Myrtaceae

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids II
Ordo: Myrtales

Familia: Myrtaceae
Subfamiliae (2): <a href="Myrtoideae.html">Myrtoideae</a> – Psiloxyloideae

Genera: Acca – Accara – Actinodium – Agonis – Algrizea – Allosyncarpia – Aluta – Amomyrtella – Amomyrtus – Angasomyrtus – Angophora – Archirhodomyrtus – Arillastrum – Astartea – Asteromyrtus – Astus – Austromyrtus – Babingtonia – Backhousia – Baeckea – Balaustion – Barongia – Basisperma – Beaufortia – Blepharocalyx – Callistemon – Calothamnus – Calycolpus – Calycorectes – Calyptranthes – Calyptrogenia – Calythropsis – Calytrix – Campomanesia – Chamelaucium – Chamguava – Cheyniana – Choricarpia – Cloezia – Conothamnus – Corymbia – Corynanthera – Cupheanthus – Curitiba – Darwinia – Decaspermum – Eremaea – Eucalyptopsis – Eucalyptus – Eugenia – Euryomyrtus – Gomidesia – Gossia – Harmogia – Heteropyxis – Hexachlamys – Homalocalyx – Homalospermum – Homoranthus – Hottea – Hypocalymma – Kania – Kanakomyrtus – Kardomia – Kjellbergiodendron – Kunzea – Lamarchea – Legrandia – Lenwebbia – Leptospermum – Lindsayomyrtus – Lithomyrtus – Lophomyrtus – Lophostemon – Luma – Lysicarpus – Malleostemon – Marlierea – Melaleuca – Metrosideros – Micromyrtus – Mitranthes – Mitrantia – Mosiera – Myrceugenia – Myrcia – Myrcianthes – Myrciaria – Myrrhinium – Myrtastrum – Myrtella – Myrteola – Myrtus – Neofabricia – Neomitranthes – Neomyrtus – Ochrosperma – Octamyrtus – Osbornia – Paramyrciaria – Pericalymma – Petraeomyrtus – Phymatocarpus – Pileanthus – Pilidiostigma – Piliocalyx – Pimenta – Pleurocalyptus – Plinia – Pseudanamomis – Psidium – Psiloxylon – Purpureostemon – Regelia – Rhodamnia – Rhodomyrtus – Rinzia – Ristantia – Sannantha – Schizocalomyrtus – Scholtzia – Seorsus – Siphoneugena – Sphaerantia – Stenostegia – Stereocaryum – Stockwellia – Syncarpia – Syzygium – Taxandria – Thaleropia – Thryptomene – Triplarina – Tristania – Tristaniopsis – Ugni – Uromyrtus – Verticordia – Welchiodendron – Whiteodendron – Xanthomyrtus – Xanthostemon

Name

Myrtaceae Juss. (1789)

Type genus: Myrtus L.

Synonyms

Heterotypic
Heteropyxidaceae Engl. & Gilg in Engl., Syllabus, ed. 8, 281. (1920) nom. cons.
Type genus: Heteropyxis Harv.
Psiloxylaceae Croizat, Principia Bot. 604, 1161, 1164. (1960)
Type genus: Psiloxylon Thouars ex Tul.

References

Jussieu, A.L. de 1789. Genera plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. 498 pp. Paris: Herissant et Theophile Barrois. BHL Reference page.
Biffin, E., Lucas, E.J., Craven, L.A., da Costa, I.R., Harrington, M.G. & Crisp, M.D. 2010. Evolution of exceptional species richness among lineages of fleshy-fruited Myrtaceae. Annals of Botany 106(1): 79–93. DOI: 10.1093/aob/mcq088
Govaerts, R. et al. 2016. Myrtaceae in World Checklist of Selected Plant Families. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the internet. Accessed: 2016 Mar. 31. Reference page.
Lucas, E.J., Belsham, S.R., Lughadha, E.N., Orlovich, D.A., Sakuragui, C.M., Chase, M.W. & Wilson, P.G. 2005. Phylogenetic patterns in the fleshy-fruited Myrtaceae–preliminary molecular evidence. Plant Systematics and Evolution, 251(1): 35–51. DOI: 10.1007/s00606-004-0164-9 Reference page.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.
Thornhill, A.H. Wilson, P.G., Drudge, J., Barrett, M.D., Hope, G.S., Craven, L.A. & Crisp, M.D. 2012. Pollen morphology of the Myrtaceae. Part 3: tribes Chamelaucieae, Leptospermeae and Lindsayomyrteae. Australian Journal of Botany 60(3): 225-259. DOI: 10.1071/BT11176 Reference page.
Tornabene, M.W. & Wagner, W.L. 2013. New combinations for Pacific endemic species: Marquesan Poaceae, and Micronesian Myrtaceae. PhytoKeys 28: 1-7. DOI: 10.3897/phytokeys.28.6139 Reference page.
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. DOI: 10.1007/s00606-004-0162-y Reference page.

