Invasive Plant and Experimental Venue Affect Tadpole Performance

Introductions of non-native predators and competitors appear to contribute to worldwide amphibian declines; however, potential negative impacts of invasive plants on habitat quality and amphibian populations have not been examined. Loss of diversity and alterations in ecosystem function associated with plant invasions may disrupt food webs, potentially leading to further declines of already threatened amphibian populations. We used a combination of small bins, mesocosms, and field experiments to examine the impacts of Eurasian purple loosestrife (Lythrum salicaria) replacing native cattails (Typha latifolia) in North American freshwater wetlands on survival, developmental rate, and diet (freshwater algae) of American toad (Bufo americanus) tadpoles. Tadpoles developed slower in L. salicaria compared to tadpoles developing in T. latifolia. This effect was consistent across experimental venues, although mesocosms showed this effect only in the second year of our study. Survival and development rates were always more variable in purple loosestrife than in cattail. In bins, tadpoles showed significantly reduced survival when raised in purple loosestrife extract and addition of leaf litter exacerbated this negative effect. Tadpole survival rates in mesocosms and field cages were not significantly different between plant species, most likely an effect of high variability among replicates. We suspect a combination of direct toxicity of high tannin concentrations in L. salicaria leaves and their indirect negative impacts on aquatic food webs are responsible for these results. Tadpole gut analyses revealed differences in algal communities among venues and between L. salicaria and T. latifolia suggesting that alterations in tadpole food quality and quantity contribute to the observed reduced tadpole performance. The replacement of native wetland plant species by L. salicaria does not represent a simple exchange of ecological equivalents and the function of invaded habitats for native species has clearly changed. While we were investigating only a single amphibian species, our results suggest that the impact of L. salicaria on ecosystem processes and aquatic food webs may be more general and likely to negatively affect other wetland species. The threats non-indigenous plants represent for amphibian populations and food webs may be underestimated, and warrant further investigation.     

Is the increased vigour of invasive weeds explained by a trade-off between growth and herbivore resistance?

Blossey and Nötzold (1995) recently hypothesised that the increased vigour of certain invasive plant species has been at the expense of defences against natural enemies. A prediction of their evolution of increased competitive ability (EICA) hypothesis is that invasive genotypes are relatively poorly defended. We tested this prediction with herbivore bioassays and with direct quantification of plant secondary metabolites comparing non-indigenous genotypes of Lythrum salicaria L. (purple loosestrife) with indigenous forms. The herbivore bioassays revealed no significant intra-specific variation in herbivore resistance between indigenous and non-indigenous hosts. The phenolic content of L. salicaria leaves was significantly higher in indigenous genotypes, as predicted by the EICA hypothesis. The average phenolic content of leaves (regardless of their origin) was, however, low, implying that the role of plant phenolics in purple loosestrife anti-herbivore defence is probably limited. It is suggested that the EICA hypothesis, as tested in the current study, does not explain the increased vigour of L. salicaria in non-indigenous habitats.     

Before, During and After: The Need for Long-term Monitoring in Invasive Plant Species Management

The invasion of non-indigenous plants is considered one of the primary threats to rare and endangered species as well as to the integrity and function of North American ecosystems. However, many of the suspected negative ecosystem impacts are based on anecdotal evidence. For example, there is almost unanimous agreement among natural resource managers of the detrimental ecological impacts of species such as Lythrum salicaria (purple loosestrife), Phragmites australis (common reed) and Alliaria petiolata (garlic mustard) but convincing documentation is scarce. Experimental and theoretical ecology predicts large ecosystem impacts of the most widespread invasive species. However, it is difficult to prioritize control of species that occur at intermediate densities. Long-term monitoring before and during the invasion as well as before, during and after any control attempts can provide valuable ecological information. In particular, it is important to understand how changes in the abundance of species influence ecosystem properties and processes which, in turn, will help guide management decisions. Ideally, this monitoring has to go beyond 'simple impacts on plant communities, involve cross-disciplinary teams of scientists and should incorporate many different taxa and their interactions. Monitoring design and data collection should be sophisticated enough to allow statistically sound data analysis. The available information will be paramount in (1) developing new political and scientific guidelines in invasive species management, (2) helping resolve potential conflicts of interest and (3) helping change public attitudes regarding growth, sale, and control of non-indigenous species.     

