AUTOTOMY IN ARIONID AND LIMACID SLUGS

The occurrence of autotomy was investigated in five terrestrialslug species. Half of the Dcroceras reticulatum and 80 per centof the Limax tenellus slugs which were stimulated mechanicallyin the tail, autotomized the tail. Anon fasciatus, Anon subfuscusand Limax cinereoniger did not show any signs of being ableto autotomize. The autotomized tail of D. rcticulatum and L.tenellus was small compared to the total body weight and alsoto the weight of many predaceous beetles. D. reticulatum andL. tenellus escaped in a few seconds after the beginning ofmechanical stimulation; the reaction time was significantlyshorter than that used by species which did not autotomize theirtail. Comparison between A. fasciatus and D. reticulatum alsoshowed that D. reticulatum sought for refuges significantlymore effectively than A. fasciatus. The exposure of D. reticulatumto the beetle Ptero-stichus mger indicated that autotomy canstop a beetle long enough to let the slug escape. It seems thatautotomy has evolved more frequently in those species whichrely to a great extent on escaping and hiding when defendingthemselves, than in species whose physiology and/or environmentdo not enable them to hide quickly after an attack. (Received 18 February 1993; accepted 19 March 1993)     

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)     

A COMPARISON OF THE LIFE CYCLES OF THE SLUGS DEROCERAS RETICULATUM (MULLER) AND ARION INTERMEDIUS NORMAND ON PERMANENT PASTURE

Life cycles of the slugs Deroceras reticulatum and Arion intermediuswere compared over a four-year period on permanent pasture nearOvingham, Northumberland. Both species were semelparous, theirlife cycles taking about one year. D. reticulatum, unlike A.intermedius and other species, had two overlapping generationsthat laid their eggs in late spring and autumn respectivelyand also had one instead of two immature phases in its lifecycle. The pattern and rates of growth for the immature phasesin the life cycle of A. intermedius differed markedly from thosedescribed under laboratory conditions. Growth in the infantilephase was delayed in the field by low winter temperatures whilesecond stage growth was usually delayed in late spring by dryconditions. The life cycle of A. intermedius appears to be synchronisedby seasonal changes in photoperiod, unlike that of D. reticulatum.Growth in D. reticulatum continued throughout the winter monthsexcept under exceptionally cold conditions and was not usuallydelayed by dry conditions in spring. The size of hermaphroditegland relative to body weight in D. reticulatum reached a maximumin mature-unmated slugs and then became progressively smalleras the slug approached the post-reproductive stage which wasrelatively brief. Exceptionally dry conditions in 1962 delayedthe development of the hermaphrodite gland, the maximum sizereached was significantly reduced and fewer eggs were laid. (Received 17 September 1987; accepted 2 January 1988)     

THE INFLUENCE OF FOOD DEPRIVATION ON THE LOCOMOTOR ACTIVITY OF SLUGS

Experimental animals used in studies of feeding behaviour areusually starved prior to observation, although little is knownof the behavioural effects of such pre-experimental treatment.This study investigated the effect of food deprivation on thelocomotor activity of two species of slugs, Deroceras reticulatumand Milax budapestensis. Locomotor activity was measured bythe quantitative assessment of mucus trails. The more surfaceactiveD. reticulatum showed a marked increase in its locomotoractivity when deprived of food, whereas the locomotor activityof M. budapestensis decreased. The implications of the resultsare discussed in relation to the ‘life-styles’ ofthe two species and the possible implications for the controlof slugs in the potato crop. ^*Present address: 59 Valley Park Drive, Clanfield, Hants. PO80PL. (Received 24 May 1986;     

Predation by the Carabid Beetles Pterostichus melanarius and Poecilus cupreus on Slugs and Slug Eggs

