Difference between revisions of "Magnoliids"

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{{Automatic taxobox
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#redirect [[:Category:Magnoliids]]
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| taxon = Magnoliids
 
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| image = Asimina triloba - pawpaw - desc-flower.jpg
 
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| image_caption = Flower of ''[[Asimina triloba]]''
 
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| subdivision_ranks = Orders
 
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| subdivision =
 
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[[Canellales]]<br/>
 
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[[Laurales]]<br/>
 
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[[Magnoliales]]<br/>
 
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[[Piperales]]
 
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| type_species = ''[[Magnolia virginiana]]''
 
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}}
 
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'''Magnoliids''' (or '''Magnoliidae''' or '''Magnolianae''') are a group of [[flowering plant]]s. Until recently, the group included about 9,000<ref name="palmer">{{cite journal | url = http://www.amjbot.org/cgi/content/full/91/10/1437 | title = The plant tree of life: an overview and some points of view | first1=Jeffrey D. |last1=Palmer |first2=Douglas E. |last2=Soltis |authorlink2=Douglas E. Soltis |first3=Mark W. |last3=Chase |authorlink3=Mark Wayne Chase | journal = American Journal of Botany | year = 2004 | volume = 91 | pages = 1437–1445 (Fig.2) | doi = 10.3732/ajb.91.10.1437 | issue=10 | pmid=21652302}}</ref> species, including [[magnolia]]s, [[nutmeg]], [[bay laurel]], [[cinnamon]], [[avocado]], [[black pepper]], [[tulip tree]] and many others.  That group is characterized by [[Merosity|trimerous]] flowers, pollen with one pore, and usually branching-veined leaves.
 
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==Classification==
 
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"Magnoliidae" is the [[botanical name]] of a [[subclass (biology)|subclass]], and "magnoliids" is an informal name that does not conform to the [[International Code of Nomenclature for algae, fungi, and plants]]. The [[wikt:circumscription|circumscription]] of a subclass will vary with the [[List of systems of plant taxonomy|taxonomic system]] being used. The only requirement is that it must include the family [[Magnoliaceae]].<ref>[[International Code of Nomenclature for algae, fungi, and plants]], Art. 16</ref> The informal name "magnoliids" is used by some researchers to avoid the confusion that recently surrounds the name "Magnoliidae". More recently,{{clarify|more recently? Since 2009?|date=February 2016}} the group{{clarify|under what name, is there a name?|date=February 2016}} has been redefined under the ''PhyloCode'' as a node-based [[clade]] comprising the [[Canellales]], [[Laurales]], [[Magnoliales]], and [[Piperales]].{{citation needed|date=February 2016}} Chase & Reveal have proposed, "Magnoliidae" as the name used for the entire group of flowering plants, and the formal name "Magnolianae" for the group of four orders are discussed here.<ref name=C&R>{{citation |author1=Chase, M.W. |authorlink1=Mark Wayne Chase |author2=Reveal, J.L. |authorlink2=James Reveal |year=2009 |title=A phylogenetic classification of the land plants to accompany APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=122–127 |doi=10.1111/j.1095-8339.2009.01002.x}}</ref>
 
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===APG system===
 
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The [[APG III system|APG III]] (2009) and its predecessor systems did not originally use formal botanical names above the [[rank (botany)|rank]] of order. Under those systems, larger clades were usually referred to by informal names, such as "magnoliids" (plural, not capitalized) or "magnoliid complex". The formal name in [[Linnean nomenclature]] was specified in a separate APG publication as the existing name "Magnolianae" [[Armen Takhtajan|Takht.]] (1967).<ref name=C&R/> The APG III recognizes a [[clade]] within the [[angiosperms]] for the magnoliids. The circumscription is:
 
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{| align="left" style="text-align:left; padding:2.5px; background:#eef"
 
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|-
 
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| style="background:#fff; padding:2.5px" |clade '''magnoliids'''
 
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: order [[Canellales]]
 
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: order [[Laurales]]
 
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: order [[Magnoliales]]
 
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: order [[Piperales]]
 
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|{{clade| style=font-size:75%;line-height:75%
 
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|label1=
 
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|1={{clade
 
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|1={{clade
 
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|1=[[Chloranthales]]
 
