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Invasion success of Invasion success of alien aquaticalien aquaticplants in their native and plants in their native and
introduced ranges. introduced ranges. ComparisonComparison between their between their
invasiveness ininvasiveness in North America and North America and in Francein France..
ThiThiéébautbaut GabrielleGabrielle
Université de Metz, UFR Sci FA, Laboratoire Biodiversité & Fonctionnement des Ecosystèmes, Avenue du Général Delestraint, 57070 METZ.
Email : thiebaut@sciences.univ-metz.fr
Acknowledgments:JN Beisel, P. Rocco and Karotsch Kevin for their helpful assistance .
Aquatic plants comprise few species worldwide,yet introductions of macrophytes represent some of the most examples ofdramatic biological invasions.
The different steps (The different steps (HegerHeger, 2001):, 2001):1.1. Immigration: Immigration: presence in the new area
2.2. Spontaneous establishmentSpontaneous establishment:: independent growth and reproduction of at least one individual
3.3. PPermanentermanent establishmentestablishment: : Population growth to Minimum Viable Population
4.4. Colonisation of new localities:Colonisation of new localities:
→ The height of each step depends on the ability of the species to overcome environmental limitations.
Generally, most Generally, most plant plant species have expanded species have expanded after a lag phaseafter a lag phase ..
L. L. hexapetalahexapetala
Ludwigia hexapetala. • Introduced in 1836 in Montpellier (France),•widespread invasive since 30 years.
The basic The basic rule rule (Williamson,1993) (Williamson,1993)
• 10% of imported plants escape to become introduced
• 10% of introduced invaders, become established,
• 10% of those established become pests.
Our objectives are to compare the distribution, the biologydistribution, the biology and the ecologyecologyof 27 spreading aquatic plants in their introduced rangesintroduced ranges and in their native native ranges.
We made the following two reciprocal comparisons:
• first European aquatic plants in France and in North America,
• and North American plants in France and in North America.
NonindigenousNonindigenous plants in North Americaplants in North America
The invasion of alien species has occurred since the mid-eighteenth century by the migration and settlement of European people on the North American continent.The documentation of introduction of each species was almost entirely based on information in the literature (e.g. Atlas North America).
IInvasinvasiveve AquaticAquatic Plants in FrancePlants in France11 invasive macrophytes in France (Muller et al.,
2004): •• 33 amphiphytesamphiphytes: Ludwigia peploides, L.
hexapetala, Myriophyllum aquaticum•• 88 hydrophyteshydrophytes plantsplants: A. caroliniana, A.
filiculoides, E. nuttallii, E. canadensis, L.minuta, L. turionifera.E. densa, L. major.9 widespread & invasive species (+100 sites /county) et 2 potentially invasives (between 11 et 100 sites/county).
Tab 1:Tab 1: Date of introduction Date of introduction in France and in in France and in NorthNorth AmericaAmerica
* native from South AmericaQ: questionable origin: considered by some to be native, but probably nonindigenous and wellnaturalized in the USN = nativeE= nonindigenous
N.America First found in USA First found in Canada France First found in FranceAzolla caroliniana N E no dataAzolla filiculoides N E 1880 ( Deux-Sèvres)Elodea nuttallii N E 1959 ( Alsace)
Group I Elodea canadensis N E 1845Lemna minuta N E 1965 (Pyrénées-Atlantiques)Lemna turionifera N E 1992 ( Alsace, Lorraine)Acorus calamus Q 1600 NButomus umbellatus E 1905 (St, Lawrence R, QUE) NCallitriche stagnalis E 1861 (New York) NGlyceria maxima E 1975 ( Wisconsin), 1956 (Ontario) NHydrocharis morus-ranae E 1932 (Ontario) NIris pseudacorus E <1980 (Washington, DC); 1961 (colombie britannique) NMarsilea quadrifolia E 1860 (Bantam Lake, CT) NMyriophyllum spicatum E 1942 (Washington, DC) 1960 (Ontario) N
Group II Myosotis scorpioides E NNasturtium officinalle E < 1826 NNajas minor E 1934 ( Hudson river) NNymphoides peltata E 1882 (Winchester, MA) ? ( Rideau river) NPhalaris arundinacea Q 1800 NPotamogeton crispus E 1859 (Wilmington, DE) <1956 (Ontario) NRorripa amphibia E 1831 NTrapa natans E <1879 (Middlessex Co,MA) NVeronica beccabunga E 1876 (Hudson Co, NJ) NEgeria densa* E 1893 (Long Island, NY); E 1961 (Manche)
Group III Ludwigia peploides* E E 1820-1830 (Montpellier)Ludwigia hexapetala* E E 1820-1830 (Montpellier)Myriophyllum aquaticum* E 1890 (Haddonfield, NJ) E 1880 ( Bordeaux)
The selected plantsThe selected plants
• 6 species coming from N. America and invasive in France (group I);
• 17 species coming from Europe and invasive in N. America (group II);
• 4 species coming from S. America and invasive in France and in N. America.
