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Chloranthales: diferenças entre revisões

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{{Info/Taxonomia
{{Info/Taxonomia
| nome = Clorantáceas
| nome = Chloranthales<br><small>Chloranthaceae</small>
| período_fóssil = {{fossil range|Barremiano|presente}}<small>[[Barremiano]] - recente</small><ref name=mobot>{{Cite web |title=Chloranthales|url=https://www.mobot.org/mobot/research/apweb/orders/chloranthalesweb.htm|access-date=2023-07-20|website=www.mobot.org}}</ref>
| cor = lightgreen
| imagem = Sarcandra_glabra1.jpg
| imagem = Sarcandra_glabra1.jpg
| imagem_legenda = ''Sarcandra glabra'' em fruto
| imagem_legenda = ''Sarcandra glabra'' em fruto
Linha 8: Linha 8:
| classe = [[Magnoliopsida]]
| classe = [[Magnoliopsida]]
| ordem = [[Chloranthales]]
| ordem = [[Chloranthales]]
| ordem_autoridade=[[Robert Brown (botanist, born 1773)|R.Br.]]<ref name="apgiii">{{Cite journal |last=Angiosperm Phylogeny Group III |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free }}</ref>
| família = '''Chloranthaceae'''
| família = '''Chloranthaceae'''
| família_autoridade = [[Robert Brown|R. Br.]] ex [[John Lindley|Lindl.]], 1821
| família_autoridade = [[R.Br.]] ''ex'' [[Sims]]<ref name="apgiii" />
| subdivisão_nome = Gêneros
| subdivisão_nome = Géneros
| subdivisão =
| subdivisão =
* ''[[Ascarina]]'' [[Johann Reinhold Forster|J.R. Forster]] & [[Georg Forster|G. Forster]], 1775
* ''[[Ascarina]]'' <small>[[Johann Reinhold Forster|J.R. Forster]] & [[Georg Forster|G. Forster]], 1775</small>
* ''[[Chloranthus]]'' [[Olof Peter Swartz|Sw.]], 1787
* ''[[Chloranthus]]'' <small>[[Olof Peter Swartz|Sw.]], 1787</small>
* ''[[Hedyosmum]]'' [[Olof Peter Swartz|Sw.]], 1788
* ''[[Hedyosmum]]'' <small>[[Olof Peter Swartz|Sw.]], 1788</small>
* ''[[Sarcandra]]'' [[George Gardner|Gardner]], 1846
* ''[[Sarcandra]]'' <small>[[George Gardner|Gardner]], 1846</small>
| sinônimos =
| sinónimos =
* Hedyosmaceae
* [[Hedyosmaceae]]
* [http://biopl-a-181.plantbio.cornell.edu/]
| sinónimos_ref=<ref>[http://biopl-a-181.plantbio.cornell.edu/ Chloranthaceae]</ref>
}}
}}
[[Ficheiro:Sarcandra glabra2.jpg|thumb|Frutos de ''S. glabra'']]
[[File:Chloranthus fortunei 2.jpg|thumb| ''[[Chloranthus fortunei]]''.]]
[[Ficheiro:Sarcandra glabra2.jpg|thumb|Frutos de ''[[Sarcandra glabra]]'']]
[[Ficheiro:Sarcandra glabra3.jpg|thumb|Flores masculinas de ''S. glabra'']]

As '''clorantáceas''' ('''Chloranthaceae''', ''nom. cons.'') são a única família de [[angiosperma]]s da ordem '''Chloranthales''' (clado independente, segundo o APG IV (2016)). É constituída por quatro [[Género (biologia)|gêneros]] com 80 [[espécie]]s, distribuídas principalmente pelo sudeste da [[Ásia]], ilhas do [[oceano Pacífico]] até o [[arquipélago das Marquesas]] e [[Nova Zelândia]], [[Madagáscar]], [[América Central|Américas central]] e do [[América do Sul|sul]] e [[Caribe]].
As '''clorantáceas''' ('''Chloranthaceae''', ''nom. cons.'') são a única família de [[angiosperma]]s da ordem '''Chloranthales''' (clado independente, segundo o APG IV (2016)). É constituída por quatro [[Género (biologia)|gêneros]] com 80 [[espécie]]s, distribuídas principalmente pelo sudeste da [[Ásia]], ilhas do [[oceano Pacífico]] até o [[arquipélago das Marquesas]] e [[Nova Zelândia]], [[Madagáscar]], [[América Central|Américas central]] e do [[América do Sul|sul]] e [[Caribe]].


