Difference between revisions of "Celastrales"

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{{Automatic taxobox
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#redirect [[:Category:Celastrales]]
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| taxon = Celastrales
 
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| image= Illustration Euonymus europaea0.jpg
 
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| image_caption = '' Euonymus europaea'', Celastraceae family
 
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| authority = [[Johann Heinrich Friedrich Link|Link]]<ref name=APGIII2009>{{Cite journal |last=Angiosperm Phylogeny Group |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 |url=http://www3.interscience.wiley.com/journal/122630309/abstract | format= PDF |accessdate=2013-07-06 |doi=10.1111/j.1095-8339.2009.00996.x }}</ref>
 
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| subdivision_ranks = Families
 
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| subdivision =
 
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[[Celastraceae]]<br/>
 
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[[Lepidobotryaceae]]
 
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}}
 
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The '''Celastrales''' are an [[order (biology)|order]] of [[flowering plant]]s found throughout the [[tropics]] and [[subtropics]], with only a few species extending far into the [[Temperateness|temperate]] regions. The 1200<ref name="kubitski2004">"Lepidobotryaceae", "Parnassiaceae", and "Celastraceae" In: Klaus Kubitzki (ed.). ''The Families and Genera of Vascular Plants'' vol. VI. Springer-Verlag: Berlin;Heidelberg, Germany. (2004). {{ISBN|978-3-540-06512-8}} (vol. VI).</ref> to 1350<ref name="celastrales">Peter F. Stevens (2001 onwards). [http://www.mobot.org/MOBOT/Research/APweb/orders/Celastralesweb.html#Celastrales Celastrales] At: [http://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny] at [http://www.mobot.org Missouri Botanical Garden]</ref> [[species]] are in about 100 [[Genus|genera]]. All but seven of these genera are in the large [[family (biology)|family]] [[Celastraceae]]. Until recently, the composition of the order and its division into families varied greatly from one author to another.
 
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== Description ==
 
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The Celastrales are a diverse order that has no conspicuous [[Synapomorphy|distinguishing characteristic]], so is consequently hard to recognize.<ref name="matthews2005">{{cite journal | doi = 10.1111/j.1095-8339.2005.00445.x | last1 = Matthews | first1 = Merran L. | last2 = Endress | first2 = Peter K. | year = 2005 | title = Comparative floral structure and systematics in Celastrales | url = | journal = Botanical Journal of the Linnean Society | volume = 149 | issue = 2| pages = 129–194 }}</ref> The [[flower]]s are usually small with a conspicuous [[nectar| nectary disk]]. The [[stipule]]s are small or rarely absent. The [[Ovule|micropyle]] has two openings and is therefore called a bistomal micropyle. Flowers with well-developed male and female parts are often [[Plant sexuality|functionally unisexual]]. The [[seed]] often has an [[aril]]. In [[bud]], the [[sepal]]s have a [[quincuncial (botany)|quincuncial]] arrangement. This means that two sepals are inside, two are outside, and the remaining sepal is half inside and half outside.
 
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== Relationships ==
 
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Perhaps the most conspicuous and unusual [[Phenotypic trait|trait]] of the Celastrales is the nectary disk, a feature that it shares with another [[Rosids|rosid]] order, [[Sapindales]]. Since the orders are not [[Plant taxonomy|closely related]], the disk must have been an independent development in each of these [[Lineage (evolution)|lines]].
 
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The Celastrales are a member of the Celastrales, [[Oxalidales]] (including [[Huaceae]]), and [[Malpighiales]] (COM) clade.<ref name="wang2009">{{Cite journal | author = Hengchang Wang, Michael J. Moore, [[Pamela S. Soltis]], Charles D. Bell, Samuel F. Brockington, Roolse Alexandre, Charles C. Davis, Maribeth Latvis, Steven R. Manchester, and [[Douglas E. Soltis]] | year = 2009 | title = Rosid radiation and the rapid rise of angiosperm-dominated forests | journal = Proceedings of the National Academy of Sciences | volume = 106 | issue = 10 | pages = 3853–3858 | doi = 10.1073/pnas.0813376106 | pmc = 2644257 | postscript = <!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}} | pmid=19223592}}</ref> of Fabidae, with Fabidae being one of the 2 groups of [[Eurosids]].<ref name="cantino2007">{{cite journal | doi = 10.2307/25065865 | last1 = Cantino | first1 = Philip D. | last2 = Doyle | first2 = James A. | last3 = Graham | first3 = Sean W. | last4 = Judd | first4 = Walter S. |authorlink4 = Walter Stephen Judd | last5 = Olmstead | first5 = Richard G. | last6 = Soltis | first6 = Douglas E. |authorlink6 = Douglas E. Soltis | last7 = Soltis | first7 = Pamela S. | authorlink7 = Pamela S. Soltis|last8 = Donoghue | first8 = Michael J. | year = 2007 | title = Towards a phylogenetic nomenclature of ''Tracheophyta''". | url = | journal = Taxon | volume = 56 | issue = 3| pages = 822–846 }}</ref>
 