Should we care about purple loosestrife? The history of an invasive plant in North America

Purple loosestrife (Lythrum salicaria L., Lythraceae) is considered one of the worst invasive plant species in the world. In this paper, I reconstruct how purple loosestrife quickly became, after a long (150 years) period of indifference, the persona non grata of North American wetlands. I then compare the portrayal of the species in newspapers (907 articles) to that supported by the scientific literature (38 peer-review papers). The depiction of purple loosestrife in scientific studies (lacking definition) is far removed from that in newspapers (alarming). Some native species likely suffer from an invasion, but stating that this plant has large negative impacts on wetlands is probably exaggerated. If purple loosestrife is not a primary cause of extinction or a major contributor to the decline of other species, but is instead an indicator of anthropogenic disturbances, the resources and efforts devoted to removing this species might be better focused on more effective means to protect wetlands against disturbances.     

Fluctuating asymmetry as an ecological indicator of heavy metal stress in Lythrum salicaria

Fluctuating asymmetry (FA), a measure of random deviation of organismal traits from perfect bilateral symmetry, can be induced by environmental and/or genetic stress. We have conducted a replicated experiment to test the effect of a locally abundant heavy metal pollutant, lead, on the FA of a common wetland invasive species, Lythrum salicaria (purple loosestrife). The exposure to lead significantly reduced the shoot length, number and length of leaves as well as the biomass, but increased FA of L. salicaria leaves. The investigation suggests that FA in L. salicaria may be used as an ecological indicator to identify environmental stress caused by certain heavy metal pollutants. However, more studies are needed for the use of FA in L. salicaria as a comprehensive tool for biomonitoring of a wider array of environmental pollutants.     

Dispersal of Galerucella pusilla and G. calmariensis via passive water transport in the Columbia River Estuary

Dispersal of biological control agents and their subsequent population growth can be a major determinant of the success of landscape-scale weed control programs. Biocontrol agents must be able to disperse across the distances between patches of host plants in order to colonize and control their targets. The presence of three species of biocontrol agents for purple loosestrife (Lythrum salicaria L.): Galerucella calmariensis L. (Coleoptera: Chrysomelidae), Galerucella pusilla Duftschmid (Coleoptera: Chrysomelidae), and Nanophyes marmoratus Goeze (Coleoptera: Brentidae), on relatively remote islands in the Columbia River Estuary (CRE) indicate that these organisms have the ability to disperse across large expanses of open flowing water to colonize remote sites. Previous studies suggest that colonization of these islands by active flight is highly unlikely; therefore, some other dispersal mechanism must be responsible for colonization. A spatial database of all known biocontrol agent release sites for purple loosestrife within 68 river kilometers of our CRE study area was developed and field surveys for biocontrol agents were conducted. A GIS was used to model dispersal distances between biocontrol agent recovery sites and the nearest conspecific release site. Tidal water flow within the CRE was assessed as a potential dispersal mechanism across the modeled distances. The ability of the biocontrol agents to withstand submersion was evaluated in field tests. Our results indicate that it is highly likely that passive water transport has been responsible for some of the long-distance open-water dispersal that would have been necessary for colonization of the remote islands where biocontrol agents were recovered.     

Detritus Quality and Locality Determines Survival and Mass,but Not Export,of Wood Frogs at Metamorphosis

Single-site experiments have demonstrated detritus quality in wetlands can have strongly negative, neutral, and even positive influences on wildlife. However, an examination of the influence of detritus quality across several regions is lacking and can provide information on whether impacts from variation in detritus quality are consistent across species with wide ranges. To address this gap in regional studies we examined effects of emergent and allochthonous detritus of different nutrient qualities on amphibians and assessed a mechanism that may contribute to potential impacts. We used aquatic mesocosms to raise wood frogs (Rana sylvatica) from two regions of the United States with whole plants from purple loosestrife (Lythrum salicaria), leaf litter from native hardwood trees, and a mixture of both. We examined several metrics of amphibian fitness and life history, including survival, number of days to metamorphosis, and size at metamorphosis. Further, we quantified whether the effects of detritus type could translate to variation in anuran biomass or standing stock of nitrogen or phosphorus export. Our results show detritus with high nutrient quality (purple loosestrife) negatively influenced survival of wood frogs, but increased size of metamorphic individuals in two different regions of the United States. Despite the decrease in survival, the increase in size of post-metamorphic anurans raised with high quality detritus resulted in anuran biomass and standing stock of N and P export being similar across treatments at both locations. These results further demonstrate the role of plant quality in shaping wetland ecosystem dynamics, and represent the first demonstration that effects are consistent within species across ecoregional boundaries.     

