Plant Life In Water
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-RESEARCH-

[Systematics and Evolution of Aquatic Plants]

  • Alisma (Alismataceae) Ito et al. (in prep.)
  • Highlight(s)
    • Q: Do hybridization and polyploidy play important roles in the evolution of Alisma?
    • A: Yes.

  • Sagittaria (Alismataceae) Ito et al. (2020)
  • Highlight(s)
    • Sagittaria is a genus of ca. 40 species in the aquatic plant family Alismataceae with a nearly global distribution, and a center of diversity in the New World. Two thirds of the known species are native to the Americas, while only a few species are distributed in Africa, Asia and Europe. A previous biogeographic analysis of the genus suggested an African origin for the genus with subsequent dispersal to North America and then to East Asia. Here we expanded the taxon sampling with a focus on the New World taxa and applied species delimitation and biogeographic analyses to revise the knowledge of the phylogeny and evolution of the genus. We obtained largely similar topologies from the chloroplast DNA and nuclear DNA (ITS) data sets. The 74 accessions sampled for our analyses were delimited into 29 species and several cryptic taxa were revealed in widely distributed species. Biogeographic analysis supported basal diversification in South America and subsequent colonization to North America and Asia.

  • Monochoria (Pontederiaceae) Ito et al. (in prep.)
  • Elatine (Elatinaceae) Ito et al. (in prep.)
  • Highlight(s)
    • Q: Can dispersal be roled out in the biogeography of Elatine as Cai et al. (2016) concluded?
    • A: No. Dispersal plays an important role in the evolution of Elatine gratioloides from Australasia.

  • Ottelia (Hydrocharitaceae) Ito et al. (2019)
  • Eriocaulon (Eriocaulaceae) Larridon et al. (2019)
  • Callitriche (Plantaginaceae) Ito et al. (2017)
  • Najas (Hydrocharitaceae) Ito et al. (2017) | Ito et al. (2018)
  • Highlight(s)
    • Q: An infrasubgeneric classification of four sections in Najas subgen. Cauinia?
    • A: Section Euvaginatae was resolved as non-monophyletic; a classification of two sections is proposed.
    • A: A revised key to sections of Najas subgen. Caulinia is provided.
    • A: Najas subgenus Caulinia section Caulinia (Tzvelev) Yu Ito, stat. nov. - Caulinia sect. Caulinia Tzvel., Novosti Sist. Vyssh. Rast. 13: 18. (1976); Tzvel., Fl. Evropeiskoi Chasti SSSR 4: 201. (1979) - Najas subgen. Caulinia gunrankedh Euvaginatae Magn., Beitr. Kenntn. Najas 57. (1870) - Najas subgen. Caulinia section Nudae Rendl., Trans. Linn. Soc. London, Bot. 399 (1899), syn. nov. (Lectotype here designated: Najas graminea Del.) - Najas subgen. Caulinia section Spathaceae Rendl., Trans. Linn. Soc. London, Bot. 398 (1899), syn. nov. (Lectotype here designated: Najas indica (Willd.) Cham.)

  • Limosella (Scrophulariaceae) Ito et al. (2017)
  • Highlight(s)
    • Q: If/which dispersal play important roles in shaping the worldwide but discontinuous distribution of Limosella?
    • A: Relatively recent continental dispersal from southern Africa to the north into Eurasia via tropical Africa.
    • A: Historical trans-oceanic dispersal from Eurasia to Australia.
    • A: Current latitudinal trans-oceanic dispersal in the Southern Hemisphere.

  • Althenia (Potamogetonaceae) Ito et al. (2016)
  • Highlight(s)
    • Q: Afro-Eurasian Althenia seems to be closely related to Australasian Lepilaena?
    • A: Lepilaena is non-monophyletic and Althenia is nested within the group.
    • A: Synapomorphies and a revised key to species of Althenia s.l. are provided.
    • A: Althenia F.Petit,, Ann. Sci. Observ. 1: 451. (1829) = Lepilaena J. Drumm. ex Harv., Hooker's J. Bot. Kew Gard. Misc. 7: 57 (1855), syn. nov.
    • A: Althenia marina (E.L. Robertson) Yu Ito, comb. nov. ί Lepilaena marina E.L. Robertson, H. B. S. Womersley, Mar. Benth. Fl. South Australia 1: 80. (1984) = Lepilaena patentifolia E.L. Robertson, Fl. S.Austral. 4: 1736. (1986), syn. nov.
    • A: Its disjunct distribution most likely originated in Australasia (Miocene), followed by dispersal to Afro-Eurasia.