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== Circumscription ==
 
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The [[Botanical name|name]] Celastrales was first used by [[Thomas Baskerville (botanist)|Thomas Baskerville]] in 1839.<ref name="baskerville1839">Thomas Baskerville. ''[https://books.google.com/books?id=F3cEAAAAQAAJ&dq=baskerville+%22affinities+of+plants%22&printsec=frontcover&source=bl&ots=eAPumlONia&sig=E4P7ib6HKR2_wPl-QqWHmqxtxLQ&hl=en&ei=bzXRSeSrEoq6tQPEleGiAw&sa=X&oi=book_result&resnum=1&ct=result Affinities of Plants: with some observations upon progressive development. page 104.]''. Taylor and Walton: Gower Street, London. (1839).</ref> In the time since Baskerville first defined the order, until the 21st century, great differences of opinion occurred about what should be included in the order and in its largest family, the [[Celastraceae]]. The family Celastraceae was the only [[clade|group]] consistently placed in the order by all authors who accepted it. Because of the ambiguity and complexity of its [[Circumscription (taxonomy)|definition]], the Celastraceae became a [[Wastebasket taxon|dumping ground]] for genera of dubious [[Systematics|affinity]]. Several genera were assigned to this family with considerable doubt about whether they really belonged there. Also, some genera that properly belong in the Celastraceae were placed elsewhere.
 
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By the end of the 20th century, ''[[Goupia]]'' and ''[[Glossopetalon|Forsellesia]]'' had been excluded from the Celastraceae and also from the Celastrales. ''Goupia'' is now in the [[Malpighiales]].<ref name="wurdack2009">{{cite journal | doi = 10.3732/ajb.0800207 | last1 = Wurdack | first1 = Kenneth J. | last2 = Davis | first2 = Charles C. | year = 2009 | title = Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm tree of life | url = | journal = American Journal of Botany | volume = 96 | issue = 8| pages = 1551–1570 | pmid=21628300}}</ref> ''Forsellesia'' is now in the [[Crossosomatales]].<ref name="thorne1978">{{cite journal | last1 = Thorne | first1 = Robert F. | last2 = Scogin | first2 = Ron | year = 1978 | title = ''Forsellesia'' Greene (''Glossopetalon'' Gray), a third genus in the Crossosomataceae (Rosinae, Rosales) | url = | journal = Aliso | volume = 9 | issue = 2| pages = 171–178 }}</ref> It continues to be the subject of a dispute about whether its [[Valid name (botany)|proper name]] is ''Forsellesia'' or ''[[Glossopetalon]]''.<ref name="sosa2007">Victoria Sosa. "Crossosomataceae" In: Klaus Kubitzki (ed.) ''The Families and Genera of Vascular Plants'' vol.IX. Springer-Verlag: Berlin,Heidelberg (2007).</ref>
 
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After being placed elsewhere, ''[[Canotia]], [[Brexia]]'', and ''[[Plagiopteron]]'' were found to belong in the Celastraceae. The family Hippocrateaceae was found to be deeply nested within the Celastraceae and is no longer recognized as a separate family.
 
