Edmund Gittenberger

The Pleistocene glaciations and the evolutionary history of the polytypic snail species Arianta arbustorum (Gastropoda, Pulmonata, Helicidae)

The evolutionary history of the snail Arianta arbustorum is controversial. This diverse, polytypic species has two distinct forms: one, with a globular shell and closed umbilicus, is found from lowland to high altitudes; the other, with a depressed shell and open umbilicus, is found at a few scattered, high altitude localities. What is the origin of these two forms? Some believe that the depressed shell is a recent, local, ecotypic adaptation to alpine environments. Others believe that this form is a relic of an ancestral condition that may have survived the Pleistocene glaciations on nunatak-like montane refugia, while the globular shell is a derived condition and its presence at high altitudes follows post-Pleistocene recolonisation. We analysed a portion of the mitochondrial gene cytochrome oxidase I for 100 snails of the species A. arbustorum, three additional Arianta species, and nine outgroup taxa from five genera, in order to understand the phylogeographic history of the species. Despite some confounding artefacts that are likely due to introgression among the morphological forms, the resulting phylogeny shows that the depressed shell is plesiomorphic, while the globular shell is derived. Moreover, their disparate histories suggest that the depressed shell variety survived the glaciations in pockets of alpine refugia, while the globular shell variety recolonised the alpine environment post-glacially.

The development and evolution of left-right asymmetry in invertebrates: Lessons from Drosophila and snails

The unique nature of body handedness, which is distinct from the anteroposterior and dorsoventral polarities, has been attracting growing interest in diverse biological disciplines. Recent research progress on the left‐right asymmetry of animal development has focused new attention on the mechanisms underlying the development and evolution of invertebrate handedness. This exploratory review of currently available information illuminates the prospective value of Drosophila and pulmonate snails for innovative new research aimed at elucidating these mechanisms. For example, findings in Drosophila and snails suggest that an actin filament–dependent mechanism may be evolutionarily conserved in protostomes. The polarity conservation of primary asymmetry across most metazoan phyla, which visceral handedness represents, indicates developmental constraint and purifying selection as possible but unexplored mechanisms. Comparative studies using Drosophila and snails, which have the great advantages of using genetic and evolutionary approaches, will accelerate our understanding of the mechanisms governing the conservation and diversity of animal handedness. Developmental Dynamics 237:3497–3515, 2008.

SELECTIVE INCREASE OF A RARE HAPLOTYPE IN A LAND SNAIL HYBRID ZONE

In hybrid zones, which are regions where genetically differentiated populations of organisms meet and produce hybrids, allozyme studies have often revealed unexpected alleles. The cause of this ‘hybrizyme’ or ‘rare allele’ phenomenon has been elusive, although it has been variously ascribed to natural selection or increased mutation rates. If the latter hypothesis is correct, selectively neutral markers should demonstrate increased variability in contrast to expressed markers such as allozymes. In this study, we screened selectively neutral variation in an intron of the calmodulin (CaM) gene in a hybrid zone between two subspecies of the Greek land snail Albinaria hippolyti. In previous allozyme studies, this hybrid zone has been shown to exhibit the rare allele phenomenon. We used a variant of the single–strand conformational polymorphism (SSCP) technique to detect seven haplotypes in both parental taxa. In the zone, one of these occurs at unexpectedly high frequencies. Since no additional mutants were found, we concluded that this is the result of selection.

Whole-Body Enantiomorphy and Maternal Inheritance of Chiral Reversal in the Pond Snail Lymnaea stagnalis

Sinistral and dextral snails have repeatedly evolved by left-right reversal of bilateral asymmetry as well as coiling direction. However, in most snail species, populations are fixed for either enantiomorph and laboratory breeding is difficult even if chiral variants are found. Thus, only few experimental models of chiral variation within species have been available to study the evolution of the primary asymmetry. We have established laboratory lines of enantiomorphs of the pond snail Lymnaea stagnalis starting from a wild population. Crossing experiments demonstrated that the primary asymmetry of L. stagnalis is determined by the maternal genotype at a single nuclear locus where the dextral allele is dominant to the sinistral allele. Field surveys revealed that the sinistral allele has persisted for at least 10 years, that is, about 10 generations. The frequency of the sinistral allele showed large fluctuations, reaching as frequent as 0.156 in estimate under the assumption of Hardy-Weinberg equilibrium. The frequency shifts suggest that selection against chiral reversal was not strong enough to counterbalance genetic drift in an ephemeral small pond. Because of the advantages as a model animal, enantiomorphs of L. stagnalis can be a unique system to study aspects of chirality in diverse biological disciplines.

Cryptic, adaptive radiation of endoparasitic snails: sibling species of Leptoconchus (Gastropoda: Coralliophilidae) in corals

Coral reefs are renowned as complex ecosystems with an extremely large biodiversity. Parasite-host relationships contribute substantially to this, but are poorly known. We describe the results of a study in which approximately 60,000 corals were searched for parasitic Leptoconchus snails (Gastropoda: Coralliophilidae) in Indo-West Pacific waters of Egypt, the Maldives, Thailand, Palau and Indonesia. We discovered an adaptive radiation of 14 snail species, each of which lives in species-specific association with one or more of 24 mushroom coral species. The 14 snail species are described as new to science under the names Leptoconchus inactiniformis sp. nov., L. inalbechi sp. nov., L. incrassa sp. nov., L. incycloseris sp. nov., L. infungites sp. nov., L. ingrandifungi sp. nov., L. ingranulosa sp. nov., L. inlimax sp. nov., L. inpileus sp. nov., L. inpleuractis sp. nov., L. inscruposa sp. nov., L. inscutaria sp. nov., L. intalpina sp. nov., and L. massini sp. nov. Their separation is based on indisputable molecular differences, whereas the rudimentary shell characters or impoverished anatomical details do not allow identification. The coral hosts also serve to distinguish the snail species, as none of the former was found to contain more than one of the latter. The complexity of coral reefs is still underrated, as is shown here by the application of DNA taxonomy as an indispensable approach to unravel cryptic radiations, which must be known in order to understand the functioning of the ecosystem.

