Leptospermoideae
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Myrtaceae (), the myrtle family, is a family of
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, ...
ous
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s placed within the order
Myrtales The Myrtales are an order of flowering plants in the malvid clade of the rosid group of dicotyledons. Well-known members of Myrtales include: myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, eucalyptus, crape myrtles, ...
. Myrtle,
pōhutukawa Pōhutukawa (''Metrosideros excelsa''), also known as the New Zealand Christmas tree, or iron tree, is a coastal evergreen tree in the Myrtus, myrtle family, Myrtaceae, that produces a brilliant display of red (or occasionally orange, yellow o ...
, bay rum tree,
clove Cloves are the aromatic flower buds of a tree in the family Myrtaceae, ''Syzygium aromaticum'' (). They are native to the Maluku Islands, or Moluccas, in Indonesia, and are commonly used as a spice, flavoring, or Aroma compound, fragrance in fin ...
,
guava Guava ( ), also known as the 'guava-pear', is a common tropical fruit cultivated in many tropical and subtropical regions. The common guava '' Psidium guajava'' (lemon guava, apple guava) is a small tree in the myrtle family (Myrtaceae), nativ ...
, acca (feijoa),
allspice Allspice, also known as Jamaica pepper, myrtle pepper, pimenta, or pimento, is the dried unripe berry of ''Pimenta dioica'', a midcanopy tree native to the Greater Antilles, southern Mexico, and Central America, now cultivated in many warm par ...
, and
eucalyptus ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...
are some notable members of this group. All species are woody, contain
essential oil An essential oil is a concentrated hydrophobic liquid containing volatile (easily evaporated at normal temperatures) chemical compounds from plants. Essential oils are also known as volatile oils, ethereal oils, aetheroleum, or simply as the ...
s, and have flower parts in multiples of four or five. The
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
are
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
, alternate to mostly opposite, simple, and usually entire (i.e., without a toothed margin). The
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s have a base number of five petals, though in several genera, the petals are minute or absent. The
stamen 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 are usually very conspicuous, brightly coloured, and numerous.


Evolutionary history

Scientists hypothesize that the family Myrtaceae arose between 60 and 56 million years ago (Mya) 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), ...
era. Pollen fossils have been sourced to the ancient supercontinent
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. The breakup of
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period (145 to 66 Mya) geographically isolated disjunct taxa and allowed for rapid speciation; in particular, genera once considered members of the now-defunct Leptospermoideae alliance are now isolated within
Oceania Oceania ( , ) is a region, geographical region including Australasia, Melanesia, Micronesia, and Polynesia. Outside of the English-speaking world, Oceania is generally considered a continent, while Mainland Australia is regarded as its co ...
. Generally, experts agree that
vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
is responsible for the differentiation of Myrtaceae taxa, except in the cases of ''
Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the greate ...
'' species now located on
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
and
New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...
, islands which may have been submerged at the time of 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 ...
differentiation.


