|Ruppia rostellata, |
R. brachypus, + R. spiralis
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.
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.
A genus-level taxonomy was briefly revised by Zhao and Wu, including the following species in the world:
- Ruppia bicarpa - Western Cape, South Africa
- 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
- Ruppia didyma - Mexico, West Indies
- Ruppia drepanensis - western + central Mediterranean
- Ruppia filifolia - southern South America, Falkland Islands
- Ruppia maritima - seashores and lakeshores around the world
- Ruppia megacarpa - Australia, New Zealand, Asia (Korea, Japan, and Russia)
- Ruppia occidentalis - Canada, USA
- Ruppia polycarpa - Australia, New Zealand (incl Chatham Islands)
- Ruppia spiralis - seashores and lakeshores around the world
- Ruppia tuberosa - Australia
|Kubitzki (ed. 1998)||Watson & Dallwitz (delta-intkey)||data.kew||APWeb (mobot.org)|
|1. Zostera L.||Zostera||Zostera L.||Zostera L. (including Heterozostera den Hartog, Macrozostera Tomlinson & Posluzny, Nanozostera Tomlinson & Posluzny, Zosterella J. K. Small)|
|2. Heterozostera den Hartog||Heterozostera||Heterozostera (Setch.) Hartog||(in Zostera)|
|3. Phyllospadix Hook.||Phyllospadix||Phyllospadix Hook.||Phyllospadix J. D. Hooker|
|1. Syringodium Kütz||Syringodium||Syringodium Kutz.||(in Cymodocea)|
|2. Halodule Endl.||Halodule||Halodule Endl.||Halodule Endlicher|
|3. Cymodocea König||Cymodocea||Cymodocea K.Koenig (including Phycoschoenus (Asch.) Nakai )||Cymodocea König (including Amphibolis Agardh ?, Syringodium Kütz. ?, Thalassodendron den Hartog ?)|
|4. Amphibolis Agardh||Amphibolis||Amphibolis C.Agardh (including Pectinella J.M.Black)||(in Cymodocea)|
|5. Thalassodendron de Hartog||(name not found)||Thalassodendron Hartog||(in Cymodocea)|
|Ruppia L.||Ruppia||(in Ruppia L. in Potamogetonaceae)||Ruppia L.|
|Posidonia König||Posidonia||Posidonia K.Koenig||Posidonia König|
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.
These plants present an anatomy non-C4 type. Seven labdanes have been identified from this genus:
- Methyl ester of (ent-12S)-15,16-Epoxy-12-hydroxy-12-oxo-8(17),13(16),14-labdatrien-19-oic acid.
- 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-yl acetate
Three steroids have been also isolated:
- 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.
- 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?)
- Kew World Checklist of Selected Plant Families
- "Ruppia". Natural Resources Conservation Service PLANTS Database. USDA. Retrieved 26 October 2015.
- 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.
- Genaust, Helmut (1976). Etymologisches Wörterbuch der botanischen Pflanzennamen ISBN 3-7643-0755-2
- Zhao L.-C., Wu Z.-Y. (2008) A review on the taxonomy and evolution of Ruppia. Journal of Systematics and Evolution 46: 467–478.
- 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
- 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
- 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
- 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
- Kubitzki (ed.) 1998. The families and genera of vascular plants, vol 4, Monocotyledons: Alismatanae and Commelinanae (except Gramineae). Springer-Verlag, Berlin.
- Watson & Dallwitz. Zosteraceae. The families of flowering plants. http://delta-intkey.com/angio/www/zosterac.htm
- 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
- 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
- Watson & Dallwitz. Potamogetonaceae. The families of flowering plants. http://delta-intkey.com/angio/www/potamoge.htm
- 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
- 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
- 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
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