Response of Macrophyte Traits to Herbivory and Neighboring Species: Integration of the Functional Trait Framework in the Context of Ecological Invasions

With the increase in the number of introduced species each year, biological invasions are considered as one of the most important environmental problems for native biodiversity. In invaded habitats, the establishment of exotic plant species depends on the abiotic and biotic environment. Herbivores and neighboring plants (native or exotic) comprise an important part of the latter. Herbivores cause trophic and non-trophic damage to focal plants, which respond to herbivory by varying their different traits quantitatively (e.g., growth rate and biomass changes) and qualitatively (e.g., variation in morphological and chemical defenses strategies affecting plant palatability). Neighboring plant species also affect functional traits and the fitness of focal plant species, thus herbivore effects on a focal plant could also depend indirectly on the palatability and defensive traits of the neighboring species inside the community. Here, in a first step toward the integration of associational susceptibility/resistance theories in the field of ecological invasion, we performed a microcosm experiment to consider the effects of an exotic crayfish on the growth rate, morphological traits and damage level of three macrophytes (two exotic, one native) growing in pairwise combinations. We found that (i) the response to herbivore presence and to neighboring species identity seemed to be species specific, and (ii) crayfish enhance the fragmentation rate of the two exotic macrophytes Ludwigia grandiflora and Egeria densa in the presence of the native macrophyte Myriophyllum spicatum, which could indirectly facilitate their invasion success. Indeed, fragmentation can increase dispersal abilities of the exotic macrophytes considered in this study as they are able to generate new plants from their fragments. However, our results showed that the interaction herbivore-neighbor species was hardly significant. Our paper presents some first results on associational resistance/susceptibility and lays the foundation for developing a general framework that combines plant community ecology and biological invasion ecology to explain invasive species success

Invasive earthworms modulate native plant trait expression and competition

Biological invasions have major impacts on a variety of ecosystems and threaten native biodiversity. Earthworms have been absent from northern parts of North America since the last ice age, but non-native earthworms were recently introduced there and are now being spread by human activities. While past work has shown that plant communities in earthworm-invaded areas change towards a lower diversity mainly dominated by grasses, the underlying mechanisms related to changes in the biotic interactions of the plants are not well understood. Here, we used a trait-based approach to study the effect of earthworms on interspecific plant competition and aboveground herbivory. We conducted a microcosm experiment in a growth chamber with a full-factorial design using three plant species native to northern North American deciduous forests, Poa palustris (grass), Symphyotrichum laeve (herb) and Vicia americana (legume), either growing in monoculture or in a mixture of three. These plant community treatments were crossed with earthworm (presence or absence) and herbivore (presence or absence) treatments. Eight out of the fourteen above- and belowground plant functional traits studied were significantly affected by earthworms, either by a general effect or in interaction with plant species identity, plant diversity level and/or herbivore presence. Earthworms increased the aboveground productivity and the number of inflorescences of the grass P. palustris. Further, earthworms and herbivores together affected root tissue density of P. palustris and the specific leaf area of V. americana. In this study, earthworm presence gave a competitive advantage to the grass species P. palustris by inducing changes in plant functional traits. Our results suggest that invasive earthworms can alter competitive and multitrophic interactions of plants, shedding light on some of the mechanisms behind invasive earthworm-induced plant community changes in northern North America forests.NE, LT and RS acknowledge funding from the German Research Foundation (DFG Ei 862/18-1). Authors also acknowledge the support of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (DFG–FZT 118, 202548816). Moreover, NE acknowledges funding by DFG (Ei 862/29-1 and Ei 862/31-1) as well as by the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant no. 677232)

Do Invasive Earthworms Affect the Functional Traits of Native Plants?

