Size matters: predation of fish eggs and larvae by native and invasive amphipods

Invasive predators can have dramatic impacts on invaded communities. Extreme declines in macroinvertebrate populations often follow killer shrimp (Dikerogammarus villosus) invasions. There are concerns over similar impacts on fish through predation of eggs and larvae, but these remain poorly quantified. We compare the predatory impact of invasive and native amphipods (D. villosus and Gammarus pulex) on fish eggs and larvae (ghost carp Cyprinus carpio and brown trout Salmo trutta) in the laboratory. We use size-matched amphipods, as well as larger D. villosus reflecting natural sizes. We quantify functional responses, and electivity amongst eggs or larvae and alternative food items (invertebrate, plant and decaying leaf). D. villosus, especially large individuals, were more likely than G. pulex to kill trout larvae. However, the magnitude of predation was low (seldom more than one larva killed over 48 h). Trout eggs were very rarely killed. In contrast, carp eggs and larvae were readily killed and consumed by all amphipod groups. Large D. villosus had maximum feeding rates 1.6–2.0 times higher than the smaller amphipods, whose functional responses did not differ. In electivity experiments with carp eggs, large D. villosus consumed the most eggs and the most food in total. However, in experiments with larvae, consumption did not differ between amphipod groups. Overall, our data suggest D. villosus will have a greater predatory impact on fish populations than G. pulex, primarily due to its larger size. Higher invader abundance could amplify this difference. The additional predatory pressure could reduce recruitment into fish populations.     

Differences in interactions of aboveground and belowground herbivores on the invasive plant <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> and native host <Emphasis Type="Italic">A. sessilis</Emphasis>

Plant invasions may result in novel plant-herbivore interactions. However, we know little about whether and how invasive plants can mediate native above- and belowground herbivore interactions. In this study, we conducted greenhouse experiments to examine the interaction between a native defoliating beetle, Cassida piperata, and a native root-knot nematode, Meloidogyne incognita, on the invasive alligator weed, Alternanthera philoxeroides. We also included their native host A. sessilis in the experiments to examine whether the patterns of above- and belowground herbivore interaction vary with host plants (invasive vs. native). We analyzed total carbon and nitrogen in leaves and roots attacked by M. incognita and C. piperata. M. incognita slightly negatively affected feeding by C. piperata on A. philoxeroides, and the leaf area damaged decreased as the number of M. incognita increased. M. incognita had a negative impact on total leaf nitrogen, but had no impact on total leaf carbon. M. incognita egg production on A. philoxeroides roots decreased as the amount of damage caused by C. piperata increased. Herbivory by C. piperata did not affect total root carbon or nitrogen. M. incognita and C. piperata did not affect each other on the native plant A. sessilis. These results suggest that invasive plants can mediate native above- and belowground herbivore interactions. The knowledge of how invasive plants affect those interactions is crucial for better understanding the impacts of biological invasions on native above- and belowground organisms.     

Opportunistic omnivory impairs our ability to predict invasive species impacts from functional response comparisons

Comparing the relationship between resource use and resource availability (i.e. the functional response, FR) between two predators can provide useful insights on their relative predatory impacts. For instance in invasion ecology, an increase in the predation pressure on local prey populations can be predicted from a significant difference in FR revealing a higher FR for the invasive predator compared to the native trophic analogue it may replace. In traditional FR experiments, the focal prey species is the only source of food. This may lead to misinterpretations with opportunistic omnivores that are able to cope with different resource availabilities in their natural environment, and whose predation rate may therefore be modulated by the presence of alternative resources. To address this question, we compared the FR of two freshwater gammarid species known to behave as opportunistic omnivores: the invasive “killer shrimp” Dikerogammarus villosus and the native Gammarus pulex, in a treatment with a focal prey species as the only food source (the water flea Daphnia magna) and in a treatment with the focal prey and an alternative food source (Carpinus betulus leaves). D. villosus showed a significantly higher FR than G. pulex with water fleas only and providing leaf litter suppressed this difference. The predatory impact of D. villosus might therefore be modulated by the relative availability of live prey compared to the alternative food sources. Increasing the realism of FR experiments through the inclusion of abundant and easily accessible alternative resources, like leaf litter for benthic invertebrates, should refine the predictions made from FR comparisons.     

