
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), E ...
(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 Pan ...
(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 (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
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 (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
, 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 Pan ...
and spread eastward during the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
;
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 in ...
of Canarium, for example, from the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
date to the Late
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
(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 ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
(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 of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
(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 Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Agg ...
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 ''Dacryodes excelsa'' is a tree native to Puerto Rico with a habitat that extends into the Lesser Antilles in the Caribbean region. Its English vernacular names include gommier and candlewood. Its Spanish Spanish might refer to: * Items from o ...
'' is a large, dominant tree found in
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and Unincorporated ...
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 In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
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 Mahogany is a straight-grained, reddish-brown timber of three tropical hardwood species of the genus ''Swietenia'', indigenous to the AmericasBridgewater, Samuel (2012). ''A Natural History of Belize: Inside the Maya Forest''. Austin: Unive ...
and
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 30 ...
. In addition, species like '' Canarium littorale,
Dacryodes costata ''Dacryodes costata'' is a tree in the family Burseraceae. The specific epithet ' is from the Latin meaning "ribbed", likely referring to the prominent veins on the leaf underside. Description ''Dacryodes costata'' grows up to tall with a trunk ...
, Santiria laevigata,'' and Santiria tomentosa from Malaysia, as well as ''
Aucoumea klaineana ''Aucoumea klaineana'' (angouma, gaboon, or okoumé) is a tree in the family Burseraceae, native to equatorial west Africa in Gabon, the Republic of the Congo, and Río Muni. It is a large hardwood tree growing to tall, rarely larger, with a ...
'' and ''
Canarium schweinfurthii ''Canarium schweinfurthii'' (commonly known as the bush candle, African olive, African elemi, or canarium), is a species of large tree native to tropical Africa.ICRAF Names in many African languages are variations of ''mupafu''. Description Bec ...
'' 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 ovatum'', the pili (Central Bikol and Filipino: ''pili'', ;), is a species of tropical tree belonging to the genus ''Canarium''. It is one of approximately 600 species in the family Burseraceae. ''C. ovatum'' are native to the Ph ...
''). 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 transparent hard protective coating or film. It is not a stain. It usually has a yellowish shade from the manufacturing process and materials used, but it may also be pigmented as desired, and is sold commercially in various ...
. The
Maya Maya may refer to: Civilizations * Maya peoples, of southern Mexico and northern Central America ** Maya civilization, the historical civilization of the Maya peoples ** Maya language, the languages of the Maya peoples * Maya (Ethiopia), a populat ...
also used a ''Bursera'' sp. to make
incense Incense is 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 ceremony. It may also be ...
. 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 state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
, Mexico, the Caribbean,
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
, and
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. 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 Tiger Balm () is an analgesic heat rub manufactured and distributed by Singaporean company Haw Par Healthcare. It is used for external pain relief. History A precursor to Tiger Balm called Ban Kin Yu ( zh, t=萬金油, l=Ten Thousand Gol ...
(containing ''Cinnamomum camphora'' of the
Lauraceae Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur ma ...
) 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, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constituti ...
, frankincense; Somalia and
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
, myrrh) and
Arabia The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plate. ...
(
Oman Oman ( ; ar, عُمَان ' ), officially the Sultanate of Oman ( ar, سلْطنةُ عُمان ), is an Arabian country located in southwestern Asia. It is situated on the southeastern coast of the Arabian Peninsula, and spans the mouth of t ...
and
Yemen Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the southern end of the Arabian Peninsula, and borders Saudi Arabia to the Saudi Arabia–Yemen border, north and ...
, 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 Pharaoh (, ; Egyptian: ''pr ꜥꜣ''; cop, , Pǝrro; Biblical Hebrew: ''Parʿō'') is the vernacular term often used by modern authors for the kings of ancient Egypt who ruled as monarchs from the First Dynasty (c. 3150 BC) until the ann ...
. 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 Ayurveda () is an alternative medicine system with historical roots in the Indian subcontinent. The theory and practice of Ayurveda is pseudoscientific. Ayurveda is heavily practiced in India and Nepal, where around 80% of the population rep ...
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 ** China *** Tibet ** Japan (Kampo) ** Korea ** Mongolia * Southeast Asia ** Cambodi ...
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 Amenorrhea is the absence of a menstrual period in a woman of reproductive age. Physiological states of amenorrhoea are seen, most commonly, during pregnancy and lactation (breastfeeding). Outside the reproductive years, there is absence of menses ...
, 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 the ...
. 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.


References


External links

{{Taxonbar, from=Q161237 Sapindales families