Invasive Slug Research Articles

An invasive slug exploits an ant-seed dispersal mutualism.

Plant-animal mutualisms, such as seed dispersal, are often vulnerable to disruption by invasive species. Here, we show for the first time how a non-ant invasive species negatively affects seed dispersal by ants. We examined the effects of several animal species that co-occur in a temperate deciduous forest-including native and invasive seed-dispersing ants (Aphaenogaster rudis and Myrmica rubra, respectively), an invasive slug (Arion subfuscus), and native rodents-on a native myrmecochorous plant, Asarum canadense. We experimentally manipulated ant, slug, and rodent access to seed depots and measured seed removal. We also video-recorded depots to determine which other taxa interact with seeds. We found that A. rudis was the main disperser of seeds and that A. subfuscus consumed elaiosomes without dispersing seeds. Rodent visitation was rare, and rodent exclusion had no significant effect on seed or elaiosome removal. We then used data obtained from laboratory and field mesocosm experiments to determine how elaiosome robbing by A. subfuscus affects seed dispersal by A. rudis and M. rubra. We found that elaiosome robbing by slugs reduced seed dispersal by ants, especially in mesocosms with A. rudis, which picks up seeds more slowly than M. rubra. Taken together, our results show that elaiosome robbing by an invasive slug reduces seed dispersal by ants, suggesting that invasive slugs can have profound negative effects on seed dispersal mutualisms.

Description of Phasmarhabditis californica n. sp. and first report of P. papillosa (Nematoda: Rhabditidae) from invasive slugs in the USA

A new species ofPhasmarhabditisand the known speciesP. papillosawere isolated from cadavers of invasive slugs in California.Phasmarhabditis papillosais the type of the genus and has not previously been reported from the Americas. Both species are characterised based on morphology, morphometrics and molecular data. Molecular phylogenies were inferred from concatenated DNA sequence alignments of nearly complete SSU and the D2-D3 domains of the LSU rDNA.Phasmarhabditis californican. sp. is characterised by a robust body, mature egg-laying specimens almost spindle-shaped when relaxed, 1.5 (1.3-1.8) mm long, tapering to a bluntly rounded anterior end with stoma about as long as lip region diam., six inner labial papillae and four outer cephalic papillae, pharynx with rounded to pyriform basal bulb, vulva located halfway along the body, hermaphroditic, didelphic, amphidelphic, and short, conoid tail constricted at one-third its length with prominent phasmids. Not a single male was found among five strains.Phasmarhabditis papillosais gonochoristic and has a longer isthmus, pyriform basal bulb and longer, dome-shaped spicate female tail constricted halfway along its length. Sequence analysis revealed unambiguous autapomorphies forP. papillosaandP. californican. sp. Phylogenetic analyses placed these two species in a monophyletic clade comprisingPhasmarhabditisspecies and other gastropod-parasitic taxa. Morphology, genetic distance, reproductive strategy and nucleotide autapomorphies support the new taxon.

Invasive slug pests and their parasites—temperature responses and potential implications of climate change

It is generally accepted that climate change is likely to disrupt host-parasite relationships, but little consideration has been given to how this may affect agricultural pests. Here, we study the potential influence of climate change on the pest slug Deroceras reticulatum and its nematode parasite Phasmarhabditis hermaphrodita. The European slug D. reticulatum is a major pest of agriculture and has become invasive in many countries throughout the world, including the Americas, Asia and Australasia. P. hermaphrodita strongly inhibits feeding and causes mortality in host slugs. The parasite is widely distributed in the slugs’ native range but is very rare in some countries where the slug is invasive, for example the USA. The influence of temperature on D. reticulatum has been studied previously but the temperature response of P. hermaphrodita is unknown. The current study showed that while host and parasite have similar optimum growth temperatures, the parasite is much less tolerant of temperatures above optimum than the host. As a result, when parasites were present, slug feeding increased significantly above the slug’s optimum of 14 up to 24 °C as the inhibitory effect of the parasites lessened. Conversely, in the absence of parasites, slug feeding significantly decreased over the same temperature range. This finding suggests that given adequate moisture, rising temperatures may increase slug problems in D. reticulatum’s European home range, but possibly reduce problems in invasive territories such as the USA. The relationship between high temperatures and poor parasite performance is supported by studying prevalence of Phasmarhabditis spp. in the UK where a significant negative relationship exists between the mean summer high temperature and parasite prevalence.