Multiple invasions in urbanized landscapes: interactions between the invasive garden ant <Emphasis Type="Italic">Lasius neglectus</Emphasis> and Japanese knotweeds (<Emphasis Type="Italic">Fallopia</Emphasis> spp.)

Urbanized landscapes are the theater of multiple simultaneous biological invasions likely to affect spread dynamics when co-occurring introduced species interact with each other. Interactions between widespread invaders call for particular attention because they are likely to be common and because non-additive outcomes of such associations might induce negative consequences (e.g., enhanced population growth increasing impacts or resistance to control). We explored the invasions of two widespread invasive taxa: the Japanese knotweed species complex Fallopia spp. and the invasive garden ant Lasius neglectus, in the urban area of Lyon (France). First, we investigated landscape habitat preferences as well as co-occurrence rates of the two species. We showed that Fallopia spp. and L. neglectus had broadly overlapping environmental preferences (measured by seven landscape variables), but their landscape co-occurrence pattern was random, indicating independent spread and non-obligatory association. Second, as Fallopia spp. produce extra-floral nectar, we estimated the amount of nectar L. neglectus used under field conditions without ant competitors. We estimated that L. neglectus collected 150–321 kg of nectar in the month of April (when nectar production is peaking) in a 1162 m² knotweed patch, an amount likely to boost ant population growth. Finally, at six patches of Fallopia spp. surveyed, herbivory levels were low (1–6% loss of leaf surface area) but no relationship between ant abundance (native and invasive) and loss of leaf surface was found. Co-occurrences of Fallopia spp. and L. neglectus are likely to become more common as both taxa colonize landscapes, which could favor the spread and invasion success of the invasive ant.     

Similar responses in morphology,growth, biomass allocation,and photosynthesis in invasive <Emphasis Type="Italic">Wedelia trilobata</Emphasis> and native congeners to CO<Subscript>2</Subscript> enrichment

Both global change and biological invasions threaten biodiversity worldwide. However, their interactions and related mechanisms are still not well elucidated. To elucidate potential traits contributing to invasiveness and whether ongoing increase in CO₂ aggravates invasions, noxious invasive Wedelia trilobata and native Wedelia urticifolia and Wedelia chinensis were compared under ambient and doubled atmospheric CO₂ concentrations in terms of growth, biomass allocation, morphology, and physiology. The invader had consistently higher leaf mass fraction (LMF) and specific leaf area than the natives, contributing to a higher leaf area ratio, and therefore to faster growth and invasiveness. The higher LMF of the invader was due to lower root mass fraction and higher fine root percent. On the other hand, the invader allocated a higher fraction of leaf nitrogen (N) to photosynthetic apparatus, which was associated with its higher photosynthetic rate, and resource use efficiency. All these traits collectively contributed to its invasiveness. CO₂ enrichment increased growth of all studied species by increasing actual photosynthesis, although it decreased photosynthetic capacities due to decreased leaf and photosynthetic N contents. Responses of the invasive and native plants to elevated CO₂ were not significantly different, indicating that the ongoing increase in CO₂ may not aggravate biological invasions, inconsistent with the prevailing results in references. Therefore, more comparative studies of related invasive and native plants are needed to elucidate whether CO₂ enrichment facilitates invasions.     

Tracking an invasion: community changes in hardwood forests following the arrival of <Emphasis Type="Italic">Amynthas agrestis</Emphasis> and <Emphasis Type="Italic">Amynthas tokioensis</Emphasis> in Wisconsin

Because Upper Midwest temperate forests lack native earthworms, the invasions of European and Asian earthworms can significantly alter soils and understory vegetation. Earthworms’ ability to increase leaf litter decay, alter nutrient cycling by mixing the organic layer with mineral soil, and decrease plant species richness leads to concern about the Asian ‘jumping earthworm’ (Amynthas agrestis and A. tokioensis) species that were recorded in the University of Wisconsin—Madison Arboretum in 2013. In 2015, we found A. agrestis and A. tokioensis in a distinct 8-ha region of a 23-ha hardwood forest surveyed in the Arboretum; by 2016 A. agrestis and A. tokioensis had spread over an additional 7 ha. Plots also contained the European earthworm species Lumbricus terrestris, L. rubellus, and Apporectodea spp., whose distributions decreased from 2015 to 2016. While leaf litter, plant species richness, and tree and shrub seedling abundance were generally reduced in areas with European earthworms, they were typically slightly increased in areas with A. agrestis and A. tokioensis versus those without. Although our results do not show substantial impacts of A. agrestis and A. tokioensis on vegetation in the initial years of invasion, the rapid replacement of European earthworms by A. agrestis and A. tokioensis suggests continued monitoring of these new invasive species is important to better understand their potential to change the Upper Midwest’s forests.     