Matrix habitat and plant damage influence colonization of purple loosestrife patches by specialist leaf-beetles

The characteristics of the matrix, that is, the unsuitable habitat connecting host-plant patches may facilitate or limit herbivore movement thus affecting their population dynamics. We evaluated the effect of matrix habitat, distance between patches, and plant damage on movement of two leaf-beetles (Galerucella calmariensis Linnaeus and G. pusilla Duft) introduced to North America as biocontrol agents of the invasive purple loosestrife (Lythrum salicaria Linnaeus). Mark-recapture/resight experiments indicated (1) that leaf-beetles are more likely to colonize purple loosestrife patches surrounded by meadow than forest; (2) that previously attacked purple loosestrife plants are more likely to be colonized by Galerucella spp. than unattacked plants, especially in the forest habitat; and (3) that leaf beetle colonization of purple loosestrife decreased with distance from release point. Low colonization rates of purple loosestrife patches embedded in forests suggest either insufficient detection or active avoidance of such habitats. Biological control programs intend to manage dispersal of specialized insect herbivores for the purpose of sufficient and sustained control of their host plants. Such management needs to be informed by knowledge of interactions of habitat structure, plant damage, and dispersal capabilities of herbivores to facilitate release programs and control at the local and regional level.     

The effects of flooding,plant traits,and predation on purple loosestrife leaf‐beetles

Classical biological weed control is based on the premise that introducing specialized natural enemies from the native range re‐establishes herbivore control of plant invaders, ultimately leading to negative population growth rates. Evidence from past biocontrol programs suggests that herbivores are not solely responsible for shaping plant demography. Diverse environmental conditions in the introduced range may not only affect demography, but also influence top‐down control of target plants. We investigated how flooding affects impacts of predators (top‐down) and plant quality (bottom‐up) on performance of two leaf‐beetles, Galerucella calmariensis L. and Galerucella pusilla Duftschmid (Coleoptera: Chrysomelidae: Galerucini), released in North America as biocontrol agents of purple loosestrife, Lythrum salicaria L. (Lythraceae). Predation and flooding regime have been linked to low leaf‐beetle recruitment at sites where insects failed to attain outbreak populations. Predator exclusion experiments at adjacent flooded and non‐flooded sites indicated a positive effect of flooding on leaf‐beetle survival for all developmental stages, whereas predator exposure had little effect. There was no difference in predation rates at sites with successful or failed purple loosestrife control, questioning the importance of predation in limiting growth and impact of these biocontrol agents’ populations. Effect of flooding on purple loosestrife quality was evaluated in a common garden study where plants were grown under different flooding treatments. Plants grown in flooded soil had higher water content and lower tannic acid concentration than plants grown in well‐drained soil. Consistent with field observations, leaf‐beetle oviposition rate and survival were higher on flooded plants. Results indicate that both bottom‐up and top‐down forces operate on Galerucella populations, yet their relative strength is mediated by flooding regime. Ignoring intricacies of plant‐herbivore and trophic interactions in the introduced range appears to be a major handicap for the improvement of weed biocontrol programs.     

Habitat and Population Effects on the Germination and Early Survival of the Invasive Weed, Lythrum salicaria L. (purple loosestrife)

