Crop monitoring to assess the risk of slug damage to winter wheat in the United Kingdom

The percentage of winter wheat seeds killed by slugs was measured, together with various risk factors, at 93 sites throughout the UK from 1987 to 1990. Estimates of slug populations at each site were obtained from traps baited with methiocarb pellets. The peak number of slugs trapped during the period from July until just before cultivation accounted for 26% of the variability in seed damage, more than any other single risk factor. A combined function of the number of slugs trapped at drilling and the percentage of fine soil aggregates in the seed-bed accounted for 21% of the variability in seed damage. Both (i) the peak number of slugs trapped before drilling, and (ii) the combined function of number of slugs trapped at drilling and the percentage of fine soil aggregates in the seed-bed, have potential for identifying fields with a negligible risk of slug damage to wheat seeds. However, further research is needed to improve the accuracy of forecasting slug damage. In particular, other forms of trap should be tested for their potential use in providing more accurate estimates of slug numbers and biomass at drilling, and techniques of assessing seed-bed conditions should be improved. The percentage of seedlings grazed by slugs was correlated with the numbers of slugs trapped before drilling, at drilling and after crop emergence.     

Effect of seed depth on slug damage to winter wheat

In a field experiment drilled at two depths on three dates in autumn 1988, with or without methiocarb pellets broadcast on the soil surface immediately after drilling, 26% of seeds of winter wheat sown at c. 20 mm depth were killed by slugs compared with only 9% of seeds sown at c. 40 mm. The protection from slug damage provided by this additional 20 mm of depth was comparable with that provided by methiocarb pellets. The effects of seed depth and pellet application did not interact and were consistent on all drilling dates. Thus, fewest seeds and seedlings were killed where methiocarb pellets were broadcast on a seed-bed with seeds sown at 40 mm depth. Intermediate damage was recorded where seeds were sown at 40 mm depth without pellets, or where pellets were broadcast on seeds sown at 20 mm depth. Most seeds and seedlings were killed where seeds were sown at 20 mm depth without pellets. Sublethal damage to seedlings was not affected by sowing depth but was reduced where pellets were broadcast immediately after sowing.     

Biological control of slugs in winter wheat using the rhabditid nematode Phasmarhabditis hermaphrodita

A field experiment on winter wheat in autumn 1991 investigated the effect of the rhabditid nematode, Phasmarhabditis hermaphrodita, applied to soil at five dose rates (10⁸ - 10¹⁰ infective larvae ha^-1) immediately after seed sowing, on slug populations and damage to seeds and seedlings. The nematode was compared with methiocarb pellets broadcast at recommended field rate immediately after drilling and no molluscicide treatment. Slug damage to wheat seeds and seedlings was assessed 6 and 13 wk after drilling. Seedling survival increased and slug grazing damage to seedlings declined linearly with increasing log nematode dose. These two measures of slug damage were combined to give an index of undamaged plant equivalents, which also increased linearly with increasing log nematode dose. ANOVA showed that, after 6 wk, there were significantly more undamaged plant equivalents on plots treated with the two highest nematode doses (3 × 10⁹ and 1 × 10¹⁰ ha^-1) than on untreated plots, but the number of undamaged plant equivalents on methiocarb-treated plots was not significantly greater than that on untreated plots. Slug populations were assessed by refuge trapping and soil sampling. Deroceras reticulatum was the commonest of several species of slugs recorded. During the first 4 wk after sowing, significantly more slugs were found under refuge traps on plots treated with certain doses of P. hermaphrodita than under traps on untreated plots and more showed signs of nematode infection than expected from the prevalence of infection in slugs from soil samples, suggesting that the presence of P. hermaphrodita altered slug behaviour. Application of P. hermaphrodita had no significant impact on numbers or biomass of slugs in soil during a 27 wk period after treatment, except after 5 wk when slug numbers were inversely related to log nematode dose. However, by this time, numbers in soil samples from untreated plots had declined to levels similar to those in plots treated with the highest dose of nematodes. During the first 5 wk after treatment, c. 20% of slugs in soil samples from untreated plots showed symptoms of nematode infection. It is suggested that this represented the background level of infection in the experimental field rather than spread of infection from treated plots. The apparent lack of impact of P. hermaphrodita on slug numbers and biomass in soil suggests that its efficacy in protecting wheat from slug damage was through inhibition of feeding by infected slugs.     

