Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

Host range and searching behaviour of Cricotopus lebetis (Diptera: Chironomidae), a tip miner of Hydrilla verticillata (Hydrocharitaceae)

A chironomid midge, Cricotopus lebetis Sublette (Diptera: Chironomidae), was discovered feeding on Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) in Crystal River, Citrus, Co., Florida, in the 1990s. Larvae of the midge mine the apical meristems of hydrilla, causing terminal branching and stunting of the plant. We investigated the fundamental host range of the midge by conducting a series of no-choice and paired-choice tests. No-choice developmental tests with neonate larvae revealed that the fundamental host range of C. lebetis included not only on hydrilla but also several other aquatic plants in different families, suggesting that this insect is not a hydrilla specialist. In paired-choice bioassays, larval colonisation of Elodea canadensis Michx. (Hydrocharitaceae) and Najas guadalupensis (Spreng.) Magnus (Najadaceae) was greater than colonisation of H. verticillata. Behavioural bioassays in a Y-tube olfactometer and in Petri dishes suggested that neonate larvae were not able to locate host plant material, whereas older larvae were successful in finding hosts. In paired-choice oviposition tests, adult females discriminated between potential oviposition sites, with greater numbers of eggs laid on E. canadensis and N. guadalupensis than on H. verticillata. This study is the first detailed account of host searching and oviposition behaviour of a phytophagous chironomid midge. The results will be used to assess the potential value of C. lebetis as a biological control agent of hydrilla.     

Present distribution of the genus Elodea in the Alsatian Upper Rhine floodplain (France) with a special focus on the expansion of Elodea nuttallii St. John during recent decades

Three species of Elodea (Elodea canadensis Michaux, E. nuttallii St John and E. ernstiae St John) have colonized Europe from the American continent. All three arrived in the Alsatian Rhine floodplain (north-eastern France) soon after their arrival in Europe, i.e. in the mid-19th century for E. canadensis, and in the mid-20th century for E. nuttallii and E. ernstiae. The paper investigates the present distribution of Elodea spp. in the floodplain by quantifying the species’ respective occurrences and by describing their habitats. The study further focuses on E. nuttallii which is presently colonizing other parts of Europe. It analyses whether it has continued to expand in the Alsatian Rhine floodplain during recent decades, and it checks whether changes in the abundance of E. nuttallii have had an impact on species richness of water plant communities. E.␣nuttallii has been found to be at present one of the most dominant and most frequent aquatic plant species in the study sector, while E. canadensis and E. ernstiae are less abundant. The species’ distributions differ with regard to water chemistry and water temperature: E. canadensis occurs in oligo-mesotrophic, rather stenothermic habitats, whereas E. nuttallii and E. ernstiae can be encountered in meso- to eutrophic sites with little or no arrival of stenothermic ground water. By comparing successive vegetation relevés from the same sites the study revealed further that the distribution of E. nuttallii has been stable in recent decades, despite local fluctuations in abundance. No relationship could be established between those fluctuations and changes in species richness or type of local plant communities. The sum of the results suggests that the expansion of E. nuttallii in the Alsatian Rhine floodplain had been completed prior to the study period. The species’ present distribution in the study sector as well as its position in local plant communities might therefore be considered a model for what can be expected to happen in areas where E.␣nuttallii has only recently arrived.     

Chlorophyll fluorometry sheds light on the role of desiccation resistance for vegetative overland dispersal of aquatic plants

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Suppression of the aquatic weed Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) by a leaf-cutting moth Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) in Jozini Dam,South Africa

A classical biological control programme against Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) was initiated in South Africa following the discovery of extensive mats in Jozini Dam in northern KwaZulu-Natal (KZN) in 2006. However, in mid-2008, high densities of a phytophagous insect, Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) were discovered on the H. verticillata infestation, causing high levels of defoliation and dieback of the weed. In order to quantify the impact of the moth and to assess the long-term threat of H. verticillata to South African water bodies, a monitoring programme was initiated in 2013. Although P. diminutalis did not demonstrate a preference for H. verticillata over a native aquatic plant, Potamogeton schweinfurthii A. Bennett (Potamogetonaceae), moth populations followed a cyclical pattern of abundance, with rapid population increases coinciding with increases in the abundance of H. verticillata. High damage levels to H. verticillata were associated with high densities of immature P. diminutalis, which consistently led to population crashes of H. verticillata. Parapoynx diminutalis appears to have great potential to contribute to the management of H. verticillata in South Africa. However, the moth's oligophagous habits will probably preclude its intentional introduction into water bodies with H. verticillata where it does not already occur.     

