Experiments on growth interactions between two invasive macrophyte species

Abstract. The success of invasive species has been attributed to the ability to displace other species by direct competition. We studied growth and possible competition between the two macrophyte species Elodea nuttallii and E. canadensis, because the former has been observed to replace the latter in the field. Additional experiments were conducted in aquaria with mixed plantings of Elodea species. Species growth was measured and competitive abilities of each species determined by applying the reciprocal yield model to mean plant weight and length. In monocultures the growth rates of the two species were similar, while in mixtures the growth rate of E. canadensis was significantly lower than that of E. nuttallii. E. canadensis was more sensitive to intraspecific than to interspecific neighbours, whereas E. nuttallii was indifferent to the presence of neighbours. Differential growth characteristics of Elodea species can explain the displacement of E. canadensis by E. nuttallii under eutrophic field conditions.     

Phenotypic plasticity as a clue for invasion success of the submerged aquatic plant Elodea nuttallii

• Two closely related alien submerged aquatic plants were introduced into Europe. The new invader (Elodea nuttallii) gradually displaced E. canadensis even at sites where the latter was well established. The aim of the study was to evaluate the combined effects of environmental factors on several phenotypic characteristics of the two Elodea species, and to relate these phenotypic characteristics to the invasion success of E. nuttallii over E. canadensis.
• In a factorial design, Elodea plants were grown in aquaria containing five different nitrogen concentrations and incubated at five different light intensities. We used six functional traits (apical shoot RGR), total shoot RGR, relative elongation, root length, lateral spread, branching degree) to measure the environmental response of the species. We calculated plasticity indices to express the phenotypic differences between species.
• Light and nitrogen jointly triggered the development of phenotypic characteristics that make E. nuttallii a more successful invader in eutrophic waters than E. canadensis. The stronger invader showed a wider range of phenotypic plasticity. The apical elongation was the main difference between the two species, with E. nuttallii being more than two times longer than E. canadensis. E. canadensis formed dense side shoots even under high shade and low nitrogen levels, whereas E. nuttallii required higher light and nitrogen levels.
• We found that under more eutrophic conditions, E. nuttallii reach the water surface sooner than E. canadensis and through intensive branching outcompetes all other plants including E. canadensis. Our findings support the theory that more successful invaders have wider phenotypic plasticity.

     

Can snail herbivory influence the outcome of competition between Elodea species?

Elodea nuttallii is more competitive than Elodea canadensis but is also more palatable to herbivores. We tested if grazing by generalist invertebrate herbivores could modify the competitive abilities of E. nuttallii, and thereby influence the outcome of the competition between the two Elodea species. The influence of snail herbivory on the competitive interactions between the two Elodea species cultivated together in indoor tanks was tested for 2 months. The presence of Lymnaea stagnalis in the tanks reduced significantly the final number of rootings, as well as the final biomass of E. nuttallii, but not the surface colonised by this species, whereas only the final number of rootings of E. canadensis was reduced. However, E. nuttallii remained the dominant species. We conclude that snail herbivory can influence the outcome of competition between Elodea species. However, snails are probably not an effective biological control agent of E. nuttallii.     

Comparative Analyses of Plastid Sequences between Native and Introduced Populations of Aquatic Weeds Elodea canadensis and E. nuttallii

Non-indigenous species (NIS) are species living outside their historic or native range. Invasive NIS often cause severe environmental impacts, and may have large economical and social consequences. Elodea (Hydrocharitaceae) is a New World genus with at least five submerged aquatic angiosperm species living in fresh water environments. Our aim was to survey the geographical distribution of cpDNA haplotypes within the native and introduced ranges of invasive aquatic weeds Elodea canadensis and E. nuttallii and to reconstruct the spreading histories of these invasive species. In order to reveal informative chloroplast (cp) genome regions for phylogeographic analyses, we compared the plastid sequences of native and introduced individuals of E. canadensis. In total, we found 235 variable sites (186 SNPs, 47 indels and two inversions) between the two plastid sequences consisting of 112,193 bp and developed primers flanking the most variable genomic areas. These 29 primer pairs were used to compare the level and pattern of intraspecific variation within E. canadensis to interspecific variation between E. canadensis and E. nuttallii. Nine potentially informative primer pairs were used to analyze the phylogeographic structure of both Elodea species, based on 70 E. canadensis and 25 E. nuttallii individuals covering native and introduced distributions. On the whole, the level of variation between the two Elodea species was 53% higher than that within E. canadensis. In our phylogeographic analysis, only a single haplotype was found in the introduced range in both species. These haplotypes H1 (E. canadensis) and A (E. nuttallii) were also widespread in the native range, covering the majority of native populations analyzed. Therefore, we were not able to identify either the geographic origin of the introduced populations or test the hypothesis of single versus multiple introductions. The divergence between E. canadensis haplotypes was surprisingly high, and future research may clarify mechanisms that structure native E. canadensis populations.     