Phytophthora survives the digestive system of the invasive slug Arion vulgaris

Decline of European beech (Fagus sylvatica) due to Phytophthora spp. has become an increasing problem in Europe, including Norway, the northernmost distribution of this beech species in the world. Observations of the invasive slug Arion vulgaris in a Phytophthora-infested beech forest in southeastern Norway, led to investigation of A. vulgaris as a potential vector for Phytophthora spp. Thus, the primary goal of this work was to reveal if Phytophthora plurivora and P. cambivora isolated from beech in Norway may survive passing through the digestive system of the slugs. The slugs were allowed to feed on hyphal mats of Phytophthora for 6 days. Faeces from the slugs were examined for the presence of mycelium using both light microscope and Phytophthora specific PCR (polymerase chain reaction). Faeces were also directly dropped or transferred onto Phytophthora selective agar to investigate if the hyphae were viable. The experiments showed that A. vulgaris slugs were capable of ingesting and excreting hyphae of P. cambivora and hyphae and oospores of P. plurivora. Hyphae from both species proved to be viable when faeces were excreted directly on the agar.

Re-description of the slug-parasitic nematode Alloionema appendiculatum Schneider, 1859 (Rhabditida: Alloionematidae)

Alloionema appendiculatum is a common juvenile parasite of many terrestrial molluscs. Its third-stage juveniles (dauers) invade the foot muscle of snails and slugs and develop into fourth-stage juveniles, which then leave the host. Later they mature and reproduce in the soil. A population of A. appendiculatum was isolated from infected individuals of the invasive slug Arion vulgaris (= A. lusitanicus), collected in the city of České Budějovice, Czech Republic, and was designated as the AL strain. This nematode is also able to reproduce on pig kidney in laboratory culture. Our isolate was compared primarily with the description published by Mengert (1953) and in some aspects with the original description published by Schneider (1859). Insufficient morphology, morphometrics, phylogeny and an absence of pictorial material encouraged us to re-describe this frequent slug parasite and to add some new information on its life-cycle. The species is characterised by the absence, in adults, of ridges in the lateral fields. At this life stage the stoma is short, narrow and approximately twice as long as it is broad. Fourth-stage female juveniles produce a mucus-like substance from the phasmids. Males possess six pairs of papillae and a single inconspicuous papilla. The nematode has both parasitic and saprophytic life-cycles. Parasitic adults are bigger than the saprophytes and have a thick, digitate tail, whereas the smaller saprophytic generation has a filiform tail. The species is amphimictic, displaying a higher proportion of females, but males are frequent.

Effect of invasive slug populations (Arion vulgaris) on grass silage. II: Microbiological quality and feed safety

This study aimed to explore how invasive slug populations of Arion vulgaris influenced the microbiological quality and animal feed safety of grass silage, and the efficiency of silage additives and wilting to control the microbiology of slug contaminated crops. The effect of four slug contamination levels, including control, of a grass crop harvested for silage production, was evaluated in laboratory scale. The crop was wilted to two dry matter (DM) levels: low (253g DM/kg) and high (372g DM/kg). Adult slugs were applied to the low DM crop corresponding to 5 (low level), 10 (medium) and 20 (high level) seven-gram sized Arion vulgaris per m2 in an assumed harvested regrowth yield of 2.5ton DM/ha. For the high DM crop, slug weights corresponding to 6 (low level), 12 (medium) and 24 (high level) slugs per m2 were applied. At low DM level, the effect of four additive treatments; control (C), inoculation with Lactobacillus plantarum (LP), a formic, propionic and benzoic acid mixture (ACID) and a chemical additive containing benzoic acid, NaNO2, hexamethylenetetramine and propionic acid (CHEM) were tested. Slugs, slug feces, grass, soil and silages were analyzed for lactic acid bacteria (LAB), Enterobacteriaceae, Listeria monocytogenes, Clostridium tyrobutyricum, molds and yeasts by cultivation methods and Clostridium botulinum type C by real-time PCR analysis.Increasing slug contamination reduced the microbial quality of silages by increasing C. tyrobutyricum levels at both silage DM levels. Only silages without slugs and silages treated with the nitrite containing additive CHEM had non-detectable mean levels of C. tyrobutyricum. Increasing slug contamination increased LAB enumerations in silages. No microbes of risk to human or animal health were detected in anaerobic silages even at the highest slug contamination.

