Abstract
Invasive species pose a major threat to native biodiversity and ecosystem integrity in many ecologically sensitive parts of the globe. Many research efforts have so far been made mainly with a focus on morphology, physiology and reproductive biology of invasive species to explain what determines the patterns of invasion. Recently, polyploidy has been reported to significantly influence plant invasiveness. Notwithstanding the profound management and conservation implications, determining the genetic basis of plant invasiveness is a challenging task for ecologists. Variation in ploidy levels of species promises to yield some useful insights in this direction and we, therefore, aimed to test the relation between polyploidy and species invasiveness. We documented chromosome number and ploidy level of 118 alien aquatic plant species including 50 alien non-invasive and 70 alien invasive species, belonging to 64 genera and 39 families distributed across various aquatic habitats of the Kashmir Himalaya at different stages of invasion. Results yielded 47 (39.83%) diploid species, 53 (44.91%) polyploids and the remaining 18 (15.25%) species were reported to have mixed ploidy, i.e., both diploids and polyploids. The invasive plant species exhibited about 40% of intraspecific ploidy polymorphism in contrast to about 28% found in non-invasive plant species. Alien invasive species (at stage IVa, IVb and V senso Colautti and MacIsaac 2004) were found to have more polyploidy (60%) than non-invasive species (at stage II and III) having only 45.23% polyploidy. Our results depict a clear pattern that non-invasive species are disproportionately more diploids with lower ploidy ratios, while invasive plant species on the contrary exhibit higher chromosome counts, thereby being predominantly polyploids. Invasion stage III with 17 diploid species represents by far the highest relative proportion of diploid species, while invasion stage V having 16 polyploid species is highest in terms of polyploid species. Regression analysis based on the stage of invasion and the ploidy status revealed that the relative proportion of polyploids on progressing stage of invasion increases significantly (p < 0.01). These results indicate that polyploidy may be one of the potential determinants of plant invasion. The implications of these results in timely prediction and better management of invasive species are discussed.
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Acknowledgements
We thank the Department of Botany; University of Kashmir, for providing research facilities. Financial support by the University Grants Commission (UGC), New Delhi, through its Indo–US project under 21st Century Knowledge Initiative to MAS is duly acknowledged.
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Appendix
Appendix
Familywise conspectus of aquatic alien plant species of Kashmir Himalayan fresh water ecosystems depicting nativity, growth form, invasion status, ploidy status and primary published source.
Families/species | Nt | Gf | Is | Chromosome status | Diploidy | Polyploidy | Primary published source |
|---|---|---|---|---|---|---|---|
Alismataceae | |||||||
Alisma gramineum Lej. | Eu | E | III | 2n = 14 | _ | Erlandsson (1946) | |
Alisma lanceolatum With | Eu | E | III | 2n = 26 | * | Bjorkqvist et al. (1969) | |
Alisma plantago-aquatica L. | Eu | E | V | 2n = 12, 10, 14 | _ | Walter (1946) | |
Sagittaria latifolia Willd. | Nam | E | III | 2n = 22 | _ | Walter (1946) | |
Sagittaria sagittifolia L. | Eu | E | V | 2n = 22, 16 | _ | Lohammar (1931) | |
Amaranthaceae | |||||||
