Spatial distribution and home-range of the pest slug Arion lusitanicus (Mollusca: Pulmonata)

The density and distribution of egg-batches, juveniles and adult Arion lusitanicus, as well as individual home-ranges, were investigated over two seasons. The study was performed on grassland which was mown twice a year and received no herbicides or pesticides, with neighbouring field and stream-side vegetation. Female-mature slugs preferred restricted areas closely neighbouring the taller vegetation of the stream-side vegetation and field, but less-favourable parts of the site were also inhabited in the second year when population densities increased and weather conditions were more favourable. It was postulated that slugs could have home-ranges, as homing behaviour is a well-known phenomenon. The application of a long-lasting individual marking technique enabled calculations of home-ranges (convex polygons) for slugs for the first time. Home-range size averaged 45.4 m² when the population density was low in 1995, and was found to be negatively correlated with density, as it decreased to 12.4 m² in 1996, when population density was high. Mating sites were found to be distinctly closer to the geometric centres of activity than the sites used for egg-deposition. Egg-batches were aggregated under the shelter traps (50 × 50 cm squares of hardboard covered with polystyrene) and the pattern of distribution of egg-batches persisted in the young hatchlings. A number of foreign A. lusitanicus were introduced into the unfavourable centre of the site, and by two to four weeks after release their distribution exactly resembled that of the local ones. In all the various aspects of behaviour investigated, large individual differences were found, suggesting a great plasticity in this generalist species. The importance of the species‘ dispersal capacity in relation to general ecological and applied aspects of control is discussed in the context of the present findings.     

The morphology of the lung of a tropical terrestrial slug Trichotoxon copleyi (Mollusca: Gastropoda: Pulmonata): A scanning and transmission electron microscopic study

The lung of the slug Trichotoxon copleyi is located in the body mantle where it opens to the outside through an adjustable pneumostome situated on the right side. The respiratory surface of the lung is profusely vascularized by anastomosing 'blood' capillaries. A thick muscular layer surrounds the gas exchange surface and may be involved in the efficient rhythmic ventilatory contraction of the lung. Externally an epithelial layer made up of large secretory cells with numerous microvilli overlies the muscular layer. The surface of the lung is made up of squamous cells with an abundance of microvilli and clustered goblet cells interspersed occasionally. The squamous cells cover the notably thin air–blood barrier which is extremely attenuated, particularly in some areas. The goblet cells contain a centrally located nucleus and numerous intracytoplasmic granules, presumably precursors of the mucus that covers the surface of the mantle cavity.
All the essential morphological prerequisites for efficient gas exchange such as an extensive surface and a thin blood–gas barrier were observed in the lung of Trichotoxon. It was concluded that this molluscan lung is structurally and functionally a true lung in all respects.     

A bioassay for screening materials influencing feeding in the field slug Deroceras reticulatum (Müller) (Mollusca: Pulmonata)

Test materials were incorporated in wheatflour pellets to measure their effect on slug feeding. The dry weights of individual pellets were measured before and after exposure for 24 h to starved slugs. The method gave consistent results, and can accommodate test materials with different physical properties. The phagostimulant effect of sucrose was confirmed. An extract of the herb tarragon (Artemisia dracunculus) was the most effective antifeedant of the materials used at the concentrations tested. The method is compared with other published techniques.     

Molecular and morphological data reveal cryptic taxonomic diversity in the terrestrial slug complex Arion subfuscus/fuscus (Mollusca, Pulmonata, Arionidae) in continental north-west Europe

The importance and abundance of cryptic species among invertebrate taxa is well documented. Nowadays, taxonomic, phylogenetic and conservation biological studies frequently use molecular markers to delineate cryptic taxa. Such studies, however, often face the problem of the differential resolution of the molecular markers and techniques involved. This issue is explored in the present study of cryptic taxa within the terrestrial slug complex Arion subfuscus/fuscus in continental north-west Europe. To this end, morphological, allozyme and mitochondrial 16S rDNA sequence data have been jointly evaluated. Using allozyme data and gonad type, two distinct groups were consistently delineated, even under sympatric conditions. The 16S rDNA data strongly supported both those groups and even suggested the presence of three distinct taxa within one of them. However, in view of: (1) the allopatric distribution of three OTUs, (2) the lack of allozyme or morphological differentiation, and (3) the extremely high degree of intraspecific mtDNA variation reported in pulmonate gastropods, they are, for the time being, not regarded as valid species under the biological species concept. By means of 16S rDNA and allozyme data, the position of type and topotype material of A. subfuscus s.s. and A. fuscus relative to the newly defined OTUs was determined, thus clarifying the nomenclature of this species complex. Additionally, gonad type proved to be a useful character for distinguishing the two species in north-west Europe.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 23–38.     