Effect of invasive slug populations (Arion vulgaris) on grass silage: I. Fermentation quality, in-silo losses and aerobic stability

This study aimed to explore how invasive slug populations of Arion vulgaris influence fermentation quality, in-silo losses and aerobic stability of grass silage, and the efficiency of silage additives and wilting to improve the quality of silages from slug contaminated crops. The effect of four levels, including control, of a slug contaminated grass crop, was evaluated in laboratory scale. The crop used was wilted to two dry matter (DM) levels: low (253g DM/kg) and high (372g DM/kg). Adult slugs were applied to the low DM crop corresponding to 5 (low level), 10 (medium) and 20 (high level) 7-g sized A. vulgaris per m2 in an assumed harvested regrowth yield of 2.5ton DM per ha. For the high DM crop, the applied slug levels corresponded to 6 (low level), 12 (medium) and 24 (high level) slugs per m2.At low DM level, the effect of four additive treatments, control (C), inoculation with Lactobacillus plantarum (LP), a formic, propionic and benzoic acid mixture (ACID) and a chemical additive containing benzoic acid, NaNO2, hexamethylenetetramine and propionic acid (CHEM) were tested. Increasing slug contamination gave increasing quality reductions both in silages containing 253 and 372g DM/kg. Compared with untreated silage, LP-treatment did not improve silage fermentation quality of contaminated crops. Treatment with ACID and CHEM, however, considerably improved the quality of heavily contaminated silages. The much higher crude protein concentration in slugs compared to grass crop made slugs a more “difficult-to-ensile” material. Wilting of the harvested crop to 372g DM/kg was not sufficient to control silage fermentation of slug contaminated crop. With contamination levels from 138 to 553g fresh slug weight/kg crop DM, efficient silage additives were able to ensure acceptable fermentation quality of grass silages.

Invasive slug populations (Arion vulgaris) as potential vectors for Clostridium botulinum.

BackgroundNorwegian meadows, including those for silage production, are recently found heavily invaded by the slug Arion vulgaris in exposed areas. As a consequence, large numbers of slugs might contaminate grass silage and cause a possible threat to animal feed quality and safety. It is well known that silage contaminated by mammalian or avian carcasses can lead to severe outbreaks of botulism among livestock. Invertebrates, especially fly-larvae (Diptera), are considered important in the transfer of Clostridium botulinum type C and its toxins among birds in wetlands. C. botulinum form highly resistant spores that could easily be consumed by the slugs during feeding. This study aimed to determine whether Arion vulgaris could hold viable C. botulinum and enrich them, which is essential knowledge for assessing the risk of botulism from slug-contaminated silage. Slug carcasses, slug feces and live slugs were tested by a quantitative real-time PCR (qPCR) method after being fed ≅ 5.8 × 104 CFU C. botulinum type C spores/slug.ResultsLow amounts of C. botulinum were detected by qPCR in six of 21 slug carcasses with an even spread throughout the 17 day long experiment. Declining amounts of C. botulinum were excreted in slug feces up to day four after the inoculated feed was given. C. botulinum was only quantified the first two days in the sampling of live slugs. The viability of C. botulinum was confirmed for all three sample types (slug carcasses, slug feces and live slugs) by visible growth in enrichment media combined with obtaining a higher quantification cycle (Cq) value than from the non-enriched samples.ConclusionsNeither dead nor live invasive Arion vulgaris slugs were shown to enrich Clostridium botulinum containing the neurotoxin type C gene in this study. Slugs excreted viable C. botulinum in their feces up to day four, but in rapidly decreasing numbers. Arion vulgaris appear not to support enrichment of C. botulinum type C.

