Editing Zingiberales

{{Short description|Order of flowering plants}}
{{expand Spanish|Zingiberales|date=February 2017}}
{{Automatic taxobox
| fossil_range = {{fossil range|80|0}}[[Late Cretaceous]] – Recent
| image = Starr 061212-2337 Alpinia purpurata.jpg
| image_caption = ''[[Alpinia purpurata]]''<br />([[Zingiberaceae]])
| taxon = Zingiberales
| authority = [[August Heinrich Rudolf Grisebach|Griseb.]]<ref name=Tropicos/><ref name=Grisebach/>
| subdivision_ranks = Families
| subdivision = {{collapsible list
| [[Canna (plant)|Cannaceae]] <small>[[Juss.]]</small> 
| [[Costaceae]] <small>[[Takenoshin Nakai|Nakai]]</small> 
| [[Heliconia]]ceae <small>[[Sydney Howard Vines|Vines]]</small> 
| [[Lowiaceae]] <small>[[Ridl.]]</small>
| [[Marantaceae]] <small>[[R.Br.]]</small>
| [[Musaceae]] <small>[[Juss.]]</small>
| [[Strelitziaceae]] <small>[[Hutch.]]</small> 
| [[Zingiberaceae]] <small>[[Ivan Martinov|Martinov]]</small>
}}
| type_genus = ''[[Zingiber]]''
| type_genus_authority = [[Mill.]]
| synonyms = * Amomales <small>[[Lindl.]]</small>
* Cannales <small>[[R.Br.]] ex [[Bercht.]] et [[J.Presl]]</small>
* Lowiales <small>Takht. ex. Reveal & Doweld</small>
* [[Scitamineae]]
* Zingiberanae <small>[[Takht.]] ex [[James Reveal|Reveal]]</small>
* Zingiberidae <small>[[Arthur Cronquist|Cronquist]]</small>
* Zingiberides
* Zingiberiflorae
| synonyms_ref = <ref name=Johansson/>{{sfn|APG|1998}}{{sfn|APG II|2003}}
| diversity = 99 genera{{citation needed|date=August 2017}}
}}
{{Infobox
|bodystyle   = width:21em
|subheader=[[Floral formula]]
|data1={{resize|120%|B X–$ ☿ {{color|red|P3+3}} {{color|olive|A3+3(1-6)}} {{color|blue|{{overline|G}}(3)}}}}<br />
 Bracteate, Zygomorphic-Asymmetric, Bisexual<br />{{color|red|Perianth: 6 tepals in 2 whorls of 3}}<br />{{color|olive|Stamens:  2 whorls of 3 frequently modified to 1–6}}<br />{{color|blue|Ovary: Inferior - 3 fused carpels}}
}}

The '''Zingiberales''' are [[flowering plant]]s forming one of four [[order (biology)|orders]] in the [[commelinids]] [[clade]] of [[Monocotyledon|monocots]], together with its [[sister group|sister]] order, [[Commelinales]]. The order includes 68 [[Genus|genera]] and 2,600 [[species]]. Zingiberales are a unique though [[plant morphology|morphologically]] diverse order that has been widely recognised as such over a long period of time. They are usually large [[herbaceous plant]]s with [[rhizome|rhizomatous]] root systems and lacking an aerial [[plant stem|stem]] except when flowering. Flowers are usually large and showy, and the [[stamen]]s are often modified ([[staminode]]s) to also form colourful petal-like structures that attract [[pollinator]]s.

Zingiberales contain eight families that are informally considered as two groups, differing in the number of fertile stamens. A "[[Banana-families|banana group]]" of four families appeared first and were named on the basis of large [[banana]]-like leaves. Later, a more genetically coherent ([[Monophyly|monophyletic]]) "[[ginger-families|ginger group]]" appeared, consisting of the remaining four families. The order, which has a [[fossil]] record, is thought to have originated in the [[Early Cretaceous]] period between 80 and 120 million years ago (Mya), most likely in [[Australia]], and [[evolutionary divergence|diverged]] relatively rapidly with the families as they are known today established by the end of the period (66 Mya). Zingiberales are found throughout the tropics (pantropical) with some extension into subtropical and temperate climates. They rely on [[insect]]s for [[pollination]], together with some [[bird]]s and small animals.

The order includes many familiar plants, and are used as ornamental plants ([[Strelitzia reginae|Bird of Paradise flower]], [[heliconia]]s, [[prayer-plant]]s), food crops ([[banana]]s, [[Plantain (cooking)|plantains]], [[arrowroot]]), spices and [[traditional medicine]]s ([[ginger]], [[cardamom]], [[turmeric]], [[galangal]], [[fingerroot]] and [[myoga]]).

== Description ==

Zingiberales are one of an [[Ecology|ecologically]] and [[plant morphology|morphologically]] diverse and species-rich [[order (biology)|order]] of [[Monocotyledon|monocots]], with one of the most distinct [[Flower|floral morphology]].{{sfn|Salvi et al|2015}}{{sfn|Deng et al|2016}}{{sfn|Kirchoff et al|2009}} They are large [[Rhizome|rhizomatous]] [[herbaceous plant]]s but lacking an aerial [[plant stem|stem]], except when flowering.<ref name=apw/>

[[Leaf|Leaves]] usually [[petiole (botany)|petiolate]] with distinct petiole and lamina, leaf arrangement [[Phyllotaxis|distichous]] (spiral in Musaceae). Venation pinnate-parallelodromous, with midrib (midvein), S-shaped lateral veins and fine transverse venation.<ref name=Johansson/>{{sfn|Simpson|2011|loc=p. 47}}

[[Flower]]s are generally large and showy, following the general monocot pattern, with [[inflorescence]]s in [[thyrse]]-like spikes, [[Floral symmetry|zygomorphic]] to asymmetric, with two [[Merosity|trimerous]] [[whorl (botany)|whorls]] of [[tepal]]s. [[Gynoecium]] tricarpellate, [[ovary (botany)|ovary]] epigynous (inferior), two trimerous [[androecial]] whorls with [[stamen]]s 6, 5 or 1. Stamens have elongated sterile filaments to which are attached anthers, distally, comprising about half of the length of the total stamen. Septal [[nectar]]ies often present.{{sfn|Kirchoff et al|2009}}{{sfn|Bartlett et al|2010}}{{sfn|Specht et al|2012}} [[Pollen]] [[Sulcus (morphology)|sulcate]] but often inaperturate (lacking apertures).