Predation on slugs, Arion lusitanicus (Mabille) and Deroceras reticulatum (Müller), and their eggs by the carabid beetles Pterostichus melanarius (Illiger) and Poecilus cupreus (L.) was investigated in the laboratory. Slugs of different size and slug eggs were offered to the beetles in petri dishes with and without several alternative prey. Pterostichus melanarius destroyed eggs of D. reticulatum and A. lusitanicus, with a clear preference for D. reticulatum eggs. The availability of some alternative prey types adversely affected feeding on D. reticulatum eggs by P. cupreus and feeding on A. lusitanicus eggs by P. melanarius. Yet consumption of D. reticulatum eggs by P. melanarius was not significantly influenced by any alternative prey. Only P. melanarius killed small D. reticulatum in the presence of alternative prey. This suggests that P. melanarius and P. cupreus may have the potential to reduce slug populations in the field by destroying slug eggs and, in the case of P. melanarius, killing freshly hatched slugs.     

Contrasting predation pressure of different ground beetles on slugs and their eggs

Slugs such as Deroceras reticulatum Müller (Pulmonata: Agriolimacidae) cause crop losses in temperate regions worldwide. Numerous species of ground beetles (Coleoptera: Carabidae) feed on slugs, but possible interactive effects are poorly known. Here, we compared predation pressure on slug eggs and slug immatures among Abax parallelepipedus Piller and Mitterpacher, Pterostichus niger Schaller, and Pterostichus melanarius Illiger. The latter was most effective against slug eggs but least effective against immature slugs compared to the other species, demonstrating some specificity of ground beetle predation on life stages. Slug egg predation of P. niger in combination with both other species was synergistic, with up to twice as many eggs consumed as for single species treatments. The effect of species combinations on immature slugs showed no differences from expectations based on single species treatments. Our results indicate that combinations of ground beetle species are favourable for the suppression of slugs in early life stages.     

Effects of metaldehyde and methiocarb on feeding preferences and survival of a slug predator (Pterostichus melanarius (F.): Carabidae,Pterostichini)

The potential for reducing slug populations in crops through predation by generalist carabid beetles is well documented. However, few studies have considered interactions between biological and chemical control agents of slugs. Laboratory assays supported previous findings that the consumption of metaldehyde by slugs (Deroceras reticulatum) leads to increased duration of feeding bouts by carabid beetles (such as Pterostichini) on sub‐lethally affected individuals. However, a similar effect was not found for Pterostichus melanarius exposed to slugs fed on the other widely applied pelleted molluscicide formulation (methiocarb). Examination of beetle survival after consumption of slugs containing molluscicides demonstrated the strong biocidal properties of methiocarb, whereas metaldehyde consumption (ingested through slug predation) did not differ from control slugs killed by freezing. Beetle avoidance of slugs containing a more toxic molluscicide compound and the interaction between slug mucus production and beetle attack rates are discussed.     

THE INFLUENCE OF AN ALTERNATIVE FOOD ON THE EFFECTIVENESS OF PROPRIETARY MOLLUSCICIDAL PELLETS AGAINST TWO SPECIES OF SLUGS

The effectiveness of three types of molluscicidal pellets againstDeroceras reticulatum (Müller) and Arion hortensis (Férussac)was assessed in the presence and absence of potato leaf discs.Pellets containing methi-ocarb were significantly more effectiveagainst both of the slug species than either of the metaldehydeformulations, in the absence of leaf discs. When leaf discswere present, methiocarb was no more effective against D. reticulatumthan were the metaldehyde formulations. Leaf discs did not interferewith the effectiveness of methiocarb against A. hortensis. Theresults are discussed in relation to past work and more recentfield studies. (Received 24 March 1986;     

Predation on the slug Deroceras reticulatum by the carabid beetles Pterostichus madidus and Nebria brevicollis in the presence of alternative prey