Diversity

Recent estimates suggest the Myrtaceae include about 5,950 species in about 132 genera. The family has a wide distribution in tropical and warm-temperate regions of the world, and is common in many of the world's
biodiversity hotspot A biodiversity hotspot is a ecoregion, biogeographic region with significant levels of biodiversity that is threatened by human habitation. Norman Myers wrote about the concept in two articles in ''The Environmentalist'' in 1988 and 1990, after ...
s. Genera with
capsular In botany, a capsule is a type of simple, dry, though rarely fleshy dehiscent fruit produced by many species of angiosperms (flowering plants). Origins and structure The capsule (Latin: ''capsula'', small box) is derived from a compound (mult ...
fruits such as ''
Eucalyptus ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...
'', ''
Corymbia ''Corymbia'', commonly known as bloodwoods, is a genus of about one hundred species of tree that, along with ''Eucalyptus'', '' Angophora'' and several smaller groups, are referred to as eucalypts. Until 1990, corymbias were included in the g ...
'', ''
Angophora ''Angophora'' is a genus of nine species of trees and shrubs in the myrtle family, Myrtaceae. Endemic to eastern Australia, they differ from other eucalypts in having juvenile and adult leaves arranged in opposite pairs, sepals reduced to projec ...
'', ''
Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the greate ...
'', and ''
Melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles, bottlebrushes or tea-trees (although the last name is also applied to species of '' Leptospermum''). They ...
'' are absent from the Americas, apart from ''
Metrosideros ''Metrosideros'' is a genus of approximately 60 trees, shrubs, and vines in the family Myrtaceae, mostly found in the Pacific region. Most of the tree forms are small, but some are exceptionally large, the New Zealand species in particular. The ...
'' in
Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
and
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. Genera with fleshy fruits have their greatest concentrations in eastern
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
and
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 ...
(the
Australasian realm The Australasian realm is one of eight biogeographic realms that is coincident with, but not (by some definitions) the same as, the geographical region of Australasia. The realm includes Australia, the island of New Guinea (comprising Papua Ne ...
) and the
Neotropic The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...
s. ''Eucalyptus'' is a dominant, nearly ubiquitous genus in the more mesic parts of Australia and extends north sporadically to the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
. ''
Eucalyptus regnans ''Eucalyptus regnans'', known variously as mountain ash (in Victoria), giant ash or swamp gum (in Tasmania), or stringy gum, is a species of very tall forest tree that is native to the Australia states of Tasmania and Victoria. It is a straigh ...
'' is the tallest flowering plant in the world. Other important Australian genera are ''
Callistemon ''Callistemon'' is a genus of shrubs in the family Myrtaceae, first described as a genus in 1814. The entire genus is endemic to Australia but widely cultivated in many other regions and naturalised in scattered locations. Their status as a se ...
'' (bottlebrushes), ''
Syzygium ''Syzygium'' () is a genus of flowering plants that belongs to the myrtle family, Myrtaceae. The genus comprises about 1200 species, and has a native range that extends from Africa and Madagascar through southern Asia east through the Pacific. I ...
'', and ''
Melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles, bottlebrushes or tea-trees (although the last name is also applied to species of '' Leptospermum''). They ...
'' (paperbarks). Species of the genus ''
Osbornia ''Osbornia'' is a monotypic genus of mangrove in the myrtle family Myrtaceae. The sole species is ''Osbornia octodonta'', commonly known as the myrtle mangrove, which inhabits coastal areas of Borneo, the Philippines, the Lesser Sunda Islands. ...
'', native to Australasia, are
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 ...
s. ''
Eugenia ''Eugenia'' is a genus of flowering plants in the myrtle family Myrtaceae. It has a worldwide, although highly uneven, distribution in tropical and subtropical regions. The bulk of the approximately 1,100 species occur in the New World tropics, ...
'', ''
Myrcia ''Myrcia'' is a genus of plants in the family Myrtaceae, containing about 765 species as of 2022. They are distributed in Central and South America, Mexico, and the Caribbean, with centers of diversity in the Brazilian Cerrado The Cerrado () ...
'', and ''
Calyptranthes ''Myrcia'' is a genus of plants in the family Myrtaceae, containing about 765 species as of 2022. They are distributed in Central and South America, Mexico, and the Caribbean, with centers of diversity in the Brazilian Cerrado The Cerrado () ...
'' are among the larger genera in the neotropics. Historically, the Myrtaceae were divided into two subfamilies. Subfamily Myrtoideae (about 75 genera) was recognized as having fleshy fruits and opposite, entire leaves. Most genera in this subfamily have one of three easily recognized types of embryos. The genera of Myrtoideae can be very difficult to distinguish in the absence of mature fruits. Myrtoideae are found worldwide in subtropical and tropical regions, with centers of diversity in the Neotropics, northeastern Australia, and Malesia. In contrast, subfamily Leptospermoideae (about 80 genera) was recognized as having dry, dehiscent fruits (capsules) and leaves arranged spirally or alternate. The Leptospermoideae are found mostly in Australasia, with a centre of diversity in Australia. Many genera in Western Australia have greatly reduced leaves and flowers typical of dryer habitats.


Taxonomy

The division of the Myrtaceae into Leptospermoideae and Myrtoideae was challenged by a number of authors, including Johnson and Briggs (1984), who identified 14 tribes or clades within Myrtaceae, and found Myrtoideae to be polyphyletic. Molecular studies by several groups of authors, as of 2008, have confirmed the baccate (fleshy) fruits evolved twice from capsular fruits and, as such, the two-subfamily classification does not accurately portray the phylogenetic history of the family. Thus, many workers are now using a recent analysis by Wilson et al. (2001) as a starting point for further analyses of the family. This study pronounced both Leptospermoideae and Myrtoideae invalid, but retained several smaller suballiances shown to be monophyletic through matK analysis. The genera '' Heteropyxis'' and '' Psiloxylon'' have been separated as separate families by many authors in the past as Heteropyxidaceae and Psiloxylaceae. However, Wilson ''et al.'' included them in Myrtaceae. These two genera are presently believed to be the earliest arising and surviving lineages of Myrtaceae. The most recent classification recognizes 17 tribes and two subfamilies, Myrtoideae and Psiloxyloideae, based on a phylogenetic analysis of
plastid A plastid is a membrane-bound organelle found in the Cell (biology), cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria. Examples of plastids include chloroplasts ...
DNA. Many new species are being described annually from throughout the range of Myrtaceae. Likewise, new genera are being described nearly yearly.