As ecosystem engineers, invasive earthworms are one of the main drivers of plant community changes in North American forests previously devoid of earthworms. One explanation for these community changes is the effects of earthworms on the reproduction, recruitment, and development of plant species. However, few studies have investigated functional trait responses of native plants to earthworm invasion to explain the mechanisms underlying community changes. In a mesocosm (Ecotron) experiment, we set up a plant community composed of two herb and two grass species commonly found in northern North American forests under two earthworm treatments (presence vs. absence). We measured earthworm effects on above- and belowground plant biomass and functional traits after 3 months of experiment. Our results showed that earthworm presence did not significantly affect plant community biomass and cover. Furthermore, only four out of the fifteen above- and belowground traits measured were affected by earthworm presence. While some traits, such as the production of ramets, the carbon and nitrogen content of leaves, responded similarly between and within functional groups in the presence or absence of earthworms, we observed opposite responses for other traits, such as height, specific leaf area, and root length within some functional groups in the presence of earthworms. Plant trait responses were thus species-specific, although the two grass species showed a more pronounced response to earthworm presence with changes in their leaf traits than herb species. Overall, earthworms affected some functional traits related to resource uptake abilities of plants and thus could change plant competition outcomes over time, which could be an explanation of plant community changes observed in invaded ecosystems

Invasive earthworms reduce chemical defense and increase herbivory and pathogen infection in native trees

Recent research shows that earthworms can alter defense traits of plants against herbivores and pathogens by affecting soil biochemistry. Yet, the effects of invasive earthworms on defense traits of native plants from previously earthworm-free ecosystems as well as the consequences for multitrophic interactions are virtually unknown. Here we use a combination of an observational study and a complementary experimental study to investigate the effects of invasive earthworms on leaf defense traits, herbivore damage and pathogen infection in two poplar tree species (Populus balsamifera and Populus tremuloides) native to North American boreal forests. Our observational study showed that earthworm invasion was associated with enhanced leaf herbivory (by leaf-chewing insects) in saplings of both tree species. However, we only detected significant shifts in the concentration of chemical defense compounds in response to earthworm invasion for P. balsamifera. Specifically, leaf phenolic concentrations, including salicinoids and catechin, were lower in P. balsamifera from earthworm-invaded sites. Our experimental study confirmed an earthworm-induced reduction in leaf defense levels in P. balsamifera for one of the defense compounds, tremulacin. The experimental study additionally showed that invasive earthworms reduced leaf dry matter content, potentially increasing leaf palatability, and enhanced susceptibility of trees to infection by a fungal pathogen, but not to aphid infestation, in the same tree species. Synthesis. Our results show that invasive earthworms can decrease the concentrations of some chemical defense compounds in P. balsamifera, which could make them susceptible to leaf-chewing insects. Such potential impacts of invasive earthworms are likely to have implications for tree survival and competition, native tree biodiversity and ecosystem functioning

Stratégies de réponse de l'espèce invasive <em></em>Ludwigia grandiflora<em></em> aux contraintes environnementales

Le succès des espèces invasives est influencé à la fois par les caractéristiques de l'espèce exotique et les caractéristiques de la communauté d'accueil. Il va donc dépendre des stratégies de réponses de ces espèces aux contraintes environnementales. Dans cette thèse, nous nous sommes intéressés à une espèce végétale : l'espèce Ludwigia grandiflora subsp. hexapetala, la Jussie à grandes fleurs. Cette espèce amphiphyte est considérée comme la plante « aquatique » la plus invasive en France. Cette thèse vise à approfondir nos connaissances sur les mécanismes d'adaptation, les stratégies de croissance et de colonisation de cette espèce face aux contraintes environnementales afin de mieux comprendre et expliquer son succès invasif. Les réponses de cette espèce face à des paramètres abiotiques et biotiques ont été étudiées via une approche par les traits fonctionnels. Pour cela, des approches expérimentales en conditions contrôlées et semi-contrôlées ont été réalisées. Nos résultats montrent que cette espèce possède une grande capacité d'adaptation aux fluctuations des paramètres abiotiques du milieu. Par ailleurs, L. grandiflora possède un taux croissance élevé, elle est capable de compenser la perte de biomasse due à la pression des herbivores et forme une dense canopée en réponse à la compétition. La stratégie de réponse de L.grandiflora aux pressions biotiques est à la fois dépendante de sa propre densité, de la densité et du statut de l'espèce voisine (indigène vs exotique), mais aussi du milieu (terrestre vs aquatique). Les stratégies de réponses de cette espèce face aux contraintes abiotiques et aux interactions biotiques pourraient expliquer son succès invasif

Biology and ecology of water primroses

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Biology and ecology of invasive water primrose: implications for ecosystem management

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Stratégies de réponses des espèces végétales aquatiques invasives

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Les élodées, un bon indicateur des contraintes environnementales?

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Réseau d’espèces invasives : effets directs et indirects sur les espèces natives

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