[[Fruit]] [[Capsule (fruit)|capsular]] or schizocarp. [[Phytochemistry]]: Often containing [[raphide]]s,<ref name=apw/><ref name=Kubitzki/>

Specific characteristics which help to distinguish this order include a herbaceous [[arborescent]] [[Plant stem|stem]], distichous [[phyllotaxy]], large petiolate leaves in which the petioles are often long, parallel and transverse venation diverging laterally from a prominent common midrib, and [[inflorescence]]s of conspicuous colorful [[bract]]s (bracteate inflorescence) and the substitution of one to five rudimentary [[wikt:staminodia|staminodia]] for fertile [[stamen]]s.<ref name=Byng/>{{sfn|Heliconia Society|2017}}{{sfn|Kress|1990}}

Leaf architecture is useful for distinguishing families within Zingiberales, based on vein pattern type, vein length per area, and other aspects of vein architecture such as angle of vein divergence, with three main types of venation recognised. These are the ''Zingiber''-type, with square to vertically elongate [[areole]]s, the ''Costus''-type, with horizontally elongate areoles and the ''Orchidantha''-type with cross veins spanning multiple parallel veins.{{sfn|Salvi et al|2015}}{{sfn|Triplett|Kirchoff|1991}}

===Apomorphies===

The [[Synapomorphy and apomorphy|apomorphies]] (derived characteristics common to a taxonomic group) are considered to be specialised isomorphic root hair cells, penni-parallel leaf venation, supervolute [[Glossary of leaf morphology|ptyxis]] (left and right halves of immature leaf lamina rolled into each other), diaphragmed air chambers in leaves and stem, presence of intracellular [[Silicon dioxide|silica]] bodies, [[Ovary (botany)|epigynous]] flowers and an inferior [[ovary (botany)|ovary]], pollen grains without distinctive aperture but with a reduced exine layer and an elaborated intine layer, nuclear endosperm development, and arillate [[seed]]s.{{sfn|Simpson|2011|loc=p. 192}}{{sfn|Stephen|2015}}{{sfn|Kress et al|2001}}

==Taxonomy==
 <blockquote>''"The Zingiberiflorae, whether treated as a separate superorder, as here, or an order in a more widely circumscribed unit, is one of the most indisputably natural suprafamilial groups."''{{sfn|Dahlgren |Clifford |Yeo|1985}}</blockquote>

=== History ===

The Zingiberales have always been considered a unique and coherent ([[Monophyly|monophyletic]]) group, although accounting for <4% of extant monocots, which has led some authors to suggest they should constitute a higher taxonomic rank than order.{{sfn|Kress et al|2001}} For a brief history of the taxonomy of this order, see [[Scitamineae]], and Kress 1990.{{sfn|Kress|1990}} They were first described by [[August Grisebach]], their [[Author citation (botany)|botanical authority]], in 1854 as Zingiberides, an order of monocotyledons, subdivided into two families, Scitamineae and Musaceae.<ref name=Grisebach/>

Based on [[plant morphology|morphological]] grounds alone, early [[list of systems of plant taxonomy|systems]], such as [[Bentham & Hooker system|Bentham and Hooker]] (1883) placed the Scitamineae as an ''Ordo'' (family) of the Epigynae alliance in the monocotyledons, incorporating both of Grisebach's families.<ref name = BH/> Later systems such as the [[Engler system]] (1903)<ref name=Engler/> and the [[Wettstein system]] (1924),{{sfn|Wettstein|1924|loc=p. 902}} also considered Scitamineae as a monocotyledon order and were influential for a long period of time. Variants included Scitaminales ([[Eugenius Warming|Warming]] 1912).{{sfn|Warming|1912}} [[Hutchinson system|Hutchinson]] (1926), although initially using Scitamineae, later followed [[Takenoshin Nakai]] (1930).<ref name=Nakai1930/>{{sfn|Nakai|1941}} in adopting Zingiberales as the name for the order (6 families) in Division Calyciferae,{{sfn|Hutchinson|1926–1934}} although credit is generally given to Nakai. This usage was followed by [[Takhtajan system|Takhtajan]] (1966) within [[Order (biology)|superorder]] [[Lilianae]]<ref name=Takhtajan/> and by [[Rolf Dahlgren|Dahlgren]] (1985) in its own superorder Zingiberiflorae.{{sfn|Dahlgren |Clifford |Yeo|1985}}

In contrast the [[Cronquist system]] retained Scitamineae as the name for this order with eight families, but organised the order in the subclass [[Zingiberidae]] of the class [[Liliopsida]] ([[monocotyledon]]s).