Orientation behavior of predaceous ground beetle species in response to volatile emissions identified from yellow starthistle damaged by an invasive slug

We investigated indirect defense in the yellow starthistle (Centaurea solstitialis)-grey garden slug (Der- oceras reticulatum)-ground beetle (Pterostichus melana- rius and Scaphinotus interruptus) system. In this host plant/ herbivore/predator system, the ground beetles are the pri- mary predator of D. reticulatum, the dominant herbivore of the highly invasive weed, C. solstitialis. The aim of our study was to examine the behavioral responses of two species of ground beetle to olfactory stimuli emitted from yellow starthistle damaged by D. reticulatum. The beetle P. melanarius showed a significant preference for the odor of damaged yellow starthistle relative to the odor of intact plants, while S. interruptus did not. Volatiles from D. re- ticulatum-damaged yellow starthistle were collected and identified as trans-b-farnesene, germacrene D, bicyclog- ermacrene, and 1,5,9-trimethyl-1,5,9-cyclododecatriene. No quantitative relationship was observed between beetle plant choice or decision time and the level of herbivory. Similarly, there was no relationship between volatile compound relative abundance and level of herbivory, suggesting that our range of leaf damage produces either undetectable semiochemicals or no variation in volatile emission.

Floral herbivory of an invasive slug on a native weed

Effects of floral herbivory (grazing flowers) by slugs on production and quality of seeds of herbaceous plants have been rarely quantified. We studied consequences of grazing by an invasive slug Arion lusitanicus (Mabille) for inflorescences of dandelion (Taraxacum officinale Weber ex Wiggers). In May 2008, a grassy site in Prague-Ruzyně (Czech Republic) was densely populated by dandelion inflorescences. Of the 40% of the flowers grazed by the slug, 70% died before seed dispersal and seed was eaten from 10% of the closed mature inflorescences. Viability of seeds from inflorescences that survived slug grazing was not affected. In addition to seedling predation, grazing of flowers may be an important source of dandelion mortality at sites where this slug is abundant.

Analysis of studies of the effectiveness of a biological method of protection for organic crops

SUMMARY As a result of intensive changes in plant production and of environmental changes in agrocenoses, certain agrophages, such as slugs, have in recent years been causing ever increasing amounts of damage to crops of agricultural, horticultural and orchard plants. As a result of the European Parliament and Council Directive adopted on 13 January 2009, the countries of Europe have been required to implement integrated plant protection. One of the principles of this protection is the implementation of strategies which minimize the use of chemical pesticides. Moreover, in accordance with decisions of the European Commission, many active substances have been or are to be withdrawn from the list of available pesticides. An example is metaldehyde, which is to be withdrawn from use in EU countries in the next few years (Commission Decision C/2008/7637 of 5 December 2008). It therefore becomes necessary to seek environmentally safe substances and to develop non-chemical methods for protecting plants against harmful slugs. An alternative to chemical molluscicides is the parasitic nematode Phasmarhabditis hermaphrodita. Studies have been made of the effectiveness of this biological method in combating Polish populations of the invasive slug species Arion lusitanicus. This was an incomplete multivariate study, i.e. not all variables could be observed for all treatments on all experimental units. Multivariate model of observations has been derived and a method of analysis has been described. The results of these studies provide a basis for the development of environmentally friendly methods of protecting plants against harmful slugs.