High phenolic content fails to deter mesograzer consumption of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> (Eurasian watermilfoil) in New England

Eurasian watermilfoil (Myriophyllum spicatum) is often considered one of the most aggressive macrophyte invaders in freshwater habitats throughout the USA. However, conditions leading to successful milfoil invasions are not well understood. This study sought to illuminate the role of herbivores in determining milfoil invasion success via the potential mechanisms of enemy release and biotic resistance. We determined feeding preferences of three herbivores native to the northeastern United States and measured macrophyte phenolic content, which may act as an herbivore feeding deterrent. We found that phenolic content in milfoil was more than two times higher than in the most abundant native macrophytes at our study sites, consistent with enemy release. However, laboratory feeding experiments demonstrated that milfoil phenolics did not deter amphipod (Hyalella azteca), snail (Physella sp.), or weevil (Euhrychiopsis lecontei) herbivory. Furthermore, amphipod consumption rates in our study were an order of magnitude higher than amphipod consumption rates reported in milfoil’s native range, contrary to the predictions of enemy release. Amphipods and snails from habitats invaded by milfoil consumed similar quantities of both milfoil and the low-phenolic native plant Elodea canadensis. In contrast, weevils consumed milfoil but not E. canadensis in choice experiments. Amphipods collected from milfoil-free habitats also readily consumed milfoil, and they consumed 2.5 times more milfoil than E. canadensis in a choice feeding trial. These results suggest that high phenolic levels do not prevent native herbivores from consuming invasive milfoil. Instead, native generalist grazers like amphipods and snails may limit milfoil proliferation and provide a measure of biotic resistance.     

Influences of the invasive tamarisk leaf beetle (<Emphasis Type="Italic">Diorhabda carinulata</Emphasis>) on avian diets along the Dolores River in Southwestern Colorado USA

The tamarisk leaf beetle (Diorhabda carinulata), introduced from Eurasia in 2001 as a biological control agent for the invasive plant Tamarix ramosissima, has spread widely throughout the western USA. With D. carinulata now very abundant, scientists and restoration managers have questioned what influence this introduced arthropod might have upon the avian component of riparian ecosystems. From 2009 through 2012 we studied the consequences of biological invasions of the introduced tamarisk shrub and tamarisk leaf beetles on the diets of native birds along the Dolores River in southwestern Colorado, USA. We examined avian foraging behavior, sampled the arthropod community, documented bird diets and the use of invasive tamarisk shrubs and tamarisk leaf beetles by birds. We documented D. carinulata abundance, on what plants the beetles occurred, and to what degree they were consumed by birds as compared to other arthropods. We hypothesized that if D. carinulata is an important new avian food source, birds should consume beetles at least in proportion to their abundance. We also hypothesized that birds should forage more in tamarisk in the late summer when tamarisk leaf beetle larvae are more abundant than in early summer, and that birds should select beetle-damaged tamarisk shrubs. We found that D. carinulata composed 24.0 percent (±?19.9–27.4%) and 35.4% biomass of all collected arthropods. From the gut contents of 188 birds (25 passerine species), only four species (n?=?11 birds) contained tamarisk leaf beetle parts. Although D. carinulata comprised one-quarter of total insect abundance, frequency of occurrence in bird gut contents was only 2.1% by abundance and 3.4% biomass. Birds used tamarisk shrubs for foraging in proportion to their availability, but foraging frequency did not increase during the late summer when more tamarisk leaf beetles were present and birds avoided beetle-damaged tamarisk shrubs. Despite D. carinulata being the most abundant arthropod in the environment, these invasive beetles were not frequently consumed by birds and seem not to provide a significant contribution to avian diets.     

