Invasion success of alien aquatic plants in their native and
Transcription
Invasion success of alien aquatic plants in their native and
Invasion success of alien aquatic plants in their native and introduced ranges. Comparison between their invasiveness in North America and in France. Thiébaut Gabrielle Université de Metz, UFR Sci FA, Laboratoire Biodiversité & Fonctionnement des Ecosystèmes, Avenue du Général Delestraint, 57070 METZ. Email : [email protected] 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 of dramatic biological invasions. The different steps (Heger, 2001): 1. Immigration: presence in the new area 2. Spontaneous establishment: independent growth and reproduction of at least one individual 3. Permanent establishment: Population growth to Minimum Viable Population 4. Colonisation of new localities: → The height of each step depends on the ability of the species to overcome environmental limitations. Generally, most plant species have expanded after a lag phase . Ludwigia hexapetala. • Introduced in 1836 in Montpellier (France), •widespread invasive since 30 years. L. hexapetala The basic rule (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 biology and the ecology of 27 spreading aquatic plants in their introduced ranges and in their 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. Material and Methods Nonindigenous plants 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). Invasive Aquatic Plants in France 11 invasive macrophytes in France (Muller et al., 2004): • 3 amphiphytes: amphiphytes Ludwigia peploides, L. hexapetala, Myriophyllum aquaticum • 8 hydrophytes plants: plants 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: Date of introduction in France and in North America Group I Group II Group III Azolla caroliniana Azolla filiculoides Elodea nuttallii Elodea canadensis Lemna minuta Lemna turionifera Acorus calamus Butomus umbellatus Callitriche stagnalis Glyceria maxima Hydrocharis morus-ranae Iris pseudacorus Marsilea quadrifolia Myriophyllum spicatum Myosotis scorpioides Nasturtium officinalle Najas minor Nymphoides peltata Phalaris arundinacea Potamogeton crispus Rorripa amphibia Trapa natans Veronica beccabunga Egeria densa* Ludwigia peploides* Ludwigia hexapetala* Myriophyllum aquaticum* N.America N N N N N N Q E E E E E E E E E E E Q E E E E E E E E First found in USA First found in Canada 1600 1905 (St, Lawrence R, QUE) 1861 (New York) 1975 ( Wisconsin), <1980 (Washington, DC); 1860 (Bantam Lake, CT) 1942 (Washington, DC) < 1826 1934 ( Hudson river) 1882 (Winchester, MA) 1859 (Wilmington, DE) 1831 <1879 (Middlessex Co,MA) 1876 (Hudson Co, NJ) 1893 (Long Island, NY); 1890 (Haddonfield, NJ) 1956 (Ontario) 1932 (Ontario) 1961 (colombie britannique) 1960 (Ontario) ? ( Rideau river) 1800 <1956 (Ontario) France E E E E E E N N N N N N N N N N N N N N N N N E E E E First found in France no data 1880 ( Deux-Sèvres) 1959 ( Alsace) 1845 1965 (Pyrénées-Atlantiques) 1992 ( Alsace, Lorraine) 1961 (Manche) 1820-1830 (Montpellier) 1820-1830 (Montpellier) 1880 ( Bordeaux) * native from South America Q: questionable origin: considered by some to be native, but probably nonindigenous and well naturalized in the US N = native E= nonindigenous The 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. Group I Group II Group III Azolla caroliniana Azolla filiculoides Elodea nuttallii Elodea canadensis Lemna minuta Lemna turionifera Acorus calamus Butomus umbellatus Callitriche stagnalis Glyceria maxima Hydrocharis morus-ranae Iris pseudacorus Marsilea quadrifolia Myriophyllum spicatum Myosotis scorpioides Nasturtium officinalle Najas minor Nymphoides peltata Phalaris arundinacea Potamogeton crispus Rorripa amphibia Trapa natans Veronica beccabunga Egeria densa* Ludwigia peploides* Ludwigia hexapetala* Myriophyllum aquaticum* Family Azollaceae Azollaceae Hydrocharitaceae Hydrocharitaceae Lemnacaeae Lemnacaeae Aracae Butomaceae Callitichaceae Poaceae Hydrocharitaceae Iridaceae Marsilaceae Haloragaceae Boraginaceae Brassicaceae Najadaceae Menyanthaceae Poaceae Potamogetonaceae Brassicaceae Trapaceae Scrophulariaceae Hydrocharitaceae Onagraceae Onagraceae Haloragaceae Growth habit Free-floating Free-floating submersed submersed Free-floating Free-floating emergent emergent submersed emergent Free-floating emergent emergent submersed emergent emergent submersed submersed emergent submersed emergent Free-floating emergent submersed submersed submersed submersed Lemna minuta P. crispus Tab.2: Main invasive plants in France and in N. America G. maxima Species traits 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. Ecological traits • • • • 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). Results Non native aquatic plants have classified in three groups: • i) restricted and invasive plants, • ii) potentially invasive and , been • iii) widespread and invasive macrophytes. Tab.3.Nonindigenous plants classification according to their invasiveness in N.America and in France Azolla caroliniana Azolla filiculoides Elodea nuttallii Group I Elodea canadensis Lemna minuta Lemna