The effects of food and density on the nocturnal behaviour ofArion ater andAriolimax columbianus (Pulmonata: Stylommatophora)

The effects of food supply and population density on the nocturnal behaviour of Arion ater and Ariolimax columbianus were investigated. Density did not significantly affect A. ater's level of activity or short-term movement, resting, or feeding. A. columbianus was more active and moved, rested, and fed more frequently when slug density was high. A. ater foraged and rested more often, but fed less when good food was unavailable. Ariolimax's only response to the food regime was to feed more when good food was available. Seasonal changes in the level of activity and behaviour of Arion were evident, whereas Ariolimax's activity and behavioural repertoire were not similarly affected. Arion ater's nightly activity appeared to be mainly food oriented, while Ariolimax columbianus seemed most responsive to slug density during its nocturnal activity periods.     

Phylogeny of slug species of the genus Arion: evidence of monophyly of Iberian endemics and of the existence of relict species in Pyrenean refuges

The Iberian Peninsula contains the majority of the Paleartic land slug species of the genus Arion, which exhibits diverse taxonomic problems. The present study investigated Arion taxonomy on the basis of analyses of the mitochondrial ND1 gene and nuclear internal transcribed spacer 1 (ITS1) sequences. The Iberian endemic species were monophyletically clustered in two divergent sister clades. The topotype specimens of Arion lusitanicus and the closely related species Arion nobrei and Arion fuligineus, as well as Arion hispanicus and Arion flagellus, were grouped into an ‘Atlantic’ clade, whereas Arion baeticus, Arion gilvus, Arion anguloi, Arion wiktori and Arion paularensis were included in a ‘Continental–Mediterranean’ clade. Calibration of mutation rate in the ND1 gene suggested that the divergence of these two clades occurred around the Pliocene–Pleistocene boundary, with subsequent speciation events during the Pleistocene. A group of ancestral and divergent endemic species with distribution centred in the Pyrenean mountain range (Arion molinae, Arion lizarrusti, Arion antrhacius and Arion iratii) arose in the Pliocene and survived through the Pleistocene in geographically confined small populations. Arion lusitanicus showed up to be polyphyletic: specimens, sampled outside the geographic range of the topotype in the north‐western Iberian Peninsula, were included in a non‐monophyletic clade together with the widely distributed species Arion ater and Arion rufus. The divergent species with a wide European distribution (Arion subfuscus, Arion hortensis, Arion fagophilus and Arion intermedius) were located in basal positions in all topologies. The evolutionary history of these slug species (highly sensitive to climatic factors, with capacity for both outcrossing and selfing, and with low dispersal ability) appears to have been moulded by Pliocene–Pleistocene climate events and by the rugged topography of southern Europe, giving rise to repeated cycles of population isolation during periods of glaciation alternating with interglacial expansions limited by geographic barriers.     

Alternative risk-taking styles: The case of time-budgeting strategies of terrestrial gastropods

The terrestrial slugs Arion ater, Limax maximus and Ariolimax columbianus have similar morphological designs but differ remarkably in their life history tactics and behavioural time budgets. The adaptive value of particular risk-taking styles was investigated using a comprehensive computer simulation model. The model allowed each species' success (growth rates, food acquisition) and costs (distance travelled, hydration deficits, injury from aggressive encounters) to be evaluated in various types of weather (benign, harsh and surprise). This was interpreted in terms of the species' life history design. In addition, each species was simulated with the behavioural strategy of the other two species substituted for its natural programming. The simulation experiments demonstrated how variations in a few simple rules could lead to divergent cost-benefit consequences. The model also illustrated that coarsely-tuned sensitivity to the environment may be better than finely-tuned responses, depending on the animal's ability to respond quickly and the degree of risk. The simulations suggested that competition has shaped the time-budgeting tactics of A. ater and L. maximus. Each species actually performed better in harsh weather, using the behavioural program of the other. L. maximus is an aggressive species with a narrow, nocturnal activity period. The concentrated activity of L. maximus may allow it to displace competitors more effectively, whereas the broader time span of A. ater's activity may be necessary to avoid L. maximus.     