== Descrição ==
== Descrição ==
[[Ficheiro:Sarcandra glabra3.jpg|thumb|left|Flores masculinas de ''S. glabra'']]

* Hábito: plantas [[Árvore|arbóreas]], às vezes grandes, [[arbusto|arbustivas]], [[subarbusto|subarbustivas]] ou [[erva|herbáceas]], aromáticas, com células de [[Óleo essencial|óleos essenciais]].
* Hábito: plantas [[Árvore|arbóreas]], às vezes grandes, [[arbusto|arbustivas]], [[subarbusto|subarbustivas]] ou [[erva|herbáceas]], aromáticas, com células de [[Óleo essencial|óleos essenciais]].
* [[Folha (botânica)|Folhas]]: perenes, conduplicadas, simples, pininérvias, secundariamente broquidódromas a eucamptódromas ou semicraspedódromas, dentadas, decussadas, [[Glabro|glabras]] ou com [[Tricoma#Tipos de tricoma|tricomas glandulares]], uni-a multisseriados, [[pecíolo]]s mais ou menos presos à base, formando uma [[bainha]] diferenciada ao longo do [[caule]], [[estípula]]s presentes, interpeciolares, na borda da bainha. [[Estômato]]s somente na epiderme inferior das folhas (parte abaxial), laterocíticos, paracíticos, enciclocíticos, às vezes anficíclicos, actinocíclicos ou tendentes a anomocíticos.
* [[Folha (botânica)|Folhas]]: perenes, conduplicadas, simples, pininérvias, secundariamente broquidódromas a eucamptódromas ou semicraspedódromas, dentadas, decussadas, [[Glabro|glabras]] ou com [[Tricoma#Tipos de tricoma|tricomas glandulares]], uni-a multisseriados, [[pecíolo]]s mais ou menos presos à base, formando uma [[bainha]] diferenciada ao longo do [[caule]], [[estípula]]s presentes, interpeciolares, na borda da bainha. [[Estômato]]s somente na epiderme inferior das folhas (parte abaxial), laterocíticos, paracíticos, enciclocíticos, às vezes anficíclicos, actinocíclicos ou tendentes a anomocíticos.
Linha 71: Linha 73:
:: ''[[Hedyosmum]]'' [[Olof Peter Swartz|Sw.]], 1788
:: ''[[Hedyosmum]]'' [[Olof Peter Swartz|Sw.]], 1788
::: [[América Central|Américas central]] e [[América do Sul|do sul]], [[Caribe]], 43 espécies; sudeste da [[Ásia]], 1 espécie.
::: [[América Central|Américas central]] e [[América do Sul|do sul]], [[Caribe]], 43 espécies; sudeste da [[Ásia]], 1 espécie.
<!----------------


'''Chloranthaceae''' {{IPAc-en|ˌ|k|l|ɔər|æ|n|ˈ|θ|eɪ|ʃ|iː}} is a [[Family (biology)|family]] of [[flowering plant]]s (angiosperms), the only family in the [[Order (biology)|order]] '''Chloranthales'''.<ref name="apgiii"/> It is not closely related to any other family of flowering plants, and is among the early-diverging lineages in the angiosperms. They are woody or weakly woody plants occurring in Southeast Asia, the Pacific, Madagascar, Central and South America, and the West Indies. The family consists of four extant genera, totalling about 77 known species according to Christenhusz and Byng in 2016.<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | doi-access = free }}</ref> Some species are used in traditional medicine. The type genus is ''[[Chloranthus]]''. The fossil record of the family, mostly represented by pollen such as ''Clavatipollenites,'' extends back to the dawn of the history of flowering plants in the [[Early Cretaceous]], and has been found on all continents.<ref>{{Cite journal|last1=Pipo|first1=M.|last2=Iglesias|first2=Ari|last3=Bodnar|first3=Josefina|date=2020|title=New vesselless angiosperm stem with a cambial variant from the Upper Cretaceous of Antarctica|url=http://www.app.pan.pl/article/item/app006972019.html|journal=Acta Palaeontologica Polonica|volume=65|doi=10.4202/app.00697.2019|doi-access=free}}</ref>