Competition between Invasive and Indigenous Species: Impact of Spatial Pattern and Developmental Stage

Elodea canadensis (indigenous) and Elodea nuttallii (invasive) were grown in experimental tanks in monocultures and mixtures in two spatial patterns (aggregated or mixed) and two developmental stages (small or large plants of E. canadensis, and small plants of E. nuttallii). Competitive interactions between the two species were assessed by monitoring the area colonised by each species, the number of rootings and biomass after 10 weeks. In monocultures the growth of E. canadensis was significantly lower than that of E. nuttallii. In mixtures the number of rootings and biomass of E. canadensis were always significantly less than those of E. nuttallii. The tank surface area colonised by E. canadensis was always significantly less than that occupied by E. nuttallii, but it was higher in the aggregated treatment, where the colonisation of E. nuttallii was lower. Therefore both spatial pattern and developmental stage of an indigenous species (E. canadensis) may influence the outcome of competition with potential invaders (E. nuttallii).     

Changes in biomass and spatial distribution of <Emphasis Type="Italic">Elodea nuttallii</Emphasis> (Planch.) St. John,an invasive submerged plant,in oligomesotrophic Lake Kizaki from 1999 to 2002

Distribution of pure Elodea nuttallii vegetation was surveyed from 1999 to 2002, immediately after the most recent expansion of the species in Lake Kizaki, Japan. During 2001 and 2002, areas of E. nuttallii vegetation rapidly diminished and the summer plant height decreased wherever the vegetation remained. The organic matter content, total phosphorus, and extracted P of the sediment from the vegetation bed were measured. A linear relationship was observed between the extracted P in the sediment and the biomass. The extracted P significantly decreased in the shallow littoral vegetation bed, where the biomass clearly diminished. A fertilization experiment using the shallow littoral sediment collected in the vegetation bed was conducted in 2001. In this experiment, apical shoots of E. nuttallii were planted in pots with fertilized sediment (nitrogen, phosphorus, and potassium additions). The growth of E. nuttallii shoots was significantly enhanced by enrichment with phosphorus alone. The ecological implication of sediment phosphorus limitation is discussed in relation to the cause of decline in the E. nuttallii population in Lake Kizaki.     

Morphological and Chemical Changes Induced by Herbivory in Three Common Aquatic Macrophytes

The Dry Matter Content (DMC), the total phenolic content, the production of new branches and the plant fragmentation were compared in three macrophyte species (Elodea canadensis, Elodea nuttallii and Myriophyllum spicatum) exposed or not to snail herbivory. Grazing significantly reduced the DMC of M. spicatum and E. canadensis, but had no effect on the DMC of E. nuttallii. The phenolic contents of Elodea species were not modified by snail herbivory, whereas that of M. spicatum significantly increased when exposed to grazers. The number of new branches produced by M. spicatum and E. canadensis plants, and the fragmentation of E. canadensis also increased in response to herbivory. Chemical defences are therefore probably constitutive in Elodea and induced in M. spicatum, and morphological changes can be related to species growth form and synthesis of phenolic compounds. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Response of <Emphasis Type="Italic">Elodea Nuttallii</Emphasis> (Planch.) H. St. John to Manual Harvesting in the North-East of France

Elodea nuttallii (Planch). H. St John is an introduced aquatic macrophyte which was first observed in France in the early 1950s. The impact of two frequencies of harvesting on the biomass and regrowth strategy of this invasive species was evaluated by assessment of morphological traits monthly from February to October 2003. The effect of this management on the floristic biodiversity was also analysed. Harvesting caused a drastic reduction of biomass of E. nuttallii. Two harvests caused almost total disappearance of E. nuttallii. Furthermore, no significant difference was observed in the architecture of E. nuttallii between an unharvested site and harvested site. In one year, harvest did not allow the development of native aquatic plants.     

Redundancy and niche differentiation among the European invasive <Emphasis Type="Italic">Elodea</Emphasis> species

Community ecologists implicitly assume redundancy when they aggregate species into functional groups. But there have been remarkably few empirical efforts to investigate the accuracy of this concept in situ. The concept of redundancy could be roughly split into two components: the ecological redundancy (similar response to environmental variations involving similar ecological processes) and the functional redundancy (similar biological trait combinations shaping similar functional processes). Both types of redundancy are tested among the 3 invasive European Elodeas. In 11 sites and during two successive years 2004–2005, the cover growth rate of each Elodea species was monthly recorded. To test ecological redundancy, cover growth rates were related to a large suite of environmental variables. To test functional redundancy, 13 biological traits involved in competitive relationships were measured each month. Firstly, the redundancy hypothesis looks problematic for Elodea ernstiae. Indeed, the later possess numerous biological traits involved in light competition and niche overlap with the other Elodeas is very low. Secondly, ecological and functional redundancy can be successfully applied to Elodea canadensis and Elodea nuttallii. They share a large suite of biological traits leading to wide niche overlaps through the growing season. And the measured environmental variables do not differentially influence their growth rates, which are, in turn, controlled by a similar group of biological traits. In this way, the different invasiveness patterns of E. canadensis and E. nuttallii could be solely due to the ecological drift and their ecological dynamic could follow neutral rules.     