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In 2000, Vincent Savolainen [[List of Latin phrases: E#et alii|et alii]] found that three families - [[Lepidobotryaceae]], [[Parnassiaceae]], and [[Celastraceae]] - were closely related.<ref name="savolainen2000">{{cite journal | doi=10.2307/4115644 | jstor=4115644 | last1=Savolainen | year = 2000 | first1=V. | last2=Fay | first2=M. F. | last3=Albach | first3=D. C. | last4=Backlund | first4=A. | last5=Van Der Bank | first5=M. | last6=Cameron | first6=K. M. | last7=Johnson | first7=S. A. | last8=Lledó | first8=M. D. | last9=Pintaud | first9=J. C.| displayauthors = 8 | title = Phylogeny of the eudicots: a nearly complete familial analysis based on ''rbcL'' gene sequences | url = | journal = Kew Bulletin | volume = 55 | issue = 2| pages = 257–309 }}</ref> They stated that these three families should constitute the order Celastrales, and this idea was accepted by the [[Angiosperm Phylogeny Group]], which later [[Synonym (taxonomy)|subsumed]] the Parnassiaceae into the Celastraceae. Savolainen and co-authors also excluded ''[[Lophopyxis]]'' from the Celastrales. ''Lophopyxis'' now constitutes a [[Monotypic taxon|monogeneric]] family in the Malpighiales.<ref name="wurdack2009"/>
 
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In 2001, in a [[molecular phylogenetic]] study of [[DNA sequence]]s, Mark Simmons and others confirmed all of these results except for the placement of ''Lophopyxis'' and the Lepidobotryaceae, which they did not sample.<ref name="simmons2001">{{cite journal | last1 = Simmons | first1 = Mark P. | last2 = Savolainen | first2 = Vincent | last3 = Clevinger | first3 = Curtis C. | last4 = Archer | first4 = Robert H. | last5 = Davis | first5 = Jerrold I. | year = 2001 | title = Phylogeny of Celastraceae Inferred from 26S Nuclear Ribosomal DNA, Phytochrome B, rbcL, atpB, and Morphology | url = | journal = Molecular Phylogenetics and Evolution | pmid = 11399146 | volume = 19 | issue = 3| pages = 353–366 | doi = 10.1006/mpev.2001.0937 }}</ref>
 
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In 2006, Li-Bing Zhang and Mark Simmons produced a [[phylogeny]] of the Celastrales based on [[Cell nucleus|nuclear]] [[ribosome|ribosomal]], and [[chloroplast]] [[DNA]].<ref name="zhang2006">{{cite journal | doi = 10.1600/036364406775971778 | last1 = Li-Bing | first1 = Zhang | last2 = Simmons | first2 = Mark P. | year = 2006 | title = Phylogeny and Delimitation of the Celastrales Inferred from Nuclear and Plastid Genes | url = | journal = Systematic Botany | volume = 31 | issue = 1| pages = 122–137 }}</ref> Their results showed that ''[[Bhesa]]'' and ''[[Perrottetia]]'' were misplaced in the Celastraceae. ''Bhesa'' is now in the [[Centroplacaceae]], a family in the Malpighiales.<ref name="wurdack2009"/> and ''Perrottetia'' is in the [[Huerteales]].<ref name="worberg2009">{{cite journal | last1 = Worberg | first1 = Andreas | last2 = Alford | first2 = Mac H. | last3 = Quandt | first3 = Dietmar | last4 = Borsch | first4 = Thomas | year = 2009 | title = Huerteales sister to Brassicales plus Malvales, and newly circumscribed to include ''Dipentodon, Gerrardina, Huertea, Perrottetia'', and ''Tapiscia'' | url = | journal = Taxon | volume = 58 | issue = 2| pages = 468–478 }}</ref> Zhang and Simmons found ''[[Pottingeria]]'' and ''[[Mortonia (plant)|Mortonia]]'' to be closely related to the families Parnassiaceae and Celastraceae, as they were then defined, but not in either of them. These two genera are therefore in the Celastrales. They found that ''[[Siphonodon]]'' and ''[[Empleuridium]]'' are proper members of the Celastraceae, removing considerable doubt about their placement there. They also showed that the small family Stackhousiaceae, consisting of three genera, is [[Paraphyly|embedded]] in the Celastraceae. Except for [[Taxon|taxa]] that were not sampled, these results were confirmed by the second phylogeny of the Celastrales, which was produced by Mark Simmons and several co-authors in 2008.<ref name="simmons2008">{{Cite journal |author1=Mark P. Simmons |author2=Jennifer J. Cappa |author3=Robert H. Archer |author4=Andrew J. Ford |author5=Dedra Eichstedt |author6=Curtis C. Clevinger | year = 2008 | title = Phylogeny of the Celastreae (Celastraceae) and the relationships of Catha edulis (qat) inferred from morphological characters and nuclear and plastid genes | journal = Molecular Phylogenetics and Evolution | volume = 48 | issue = 2 | pages = 745–757 | doi = 10.1016/j.ympev.2008.04.039 | pmid = 18550389 | postscript = <!-- Bot inserted parameter. Either remove it; or change its value to "." for the cite to end in a ".", as necessary. -->{{inconsistent citations}} }}</ref>
 