1 Slugs are important pests in many agricultural crops and potential biological control agents are being studied as an alternative to molluscicides. Carabid beetles may be able to reduce slug populations, but their role as control agents may be influenced by the presence of alternative prey. 2 Attacks on the pest slug Deroceras reticulatum (Müller) by the carabid beetles Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) were investigated in the presence of alternative prey (earthworms and Calliphora fly larvae). Consumption of slug eggs and aphids was also investigated. 3 All five prey types were consumed to varying degrees during the experiments. Both beetle species showed a significant preference for Calliphora larvae over slugs. Pterostichus madidus showed a significant preference for earthworms over slugs. No preference was shown between earthworms or Calliphora larvae by P. madidus females or N. brevicollis. However, P. madidus males showed a significant preference for Calliphora larvae over worms. Pterostichus madidus showed no preference between slug eggs and aphids; N. brevicollis showed a significant preference for aphids over slug eggs. 4 The results from this study indicate that generalist beetles will often attack other prey in preferences to adult slugs. Slugs may not be preferred because of their mucus. Other prey items occur frequently in arable soils and generalist carabids may ignore slugs altogether and may only feed on them when slug density is high or other prey are unavailable.     

Molecular identification of predation by carabid beetles on exotic and native slugs in a strawberry agroecosystem

Generalist predators have the capacity to exert significant pressure on prey populations. However, integrating them into biological control programs relies on a detailed understanding of their foraging behavior and the levels of trophic connectedness with pest species. Carabid beetles are important predators of slugs, pests of agricultural, floricultural and horticultural crops worldwide, but these interactions have been rarely studied outside the Western Palearctic ecozone. Diagnostic molecular gut-content analysis was used to examine the strength of trophic pathways between a community of carabid beetles and two slug species, the exotic Deroceras reticulatum and native Deroceras laeve, in strawberry agroecosystems. Strawberries were grown according to standard horticultural practices for central Kentucky, following traditional bare ground planting or with the addition of detrital subsidies, to quantify the impact of habitat management on the abundance of pests and the strength of these trophic pathways. Following laboratory characterization of species-specific molecular markers targeting both Deroceras species, carabid beetles collected from a strawberry agroecosystem were screened for slug DNA. Field collections revealed important food web pathways existed between Harpalus pensylvanicus and D. reticulatum, with 7.2% screening positive for these prey yet none screening positive for D. laeve. In contrast, Chlaenius tricolor was found to feed on both slug species in the field, with 16% screening positive for both Deroceras. Despite below normal rainfall limiting slug densities in the field, the results presented here reveal the potential importance of carabid beetles in slug population dynamics in the Nearctic.     

DOES TANDONIA BUDAPESTENIS (MOLLUSCA: PULMONATA) CONTAIN TOXINS? EVIDENCE FROM FEEDING TRIALS WITH THE SLUG PREDATOR PTEROSTICHUS MELANARIUS (COLEOPTERA: CARABIDAE)

Slugs are known to be killed and consumed by a range of invertebrateand vertebrate predators in the field. Carabid beetles (Coleoptera:Carabidae) in particular are major natural enemies of slugs,and have been shown to be capable of controlling certain speciesin a crop environment. This paper reports experiments to investigatethe effects, on the carabid Pterostichus melanarius, of feedingon the milacid slug Tandonia budapestensis. The slugs provedto be toxic, 50% of the beetles dying within two days. Mortalityof beetles fed on two other species of slug, Deroceras reticulatumand Arion distinctus, was not significantly different from thatfor beetles fed upon a control prey (Calliphora vomitoria).These results represent the only known case of a European slugproving to be toxic to potential predators, and is one of avery small number of reported instances of possible toxicityamongst terrestrial gastropods. The possibility that the orangeline down the dorsal keel of T. budapestensis may act as warningcoloration to birds is discussed. (Received 4 December 1996; accepted 5 March 1997)     

Spatiotemporal Analysis of Predation by Carabid Beetles (Carabidae) on Nematode Infected and Uninfected Slugs in the Field