Arsenic species uptake and translocation in Elodea canadensis

Abstract

The capacity of Elodea canadensis to phytofiltrate arsenic species from water was evaluated. Plants were adapted to tap water and supplemented with 15 and 250?µg L^?1 of As. Inorganic arsenic species (As III, As V), and organic arsenic compounds: monomethylarsonate (MMA) and dimethylarsinate (DMA) were analyzed. Sampling was carried out at different times after exposure in culture water and plant organs. Plants exposed to 15?µg L^?1 of As concentration showed no significant difference on As concentration (95% confidence level) in their organs compared to controls. When plants were exposed to 250?µg L^?1 of As concentration, a significant increase of As concentration in plant organs was observed. After 1?h exposure, plants reduce 63.16% the As concentration in the culture water, with a bioaccumulation factor (BF) of 4.3. Under these conditions, E. canadensis accumulate As V in roots and do not translocate it to stems (transfer factor <1). MMA was determined in stems and leaves. E. canadensis effectively phytofiltrate As from tap water of a city located in an arsenic endemic area from concentrations of 36?µg L^?1 to undetectable levels (10?ng L^?1).     

Morphological and Morphometrical Variations of Selected Rotifer Species in Response to Predation: a Seasonal Study of Selected Brachionid Species from Lake Xochimilco (Mexico)

Allelopathy of filamentous green algae (FGA) has been less studied than that of macrophytes. Little Budworth Pool, Cheshire, UK is a small, shallow, clear-water lake with high TP concentrations, very high NO_3-N concentrations, only moderate phytoplankton density, high FGA growth (mainly Spirogyra sp.) and no submerged plants. Experiments were carried out to test the possible allelopathic effects of Spirogyra on the phytoplankton of this lake and on a submerged plant Elodea nuttallii. Changes in phytoplankton growth, phytoplankton species dynamics and species composition were apparently not influenced by allelopathy of live or decaying Spirogyra. A shift from diatom (Cyclotella sp) – cryptomonad (Chroomonas acuta and Cryptomonas erosa) dominance to Chlorococcales (Micractinium pusillum, Monoraphidium contortum and Scenedesmus opoliensis) – Volvocales (Chlorogonium elongatum and Pandorina morum) dominance was recorded in both control and FGA treatments, suggesting an effect of nutrient enrichment. Nutrient concentrations and differences in competitiveness among phytoplankton species can also explain differences in their growth rates in Spirogyra filtrate. Spirogyra also did not influence apex number per plant, shoot length or growth rate of E. nuttallii. This FGA species probably cannot control phytoplankton or E. nuttallii growth in nutrient rich conditions through allelopathy.     

Long‐term allelopathic control of phytoplankton by the submerged macrophyte Elodea nuttallii

相似文献   

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.     

Allelopathic activity of Elodea canadensis and Elodea nuttallii against epiphytes and phytoplankton

Elodea nuttallii and Elodea canadensis have both been introduced from North America to Europe. They are now common in many water bodies where they often form dominating stands. It was suggested that negative relationships between Elodea and phytoplankton or epiphytic covers exist, probably due to the release of growth inhibiting allelochemicals. This would be an effective strategy to avoid light limitation caused by algae and cyanobacteria. We investigated the allelopathic potential of both E. nuttallii and E. canadensis against different photoautotrophs, focussing on epiphytic algae and cyanobacteria isolated from different submersed macrophytes and culture strains. Methanolic extracts of both species inhibited the growth of most of these organisms. Only a culture strain of Scenedesmus brevispina was stimulated. Further separation of extracts yielded several active fractions, indicating that hydrophilic and slightly lipophilic compounds were responsible for growth reduction. At least some of the activity seems to be related to phenolic substances, but flavonoids in these species are inactive. Since growth declined also in a moderately lipophilic fraction of culture filtrate of E. nuttallii, we assume that active compounds were exuded in the water. Allelopathy might thus be relevant in situ and suppress cyanobacteria and algae. We furthermore found differences in the susceptibility of target organisms, which could (1) at least partly be a result of adaptation to the respective host plants and (2) indicate that allelopathic interference might reduce the abundance of some species, especially cyanobacteria, in epiphytic biofilms.     

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.     