Biocontrol of Slugs in Protected Lettuce Using the Rhabditid Nematode Phasmarhabditis hermaphrodita

In two experiments, the rhabditid nematode Phasmarhabditis hermaphrodita, a parasite of slugs, was cultured in vitro and applied as a drench to soil at four dose rates (3 108, 1 109, 3 109 and 1 1010 ha-1) 1 or 4 days before planting lettuce seedlings in a polythene tunnel. The effects of the four nematode doses on slug damage during the first 3 weeks after planting and on the numbers of slugs found within and below lettuce plants at harvest were measured. Results were then compared with untreated plots and with plots where methiocarb pellets were applied at the recommended field rate. In the first experiment, methiocarb pellets significantly reduced the percentage of plants damaged by slugs, but the nematode did not. In the second experiment, methiocarb pellets and the second highest dose of nematodes significantly reduced the percentage of plants damaged by slugs. The different effect of the nematode in the two experiments may have resulted from differences in the timing of nematode application and/or differences in the pattern of slug damage between experiments. At the end of the first experiment, the highest two doses of nematodes and methiocarb pellets had significantly reduced the number of slugs found within lettuce plants at harvest and on the soil surface below the plants. At the end of the second experiment, analysis of variance showed no significant effects of any treatment on slug numbers or biomass, but regression analysis showed significant negative relationships between nematode dose and total slug numbers, numbers of Arion ater agg. and biomass of Deroceras reticulatum. In both experiments, increasing nematode dose significantly reduced the numbers of slugs found contaminating the harvested lettuce. At the end of the second experiment, the mean weight of individuals of A. ater agg. increased with rising nematode dose.     

Seedling herbivory by slugs in a willow hybrid system: developmental changes in damage, chemical defense, and plant performance

We evaluated feeding preference and damage by the slug, Arion subfuscus, on seedlings of two willow species, Salix sericea and S. eriocephala, and their F₁ interspecific hybrids. Trays of seedlings were placed in the field and excised leaves were presented to slugs in choice tests. Slugs preferred feeding on and caused the most damage to S. eriocephala seedlings. S. sericea seedlings were least preferred and least damaged. F₁ hybrid seedlings were intermediate in preference and damage. Slug preference of and damage to these seedlings decreased over time, suggesting developmental changes in resistance. Seedlings were sampled for phenolic glycoside and tannin chemistry weekly to coincide with the field and laboratory experiments. Concentrations of phenolic glycosides and tannins increased linearly with seedling age, coincident with changes in slug preference and damage, indicating a developmental change in defense. Slug deterrence was not detected at low concentrations of salicortin when painted on leaves or discs, but both salicortin and condensed tannins deterred slug feeding at concentrations between 50 and 100 mg/g, levels found in adult willows. Seedling performance was related to damage inflicted by slugs. Due to lower levels of damage when exposed to slugs in the field, S. sericea plants had significantly greater biomass than S. eriocephala plants. Biomass of F₁ hybrids was equal to S. sericea when damaged. However, undamaged S. eriocephala and F₁ hybrid plants had the greatest biomass. Because F₁ hybrid seedlings performed as well as the most fit parent in all cases, slugs could be an important selective factor favoring introgression of defensive traits between these willow species.     