Trait-based plant mortality and preference for native versus non-native seedlings by invasive slug and snail herbivores in Hawaii

Non-native herbivores may alter plant communities through their preferential consumption of seedlings of different species. We assessed seedling herbivory by two invasive gastropod species in Hawaii, the giant African snail (Achatina fulica) and the Cuban brown slug (Veronicella cubensis). We hypothesized that six native species would suffer greater gastropod herbivory than four non-native species, and that species with short stature, thin leaves, and lacking physical defenses would suffer the greatest mortality from gastropods. Herbivory was measured during 13-day preference trials using enclosures that each contained four different woody species (two native, two non-native) and were assigned to one of three treatments: giant African snail, Cuban brown slug, or control (no gastropod). Discriminant function analysis was used to predict gastropod-induced seed- ling mortality from a suite of seedling characteristics. Native species did not always experience greater herbivory than non-natives species, and seedling mortality was 0-100 %. Native Pipturus albidus and Clermontia parviflora suffered 100 % mortality from V. cubensis herbivory, and P. albidus, Psychotria hawaiiensis, and Myrsine lessertiana suffered C80 % mortality from A. fulica. Two non-natives (Fraxinus uhdei, Clidemia hirta), and two natives (Metrosideros polymorpha, Diospyros sandwicensis), suffered little damage and no mortality. Non-native Ardisia elliptica suffered 10-30 % gastropod mortality, and non-native Psidium cattleianum mortality was 0-50 %. Leaf thickness best predicted species mortality caused by slugs and snails; some thicker-leaved species suffered most. Invasive snails and slugs threaten some native and non-native seedlings by directly consuming them. Current and future plant community structure in Hawaii may in part reflect the feeding preferences of invasive gastropods.

Phasmarhabditis hermaphrodita (Nematoda: Rhabditidae), a potential biocontrol agent isolated for the first time from invasive slugs in North America

Phasmarhabditis hermaphroditais reported for the first time in North America from cadavers of the invasive slug speciesDeroceras reticulatum,D. laeveandLehmannia valentianacollected from three different locations in California, USA. Four isolates were identified using combined morphology, morphometrics and molecular sequence data for complete internal transcribed spacer (ITS-1, 5.8S, ITS-2), D2-D3 expansion segments of the large subunit (LSU or 28S) and nearly complete small subunit (SSU or 18S) ribosomal DNA. Extremely low sequence variations in the COI gene of the mitochondria were observed among US isolates as well as between US isolates and the two UK sequences. The occurrence ofP. hermaphroditain North America has regulatory implications for potential biological control strategies against non-native gastropod species that are pests in ornamental and agricultural cultivation on this continent. The D2-D3 sequence of the LSU rDNA is new for the species.

Gastropod Seed Dispersal: An Invasive Slug Destroys Far More Seeds in Its Gut than Native Gastropods

Seed dispersal is one of the most important mechanisms shaping biodiversity, and animals are one of the key dispersal vectors. Animal seed dispersal can directly or indirectly be altered by invasive organisms through the establishment of new or the disruption of existing seed dispersal interactions. So far it is known for a few gastropod species that they ingest and defecate viable plant seeds and consequently act as seed dispersers, referred to as gastropodochory. In a multi-species experiment, consisting of five different plant species and four different gastropod species, we tested with a fully crossed design whether gastropodochory is a general mechanism across native gastropod species, and whether it is altered by the invasive alien slug species Arion lusitanicus. Specifically, we hypothesized that a) native gastropod species consume the seeds from all tested plant species in equal numbers (have no preference), b) the voracious invasive alien slug A. lusitanicus – similarly to its herbivore behaviour – consumes a higher amount of seeds than native gastropods, and that c) seed viability is equal among different gastropod species after gut passage. As expected all tested gastropod species consumed all tested plant species. Against our expectation there was a difference in the amount of consumed seeds, with the largest and native mollusk Helix pomatia consuming most seeds, followed by the invasive slug and the other gastropods. Seed damage and germination rates did not differ after gut passage through different native species, but seed damage was significantly higher after gut passage through the invasive slug A. lusitanicus, and their germination rates were significantly reduced.