=== Modern era ===

Using [[molecular phylogenetics]], which was first applied to the order in 1993,{{sfn|Smith et al|1993}} the [[Angiosperm Phylogeny Group]] (APG) system (1998),{{sfn|APG|1998}} (which generally followed Dahlgren, but with fewer divisions) confirmed the position of Zingiberales as a monophyletic order within the monocots, placing it in the [[commelinoid]] clade, as [[sister group]] to [[Commelinales]], which Dahlgren had treated within a separate superorder. This was an ordinal system that did not examine subordinal structure. The 2003 revision ([[APG II]]) changed commelinoid to commelinid, but not the relationships,{{sfn|APG II|2003}} and this remained unchanged in the subsequent 2009 [[APG III system]]{{sfn|APG III|2009}} and 2016 [[APG IV system]] without addressing interfamilial relationships.{{sfn|APG IV|2016}}

{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
| style="background:#fff; padding:2.5px" | clade monocots :
* clade [[commelinids]]:
::* order [[Arecales]] (palms)
::* order [[Commelinales]] (spiderwort, water hyacinth)
::* order [[Poales]] (grasses, rushes, bromeliads)
::* order '''Zingiberales''' <small>[[August Heinrich Rudolf Grisebach|Griseb.]]</small>(gingers, banana)
|{{cladogram | title= {{anchor|Clad}}Cladogram: [[phylogenetics|Phylogeny]] of the commelinids (APG IV){{sfn|APG IV|2016}}|style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1='''[[commelinids]]'''
|1={{clade|style=
    |1=[[Arecales]]
    |2=[[Poales]]
    |3={{clade
        |1={{clade
            |1='''Zingiberales'''
            |2=[[Commelinales]]
            }}
        }}
    }}

}}}}
|}
{{clear}}

=== Subdivision ===

The order, which now has more than 2,600 species, distributed in 68 genera over eight families, has been subdivided from early times. In the [[Bentham & Hooker system]] (1883), their Ordo Scitamineae had four [[tribe (biology)|tribes]]: Zingibereae, Maranteae, Canneae, and Museae. These have become progressively divided to form the modern [[Phylogenetics|phyletic]] classification into the following monophyletic families: Zingiberaceae (gingers), Musaceae (bananas), Heliconiaceae (heliconias), Strelitziaceae (bird-of-paradise), Costaceae (spiral gingers), Cannaceae (canna lilies), Marantaceae (prayer plants), and Lowiaceae (Orchidantha).{{sfn|Salvi et al|2015}}{{sfn|Kress et al|2001}} The [[APG II system]] (2003) provided a classification of families for the first time, retaining Kress's eight families.{{sfn|APG II|2003}}

;Families (genera/species){{sfn|Christenhusz|Byng|2016}}
* order Zingiberales <small>Griseb.</small>
** family Cannaceae <small>[[Juss.]]</small> (1/10 ''[[Canna (plant)|Canna]]'' cannas)
** family [[Costaceae]] <small>[[Takenoshin Nakai|Nakai]]</small> (7/143 ''e.g.'' ''[[Costus]]'' spiral gingers)
** family [[Heliconia]]ceae <small>[[Sydney Howard Vines|Vines]]</small> (1/194 ''[[Heliconia]]'' heliconias)
** family Lowiaceae <small>[[Ridl.]]</small> (1/18 ''[[Orchidantha]]'')
** family [[Marantaceae]] <small>[[R.Br.]]</small> (29/570 ''e.g.'' ''[[Maranta (plant)|Maranta]]'' prayer plants)
** family [[Musaceae]] <small>[[Juss.]]</small> (3/91 ''e.g.'' ''[[Musa (genus)|Musa]]'' bananas)
** family [[Strelitziaceae]] <small>[[Hutch.]]</small> (3/7 ''e.g.'' ''[[Strelitzia]]'' birds of paradise)
** family [[Zingiberaceae]] <small>[[Ivan Martinov|Martinov]]</small> (50/1,600 ''e.g.'' ''[[Zingiber]]'' gingers)

<gallery mode="packed" caption="Families of Zingiberales">
Image:Canna Coccinea 2003072001.jpg|''[[Canna coccinea]]''<br />([[Canna (plant)|Cannaceae]])
Image:Tapeinochilos ananassae2.jpg|''[[Tapeinochilos ananassae]]''<br />([[Costaceae]])
Image:Heliconia latispatha (Starwiz).jpg|''[[Heliconia latispatha]]''<br />([[Heliconia]]ceae)
Image:Orchidantha maxillarioides - Botanischer Garten - Heidelberg, Germany - DSC01154.jpg |''[[Orchidantha|Orchidantha maxillarioides]]''<br />([[Lowiaceae]])
Image:Maranta leuconeura3.jpg|''[[Maranta leuconeura]]''<br />([[Marantaceae]])
Image:Musa rosea (11136399706).jpg|''[[Musa rosea ]]''<br />([[Musaceae]])
Image:Strelitzia reginae flower 2016022.jpg|''[[Strelitzia reginae ]]''<br />([[Strelitziaceae]])
Image:Tropical plant hilo5.jpg|''[[Etlingera elatior]]''<br />([[Zingiberaceae]])
</gallery>

Based on morphology alone, the Zingiberales have been considered to form two main groups, each with four families, utilising the number of fertile stamens;

;[[Banana-families]]
* [[Musaceae]], [[Strelitziaceae]], [[Orchidantha|Lowiaceae]], [[Heliconia]]ceae. A [[Paraphyly|paraphyletic]] basal assemblage with 5 or (rarely) 6 fertile stamens at maturity, arranged in as trimerous inner and outer whorls. In those with five stamina, the sixth may regress and be absent (Strelitziaceae and Lowiaceae, some Musaceae) or develop as an infertile staminode (Heliconiaceae, some Musaceae). Petals and stamens are often fused at the base to form a [[Hypanthium|floral tube]]. These are known as the banana-families or the bananas on the basis of large banana-like leaves. For this reason these four families were previously all included in Musaceae, but the exact relationship between them remains somewhat uncertain;{{sfn|Kirchoff et al|2009}}{{sfn|Bartlett et al|2010}}{{sfn|Kress et al|2001}}

