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Myrtaceae

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Myrtaceae
Temporal range: Santonian - recent[1][2]
Myrtus communis foliage and flowers
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Myrtales
Family: Myrtaceae
Juss.[3]
Genera

About 130; see list

Pimenta dioica

Myrtaceae (/mərˈteɪsiˌaɪ, -siːˌiː/), the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All species are woody, contain essential oils, and have flower parts in multiples of four or five. The leaves are evergreen, alternate to mostly opposite, simple, and usually entire (i.e., without a toothed margin). The flowers have a base number of five petals, though in several genera, the petals are minute or absent. The stamens are usually very conspicuous, brightly coloured, and numerous.

Evolutionary history

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Scientists hypothesize that the family Myrtaceae arose between 60 and 56 million years ago (Mya) during the Paleocene era. Pollen fossils have been sourced to the ancient supercontinent Gondwana.[4] The breakup of Gondwana during the Cretaceous 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.[5] Generally, experts agree that vicariance is responsible for the differentiation of Myrtaceae taxa, except in the cases of Leptospermum species now located on New Zealand and New Caledonia, islands which may have been submerged at the time of late Eocene differentiation.[4]

Diversity

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Recent estimates suggest the Myrtaceae include about 5,950 species in about 132 genera.[6][7] 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 hotspots. Genera with capsular fruits such as Eucalyptus, Corymbia, Angophora, Leptospermum, and Melaleuca are absent from the Americas, apart from Metrosideros in Chile and Argentina. Genera with fleshy fruits have their greatest concentrations in eastern Australia and Malesia (the Australasian realm) and the Neotropics. Eucalyptus is a dominant, nearly ubiquitous genus in the more mesic parts of Australia and extends north sporadically to the Philippines. Eucalyptus regnans is the tallest flowering plant in the world. Other important Australian genera are Callistemon (bottlebrushes), Syzygium, and Melaleuca (paperbarks). Species of the genus Osbornia, native to Australasia, are mangroves. Eugenia, Myrcia, and Calyptranthes are among the larger genera in the neotropics.

Syzygium samarangense, with a cross-section of the fruit

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 more xeric habitats.

Taxonomy

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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.[8] 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.[9]

The genera Heteropyxis and Psiloxylon have been separated as separate families by many authors in the past as Heteropyxidaceae and Psiloxylaceae.[10][8] However, Wilson et al.[9] 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 DNA.[11]

Many new species are being described annually from throughout the range of Myrtaceae. Likewise, new genera are being described nearly yearly.

Classification

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Following Wilson (2011)[12]

Subfamily Psiloxyloideae

Subfamily Myrtoideae

Genera

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127 genera are currently accepted:[13]

Ecology

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Myrtaceae is foraged by many stingless bees, especially by species such as Melipona bicolor 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, resulting in mellitochory as the seeds get stuck onto the corbiculae 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, but could occur in other neighbouring countries where some Corymbia species are native.[15]

Weevils in the tribe Cryptoplini mostly use Myrtaceae as hosts. Their larvae can develop in flower and fruit buds, or in galls (often galls already formed by other insects).[16]

References

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  1. ^ "Myrtales". www.mobot.org. Retrieved 2023-07-20.
  2. ^ Biffin, Ed; Lucas, Eve J.; Craven, Lyn A.; Ribeiro da Costa, Itayguara; Harrington, Mark G.; Crisp, Michael D. (2010-05-11). "Evolution of exceptional species richness among lineages of fleshy-fruited Myrtaceae". Annals of Botany. 106 (1): 79–93. doi:10.1093/aob/mcq088. ISSN 1095-8290. PMC 2889796. PMID 20462850.
  3. ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
  4. ^ a b Thornhill, Andrew H.; Ho, Simon Y.W.; Külheim, Carsten; Crisp, Michael D. (December 2015). "Interpreting the modern distribution of Myrtaceae using a dated molecular phylogeny". Molecular Phylogenetics and Evolution. 93: 29–43. doi:10.1016/j.ympev.2015.07.007. ISSN 1055-7903. PMID 26211451. S2CID 21263153.
  5. ^ Sytsma, Kenneth J.; Litt, Amy; Zjhra, Michelle L.; Chris Pires, J.; Nepokroeff, Molly; Conti, Elena; Walker, Jay; Wilson, Peter G. (July 2004). "Clades, Clocks, and Continents: Historical and Biogeographical Analysis of Myrtaceae, Vochysiaceae, and Relatives in the Southern Hemisphere". International Journal of Plant Sciences. 165 (S4): S85–S105. doi:10.1086/421066. ISSN 1058-5893. S2CID 62825431.
  6. ^ Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3). Magnolia Press: 201–217. doi:10.11646/phytotaxa.261.3.1.
  7. ^ Govaerts, R. et al. (12 additional authors). 2008. World Checklist of Myrtaceae. Royal Botanic Gardens, Kew. xv + 455 pp.
  8. ^ a b Johnson, L. A. S.; Briggs, B. G. (1984). "Myrtales and Myrtaceae-A Phylogenetic Analysis". Annals of the Missouri Botanical Garden. 71 (3): 700. doi:10.2307/2399159. ISSN 0026-6493. JSTOR 2399159.
  9. ^ a b Wilson, Peter G.; O'Brien, Marcelle M.; Gadek, Paul A.; Quinn, Christopher J. (2001). "Myrtaceae revisited: a reassessment of infrafamilial groups". American Journal of Botany. 88 (11): 2013–2025. doi:10.2307/3558428. JSTOR 3558428. PMID 21669634.
  10. ^ Sytsma, Kenneth J. and Amy Litt. 2002. Tropical disjunctions in and among the Myrtaceae clade (Myrtaceae, Heteropyxidaceae, Psiloxylaceae, Vochysiaceae): Gondwanan vicariance or dispersal? (Abstract). Botany 2002 Conference, University of Wisconsin, Madison, Wisconsin, August 4–7, 2002.
  11. ^ Wilson, P. G.; O'Brien, M. M.; Heslewood, M. M.; Quinn, C. J. (2005). "Relationships within Myrtaceae sensu lato based on a matK phylogeny". Plant Systematics and Evolution. 251 (1): 3–19. Bibcode:2005PSyEv.251....3W. doi:10.1007/s00606-004-0162-y. S2CID 23470845.
  12. ^ 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, 2011.
  13. ^ Myrtaceae Juss. Plants of the World Online. Retrieved 30 June 2023.
  14. ^ Pigg, K. B.; Stockey, R. A.; Maxwell, S. L. (1993). ""Paleomyrtinaea", a new genus of permineralized myrtaceous fruits and seeds from the Eocene of British Columbia and Paleocene of North Dakota". Canadian Journal of Botany. 71 (1): 1–9. doi:10.1139/b93-001.
  15. ^ Hilário, S.D.; Imperatriz-Fonseca, V.L. (2009). "Pollen foraging in colonies of Melipona bicolor (Apidae, Meliponini): effects of season, colony size and queen number". Genetics and Molecular Research. 8 (2): 664–671. doi:10.4238/vol8-2kerr029. PMID 19554765.
  16. ^ Jennings, Debbie; Oberprieler, Rolf (2018-07-23). "A Review of the Tribe Cryptoplini (Coleoptera: Curculioninae), with Revision of the Genus Menechirus Hartmann, 1901 and Description of a New Genus Associated with Macadamia". Diversity. 10 (3): 71. doi:10.3390/d10030071. ISSN 1424-2818.
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