==Description==
Chloranthaceae are [[Essential oil|fragrant]] [[shrub]]s or [[herbaceous plant]]s, that only produce side branches on the new growth.{{explain|date=February 2023}} The stems are mostly cylindrical, with solid [[Plant stem|internodes]], thickened [[Plant stem|nodes]] in many species, that carry evergreen leaves [[Leaf#Arrangement on the stem|arranged in pairs on opposite sides]] of the stem, with [[stipule]]s that have merged with that of the opposing leaf. The small [[flower]]s are seated directly on the axis of the [[inflorescence]]. [[Petal]]s are absent in this family, and sometimes so are [[sepal]]s. The flowers can be either hermaphrodite or of separate sexes. The fruit is a [[drupe]] or [[Berry (botany)|berry]], consisting of one [[carpel]].<ref name=MOBOT>{{cite web|website= Angiosperm Phylogeny Website|title= Chloranthaceae|author= Stevens, P.F.|year= 2001|orig-year= onwards|url= http://www.mobot.org/MOBOT/research/APweb/orders/chloranthalesweb.htm#Chloranthaceae|access-date= 2016-06-12}}</ref>

===Differences between the genera===
The four genera assigned to this family can be distinguished from each other by the following characters:<!--Note: wiki-linking the genera again, as it seems to improve article navigation-->

''[[Sarcandra]]'' species are shrubs whose wood lacks [[Vessel element|vessels]]. They have [[perfect flower|bisexual flowers]], with only one, club-shaped, [[stamen]], in which the connective tissue (between the [[anther]] lobes which carry the pollen) is wide, and with a smooth and moist [[Stigma (botany)|stigma]]. This genus has four species, which occur in Malaysia, China, Indochina, Japan, India, and Sri Lanka.<br>

''[[Chloranthus]]'' species are dwarf shrubs or [[herbaceous plant]]s, with [[xylem]] that contains vessels. Flowers are bisexual, each of which bears three stamens on straight filaments with three anther lobes and a wide [[Stamen#Morphology and terminology|connective]], and with a smooth and moist [[Stigma (botany)|stigma]]. The 20 species occur in southern and eastern Asia.<br>

''[[Ascarina]]'' has separate male and female flowers. The male flowers are subtended by two [[bract]]s and have between one and five stamens, in which the connective is not widened. The female flower is without bracts, the stigma is dry and covered in [[Stigma (botany)#Description|papillae]]. The fruit is a [[drupe]]-like [[berry (botany)|berry]]. There are 12 species, which occur on islands in the Pacific and insular South-East Asia, from New Zealand and the [[Marquesas]] to [[Borneo]], and on [[Madagascar]].<br>

''[[Hedyosmum]]'' has separate male and female flowers. The male flowers are without bracts and have one stamen, in which the connective is not widened. The female flower is without bracts; the stigma is dry and covered in [[Stigma (botany)#Description|papillae]]. Female flowers have a 3-lobed [[Sepal|calyx]]. The fruit, a drupe, has a kernel with a hard and woody shell. There are 43 species found in Latin America, including the [[Antilles]], as well as one species found in Southeast Asia.