;[[Ginger-families]]
* [[Zingiberaceae]], [[Costaceae]], [[Canna (plant)|Cannaceae]], [[Marantaceae]] (the gingers). A [[monophyletic]] derived terminal clade with the number of fertile stamens reduced to one (Zingiberaceae, Costaceae) or to one half, with a single [[theca]] (Cannaceae, Marantaceae). The remaining components of the androecium develop as four or five elaborate petaloid staminodia, highly modified from sterile stamens. This group may have one (Cannaceae, Marantaceae) or two (Zingiberaceae, Costaceae) anther pollen sacs. The infertile stamina are [[homology (biology)|homologous]] with the fertile stamina in the bananas and other monocots) but assume the structure and function of [[petal]]s as pollinator attraction. This group demonstrate complex patterns of fusion among their floral organs including the staminodes. In Zingiberaceae and Costaceae the staminodes fuse to form a staminodial labellum which provides much of the floral display. In general the flowers of this group display higher degrees of organ fusion and specialisation.{{sfn|Kirchoff et al|2009}}{{sfn|Bartlett et al|2010}}{{sfn|Kress et al|2001}}

==== Phylogeny ====
Using combined morphology and molecular data, Kress and colleagues (1993, 1995, 2001) confirmed the broad separation into two clades based on morphology alone, and produced an infraordinal structure.{{sfn|Kress et al|2001}}{{sfn|Smith et al|1993}}{{sfn|Kress|1995}} In this scheme, they divided the families of the order as follows, with the ginger group as one suborder, and the banana group divided amongst three separate suborders:{{sfn|Kress|Specht|2005}}

Suborder [[Ginger-families|Zingiberineae]] <small>Kress</small> ("gingers"; 2 superfamilies)
* Superfamily Cannariae <small>Kress</small>
** Cannaceae <small>A.L. Jussieu</small>
** Marantaceae <small>Petersen</small>
* Superfamily Zingiberariae <small>Kress</small>
** Costaceae <small>Nakai</small>
** Zingiberaceae <small>Lindley</small>
Suborder Strelitziineae <small>Kress </small>
** Lowiaceae <small>Ridley</small>
** Strelitziaceae <small>Hutchinson</small>
Suborder Musineae <small>Kress</small>
** Musaceae <small>A.L. Jussieu</small>
Suborder Heliconiineae <small>Kress</small>
** Heliconiaceae <small>Nakai</small>

While the two sister family groups that constitute the Zingiberineae ([[#CladI|Cladogram I]]) and also the basal Strelitziineae (Strelitziaceae-Lowiaceae) sister group ([[#CladII|Cladogram II]]) were strongly supported, the position of Musaceae and Heliconiaceae were not. In the above model (Model 1), Musaceae appears as the first branching family, and Heliconiaceae placed as sister to the Zingiberineae ([[#CladIII|Cladogram III]]). Other studies placed these and the Strelitziaceae-Lowiaceae sister group in a [[Speciation#Genetics|trichotomy]] with the remaining families.{{sfn|Givnish et al.|2006}} While a revision of the first model (Model 2) placed Heliconiaceae as the first branching lineage with Musaceae in a sister relationship with Strelitziineae, which in turn was sister to Zingiberineae ([[#CladIV|Cladogram IV]]).{{sfn|Barrett et al|2014}} A third model (Model 3) supports Musaceae as the basal group but places Heliconiaceae as sister to Strelitziineae ([[#CladV|Cladogram V]]).{{sfn|Deng et al|2016}} The failure to resolve the ancient rapid divergences of this order with multi-gene phylogenies and plastid data has important implications for understanding the evolution of characteristics.{{sfn|Barrett et al|2014}}{{sfn|Sass et al|2016}}

{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
|{{cladogram | title= {{anchor|CladI}}Cladogram I: [[phylogenetics|Phylogeny]] of Suborder [[Ginger-families|Zingiberineae]] (gingers)
|style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1=Zingiberineae
|1={{clade
|label1=Zingiberariae
|1={{clade
    |1=Zingiberaceae
    |2=Costaceae}}
|label2=Cannariae
|2={{clade
    |1=Cannaceae
    |2=Marantaceae}}
    }}
}}}}
|}
{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
|{{cladogram | title= {{anchor|CladII}}Cladogram II: Phylogeny of Suborder Strelitziineae
|style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1=Strelitziineae
|1={{clade
|1={{clade
    |1=Lowiaceae
    |2=Strelitziaceae}}
}}}}}}
|}
{{clear}}
{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
|{{cladogram
| title= {{anchor|CladIII}}Cladogram III: Phylogeny of Zingiberales (Model 1){{sfn|Kress et al|2001}}
| caption= Dashed line -- indicates weak support
| style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1=Zingiberales
|1={{clade
 |1={{clade
  |1={{clade
     |1=Zingiberineae
     |2=Heliconiaceae|state2=dashed}}
  |2=Strelitziineae}}
|2=Musaceae|state2=dashed}}
}}
}}
|}
{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
|{{cladogram 
| title= {{anchor|CladIV}}Cladogram IV: Phylogeny of Zingiberales (Model 2){{sfn|Barrett et al|2014}}{{sfn|Barrett et al|2016}}
| caption= Dashed line -- indicates weak support
|style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1=Zingiberales
|1={{Clade
 |1={{clade
    |1=Zingiberineae
    |2={{clade
|      1=Musaceae|state1=dashed
|      2=Strelitziineae}}
}}
 |2=Heliconiaceae|state2=dashed}}
}}}}
|}
{|align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
|{{cladogram 
| title= {{anchor|CladV}}Cladogram V: Phylogeny of Zingiberales (Model 3){{sfn|Deng et al|2016}}
| caption= Dashed line -- indicates weak support
|style=background:#eef
| align=center
| cladogram={{clade|style=font-size:95%;line-height:75%; background:#eef
|label1=Zingiberales
|1={{Clade
|1={{clade
    |1=Zingiberineae
    |2={{clade
|      1=Heliconiaceae|state1=dashed
|      2=Strelitziineae}}
}}
|2=Musaceae|state2=dashed}}
}}}}
|}
{{clear}}
Finally in 2016 Sass and colleagues, using multiplexed exon capture were able to resolve the entire phylogenetic tree with high support. This confirmed the place of Musaceae as sister to the remaining families, confirming Model 3.{{sfn|Sass et al|2016}}
{{cladogram
| title= {{anchor|CladVI}}Cladogram VI: Phylogeny of Zingiberales<br />(Final Model){{sfn|Sass et al|2016}}
| align=center
| cladogram={{clade|style=font-size:100%;line-height:85%
|label1=Zingiberales
|1={{clade
 |1={{clade
  |1={{clade
   |label1=[[Ginger-families|Zingiberineae]]
   |1={{clade
      |label1=Zingiberariae
      |1={{clade
         |1=[[Zingiberaceae]]
         |2=[[Costaceae]]}}
      |label2=Cannariae
      |2={{clade
         |1=[[Canna (plant)|Cannaceae]]
         |2=[[Marantaceae]]}}}}
   |2={{clade
    |label1=Strelitziineae 
    |1={{clade
       |1= [[Lowiaceae]]
       |2= [[Strelitziaceae]]}}
    |2=[[Heliconia]]ceae}}
}}}}
|2={{clade
|1=[[Musaceae]]}}}}
}}
}}