Amaranthus lividus L. | As, Af, Sam | E | II | 2n = 34 | _ | Bao-Hua et al. (2002) | |
Alternanthera sessilis (L.) R. Br. ex DC. | Sam | E | IVb | 2n = 34, 96 | _ | * | Jansen (2004) |
Apiaceae | |||||||
Berula erecta (Huds.) Coville | Eu | E | III | 2n = 18 | _ | Hamal et al. (1986) | |
Araceae | |||||||
Acorus calamus L. | As | E | II | 2n = 24, 36, 48, 72 | _ | * | Ogra et al. (2009) |
Asteraceae | |||||||
Bidens cernua L. | Nam | E | IVa | 2n = 24, 48 | _ | * | Crowel and Parker (1981) |
Bidens tripartita L. | Eu | E | II | 2n = 24, 48, 72 | _ | * | Tanahara et al. (2010) |
Azollaceae | |||||||
Azolla cristata Kaulf. | Nam | FF | III | 2n = 44, 66, 88 | * | Stergianou and Fowler (1990) | |
Boraginaceae | |||||||
Myosotis caespitosa Schultz | Nam | E | IVb | 2n = 44, 48, 88 | _ | * | Stepankova (1993) |
Myosotis scorpioides L. | Eu | E | V | 2n = 22 | _ | Stepankova (1993) | |
Brassicaceae | |||||||
Barbarea intermedia Boreau | Eu | E | IVa | 2n = 16 | _ | Marian et al. (2008) | |
Barbarea vulgaris W. T. Aiton | Eu | E | IVa | 2n = 18 | _ | Marian et al. (2008) | |
Cardamine flexuosa With | Eu | E | IVb | 2n = 32 | * | Lihova et al. (2006) | |
Cardamine hirsuta L. | Eu | E | IVb | 2n = 16 | _ | Thurling (1968) | |
Nasturtium officinale W. T. | Eu | E | IVb | 2n = 32 | * | Walter et al. (2006) | |
Rorippa islandica (Oeder) Borbás | Eu | E | IVb | 2n = 16 | _ | Bleeker et al. (2007) | |
Butomus umbellatus L. | Eu | E | V | 2n = 26, 39 | _ | * | Anna and Vlasta (1993) |
Callitrichaceae | |||||||
Callitriche stagnalis Scop | Eu | E | IVb | 2n = 10 | _ | Alina et al. (2007) | |
Caryophylaceae | |||||||
Myosoton aquaticum (L.) Moench | Eu | E | III | 2n = 28 | _ | Fernandes and Leitao (1971) | |
Sagina saginoides (L.) H.Karst. | Eu | E | IVb | 2n = 22 | _ | Mardon et al. (2005) | |
Ceratophyllaceae | |||||||
Ceratophyllum demersum L. | Eu | S | V | 2n = 24, 38, 40, 48 | _ | * | Love et al. (1956) |
Cyperaceae | |||||||
Carex diluta Bieb. | Eu | E | V | 2n = 56 | * | Favarger et al. (1979) | |
Cladium mariscus (L.) Pohl | Eu | E | III | 2n = 18 | _ | Favarger et al. (1979) | |
Cyperus difformis L | Eu | E | V | 2n = 36 | * | Kumar and Subramaniam (1986) | |
Cyperus flabelliformis Rottb | Af | E | IVa | 2n = 30 | * | Bir et al. (1992) | |
Cyperus fuscus L. | Eu | E | IVb | 2n = 36, 48, 72 | _ | * | Subramanian (1988) |
Cyperus globosus L. | Af, Eu | E | V | 2n = 40 | * | Bir et al. (1992) | |
Cyperus glomeratus L. | Eu | E | V | 2n = 60 | * | Bir et al. (1992) | |
Cyperus iria L. | As, Af | E | IVb | n-56, 64 | _ | Bir et al. (1992) | |
Cyperus pumilus L. | Au | E | IVa | n-45 | _ | Bir et al. (1992) | |
Cyperus rotundus L. | Eu | E | V | 2n = 42–80, 108 | * | Bir et al. (1992) | |
Cyperus sanguinolentus L | NAm,SAm | E | III | n-40, 56 | _ | Bir et al. (1992) | |
Eleocharis atropurpurea (Retz.) | SAm | E | III | 2n = 20 | _ | Yano et al. (2004) | |
Eleocharis acicularis (L.) Roem. & Schult. | NAm,SAm | E | III | 2n = 20 | _ | Zedek et al. (2010) | |
Eleocharis palustris (L.) Roem. & Schult. | Eu | E | III | 2n = 16 | _ | Tanaka (1948) | |
Eleocharis parishii Britton | Nam | E | II | n-5 | _ | Tanaka (1948) | |
Eleocharis pauciflora Link | Nam | E | II | 2n = 76 | * | Roalson (2008) | |
Fimbristylis dichotoma (L.) Vahl. | Af, As | E | IVb | 2n = 10 | _ | Da Silva et al. (2008) | |
Scirpus juncoides Roxb | NAm | E | II | 2n = 74 | * | Yano and Hoshino (2005) | |
Scirpus martimus L. | Eu | E | II | 2n = 86, 104, 110 | _ | * | Hakansson (1928) |
Scirpus triqueter L. | Eu | E | IVa | 2n = 40, 42 | * | Da Silva et al. (2008) | |
Elatinaceae | |||||||