==Taxonomy==

The Chloranthaceae have been recognised as a family in most classifications but without clear relatives. [[Molecular systematics]] studies have shown that it is not closely related to any other family and is among the [[Flowering_plant#Phylogeny|early-diverging lineages in the angiosperms]]. In particular, it is neither a [[eudicot]] nor a [[monocot]]. Fossils assigned to Chloranthaceae, or closely related to the family, are among the oldest angiosperms known.<ref>{{Cite journal|last1=Herendeen|first1=Patrick S.|last2=Friis|first2=Else Marie |last3=Pedersen |first3=Kaj Raunsgaard|last4=Crane|first4=Peter R.|date=2017-03-03|title=Palaeobotanical redux: revisiting the age of the angiosperms |url=https://www.nature.com/articles/nplants201715|journal=Nature Plants |volume=3|issue=3 |page=17015 |doi=10.1038/nplants.2017.15 |pmid=28260783 |s2cid=205458714 |issn=2055-0278}}</ref><ref>{{Cite journal|last1=Doyle|first1=James A.|last2=Endress |first2=Peter K. |date=2018-06-01|title=Phylogenetic Analyses of Cretaceous Fossils Related to Chloranthaceae and their Evolutionary Implications |journal=The Botanical Review|language=en|volume=84 |issue=2|pages=156–202 |doi=10.1007/s12229-018-9197-6 |s2cid=46980346 |issn=1874-9372 |url=https://escholarship.org/uc/item/0d58r5r0}}</ref><ref>{{Cite journal|last1=Friis|first1=Else Marie|last2=Crane|first2=Peter R. |last3=Pedersen |first3=Kaj Raunsgaard|date=2019-03-01|title=Hedyosmum-Like Fossils in the Early Cretaceous Diversification of Angiosperms |journal=International Journal of Plant Sciences|volume=180|issue=3|pages=232–239|doi=10.1086/701819|s2cid=91649388 |issn=1058-5893 |url=https://www.dora.lib4ri.ch/psi/islandora/object/psi%3A20566}}</ref> The [[APG II system]] (2003) left the family unplaced as to order, but the [[APG III system]] (2009) accepted [[Chloranthales]], [[monotypy|containing only this family]].<ref name="apgiii"/> The cladogram below, from the [[APG IV system]] (2016), shows the Chloranthales in a [[Speciation#Genetics|trichotomy]] with the [[magnoliids]] and the monocot-Ceratophyllales-dicot [[clade]]. Earlier, the order was grouped with magnoliids, but studies in 2014 did not support this placement, leaving its phylogeny unclear.<ref name=APG4>{{Cite journal |last=Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=Botanical Journal of the Linnean Society |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |doi-access=free }}</ref> A 2021 study sequenced the ''Chloranthus'' genome and found Chloranthales as sister to magnoliids.<ref name="Guo2021">{{cite journal |last1=Guo |first1=Xing |title=Chloranthus genome provides insights into the early diversification of angiosperms |journal=Nature Communications |date=26 November 2021 |volume=12 |issue=1 |page=6930 |doi=10.1038/s41467-021-26922-4 |pmid=34836973 |pmc=8626473 |doi-access=free }}</ref>


{{clade |style=font-size:100%;line-height:110%;
|label1=[[angiosperms]]
|1={{clade
|1=[[Amborellales]]
|2={{clade
|1=[[Nymphaeales]]
|2={{clade
|1=[[Austrobaileyales]]
|2={{clade
|1={{clade
|1=[[magnoliids]]
|2='''Chloranthales'''
}}
|2={{clade
|1=[[monocots]]
|2={{clade
|1=[[Ceratophyllales]]
|2=[[eudicots]]
}}
}}
}}
}}
}}
}}
}}

A 2004 study based on comparisons of homologous DNA fragments indicated that both the family Chloranthaceae and its extant genera ''Ascarina'', ''Chloranthus'', ''Hedyosmum'' and ''Sarcandra'' are probably [[Monophyly|monophyletic]], with ''Hedyosmum'' being the first to diverge from the rest, and ''Ascarina'' being the [[sister group]] of the clade consisting of ''Sarcandra'' and ''Chloranthus''. {{As of|2016|June}}, these four extant genera are recognized, and insights into their relationships are expressed in the [[cladogram|tree]] below:<ref name=MOBOT /><ref>{{cite journal|first1= Helena|last1= Eklund|first2= James A.|last2= Doyle|first3= Patrick S.|last3= Herendeen|year= 2004|title= Morphological Phylogenetic Analysis of Living and Fossil Chloranthaceae|journal= International Journal of Plant Sciences|volume= 165|issue= 1|pages= 107–151|jstor= 10.1086/380987|doi=10.1086/380987|s2cid= 84746116}}</ref>

{{clade |style=font-size:100%;line-height:110%;
|label1='''Chloranthaceae'''
|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1=''[[Chloranthus]]''
|2=''[[Sarcandra]]''
}}
|2=''[[Ascarina]]''
}}
|2=''[[Hedyosmum]]''
}}
}}
}}

The extinct genus ''[[Chloranthistemon]]'' also belongs to this family.<ref>{{cite journal | last1 = Herendeen | first1 = P. S. | last2 = Crepet | first2 = W. L. | last3 = Nixon | first3 = K. C. | year = 1993 | title = Chloranthus-like stamens from the Upper Cretaceous of New Jersey | journal = American Journal of Botany | volume = 80 | issue = 8| pages = 865–871 | doi=10.1002/j.1537-2197.1993.tb15306.x}}</ref>

===Historical classifications===
The [[Cronquist system]] (1981) assigned the family
: to the order [[Piperales]]
:: in subclass [[Magnoliids|Magnoliidae]]
::: in class [[Magnoliopsida]] [=[[dicotyledons]]]
:::: of division [[Magnoliophyta]] [=angiosperms].