In this study, we examined how environmental factors and differences among populations of Lythrum salicaria L. (Lythraceae; purple loosestrife) influenced the establishment of this species into both its preferred wetland habitat and secondary upland habitat. Seeds and seedlings from five parents from each of the three populations were planted in each wetland and upland of two sites in both disturbed and undisturbed plots. More seeds germinated in wetland than in upland sites (31% as compared to 11%) and, although a disturbance resulted in greater germination (23% as compared to 17% in undisturbed plots), there was no interaction with habitats. The results of a simultaneous greenhouse germination study in which seeds were planted under saturated and drained conditions suggest that the higher germination observed in wetland sites was due to the higher moisture present at such sites (79% vs. 45% in saturated and drained conditions, respectively). Disturbance enhanced both germination and growth of L. salicaria and this was especially so for seedlings. Dry weights of seedlings from disturbed plots were five times greater than those from undisturbed plots ( =9.2 vs. 1.8g in disturbed and undisturbed conditions, respectively). In addition to environmental factors, population differences are likely to play an important role in the spread of L. salicaria as we found differences between the populations in both life stages examined. One of the populations germinated equally well in both moisture treatments. In our seedling establishment experiment, one of the populations produced both the smallest plants and the most and earliest flowers, suggesting superior reproduction.     

Restoration of Freshwater Intertidal Habitat Functions at Spencer Island, Everett, Washington

In November 1994 dikes were breached around Spencer Island, restoring tidal inundation and connections to the Snohomish River estuary, Washington. Approximately 23.7 ha (58.5 ac) of palustrine wetlands previously dominated by Phalaris arundinacea (reed canarygrass) now experience diurnal tides and are in the process of transition to a freshwater tidal system. It was expected that brackish water would accompany the return of tidal influence to the site, but post‐project monitoring has revealed little evidence of salinity. Pre‐ and post‐project monitoring of changes in habitat function included aerial photography, vegetation and fish sampling, and benthic prey studies. To date site changes include (1) die back of pre‐project vegetation, development of tidal mudflat, and emergent wetland habitats, with recruitment of vegetation typical of freshwater tidal wetlands; (2) presence of juvenile coho, chum, and chinook salmon that feed on invertebrate prey typical of the site; (3) presence of three distinct benthic invertebrate assemblages in the project area; and (4) some invasion by Lythrum salicaria (purple loosestrife). The unexpected freshwater conditions, the lack of published information about tidal oligohaline marshes in the Pacific Northwest, the use of the site by endangered salmonid species, and the invasion by an undesired plant species underscore the importance of long‐term monitoring at the site.     

Potential of aquatic macrophytes and artificial floating island for removing contaminants

Eutrophication is a major environmental issue that mankind is facing today as a result of rapid development. To reduce the eutrophication problem, we are testing an innovative artificial floating islands (AFIs) approach. AFIs include aquatic plants composed of emergent and floating macrophytes. In this experimental approach, two aquatic plants were selected to compare with the control group in order to evaluate the capacity of AFIs. These two emergent macrophytes were evaluated for a 40-day timeframe to examine their ability to purify waste water. The results showed that an AFI with purple loosestrife (Lythrum salicaria Linn.) and one with yellow-flowered iris (Iris wilsonii) have a strong ability to remove nitrogen, phosphorus, and other pollutants from water bodies. The pollutant removal rates of the AFIs with purple loosestrife and yellow-flowered iris were almost over 50%. The chemical oxygen demand (COD), total nitrogen (TN), and total phosphorus (TP) removal rates of the AFI with purple loosestrife were about 75%, 57%, 71%, respectively. At the same time, the COD, TN, and TP removal rates of the AFI with yellow-flowered iris were 60%, 49%, and 68%, respectively. AFIs with aeration are also a good way to remove pollutants. This study reveals that AFIs can be incorporated into wetlands to reduce the effects of water contamination and help strengthen wetland restoration.     

Quantifying the effects of distance and conspecifics on colonization: experiments and models using the loosestrife leaf beetle, Galerucella calmariensis

The ability of an insect to disperse to new habitat patches is difficult to quantify, but key to the establishment and persistence of populations. In this study, we examined dispersal of the phytophagous chrysomelid beetle, Galerucella calmariensis, which is currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), an aggressive wetland weed. We used a mark, release, and recapture approach to determine how rates of colonization of host patches by this beetle are influenced by the distance of the patch from the source of dispersers, and by the presence of conspecifics at the patch. We released color-coded beetles at six distances from a long, linear patch of purple loosestrife that was divided into segments with and without conspecifics. We observed initial flight directions as beetles left the release points and collected all beetles that settled at the target patch. We found a bias in initial flight toward the target for distances up to 50 m. Over the 7 days of the experiment, beetles arrived at the target from all release points, including the farthest release point, 847 m away. G. calmariensis was strongly attracted to conspecifics when settling after dispersal; 86% of the 582 recovered beetles came from the segments inhabited by conspecifics. The probability of an individual arriving at the patch declined steeply with release distance. This relationship fits a model in which beetles move in a random direction and stop if they intercept the target patch, and where beetles are lost at a constant rate with distance travelled. The dispersal and patch-colonizing behavior of G. calmariensis is likely to have important consequences for the biological control program against purple loosestrife. Received: 23 January 1996 / Accepted:30 September 1996     