Seasonal variability in the palatability of freshwater macrophytes: a case study

The pond snail Lymnaea stagnalis (L.) was used for a laboratory assessment of seasonal variation in palatability of three freshwater macrophytes: Potamogeton lucens, Elodea canadensis and E. nuttallii. For each species, 2–5 populations were investigated in spring and in summer. Preliminary results showed that the feeding rate of similarly-aged snails bred under standard conditions was stable over time. In contrast, snail feeding rate on the three macrophyte species decreased from spring to summer, which was therefore interpreted as a decrease in plant palatability. This decrease was probably due to tissue maturation, as suggested by the concomitant increase in the dry matter content of leaves of the three species. The high palatability of the species studied during the spring may prove detrimental in cases of strong herbivore pressure, and could have consequences for macrophyte distribution among aquatic habitats.     

Resistance to herbivory of two populations of <Emphasis Type="Italic">Elodea</Emphasis><Emphasis Type="Italic">canadensis</Emphasis> Michaux and <Emphasis Type="Italic">Elodea nuttallii</Emphasis> Planchon St. John

In this article, we compared the resistance of two introduced populations of Elodea nuttallii and Elodea canadensis to two different herbivores. Samples were collected from the River Rhine and River Rhône in eastern France. The two populations of E. nuttallii differed in their introduction history, whereas E. canadensis was introduced at the same time in the two sites. The Daily Food Consumption (DFC) rates of the two macrophyte populations were evaluated in no-choice experiments using the scraper Lymnaea stagnalis and the shredder Gammarus roeseli. At the same time, we assessed four plant traits: dry matter content (DMC), total nitrogen content, carbon/nitrogen ratio and total phenolic content. The two populations of E. canadensis were consumed at low levels by both the herbivores. L. stagnalis showed a higher DFC on the Rhône population of E. nuttallii than on the Rhine population. No significant difference between the two populations was established with G. roeseli, but the level of DFC was high. This result demonstrates that the assessment of plant palatability should be carried out with several generalist herbivores belonging to various feeding groups (e.g. scrapers or shredders). Although the Rhône population of E. nuttallii had higher levels of phenols than the other populations, it was consistently consumed in greater quantities than E. canadensis. Neither the phenolic contents were not effective against these herbivores, nor the levels of phenolics too low to induce an efficient resistance. The higher DMC and the lower DFC of the two populations of E. canadensis suggest that this introduced plant has co-evolved with indigenous enemies in the introduced range.     

The effects of floating mats of Azolla filiculoides Lam. and Lemna minuta Kunth on the growth of submerged macrophytes

In laboratory experiments, the growth characteristics of the submerged species Elodea nuttallii (Planch.) St. John and Potamogeton crispus L. were assessed in the presence and absence of floating mats of Azolla filiculoides Lam. and Lemna minuta Kunth. Light penetration and the development of pH and dissolved oxygen differences were monitored. The growth of P. crispus was suppressed much more than that of E. nuttallii and the effects of A. filiculoides were more severe than those of L. minuta. Findings are related to possible field responses of submerged plants under floating mats, especially their abilities to compensate for the potential suppressive effects of floating mats under natural conditions.     

Studies of Elodea nuttallii grown under photorespiratory conditions. III. Quantitative cytological characteristics

Abstract. Elodea nuttallii was grown in primary effluent from domestic wastewater under conditions in which CO₂ may be limiting. However, high photosynthetic inorganic carbon uptake rates have been reported for Elodea under these conditions. In order to determine if leaf cell structure showed modifications which would support proposed models of a plasma membrane (PM) bicarbonate transport system and observed high photo-synthetic rates, leaf cell ultrastructure was analysed using quantitative techniques (stereology). A Fold Index (FI) calculated for the PM showed that infolding increased surface area to 2.15 times that of an idealized cell of the same shape and size. Association Indices (Sa) showed a significant association of the mitochondria with the PM. These observations support models for an ATPase-driven HCO₃-cation cotransport system in Elodea cells. High chloroplast thylakoid surface density values (Sv) were similar to C₄:monocots and indicated high light-gathering potential. The granal/stromal (g/s), granal/cristae (g/c), and stromal/cristae (s/c) membrane ratios were similar to those of C₃ plant cells. Thus, Elodea chloroplasts and cells exhibited some structural features similar to both C₃ and C₄ plants. Membrane-bound inclusion bodies, which occupied 1% of the cell volume, were also observed in the leaf cells. Cell walls and nucleoid regions were absent in these bodies and X-ray (EDAX) analysis failed to detect any element (above the resolution limit of atomic no. 12) in substantial quantities. Inclusion bodies were observed in both the cytoplasm and the periplast and remain unidentified. A model is proposed for Elodea using an ATPase proton pump in the plasma membrane which extrudes protons into the periplast space between the cell wall and plasma membrane. This proton gradient is coupled to a bicarbonate symport.