The [[Thorne system (1992)]] placed it
: in the order [[Magnoliales]], which was assigned
:: to superorder [[Magnolianae]]
::: in subclass Magnoliideae [=dicotyledons],
:::: in class Magnoliopsida [=angiosperms].

The [[Dahlgren system]] raised the family to be
: its own order [[Chloranthales]], which was assigned
:: to superorder Magnolianae
::: in subclass Magnoliideae [=dicotyledons],
:::: in class Magnoliopsida [=angiosperms].

-------------->
==Referências==
==Referências==
{{Reflist}}
==Bibliografia==
* {{((en))}} Angiosperm Phylogeny Group (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. ''Botanical Journal of the Linnean Society'' '''141''': 399-436. (Disponível online: [https://web.archive.org/web/20071222063800/http://www.blackwell-synergy.com/links/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/full/ Texto completo (HTML)] | [https://web.archive.org/web/20190912214437/http://www.blackwell-synergy.com/links/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/pdf Texto completo (PDF)])
* {{((en))}} Angiosperm Phylogeny Group (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. ''Botanical Journal of the Linnean Society'' '''141''': 399-436. (Disponível online: [https://web.archive.org/web/20071222063800/http://www.blackwell-synergy.com/links/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/full/ Texto completo (HTML)] | [https://web.archive.org/web/20190912214437/http://www.blackwell-synergy.com/links/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/pdf Texto completo (PDF)])
* {{((en))}} [https://web.archive.org/web/20060927212130/http://www.itis.usda.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=18063 Classificação das dicotiledóneas - ITIS 18063]
* {{((en))}} [https://web.archive.org/web/20060927212130/http://www.itis.usda.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=18063 Classificação das dicotiledóneas - ITIS 18063]
Linha 79: Linha 171:
* Todzia, C.A. 1993. Chloranthaceae. En: Kubitzki, K., Rohwer, J.G. & Bittrich, V. (Editores). The Families and Genera of Vascular Plants. II. Flowering Plants - Dicotyledons. Springer-Verlag.
* Todzia, C.A. 1993. Chloranthaceae. En: Kubitzki, K., Rohwer, J.G. & Bittrich, V. (Editores). The Families and Genera of Vascular Plants. II. Flowering Plants - Dicotyledons. Springer-Verlag.
* Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 29th July 2006. http://delta-intkey.com
* Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 29th July 2006. http://delta-intkey.com

== Ligações externas ==
== Ligações externas ==
{{commonscat|Chloranthaceae}}
{{Correlatos
|commonscat = Chloranthaceae
{{Wikispecies|Chloranthaceae}}
|wikispecies = Chloranthaceae
}}
* [http://digitalis.mobot.org/mrsid/073/07300003_002.jpg ''Chloranthus japonicus'']
* [http://digitalis.mobot.org/mrsid/073/07300003_002.jpg ''Chloranthus japonicus'']
* [http://digitalis.mobot.org/mrsid/073/MOA-02053_001.jpg Inflorescencia de ''Hedyosmum racemosum'']
* [http://digitalis.mobot.org/mrsid/073/MOA-02053_001.jpg Inflorescencia de ''Hedyosmum racemosum'']
* [http://www.phylodiversity.net/borneo/delta/Itemscan/genascar.gif ''Ascarina philippinensis'']
* [http://www.phylodiversity.net/borneo/delta/Itemscan/genascar.gif ''Ascarina philippinensis'']
* [http://www.mobot.org/MOBOT/Research/APweb/maps/chloranthaceaemap.gif Mapa]
* [http://www.mobot.org/MOBOT/Research/APweb/maps/chloranthaceaemap.gif Mapa]
* [http://delta-intkey.com/angio/www/chlorant.htm Chloranthaceae] in L. Watson and M.J. Dallwitz (1992 onwards) ''[https://web.archive.org/web/20101213041459/http://delta-intkey.com/angio/ The families of flowering plants:] descriptions, illustrations, identification, information retrieval''. Version: 3 May 2006. https://web.archive.org/web/20070103200438/http://delta-intkey.com/.
* [http://www.efloras.org/florataxon.aspx?flora_id=201&taxon_id=10187 ''Andes Trees'']
* [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=16737&lvl=3&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]
* [https://web.archive.org/web/20070930183621/http://www.alpine-plants-jp.com/art/index_senryouka.htm Images of ''Chloranthus''] Flavon's art gallery: Chloranthaceae
* [https://web.archive.org/web/20070928202208/http://www.mygarden.me.uk/Chloranthus%20oldhamii%20BSWJ6788.jpg Image of ''Chloranthus oldhamii'']
* [http://planteliste.net/slides/chloranthus_sp.html Image of ''Chloranthus serratus'']
* [https://web.archive.org/web/20070928075927/http://biotech.tipo.gov.tw/plantjpg/1/Sarcandra%20glabra.jpg Images of ''Sarcandra glabra'']