Differential invasion success in aquatic invasive species: the role of within- and among-population genetic diversity

Despite a well-developed theoretical basis for the role of genetic diversity in the colonization process, contemporary investigations of genetic diversity in biological invasions have downplayed its importance. Observed reductions in genetic diversity have been argued to have a limited effect on the success of establishment and impact based on empirical studies; however, those studies rarely include assessment of failed or comparatively less-successful biological invasions. We address this gap by comparing genetic diversity at microsatellite loci for taxonomically and geographically paired aquatic invasive species. Our four species pairs contain one highly successful and one less-successful invasive species (Gobies: Neogobius melanostomus, Proterorhinus semilunaris; waterfleas: Bythotrephes longimanus, Cercopagis pengoi; oysters: Crassostrea gigas, Crassostrea virginica; tunicates: Bortylloides violaceous, Botryllus schlosseri). We genotyped 2717 individuals across all species from multiple locations in multiple years and explicitly test whether genetic diversity is lower for less-successful biological invaders within each species pair. We demonstrate that, for gobies and tunicates, reduced allelic diversity is associated with lower success of invasion. We also found that less-successful invasive species tend to have greater divergence among populations. This suggests that intraspecific hybridization may be acting to convert among-population variation to within-population variation for highly successful invasive species and buffering any loss of diversity. While our findings highlight the species-specific nature of the effects of genetic diversity on invasion success, they do support the use of genetic diversity information in the management of current species invasions and in the risk assessment of potential future invaders.     

Do associations between native and invasive plants provide signals of invasive impacts?

Do invasive plant species act more as “passengers” or drivers of ecological change in native plant communities? Snapshot studies based on correlations at the site scale ignore longer-term dynamics and variation in how particular invaders affect particular native species. We analyzed patterns of co-occurrence between three invading species (Alliaria petiolata, Lonicera x bella, and Rhamnus cathartica) and 70 native plant species in 94 southern Wisconsin forests at two scales to test four hypotheses. Surveys at these sites in the 1950s and again in the 2000s allowed us to assess how initial plant diversity and site conditions affected subsequent patterns of invasion. Sites with more native species in the 1950s experienced fewer invasions of Lonicera and Rhamnus. However, this result may reflect the fact that more fragmented habitat patches supported both fewer species in the 1950s and more invasions. At the site-level, few negative correlations exist between invasive and native species’ abundances. Sites with higher Alliaria densities in the 2000s, however, support fewer native species and lower populations of several declining natives. Rhamnus-invaded sites support lower populations of two increasing species. Association (C-score) analyses detect more associations and more negative associations at the 1 m² scale than at the site scale. Most strong associations between invasive and increasing native species are positive while those with declining natives are often negative. Species restricted to specialized habitats rarely co-occur with invaders. Alliaria has more negative associations at fragmented sites where it is more abundant and invasions may be older. Fine-scale invasive-native associations were stronger, easier to detect, and less consistent than those detectable at the site-level. Thus, screening large numbers of local associations using observational data may allow us to identify particular invasive-native interactions worth further investigation. Although invading plants sometimes act as passive passengers, increasing in tandem with certain native plants in response to disturbed fragmented habitats, they may also contribute to the declines we observe in many native species. Monitoring invasions would allow us to assess whether local associations serve to predict subsequent invasive species impacts.     

Heritable bacterial endosymbionts in native and invasive populations of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis Pallas (1773) (Coleoptera: Coccinellidae) is the well-studied system of invasive insect species. Native and invasive parts of the area of H. axyridis are isolated geographically. We studied the species composition and the distribution of bacterial symbionts Spiroplasma and Rickettsia in seven localities of the native area and six localities of the invasive area of H. axyridis. Rickettsia was detected in H. axyridis populations for the first time. We found that the proportion of beetles infected with Rickettsia in native and invasive populations of H. axyridis is about 0.03. Spiroplasma was found only in native populations of H. axyridis. The proportion of infected individuals with Spiroplasma in native populations of H. axyridis is about 0.08. All studied native populations of H. axyridis are infected with Spiroplasma, while all invasive populations are not. We discuss the possible influence of Spiroplasma and Rickettsia in the formation of invasive populations of H. axyridis.     