An electrophoretic characterisation of three paratypes of Arion fagophilus De Winter, 1986, with notes on the subgeneric division of the genus Arion Férussac, 1819 (Mollusca, Pulmonata)

Three paratypes of Arion fagophilus were electrophoretically compared with nine other West European species of the genus Arion. Esterases, glutamate-oxaloacetate-transaminase, a-amylase, superoxide dismutase, phosphoglucomutase, fumarase, α-glycerophosphate dehydrogenase and phosphogluconate dehydrogenase were screened in the hepatopancreas and general proteins in the albumen gland. A. fagophilus appears to be a very well defined species, which may be assigned to the subgenus Kobeltia. The subgeneric division of the genus Arion is preliminary discussed with respect to electrophoretic and chromosomal data. It is concluded to retain four subgenera, which may be grouped two by two: Mesarion and Arion s. s. on one hand, and Kobeltia and Carinarion on the other. It is remarked that the electrophoretic differences between Norwegian and Belgian A. ater suggest a separation into two taxa.     

Immunocytochemical Localisation of Glycosaminoglycan-Like Activity in the Mucus and Associated Tissues of Terrestrial Slug Species (Gastropoda: Pulmonata)

Indirect immunofluorescence assays were conducted on cryotome sections of four terrestrial slug species from three distinct phylogenetic groups, Arion ater (L.), Arion hortensis (Férussac), Tandonia (Milax) budapestensis (Hazay), and Deroceras reticulatum (Müller) using monoclonal antibodies for two glycosaminoglycans (GAGs), heparan sulphate, and chondroitin sulphate. Specific staining for a heparan sulphate-like component was demonstrated in the foot and tail regions of each species and was particularly intense in A. ater and A. hortensis, notably in the epidermis and associated mucus-like material, and in mucus-like material from the pedal gland region of the latter species. Subsequent studies with A. ater confirmed the presence of heparan-sulphate–like activity in the caudal gland duct region. No evidence of specific staining for chondroitin sulphate-like activity was found in any of the slug species. This work suggests that a specific GAG, or a group of closely related GAGs, is a common component of mucus in a range of slug species and of different types of mucus, including trail (pedal) mucus and the more viscous mucus produced by the caudal gland.     

Dispersal movements of non-native and native terrestrial slugs in an urban environment

Animal movement varies from undirected dispersal to directed migration. Movement rates may have implications for conservation and resource management, as well as pest control, and they play a key role in invasion success. In slugs, long-distance dispersal is typically passive, whereas active movement is critical for local dispersal and determines access to resources such as food and shelter. Telemetry has recently been used to study individual slug movements in the wild, whereas movement in arena tests has explored mechanisms of interspecific competition and invasiveness in slugs. Studies that relate the performance of individual slugs in arena tests to their post-release behavior in nature are lacking. We measured individual short-term movement speed of commonly occurring native and non-native slugs of the genera Arion and Limax in arena tests and tracked their post-release dispersal movements in a garden by PIT telemetry. We demonstrate clear differences in movement behavior among the species, but non-native slugs did not display higher movement rates than their native congeners. In the arena test, slugs of the genus Limax displayed a higher short-term speed than slugs of the genus Arion, whereas in the field, individuals of Limax maximus showed lower dispersal rates compared to the other slug species. Moreover, there was a positive correlation between short-term speed in the arena test and movement in the field among individuals of L. cinereoniger, indicating the possible existence of behavioral syndromes in slugs, which may link movement ecology, animal personality, and the invasion ecology of pest species.     

Consequences of competition on the reproduction and mortality of three species of terrestrial slugs

The interaction of three species of terrestrial slugs was studied in six field cages containing monocultures and 2-way combinations. A. columbianus and A. ater were non-aggressive species and L. maximus was highly aggressive during the summer. All three species were similar in body design and resource utilization. Reproduction (egg size, eggs/batch, number of batches, wet-weight productivity) and mortality were monitored through the 1977 season. L. maximus drastically reduced reproductive success of the non-aggressive slugs and increased their mortality. There were also significant interactions between the non-aggressive species. Compared to other invertebrates, the life history characteristics of A. columbianus are strongly K-selected and probably evolved through intraspecific exploitative competition in predictable, stable forest habitats. A. ater is best classified as a refuging r-selected species adapted to less predictable, unstable field environments. L. maximus appears to be α-selected and its overall design is suited to density-dependent regulation in fields.     

