View source for Chloranthaceae

{{Short description|Family of flowering plants}}
{{Automatic taxobox
| fossil_range = {{fossil range|Barremian|present|[[Barremian]] - recent|ref=<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>}}
| image = Sarcandra_glabra2.jpg
| image_caption = ''[[Sarcandra glabra]]''
| image2 = Chloranthus fortunei 2.jpg
| image2_caption = ''[[Chloranthus fortunei]]''
| parent_authority = [[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 |hdl=10654/18083 |hdl-access=free }}</ref>
| taxon = Chloranthaceae
| authority = R.Br. ex Sims<ref name="apgiii" />
| subdivision_ranks = Genera
| subdivision = * ''[[Ascarina]]'' <small>J.R.Forst. & G.Forst.</small>
* ''[[Chloranthus]]'' <small>Sw.</small>
* ''[[Hedyosmum]]'' <small>Sw.</small>
* ''[[Sarcandra]]'' <small>Gardner</small>
}}

'''Chloranthaceae''' ({{IPAc-en|k|l|ɔər|æ|n|ˈ|θ|eɪ|s|iː|.|i}} {{respell|klor|ann|THAY|see|ee}})<ref>{{Cite Merriam-Webster|Chloranthaceae|access-date=2024-11-04}}</ref> 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|hdl=11336/136234|hdl-access=free}}</ref>

==Description==
Chloranthaceae are [[Essential oil|fragrant]] [[shrub]]s or [[herbaceous plant]]s, that only produce new 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.

''[[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.

''[[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]].

''[[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].

==References==
{{Reflist}}

==External links==
* [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'']

{{Angiosperm orders}}
{{Angiosperm families}}
{{Taxonbar|from=Q133208}}
{{Authority control}}

[[Category:Chloranthaceae| ]]
[[Category:Angiosperm families]]