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|2={{clade
 
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|label1='''magnoliids'''
 
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|1={{clade
 
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|1={{clade
 
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|1=[[Canellales]]
 
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|2=[[Piperales]]
 
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}}
 
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|2={{clade
 
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|1=[[Laurales]]
 
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|2=[[Magnoliales]]
 
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}}
 
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}}
 
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}}
 
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}}
 
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|2={{clade
 
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|1=[[monocots]]
 
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|2={{clade
 
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|1=[[Ceratophyllales]]
 
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|2=[[eudicots]]
 
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}}
 
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}}
 
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}}
 
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}}
 
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|-style="font-size:90%;"
 
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| colspan=2 | The current [[phylogenetics|phylogeny]] and composition of the magnoliids.<ref>{{cite journal | doi=10.1111/j.1095-8339.2009.00996.x | author=The 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 | pages=105–121 }}</ref><ref>{{cite journal | last1=Soltis | first1=P. S. |authorlink1 = Pamela S. Soltis |last2=Soltis |first2=Douglas E. |authorlink2=Douglas E. Soltis | year=2004 | title=The origin and diversification of Angiosperms | journal= American Journal of Botany | volume=91 | pages=1614–1626 |url= http://www.amjbot.org/cgi/content/abstract/91/10/1614 | doi=10.3732/ajb.91.10.1614 | issue=10 | pmid=21652312}}</ref>
 
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|}{{Clear}}
 
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The clade includes most of the basal groups of the angiosperms.  This clade was formally named Magnoliidae in 2007 under provisions of the ''[[PhyloCode]]''.<ref name="Cantino">{{cite journal | last= Cantino | first= Philip D. |author2=James A. Doyle |author3=Sean W. Graham |author4=Walter S. Judd |authorlink4=Walter Stephen Judd |author5=Richard G. Olmstead |author6=Douglas E. Soltis |authorlink6=Douglas E. Soltis |author7=Pamela S. Soltis |authorlink7 = Pamela S. Soltis|author8=Michael J. Donoghue  | year=2007 | title= Towards a phylogenetic nomenclature of ''Tracheophyta'' | journal=Taxon | volume=56 | issue=3 | pages= E1–E44 | doi=10.2307/25065865}}</ref>
 
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===Cronquist system===
 
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[[File:Magnolia obovata 02.jpg|thumb|right|240px|Flower of ''[[Magnolia obovata]]'', showing multiple [[petal]]s, [[stamen]]s, and [[pistil]]s.]]
 
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The [[Cronquist system]] (1981) used the name Magnoliidae for one of six subclasses (within class [[Magnoliopsida]] = [[dicotyledons]]). In the original version of this system the circumscription was:<ref name="Cronquist">{{cite book | last=Cronquist | first=Arthur | year=1981 | title=An Integrated System of Classification of Flowering Plants | location=New York | publisher=Columbia University Press | isbn=0-231-03880-1 }}</ref>
 
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* Subclass Magnoliidae :
 
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*: Order [[Aristolochiales]]
 
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*: Order [[Illiciales]]
 
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*: Order [[Laurales]]
 
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*: Order [[Magnoliales]]
 
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*: Order [[Nymphaeales]]
 
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*: Order [[Papaverales]]
 
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*: Order [[Piperales]]
 
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*: Order [[Ranunculales]]
 
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===Dahlgren and Thorne systems===
 
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Both [[Dahlgren system|Dahlgren]] and [[Thorne system (1992)|Thorne]] classified the magnoliids (''sensu'' APG) in superorder '''Magnolianae''', rather than as a subclass.<ref name="Thorne 1992"/>  In their systems, the name Magnoliidae is used for a much larger group including all [[dicotyledons]]. This is also the case in some of the systems derived from the Cronquist system.
 
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Dahlgren divided his Magnolianae into ten [[order (biology)|orders]], more than other systems of the time, and unlike Cronquist and Thorne, he did not include the [[Piperales]].<ref name="Dahlgren"/>  Thorne grouped most of his Magnolianae into two large orders, [[Magnoliales]] and [[Berberidales]], although his Magnoliales was divided into suborders along lines similar to the ordinal groupings used by both Cronquist and Dahlgren. Thorne revised his system in 2000, restricting the name Magnoliidae to include only the Magnolianae, Nymphaeanae, and Rafflesianae, and removing the Berberidales and other previously included groups to his subclass Ranunculidae.<ref name="Thorne 2000"/> This revised system diverges from the Cronquist system, but agrees more closely with the circumscription later published under APG II.
 