Elatine triandra Schkuhr | Eu | S | IVa | 2n = 40, 36 | * | Frisendahl et al. (1927) | |
Myriophyllum verticillatum L. | Eu | S | V | 2n = 28 | * | Taylor and Mulligan (1968) | |
Myriophyllum spicatum L. | Eu | S | V | 2n = 42 | * | Aiken et al. (1978) | |
Myriophyllum aquaticum (Vell.) Verdc. | SAM | S | V | 2n = 14 | _ | Love et al. (1961) | |
Hippuris vulgaris L. | Eu | E | IVa | 2n = 16, 30, 32 | _ | * | Marhold and Breitwieser (2010) |
Hydrocharitaceae | |||||||
Hydrilla verticillata (L.f.) Royle | Eu | S | V | 2n = 16 | _ | Pieterse et al. (1985) | |
Hydrocharis dubia (Blume) Backer | As | E | IVb | 2n = 16 | _ | Goldblatt and Johnson (1991) | |
Vallisneria spiralis L. | Eu | FF | III | 2n = 20, 30, 40 | _ | * | Goldblatt and Johnson (1991) |
Juncaceae | |||||||
Juncus articulatus L. | Eu | E | V | 2n = 80 | * | Harriman and Redmond (1976) | |
Juncus bufonius L. | Eu | E | IVb | 2n = 30, 34 | * | Tylor and Mulligon (1968) | |
Juncus effusus L. | Eu | E | IVb | 2n = 40, 42, 80 | * | Harriman et al. (1975) | |
Juncus inflexus L. | Eu | E | II | 2n = 40 | _ | Harriman et al. (1975) | |
Juncaginaceae | |||||||
Triglochin palustris L. | Eu | E | III | 2n = 24 | _ | Dawson (2008) | |
Labiatae (Lamiaceae) | |||||||
Mentha aquatica L. | Eu | E | III | 2n = 96 | * | Murray et al. (1972) | |
Mentha arvensis L. | Eu | E | III | 2n = 72 | * | Tucker and Fairbrothers (1981) | |
Mentha longifolia (L.) Huds | Eu | E | V | 2n = 24 | _ | Tucker and Chambers (2002) | |
Mentha piperita L. | Eu | E | IVb | 2n = 144 | * | Bhat Savithri et al. (2002) | |
Mentha spicata L. | NAm | E | II | 2n = 48 | * | Murray et al. (1972) | |
Lycopus europaeus L. | Eu | E | IVa | 2n = 22 | _ | Kondo and Motohashi (2009) | |
Lemnaceae | |||||||
Lemna gibba L. | Eu | FF | IVa | 2n = 40, 50, 60, 80 | _ | * | Daniel et al. (2005) |
Lemna minor L. | Eu | FF | V | 2n = 126 | * | Daniel et al. (2005) | |
Lemna turionifera Landolt | NAm | FF | II | 2n = 40, 42, 50, 80 | * | Urbanska-worytkiewicz (1980) | |
Lemna trisulca L. | Eu | FF | II | 2n = 20, 40, 60, 80 | _ | * | Urbanska-worytkiewicz (1975) |
Spirodela polyrhiza (L.) Schleid | Eu | FF | V | 2n = 80 | * | Urbanska-worytkiewicz (1975) | |
Wolffia arrhiza (L.) Horkel ex Wimm. | Eu | FF | III | 2n = 42 | _ | Urbanska-worytkiewicz (1975) | |
Utricularia aurea Lour. | As, Au | S | III | 2n = 80 | * | Subramanyam (1988) | |
Lentibulariacea | |||||||
Ammannia auriculata Willd. | As, Au | E | III | n-16 | _ | Shirley et al. (2011) | |
Ammannia baccifera L. | As | E | III | 2n = 24 | _ | Shirley et al. (2011) | |
Lythrum salicaria L. | Eu | E | III | 2n = 30 | _ | * | Urbanska-worytkiewicz (1975) |
Rotala densiflora (Willd.) Koehne | As, Au | E | IVb | 2n = 32 | _ | Urbanska-worytkiewicz (1975) | |
Marsiliaceae | |||||||
Marsilia quadrifolia L. | Eu | RF | V | 2n = 40 | _ | Marcon et al. (2003). | |
Menyanthaceae | |||||||
Menyanthes trifoliata L. | Eu | E | III | 2n = 54 | _ | * | Peruzzi et al. (2005) |
Nymphoides peltata (S.G.Gmel.) Kuntze | Eu | RF | V | 2n = 24, 54, 56 | _ | * | Shigenobu and Tanaka (1980) |
Nelumbonaceae | |||||||
Nelumbo nucifera Gaertn. | As, Eu | RF | V | 2n = 16 | _ | Huang (1992) | |
Nymphaeaceae | |||||||
Nymphaea alba L | Eu | RF | IVb | 2n = 48, 56, 64, 84 | * | Pellicer (2013) | |
Nymphaea lotus L. | Eu | RF | II | 2n = 28, 56, 84 | _ | * | Pellicer (2013) |
Nymphaea mexicana Zucc. | NAm | RF | III | 2n = 56 | * | Diao et al. (2006) | |
Nymphaea tetragona Georgi | Eu | RF | III | 2n = 28 | _ | Diao et al. (2006) | |
Nymphaea tuberosa Paine. | NAm | RF | III | 2n = 84 | * | Pellicer (2013) | |
Onagraceae | |||||||
Epilobium palustre L. | Eu | E | IVa | 2n = 36 | _ | Krasnikov et al. (1990) | |