===Evolution ===

The common ancestor of the Zingiberales together with those of its sister order, the [[Commelinales]], is estimated to have originated 158 Mya (million years ago) in the [[Early Cretaceous]], with separation of the two orders between 80 and 124 Mya, and with rapid [[evolutionary radiation|radiation]] into the major lineages in the mid Cretaceous ca. 60–100 Mya and six of the eight families established by the end of the Cretaceous,{{sfn|Kress|Specht|2005}}{{sfn|Sass et al|2016}} Estimates of [[crown group]] age (most recent common ancestor of the sampled species of the clade of interest) vary widely between 34 and 110 Mya, and may need revision in the light of developing knowledge of the [[phylogenetic tree|topology]] of the order.<ref name=apw/>{{sfn|Sass et al|2016}}

Fossil-calibrated molecular estimates suggest a date of 110 to 80 Mya for the diversification of primary lineages.{{sfn|Barrett et al|2014}} If Musaceae is the stem family of the order, as seems likely this places the origin of Zingiberales ca. 124&nbsp;Mya, with diversification occurring ca. 110 Mya in the middle Cretaceous. That origin (124 Mya) was congruent with the breakup of the southern land mass, [[Gondwana]].{{sfn|Deng et al|2016}}

Probably the ancestral Zingiberales were distributed in tropical Gondwanal and encompassing present-day Americas, Africa, and Southeast Asia. Within this land mass, Australia seems the most likely ancestral area, with subsequent dispersals between Africa and neotropical America. Earlier studies had implied Southeast Asia as the origin.{{sfn|Kress|Specht|2005}} The current distribution of the Zingiberales seems to be a product of numerous secondary and tertiary dispersal events between the major tropical regions of the world.{{sfn|Deng et al|2016}}

Zingiberales demonstrate an evolutionary trend in the [[ontogeny]] of the [[perianth]] (sepals and petals). The appearance of a dimorphic perianth (in which petals and sepals differ in appearance) is variable throughout the commelinid monocots, with a transition from an undifferentiated monomorphic perianth to a dimorphic one occurring independently in the two sister orders, Commelinales and the Zingiberales.{{sfn|Specht et al|2012}} The evolution of floral morphology within Zingiberales demonstrates a marked correlation between the reduction of the number of fertile stamens, and increased petaloidy.{{sfn|Deng et al|2016}} The ancestral Zingiberales flower is thought to have had 5–6 fertile stamens, following which the staminode evolved in the lineage leading to Heliconiaceae+Zingiberineae, finally leading to 2–5 staminodia dominating the floral display.{{sfn|Specht et al|2012}}

The phylogenetic diversification and biogeographic dispersal of the Zingiberales was, in part, driven by the evolutionary radiation and diversification of their associated animal pollinators, which include [[bat]]s, [[bird]]s, non-flying [[mammal]]s and [[insect]]s.{{sfn|Kress|Specht|2005}} Six of the eight families of the Zingiberales contain taxa specialised for pollination by vertebrates, which appears to be the [[Cladistics|plesiomorphic]] state in the order.{{sfn|Kress|Specht|2005}} Of these six families two are exclusively vertebrate-pollinated ([[Strelitziaceae]], [[Heliconia]]ceae).{{sfn|Kress|Specht|2005}} Pollination by insects also occurs in six families with one ([[Marantaceae]]) or possibly two ([[Orchidantha|Lowiaceae]]) families predominantly specialised for insect visitors.{{sfn|Kress|Specht|2005}}

==== Fossil record ====

Seed and fruit fossils of the Zingiberales appear in the [[Santonian]] of the Late Cretaceous, ca. 85&nbsp;Mya, and thought to represent Musaceae and Zingiberaceae, but the place of the oldest Zingiberales fossil, ''Spirematospermum'' with its distinctive seeds remains uncertain, but is most likely Zingiberaceae.<ref name=apw/> The leaf fossil record for Zingiberales also extends back to the Late Cretaceous.{{sfn|Deng et al|2016}} Other fossil records include rhizomes and [[phytolith]]s.{{sfn|Barrett et al|2014}}{{sfn|Sass et al|2016}}

=== Diversity and biogeography ===

The four families of the basal banana group exhibit less taxonomic diversity than the terminal ginger group. The three genera of Musaceae are distributed in tropical Asia and Africa, although fossil evidence reveals their presence in North America and Africa in the [[Tertiary]], while the three genera of Strelitziaceae are [[Allopatric speciation|allopatric]] (isolated from each other) being found in Madagascar, southern Africa and the Amazon Basin respectively and Lowiaceae occurs in Southeast Asia. The largest family in this group, the Heliconiaceae is primarily [[neotropical]], but also occurs in the Pacific from Samoa to Sulawesi.{{sfn|Kress|Specht|2005}}