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.     

Growth of Hydrilla verticillata (L.f.) Royle under controlled conditions

Aquatic macrophytes are important resources for the maintenance of trophic chains and in biogeochemical processes, but they can also be deleterious for several uses if present in excess. Hydrilla verticillata was found in the Paraná River (Brazil) after 2005, which requires monitoring owing to the invasive potential of this species. In this study, we measured the growth of H. verticillata under controlled conditions and compared the growth dynamics for the two development strategies (branch and tuber). We show that this species has great potential to develop in tropical (Brazilian) aquatic ecosystems. The parameters from the modelling of the growth kinetics indicated a doubling time of 19.8 days for H. verticillata growing from stems; however, the growth from tubers were much faster, with doubling times ranging from 2.5 to 11 days. The delay for the tubers to sprout caused a decrease in the number of branches of H. verticillata stems. From the growth parameters obtained from the experiments under controlled conditions, we concluded that the high temperature and light availability in most South American reservoirs (including the Porto Primavera Reservoir where it was first recorded) are suitable for H. verticillata to compete and probably displace other native aquatic macrophytes, such as Egeria najas, Egeria densa, and Cerathophyllum demersum. This is a matter of concern because these and other submersed species are commonly found in several natural and man-made South-American aquatic ecosystems, where they are key for biodiversity maintenance.     

凤眼莲入侵程度对金鱼藻和黑藻生长及种间关系的影响

湿地生态系统中凤眼莲（Eichhornia crassipes）入侵造成湿地植物群落结构退化及功能崩溃,直接影响沉水植物的生长繁殖及初级生产力。目前关于凤眼莲的入侵机制有一定的研究,而关于凤眼莲入侵程度对沉水植物金鱼藻（Ceratophyllum demersum）和黑藻（Hydrilla verticillate）生长及种间关系的影响相对缺乏。以外来入侵植物凤眼莲,沉水植物金鱼藻和黑藻为研究对象,设计凤眼莲入侵程度（无入侵,轻度入侵对应盖度25%,重度入侵对应盖度75%）交叉定植方式（黑藻单种模式、金鱼藻单种模式,金鱼藻和黑藻混种模式）的控制实验,探究凤眼莲入侵强度对沉水植物金鱼藻和黑藻生长及种间关系的影响。结果表明,凤眼莲入侵程度显著降低了金鱼藻的生物量、分枝数;黑藻的株高、分枝数和分节数。无凤眼莲入侵时,两种沉水植物生物量均最大,两者种间竞争关系较强;随凤眼莲入侵盖度增加,两种沉水植物的生物量先急剧降低后略微增加,种间关系经过微弱促进后又变为竞争作用,其中黑藻表现出明显的竞争优势。此外,凤眼莲入侵略微降低了水体中的总氮、总磷含量。结构方程模型分析结果表明凤眼莲入侵以及水体总氮、总磷等水体理化性质对沉水植物生长均有显著负向影响（P<0.05）,且水体理化性质对沉水植物生长的影响强于凤眼莲入侵。总之,凤眼莲入侵显著降低了金鱼藻和黑藻生长繁殖,随着凤眼莲入侵程度增加,两种沉水植物种间关系由竞争转变为促进再转变为竞争。研究结果为凤眼莲入侵有效控制及湿地沉水植被的恢复与重建提供了一定的理论依据和技术支撑。     

Complete development on Elodea canadensis (Hydrocharitaceae) eliminates Hydrellia egeriae (Diptera,Ephydridae) as a candidate biological control agent of Egeria densa (Hydrocharitaceae) in the U.S.A.

The leaf-mining fly Hydrellia egeriae Rodrigues-Junior (Diptera: Ephydridae) was evaluated under quarantine conditions as a candidate biological control agent of the invasive aquatic weed Egeria densa Planchon (Hydrocharitaceae) in California, U.S.A. The objective of this study was to test the fly’s feeding preference for E. densa as compared to the North American native Elodea canadensis Michaux (Hydrocharitaceae), an important benchmark species. No choice testing revealed that H. egeriae can complete development on E. canadensis and choice testing indicated the fly presents little oviposition discrimination between E. densa and E. canadensis. It is concluded that the host range of H. egeriae is too broad for use as a biological control agent of E. densa in the U.S.A.     