The dynamics of predation on parasites within prey has received relatively little attention despite the profound effects this is likely to have on both prey and parasite numbers and hence on biological control programmes where parasites are employed. The nematode Phasmarhabditis hermaphrodita is a commercially available biological agent against slugs. Predation on these slugs may, at the same time, result in intraguild predation on slug-parasitic nematodes. This study describes, for the first time, predation by carabid beetles on slugs and their nematode parasites on both spatial and temporal scales, using PCR-based methods. The highest nematode infection levels were found in the slugs Deroceras reticulatum and Arion silvaticus. Numbers of infected slugs decreased over time and no infected slugs were found four months after nematode application. The density of the most abundant slug, the invasive Arion vulgaris, was positively related to the activity-density of the carabid beetle, Carabus nemoralis. Predation on slugs was density and size related, with highest predation levels also on A. vulgaris. Predation on A. vulgaris decreased significantly in summer when these slugs were larger than one gram. Predation by C. nemoralis on slugs was opportunistic, without any preferences for specific species. Intraguild predation on the nematodes was low, suggesting that carabid beetles such as C. nemoralis probably do not have a significant impact on the success of biological control using P. hermaphrodita.     

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.     

Pathogenicity of Moraxella osloensis, a Bacterium Associated with the Nematode Phasmarhabditis hermaphrodita, to the Slug Deroceras reticulatum

Moraxella osloensis, a gram-negative bacterium, is associated with Phasmarhabditis hermaphrodita, a nematode parasite of slugs. This bacterium-feeding nematode has potential for the biological control of slugs, especially the grey garden slug, Deroceras reticulatum. Infective juveniles of P. hermaphrodita invade the shell cavity of the slug, develop into self-fertilizing hermaphrodites, and produce progeny, resulting in host death. However, the role of the associated bacterium in the pathogenicity of the nematode to the slug is unknown. We discovered that M. osloensis alone is pathogenic to D. reticulatum after injection into the shell cavity or hemocoel of the slug. The bacteria from 60-h cultures were more pathogenic than the bacteria from 40-h cultures, as indicated by the higher and more rapid mortality of the slugs injected with the former. Coinjection of penicillin and streptomycin with the 60-h bacterial culture reduced its pathogenicity to the slug. Further work suggested that the reduction and loss of pathogenicity of the aged infective juveniles of P. hermaphrodita to D. reticulatum result from the loss of M. osloensis from the aged nematodes. Also, axenic J1/J2 nematodes were nonpathogenic after injection into the shell cavity. Therefore, we conclude that the bacterium is the sole killing agent of D. reticulatum in the nematode-bacterium complex and that P. hermaphrodita acts only as a vector to transport the bacterium into the shell cavity of the slug. The identification of the toxic metabolites produced by M. osloensis is being pursued.     

TESTING MOLLUSCICIDES AGAINST SLUGS -- THE IMPORTANCE OF ASSESSING THE RESIDUAL POPULATION

The field performance of molluscicidal baits used against slugpests of crops is normally measured by the numbers of dead animalsfound on the surface of plots which have been treated with differentbaits. This paper presents evidence that although some materialsseem to give high kills of the Grey Field Slug, Deroceras reticulatum(Müll.), the numbers of slugs left behind on the plotsafter such treatments, the residual population, are the sameas other plots where apparently fewer slugs have been killed.It thus seems likely that the field performance of baits containingmethiocarb and metaldehyde are very similar, both materialsproducing a similar reduction in the adult slug population. Field trials of mollusdcides should always attempt to measurethe residual population left behind on the plots and this wouldthen eliminate apparent errors in the interpretation of resultscaused by the dissimilar modes of action of the various materialsused as molluscicides. (Received 8 June 1982;     

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.     

DYE-MARKING SLUGS

The problems of marking slugs are discussed and a dye-markingtechnique using a Panjet dental inoculator tested for markingthe grey field slug Deroceras reticulatum (Müller) in thelaboratory. The mortalities of a group of 50 marked slugs anda group of 50 unmarked slugs are compared. Results of anotherexperiment to compare the Panjet dye marking technique withfreeze branding are also presented. The reaction of slugs to the marking techniques are described,the possibilities for future use of the technique are discussedansd the case of Panjet marked slugs for ecological and populationstudies is recommended. (Received 31 October 1985;     