Consequences of competition on the time budgets,growth and distributions of three species of terrestrial slugs

Summary The behavioural time budgets of two non-aggressive slugs and one aggreessive species were evaluated in monocultures or in 2-way combinations in field cages. The seasonal time budgets ofL. maximus, A. ater andA. columbianus in monoculture were remarkably similar despite their strong differences in physiological resource allocation. Similarity in physical design and resource utilization appeared to be more important determinants of behaviour. The time budgets showed distinct seasonal patterns associated with underlying physiological demands for reserve accumulation, sperm acquisition and oviposition.A. ater andL. maximus displayed a definite dispersal phase associated with femalephase maturation. In combination,A. ater andA. columbianus had no detectable effect on one another’s time budgets.L. maximus disrupted nearly every category of behaviour in these non-aggressive species, but its own time budget was not significantly different in the monoculture or in the two combinations, except for minor effects on resting and sexual behaviour.L. maximus appeared to be adapted to maximize behavioural disturbance of interspecific competitors while minimizing intraspecific effects.     

FEEDING AND GROWTH RESPONSES TO COPPER, ZINC, MERCURY AND LEAD IN THE TERRESTRIAL GASTROPOD ARIONATER (LINNE)

Food consumption and growth variations related to several dosagesof heavy metals (Cu, Zn, Hg and Pb) in diet were studied inthe terrestrial slug Arion ater for 27 days of experimentaltreatment. The work reported here is part of a larger project to investigatethe histological effects of metal exposure in pulmonate gastropods. Graphics on food consumption and growth vs. time for each metal,and regression for each behaviour observed are presented. (Received 30 May 1985;     

Mini-plot field experiments on slug control using biological and chemical control agents

In three field experiments, the rhabditid nematode Phasmarhabditis hermaphrodita was applied one or more times at the standard rate (3 × 10⁹ ha^?1) or half the standard rate to protect crops from slug damage under experimental conditions. Expt 1 was done in a field planted with the ornamental Polygonatum japonica. The treatments were: infective juveniles of the nematode at the standard field rate, metaldehyde pellets at the recommended field rate, and ioxynil (a herbicide with molluscicidal properties) at 90 mg m^?2. The treatments were repeated every 2 wk. Arion ater agg. caused most of the damage to P. japonica. There were no significant differences in damage between treatments during the 3 wk after first application, but plants on plots treated with metaldehyde or nematodes had significantly less damage than plants on untreated plots in the fourth and fifth weeks. Expts 2 and 3 were done on the same site, the first with leaf beet and the second with lettuce. The treatments in these experiments were: nematodes applied to the planted area at the standard field rate 3 days prior to planting, with or without previous application of cow manure; nematodes at half standard rate applied twice, 6 days apart, to the planted area or to the surrounding area; metaldehyde pellets and iron phosphate pellets, both applied at the recommended rate to the planted area immediately after planting. In both experiments, the two chemical molluscicides and nematodes applied once to the planted area at the standard field rate without previous application of cow manure, or twice at half standard rate, were able to reduce slug damage. Nematodes applied after manure did not reduce slug damage. None of the treatments reduced the numbers of slugs contaminating the harvested plants. Slug populations were assessed by means of soil sampling before and after Expts 2 and 3. Only after Expt 3 was there a significant effect of treatment on slug numbers, with significantly fewer in metaldehyde treated plots than in untreated plots.     

How can two soft bodied animals be precisely connected? a miniature quick‐connect system in the slugs,Arion lusitanicus and Arion rufus

Among stylommatophoran gastropods, many species have simplified or reduced their copulatory organs, for example, within the Arionidae, many species lack penes. In this study, I ask two questions 1) How are soft bodied slugs which do not possess a penis connected during copulation? and 2) Is there a mechanical isolating barrier between related sympatric slug species? Observations on the mating behavior and the functional morphology of the distal genital apparatus were made in a mixed population of Arion lusitanicus and Arion rufus. The investigated Arion species exhibit an elaborate copulation process with a quick genital coupling. Prior to full eversion of the distal genitalia, the genital coupling proceeds inside the atrium cavity of one of the partners. This is in contrast to the symmetrical mutual eversion in penis‐bearing species. The donor–recipient channels are tightly connected to one after another and fit precisely. During copulation, the jelly‐coated spermatophore of the donor is pressed out into the connected channel of the partner, where it is implanted only with its frontal part. In the field, successful interspecific matings in terms of spermatophore transfer were rarely observed. The observations presented indicate a mechanical barrier which may profoundly influence the intraspecific and interspecific mating success. J. Morphol. 276:631–648, 2015. © 2014 Wiley Periodicals, Inc.     

