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Ruppia
Ruppia rostellata, R. brachypus, + R. spiralis
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Monocots
Order:	Alismatales
Family:	Ruppiaceae Horan.
Genus:	Ruppia L.
Synonyms
Bucafer Adans. Buccaferrea P.Micheli ex Petagna Dzieduszyckia Rehmann

Ruppia, also known as the widgeonweeds, ditch grasses or widgeon grass, is the only extant genus in the family Ruppiaceae, with eight known species. These are aquatic plants widespread over much of the world. The genus name honours Heinrich Bernhard Rupp, a German botanist (1688–1719). They are widespread outside of frigid zones and the tropics.

Description

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The leaf is simple and not rhizomatous. They can be annual (commonly) or perennial (rarely); stem growth is conspicuously sympodial, but sometimes is not. These species are adapted to be in brackish water (and salt marshes). The leaves are small or medium-sized. Their disposition can be alternate, opposite, or whorled (usually alternate except when subtending an inflorescence). Even, lamina keep entire and are setaceous or linear. The leaf just shows one vein without cross-venules. Stomata are not present. The mesophyll leaks calcium oxalate crystals. The minor leaf veins do not present phloem transfer cells and leaks vessels.

These plants have stems without secondary thickening and xylem without vessels. The sieve-tube plastids are P-type. The root xylem does not present vessels.

These plants are hermaphroditic, with anemophilous or hydrophilous pollination. The flowers are ebracteate, small, and regular. Commonly, the flowers are aggregated in ‘inflorescences’, but sometimes they are solitary. Often, they grow in racemes, spikes, or umbels. The scapiflorous inflorescences are terminal, in short spikes, or subumbelliform racemes, sometimes one- or few-flowered. They do not have hypogynous disks. These flowers do not have perianth absent, except when small staminal appendages are regarded as perianth segments. The androecial members are all equal. The androecium just presents two fertile stamens with sessile anthers dehiscing by longitudinal slits. The pollen is polysiphonous and its grains are three-celled and nonaperturate. The gynoecium (2–)4(−16) is superior, carpelled, and euapocarpous. The carpel is not stylate, apically stigmatic with the stigma peltate, or umbonate. These flowers only present one ovule pendulous, nonarillate, campylotropous, bitegmic, and crassinucellate. The placentation is apical and embryo-sac development is of the polygonum type. Before fertilization, they fuse polar nuclei. The fruit is drupaceous and fleshy, forming an aggregate. The fruiting carpel is indehiscent, commonly on a long, spirally twisted peduncle, with each drupelet becoming very long-stalked. The fruit contains one nonendospermic seed with starch. The embryo can be straight or slightly curved. Membranous testa do not have phytomelan.

Taxonomy

The Cronquist system of 1981 placed the family in order Najadales of subclass Alismatidae in class Liliopsida in division Magnoliophyta .

The APG II system of 2003 (unchanged from the APG system of 1998) does recognize such a family and places it in the order Alismatales, in the clade monocots.

According to the AP-Website the family is doubtfully distinct from the family Cymodoceaceae: the plants in the three families Cymodoceaceae, Posidoniaceae, and Ruppiaceae form a monophyletic group.

A genus-level taxonomy was briefly revised by Zhao and Wu, including the following species in the world:

species

1. Ruppia bicarpa – Western Cape, South Africa
2. Ruppia cirrhosa – temperate regions: Europe, Asia, north + south (but not tropical) Africa, North America, West Indies, Argentina. The name is a homotypic synonym of R. maritima.
3. Ruppia didyma – Mexico, West Indies
4. Ruppia drepanensis – western + central Mediterranean
5. Ruppia filifolia – southern South America, Falkland Islands
6. Ruppia maritima – seashores and lakeshores around the world
7. Ruppia megacarpa – Australia, New Zealand, Asia (Korea, Japan, and Russia)
8. Ruppia occidentalis – Canada, USA
9. Ruppia polycarpa – Australia, New Zealand (including Chatham Islands)
10. Ruppia spiralis – seashores and lakeshores around the world
11. Ruppia tuberosa – Australia

Marine grasses families: Zosteraceae, Cymodoceaceae, Ruppiaceae and Posidoniaceae. Related families: Potamogetonaceae, Zannichelliaceae (not consistently).

Phylogeny and evolution

The first molecular phylogeny of the monogeneric family discerned three distinct species, R. tuberosa, R. megacarpa, and R. polycarpa, and one species complex comprising six lineages. The species complex, named R. maritima complex, was later updated as a group of eight lineages. These studies revealed that multiple hybridization and polyploidy events as well as chloroplast capture have occurred in the evolution of the genus.