Seasonal and long-term changes in surface cover of aquatic plants in a shallow pond,Ojaga-ike,Chiba, Japan

Seasonal changes in surface cover and long-term floral changes in aquatic plants of a shallow eutrophic pond, Ojaga-ike, as well as the phenology of each species, were studied. Out of ten floating-leaved and submerged plants, the following five species showed a change in surface cover during 1978: Potamogeton crispus and Elodea nuttallii occupied 53.5% of the total pond surface in May, Trapa natans and Nelumbo nucifera 79.1% in July, N. nucifera and Hydrilla verticillata 62.0% in September and N. nucifera 41.2% in November. In addition to dramatic seasonal changes in surface cover, long-term floral changes occurred in this pond from 1971 onwards. While Nymphoides indica, Myriophyllum spicatum and H. verticillata decreased in number and in distribution ranges between 1971 and 1978, two newly established plants, N. nucifera and E. nuttallii, have now become dominant. The alteration in dominance may be due to water level fluctuations and interspecific competition for light. Seasonal changes in water quality during the study period are also summarized.     

Habitat characteristics of an alien species <Emphasis Type="Italic">Elodea canadensis</Emphasis> in Slovenian watercourses

The distribution, abundance and habitat characteristics of an alien species, Elodea canadensis, were surveyed in watercourses in Slovenia. The accompanying plant community was also examined. Distribution and abundance of macrophytes were assessed in reaches of different length, and habitat assessment is based on 12 parameters of the Riparian, Channel, and Environmental (RCE) Inventory. E. canadensis thrived in 132 out of 1,227 reaches examined and in 12 of the 39 watercourses surveyed. It grew in the company of a variety of species, most frequently with different species of Potamogeton. It was rarely found as the prevailing, and never as the only species in any reach. Canonical correspondence analysis (CCA) of reaches with E. canadensis revealed that the presence of detritus, retention structures and properties of the riparian zone explained most of the variance in community composition. Habitat analysis of well-established stands of E. canadensis showed that its preferred habitats were watercourses flowing through agricultural landscape, with a narrow, more or less disturbed riparian zone, with moderate presence of retention structures, and with sediment that was a mixture of gravel, sand and silt with either coarse or fine organic matter. It was not found in the parts of streams with the most dynamic flow, and was absent from watercourses in the karst region, due to the frequent and extreme water level fluctuations. The alien species E. canadensis did not express its invasive character in heterogeneous watercourses with rich macrophyte communities.     

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.

     

The effects of nitrate loading on the invasive macrophyte Hydrilla verticillata and two common, native macrophytes in Florida

Rising nitrate concentrations in the water column and the spread of invasive, non-native macrophytes are two major threats to Florida's oligotrophic, freshwater ecosystems. We used a replicated mesocosm experiment to test the effects of elevated nitrate concentrations in the water on the growth of the invasive macrophyte Hydrilla verticillata and two common, native submerged macrophytes Vallisneria americana and Sagittaria kurziana. Results from this study indicate that nitrate concentrations of 1.0 mg L⁻¹ NO₃-N in the water increased the final dry-weight biomass of H. verticillata by 2.75 times, while having no statistical effect on the growth of the two native species. Additionally, H. verticillata grew at a faster rate than the two native species in the low nitrate treatments accounting for 82% of the total biomass, indicating that it may have the capacity to invade relatively pristine communities. In waters where nitrate concentrations continue to rise, the cost of control efforts for H. verticillata may substantially increase in the future.     

Community collocation of four submerged macrophytes on two kinds of sediments in Lake Taihu, China

To explore a method for rapid restoration and artificial regulation of submerged macrophytes in large-scale restoration of eutrophic lakes, the succession and the biodiversity changes of four communities composed of four native, common submerged macrophytes, Hydrilla verticillata, Potamogeton malaianus, Vallisneria spiralis and Najas marina, on two kinds of sediments were investigated. Under low light intensity (reduced by 99%), the plant biomass changed with seasonal changes, plant competition, and environmental stress. The competitive capability for light differed in the four species due to different shoot height and tiller number. After 405 days of transplantation, H. verticillata became dominant in all communities. The biomass of H. verticillata, with strong ability to endure low water light environment, accounted for more than 90% of the total community biomass, and P. malaianus had only weak growth, while V. spiralis and N. marina almost disappeared. Based on livability and biomass of submerged macrophytes on two sediment types, brown clay sediment appeared to be more favorable for the settlement of the plants, while fertile sludge sediment was suitable for vegetative growth. In conclusion, the improvement of habitats and the selection of appropriate plant species are of the greatest importance for ecological restoration of the aquatic ecosystem.     