Vernacular names
español: Mirtáceas
suomi: Myrttikasvit
Nordfriisk: Myrtenplaanten
français: Myrtacées
日本語: フトモモ科
македонски: Мирти
Nederlands: Mirteachtigen
русский: Миртовые
Türkçe: Mersingiller
українська: Миртові
中文: 桃金娘科

Myrtaceae or 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

Scientists hypothesize that the family Myrtaceae arose between sixty and fifty-six million years ago during the Paleocene era. Pollen fossils have been sourced to the ancient supercontinent Gondwana.[2] 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.[3] 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.[2]

Diversity

Recent estimates suggest the Myrtaceae include approximately 5,950 species in about 132 genera.[4][5] 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

The division of 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.[6] 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.[7]

The genera Heteropyxis and Psiloxylon have been separated as separate families by many authors in the past as Heteropyxidaceae and Psiloxylaceae.[8][6] However, Wilson et al.[7] 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.[9]

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)[10]

Subfamily Psiloxyloideae

tribe Psiloxyleae
tribe Heteropyxideae

Subfamily Myrtoideae

tribe Xanthostemoneae
tribe Lophostemoneae
tribe Osbornieae
tribe Melaleuceae
tribe Kanieae
tribe Backhousieae
tribe Metrosidereae
tribe Tristanieae
tribe Syzygieae
tribe Myrteae
tribe Eucalypteae
tribe Syncarpieae
tribe Lindsayomyrteae
tribe Leptospermeae
tribe Chamelaucieae

Genera

Abbevillea
Actinodium
Agonis
Algrizea
Allosyncarpia
Aluta
Amomyrtella
Amomyrtus
Angasomyrtus
Angophora
Archirhodomyrtus
Arillastrum
Astartea
Asteromyrtus
Astus
Austromyrtus
Babingtonia
Backhousia
Baeckea
Balaustion
Barongia
Basisperma
Beaufortia
Blepharocalyx
Callistemon
Calothamnus
Calycolpus
Calycorectes
Calyptranthes
Calyptrogenia
Calytrix
Campomanesia
Chamelaucium
Chamguava
Cheyniana
Choricarpia
Cleistocalyx
Cloezia
Conothamnus
Corymbia
Corynanthera
Curitiba
Cyathostemon
Darwinia
Decaspermum
Enekbatus
Eremaea
Ericomyrtus
Eucalyptopsis
Eucalyptus
Eugenia
Euryomyrtus
Gomidesia
Gossia
Harmogia
Heteropyxis
Hexachlamys
Homalocalyx
Homalospermum
Homoranthus
Hottea
Hypocalymma
Kanakomyrtus
Kania
Kardomia
Kjellbergiodendron
Kunzea
Lamarchea
Legrandia
Lenwebbia
Leptospermum
Lindsayomyrtus
Lithomyrtus
Lophomyrtus
Lophostemon
Luma
Lysicarpus
Malleostemon
Marlierea
Melaleuca
Meteoromyrtus
Metrosideros
Micromyrtus
Mitranthes
Mitrantia
Mosiera
Myrceugenia
Myrcia
Myrcianthes
Myrciaria
Myrrhinium
Myrtastrum
Myrtella
Myrteola
Myrtus
Neofabricia
Neomitranthes
Neomyrtus
Ochrosperma
Octamyrtus
Osbornia
†Paleomyrtinaea[11]
Paragonis
Pericalymma
Phymatocarpus
Pileanthus
Pilidiostigma
Pimenta
Pleurocalyptus
Plinia
Pseudanamomis
Psidium
Psiloxylon
Purpureostemon
Regelia
Rhodamnia
Rhodomyrtus
Rinzia
Ristantia
Sannantha
Scholtzia
Seorsus
Siphoneugena
Sphaerantia
Stenostegia
Stereocaryum
Stockwellia
Syncarpia
Syzygium
Taxandria
Thaleropia
Thryptomene
Triplarina
Tristania
Tristaniopsis
Ugni
Uromyrtus
Verticordia
Waterhousea
Welchiodendron
Whiteodendron
Xanthomyrtus
Xanthostemon

Foraging

Myrtaceae is foraged by many stingless bees, especially by the species Melipona bicolor which gather pollen from this plant family.[12]
References

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.
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.
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.
Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. Magnolia Press. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
Govaerts, R. et al. (12 additional authors). 2008. World Checklist of Myrtaceae. Royal Botanic Gardens, Kew. xv + 455 pp.
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.
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.
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.
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: 3–19. doi:10.1007/s00606-004-0162-y. S2CID 23470845.
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.
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.
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.