Lemna Lemna minutaminuta
P. crispus
G. maxima
Family Growth habitAzolla caroliniana Azollaceae Free-floatingAzolla filiculoides Azollaceae Free-floatingElodea nuttallii Hydrocharitaceae submersed
Group I Elodea canadensis Hydrocharitaceae submersedLemna minuta Lemnacaeae Free-floatingLemna turionifera Lemnacaeae Free-floatingAcorus calamus Aracae emergentButomus umbellatus Butomaceae emergentCallitriche stagnalis Callitichaceae submersedGlyceria maxima Poaceae emergentHydrocharis morus-ranae Hydrocharitaceae Free-floatingIris pseudacorus Iridaceae emergentMarsilea quadrifolia Marsilaceae emergentMyriophyllum spicatum Haloragaceae submersed
Group II Myosotis scorpioides Boraginaceae emergentNasturtium officinalle Brassicaceae emergentNajas minor Najadaceae submersedNymphoides peltata Menyanthaceae submersedPhalaris arundinacea Poaceae emergentPotamogeton crispus Potamogetonaceae submersedRorripa amphibia Brassicaceae emergentTrapa natans Trapaceae Free-floatingVeronica beccabunga Scrophulariaceae emergentEgeria densa* Hydrocharitaceae submersed
Group III Ludwigia peploides* Onagraceae submersedLudwigia hexapetala* Onagraceae submersedMyriophyllum aquaticum* Haloragaceae submersed
Tab.2: Main invasive plants in France and in N. America
SpeciesSpecies traitstraits
12 biological traits were selected.The documentation of species traits was
based on literature data.
Among them: • Size• Leaf size: length, width• Flower width• Reproduction: winter buds, seeds, rhizome,
stolons….• Dissemination: water, animals, wind, human
activities• Growth form: free, anchored and submersed,
anchored and support tissue, floating leaves• Biological type: aquatic, helophyte, amphiphyte.
EcologicalEcological traitstraits
• Water pH• Water velocity• Water trophic requirements• Light requirements.
The available information on species traits was structured using MCA on fuzzy table with a scale of 0 (no affinity of a species) to 1 (affinity of a species).
Non native aquatic plants have been classified in three groups:
• i) restricted restricted and invasive plants, • ii) potentiallypotentially invasive and ,• iii) widespreadwidespread and invasive macrophytes.
Tab.3.NTab.3.Nonindigenousonindigenous plants classification according to plants classification according to their invasiveness intheir invasiveness in N.N.America and in America and in FranceFrance
Invasive potentially widespread restricted invasive & invasive
Azolla caroliniana xAzolla filiculoides xElodea nuttallii x
Group I Elodea canadensis xLemna minuta xLemna turionifera xAcorus calamus x (NE, Central USA))Butomus umbellatus x (Connecticut) x (Eastern region USA, Canada)Callitriche stagnalis x (Canada, USA) x (Connecticut)Glyceria maxima x (Eastern region USA) x(Canada)Hydrocharis morus-ranae x (Eastern region USA)Iris pseudacorus xMarsilea quadrifolia x (Eastern region USA) x (Connecticut)Myriophyllum spicatum x
Group II Myosotis scorpioides x (Connecticut) x (Canada)Nasturtium officinalle x (Eastern region USA) xNajas minor x (Eastern region USA)Nymphoides peltata x x (Eastern region USA)Phalaris arundinacea xPotamogeton crispus xRorripa amphibia x (Eastern region USA) x (Canada)Typha angustifolia xTrapa natans x x (Eastern region USA)Veronica beccabunga xEgeria densa* x (Connecticut)) x (Eastern region USA, France)
Group III Ludwigia peploides* x (France)Ludwigia hexapetala* x (France)Myriophyllum aquaticum* x (Eastern region USA) x (Connecticut) x (France)
MostMost nonindigenous aquatic species are widespread or potentially invasivewidespread or potentially invasive in their introduced country, and only few alien species are known from a few occurrences.
SomeSome species are viewed as restrictedrestricted in some areas (e.g. G. maxima in USA) and are considered as widespread elsewhere (in Canada).
E. nuttalliiE. nuttallii in France (in France (alienalien speciesspecies))
Invasive Invasive andand widespreadwidespread in in NorthNorthof France,of France,GermanyGermany,,EnglandEngland, , BelgiumBelgium..
Rare in Rare in SouthSouth ofof France.France.
Elodea nuttalliiElodea nuttallii in in N.AmericaN.America (native)(native)
No instance in N. America where E.nuttallii or E. canadensis is weedy.