{{Taxonbar|from=Q133208}}
{{APG IV}}
{{APG IV}}
{{Bases de dados taxonómicos}}


[[Categoria:Chloranthaceae| ]]
[[Categoria:Chloranthaceae| ]]

Revisão das 15h49min de 13 de setembro de 2023

Como ler uma infocaixa de taxonomiaChloranthales
Chloranthaceae
Ocorrência: Barremiano–presente

Barremiano - recente[1]

Sarcandra glabra em fruto
Sarcandra glabra em fruto
Classificação científica
Reino: Plantae
Divisão: Magnoliophyta
Classe: Magnoliopsida
Ordem: Chloranthales
R.Br.[2]
Família: Chloranthaceae
R.Br. ex Sims[2]
Géneros
Sinónimos[3]
Chloranthus fortunei.
Frutos de Sarcandra glabra
Flores masculinas de S. glabra

As clorantáceas (Chloranthaceae, nom. cons.) são a única família de angiospermas da ordem Chloranthales (clado independente, segundo o APG IV (2016)). É constituída por quatro gêneros com 80 espécies, distribuídas principalmente pelo sudeste da Ásia, ilhas do oceano Pacífico até o arquipélago das Marquesas e Nova Zelândia, Madagáscar, Américas central e do sul e Caribe.

Descrição

Ecologia

São encontradas nas florestas úmidas desde as altitudes mais baixas até o alto das montanhas. As espécies de Hedyosmum e Ascarina apresentam aparentemente polinização anemófila, por possuírem flores sem cheiro e apresentarem longa inflorescência masculina com anteras grandes e estigmas grandes e úmidos. As espécies de Chloranthus e Sarcandra, ao contrário, com flores geralmente perfumadas, hermafroditas e de anteras pequenas, com estigmas pequenos e secos, são provavelmente entomófilas. Os frutos parecem apresentar ornitocoria.

Fitoquímica

Grande presença de óleos essenciais em idioblastos, lactonas sesquiterpênicas (do tipo germanacrólido, eudesmanólido), derivados O- e C-glicosílados de flavonas e flavonóis comuns, cumarina e N-(feniletil)-cinamoil-amidas presentes. Proantocianidinas ausentes.

Usos

Algunas espécies produzem substâncias de uso na farmacopeia traditional e moderna.

Posição sistemática

A posição sistemática desta família tem sido objeto de muita controvérsia. Em classificações recentes, tem sido colocada em Piperales, Magnoliales ou Laurales, mas considera-se que, caso exista, a maior relação parece ocorrer com Trimeniaceae, baseado em várias características. Considera-se que Sarcandra e Chloranthus estão evolutivamente mais relacionadas entre si morfológicamente que com as outras dos gêneros, que, por sua vez, formariam um outro par evolutivamente relacionado. Entretanto, as análises moleculares preconizam uma relação do tipo (Hedyosmum(Ascarina(Chloranthus, Sarcandra))). O APW (Angiosperm Phylogeny Website) considera que trata-se da única família da ordem Chloranthales (cf. AP-website).