The influence of below ground herbivory and plant competition on growth and biomass allocation of purple loosestrife

Experiments investigating plant-herbivore interactions have primarily focused on above-ground herbivory, with occasional studies evaluating the effect of below-ground herbivores on plant performance. This study investigated the growth of the wetland perennial Lythrum salicaria (purple loosestrife) under three levels of root herbivory by the weevil Hylobiustransversovittatus and three levels of plant competition by the grass Phleumpratense in a common garden. Plant growth, flowering phenology, and biomass allocation patterns of purple loosestrife were recorded for two growing seasons. During the first year, root herbivory reduced plant height; plant competition delayed flowering; and the interaction of root herbivory and plant competition resulted in reductions in plant height, shoot weight and total dry biomass. Plant competition or larval feeding did not affect the biomass allocation pattern in the first year. These results indicate the importance of interactions of plant competition and herbivory in reducing plant performance – at least during the establishment period of purple loosestrife. In the second growing season, root herbivory reduced plant height, biomass of all plant parts, delayed and shortened the flowering period, and changed the biomass allocation patterns. Plant competition delayed flowering and reduced the dry weight of fine roots. The interaction of root herbivory and plant competition delayed flowering. Root herbivory was more important than plant competition in reducing the performance of established purple loosestrife plants. This was due, in part, to intense intraspecific competition among the grass individuals effectively preventing shoot elongation of P. pratense and resulting in a carpet like growth. Received: 3 April 1997 / Accepted: 27 July 1997     

The implications of accepting untestedhypotheses: a review of the effectsof purple loosestrife (Lythrum salicaria)in North America

The acceptance of poorly tested hypotheses has adverse scientific consequences, and may have adverse ecological and social consequences. The hypothesis that purple loosestrife (Lythrum salicaria) has deleterious effects on North American wetlands is an example. We traced the history of purple loosestrife and its control in North America and found little scientific evidence consistent with the hypothesis that purple loosestrife has deleterious effects. The most commonly cited study of the effects of purple loosestrife on native flora and fauna produced inconclusive results. The general acceptance of this hypothesis, however, has resulted in the introduction of nonindigenous insects for biological control. Efforts to control purple loosestrife may be misplayed and may have long-term ecological consequences if loosestrife does not have the impact it is believed to have. The acceptance of this hypothesis using scientific justifications may affect future scientific credibil ity. Careful evaluation of the precautionary principle is necessary when considering the control of nonindigenous organisms.     

Comparison of created and natural freshwater emergent wetlands in Connecticut (USA)

Five three- to four-year old created palustrine/emergent wetland sites were compared with five nearby natural wetlands of comparable size and type. Hydrologic, soil and vegetation data were compiled over a nearly two-year period (1988-90). Created sites, which were located along major highways, exhibited more open water, greater water depth, and greater fluctuation in water depth than natural wetlands. Typical wetland soils exhibiting mottling and organic accumulation were wanting in created sites as compared with natural sites. Typha latifolia (common cattail) was the characteristic emergent vegetation at created sites, whereas a more diverse mosaic of emergent wetland species was often associated with Typha at the natural sites. Species richness was slightly higher in created (22–45) vs. natural (20–39) wetlands, but the mean difference (33 vs. 30) was not significant. Nearly half (44%) of the 54 wetland taxa found at the various study sites were more frequently recorded at created than natural wetlands. The presence of mycorrhizae in roots of Typha angustifolia (narrow-leaved cattail) and Phragmites australis (common reed) was greater at created than natural wetlands, which may be related to differential nutrient availability. Wildlife use at all sites ranged from occasional to rare, with more sightings of different species in the natural (39) than created (29) wetlands. The presence of P. australis and introduced Lythrum salicaria (purple loosestrife) may pose a threat to future species richness at the created sites. One created site has permanent flow-through hydrology, and its vegetation and wildlife somewhat mimic a natural wetland; however, the presence of P. australis and its potential spread pose an uncertain future for this site. This study suggests the possibility of creating small palustrine/emergent wetlands having certain functions associated with natural wetlands, such as flood water storage, sediment accretion and wildlife habitat. It is premature to evaluate fully the outcome of these wetland creation efforts. A decade or more is needed, emphasizing the importance of long term monitoring and the need to establish demonstration areas.     

