Life on the edge: a hybrid zone in AZbinaria hippolyti (Gastropoda: Clausiliidae) from Crete

The hybrid zone between the Greek land snails Albinaria hippolyti aphrodite and Albinaria hippolyti harmonia runs exactly along the edge of a gorge. At two localities, we analysed patterns of variation in three enzyme loci, six conchological characters, and two anatomical characters across the zone. Clines in the various characters are largely coincident and concordant. They are narrow (approximately 30 m wide) where the ecological transition is abrupt, and wider (approximately 120 m wide) where a valley cuts into the gorge, making the environmental change more gradual. In spite of the obvious correlation with an ecological transition, we do not believe the zone is maintained by a selection gradient. Instead, direct and indirect evidence for hybrid disadvantage and evidence for directional dispersal at the edge of the gorge suggest that it is a tension zone, held at the ecotone by differential dispersal of at least one of the parental types. We speculate on the possibility that reinforcement may act in the hybrid zone. Our data on dispersal and reproductive isolating mechanisms lead us to believe that reinforcement may only be possible at localities where the hybrid zone is very narrow.     

Systematic revision of the genus Everettia Godwin‐Austen, 1891 (Mollusca: Gastropoda: Dyakiidae) in Sabah,northern Borneo

The species of the snail genus Everettia in the Malaysian state of Sabah are superficially similar and difficult to distinguish by their shells. This paper presents new data on the taxonomy and distribution of Everettia in Sabah that have accumulated since the revision by Godwin‐Austen in 1891. By using morphological and molecular phylogenetic approaches, we reveal at least seventeen species of Everettia in Sabah, of which eleven are new to science, namely: Everettia layanglayang sp. nov. , Everettia lapidini sp. nov. , Everettia paulbasintali sp. nov. , Everettia occidentalis sp. nov. , Everettia jasilini sp. nov. , Everettia safriei sp. nov. , Everettia interior sp. nov. , Everettia jucundior sp. nov. , Everettia planispira sp. nov. , Everettia monticola sp. nov. , and Everettia dominiki sp. nov. , and one new subspecies, namely, Everettia corrugata williamsi ssp. nov. Phylogenetic analysis of mitochondrial COI and 16S, and nuclear ITS‐1 sequences demonstrates the monophyly of most of the morphologically well‐defined species. Our results show that certain aspects of classical morphology‐based taxonomy for Everettia species, especially with regard to the unique combination of shell surface sculptures, animal head colour, and mantle pigmentation, are solid. A dichotomous key to the Sabah species and subspecies of Everettia is provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 515–550.     

Phylogeography of the land snail Albinaria hippolyti (Pulmonata: Clausiliidae) from Crete, inferred from ITS-1 sequences

The polytypic Cretan land snail Albinaria hippolyti has a range that is partly fragmented and partly subdivided by hybrid zones. For this reason, it has served as a model species for investigating speciation and radiation in Mediterranean Clausiliidae. The first internal transcribed spacer ( ITS-1 ) of the nuclear ribosomal DNA was sequenced in 20 populations of A. hippolyti and phylogenetically analysed using maximum parsimony. We employed a novel method involving logarithmic weighting of gaps and topological constraints based on bootstrap values. The resulting phylogeography suggests that the species has undergone a recent cycle of range expansion and range reduction. Speciation cannot be linked to major geological vicariance events in the Miocene and Pliocene, as has been suggested previously. The subspecies A. h. arthuriana appears unrelated to other A. hippolyti subspecies, which supports recent suggestions, based on morphology, to regard it as a separate species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 317–326.     

Morphological and molecular phylogenetics in the genus Leptopilina (Hymenoptera: Cynipoidea: Eucoilidae)

We have conducted cladistic analyses of the genus Leptopilina , a group of Drosophila parasitoids studied intensively by (behavioural) ecologists. Twenty-three morphological characters were scored in ten Leptopilina and two outgroup species. At the same time, DNA sequences for the second ribosomal internal transcribed spacer (ITS2) were gathered for eight Leptopilina and one outgroup species. Both data sets yielded phylogenetic trees which were largely compatible. A 'total evidence' analysis resulted in a single tree that provides a relatively robust phylogenetic reconstruction of the group, which may serve as a basis for historically interpreting the distribution of ecological and behavioural traits.     

Reproductive character displacement by inversion of coiling in clausiliid snails (Gastropoda, Pulmonata)

In land snails, a change in the direction of coiling, being associated with a shift in the position of the genital apparatus, may act as a barrier against hybridization between sympatric species. Putative reproductive character displacement by an inversion in chirality has been reported in only a few land snails, based on observations in the field and interbreeding experiments. In this study, we present a new case of possible reproductive character displacement in the direction of coiling, in the clausiliid snail Isabellaria dextrorsa . This species is dextral, in contrast with its nearest relatives, including I. torifera and I. lophauchena , which share plesiomorphic sinistral coiling. Whereas I. dextrorsa occurs in sympatry and even syntopically with I. lophauchena throughout most of its range, the sinistral species have a mosaic distribution. Phylogenetic analyses of mitochondrial cytochrome c oxidase subunit I (COI) sequences demonstrated that I. dextrorsa constitutes a clade with I. torifera . In this clade, a shift in coiling direction occurred at least twice, maybe triggered by the presence of a sympatric congeneric sinistral species. The analyses separated the sequences of all I. dextrorsa samples from those of sympatric and syntopic I. lophauchena samples. The failure to demonstrate gene flow between these species is consistent with the hypothesis of genetic isolation by reproductive character displacement.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 155–164.     

Small‐scale genetic structuring in a tropical cave snail and admixture with its above‐ground sister species†

We analyse the phylogeographical structure in the cave snail Georissa filiasaulae Haase & Schilthuizen, 2007 (Gastropoda: Hydrocenidae) and its above‐ground sister species G. saulae (van Benthem‐Jutting, 1966) at limestone outcrops in Sabah, Malaysian Borneo. Morphometric and 16S mitochondrial DNA data for some 220 individuals reveal strong morphological differentiation, despite ongoing unidirectional gene flow from the epigean into the hypogean environment, strong, small‐scale genetic structuring within the cave and underground dispersal between caves that were previously thought to be isolated. We discuss these results – which constitute the first phylogeographical analysis of a terrestrial cave snail – in the light of speciation in cave organisms and across ecotones in general. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 727–740.     

Reproductive isolation in snails of the genus Albinaria (Gastropoda: Glausiliidae)

Parapatric, morphologically characterized forms of the Mediterranean snail genus Albinaria have traditionally been regarded as biological species. Recently, this view has been challenged on the basis of small interspecific genetic distances and cross-breeding experiments. In order to test whether traditionally recognized species are reproductively isolated lineages, two cases of syntopy were analysed by means of isozyme electrophoresis. The data suggest that complete reproductive isolation is present. In one of the two cases, ecological differentiation on a small spatial scale was observed.