Light energy partitioning,photosynthetic efficiency and biomass allocation in invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> and native <Emphasis Type="Italic">Quercus petraea</Emphasis> in relation to light environment,competition and allelopathy

This study addressed whether competition under different light environments was reflected by changes in leaf absorbed light energy partitioning, photosynthetic efficiency, relative growth rate and biomass allocation in invasive and native competitors. Additionally, a potential allelopathic effect of mulching with invasive Prunus serotina leaves on native Quercus petraea growth and photosynthesis was tested. The effect of light environment on leaf absorbed light energy partitioning and photosynthetic characteristics was more pronounced than the effects of interspecific competition and allelopathy. The quantum yield of PSII of invasive P. serotina increased in the presence of a competitor, indicating a higher plasticity in energy partitioning for the invasive over the native Q. petraea, giving it a competitive advantage. The most striking difference between the two study species was the higher crown-level net CO₂ assimilation rates (A_crown) of P. serotina compared with Q. petraea. At the juvenile life stage, higher relative growth rate and higher biomass allocation to foliage allowed P. serotina to absorb and use light energy for photosynthesis more efficiently than Q. petraea. Species-specific strategies of growth, biomass allocation, light energy partitioning and photosynthetic efficiency varied with the light environment and gave an advantage to the invader over its native competitor in competition for light. However, higher biomass allocation to roots in Q. petraea allows for greater belowground competition for water and nutrients as compared to P. serotina. This niche differentiation may compensate for the lower aboveground competitiveness of the native species and explain its ability to co-occur with the invasive competitor in natural forest settings.     

Fast seedling root growth leads to competitive superiority of invasive plants

The competitive superiority of invasive plants plays a key role in the process of plant invasions, enabling invasive plants to overcome the resistance of local plant communities. Fast aboveground growth and high densities lead to the competitive superiority of invasive species in the competition for light. However, little is understood of the role belowground root competition may play in invasion. We conducted an experiment to test the effect of root growth on the performance of an invasive shrub Cassia alata, a naturalized, non-invasive shrub Corchorus capsularis, and a native shrub Desmodium reticulatum. We compared seedling growth of the three species and their competitive ability in situ. The roots of the C. alata seedlings grew much faster than those of C. capsularis and D. reticulatum during the entire growth period although C. alata had shorter shoots than D. reticulatum. Furthermore, C. alata showed an apparent competition advantage compared to the other two species as evidenced by less biomass reduction in intraspecific competition and higher competitive effects in interspecific competition. Our study reveals that fast seedling root growth may be important in explaining the competitive advantages of invasive plants. Future studies should pay more attention to the belowground traits of invasive plants, the trade-off between shoot and root growth, and the role of root competition in affecting the population dynamics of invasive plants and the structures of invaded communities.     

Differences in physiological traits and resistances of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> after herbivory by generalists and specialists

Many researchers have surveyed damages caused by natural enemies of invasive plants in both native and introduced ranges to test the enemy release hypothesis. In this study, we report our findings on the physiological and morphological impacts of a co-evolved specialist insect (Agasicles hygrophila) and two generalist insects (Atractomorpha sinensis and Hymenia recurvalis) in introduced ranges on an invasive plant, Alternanthera philoxeroides, in both field trials and controlled environments. The resistance of A. philoxeroides against the generalists and the specialist was also studied. We obtained consistent results in both the field trials and the controlled treatments: both the generalists and the specialist decreased leaf biomass, photosynthesis, leaf nitrogen content, and total leaf non-structural carbohydrate content in A. philoxeroides. However, the specialist decreased leaf mass, photosynthesis, and leaf nitrogen content more acutely than the generalists. Moreover, A. philoxeroides increased both leaf lignin and cellulose concentrations upon the generalists’ attack but only increased cellulose concentration in response to the specialist. Our results showed that even under the same population density, the co-evolved specialists from original ranges caused more severe morphological and physiological damage to A. philoxeroides than the generalists in introduced ranges. This revealed that invasive plants released some herbivory stress before their co-evolved specialists were introduced, which may contribute to the superior performance of invasive plants in introduced regions.     