Fósseis

Androceus fossilizados são preservados do Albiano superior no leste da América do Norte e do Santoniano superior-Campaniano no sul da Suécia (Chloranthistemon) intermediários entre Sarcandra e Chloranthus. As cutículas foliares de angiospermas do Cretáceo inferior parecem corresponder a ancestrais das atuais Chloranthaceae. Também têm sido atribuído a esta família alguns dos pólens fossilizados mais antigos conhecidos (Cretáceo inferior): Clavatipollenites do Barremiano, Asteropollis e Stephanocolpites do Albiano (até 125 milhões de anos). A abundância de Clavatipollenites e Asteropollis no Cretáceo inferior médio pode indicar que esses antepassados das Chloranthaceae foram os primeiros angiospermas anemófilos. O pólen de Asteropollis é atribuído a Hedyosmum.

Táxons incluídos

Introdução teórica em Taxonomia

Os quatro gêneros da família são os seguintes:

Sarcandra Gardner, 1846
Malásia, China, Indochina, Japão, Índia, Sri Lanka. 4 espécies.
  • Ervas ou subarbustos. Caules com vasos na xilema. Estames 3, presos em uma escama trilobulada. Pólen 4-6-colpado.
Chloranthus Sw., 1787
Ásia; 20 espécies.
  • Flores unissexuadas. Conectivo não expandido. Pólen monossulcado ou 5-6-tomossulcado. Estigmas papilosos, secos. Geralmente arbustos ou árvores, raramente ervas.
Ascarina J.R. Forster & G. Forster, 1775
Ilhas do Pacífico, desde Nova Zelândia e o arquipélago das Marquesas até Bornéu, e Madagáscar; 12 espécies.
  • Flores masculinas sem brácteas. Estame 1. Flores femininas com perianto trilobulado. Fruto em drupa.
Hedyosmum Sw., 1788
Américas central e do sul, Caribe, 43 espécies; sudeste da Ásia, 1 espécie.

Sarcandra species are shrubs whose wood lacks vessels. They have bisexual flowers, with only one, club-shaped, stamen, in which the connective tissue (between the anther lobes which carry the pollen) is wide, and with a smooth and moist stigma. This genus has four species, which occur in Malaysia, China, Indochina, Japan, India, and Sri Lanka.

Chloranthus species are dwarf shrubs or herbaceous plants, with xylem that contains vessels. Flowers are bisexual, each of which bears three stamens on straight filaments with three anther lobes and a wide connective, and with a smooth and moist stigma. The 20 species occur in southern and eastern Asia.

Ascarina has separate male and female flowers. The male flowers are subtended by two bracts and have between one and five stamens, in which the connective is not widened. The female flower is without bracts, the stigma is dry and covered in papillae. The fruit is a drupe-like berry. There are 12 species, which occur on islands in the Pacific and insular South-East Asia, from New Zealand and the Marquesas to Borneo, and on Madagascar.

Hedyosmum has separate male and female flowers. The male flowers are without bracts and have one stamen, in which the connective is not widened. The female flower is without bracts; the stigma is dry and covered in papillae. Female flowers have a 3-lobed calyx. The fruit, a drupe, has a kernel with a hard and woody shell. There are 43 species found in Latin America, including the Antilles, as well as one species found in Southeast Asia.

Taxonomy

The Chloranthaceae have been recognised as a family in most classifications but without clear relatives. Molecular systematics studies have shown that it is not closely related to any other family and is among the early-diverging lineages in the angiosperms. In particular, it is neither a eudicot nor a monocot. Fossils assigned to Chloranthaceae, or closely related to the family, are among the oldest angiosperms known.[4][5][6] The APG II system (2003) left the family unplaced as to order, but the APG III system (2009) accepted Chloranthales, containing only this family.[2] The cladogram below, from the APG IV system (2016), shows the Chloranthales in a trichotomy with the magnoliids and the monocot-Ceratophyllales-dicot clade. Earlier, the order was grouped with magnoliids, but studies in 2014 did not support this placement, leaving its phylogeny unclear.[7] A 2021 study sequenced the Chloranthus genome and found Chloranthales as sister to magnoliids.[8]


angiosperms
 

Amborellales

 
 
 

Nymphaeales

 
 
 

Austrobaileyales

 
 