Classification

Following Wilson (2011) Subfamily Psiloxyloideae * tribe Psiloxyleae * tribe Heteropyxideae Subfamily Myrtoideae * tribe
Xanthostemoneae Xanthostemoneae is a tribe in the plant family Myrtaceae.Wilson, P. G. (2011) Myrtaceae. In The Families and Genera of Vascular Plants. Volume X. Sapindales, Cucurbitales, Myrtaceae, edited by K. Kubitzki, X:212–71. Heidelberg: Springer-Verlag, ...
* tribe Lophostemoneae * tribe Osbornieae * tribe Melaleuceae * tribe
Kanieae Kanieae is a tribe in the plant family MyrtaceaeWilson, P. G. (2011) Myrtaceae. In The Families and Genera of Vascular Plants. Volume X. Sapindales, Cucurbitales, Myrtaceae, edited by K. Kubitzki, X:212–71. Heidelberg: Springer-Verlag, 2011. fr ...
* tribe Backhousieae * tribe Metrosidereae * tribe Tristanieae * tribe Syzygieae * tribe
Myrteae Myrteae is the largest tribe in the plant family Myrtaceae. It includes most of the species of the family that have fleshy fruits. Well-known members include edible fruit such as Feijoa sellowiana, feijoa (''Feijoa sellowiana''), Psidium guajava ...
* tribe
Eucalypteae Eucalypteae is a large tribe of flowering plants in the family Myrtaceae; members of this tribe are known as eucalypts. In Australia the genera ''Angophora'', ''Corymbia'', and ''Eucalyptus'' are commonly known as gum trees, for the sticky substa ...
* tribe Syncarpieae * tribe Lindsayomyrteae * tribe
Leptospermeae Leptospermeae is a tribe in the plant family MyrtaceaeWilson, P. G. (2011) Myrtaceae. In The Families and Genera of Vascular Plants. Volume X. Sapindales, Cucurbitales, Myrtaceae, edited by K. Kubitzki, X:212–71. Heidelberg: Springer-Verlag, 20 ...
* tribe
Chamelaucieae Chamelaucieae is a tribe of flowering plants within the family Myrtaceae, mostly from Australia, with a few species in New Caledonia and south-east Asia. Genera include:Barbara L. Rye, Peter G. Wilson, Margaret M. Heslewood, Andrew J. Perkins, K ...


Genera

,
Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
accepts the following 130 genera:


Ecology

Myrtaceae is foraged by many stingless bees, especially by species such as ''
Melipona bicolor ''Melipona bicolor'' , commonly known as Guaraipo or Guarupu, is a Eusociality, eusocial bee found primarily in South America. It is an inhabitant of the Araucaria moist forests, Araucaria Forest and the Atlantic Rainforest, and is most commonly ...
'' which gather pollen from this plant family. Some Australian species such as '' Tetragonula hockingsi'' and '' T. carbonaria'' are also known to collect resin from the mature seed pods of ''
Corymbia torelliana ''Corymbia torelliana'', commonly known as cadaghi or cadaga, is a species of tree that is Endemism, endemic to north Queensland. It has smooth, greenish grey to white bark, rough at the base of older trees, egg-shaped, heart-shaped or lance-sh ...
'', resulting in '' mellitochory'' as the seeds get stuck onto the
corbiculae The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of the female of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive where it is used as food by the colo ...
of the bees and sometimes are successfully disposed of by colony members that remove them. But usually, they get stuck in the hives or near hive entrances instead, hence also making it a minor nuisance for some keepers as they can take up a lot of space. Fortunately, this is only known to occur in the eastern areas of
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, but could occur in other neighbouring countries where some ''
Corymbia ''Corymbia'', commonly known as bloodwoods, is a genus of about one hundred species of tree that, along with ''Eucalyptus'', '' Angophora'' and several smaller groups, are referred to as eucalypts. Until 1990, corymbias were included in the g ...
'' species are native.
Weevils Weevils are beetles belonging to the Taxonomic rank, superfamily Curculionoidea, known for their elongated snouts. They are usually small – less than in length – and Herbivore, herbivorous. Approximately 97,000 species of weevils are known. ...
in the tribe Cryptoplini mostly use Myrtaceae as hosts. Their larvae can develop in flower and fruit buds, or in
galls Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or wart ...
(often galls already formed by other insects).


References


External links

* {{Authority control Myrtales families