Of the four Zingiberineae (gingers) families, three (Zingiberaceae, Costaceae, Marantaceae) are pantropical. The fourth, Cannaceae is restricted to the New World, although widely cultivated. This suborder contains the two largest families (Zingiberaceae and Marantaceae) and the largest number of species.{{sfn|Kress|Specht|2005}}

== Distribution and habitat ==

Zingiberales are [[pantropical]] and occur predominantly in the wet tropical regions of Asia, Africa, and the Americas, occupying nearly all tropical wet lowlands or middle elevation forests as part of the [[understory]] flora.{{sfn|Salvi et al|2015}} In addition five genera from three of the Zingiberineae families, including ''[[canna (plant)|Canna]]'' extend into subtropical and temperate regions. Of the eight families, Heliconiaceae, Marantaceae, and Costaceae are predominantly neotropical and Zingiberaceae most prevalent in Southeast Asian wet understory habitats. These are mainly small to medium-sized herbaceous taxa or vines. While some herbaceous Zingiberaceae such as ''[[Alpinia|Alpinia boia]]'' can attain a height of ten metres, only one species is a true [[canopy (biology)|canopy]] plant (''[[Ravenala]] madagascariensis'' – Strelitziaceae). The latter, a Madagascar [[endemic]], has thick, palm-like trunks which push the fan-shaped crown of leaves up into the top layers of the forest. Some Zingiberales prefer a greater degree of light and are found in forest glades or margins, or in open secondary growth along streams and rivers.{{sfn|Kress|Specht|2005}}

The large family Zingiberaceae has a number of subfamilies, one of which, Zingiberoideae, has members that have adapted to [[Southeast Asia]]'s [[monsoonal]] climates. They do so by becoming [[dormancy|dormant]] in the dry season, which may last four to six months, shedding all above ground parts, existing only as underground fleshy underground rhizomes, some of which have starch rich [[tubers]]. With the onset of the wet season, they send up shoots and complete their life cycle during this time. Some taxa within Marantaceae, Costaceae, and Musaceae also occur in these habitats and have adapted in this way, but no Zingiberales occur in true [[desert]] regions.

In contrast some Zingiberales, including taxa from Marantaceae, Heliconiaceae, Cannaceae (e.g. ''[[Canna glauca]]'') have adopted an [[aquatic ecosystem|aquatic]] habitat and are found along river margins, ponds, and swampy areas, with their rhizomes rooted underwater.{{sfn|Kress|Specht|2005}}

== Ecology ==

Zingiberales forms a major component of tropical and subtropical ecosystems. Many of these plants have formed specialised [[pollination]] relationships with [[mammal]]s (e.g., [[bat]]s and [[lemur]]s), [[bird]]s and [[insect]]s (e.g., [[bee]]s, [[dung beetle]]s, [[moth]]s and [[Butterfly|butterflies]]) through alterations in floral form.{{sfn|Bartlett et al|2010}}

== Uses ==

Many Zingiberales are [[horticulture|horticulturally]] important and grown as [[ornamental plant]]s, e.g., ''[[Heliconia]]'' (false [[bird-of-paradise]]), ''[[Strelitzia]]'' (bird-of-paradise), ''[[Maranta (plant)|Maranta]]'' (prayer plants) and ''[[Canna (plant)|Canna]]''. Others are crop plants with [[culinary]] usage, e.g., ''[[Musa (genus)|Musa]]'' (bananas, plantains) and ''[[Zingiber]]'' (ginger). Zingiberales also include sources of [[traditional medicine]]s and [[spice]]s, ''e.g.'' [[Alpinieae]] such as ''[[Elettaria]]'' and ''[[Amomum]]'' ([[cardamom]]) and [[galanga]] and ''[[Curcuma]]'' ([[turmeric]]).{{sfn|Bartlett et al|2010}}{{sfn|Heliconia Society|2017}}

== See also ==
* [[Scitamineae]]

==References==
{{Reflist|20em|refs=

<ref name = BH>{{harvnb|Bentham|Hooker|1862–1883|loc=[https://www.biodiversitylibrary.org/item/14690#page/197/mode/1up Scitamineae pp. 636–657]}}</ref>

<ref name=Byng>{{harvnb|Byng|2014|loc=[https://books.google.com/books?id=yoLaBAAAQBAJ&pg=PA101 Zingiberales]}}</ref>

<ref name=Engler>{{harvnb|Engler|1903|loc=[https://www.biodiversitylibrary.org/item/63778#page/135/mode/1up Scitamineae]}}</ref>

<ref name=Grisebach>{{harvnb|Grisebach|1854|loc=[https://archive.org/stream/grundrissdersys00grisgoog#page/n169/mode/2up Zingiberides p. 167]}}</ref>

<ref name=Johansson>{{harvnb|Johansson|2013|loc=[http://angio.bergianska.se/monocots/Cannales/Cannales.html Cannales]}}
</ref>

<ref name=Kubitzki>{{harvnb|Kubitzki|1998|loc=[https://books.google.com/books?id=bNQDsSmx2roC&pg=PA2 Zingiberales pp. 2ff.]}}</ref>

<ref name=Nakai1930>{{harvnb|Nakai|1930|loc=p. 49}}</ref>

<ref name=Takhtajan>{{harvnb|Takhtajan|1966|loc=[https://books.google.com/books?id=xljPEon9Qy0C&pg=PA498 Zingiberales p. 498]}}</ref>

<ref name=apw>{{harvnb|Stevens|2017|loc=[http://www.mobot.org/mobot/research/apweb/orders/zingiberalesweb.htm#Zingiberales Zingiberales]}}</ref>

<ref name=Tropicos>{{harvnb|Tropicos|2015|loc=[http://legacy.tropicos.org/Name/43000104 Zingiberales]}}</ref>
}}

