Alismatales

Alismatales
Alisma plantago-aquatica
Scientific classification
Kingdom:	Plantae
Clade:	Angiosperms
Clade:	Monocots
Order:	Alismatales R.Br. ex Bercht. & J.Presl
Families
See Classification

Snake lily (Dracunculus vulgaris) of Araceae family in Crete, Greece.

Ottelia alismoides from family Hydrocharitaceae in Hyderabad, India.

The Alismatales (alismatids) are an order of flowering plants including about 4500 species. Plants assigned to this order are mostly tropical or aquatic. Some grow in fresh water, some in marine habitats.

Description

The Alismatales comprise herbaceous flowering plants of aquatic and marshy habitats, and the only monocots known to have green embryos other than the Amaryllidaceae. They also include the only marine angiosperms. The flowers are usually arranged in inflorescences, and the mature seeds lack endosperm.

Both marine and freshwater forms include those with staminate flowers that detach from the parent plant and float to the surface where they become pollinated. In others, pollination occurs underwater, where pollen may form elongated strands, increasing chance of success. Most aquatic species have a totally submerged juvenile phase, and flowers are either floating or emergent. Vegetation may be totally submersed, have floating leaves, or protrude from the water. Collectively, they are commonly known as "water plantain".

Taxonomy

The Alismatales contain about 165 genera in 13 families, with a cosmopolitan distribution. Phylogenetically, they are basal monocots, diverging early in evolution relative to the lilioid and commelinid monocot lineages. Together with the Acorales, the Alismatales are referred to informally as the alismatid monocots.

Early systems

The Cronquist system (1981) places the Alismatales in subclass Alismatidae, class Liliopsida [= monocotyledons] and includes only three families as shown:

• Alismataceae
• Butomaceae
• Limnocharitaceae

Cronquist's subclass Alismatidae conformed fairly closely to the order Alismatales as defined by APG, minus the Araceae.

The Dahlgren system places the Alismatales in the superorder Alismatanae in the subclass Liliidae [= monocotyledons] in the class Magnoliopsida [= angiosperms] with the following families included:

• Alismataceae
• Aponogetonaceae
• Butomaceae
• Hydrocharitaceae
• Limnocharitaceae

In Tahktajan's classification (1997), the order Alismatales contains only the Alismataceae and Limnocharitaceae, making it equivalent to the Alismataceae as revised in APG-III. Other families included in the Alismatates as currently defined are here distributed among 10 additional orders, all of which are assigned, with the following exception, to the Subclass Alismatidae. Araceae in Tahktajan 1997 is assigned to the Arales and placed in the Subclass Aridae; Tofieldiaceae to the Melanthiales and placed in the Liliidae.

Angiosperm Phylogeny Group

The Angiosperm Phylogeny Group system (APG) of 1998 and APG II (2003) assigned the Alismatales to the monocots, which may be thought of as an unranked clade containing the families listed below. The biggest departure from earlier systems (see below) is the inclusion of family Araceae. By its inclusion, the order has grown enormously in number of species. The family Araceae alone accounts for about a hundred genera, totaling over two thousand species. The rest of the families together contain only about five hundred species, many of which are in very small families.

The APG III system (2009) differs only in that the Limnocharitaceae are combined with the Alismataceae; it was also suggested that the genus Maundia (of the Juncaginaceae) could be separated into a monogeneric family, the Maundiaceae, but the authors noted that more study was necessary before the Maundiaceae could be recognized.

• order Alismatales sensu APG III

  family Alismataceae (including Limnocharitaceae)

  family Aponogetonaceae

  family Araceae

  family Butomaceae

  family Cymodoceaceae

  family Hydrocharitaceae

  family Juncaginaceae

  family Posidoniaceae

  family Potamogetonaceae

  family Ruppiaceae

  family Scheuchzeriaceae

  family Tofieldiaceae

  family Zosteraceae

In APG IV (2016), it was decided that evidence was sufficient to elevate Maundia to family level as the monogeneric Maundiaceae. The authors considered including a number of the smaller orders within the Juncaginaceae, but an online survey of botanists and other users found little support for this "lumping" approach. Consequently, the family structure for APG IV is:

• family Alismataceae (including Limnocharitaceae)

  family Aponogetonaceae

  family Araceae

  family Butomaceae

  family Cymodoceaceae

  family Hydrocharitaceae

  family Juncaginaceae

  family Maundiaceae

  family Posidoniaceae

  family Potamogetonaceae

  family Ruppiaceae

  family Scheuchzeriaceae

  family Tofieldiaceae

  family Zosteraceae

Phylogeny

Cladogram showing the orders of monocots (Lilianae sensu Chase & Reveal) based on molecular phylogenetic evidence:

Lilianae sensu Chase & Reveal  Acorales   Alismatales   Petrosaviales   Dioscoreales   Pandanales   Liliales   Asparagales  commelinids  Dasypogonaceae  Arecales  Poales  Zingiberales  Commelinales

Lilioid monocots Alismatid monocots

Further reading

• B. C. J. du Mortier 1829. Analyse des Familles de Plantes : avec l'indication des principaux genres qui s'y rattachent. Imprimerie de J. Casterman, Tournay
• W. S. Judd, C. S. Campbell, E. A. Kellogg, P. F. Stevens, M. J. Donoghue, 2002. Plant Systematics: A Phylogenetic Approach, 2nd edition. Sinauer Associates, Sunderland, Massachusetts ISBN 0-87893-403-0.
• Wilkin, Paul; Mayo, Simon J, eds. (2013). Early events in monocot evolution. Cambridge: Cambridge University Press. ISBN 978-1-107-01276-9. Retrieved 9 December 2015.
• Royal Botanic Gardens, Kew (2016), Monocots I: General Alismatids & Lilioids
• Chase, Mark W; Reveal, James L (2009), "A phylogenetic classification of the land plants to accompany APG III" (PDF), Botanical Journal of the Linnean Society, 161 (2): 122–127, doi:10.1111/j.1095-8339.2009.01002.x
• Les, Donald H; Tippery, Nicholas P. In time and with water ... the systematics of alismatid monocotyledons (PDF). pp. 118–164., in
• Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV" (PDF). Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. Retrieved 11 June 2016.
• Christenhusz, Maarten J.M.; Vorontsova, Maria S.; Fay, Michael F. & Chase, Mark W. (August 2015), "Results from an online survey of family delimitation in angiosperms and ferns: recommendations to the Angiosperm Phylogeny Group for thorny problems in plant classification", Botanical Journal of the Linnean Society, 178 (4): 501–528, doi:10.1111/boj.12285