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''[[Nicobariodendron]] sleumeri'', the only member of its genus, continues to be an enigma. It is a small tree from the [[Andaman and Nicobar Islands]] of India. Little is known of it and it has never been sampled for DNA. It is generally thought to belong in the Celastrales,<ref name="celastrales"/> but this is not a certainty. It is one of the five taxa placed ''[[incertae sedis]]'' in the angiosperms in the [[APG III system]] of [[Biological classification|classification]].<ref name=APGIII2009/>
 
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== Families ==
 
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The Celastrales have been divided into families in various ways. In their [[APG II]] classification in 2003, the [[Angiosperm Phylogeny Group]] recognized three families in the Celastrales - [[Lepidobotryaceae]], [[Parnassiaceae]], and [[Celastraceae]]. When they revised their classification in 2009, they recognized only two families because ''Pottingeria'' and the two genera of Parnassiaceae were transferred to the Celastraceae. ''Nicobariodendron'' became one of the five taxa placed ''incertae sedis'' in the angiosperms.
 
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In the 2006 phylogeny, ''Nicobariodendron'' was not sampled, but those species that were sampled fell into two strongly supported clades. One was a small clade consisting only of the family Lepidobotryaceae. Its [[sister taxon|sister]] was a very large clade containing the rest of the order. The large clade consisted of five strongly supported groups. These are the family Parnassiaceae, the genus ''Pottingeria'', the genus ''[[Mortonia (plant)|Mortonia]]'' (in the Celastraceae), and a pair of genera from the Celastraceae (''[[Quetzalia]]'' and ''[[Zinowiewia]]''), and the rest of the Celastraceae. No relationships were resolved among these groups.
 
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In 2008, Simmons and others produced a phylogeny of the Celastrales that achieved better resolution than the 2006 study by sampling more species and more DNA. They found the same [[Polytomy|pentatomy]] of five strongly supported groups that the previous study had found, but only weak to moderate support for any relationships between the five groups.<ref name="simmons2008"/> In the APG III system, the Celastraceae family was expanded to consist of these five groups. No one has yet published an intrafamilial classification for the expanded Celastraceae.<ref name=APGIII2009/>
 
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== Phylogeny ==
 
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The following [[phylogenetic tree]] was made by combining parts of three different trees.<ref name="simmons2001"/><ref name="zhang2006"/><ref name="simmons2008"/> Bootstrap support is 100% except where shown. Branches with less than 50% bootstrap support are collapsed. The clade numbers are from Simmons ''et al.'' (2008).<ref name="simmons2008"/>
 
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{{clade | style=font-size:75%;line-height:100%
 
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|label1='''Celastrales'''&nbsp;&nbsp;
 
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|1={{clade
 
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|label1=[[Lepidobotryaceae]]
 
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|1={{clade
 
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  |1=''[[Lepidobotrys]]
 
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  |2=''[[Ruptiliocarpon]]
 
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  }}
 
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|2={{clade
 
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  |1={{clade
 
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      |1=''[[Lepuropetalon]]
 
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      |2=''[[Parnassia]]
 
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      }}
 
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  |label2=68&nbsp;
 
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  |2={{clade
 
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      |1=''[[Pottingeria]]
 
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      |2=[[Mortonia (plant)|''Mortonia'']]
 
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      }}
 
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  |label3=56&nbsp;
 
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  |3={{clade
 
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      |1={{clade
 
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        |1=''[[Quetzalia]]
 
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        |2=''[[Zinowiewia]]
 
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        }}
 
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      |2={{clade
 
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        |label1=CLADE&nbsp;'''1'''&nbsp;
 
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        |1={{clade
 
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            |1={{clade
 
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              |1=''[[Peripterygia]]
 
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              |2=''[[Siphonodon]]
 
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              }}
 
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            |label2=80&nbsp;
 
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            |2={{clade
 
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              |1=''[[Dicarpellum]]
 
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              |2={{clade
 
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                  |1=''[[Tripterococcus]]
 
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                  |2={{clade
 
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                    |1=[[Macgregoria (plant)|''Macgregoria'']]
 
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                    |2=''[[Stackhousia]]
 
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                    }}
 
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                                  }}
 
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              |3={{clade
 
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                  |1=''[[Menepetalum]]
 