 
 

magnoliids

 
 

Chloranthales

 
 
 
 

monocots

 
 
 

Ceratophyllales

 
 

eudicots

 
 
 
 
 
 
 

A 2004 study based on comparisons of homologous DNA fragments indicated that both the family Chloranthaceae and its extant genera Ascarina, Chloranthus, Hedyosmum and Sarcandra are probably monophyletic, with Hedyosmum being the first to diverge from the rest, and Ascarina being the sister group of the clade consisting of Sarcandra and Chloranthus. Desde O primeiro parâmetro é necessário, mas foi fornecido incorretamente! de 2016 (2016 -Erro de expressão: Palavra "june" não reconhecida), these four extant genera are recognized, and insights into their relationships are expressed in the tree below:[9][10]

Chloranthaceae
 
 
 
 

Chloranthus

 
 

Sarcandra

 
 
 

Ascarina

 
 
 

Hedyosmum

 
 
 

The extinct genus Chloranthistemon also belongs to this family.[11]

Historical classifications

The Cronquist system (1981) assigned the family

to the order Piperales
in subclass Magnoliidae
in class Magnoliopsida [=dicotyledons]
of division Magnoliophyta [=angiosperms].

The Thorne system (1992) placed it

in the order Magnoliales, which was assigned
to superorder Magnolianae
in subclass Magnoliideae [=dicotyledons],
in class Magnoliopsida [=angiosperms].

The Dahlgren system raised the family to be

its own order Chloranthales, which was assigned
to superorder Magnolianae
in subclass Magnoliideae [=dicotyledons],
in class Magnoliopsida [=angiosperms].

>

Referências

  1. ↑ «Chloranthales». www.mobot.org. Consultado em 20 de julho de 2023 
  2. ↑ a b c Angiosperm Phylogeny Group III (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.xAcessível livremente 
  3. ↑ Chloranthaceae
  4. ↑ Herendeen, Patrick S.; Friis, Else Marie; Pedersen, Kaj Raunsgaard; Crane, Peter R. (3 de março de 2017). «Palaeobotanical redux: revisiting the age of the angiosperms». Nature Plants. 3 (3): 17015. ISSN 2055-0278. PMID 28260783. doi:10.1038/nplants.2017.15 
  5. ↑ Doyle, James A.; Endress, Peter K. (1 de junho de 2018). «Phylogenetic Analyses of Cretaceous Fossils Related to Chloranthaceae and their Evolutionary Implications». The Botanical Review (em inglês). 84 (2): 156–202. ISSN 1874-9372. doi:10.1007/s12229-018-9197-6 
  6. ↑ Friis, Else Marie; Crane, Peter R.; Pedersen, Kaj Raunsgaard (1 de março de 2019). «Hedyosmum-Like Fossils in the Early Cretaceous Diversification of Angiosperms». International Journal of Plant Sciences. 180 (3): 232–239. ISSN 1058-5893. doi:10.1086/701819 
  7. ↑ Angiosperm Phylogeny Group (2016). «An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV». Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385Acessível livremente 
  8. ↑ Guo, Xing (26 November 2021). «Chloranthus genome provides insights into the early diversification of angiosperms». Nature Communications. 12 (1): 6930. PMC 8626473Acessível livremente. PMID 34836973. doi:10.1038/s41467-021-26922-4Acessível livremente  Verifique data em: |data= (ajuda)
  9. ↑ Erro de citação: Etiqueta <ref> inválida; não foi fornecido texto para as refs de nome MOBOT
  10. ↑ Eklund, Helena; Doyle, James A.; Herendeen, Patrick S. (2004). «Morphological Phylogenetic Analysis of Living and Fossil Chloranthaceae». International Journal of Plant Sciences. 165 (1): 107–151. JSTOR 10.1086/380987. doi:10.1086/380987 
  11. ↑ Herendeen, P. S.; Crepet, W. L.; Nixon, K. C. (1993). «Chloranthus-like stamens from the Upper Cretaceous of New Jersey». American Journal of Botany. 80 (8): 865–871. doi:10.1002/j.1537-2197.1993.tb15306.x 

Bibliografia

Ligações externas

O Commons possui uma categoria com imagens e outros ficheiros sobre Chloranthales
Wikispecies
Wikispecies
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