A method to study slug predation on seedlings in the field

Dandelion is a common Asteraceae species that populates disturbed sites and gaps within swards where it becomes an important competitor of grasses. The natural control against dandelion includes seedling predation, with slugs, particularly Arion lusitanicus, being the most important in the Czech Republic. However, the study of slug seedling consumption is difficult because naturally established seedlings are not always available. Therefore, we developed a method of exposing laboratory‐grown seedlings as bait for slug predation. Dandelion seeds were sown in plastic cups containing a bottom layer of moist substrate. The emerged seedlings were thinned to 20 and displayed in an open area. During 2008–2010, the seedling baits were placed at 15 sites at 1 month intervals throughout the dandelion vegetative season, in parallel with plasticine baits that monitored slug feeding activity. For each 1 month interval, seedling survival was observed for a period of 8 days, and the estimated time to death was calculated; the percentage of surviving seedlings was then recorded. This method of seedling presentation demonstrated that local and temporal variation in seedling survival is correlated with slug feeding activity. The advantage of this technique is that the seedlings in baits may be presented at any place and time, as required by the experimental design; however, we found that the estimated time to seedling death was shorter for the exposed baits than for the naturally established seedlings. The method is suitable for the study of plant species other than dandelion, and also for aims other than the study of spatiotemporal trends in seedling consumption by slugs.     

Spatial sampling to detect slug abundance in an arable field

Overuse of molluscicides by farmers in arable systems can lead to environmental and product contamination. Here we assess a simple and inexpensive surface trapping method for monitoring populations of slugs (Deroceras reticulatum and Arion intermedius). This method was biased against small slugs, and against A. intermedius, when compared to direct soil sampling. Regression was used to model the relationship between the results of surface trapping and soil sampling methods. Spatial Analysis by Distance IndicEs (SADIE) algorithms were used to describe the spatial relationships between the two sets of samples. Using both traditional statistical methods and spatial statistics, the spatial information collected from surface traps was sufficient to identify patches and gaps in slug numbers and possibly to allow the spot application of slug control, and thus provide land managers who experience slug damage with a way of reducing molluscicides use, whilst maintaining slug control. Further improvements and applications of the model are discussed.     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.     

Damage to oilseed rape seedlings by the field slug Deroceras reticulatum in relation to glucosinolate concentration

Seedlings of nine commercial cultivars of oilseed rape were exposed to the field slug Deroceras reticulatum immediately after sowing in compost in trays. There was a small reduction in seedling numbers in the presence of slugs which was not related to glucosinolate concentration in seeds or seedlings. However, the number and leaf-area of seedlings with damage symptoms were strongly and inversely related to the total concentration of glucosinolates in seeds and one wk-old seedlings. The presence of barley seedlings as alternative food did not significantly affect this relationship. The glucosinolate concentration of seeds was closely correlated with that of wk-old seedlings. Analysis of individual glucosinolates in four cultivars spanning the range of concentrations found, showed that the concentration of most components declined as total glucosinolate concentration decreased. However, 2-phenyl ethyl-glucosinolate (gluconasturtiin) concentration tended to increase in seeds and 3-indolyl methyl-glucosinolate (glucobrassicin) increased in seedlings as total glucosinolate concentration decreased. Damage by slugs was inversely related to the concentration of those glucosinolates which decreased and was positively correlated with the two compounds which increased as total glucosinolate concentration decreased. The results support the hypothesis that glucosinolates in brassicas protect them from polyphagous herbivores, and, in particular, that an important function of glucosinolates in rape seeds is to protect seedlings from slugs. As glucosinolate concentrations of oilseed rape cultivars continue to decline, so the risk of slug damage to seedlings may well increase.     

SLUG DAMAGE AND NUMBER OF SLUGS (GASTROPODA: PULMONATA) IN WINTER WHEAT IN FIELDS WITH SOWN WILDFLOWER STRIPS

Previous observations have shown that slugs found favourableconditions for reproduction in wildflower strips and that theycaused high damage to oilseed rape adjacent to the strips. Inthe current study slug numbers and damage were estimated inwildflower strips and at different distances from the stripsinto winter wheat crops, from 1994 to 1996. Slug damage wasestimated using an index of defoliation and slug numbers andactivity were measured using wheat bran traps. Investigationsbegan when winter wheat seedlings emerged and lasted for fiveweeks. Slug damage was never severe in any of the fields studied.In most fields, slug damage was higher close to the wildflowerstrips than at greater distances from the strips. The decliningslug damage with increasing distances from the wildflowers stripswas related to the distribution pattern of juvenile Arion lusitanicus,suggesting that this species was responsible for the higherdamage near the strips. Other slug species (Deroceras reticulatum,Arion fasciatus, Deroceras laeve) were more or less evenly distributedover the field or were almost entirely confined to the wildflowerstrips. Results showed that winter wheat was not at risk fromslug feeding in spite of the establishment of wildflower stripsand that therefore the application of molluscicide along thestrips is unnecessary. (Received 23 June 1997; accepted 17 October 1997)     