  • Ruppia (Ruppiaceae) Ito et al. (2010) | Ito et al. (2013) | Ito et al. (2015) | Ito et al. (2017)
  • Highlight(s)
    • Q: The cosmopolitan genus Ruppia with one species? two? many more?
    • A: Four species and one species complex comprising of nine entities.
    • A: Ruppia bicarpa Yu Ito et Muasya, sp. nov.---TYPE: SOUTH AFRICA. Western Cape: Cape Town Dist., Jakkalsfontein Private Nature Reserve, 33‹24'09"S, 18‹15'07"E, Brackish pool, 12 Nov 2014, Y. Ito & A.M. Muasya YI01979 (holotype: BOL!; isotype: TNS!)
    • Q: Polyploidy event(s) in Ruppia occur once? twise? many times?
    • A: Tetraploidy evolves at least twise ("Tetraploid_ƒΏ" and "Tetraploid_ƒΑ").
    • A: "Tetraploid_ƒΐ" is best explained by chloroplast capture between "Tetraploid_ƒΏ" and "Tetraploid_ƒΑ".
    • Q: Morphologically ambiguous collections to be explained by hybridization?
    • A: "Triploid" occurs between "Occidentalis" and "Tetraploid_ƒΏ".
    • A: Ruppia megacarpa and "Occidentalis" hybridized in Hokkaido, Japan.
    • Q: Anything else?
    • A: Disjunct distribution between Australasia and East Asia (Ruppia megacarpa and "Tetraploid_ƒΏ")
    • A: Siblings: "Utahensis" (inland salinity water) and "Occidentalis" (inland fresh alkaline water).
    • Q: Nomenclature notes:
    • A: Ruppia maritima L., Sp. Pl. 127. 1753 ? Lectotype (designated by Setchell in Proc. Calif. Acad. Sci. ser. 4, 25: 470. 1946): [illustration in] Micheli, Nov. Pl. Gen., pl. 72, tab. 35. 1729 - Buccaferra cirrhosa Petagna, Instit. Bot. 5: 1826. 1787 (syn. nov.) - Ruppia cirrhosa (Petagna) Grande in Bull. Orto Bot. Regia Univ. Napoli 5: 58. 1918 (syn. nov.) Lectotype (designated here): [illustration in] Micheli, Nov. Pl. Gen., pl. 72, tab. 35. 1729
    • A: Ruppia spiralis L. ex Dumort., Fl. Berg. 164. 1827 ? Lectotype (designated here): origin unknown (LINN 176.2 [digital image!])

  • Sparganium (Typhaceae) Ito et al. (2016)
  • Highlight(s)
    • Q: Varieties of Sparganium emersum seem not to be close each other?
    • A: Sparganium emersum s.l. revised to be non-monophyletic; S. acaule from North America resurrected.
    • Q: Morphologically ambiguous collections to be explained by hybridization?
    • A: Sparganium acaule ~ S. fluctuans hybridized in North America
    • A: Sparganium emersum ~ S. glomeratum hybridized in Europe
    • A: Sparganium fallax ~ S. japonicum hybridized in Japan
    • A: Sparganium glomeratum ~ S. gramineum hybridized in Japan
    • Q: Anything else?
    • A: Siblings: Sparganium androcladum (emergent) and S. fluctuans-S. gramineus (floating-leaved).
    • A: Siblings: Sparganium acaule (emergent) and S. glomeratum (amphibium).
    • A: Siblings: Sparganium emersum (amphibium) and S. angustifolium (floating-leaved).

  • Potamogeton (Potamogetonaceae) Ito et al. (2007) | Ito & Tanaka (2013) | Ito et al. (2014) | Ito et al. (2016)
  • Highlight(s)
    • Q: Morphologically ambiguous collections to be explained by hybridization?
    • A: Potamogeton ~inbaensis occurs between P. wrightii and P. lucens in China and Japan.
    • A: Potamogeton distinctus ~ P. nodosus hybridized in Myanmar.
    • A: Potamogeton ~malainoides occurs between P. wrightii and P. distinctus in Myanmar.
    • A: Potamogeton pusillus s.l. ~ P. illinoensis hybridized in Argentina.