Locomotor activity of two sympatric slugs: implications for the invasion success of terrestrial invertebrates

A central focus of invasion biology is to identify the traits that predict which introduced species will become invasive. Behavioral traits related to locomotor activity most likely play a pivotal role in determining a species' invasion success but have rarely been studied, particularly in terrestrial invertebrates. Here, we experimentally investigated the small‐scale locomotor activity of two slug species with divergent invasion success in Europe, the highly invasive slug, Arion lusitanicus, and the closely related, non‐invasive and native slug, Arion rufus. To do so, we used a multi‐state capture‐mark‐recapture approach, and hypothesized that the invasive slug has a higher moving rate (keeps on moving) and leaving rate (leaves more frequently known places). A total of 221 invasive and 241 non‐invasive slugs were individually marked using magnetic transponders and released in three study sites differing in habitat type. The slugs were recaptured using shelter traps, and moving and leaving rates were estimated. Both rates were significantly higher for the invasive slug, demonstrating a higher locomotor activity which might partly explain its invasion success. Our results provide evidence for the recently suggested idea that locomotor activity might be an important trait underlying animal invasions using for the first time terrestrial invertebrates.

Herbivory of an invasive slug in a model grassland community can be affected by earthworms and mycorrhizal fungi.

Invasion of non-native species is among the top threats for the biodiversity and functioning of native and agricultural ecosystems worldwide. We investigated whether the herbivory of the slug Arion vulgaris (formerly Arion lusitanicus; Gastropoda), that is listed among the 100 worst alien species in Europe, is affected by soil organisms commonly present in terrestrial ecosystems (i.e. earthworms—Annelida: Lumbricidae and arbuscular mycorrhizal fungi—AMF, Glomerales). We hypothesized that slug herbivory would be affected by soil organisms via altered plant nutrient availability and plant quality. In a greenhouse experiment, we created a simple plant community consisting of a grass, a forb, and a legume species and inoculated these systems with either two earthworm species and/or four AMF taxa. Slugs were introduced after plants were established. Earthworms significantly reduced total slug herbivory in AMF-inoculated plant communities (P = 0.013). Across plant species, earthworms increased leaf total N and secondary metabolites, AMF decreased leaf thickness. Mycorrhizae induced a shift in slug feeding preference from non-legumes to legumes; the grass was generally avoided by slugs. AMF effects on legume herbivory can partly be explained by the AMF-induced increase in total N and decrease in C/N ratio; earthworm effects are less clear as no worm-induced alterations of legume plant chemistry were observed. The presence of earthworms increased average AMF colonization of plant roots by 140 % (P < 0.001). Total shoot mass was significantly increased by AMF (P < 0.001). These data suggest that the feeding behavior of this invasive slug is altered by a belowground control of plant chemical quality and community structure.

Distribution and spread of the invasive slug Arion vulgaris Moquin-Tandon in Norway

The present distribution of the invasive slug Arion vulgaris Moquin-Tandon was studied in Norway. This important pest species has spread to many parts of Europe during the last decades, inflicting damage to agriculture and domestic gardens. It was first recorded in Norway in 1988, and has since spread to many parts of the country and is now recorded in 192 municipalities. We surveyed the current distribution by sampling and gathering species records in cooperation with garden societies and local authorities. Based on these records, we present distributional data as well as relative predictions of future distributions based on geoclimatic parameters. Currently, A. vulgaris covers most of coastal southern Norway while it shows a patchy distribution in northern Norway, recorded as far north as Finnsnes in Troms County.

Herbivory of an invasive slug is affected by earthworms and the composition of plant communities.