Orchidaceae | |||||||
Spiranthes lancea (Thunb.) Baker | Eu | E | II | 2n = 72 | * | Lou (2004) | |
Poaceae | |||||||
Echinochloa colonum (L.) Link | As | E | III | 2n = 36, 48, 54 | * | Gould and Soderstrom (1967) | |
Echinochloa crus-galli (L.) P.Beauv. | Eu | E | IVa | 2n = 54 | * | Gould and Soderstrom (1967) | |
Paspalum paspalodes (Michx.) Scribn. | SAm | E | III | n-30 | * | Gould and Soderstrom (1967) | |
Phalaris arundinacea L. | Eu | E | IVb | 2n = 28, 42 | * | McWilliam & Neal-Smith (1962) | |
Phragmites australis (Cav.) Trin. ex Steud. | Eu | E | V | 2n = 96 | * | Connor et al. (1998) | |
Polygonaceae | |||||||
Polygonum hydropiper L. | Eu | RF | V | 2n = 20, 22 | _ | Love (1942) | |
Rumex chalepensis Mill | As | E | III | 2n = 70, 90 | * | Love and Love (1975) | |
Rumex conglomeratus Murray | Eu | E | II | 2n = 20 | _ | Love and Love (1975) | |
Rumex dentatus L. | Eu | E | IVb | 2n = 40 | * | Love (1967) | |
Potamogetonaceae | |||||||
Potamogeton crispus L | Eu | S | V | 2n = 50, 52, 56, 72, 78 | * | Jankun et al. (1989) | |
Potamogeton filiformis Pers. | Eu | RF | IVb | 2n = 66, 78 | * | Sharma and Chatterjee (1967) | |
Potamogeton fluitans Roth | Eu | RF | IVb | 2n = 52 | * | Yurtsev and Zhukova (1975) | |
Potamogeton natans L. | Eu | S | IVb | 2n = 42, 52 | .* | Kuleszanka (1934) | |
Potamogeton pectinatus L | Eu | RF | V | 2n = 80, 70–85, 87 | * | Kalkman and Van Wijk (1984) | |
Potamogeton perfoliatus L. | Eu | S | II | 2n = 26, 40, 52, 78 | * | Yurtsev et al. (1975) | |
Potamogeton pusillus L | Eu | S | III | 2n = 26 | _ | Arohonka et al. (1982) | |
P. berchtoldii Fieber | Eu | RF | Ivb | 2n = 26, 28 | * | Palmgren (1939) | |
P. nodosus Poir. | Eu | RF | V | 2n = 52 | * | Ottonello et al. (1985) | |
P. lucens L. | Eu | S | IVa | 2n = 52 | * | Hollingsworth et al. (1998) | |
P. trichoides Cham. & Schltdl. | Af | RF | II | 2n = 26 | - | Palmgren (1939) | |
P. indicus Roxb. | As | S | III | 2n = 42, 44, 46, 50, 56 | * | Misra (1972) | |
P. amblyphyllus | NAm | S | IVa | 2n = 52, 84 | * | Love and Love (1982) | |
P. wrightii | As | FF | IVa | 2n = 52 | * | Ganie (2010) | |
Primulaceae | |||||||
Primula sp. | NAm | RF | III | 2n = 44 | * | Maged (2006) | |
Ranunculaceae | |||||||
Caltha alba K. Jacq | Eu | E | II | 2n = 32 | _ | Kumar et al. (2012) | |
Rananculus aquatilis L. | Eu | RF | IVb | 2n = 16, 24, 32, 48 | _ | * | Love et al. (1982) |
Rananculus lingua L. | Eu | RF | III | 2n = 64, 128 | * | Love et al. (1982) | |
Salviniaceae | |||||||
Salvinia natans All. | Eu | FF | V | 2n = 18 | _ | Tryon and Tryon (1982) | |
Scrophulariaceae | |||||||
Veronica beccabunga L. | Eu | E | III | 2n = 18 | _ | Rychlewski and Jankun (1972) | |
Sparganiaceae | |||||||
Sparganium erectum Huds | Eu | E | V | 2n = 30 | _ | Love and Love (1942) | |
Typhaceae | |||||||
Typha angustata Bory & Chaub. | Eu, NAm | E | V | 2n = 60 | * | Heiser Charles (1948) | |
Zannichelliaceae | |||||||
Zannichellia palustris L. | Eu | S | III | 2n = 24 | – | Vierssen (1982) | |
As Asia, Au Australia, Af Africa, Eu Europe, Nam North America, Sam South America, E emergent, S submerged, RF rooted free floating, FF free floating, NT native, GF growth form, Is invasion status, * polyploidy, - diploidy
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Wani, G.A., Shah, M.A., Reshi, Z.A. et al. Polyploidy determines the stage of invasion: clues from Kashmir Himalayan aquatic flora. Acta Physiol Plant 40, 58 (2018). https://doi.org/10.1007/s11738-018-2629-4
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DOI: https://doi.org/10.1007/s11738-018-2629-4