  • Isolepis (Cyperaceae) Ito et al. (2016) | Yano et al. (2016)
  • Highlight(s)
    • Q: Varieties of Isolepis fluitans seem not to be close each other?
    • A: Isolepis fluitans s.l. revised to be non-monophyletic; I. lenticularis from Australasia resurrected.
    • Q: Morphologically ambiguous collections to be explained by hybridization?
    • A: Isolepis crassiuscula ~ I. lenticularis hybridized in New Zealand.
    • Q: Anything else?
    • A: Colonization into Australasia from South Africa occurred twise, followed by cladogenesis radiation
    • A: Isolepis crassiuscula distributed disjunctively between Australasia and East Asia
    • A: Isolepis fluitans distributed disjunctively between Europe and South Africa

  • Halodule (Cymodoceaceae) Ito & Tanaka (2011)
  • Highlight(s)
    • Q: Morphologically ambiguous collections to be explained by hybridization?
    • A: Halodule uninervis x I. pinifolia hybridized in Okinawa, Japan.


[Aquatic Plants Inventory]


  • East Asia: Japan (Ito et al. 2014b) PDF
  • New record(s)
    • Ruppia megacarpa.

  • East Asia: S. Korea (Ito et al. 2007) (Ito et al. 2014b) PDF
  • New record(s)
    • Ruppia megacarpa.
    Observed taxa
    • Bolboschoenus fluviatilis, Brasenia schreberi, Callitriche palustris, Lemna minor, Myriophyllum spicatum, Najas marina, Nymphaea tetragona, Nuphar japonicum, Potamogeton distinctus, Potamogeton fryeri, Potamogeton natans, Potamogeton octandrus, Potamogeton panormitanus, Potamogeton whightii, Phyllospadix japonicus, Ruppia maritima, Salicornia europaea, Schoenoplectus validus, Sparganium erectum, Spirodela polyrhiza, Stuckenia pectinata, Suaeda maritima, Trapa japonica, Triglochin maritimum, Utricularia australis, Zannichellia palustris, Zostera marina.

  • East Asia: Russian Far East (Ito & Skriptsova 2008) (Ito et al. 2014b) PDF
  • New record(s)
    • Ruppia megacarpa.
    Observed taxa
    • Alisma orientale, Cicuta virosa, Ceratophyllum demersum, Eleocharis mamillata, Equisetum fluviatile, Hydrilla verticillata, Monochoria korsakowii, Myriophyllum spicatum, Lamprothamnium sp., Lemna minor, Lemna trisula, Lythrum salicaria, Nymphaea tetragona, Penthorum chinense, Phyllospadix iwatensis, Potamogeton cristatus, Potamogeton macckianus, Potamogeton obtusifolius, Potamogeton oxyphyllus, Potamogeton perfoliatus, Potamogeton panormitanus, Ruppia maritima, Ruppia occidentalis, Sagittalia trifolia, Salicornia europaea, Schoenoplectus validus, Schoenoplectus mucronatus, Sparganium erectum, Sparganium glomeratum, Spirodela polyrhiza, Stuckenia pectinata, Suaeda malacosperma, Thacla natans, Trapa japonica, Trapella sinensis, Typha latifolia, Utricularia vulgaris, Zostera asiatica, Zostera japonica, Zostera marina.

  • Southeast Asia: Cambodia (Ito et al. (in prep.))

    New record(s)
    • Nechamandra alternifolia.
    Observed taxa

  • Southeast Asia: Myanmar (Ito et al. 2009; Ito & Barfod 2014; Ito et al. 2014a; Ito et al. 2014c)

    New record(s)
    • Blyxa aubertii var. echinosperma, Caldesia parnassifolia, Lemna trisulca, Nechamandra alternifolia, Najas tenuis, Potamogeton distinctus, Potamogeton distinctus ~ P. nodosus, Potamogeton maackianus, Potamogeton ~malainoides, Potamogeton octandrus.
    Observed taxa

  • Southeast Asia: Thailand (Ito 2013c; Ito 2014; Ito et al. 2014a; Ito 2016) PDF
  • New record(s)
    • Nechamandra alternifolia, Potamogeton octandrus, Najas marina.
    Observed taxa
    • Potamogeton distinctus, Ruppia maritima.