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)     

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).     

Effects of Desiccation and Sediment Type on Early Regeneration of Plant Fragments of Three Species of Aquatic Macrophytes

We tested the effects of desiccation and type of sediment (clay or sand) on the regeneration of fragments of the apical shoots of three species of Hydrocharitaceae: Egeria najas, E. densa and Hydrilla verticillata. The shoots were left to dry on clay or sand from zero to four days, and were then returned to aquaria containing water. To approximate natural conditions, the aquaria containing clay had higher turbidity and nutrient (P and N) concentrations than did the aquaria with sand. All species dried faster on sand substrate, which led to lower regeneration (in terms of dry weight, length, sprout and root formation) in this treatment. H. verticillata fragments elongated faster than the other species, but E. najas was the most successful species (in terms of increase in dry weight) in the sand treatment. Our results indicate that the exotic H. verticillata has a competitive advantage, at least in its early stages of regeneration, over the other two, native species, especially in more eutrophic and turbid habitats; whereas E. najas fragments have a competitive advantage in less‐turbid, oligotrophic and sand‐dominated sites. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Faunal influences on New Zealand seed dispersal characteristics

A range of distinctive dispersal features have been recognised within the New Zealand flora, and a wide range of fauna are involved in the dispersal of seed in New Zealand, either by consuming fruit or seed, or as transporters of adhesive seed. In this study the composition of New Zealand’s zoochorous fauna (except insects) was examined using both trait matching within environmental, morphological and behavioural variables, and compared to the trait pattern of the groups of plant species they disperse. The importance of the different dispersal groups to the plant species they disperse varies with habitat, landform, region of New Zealand, foraging behaviour, and morphology. Over half of New Zealand’s vertebrate fauna are involved in fruit dispersal, though only 6% are considered frugivorous—the remainder include varying quantities of insects and other plant material in their diets. Flighted species are over-represented in wooded environments and higher strata and flightless species predominate in low alpine and grassland habitats. The frugivore-fruiting plant interaction group shows some indications of ecological generalism as frugivorous species consume a range of fruit sizes across all vegetation strata and fruit-bearing plants have lower species diversity and occupy a wide range of habitats. Granivores are over-represented in wetland habitats and the eastern South Island. The importance of species which unintentionally disperse adhesive seed depends on whether they are volant (higher importance in coastal environments) or flightless (higher importance dry grasslands and in low alpine areas). A subgroup of birds, such as the ratite Apteryx spp. and the now extinct Dinornithiform moa, with loose feathers (“velcro” species) are over-represented in lower vegetation strata and this matches the zone where many attachment-dispersed plant species present their seed.     

Comparative analysis of growth and physio-biochemical responses of Hydrilla verticillata to different sediments in freshwater microcosms

The long-term effect of Hydrilla verticillata to different dredged sediments was investigated in laboratory aquatic microcosms. Two kinds of lake sediments, i.e., fertile sludge and brown clay, which were acquired by deep or slight dredging and differed in their nutrients levels and particle sizes, were used as growth substrate. One year after H. verticillata seedlings were transplanted into the microcosms, growth and physio-biochemical characters of the plants were estimated. In our study, sludge sediment eventually promoted the increase of the biomass, shoot length, bifurcation, nitrogen, phosphorus, soluble protein content of H. verticillata, but not root number and root mass ratio. Additionally, the soluble sugar content was significantly lower in summer (August) in the sludge sediment. Sediment nutrients also increased photosynthetic pigment content, while POD and SOD activities were not affected. The fertile sludge and brown clay formed by different dredging are quite different, but both sediments could meet the requirement of H. verticillata during early growth stage. The fertile sediment after moderate dredge-up fit the growth of H. verticillata. The brown clay formed after excessive dredging showed a lack of nutrients and therefore led to the weak growth of the aquatic plants. It is concluded that moderate dredging was suitable for H. verticillata growth and for absorbing nitrogen and phosphorus from eutrophic fresh water. The experiment also suggested that submersed aquatic plant restoration in freshwater can be promoted by human assistance.