SOME OBSERVATIONS ON DAMAGE TO POTATOES BY SLUGS

The garden slug, Arion hortensis Fér., and the keeled slugs, Milax spp., are the slugs mainly responsible for primary damage to potato tubers in the south-west of England. A variety trial clearly demonstrated the existence of significant differences in susceptibility to slug attack between the three varieties tested. Arran Banner was the most damaged variety, Arran Peak the least, while Majestic occupied an intermediate position. Varietal differences in the amount and time of damage to tubers were associated with differences in the dates of maturity, and little damage occurred prior to the maturing of the tuber. The results confirm the need for prompt lifting of the potato crop on maturity, in order to avoid damage by slugs.     

The rhabditid nematode Phasmarhabditis hermaphrodita as a potential biological control agent for slugs

A nematode, Phasmarhabditis hermaphrodita, known to be associated with slugs but not previously thought to be parasitic, was shown to be a parasite capable of killing the pest slug Deroceras reticulatum. The parasite infects slugs in the area beneath the mantle surrounding the shell, causing a disease with characteristic symptoms, particularly swelling of the mantle. Infection leads to death of the slug, usually between seven and 21 days afterwards. The nematode then spreads and multiplies in the cadaver. In an experiment where individual D. reticulatum were exposed to different numbers of P. hermaphrodita, a significant positive relationship was found between nematode dose and slug mortality. In two experiments on host range, the nematode was found to infect and kill all pest slug species tested: Deroceras caruanae, Arion distinctus, Arion silvaticus, Arion intermedius, Arion ater, Tandonia sowerbyi and T. budapestensis, in addition to D. reticulatum.     

Slugs, willow seedlings and nutrient fertilization: intrinsic vigor inversely affects palatability

This study evaluates how preference by a generalist slug herbivore Arion subfuscus changes inversely with seedling size across three levels of fertilization for three full‐sib families of willow seedlings. We analyzed seedlings for condensed tannin and protein concentration, and related these data to changes in palatability. In preference tests over time, leaf discs from more fertilized seedlings experienced an extended window of vulnerability compared to discs from less fertilized seedlings, which were also more tannin‐rich. In a whole seedling selection study, slugs readily attacked smaller seedlings (<5 cm) but rarely attacked taller seedlings (>10 cm). However, a general difference in risk of damage close to 50% existed when comparing shorter and taller individuals within each family and level of fertilizer. The decrease in palatability with height of the seedlings was positively correlated with an increase in condensed tannin concentration. We found no effect of seedling size on protein concentration. Akaiki index criterion model comparisons suggested that only main effects were important for explaining seedling choice by slugs as well as the ratio between proteins and condensed tannins. Seedling size, had the largest effect, followed by fertilizer level and family. Surprisingly, seedling size and fertilizer treatment had opposite effects on palatability to slugs. Size decreased probability of damage, whereas fertilization extended the window of susceptibility. Because the seedlings were even‐aged, differences in size are interpreted as differences in growth rate or vigor. The positive phenotypic correlation found between size and tannin production in the less preferred willow seedlings confirms that several plant defense traits may be selected for simultaneously, because fast growth may allow an early development of plant defenses. We discuss these results in the light of plant‐defense theories that predict a negative correlation between the allocation to growth and the production of secondary defense compounds.     