Meta-analysis reveals asymmetric reduction in the genetic diversity of introduced populations of exotic insects

Factors promoting the invasion success of introduced populations have been receiving increased attention in studies of biological invasions. Previous reports have indicated that successful invasions may be attributable to reduced genetic diversity in the invasive species. However, there is large variation in the magnitude and direction of the impact of exotic species that have remained unexplained. Here, we present a structured meta-analysis of papers investigating the genetic diversity of native and introduced populations of exotic insects using nuclear microsatellites and mitochondrial DNA sequences. The results indicate that invasion by exotic insects had an overall reducing effect on the genetic diversity of the invading population, with nonzero effect sizes for the number of alleles (NA), observed heterozygosity (Ho), expected heterozygosity (He) and nucleotide diversity (Nd). However, when analyzing different orders (e.g., Lepidoptera, Hemiptera), the effect sizes of NA, Ho and Nd in Lepidoptera were found to bracket zero, as did the effect size of He in Hemiptera. These results suggest an asymmetric reduction in the genetic diversity of introduced populations of exotic insects, indicating diverse mechanisms underlying their successful invasion.     

Substrate preferences of coexisting invasive amphipods, <Emphasis Type="Italic">Dikerogammarus villosus</Emphasis> and <Emphasis Type="Italic">Dikerogammarus haemobaphes</Emphasis>, under field and laboratory conditions

Two Ponto-Caspian amphipods, Dikerogammarus villosus and Dikerogammarus haemobaphes, have expanded their geographical ranges from eastern Europe into Great Britain in recent years. This study represents one of the first examining the distribution and habitat preferences of coexisting populations of D. haemobaphes and D. villosus via field and laboratory experiments in the UK. Field surveys of a recently invaded lowland reservoir in the UK are complimented with ex situ laboratory mesocosm experiments examining the substrate preferences of coexisting populations of D. villosus and D. haemobaphes. Results from the field study indicated that D. haemobaphes dominated the macroinvertebrate community within the reservoir and demonstrated a strong affinity for large cobble and artificial substrates. D. villosus occurred at lower abundances but displayed a strong preference for coarse cobble substrates. A third invasive amphipod, Crangonyx pseudogracilis, was largely confined to sand/silt habitats. Laboratory mesocosm experiments clearly supported the field observations of D. villosus and D. haemobaphes with both species demonstrating a preference for cobble substrates. Results from the study highlight the importance of characterising physical habitat when investigating biological invasions and suggest that habitat availability may influence the extent and speed at which range expansion of new amphipod invaders occurs.     

Taking a detour: invasion of an octocoral into the Tropical Eastern Pacific

The tropical snowflake octocoral Carijoa riisei, which is thought to be native to the Indo-Pacific biogeographical region, has been increasingly reported from the Colombian Tropical Eastern Pacific (TEP) over the past two decades. Massive mortalities of native octocorals, particularly in Pacifigorgia spp. and Muricea spp., were observed due to C. riisei overgrowth. However, the area of origin of TEP C. riisei remains unknown and its potential invasive status has not been addressed yet. We evaluated geographical scenarios for the colonization of the Colombian TEP by conducting phylogeographical analyses based on nuclear and mitochondrial sequences of 306 individual specimens from across the species’ (native/non-native) range and applying hypothesis-specific operational criteria. Additionally, we assessed whether C. riisei has to be considered an invasive species based on the previously proposed ‘unified framework for biological invasions’. Our results showed relatively high genetic differentiation between Colombian TEP populations, on the one side, and Indo-Pacific and Hawaiian populations, on the other side. In contrast, we could not identify genetic differentiation and significant isolation by distance (IBD) between Colombian TEP and Tropical Atlantic populations. C. riisei might have been introduced from the Atlantic into the Colombian TEP, possibly via the Panama Canal. Based on the criteria of the ‘unified framework for biological invasions’, we also conclude that this octocoral constitutes an invasive species. Our study may serve as a basis for establishing strategies to protect native species from one of the very few invasive coral species worldwide.     