Phytochemistry

These plants present an anatomy non-C4 type. Seven labdanes have been identified from this genus:

• ent-14,15-Dinor-8(17)-labden-13-one
• Methyl ester of (ent-12S)-15,16-Epoxy-12-hydroxy-12-oxo-8(17),13(16),14-labdatrien-19-oic acid.
• (-)-15,16-epoxy-8(17),13(16),14-labdatrien-19-ol.
• Methyl ester of (-)-15,16-epoxy-8(17),13(16),14-labdatrien-19-oic acid.
• (-)-15,16-Epoxy-8(17),13(16),14-labdatrien-19-al.
• (-)-15,16-Epoxy-8(17),13(16),14-labdatrien-19-yl acetate
• (ent-13E)-8(17),13-Labdadien-15-ol

Three steroids have been also isolated:

• (3β,5α,6β,7α,22E,24R)-Ergosta-8(14),22-diene-3,6,7-triol.
• (3β,5α,6β,7α,22E,24R)-Ergosta-8,22-diene-3,6,7-triol
• (24R)-Ergost-4-ene-3,6-dione.

References

1. Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
2. painting by the Swedish botanist C. A. M. Lindman (1856–1928), taken from his book(s) Bilder ur Nordens Flora (first edition published 1901–1905, supplemented edition 1917–1926?)
3. ^ Kew World Checklist of Selected Plant Families
4. NRCS. "Ruppia". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 26 October 2015.
5. Christenhusz, M. J. M. & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
6. Genaust, Helmut (1976). Etymologisches Wörterbuch der botanischen Pflanzennamen ISBN 3-7643-0755-2
7. Zhao L.-C., Wu Z.-Y. (2008) A review on the taxonomy and evolution of Ruppia. Journal of Systematics and Evolution 46: 467–478.
8. Ito, Y., Nr. Tanaka, T. Ohi-Toma, J. Murata, and A.M. Muasya (2015) Phylogeny of Ruppia (Ruppiaceae) revisited: Molecular and morphological evidence for a new species from Western Cape, South Africa. Systematic Botany 40(4): 942–949. doi: 10.1600/036364415X689988
9. ^ Ito, Y., T. Ohi-Toma, C. Nepi, A. Santangelo, A. Stinca, N. Tanaka, & J. Murata (2017) Towards a better understanding of the Ruppia maritima complex (Ruppiaceae): Notes on the correct application and typification of the names R. cirrhosa and R. spiralis Taxon 66: 167–171
10. ^ Ito, Y., T. Ohi-Toma, J. Murata & Nr. Tanaka (2010) Hybridization and polyploidy of an aquatic plant, Ruppia (Ruppiaceae), inferred from plastid and nuclear DNA phylogenies American Journal of Botany 97: 1156–1167
11. Ito, Y., T. Ohi-Toma, A. V. Skriptsova, M. Sasagawa, Nr. Tanaka, and J. Murata (2014) Ruppia megacarpa (Ruppiaceae): a new species to the floras of Japan, Korea, and Russia. Botanica Pacofica 3: 49–52
12. ^ Kubitzki (ed.) 1998. The families and genera of vascular plants, vol 4, Monocotyledons: Alismatanae and Commelinanae (except Gramineae). Springer-Verlag, Berlin.
13. Watson & Dallwitz. Zosteraceae. The families of flowering plants. http://delta-intkey.com/angio/www/zosterac.htm Archived 2017-07-09 at the Wayback Machine
14. Vascular Plant Families and Genera. List of genera in family CYMODOCEACEAE (accessed 2016-06-02) http://data.kew.org/cgi-bin/vpfg1992/genlist.pl?CYMODOCEACEAE
15. VASCULAR PLANT FAMILIES and GENERA. List of Genera in CYMODOCEACEAE (accessed 2016-06-02) http://www.mobot.org/mobot/research/apweb/orders/alismatalesweb.htm#Cymodoceaceael
16. Watson & Dallwitz. Potamogetonaceae. The families of flowering plants. http://delta-intkey.com/angio/www/potamoge.htm Archived 2021-04-17 at the Wayback Machine
17. Vascular Plant Families and Genera. List of genera in family POTAMOGETONACEAE (accessed 2016-06-02) http://data.kew.org/cgi-bin/vpfg1992/genlist.pl?POTAMOGETONACEAE
18. VASCULAR PLANT FAMILIES and GENERA. List of Genera in POTAMOGETONACEAE (accessed 2016-06-02) http://www.mobot.org/mobot/research/apweb/orders/alismatalesweb.htm#Potamogetonaceae
19. Ito, Y., T. Ohi-Toma, J. Murata & Nr. Tanaka (2013) Comprehensive phylogenetic analyses of the Ruppia maritima complex focusing on taxa from the Mediterranean Journal of Plant Research 126: 753–762

External links

• Ruppiaceae in the Flora of North America
• NCBI Taxonomy Browser
• links at CSDL, Texas

• Ruppia
• Brackish water plants
• Alismatales genera