SLUG DAMAGE AND NUMBERS OF SLUGS IN OILSEED RAPE BORDERING ON GRASS STRIPS

In 1995, slug damage and numbers of slugs were estimated intwo grass strips and adjacent rape fields. Investigations beganas soon as rape seedlings emerged and lasted for five weeks.Slug damage to rape plants 1 m from the grass strips was significantlyhigher than at greater distances from the strips. Derocerasreticulatum was the most abundant slug species recorded in bothgrass strips and adjacent rape fields. Arion lusitanicus andArion fasciatus were much less abundant than D. reticulatum.In one field, D. reticulatum declined steadily with increasingdistance from the grass strips and therefore appeared to havecaused the majority of severe damage to rape plants close tothe strips. This finding was surprising because until now severeslug damage in oilseed rape beside semi-natural habitats hasbeen observed only where A. lusitanicus was abundant. (Received 12 November 1997; accepted 26 January 1998)     

Daily activity rhythm and use of shelter in the slugs Deroceras reticulation and Arion distinctus under laboratory conditions

Daily activity of slugs, Deroceras reticulatum and Arion distinctus, was studied in the laboratory in relation to the presence of shelters so as to define optimal conditions for using traps to forecast crop damage. Under constant temperature and humidity, activity of slugs began with the lights off. Its arrest occurred after a period of activity with the lights on. Slugs rested most frequently under a shelter on bare earth and rarely used the same shelter for more than two consecutive days. When food was placed under the shelter, the number of slugs found under the shelter was more constant throughout the 24 h period, but lower during diurnal rest. The presence of 4% methiocarb pellets under the shelter led to an increased occupation by slugs, due to the poisoning of some of them. Acts of aggression were more numerous in A. distinctus than in D. reticulatum. In D. reticulatum the smallest slug was the least aggressive and suffered more attacks from the other slugs. In A. distinctus the numbers of attacks suffered or provoked varied little with the size of the slug, but the smallest slug showed less interactions with other slugs.     

Tracking predator density dependence and subterranean predation by carabid larvae on slugs using monoclonal antibodies

1. Subterranean carabid larvae are more numerous than surface‐active adults, yet very little is known about their ecological significance, dietary preferences or ability to regulate populations of prey species, particularly pests. Part of the reason for this is that predator–prey interactions beneath the soil are almost impossible to observe. 2. Extensive field studies have shown that adult Pterostichus melanarius (Illiger) can affect the temporal and spatial dynamics of their slug prey. However, if larvae too are feeding on slugs, this could radically affect overall predator–prey dynamics. 3. We tested the hypotheses that P. melanarius larvae would kill and consume two slug species, Deroceras reticulatum Müller and Arion intermedius Normand, under laboratory and semi‐field conditions, and that there would be no significant difference in rates of predation on these slug species. 4. A new monoclonal antibody was developed that was capable of detecting the presence of slug proteins in the guts of P. melanarius larvae. 5. Pterostichus melanarius larvae killed both A. intermedius and D. reticulatum in the laboratory, feeding to a greater extent, and growing more rapidly, on the latter. The larvae were equally effective at reducing numbers of both slug species in a crop of wheat grown in semi‐field mini plots, but predation was affected by density‐dependent intra‐specific competition amongst the beetle larvae. 6. Future modelling of the dynamic interactions between carabids and slugs will need to take into account predation by larvae.     

Molluscicidal and slug-repellent properties of anaerobically digested organic matter

Slug problems in arable crops and vegetables have increased drastically during the past few decades. Observations on slug damage to oilseed rape suggested that fresh, anaerobically digested organic material from a biogas production plant is molluscicidal. To find out whether digested matter can be used for the control of agricultural pest slugs, a series of exeriments were carried out. The laboratory experiments demonstrated strong mollusc repellent and molluscicidal effects of digested organic matter against the three most important pest slugs of Switzerland, Arion lusitanicus, A. distinctus and Deroceras reticulatum. The effects were restricted to fresh digested matter and were rapidly lost when the material was stored, and also after application in the field. In the field experiment, fresh digested matter greatly reduced slug damage to lettuce in comparison with the untreated plots. At present, the chemical nature of the molluscicidal compound(s) in digested matter is unknown, but environmental pollutants such as heavy metals can be ruled out. Current research aims at a new formulation which is easier to apply and has a longer-lasting molluscicidal or slug-repellent effect, and at optimising the dosage and number of applications.     