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                  |2=''[[Psammomoya]]
 
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                  }}
 
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              |4=''[[Denhamia]]
 
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              }}
 
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            }}
 
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        |label2=87&nbsp;
 
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        |2={{clade
 
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            |label1=CLADE&nbsp;'''2'''&nbsp;
 
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            |1={{clade
 
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              |1=''[[Maytenus]]
 
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              |label2=53&nbsp;
 
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              |2={{clade
 
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                  |1=''[[Gyminda]]
 
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                  |label2=89&nbsp;
 
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                  |2={{clade
 
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                    |1={{clade
 
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                        |1=''[[Tripterygium]]
 
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                        |2=''[[Celastrus]]
 
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                        }}
 
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                    |label2=99&nbsp;
 
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                    |2={{clade
 
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                        |1=''[[Paxistima]]
 
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                        |2=''[[Crossopetalum]]
 
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                        |3=''[[Canotia]]
 
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                        |4=''[[Euonymus]]
 
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                        }}
 
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                    }}
 
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                  }}
 
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              }}
 
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            |label2=CLADE&nbsp;'''3'''&nbsp; 89
 
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            |2={{clade
 
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              |label1=CLADE&nbsp;'''4'''&nbsp;
 
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              |1={{clade
 
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                  |1=''[[Empleuridium]]
 
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                  |label2=72&nbsp;
 
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                  |2={{clade
 
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                    |1=''[[Pterocelastrus]]
 
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                    |label2=69&nbsp;
 
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                    |2={{clade
 
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                        |1=''[[Mystroxylon]]
 
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                        |2=''[[Robsonodendron]]
 
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                        }}
 
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                    }}
 
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                  }}
 
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              |label2=CLADE&nbsp;'''5'''&nbsp; 50
 
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              |2={{clade
 
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                  |1=''[[Salaciopsis]]
 
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                  |label2=CLADE&nbsp;'''6'''&nbsp;
 
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                  |2={{clade
 
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                    |label1=96&nbsp;
 
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                    |1={{clade
 
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                        |1=[[Catha (plant)|''Catha'']]
 
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                        |2={{clade
 
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                          |1=''[[Hartogiella]]
 
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                          |label2=&nbsp;&nbsp;&nbsp;
 
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                          |2={{clade
 
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                              |1=[[Cassine (plant)|''Cassine'']]
 
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                              |2=[[Maurocenia]]
 
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                              }}
 
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                          }}
 
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                        }}
 
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                    |label2=98&nbsp;
 
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                    |2={{clade
 
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                        |1=''[[Lydenburgia]]
 
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                        |2=''[[Gymnosporia]]
 
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                        }}
 
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                    }}
 
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                  |label3=CLADE&nbsp;7&nbsp;
 
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                  |3={{clade
 
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                    |label1=&nbsp;&nbsp;&nbsp;
 
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                    |1={{clade
 
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                        |label1=&nbsp;&nbsp;&nbsp;
 
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                        |1={{clade
 
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                          |1=''[[Polycardia]]
 
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                          |2=''[[Brexia]]
 
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                          }}
 
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                        |label2=&nbsp;&nbsp;&nbsp;
 
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                        |2={{clade
 
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                          |1=''[[Pleurostylia]]
 
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                          |label2=&nbsp;&nbsp;&nbsp;
 
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                          |2={{clade
 
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                              |1=''[[Elaeodendron]]
 
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                              |2=''[[Pseudocatha]]
 
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                              }}
 
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                          }}
 
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                        }}
 
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                    |2=''[[Kokoona]]
 
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                        |3={{clade
 
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                          |1={{clade
 
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                              |1=''[[Lophopetalon]]
 
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                              |2={{clade
 
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                                |1=[[Salacia (plant)|''Salacia'']]
 
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                                |2=''[[Tontelea]]
 
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                                }}
 
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                              }}
 
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                          |2={{clade
 
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                              |1=''[[Plagiopteron]]
 
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                              |2=''[[Hippocratea]]
 
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                              |3=''[[Pristimera]]
 
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                              |4=''[[Loeseneriella]]
 
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                              }}
 
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                          }}
 
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                    }}
 
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}}
 
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{{Commons}}
 
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== References ==
 
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{{reflist}}
 
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{{taxonbar}}
 
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[[Category:Celastrales| ]]
 
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[[Category:Rosid orders]]
 
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[[Category:Angiosperm orders]]
 

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