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.     

The role of genetic and chemical variation of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> seedlings in influencing slug herbivory

This study investigated the genetic and chemical basis of resistance of Pinus sylvestris seedlings to herbivory by a generalist mollusc, Arion ater. Using feeding trials with captive animals, we examined selective herbivory by A. ater of young P. sylvestris seedlings of different genotypes and correlated preferences with seedling monoterpene levels. We also investigated the feeding responses of A. ater to artificial diets laced with two monoterpenes, Δ³-carene and α-pinene. Logistic regression indicated that two factors were the best predictors of whether seedlings in the trial would be consumed. Individual slug variation (replicates) was the most significant factor in the model; however, α-pinene concentration (also representing β-pinene, Δ³-carene and total monoterpenes due to multicollinearity) of needles was also a significant factor. While A. ater did not select seedlings on the basis of family, seedlings not eaten were significantly higher in levels of α-pinene compared to seedlings that were consumed. We also demonstrated significant genetic variation in α-pinene concentration of seedlings between different families of P. sylvestris. Nitrogen and three morphological seedling characteristics (stem length, needle length and stem diameter) also showed significant genetic variation between P. sylvestris families. Artificial diets laced with high (5 mg g⁻¹ dry matter) quantities of either Δ³-carene or α-pinene, were eaten significantly less than control diets with no added monoterpenes, supporting the results of the seedling feeding trial. This study demonstrates that A. ater selectively feed on P. sylvestris seedlings and that this selection is based, in part, on the monoterpene concentration of seedlings. These results, coupled with significant genetic variation in α-pinene concentration of seedlings and evidence that slug herbivory is detrimental to P. sylvestris fitness, are discussed as possible evidence for A. ater as a selective force on the evolution of defensive chemistry in P. sylvestris.     

The genetic dynamics of the rapid and recent colonization of Denmark by <Emphasis Type="Italic">Arion lusitanicus</Emphasis> (Mollusca,Pulmonata, Arionidae)

We describe the genetic dynamics of the recent establishment of the ‘Iberian slug’, Arion lusitanicus J. Mabille 1868, in Denmark and compare its population structure to two other members of the ‘large Arion complex’, Arion ater ater, native to Denmark, and Arion ater rufus, introduced into Denmark in the early 1900s. Assaying allozyme polymorphism at seven enzyme loci, we found that: (1) None of the three taxa reproduce primarily by self-fertilization. Differences among loci and colonies in the pattern of deviation from Hardy–Weinberg equilibrium are most consistent with isolate mixing and perhaps with low amounts of selfing. (2) For both A. lusitanicus and A. a. rufus, gene diversity is lower in Danish colonies than in southern German colonies, implying population bottlenecks in the establishment of Danish colonies. (3) Significant linkage disequilibrium values usually involve the same three loci, viz. PGI, MDH-1 and MDH-2, suggesting physical linkage among these loci. (4) For both A. a. rufus and A. lusitanicus, the overall gene frequencies from Denmark and southern Germany are homogeneous, while variation among colonies within these regions ranges from around 15 to 28% for the three taxa. This indicates strong, local population genetic subdivision but with little restriction to gene flow from possible source areas. The heterogeneity in measures of diversity and differentiation indicates that population structure for all three taxa is dominated by ongoing founder effects, local extinction/colonisation dynamics, and genetic drift processes.     