turionifera Acorus calamus Butomus umbellatus Callitriche stagnalis Glyceria maxima Hydrocharis morus-ranae Iris pseudacorus Marsilea quadrifolia Myriophyllum spicatum Group II Myosotis scorpioides Nasturtium officinalle Najas minor Nymphoides peltata Phalaris arundinacea Potamogeton crispus Rorripa amphibia Typha angustifolia Trapa natans Veronica beccabunga Egeria densa* Group III Ludwigia peploides* Ludwigia hexapetala* Myriophyllum aquaticum* Invasive restricted x potentially invasive widespread & invasive x x x x x x (Canada, USA) x (Eastern region USA) x (Eastern region USA) x x (Eastern region USA) x x (Connecticut) x (Connecticut) x (NE, Central USA)) x (Eastern region USA, Canada) x(Canada) x (Connecticut) x (Connecticut) x (Eastern region USA) x (Eastern region USA) x (Eastern region USA) x x x (Eastern region USA) x (Connecticut)) x x (Canada) x x (Canada) x x x x (Eastern region USA, France) x (Eastern region USA) x (France) x (Eastern region USA) x (France) x (Connecticut) x (France) Most nonindigenous aquatic species are widespread or potentially invasive in their introduced country, and only few alien species are known from a few occurrences. Some species are viewed as restricted in some areas (e.g. G. maxima in USA) and are considered as widespread elsewhere (in Canada). E. nuttallii in France (alien species) Invasive and widespread in North of France,Germany,England, Belgium. Rare in South of France. Elodea nuttallii in N.America (native) No instance in N. E. America where nuttallii or E. canadensis is weedy. E. nuttallii Pond Troy (NYS) Trapa natans in N.America (non indigenous area) T. natans in Cohoes Lake (NYS) Widespread & invasive in Eastern region of USA - - endangered in much of Europe (native) Widespread species in France and in N. America Native from South America Native from Europe Myriophyllum spicatum Relationships among species traits Traits modalities are mainly distributed along the F1 and F2 axes (17, 0 and 11.6% of 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. Distribution of modalities on the F1xF2 planes Size Leaf length3-5cm >250cm 50-70cm 150-250cm 20-50cm 70-150cm Leaf width >40mm 1-3cm 0.51 0.75 5-20cm <=20cm 20-40mm 0.62 4-12mm <=2m m 2-4mm <=1cm 12-20mm >80cm Flower width 0.22 >10mm 3-10mm 20-80cm Reproduction Dissemination fragmentation stolons 0.15 0.66 seeds 0.34 human dissemination winterbuds hydrochory 0.40 zoochory <=3mm other 0.12 anemochory rhizomes F1: biology axis F2: distribution axis 0.15 0.37 0.05 Growth form amphyphyte Water velocity floating leaves Biological Type 0.70 hydrophyte support tissue anchored submers ed helophyte Trophic level still 0.03 pH Light 1.3 -1.3 oligotrophe eutrophe low velocity free 0.05 0.66 0.02 running 0.57 0.05 alkaline acid mesotrophe 0.05 unshaded 0.05 neutral 0.04 0.03 1.4 -1.9 shade shade 50% 0.11 In bold: correlation ratios for each trait species indicated on the axes Main axes are not related to ecological traits but related to morphology. → Uniformity of the aquatic environment allows macrophyte to occupy very large ranges. Traits comparison between N.America and France 0.69 -0.96 0.79 -1.4 1 2 No difference between traits of species in France and in North America. 1: North America, 2: France Traits comparison between France and N. America The co-structure between species traits in France and in North America varies according to the species. • Helophytes (I. pseudacorus, V. beccabunga…) have different species traits in NA and France; • Free-floating species have relatively short vectors; • Aquatic species or amphiphytes widespread in the 2 areas (e.g.M. spicatum) have a high co-structure between traits in France and in N. America. Traits comparison between France and N. America Lpep Noff 0.86 -0.84 2 Maqu 1.1 -1.1 Tnat Ramp Msco 3 Eden Lgra Npel Vbec Pcri 3 groups : 1.Helophyte 2.Amphiphyte 3.Aquatic plants. plants Mspi Hmor Enut Csta Ecan Nmar 1 F1: 17% Nmin Ltur Lminu Lmino Afil Mqua Acar Ipseu Paru Bumb Acal F2: 12% Gmax M. aquaticum Conclusions • Invasive aquatic plants when introduced into nonindigenous habitats were characterized by: – a high plasticity – a broad distribution. • The architecture of invasive plants in non 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 all the following characteristics: 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. Ludwigia uruguyensis in Britain (France) Propagules Knowledge of strategies of reproduction, dispersion and competition among aquatic plants in their native area remains fragmentary . Physiological traits are unknown In addition, decline of widespread species (e.g. M. spicatum in USA and E. canadensis in France) was observed in some places. Myriophyllum spicatum E. canadensis → Difficult to predict macrophyte invasion. Further studies required for predicting macrophyte invasions. To explain the mechanisms of invasions, we need : 1. to improve our knowledge on the characteristics of invading species, and 2. on those of the ecosystems invaded 3. to investigate the relationships between these 2 factors, 4. or to study the invasion process in time.