  • South Asia: India (Ito 2013b) PDF
  • Observed taxa
    • Pteridophytes: Marsilea minuta (Marsileaceae), Acrostichum aureum, Ceratopteris thalictroides (Pteridaceae), Salvinia natans (Salviniaceae); Angiosperms: Caldesia parnassifolia (Alismataceae), Aponogeton natans, A. undulatus (Aponogetonaceae), Cryptocoryne spiralis (Araceae), Callitriche stagnalis (Callitrichaceae), Drosera indica (Droseraceae), Eriocaulon sp. (Eriocaulaceae), Aeschynomene indica (Fabaceae), Myriophyllum indicum (Haloragaceae), Blyxa aubertii, Najas indica, Otteria alismoides (Hydrocharitaceae), Utricularia aurea, U. bifida, U. graminifolia, U. striatula (Lentibulariaceae), Nymphaea nouchalii (Nymphaeaceae), Zeylanidium lichenoides (Podostemaceae), Monochoria vaginalis (Pontederiaceae), Ruppia maritima (Ruppiaceae), Limophylla anomala, L. heterophila, L. indica (Scrophulariaceae), Trapa natans (Trapaceae).

  • Africa: Western Cape (Ito et al. 2015) PDF
  • New species
    • Ruppia bicarpa.
    Observed taxa
    • Althenia filiformis, Pseudalthenia aschersoniana.

  • Oceania: Australia and New Zealand (Ito in prep.) PDF

  • Europe: the Mediterranean (Ito 2012a) PDF
  • Observed taxa
    • Alisma plantago-aquatica, Althenia filiformis, Balderia ranunculoides, Callitriche brutia, Callitriche lusitanica, C. palustris, Cardamine lyrata, Damasonium alisma, Eleocharis mamillata , Eleocharis uniglumis, Isolepis fluitans, Juncus acutus, Juncus heterophyllus, Juncus tenuis, Lemna minor, Myriophyllum alternifolium, Posidonia australis, Potamogeton pectinatus, Ranunculus peltatus, Ranunculus trichophyllus, Ruppia cirrhosa, Ruppia drepanensis, Ruppia maritima, Sarcocornia sp., Triglochin bulbosum, Triglochin maritimum, Typha latifolia, Zannichellia major, Zannichellia obtusifolius, Zannichellia palustris.

  • Europe: Peri-Baltic sea (Ito 2012b) PDF
  • Observed taxa
    • Alisma lanceolatum, Alisma plantago-aquatica, Butomus umbellatus, Callitriche sp., Ceratophyllum demersum, Elatine alsinastrum, Elatine hungarica, Eleocharis mamillata, Hydrocharis morsus-ranae, Lemna minor, Lemna trisulca, Limosella aquatica, Lobelia dortmanna, Myriophyllum spicatum, Myriophyllum verticillatum, Najas marina, Nuphar lutea, Polygonum amphibium, Potamogeton coloratus, Potamogeton compressus, P. crispus, Potamogeton lucens, Potamogeton natans, Potamogeton perfoliatus, Ranunculus peltatus, Ruppia cirrhosa, Ruppia maritima, Sagittaria natans, Sagittaria sagitifolia, Sparganium erectum, Stratiotes aloides, Suaeda maritima, Trapa natans, Triglochin maritimum, Typha latifolia, Utricularia intermedia, Utricularia vulgaris, Zannichellia major, Zannichellia palustris, Zostera noltii.

  • North America: Canadian Great Plains (Ito & Cota-Sanchez 2014) PDF
  • New record(s)
    • Sparganium glomeratum.
    Observed taxa
    • Sparganium americanum, Sparganium fluctuans, Sparganium hyperboreum

  • North America: East and Central (Ito 2012c) PDF
  • Observed taxa
    • Callitriche verna, Chara sp. Eleocharis pauciflora, Elodea canadensis, Lemna trisulca, Limosella aquatica, Myriophyllum spicatum, Phyllospadix serrulatus, Potamogeton pectinatus, Potamogeton pusillus subsp. pusillus, Potamogeton pusillus subsp. tenuissimus, Ranunculus aquatilis var. diffuses, Ranunculus cymbalaria, Ruppia maritima, Ruppia occidentalis, Salicornia europaea, Spirodela polyrhiza, Suaeda maritima, Triglochin malitimum.

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