Long‐distance dispersal of non‐native pine bark beetles from host resources

1. Dispersal and host detection are behaviours promoting the spread of invading populations in a landscape matrix. In fragmented landscapes, the spatial arrangement of habitat structure affects the dispersal success of organisms. 2. The aim of the present study was to determine the long distance dispersal capabilities of two non‐native pine bark beetles (Hylurgus ligniperda and Hylastes ater) in a modified and fragmented landscape with non‐native pine trees. The role of pine density in relation to the abundance of dispersing beetles was also investigated. 3. This study took place in the Southern Alps, New Zealand. A network of insect panel traps was installed in remote valleys at known distances from pine resources (plantations or windbreaks). Beetle abundance was compared with spatially weighted estimates of nearby pine plantations and pine windbreaks. 4. Both beetles were found ≥25 km from the nearest host patch, indicating strong dispersal and host detection capabilities. Small pine patches appear to serve as stepping stones, promoting spread through the landscape. Hylurgus ligniperda (F.) abundance had a strong inverse association with pine plantations and windbreaks, whereas H. ater abundance was not correlated with distance to pine plantations but positively correlated with distance to pine windbreaks, probably reflecting differences in biology and niche preferences. Host availability and dispersed beetle abundance are the proposed limiting factors impeding the spread of these beetles. 5. These mechanistic insights into the spread and persistence of H. ater and H. ligniperda in a fragmented landscape provide ecologists and land managers with a better understanding of factors leading to successful invasion events, particularly in relation to the importance of long‐distance dispersal ability and the distribution and size of host patches.     

Aerial dispersal of the two-spotted spider mite (Tetranychus urticae) from field corn

Field data from 48 plots monitored during diverse weather conditions in two separate years were subjected to multiple regression analysis to determine which factors were related to spider-mite aerial dispersal. With the number of aerially dispersing mites as the dependent variable, partial regression coefficients (b) for mite population density and percent corn-leaf area infested with mites were positive, while those for percentNeozygites-infected mites and hours per week 90% r.h. were negative. When an aerial dispersal index (number of aerially dispersing mites/mites per plant) was used as the dependent variable, the partial regression coefficient for percent leaf area infested was positive, while coefficients for hours per week 90% r.h. and percentNeozygites-infected mites were negative. Mite aerial dispersal was greatest in predator-suppressed field plots under dry weather conditions. Mite aerial dispersal was substantially reduced in plots where moist weather conditions induced epizootics ofNeozygites floridana before corn plants became entirely infested with mites.     

Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

Fish predation and offspring survival in the prosobranch snail Viviparus ater

In this study we identified some of the predators of the freshwater snail Viviparus ater and estimated offspring survival to the end of the first summer in a natural population. Newborn V. ater were eaten by the fish Barbus barbus, Rutilus rutilus, Scardinius erythrophtalmus and Tinca tinca. Out of 137 guts of Abramis brama caught in Lake Zürich 1 contained shell fragments and an operculum of a newborn V. ater. On a 40×40 m grid near Goldbach, Lake Zürich, we counted 678 adult females of V. ater in June 1988, which together gave birth to approximately 13 300 offspring throughout the summer. In October we found 1348 V. ater of age class 0 on the grid, i.e. approximately 10% of the year's young had survived to the end of their first summer. The grid had been divided into two sections, A and B. In section A, 72 carthenware tiles had been placed as shelters for V. ater. Offspring survival was slightly but significantly higher here (10.9%) than in section B (9.6%). The observation that offspring survival in the natural habitat was 1 order of magnitude lower than in cages suggests that predation is an important cause of mortality of newborn V. ater. Average population density was 2 individuals m^–2 There was a migration from shallow (1–4 m) to deeper water (5–9 m) in September. The average distance between weekly recordings of tagged V. ater was 4.7 m in males and 3.0 m in females.     

A HISTOLOGICAL SURVEY OF GONADAL DEVELOPMENT IN ARION ATER L. (MOLLUSCA, PULMONATA)

A method of breeding the slug Arion ater L. in the laboratorywas established. Measurements were made at monthly intervalsof body weight, weight and volume of the hermaphrodite gland,diameter and number of acini, oocytes and other cell types insidethe acini in field and laboratory-reared animals. Male and femalegametes grow simultaneously in the hermaphrodite gland of fieldanimals during the year. High humidity and low temperature causedinhibition of spermatogenesis and increase in the number offemale gametes in laboratory-reared animals. The average rateof increase in diameter of the oocytes of field samples was10 µm per month and in laboratory samples 9 µm permonth. ^*Present address: Department of Biology, School of Sciences,Ferdowsi University, Mashhad, Iran. (Received 30 September 1977;