The 'rare allele phenomenon' in a ribosomal spacer.

We describe the increased frequency of a particular length variant of the internal transcribed spacer 1 (ITS‐1) of the ribosomal DNA in a hybrid zone of the land snail Albinaria hippolyti. The phenomenon that normally rare alleles or other markers can increase in frequency in the centre of hybrid zones is not new. Under the term ‘hybrizyme’ or ‘rare allele’ phenomenon it has been recorded in many organisms and different genetic markers. However, this is the first time that it has been found in a multicopy locus. On the one hand, the pattern fits well with the view that purifying selection in hybrid populations works on many loci across the genome and should thus have its effect on many independent molecular markers. On the other hand, the results are puzzling, given that the multiple copies of rDNA are not expected to respond in unison. We suggest two possible explanations for these conflicting observations.

Parallel evolution of an sAat -‘hybrizyme’ in hybrid zones in Albinaria hippolyti (Boettger)

Populations of the land snail Albinaria hippolyti from Crete were sampled across two hybrid zones separating A. h. aphrodite from A. h. holtzi and A. h. harmonia and studied by means of starch gel electrophoresis. At both sites, frequencies of an otherwise rare allele of sAat reached 0.70 and 0.18, respectively, in the centres of the hybrid zones. It is argued that the allele is deleterious and that it is maintained in the zones by a balance between elevated mutation rates (at least 1.4 × 10−4 and 3.2 × 10−4, respectively) and selection (at least 2 × 10−4 and 1.8 × 10−3, respectively). The observed parallellism may be the result of constraints on the numbers of metabolically active sAat variants possible.

Chiral Speciation in Terrestrial Pulmonate Snails

On the basis of data in the literature, the percentages of dextral versus sinistral species of snails have been calculated for western Europe, Turkey, North America (north of Mexico), and Japan. When the family of Clausiliidae is represented, about a quarter of all snail species may be sinistral, whereas less than one per cent of the species may be sinistral where that family does not occur. The number of single-gene speciation events on the basis of chirality, resulting in the origin of mirror image species, is not closely linked to the percentage of sinistral versus dextral species in a particular region. Turkey is nevertheless exceptional by both a high percentage of sinistral species and a high number of speciation events resulting in mirror image species. Shell morphology and genetic background may influence the ease of chirality-linked speciation, whereas sinistrality may additionally be selected against by internal selection. For the Clausiliidae, the fossil record and the recent fauna suggest that successful reversals in coiling direction occurred with a frequency of once every three to four million years.

INTERNAL SELECTION AGAINST THE EVOLUTION OF LEFT–RIGHT REVERSAL

Among metazoan species, left–right reversals in primary asymmetry have rarely gone to fixation. This suggests that a general mechanism suppresses the evolution of polarity reversal. Most metazoans appear externally symmetric and reproduce by external fertilization or copulation with genitalia located in the midline. Thus, reversal should generate little exogenous disadvantage when interacting with the external environment or in mating with the common wild‐type. Accordingly, an endogenously caused fitness reduction may be responsible for the general absence of reversed species. However, how this selection operates is little understood. Phenotypic changes associated with reversal are usually inseparable from zygotic pleiotropy. By exploiting hermaphroditism and the maternal inheritance of left–right polarity, we generated dextral and sinistral snails that share the same zygotic genotype. Before hatching, these sinistrals developed lethal morphological anomalies more frequently than dextrals. Their shell shape at maturity differed from the mirror image of the dextral shell. These interchiral differences demonstrate pleiotropy in maternal effects of the polarity or linked genes. Variation in interchiral differences between parental crosses suggests the presence of epistatic variation in relative performance of sinistrals. Our results show that internal selection operates against polarity reversal, and we suggest that this is due to changes in blastomere configuration.

Islands from a Snail's Perspective

Islands are broadly defined as inhabitable areas surrounded by a hostile environment, which makes island a relational term. The various kinds of islands from a snail’s perspective are listed, with examples of species occurring in those places. Isolation may be brought about by surrounding water, ice, or a variety of other ecological factors. Palaeoislands have existed in the geological past but ended their existence afterwards, for example by a lowering of sea level. Since there are by definition no contact zones between islands, a species or subspecies concept based on reproductive isolation under natural circumstances cannot be applied there. It is concluded still that archipelagos constitute the most important reservoir of gastropod diversity, with a high degree of endemism. Some predictions of evolutionary theory about speciation and adaptation are tested in an island setting, with also palaeoislands taken into account. Allopatry may result in differentiation and speciation, but whether, when, and how this happens differs considerably among the taxa. The adaptation to a particular island habitat, for example a cave, may follow quite different routes, probably because of the random mutations that enable the process. W. Renema (ed.), Biogeography, Time, and Place: Distributions, Barriers, and Islands, 347–363