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===Comparison table===
 
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Comparison of classification systems is often difficult.  Two authors may apply the same name to groups with different composition of members; for example, Dahlgren's Magnoliidae includes all [[dicot]]s, whereas Cronquists' Magnoliidae is only one of five dicot groups.  Two authors may also describe the same group with nearly identical composition, but each may then apply a different name to that group or place the group at a different [[taxonomic rank]].  For example, the composition of Cronquist's ''subclass'' Magnoliidae is nearly the same as Thorne's (1992) ''superorder'' Magnolianae, despite the difference in taxonomic rank.
 
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Because of these difficulties and others, the synoptic table below imprecisely compares the definition of "magnoliid" groups in the systems of four authors.  For each system, only orders are named in the table.  All orders included by a particular author are listed and linked in that column.  When a taxon is not included by that author, but was included by an author in another column, that item appears in unlinked italics and indicates remote placement.  The sequence of each system has been altered from its publication in order to pair corresponding taxa between columns.
 
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{| class="wikitable" align="left" style="margin-left:1em"
 
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|+'''Comparison of the magnoliids across five systems'''
 
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!style="background:lightgreen" align="center" width="160px" | [[APG II system]]<ref name="APG2">{{cite journal | journal=Botanical Journal of the Linnean Society | volume=141 | pages=399–436 | author=Angiosperm Phylogeny Group | year=2003 | title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II | url=http://www.blackwell-synergy.com/links/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/full/ | doi=10.1046/j.1095-8339.2003.t01-1-00158.x }}</ref><br/>magnoliids
 
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!style="background:lightgreen" align="center" width="160px" | [[Cronquist system]]<ref name="Cronquist"/><br/>Magnoliidae
 
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!style="background:lightgreen" align="center" width="160px" | [[Dahlgren system]]<ref name="Dahlgren">{{cite journal | last=Dahlgren | first=R.M.T. | year=1980 | title = A revised system of classification of angiosperms | journal= Botanical Journal of the Linnean Society | volume=80 | issue=2 | pages=91–124 | doi = 10.1111/j.1095-8339.1980.tb01661.x }}</ref><br/>Magnolianae
 
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!style="background:lightgreen" align="center" width="160px" | [[Thorne system (1992)]]<ref name="Thorne 1992">{{cite journal | last=Thorne | first=R. F. | year=1992 | title= Classification and geography of the flowering plants | journal= Botanical Review | volume=58 | pages=225–348 | doi= 10.1007/BF02858611}}</ref><br/>Magnolianae
 
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!style="background:lightgreen" align="center" width="160px" | Thorne system (2000)<ref name="Thorne 2000">{{cite journal | last=Thorne | first=R. F. | year=2000 | title= The classification and geography of the flowering plants: Dicotyledons of the class Angiospermae | journal= Botanical Review | volume=66 | issue=4 | pages=441–647 |doi= 10.1007/BF02869011 }}</ref><br/>Magnolianae
 
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|-
 
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|-
 
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| [[Laurales]]
 
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| [[Laurales]]
 
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| [[Laurales]]
 
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| rowspan=9 | [[Magnoliales]]
 
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| rowspan=9 | [[Magnoliales]]
 
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|-
 
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| rowspan=2 | [[Magnoliales]]
 
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| rowspan=4 | [[Magnoliales]]
 
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| [[Magnoliales]]
 
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|-
 
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| [[Annonales]]
 
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|-
 
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| [[Canellales]]
 
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| [[Winterales]]
 
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|-
 
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| rowspan=3 | [[Piperales]]
 
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| [[Lactoridales]]
 
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|-
 
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| [[Aristolochiales]]
 
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| [[Aristolochiales]]
 
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|-
 
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| rowspan=2 | [[Piperales]]
 
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| ''Piperales in Nymphaeanae''
 
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|-
 
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| rowspan=5 | ''unplaced or in basal clades''
 
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| [[Chloranthales]]
 
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|-
 
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| [[Illiciales]]
 
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| [[Illiciales]]
 