Extreme mtDNA divergences in a terrestrial slug (Gastropoda, Pulmonata, Arionidae): accelerated evolution, allopatric divergence and secondary contact

Extremely high levels of intraspecific mtDNA differences in pulmonate gastropods have been reported repeatedly and several hypotheses to explain them have been postulated. We studied the phylogeny and phylogeography of 51 populations (n = 843) of the highly polymorphic terrestrial slug Arion subfuscus (Draparnaud, 1805) across its native distribution range in Western Europe. By combining the analysis of single stranded conformation polymorphisms (SSCP) and nucleotide sequencing, we obtained individual sequence data for a fragment of the mitochondrial 16S rDNA and a fragment of the nuclear ITS1. Additionally, five polymorphic allozyme loci were scored. Based on the 16S rDNA phylogeny, five monophyletic haplotype groups with sequence divergences of 9-21% were found. Despite this deep mitochondrial divergence, the haplotype groups were not monophyletic for the nuclear ITS1 fragment and haplotype group-specific allozyme alleles were not found. Although there is evidence for an accelerated mtDNA clock, the divergence among the haplotype groups is older than the Pleistocene and their current allopatric ranges probably reflect allopatric divergence and glacial survival in separate refugia from which different post-glacial colonization routes were established. A range-overlap of two mtDNA groups (S1 and S2, 21% sequence divergence) stretched from Central France and Belgium up to the North of the British Isles. The nuclear data suggest that this secondary contact resulted in hybridization between the allopatrically diverged groups. Therefore, it seems that, at least for two of the groups, the deep mtDNA divergence was only partially accompanied by the formation of reproductive isolation.     

Intracisternal polycylinders: A cytoplasmic structure in cells of the terrestrial slug Arion empiricorum férussac (Pulmonata,Stylommatophora)

Study of the digestive organs of the slug Arion empiricorum with the electron microscopy has revealed cytoplasmic structures that we call intracisternal polycylinders (ICPC). They consist of cylinders of cytoplasm (about 550 A in diameter) arranged in sheafs within cisterns of the endoplasmic reticulum. They appear in different cell types, being most common in the digestive epithelium of the midgut. Their morphology and apparent association with other cytoplasmic organelles such as mitochondria, peroxisomes, multivesicular and residual bodies suggests that the ICPC might be involved in exchange, transport and oxidation processes, contributing to the excretory function at a subcellular level.     

Arion transsylvanus (Mollusca,Pulmonata, Arionidae): rediscovery of a cryptic species

Jordaens, K., Pinceel, J., Van Houtte, N., Breugelmans, K. & Backeljau, T. (2010). Arion transsylvanus (Mollusca, Pulmonata, Arionidae): rediscovery of a cryptic species. —Zoologica Scripta, 39, 343–362. Cryptic species are abundant among invertebrates and are often hard to recognise. Molecular markers are an extremely useful tool to delineate cryptic taxa, although they should be applied with caution because different genes and techniques may yield different outcomes. We illustrate how cross‐validation by molecular and morphological data can be applied to optimise taxonomic interpretations when cryptic species are involved. This is performed for the terrestrial slug Arion subfuscus species complex which represents a historical ‘taxonomic garbage can’. Gonad morphology, allozymes and mtDNA data consistently showed that slugs from Romania and a location in E Poland represent a strongly differentiated taxon within this complex. These slugs are therefore formally redescribed and assigned to A. transsylvanus Simroth 1885 ; a forgotten nominal taxon from Transylvania. Diagnostic characters, including DNA sequences for the mitochondrial 16S rDNA are presented. Animals with the morphology of A. brunneus Lehmann 1862 , a nominal taxon which has also been reported from Romania, have the gonad type, allozyme alleles and 16S rDNA haplotypes of either A. fuscus, A. subfuscus or A. transsylvanus. Therefore, A. brunneus is regarded as a colour morph shared by several species and hence, A. transsylvanus is probably the only A. subfuscus‐like species in the Romanian Carpathians.     

The comparative anatomy and systematics of Jamaican Urocoptidae (Mollusca: Pulmonata)