E. nuttallii
Pond Troy (NYS)
TrapaTrapa natansnatans in in N.AmericaN.America (non (non indigenousindigenous area)area)
- Widespread & invasive in Eastern region of USA
- endangered in much of Europe(native)
T. natans in Cohoes Lake (NYS)
WidespreadWidespread speciesspecies iin Francen France andand in N. in N. AmericaAmerica
Myriophyllum Myriophyllum spicatumspicatum
Native from South AmericaNative from Europe
Relationships among speciesRelationships among species traitstraits
Traits modalities are mainly distributed along Traits modalities are mainly distributed along the F1 and F2 axes (17, 0 and 11.6% of the F1 and F2 axes (17, 0 and 11.6% of total inertia)total inertia)– F1 is mainly related to potential size, leaf size
(length, width), biological type, growth form and reproduction
– F2 is related to leaf length, biological type and to a minor extend with light requirements.
<=20cm20-50cm
50-70cm
70-150cm
150-250cm
>250cmSize
<=1cm
1-3cm3-5cm
5-20cm
20-80cm
>80cm
Leaf length
<=2mm
2-4mm
4-12mm
12-20mm
20-40mm>40mmLeaf width
<=3mm
3-10mm>10mm
Flower widthfragmentation
rhizomes
stolons
winterbudsseeds
other
Reproduction
hydrochory
zoochory
anemochory
human dissemination
Dissemination
helophyte
hydrophyte
amphyphyte
Biological Type
freeanchored
floating leaves
support tissue
submersed
Growth form
stilllow velocity
running
Water velocity
oligotrophe
mesotropheeutrophe
Trophic level
acid
neutral
alkaline
pH
shade
shade 50%
unshaded
Light
-1.9
1.3-1.3 1.4
0.750.51
0.66
0.62
0.70
0.66
0.57
In bold: correlation ratios for each trait species indicated on the axes
Distribution of Distribution of modalitiesmodalities on on the F1xF2 the F1xF2 planesplanes
0.22 0.34
0.15
0.15
0.05 0.03
0.02
0.050.37
0.40 0.12
0.050.050.05
0.040.03
0.11
F1: biologyaxis
F2: distribution axis
Main axes are not related to ecological traits but related to morphology.
→ Uniformity of the aquatic environment Uniformity of the aquatic environment allows allows macrophytemacrophyte to occupy very large to occupy very large ranges.ranges.
1 2
-1.4
0.69-0.96 0.79
Traits Traits comparisoncomparison betweenbetween N.AmericaN.America andand FranceFrance
No difference between traits of species in France and in North America.
1: 1: NorthNorth AmericaAmerica, , 2: France2: France
Traits Traits comparisoncomparison betweenbetween France France andand N. N. AmericaAmerica
The coThe co--structure between species traits in structure between species traits in France and in North America varies France and in North America varies according according to theto the species.species.
•• HelophytesHelophytes (I. pseudacorus, V. beccabunga…) have different species traits in NA and France;
•• FreeFree--floatingfloating species have relatively short vectors;•• Aquatic species or Aquatic species or amphiphytesamphiphytes widespread in the 2
areas (e.g.M. spicatum) have a high co-structure between traits in France and in N. America.
Traits Traits comparisoncomparison betweenbetween France France andand N. N. AmericaAmerica
Acal
Acar
Afil
Bumb
Csta
Eden
EnutEcan
Gmax
Hmor
Ipseu
LminoLminu
Ltur
Lgra
Lpep
Mqua
Msco
Maqu
Mspi
Nmar Nmin
Noff
Npel
Paru
Pcri
Ramp
Tnat
Vbec
-1.1
0.86-0.84 1.1
3 groups :1.Helophyte2.Amphiphyte2.Amphiphyte3.Aquatic plants3.Aquatic plants.
33
1
2
MM. . aquaticumaquaticum
F1: 17%
F2: 12%
•• Invasive aquatic plants when introduced Invasive aquatic plants when introduced into into nonindigenousnonindigenous habitats were habitats were characterizedcharacterized by:by:–– a high plasticity– a broad distribution..
•• The architecture of invasive plants in non The architecture of invasive plants in non indigenous and in native area:indigenous and in native area:– Morphology of emergent species differed in
France and in North America.– Submersed and floating plants are very
plastic in the two areas. No difference.
The majority of invasive species share some or The majority of invasive species share some or all the following characteristicsall the following characteristics:
•short life cycle,
•rapid growth rate,
•high level of energy allocated to reproduction,
•efficient dispersal mechanisms,
•high population growth rate,
• and flexible use of environmental resources.
Knowledge of strategies of reproduction, dispersion and competition among aquatic plants in their native area remains fragmentary .
Physiological traits are unknown
In addition, dIn addition, declineecline of widespread species of widespread species (e.g. (e.g. MM. . spicatumspicatum in USA and in USA and E. E. canadensiscanadensis in France) in France) waswas observed in observed in some places. some places.
E. canadensisE. canadensisMyriophyllum spicatumMyriophyllum spicatum
→ Difficult to predict macrophyte invasion.
Further studies required for predictingFurther studies required for predictingmacrophytemacrophyte invasions.invasions.
To explain the mechanisms of invasions, we To explain the mechanisms of invasions, we need :need :
1. to improve our knowledge on the characteristics of invading species, and
2. on those of the ecosystems invaded3. to investigate the relationships between these 2
factors,4. or to study the invasion process in time.
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