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|-
 
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| ''in Rosidae''
 
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| [[Rafflesiales]]
 
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| ''in Rafflesianae''
 
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| ''in Rafflesianae''
 
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|-
 
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| rowspan=3 | [[Nymphaeales]]
 
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| rowspan=2 | ''in Nymphaeanae''
 
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| ''in Nymphaeanae''
 
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| ''in Nymphaeanae''
 
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|-
 
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| [[Ceratophyllales]]
 
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| rowspan=5 | ''in Ranunculidae''
 
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|-
 
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| rowspan=4 | ''placed in eudicot clade''
 
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| [[Nelumbonales]]
 
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| [[Nelumbonales]]
 
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|-
 
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| [[Ranunculales]]
 
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| rowspan=2 | ''in Ranunculanae''
 
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| rowspan=2 | [[Berberidales]]
 
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|-
 
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| [[Papaverales]]
 
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|-
 
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| ''in Dilleniidae''
 
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| ''in Theanae''
 
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| [[Paeoniales]]
 
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|}
 
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{{Clear}}
 
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==Economic uses==
 
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The magnoliids is a large group of plants, with many species that are economically important as food, drugs, perfumes, timber, and as ornamentals, among many other uses.
 
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[[File:Persea americana 2.jpg|thumb|right|The [[avocado]] has been cultivated in the Americas for thousands of years.]]
 
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One widely cultivated magnoliid [[fruit]] is the [[avocado]] (''Persea americana''), which is believed to have been cultivated in [[Mexico]] and [[Central America]] for nearly 10,000 years.<ref name="EB">{{cite encyclopedia | title = Angiosperms | encyclopedia = The New Encyclopædia Britannica | volume = 13 | pages = 634–645 | year = 1994 }}</ref>  Now grown throughout the American tropics, it probably originates from the [[Chiapas]] region of Mexico or [[Guatemala]], where "wild" avocados may still be found.<ref>{{cite journal | last=Kopp | first=Lucille E. | year=1966 | title=A taxonomic revision of the genus ''Persea'' in the Western Hemisphere. (''Persea''-Lauraceae) | journal=Memoirs of the New York Botanical Garden | volume=14 | issue=1 | pages=1–117 }}</ref> The soft pulp of the fruit is eaten fresh or mashed into [[guacamole]].  The ancient peoples of Central America were also the first to cultivate several fruit-bearing species of ''[[Annona]]''.<ref name="Cronquist"/>  These include the [[custard-apple]] (''A. reticulata''), [[soursop]] (''A. muricata''), sweetsop or [[sugar-apple]] (''A. squamosa''), and the [[cherimoya]] (''A. cherimola'').  Both soursop and sweetsop now are widely grown for their fruits in the Old World as well.<ref name="Heywood">{{cite book | last=Heywood | first=V. H. (ed.) | year=1993 | title= Flowering Plants of the World | edition= updated | location=New York | publisher=Oxford University Press | isbn=0-19-521037-9 | pages=27–42 }}</ref>
 
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Some members of the magnoliids have served as important food additives.  Oil of [[sassafras]] was formerly used as a key flavoring in both [[root beer]] and in [[Sarsaparilla (soft drink)|sarsaparilla]].<ref>{{cite book | last=Hester |author2=Roy M. Harrison | year=2001 | title=Food safety and food quality | publisher=Royal Society of Chemistry | isbn=0-85404-270-9 |pages=118|first=R. E. }}</ref>  The primary ingredient responsible for the oil's flavor is [[safrole]], but it is no longer used in either the [[United States]] or [[Canada]].  Both nations banned the use of safrole as a food additive in 1960 as a result of studies that demonstrated safrole promoted [[liver]] damage and [[tumor]]s in mice.<ref name="Hayes">{{cite book | last=Hayes | first=Andrew Wallace | year=2001 | title= Principles and Methods of Toxicology | edition=4th | publisher=CRC Press | isbn=1-56032-814-2 | pages=518 }}</ref>  Consumption of more than a minute quantity of the oil causes [[nausea]], vomiting, hallucinations, and shallow rapid breathing. It is very toxic, and can severely damage the kidneys.<ref>{{cite web | title=Sassafras oil overdose |  work=New York Times |
 