Jamaican Urocoptidae can be assigned to three subfamilies, the Urocoptinae Pilsbry 1902, Apominae Paul 1982 and Microceraminae Pilsbry 1902, on shell and anatomical characters. Urocoptinae are characterized by a cylindrical or conical decollate shell; genitalia with an elongate spermatheca and no penial retractor muscle; the left ocular retractor arising from the pharyngeal retractor and both oculars divided anteriorly; a short or long radula (L/W ≏ 5 or ≏ 10) with V-shaped rows of teeth, no marginals and the mesocones and ectocones of laterals widely separated in alternating rows; and include the genera Urocoptis Beck, Anoma Albers and Spirostemma Pilsbry & Vanatta. The Apominae are characterized by elongate-cylindrical decollate shells; genitalia with a globular spermatheca and a penial retractor muscle; both ocular retractors arising from the pharyngeal and both divided anteriorly; an extremely long radula (L/W= 25–45) with narrow, weakly-lobed centrals flanked by two enormous laterals with ectocones reduced or absent, and 4–9 marginals; and include the genera Apoma Beck, Mychostoma Albers, Geoscala Pilsbry & Vanatta and Simplicervix Pilsbry. The type species of the type genera of both subfamilies occur in Jamaica and their anatomy is described. Jamaican Microceraminae include one species, Microceramus gossei (Pfeiffer) which has a conical, entire shell; genitalia with an oval spermatheca and no penial retractor muscle; neither ocular retractor arising from the pharyngeal and neither branched anteriorly; a short radula (L/W ≏ 5) with straight rows of lateral teeth and no marginals. Cladistic analysis suggests that Microceramus is the most primitive genus; that within the Urocoptinae Anoma and Spirostemma are more closely related to each other than either is to Urocoptis ; and that within the Apominae, Apoma is less closely related to Mychostoma, Geoscala and Simplicervix than they are to each other.     

A photoperiod gene regulates vitellogenesis in Lymnaea peregra (Mollusca: Gastropoda: Pulmonata)

Summary

Populations of the fresh-water snail Lymnaea peregra from different localities vary in their capacity to lay eggs, responding either to long day or to short day photoperiods. These responses are genetically determined by a single gene locus with the long day allele dominant over the short day allele. Although previtellogenic stages of oogenesis are not affected by an inhibitory photoperiod, vitellogenesis is essentially shut down. The volumes of the dorsal bodies, which are endocrine organs that mediate vitellogenesis, are significantly smaller in snails maintained in an inhibitory photoperiod. Implantation of cerebral ganglia with attached dorsal bodies, both from egg laying animals and from dormant animals that have the potential to lay eggs in the postoperative photoperiod, into snails that were inhibited by insufficient photoperiod resulted in a significant increase in vitellogenesis in the ovotestes of the hosts. An increase in host dorsal body volume was also observed. The photoperiod gene appears to control some aspect of the cerebral ganglion that activates the dorsal body to produce a hormone that stimulates vitellogenesis in the ovotestes. Photoperiodic control of reproduction may be a factor contributing to the diverse life history strategies observed in this species.     

Breeding system and population genetic structure in philomycid slugs (Mollusca: Pulmonata)

Four species of philomycid slugs were collected at 13 sites in Tennessee and one site in Virginia. A total of 361 individuals were examined electrophoretically at 11 allozyme loci. Outcrossing was the primary breeding in all populations of all four species. Genetic similarities among demes, among species and among genera are similar to values obtained in studies of other animals with similar vagility and demography and are in good agreement with systematics based on morphological characters. In the species examined in greatest detail, Philomycus carolinianus , there was substantial genetic differentiation among demes. This differentiation appears to result from genetic drift and restricted gene flow. Genetic similarity between P. carolinianus demes did not correlate with the geographic distance between demes and hierarchical F-statistic analysis showed more than half of the differentiation among demes to result from differentiation at the smallest geographic scale. The Tennessee River system also appears to be a major barrier to gene flow among the P. carolinianus demes sampled.     

Ultrastructure of the osphradium of Siphonaria grisea (Mollusca, Pulmonata)

By means of scanning and transmissive electron microscopic methods osphradium of Siphonaria grisea has been studied. The osphradium of the animal is presented as a small torulus formed by supporting ciliated cells. Among them bodies of receptory cells are situated; they are of smaller size and decorated with a bundle of microvilli. Central processes of the receptory cells, penetrating through a thick layer of the connective tissue, reach the subepithelial neural trunk. Concentration of the neural cells in the periphery of the trunk is small, and in the central area no chemical synapses are revealed. The data presented demonstrate a primitive structure of the osphradial chemoreceptory organ in Siphonaria.     

Biogeography of the Helicoidea (Mollusca: Gastropoda: Pulmonata): land snails with a Pangean distribution

The biogeography of helicoid land snails was investigated using cladistic methods. Parsimony analysis under Assumption 0 yielded twelve area cladograms (length=25, c.i.=0.76, r.i.=0.86). The pattern of vicariance for the Helicoidea indicated that families originated with the break up of eastern Gondwana and Laurasia between the late Mesozoic and mid-Tertiary, and possible vicariance events are identified. It is proposed that Asian terranes, located between India and Australia, maintained contact with northern Australia until the late Cretaceous, which is later than is suggested in current palaeogeographical hypotheses.