Parasitism may alter functional response comparisons: a case study on the killer shrimp <Emphasis Type="Italic">Dikerogammarus villosus</Emphasis> and two non-invasive gammarids

The Ponto-Caspian freshwater amphipod Dikerogammarus villosus has colonized most of the water bodies of continental Europe where it causes strong structural alterations in recipient communities that can lead to changes in ecosystem-level processes, mainly because of a strong predatory behaviour. Most of the D. villosus populations from the invaded range have been found infected with the co-introduced microsporidian parasite Cucumispora dikerogammari, known to decrease the predation rate of its host. Infection might thus mitigate the ecological impact of D. villosus and we wanted to test this assumption using the comparative functional response approach. We compared the relationship between resource use and resource availability (i.e. the functional response, FR) of D. villosus, either with infected individuals or not, to that of two non-invasive gammarids: Gammarus pulex and Echinogammarus berilloni. With infected individuals included, D. villosus displayed a higher FR than the two non-invasive gammarids. Although this effect was not significant, C. dikerogammari infection tended to alter the FR of D. villosus with a slight decrease in attack rate and handling time, resulting in a less steep initial slope and a higher asymptote, respectively. Removing infected D. villosus from the dataset did not affect the FR comparison with G. pulex but suppressed the difference in FR with E. berilloni. Although we cannot exclude the role of sample size reduction in this effect, this suggests that C. dikerogammari infection might increase the predation pressure on local prey populations in case of species replacement between D. villosus and E. berilloni. From a more general perspective, our study illustrates how parasites may alter our capacity to predict invasive species impacts from FR comparisons.     

Local distribution of native and invasive earthworms and effects on a native salamander

North America is home to both native and invasive earthworms acting as ecosystem engineers as they build burrows that can serve as habitat for other species or otherwise alter soil structure, affecting nutrient cycling and other ecosystem processes. Here I determine where and what earthworm species commonly occur in my study area, and compare effects of native and invasive earthworms on the common woodland salamander, Plethodon cinereus, in field surveys and laboratory experiments. The native earthworm Eisenoides carolinensis was the most common earthworm, followed by two invasive species Dendrobaena octaedra and Octolasion tyrtaeum. The presence of O. tyrtaeum was associated with a narrower O-horizon (i.e., organic layer in the soil). Using structural equation modeling to explore direct and indirect pathways of these three most common earthworm species on salamanders, I found O. tyrtaeum occurrence was negatively correlated with nighttime salamander counts, a proxy for total salamander numbers, mediated by negative effects on O-horizon depth and microinvertebrate numbers. In the laboratory, O. tyrtaeum and D. octaedra consumed more leaf litter per gram of earthworm per day than the native E. carolinensis. However, salamanders consumed earthworms and used burrows of all native and invasive species of earthworms similarly. The potential for negative indirect effects of the invasive earthworm O. tyrtaeum on P. cinereus was demonstrated both in the field and laboratory, highlighting that seemingly small differences between native and invasive ecosystem engineers have the potential to significantly alter the effects of these closely related native and invasive organisms.     

The extended phenology of <Emphasis Type="Italic">Spartina</Emphasis> invasion alters a native herbivorous insect’s abundance and diet in a Chinese salt marsh

Plant invasions can alter the trophic interactions of invaded ecosystems because of phenological differences between native and invasive plants that may affect the population dynamics and diets of indigenous arthropod herbivores. This issue, however, has seldom been studied. We here report on how abundance and diet of a local tussock moth (Laelia coenosa) are affected by the invasive plant Spartina alterniflora in a Chinese salt marsh previously dominated by the moth’s native host plant, Phragmites australis. We monitored the population dynamics of L. coenosa from four types of hosts: (1) Phragmites in its monoculture, (2) Spartina in its monoculture, and either (3) Phragmites, or (4) Spartina in Phragmites–Spartina mixtures. Additionally, we tested the diet of L. coenosa from the mixtures with isotope analysis. We found that the larval densities of L. coenosa were similar on Spartina and Phragmites in their respective monocultures and mixtures in summer but were greater on Spartina than on Phragmites in autumn. Stable isotope analysis showed that Spartina was a food resource for L. coenosa. The change in the insect’s population dynamics associated with Spartina invasion might be caused by phenological differences between Spartina and Phragmites in that Spartina has a longer growing season than Phragmites. Our study indicates that the extended phenology of Spartina invasion has altered the abundance and diet of the indigenous herbivorous insect (L. coenosa) previously feeding on native Phragmites. We predict such alternation may increase the consuming pressure to native plants via apparent competition, and thereby may facilitate the further invasion of the exotic plants in the salt marsh.