BackgroundBiodiversity loss and species invasions are among the most important human-induced global changes. Moreover, these two processes are interlinked as ecosystem invasibility is considered to increase with decreasing biodiversity. In temperate grasslands, earthworms serve as important ecosystem engineers making up the majority of soil faunal biomass. Herbivore behaviour has been shown to be affected by earthworms, however it is unclear whether these effects differ with the composition of plant communities. To test this we conducted a mesocosm experiment where we added earthworms (Annelida: Lumbricidae) to planted grassland communities with different plant species composition (3 vs. 12 plant spp.). Plant communities had equal plant densities and ratios of the functional groups grasses, non-leguminous forbs and legumes. Later, Arion vulgaris slugs (formerly known as A. lusitanicus; Gastropoda: Arionidae) were added and allowed to freely choose among the available plant species. This slug species is listed among the 100 worst alien species in Europe. We hypothesized that (i) the food choice of slugs would be altered by earthworms’ specific effects on the growth and nutrient content of plant species, (ii) slug herbivory will be less affected by earthworms in plant communities containing more plant species than in those with fewer plant species because of a more readily utilization of plant resources making the impacts of earthworms less pronounced.ResultsSlug herbivory was significantly affected by both earthworms and plant species composition. Slugs damaged 60% less leaves when earthworms were present, regardless of the species composition of the plant communities. Percent leaf area consumed by slugs was 40% lower in communities containing 12 plant species; in communities containing only three species earthworms increased slug leaf area consumption. Grasses were generally avoided by slugs. Leaf length and number of tillers was increased in mesocosms containing more plant species but little influenced by earthworms. Overall shoot biomass was decreased, root biomass increased in plant communities with more plant species. Earthworms decreased total shoot biomass in mesocosms with more plant species but did not affect biomass production of individual functional groups. Plant nitrogen concentrations across three focus species were 18% higher when earthworms were present; composition of plant communities did not affect plant quality.ConclusionsGiven the important role that both herbivores and earthworms play in structuring plant communities the implications of belowground-aboveground linkages should more broadly be considered when investigating global change effects on ecosystems.

Predation by carabid beetles on the invasive slugArion vulgarisin an agricultural semi-field experiment

Arion vulgaris Moquin-Tandon 1855 is one of the most important invasive species in Europe, affecting both biodiversity and agriculture. The species is spreading in many parts of Europe, inflicting severe damage to horticultural plants and cultivated crops partly due to a lack of satisfactory and effective management solutions. Molluscicides have traditionally been used to manage slug densities, although the effects are variable and some have severe side-effects on other biota. Thus, there is a need to explore potential alternatives such as biological control. The nematode Phasmarhabditis hermaphrodita is the only biological agent that has been applied commercially so far. However, other biological control agents such as carabid beetles have also been found to be promising. In addition, some carabid species have been shown to feed on A. vulgaris in the field as well as in the laboratory. Two species in particular have been found to be important predators of A. vulgaris, and these species are also common in agricultural environments: Pterostichus melanarius and Carabus nemoralis. This study is the first to use semi-field experiments in a strawberry field, manipulating densities, to investigate how P. melanarius and C. nemoralis affect densities of A. vulgaris eggs and juveniles, respectively. Gut contents of C. nemoralis were analysed using multiplex PCR methods to detect DNA of juvenile slugs. Results show that both P. melanarius and C. nemoralis significantly affect densities of slug eggs and juvenile slugs under semi-field conditions and that C. nemoralis seems to prefer slugs smaller than one gram. Carabus nemoralis seems to be especially promising in reducing densities of A. vulgaris, and future studies should investigate the potential of using this species as a biological control agent.

Jack-of-all-trades: phenotypic plasticity facilitates the invasion of an alien slug species

Invasive alien species might benefit from phenotypic plasticity by being able to (i) maintain fitness in stressful environments ('robust'), (ii) increase fitness in favourable environments ('opportunistic'), or (iii) combine both abilities ('robust and opportunistic'). Here, we applied this framework, for the first time, to an animal, the invasive slug, Arion lusitanicus, and tested (i) whether it has a more adaptive phenotypic plasticity compared with a congeneric native slug, Arion fuscus, and (ii) whether it is robust, opportunistic or both. During one year, we exposed specimens of both species to a range of temperatures along an altitudinal gradient (700-2400 m a.s.l.) and to high and low food levels, and we compared the responsiveness of two fitness traits: survival and egg production. During summer, the invasive species had a more adaptive phenotypic plasticity, and at high temperatures and low food levels, it survived better and produced more eggs than A. fuscus, representing the robust phenotype. During winter, A. lusitanicus displayed a less adaptive phenotype than A. fuscus. We show that the framework developed for plants is also very useful for a better mechanistic understanding of animal invasions. Warmer summers and milder winters might lead to an expansion of this invasive species to higher altitudes and enhance its spread in the lowlands, supporting the concern that global climate change will increase biological invasions.