Comparison of life history traits between invasive and native populations of purple loosestrife (Lythrum salicaria) using nonlinear mixed effects model

We compared growth patterns of invasive and native populations of purple loosestrife (Lythrum salicaria) while varying water and nutrient levels. We examined three life-history traits (height, number of branches, and the size of largest leaf) during the growth period adopting a nonlinear mixed effects model. Invasive populations were found to be slower in shoot elongation but grew to be taller than native populations. Invasive populations produced more branches than natives only in the high water, high nutrient treatment. Invasive populations had a similar increase in the size of the largest leaf compared to natives, but ultimately produced a greater size of largest leaf than natives. Invasive populations were found to display a greater vegetative expansion, but this was not strongly affected by our treatments.     

Comparison of life history traits between invasive and native populations of purple loosestrife (Lythrum salicaria) using nonlinear mixed effects model

We compared growth patterns of invasive and native populations of purple loosestrife (Lythrum salicaria) while varying water and nutrient levels. We examined three life-history traits (height, number of branches, and the size of largest leaf) during the growth period adopting a nonlinear mixed effects model. Invasive populations were found to be slower in shoot elongation but grew to be taller than native populations. Invasive populations produced more branches than natives only in the high water, high nutrient treatment. Invasive populations had a similar increase in the size of the largest leaf compared to natives, but ultimately produced a greater size of largest leaf than natives. Invasive populations were found to display a greater vegetative expansion, but this was not strongly affected by our treatments.     

Experimental Invasions Using Biological Control Introductions: The Influence of Release Size on the Chance of Population Establishment

Introductions of biological control organisms offer a unique opportunity to experimentally study the process of invasion by exotic species. I used two chrysomelid beetles, Galerucella calmariensis and Galerucella pusilla, which are currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), to determine how the initial size of a release affects the probability that the introduced population grows and persists. I released both species into stands of their host plant at 36 sites scattered throughout central New York State using four release sizes: 20, 60, 180, and 540. I returned to these sites over the next 3 years to census the populations. For both species, the probability of population establishment increased with release size. Population growth rates also depended positively on release size. The implication from these results is that the demographic factors whose influence depends on population size or density such as demographic stochasticity, Allee effects, and genetics play important roles in the establishment of invading populations. A second set of releases was used to determine if it was at all possible for a single gravid female to found a population. Out of twenty individual females released, one female (a G. calmariensis) founded a population that persisted until the end of the study (3 generations). This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of an invasive plant transcend ecosystem boundaries through a dragonfly-mediated trophic pathway

Trophic interactions can strongly influence the structure and function of terrestrial and aquatic communities through top-down and bottom-up processes. Species with life stages in both terrestrial and aquatic systems may be particularly likely to link the effects of trophic interactions across ecosystem boundaries. Using experimental wetlands planted with purple loosestrife (Lythrum salicaria), we tested the degree to which the bottom-up effects of floral density of this invasive plant could trigger a chain of interactions, changing the behavior of terrestrial flying insect prey and predators and ultimately cascading through top-down interactions to alter lower trophic levels in the aquatic community. The results of our experiment support the linkage of terrestrial and aquatic food webs through this hypothesized pathway, with high loosestrife floral density treatments attracting high levels of visiting insect pollinators and predatory adult dragonflies. High floral densities were also associated with increased adult dragonfly oviposition and subsequently high larval dragonfly abundance in the aquatic community. Finally, high-flower treatments were coupled with changes in zooplankton species richness and shifts in the composition of zooplankton communities. Through changes in animal behavior and trophic interactions in terrestrial and aquatic systems, this work illustrates the broad and potentially cryptic effects of invasive species, and provides additional compelling motivation for ecologists to conduct investigations that cross traditional ecosystem boundaries.