Size-specific susceptibility of the pest slugs Deroceras reticulatum and Arion lusitanicus to the nematode biocontrol agent Phasmarhabditis hermaphrodita

The nematode Phasmarhabditis hermaphrodita is a commercially available biocontrol agent against slugs. This product is especially interesting for use in organic farming, where products containing metaldehyde or carbamates cannot be used for controlling pest slugs. We investigated the potential of P. hermaphrodita for the control of the pest slugs Deroceras reticulatum and Arion lusitanicus. These two species are the most harmful slug pests in Switzerland. At different times of the year, we collected slug specimens of different weight and assessed their susceptibility to P. hermaphrodita in the laboratory. Batches of five slugs were subjected to five different doses of nematodes plus an untreated control and replicated three times. During six weeks, feeding and survival of the slugs were recorded. D. reticulatum was strongly affected by increasing nematode doses, irrespective of the slugs' body weight. In small specimens of A. lusitanicus, feeding and survival were strongly affected by the nematodes, while larger specimens remained almost unaffected. Because A. lusitanicus has an asynchronous development in Switzerland, it seems difficult to control the entire population with a single nematode application. To what extent nematodes will be used in practice for slug control depends on their effectivity against the pest slugs of major importance, on the longevity of the molluscicidal effect and on the price of nematodes.     

Feeding deterrent effect of carvone, a compound from caraway seeds, on the slug Arion lusitanicus

The feeding deterrent effect of carvone on the slug Arion lusitanicus was investigated. Carvone, a natural compound from caraway seeds, was incorporated into mulch to reduce its inherent volatility. In a laboratory choice experiment, boxes were filled on one side with carvone‐treated mulch and on the other side with untreated mulch. At carvone concentrations ranging from 0.03–0.75 ml litre^?1 mulch, slugs ate significantly more lettuce on the untreated side. In a laboratory based no‐choice experiment, carvone concentrations of 0.25 and 0.75 ml litre^?1 mulch significantly reduced slug feeding in comparison with the untreated control. Moreover at the highest concentration of carvone (0.75 ml litre^?1 mulch) 50% mortality was recorded over a period of 5 days, indicating a clear molluscicidal effect. Due to its volatility carvone did not decrease plant defoliation by A. lusitanicus when applied directly onto lettuce. Subsequent field evaluation showed 0.75 ml litre^?1 mulch to partially reduce slug feeding damage. However, this effect was not sufficient to significantly increase lettuce yield. The incorporation of a higher carvone concentration into mulch is still to be tested to confirm whether carvone‐treated mulch can be recommended as an effective alternative approach to chemical slug control.     

Mini‐plot field experiments using the rhabditid nematode Phasmarhabditis hermaphrodite for biocontrol of slugs

The nematode Phasmarhabditis hermaphrodita, a parasite of slugs, was cultured in vitro and applied as a drench in two outdoor mini‐plot field experiments to test the capacity of the nematode to protect Chinese cabbage seedlings and wheat seeds from damage by the field slug Deroceras reticulatum. The first experiment compared a single dose of nematodes (2 X 10¹⁰ ha^‐1) with methiocarb pellets added at the recommended field rate (5.5 kg.product ha^‐1 ) and untreated plots. Plots treated with either nematodes or methiocarb pellets had significantly less slug damage than untreated plots and, from the third week onwards, there was significantly less slug damage on plots treated with nematodes than on methiocarb‐treated plots. At the end of the experiment, 6 weeks after treatment, both slug numbers and biomass were significantly higher in untreated plots than in either methiocarb‐treated or nematode‐treated plots. In the second experiment, six nematode doses ranging from 1 X 10 ⁸ to 2 X 10¹⁰ ha^‐1 were compared with a standard rate application of methiocarb pellets and untreated plots. Plant protection improved with increasing nematode dose between 1 X10⁸ and 8 X 10⁸ha^‐1, but showed little or no further improvement at higher doses. Plant protection similar to that given by methiocarb pellets was provided by nematode doses of 8 X 10⁸ ha^‐1 and above.