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* {{cite book|last1=Kress|first1=W.J.|last2=Specht|first2=C.D.|title=Between Cancer and Capricorn: Phylogeny, evolution and ecology of the primarily tropical Zingiberales.|pages=459–478|url=http://spechtlab.berkeley.edu/sites/default/files/spechtlab/publications/12%20Kress%20and%20Specht%202005.pdf|ref={{harvid|Kress|Specht|2005}}}}, in {{harvtxt|Friis|Balslev|2005}}
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* {{cite book|last1=Warming|first1=Eugenius|author-link=Eugenius Warming|title=Frøplanterne (Spermatofyter)|date=1912|publisher=Gyldendalske Boghandel Nordisk Forlag|location=Kjøbenhavn|language=da|url=https://books.google.com/books?id=TVU-AQAAMAAJ|access-date=28 December 2015}}
* {{cite book| first= Richard|last= Wettstein|author-link=Richard Wettstein|year = 1924|edition=3rd|title = Handbuch der Systematischen Botanik 2 vols.|url=http://biolib.mpipz.mpg.de/library/authors/author_00267_de.html|access-date=15 April 2015}}

=== Articles ===

* {{cite journal|last1=Barrett|first1=C. F.|last2=Specht|first2=C. D.|last3=Leebens-Mack|first3=J.|last4=Stevenson|first4=D. W.|last5=Zomlefer|first5=W. B.|last6=Davis|first6=J. I.|title=Resolving ancient radiations: can complete plastid gene sets elucidate deep relationships among the tropical gingers (Zingiberales)?|journal=[[Annals of Botany]]|date=25 November 2013|volume=113|issue=1|pages=119–133|doi=10.1093/aob/mct264|pmid=24280362|ref={{harvid|Barrett et al|2014}}|pmc=3864734}}
* {{cite journal|last1=Barrett|first1=Craig F.|last2=Baker|first2=William J.|last3=Comer|first3=Jason R.|last4=Conran|first4=John G.|last5=Lahmeyer|first5=Sean C.|last6=Leebens-Mack|first6=James H.|last7=Li|first7=Jeff|last8=Lim|first8=Gwynne S.|last9=Mayfield-Jones|first9=Dustin R.|last10=Perez|first10=Leticia|last11=Medina|first11=Jesus|last12=Pires|first12=J. Chris|last13=Santos|first13=Cristian|last14=Wm. Stevenson|first14=Dennis|last15=Zomlefer|first15=Wendy B.|last16=Davis|first16=Jerrold I.|title=Plastid genomes reveal support for deep phylogenetic relationships and extensive rate variation among palms and other commelinid monocots|journal=New Phytologist|date=January 2016|volume=209|issue=2|pages=855–870|doi=10.1111/nph.13617|pmid=26350789|ref={{harvid|Barrett et al|2016}}|doi-access=free}}
* {{cite journal|last1=Bartlett|first1=Madelaine E.|last2=Specht|first2=Chelsea D.|title=Evidence for the involvement of GLOBOSA-like gene duplications and expression divergence in the evolution of floral morphology in the Zingiberales|journal=[[New Phytologist]]|date=July 2010|volume=187|issue=2|pages=521–541|doi=10.1111/j.1469-8137.2010.03279.x|pmid=20456055|ref={{harvid|Bartlett et al|2010}}|doi-access=free}}
* {{cite journal |last1=Christenhusz|first1=Maarten JM|last2=Byng|first2= J. W.  |author-link1=Maarten Christenhusz|author-link2=James W. Byng|name-list-style=amp | 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 }}
* {{cite journal|last1=Deng|first1=Jiabin|last2=Gao|first2=Gang|last3=Zhang|first3=Yan|last4=He|first4=Fengmei|last5=Luo|first5=Xuqiang|last6=Zhang|first6=Fengtai|last7=Liao|first7=Xinrong|last8=Ahmad|first8=Khawaja Shafique|last9=Yang|first9=Ruiwu|title=Phylogenetic and ancestral area reconstruction of Zingiberales from plastid genomes|journal=[[Biochemical Systematics and Ecology]]|date=June 2016|volume=66|pages=123–128|doi=10.1016/j.bse.2016.03.013|ref={{harvid|Deng et al|2016}}}}
* {{cite journal|last1=Kirchoff|first1=B. K.|last2=Lagomarsino|first2=L. P.|last3=Newman|first3=W. H.|last4=Bartlett|first4=M. E.|last5=Specht|first5=C. D.|title=Early floral development of ''Heliconia latispatha'' (Heliconiaceae), a key taxon for understanding the evolution of flower development in the Zingiberales|journal=[[American Journal of Botany]]|date=5 February 2009|volume=96|issue=3|pages=580–593|doi=10.3732/ajb.0800305|pmid=21628214|ref={{harvid|Kirchoff et al|2009}}|doi-access=free}}
* {{cite journal|last1=Kress|first1=W. John|title=The Phylogeny and Classification of the Zingiberales|journal=[[Annals of the Missouri Botanical Garden]]|date=1990|volume=77|issue=4|pages=698–721|doi=10.2307/2399669|jstor=2399669|url=https://www.biodiversitylibrary.org/part/7588}}
* {{cite journal|last1=Kress|first1=W. John|last2=Prince|first2=Linda M.|last3=Hahn|first3=William J.|last4=Zimmer|first4=Elizabeth A.|title=Unraveling the Evolutionary Radiation of the Families of the Zingiberales Using Morphological and Molecular Evidence|journal=[[Syst. Biol.]]|date=2001|volume=50|issue=6|pages=926–944|ref={{harvid|Kress et al|2001}}|doi=10.1080/106351501753462885|pmid=12116641|citeseerx=10.1.1.977.833|s2cid=24179401 }}
* {{cite journal|last1=Salvi |first1=Amanda M. |last2=Smith|first2=Selena Y.|last3=Benedict|first3=John C.|last4=Leong-Škorničková|first4=Jana|last5=Specht|first5=Chelsea | title=  Re-Examining Zingiberales Fossils Using Leaf Venation|url=http://2015.botanyconference.org/engine/search/index.php?func=detail&aid=769|journal=Botany |date=27 July 2015|issue=15004|format=Abstract of oral presentation|ref={{harvid|Salvi et al|2015}}}}
* {{cite journal|last1=Sass|first1=C|last2=Iles|first2=WJ|last3=Barrett|first3=CF|last4=Smith|first4=SY|last5=Specht|first5=CD|title=Revisiting the Zingiberales: using multiplexed exon capture to resolve ancient and recent phylogenetic splits in a charismatic plant lineage.|journal=[[PeerJ]]|date=21 January 2016|volume=4|pages=e1584|pmid=26819846|doi=10.7717/peerj.1584|ref={{harvid|Sass et al|2016}}|pmc=4727956|doi-access=free}}
* {{cite journal|last1=Smith|first1=James F.|last2=Kress|first2=W.John|last3=Zimmer|first3=Elizabeth A.|title=Phylogenetic analysis of the Zingiberales based on ''rbc''L sequences|url=http://scholarworks.boisestate.edu/cgi/viewcontent.cgi?article=1042&context=bio_facpubs|journal=[[Annals of the Missouri Botanical Garden]] |year=1993 |volume=80 |issue=3 |pages=620–630|ref={{harvid|Smith et al|1993}} |doi=10.2307/2399850|jstor=2399850}}
* {{cite journal|last1=Specht|first1=Chelsea D.|last2=Yockteng|first2=Roxana|last3=Almeida|first3=Ana Maria|last4=Kirchoff|first4=Bruce K.|last5=Kress|first5=W. John|title=Homoplasy, Pollination, and Emerging Complexity During the Evolution of Floral Development in the Tropical Gingers (Zingiberales)|journal=The Botanical Review|date=18 December 2012|volume=78|issue=4|pages=440–462|doi=10.1007/s12229-012-9111-6|s2cid=15781695|ref={{harvid|Specht et al|2012}}|url=http://libres.uncg.edu/ir/uncg/f/B_Kirchoff_Homoplasy_2012.pdf}}
* {{cite journal|last1=Tomlinson|first1=P. B.|author-link=P.B. Tomlinson|title=Phylogeny of the Scitamineae - Morphological and Anatomical Considerations|journal=[[Evolution (journal)|Evolution]]|date=1 January 1962|volume=16|issue=2|pages=192–213|doi=10.2307/2406197|jstor=2406197}}
* {{cite journal|last1=Triplett|first1=J. K.|last2=Kirchoff|first2=B. K.|title=Lamina architecture and anatomy in the Heliconiaceae and Musaceae (Zingiberales)|journal=[[Canadian Journal of Botany]]|date=1991|volume=69|issue=4|pages=887–900|url=http://gerontology.wp.uncg.edu/wp-content/uploads/sites/58/2014/09/12-Triplett1991.pdf|doi=10.1139/b91-115}}