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url=http://health.nytimes.com/health/guides/poison/sassafras-oil-overdose/overview.html | accessdate=2008-07-12 }}</ref> In addition to its former use as a food additive, safrole from either ''Sassafras'' or ''[[Ocotea cymbarum]]'' is also the primary precursor for synthesis of MDMA ([[methylenedioxymethamphetamine]]), commonly known as the drug ecstasy.<ref>{{cite journal | year=2005 | title= MDMA and MDA producers using Ocotea cymbarum as a precursor | url=http://www.usdoj.gov/dea/programs/forensicsci/microgram/mg1105/mg1105.html | journal=Microgram Bulletin | volume=XXXVIII | issue=11 }}</ref>
 
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[[File:Myris fragr Fr 080112-3294 ltn.jpg|thumb|left|[[Nutmeg]] fruits are a source of the [[hallucinogen]] [[myristicin]].]]
 
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Other magnoliids also are known for their narcotic, hallucinogenic, or paralytic properties.  The [[Polynesian culture|Polynesian]] beverage [[kava]] is prepared from the pulverized roots of ''Piper methysticum'', and has both sedative and [[narcotic]] properties.<ref name="Heywood"/>  It is used throughout the Pacific in social gatherings or after work to relax.  Likewise, some native peoples of the [[Amazon Rainforest|Amazon]] take a [[hallucinogen]]ic [[Herbal smokeless tobacco|snuff]] made from the dried and powdered fluid exuded from the bark of ''[[Virola]]'' trees.<ref name="Cronquist"/> Another hallucinogenic compound, [[myristicin]], comes from the spice nutmeg.<ref>{{cite journal | last=Shulgin | first=Alexander T. | date=1966-04-23 | title=Possible implication of myristicin as a psychotropic substance | journal=Nature | volume=210 | pages=380–384 | doi=10.1038/210380a0 | pmid=5336379 | issue=5034 }}</ref>  As with safrole, ingestion of nutmeg in quantities can lead to hallucinations, nausea, and vomiting, with symptoms lasting several days.<ref>{{cite journal | last=Panayotopoulos | first=D. J. |author2=D. D. Chisholm | year=1970 | title= Hallucinogenic effect of nutmeg | journal=British Medical Journal | volume=1 | issue=5698 | pages=754 | pmid=5440555 | doi=10.1136/bmj.1.5698.754-b | pmc=1699804 }}</ref>  A more severe reaction comes from poisoning by [[rodiasine]] and [[demethylrodiasine]], the active ingredients in fruit extract from ''[[Chlorocardium venenosum]]''.  These chemicals paralyze muscles and nerves, resulting in [[tetanus]]-like reactions in animals. The [[Cofán]] peoples of westernmost Amazon in [[Colombia]] and [[Ecuador]] use the compound as a poison to tip their arrows in hunting.<ref>{{cite journal | last=Kostermans | first=A. J. |author2=Homer V. Pinkley |author3=William L Stern  | year=1969 | title=A new Amazonian arrow poison: ''Ocotea venenosa'' | journal=Botanical Museum Leaflets, Harvard University | volume=22 | issue=7 | pages=241–252 }}</ref>
 