The Influence of Mucus Production by the Slug,Deroceras reticulatum,on Predation by Pterostichus madidus and Nebria brevicollis (Coleoptera: Carabidae)

Slugs are important pests in many agricultural crops and potential biological control agents are being studied as an alternative to molluscicides. Carabid beetles may be able to reduce slug populations, but the defence mucus of slugs may deter some predator attacks. Interactions between the carabids Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) with healthy and 'stressed' (unable to produce defence mucus) Deroceras reticulatum (Müller) were investigated in laboratory conditions. Both beetle species consumed significantly more stressed slugs than controls. Defence mucus production by control slugs hampered beetle attacks. These generalist beetle species did not direct their attacks at vulnerable parts of the prey as equal numbers of contacts were made on the slug head, mantle and tail. Blowfly ( Calliphora sp.) larvae are readily consumed by P. madidus and N. brevicollis . Calliphora larvae coated in slug defence mucus were less acceptable to both beetle species compared with control larvae. Results indicate that these generalist beetle species are unable to overcome the defence mucus production of healthy slugs. Slugs sub-lethally poisoned by molluscicides may be a more suitable prey item due to a reduction in defence mucus production.     

FOOD-INDUCED BODY PIGMENTATION QUESTIONS THE TAXONOMIC VALUE OF COLOUR IN THE SELF-FERTILIZING SLUG CARINARION SPP

Body pigmentation is a popular taxonomic marker in slugs to discriminateclosely related species. However, the genetic background ofbody pigmentation is known only for a few species, while inmany others body pigmentation is influenced by age, food and/orclimate. In this study, we investigated the effects of differentfood items on body pigmentation expression in two selfing pulmonategastropods, Arion (Carinarion) silvaticus and Arion (Carinarion) fasciatus.Both species mainly differ in the distribution of yellow-orangegranules on the body, which in A. fasciatus are concentratedin lateral bands, and in A. silvaticus are evenly scattered.Animals were raised individually under the same conditions, whilethey laid eggs as a consequence of selfing. This F₁ generation wasafterwards divided into two groups, which were fed with different fooditems. A diet of carrot, lettuce or paper had no effect on the distributionof the yellow-orange granules in A. silvaticus, but provokeda loss of the yellow-orange lateral bands in A. fasciatusso that externally these F₁ specimens became similar to A. silvaticus.In both species, a diet of nettle resulted in a strong yellow-orangepigmentation, which often formed yellow-orange lateral bands.These results indicate that food can probably influence the'species-specific' body pigmentation in Carinarion, and thusquestion the reliability of colour traits to distinguish A.silvaticus and A. fasciatus. (Received 5 May 2000; accepted 29 September 2000)     

Predation by the carabid beetle Harpalus rufipes on the pest slug Deroceras reticulatum in the laboratory

The Harpalini species Harpalus rufipes, as many other generalist carabids, consume a wide variety of prey and it is known to feed on pest slugs such as the grey field slug Deroceras reticulatum, but quantitative data about the predatory activity of H. rufipes on slugs are very scarce. In laboratory experiments, we assessed the capability of male H. rufipes to kill eggs and different‐sized slugs of the pest species D. reticulatum in either the absence or the presence of alternative live prey (dipteran larvae and aphids). We also investigated the preference of H. rufipes for eggs and hatchlings of D. reticulatum in a choice experiment. H. rufipes killed considerable amounts of eggs and small juveniles (≤5.0 mg) of D. reticulatum, both in no‐choice and in choice situations. Medium‐sized juvenile slugs (10–20 mg) were seldom killed only in no‐choice situations, and no large juveniles (50–60 mg) were killed. Dipteran larvae and aphids were killed also in no‐choice and in choice situations. The type of alternative prey presented with slug eggs affected the survival of the eggs to H. rufipes predation. The presence of dipteran larvae as alternative prey did not affect the survival of juvenile slugs. When eggs and small juvenile slugs were offered together, the survivals of both items were similar. The obtained results under laboratory conditions suggest that the generalist predator H. rufipes might realise an important contribution to the control of pest slugs.