=== APG ===

* {{cite journal |last=APG |author-link=Angiosperm Phylogeny Group|title=An ordinal classification for the families of flowering plants |journal=[[Annals of the Missouri Botanical Garden]] |year=1998 |volume=85 |issue=4 |pages=531–553 |jstor=2992015 |doi=10.2307/2992015|url=https://www.biodiversitylibrary.org/part/2234 }}
* {{cite journal |last=APG II |author-link=Angiosperm Phylogeny Group|title=An Update of the Angiosperm Phylogeny Group Classification for the orders and families of flowering plants: APG II |journal=[[Botanical Journal of the Linnean Society]] |year=2003 |volume=141 |issue=4 |pages=399–436 |doi=10.1046/j.1095-8339.2003.t01-1-00158.x |doi-access=free }}
* {{cite journal |last = APG III |author-link=Angiosperm Phylogeny Group|title=An Update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |year = 2009 |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 }}
* {{cite journal |last = APG IV |author-link=Angiosperm Phylogeny Group|title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |year=2016 |journal=[[Botanical Journal of the Linnean Society]] |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |doi-access=free }}

=== Websites ===

* {{cite web|title=Tropicos|url=http://www.tropicos.org/Home.aspx|publisher=[[Missouri Botanical Garden]]|access-date=30 December 2015|date=2015|ref={{harvid|Tropicos|2015}}}}
* {{cite web|title=Zingiberales|url=http://www.heliconiasocietypr.org/zingiberales.htm|publisher=Heliconia Society of Puerto Rico|access-date=22 February 2017|ref={{harvid|Heliconia Society|2017}}}}
* {{cite web|last1=Johansson|first1=Jan Thomas|title=The phylogeny of angiosperms|url=http://angio.bergianska.se/|access-date=23 February 2017|date=2013}}
* {{cite web|last1=Stephen|first1=Mark|title=Zingiberales|url=https://prezi.com/xu8gygckwlwk/zingiberales/|publisher=Prezi|access-date=23 February 2017|format=Graphic presentation|date=14 July 2015}}
* {{cite web |last=Stevens |first=P.F. |author-link=Peter F. Stevens|date=2017|orig-year= 2001 |title=Angiosperm Phylogeny Website |publisher=[[Missouri Botanical Garden]]|url=http://www.mobot.org/mobot/research/APWeb/orders/acoralesweb.htm#Monocots|access-date=31 January 2017}} (''see also'' [[Angiosperm Phylogeny Website]])
{{Refend}}

==External links==
{{Commons category|Zingiberales}}
{{Wikispecies}}
* [https://www.inaturalist.org/observations?place_id=any&subview=map&taxon_id=48318 Zingiberales observations on iNaturalist]

{{Monocotyledons}}
{{Zingiberales}}
{{Taxonbar|from=Q203779}}
{{Authority control}}

[[Category:Zingiberales| ]]
[[Category:Angiosperm orders]]
[[Category:Extant Campanian first appearances]]
[[Category:Pantropical flora]]