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Not all the effects of chemical compounds in the magnoliids are detrimental.  In previous centuries, sailors would use Winter's Bark from the [[South America]]n tree ''[[Drimys winteri]]'' to ward off the vitamin-deficiency of [[scurvy]].<ref name="Heywood"/> Today, [[benzoyl]] is extracted from ''[[Lindera benzoin]]'' (common spicebush) for use as a food additive and skin medicine, due to its anti-bacterial and anti-fungal properties.<ref>{{cite book | last=Zomlefer | first=Wendy B. | year=1994 | title= Guide to Flowering Plant Families | location=Chapel Hill | publisher=University of North Carolina Press | isbn=0-8078-2160-8 | pages=29–39 }}</ref> Drugs extracted from the bark of ''[[Magnolia]]'' have long been used in traditional Chinese medicine.  Scientific investigation of [[magnolol]] and [[honokiol]] have shown promise for their use in dental health.  Both compounds demonstrate effective anti-bacterial activity against the bacteria responsible for [[halitosis|bad breath]] and [[dental caries]].<ref>{{cite journal | last=Greenberg | first=M |author2=P. Urnezis |author3=M. Tian  | year=2007 | title=Compressed mints and chewing gum containing magnolia bark extract are effective against bacteria responsible for oral malodor | journal=Journal of Agricultural and Food Chemistry | volume=55 | issue=23 | pages=9465–9469 | doi=10.1021/jf072122h | pmid=17949053 }}</ref><ref>{{cite journal | last=Chang | first=B |author2=Lee Y |author3=Ku Y |author4=Bae K |author5=Chung C.  | year=1998 | title=Antimicrobial activity of magnolol and honokiol against periodontopathic microorganisms | journal=Planta Medica | volume=64 | issue=4 | pages=367–369 | doi=10.1055/s-2006-957453 | pmid=9619121 }}</ref>  Several members of the family [[Annonaceae]] are also under investigation for uses of a group of chemicals called [[acetogenin]]s.  The first acetogenin discovered was [[uvaricin]], which has anti-[[leukemic]] properties when used in living organisms.  Other acetogenins have been discovered with anti-[[malaria]]l and anti-tumor properties, and some even inhibit [[HIV]] replication in laboratory studies.<ref name="Pilar">{{cite book | last = Pilar Rauter | first = Amélia |author2=A. F. Dos Santos |author3=A. E. G. Santana | title = Natural Products in the New Millennium: Prospects and Industrial Application | url = https://books.google.com/books?id=4rrC7c_6OUoC | year = 2002 | publisher = [[Springer Science+Business Media]] | isbn = 1-4020-1047-8 | pages = 540 pages | chapter = Toxicity of Some species of ''Annona'' Toward ''Artemia Salina'' Leach and ''Biomphalaria Glabrata'' Say | chapterurl = https://books.google.com/books?id=4rrC7c_6OUoC&pg=PA264&lpg=PA264&source=web&ots=GVwQsxA_oK&sig=9U5mL2oGo14l_K6XnC8wb8k1a_M&hl=en | accessdate = 2008-07-12 }}</ref>
 
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Many magnoliid species produce [[essential oil]]s in their leaves, bark, or wood.  The tree ''[[Virola surinamensis]]'' (Brazilian "nutmeg") contains [[trimyristin]], which is extracted in the form of a fat and used in [[soap]]s and [[candle]]s, as well as in [[shortening]]s.<ref>{{cite journal | last= Pereira Pinto | first=Gerson | year=1951 | title= Contribuição ao estudo químico do Sêbo de Ucuúba | journal=Boletim Técnico do Instituto Agronômico do Norte | volume=23 | pages=1–63 }}</ref>  Other fragrant volatile oils are extracted from ''[[Aniba rosaeodora]]'' ([[bois-de-rose oil]]), ''[[Cinnamomum porrectum]]'', ''[[Cinnamomum cassia]]'', and ''[[Litsea odorifera]]'' for scenting soaps.<ref>{{cite journal | last= Kostermans | first= A. J. G. H. | year=1957 | title=Lauraceae | journal=Communication of the Forest Research Institute, Indonesia | volume=57 | pages=1–64 }}</ref> [[Perfume]]s also are made from some of these oils; [[ylang-ylang]] comes from the flowers of ''[[Cananga odorata]]'', and is used by [[Arab]] and [[Swahili people|Swahili]] women.<ref name="Heywood"/>  A compound called [[nutmeg butter]] is produced from the same tree as the spice of that name, but the sweet-smelling "butter" is used in perfumery or as a [[lubricant]] rather than as a food.
 
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Magnoliids are also important sources of spices and herbs used to flavor food, including the spices [[black pepper]], [[cinnamon]] and [[nutmeg]], and the herb [[bay laurel]].
 
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==See also==
 
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* [[Eudicots]]
 
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* [[Monocotyledon]]
 
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==References==
 
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{{Reflist|2}}
 
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==External links==
 
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{{Wikispecies|Magnoliids}}
 
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{{Commons category multi|Magnoliidae|magnoliids}}
 
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* [http://tolweb.org/magnoliids/20670 Tree of Life Magnoliids]
 
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{{taxonbar}}
 
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[[Category:Magnoliids| 01]]
 
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[[Category:Angiosperms]]
 
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[[Category:Eukaryote unranked clades]]
 

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