ACCELERATED EVOLUTION OF LAND SNAILS MANDARINA IN THE OCEANIC BONIN ISLANDS: EVIDENCE FROM MITOCHONDRIAL DNA SEQUENCES

An endemic land snail genus Mandarina of the oceanic Bonin (Ogasawara) Islands shows exceptionally rapid evolution not only of morphological and ecological traits, but of DNA sequence. A phylogenetic relationship based on mitochondrial DNA (mtDNA) sequences suggests that morphological differences equivalent to the differences between families were produced between Mandarina and its ancestor during the Pleistocene. The inferred phylogeny shows that species with similar morphologies and life habitats appeared repeatedly and independently in different lineages and islands at different times. Sequential adaptive radiations occurred in different islands of the Bonin Islands and species occupying arboreal, semiarboreal, and terrestrial habitat arose independently in each island. Because of a close relationship between shell morphology and life habitat, independent evolution of the same life habitat in different islands created species possesing the same shell morphology in different islands and lineages. This rapid evolution produced some incongruences between phylogenetic relationship and species taxonomy. Levels of sequence divergence of mtDNA among the species of Mandarina is extremely high. The maximum level of sequence divergence at 16S and 12S ribosomal RNA sequence within Mandarina are 18.7% and 17.7%, respectively, and this suggests that evolution of mtDNA of Mandarina is extremely rapid, more than 20 times faster than the standard rate in other animals. The present examination reveals that evolution of morphological and ecological traits occurs at extremely high rates in the time of adaptive radiation, especially in fragmented environments.     

Long‐term stasis and short‐term divergence in the phenotypes of microsnails on oceanic islands

Phenotypic divergence is often unrelated to genotypic divergence. An extreme example is rapid phenotypic differentiation despite genetic similarity. Another extreme is morphological stasis despite substantial genetic divergence. These opposite patterns have been viewed as reflecting opposite properties of the lineages. In this study, phenotypic radiation accompanied by both rapid divergence and long‐term conservatism is documented in the inferred molecular phylogeny of the micro land snails Cavernacmella (Assimineidae) on the Ogasawara Islands. The populations of Cavernacmella on the Sekimon limestone outcrop of Hahajima Island showed marked divergence in shell morphology. Within this area, one lineage diversified into types with elongated turret shells, conical shells and flat disc‐like shells without substantial genetic differentiation. Additionally, a co‐occurring species with these types developed a much larger shell size. Moreover, a lineage adapted to live inside caves in this area. In contrast, populations in the other areas exhibited no morphological differences despite high genetic divergence among populations. Accordingly, the phenotypic evolution of Cavernacmella in Ogasawara is characterized by a pattern of long‐term stasis and periodic bursts of change. This pattern suggests that even lineages with phenotypic conservatism could shift to an alternative state allowing rapid phenotypic divergence.     

Disentangling the effects of intraspecies variability,phylogeny, space,and climate on the evolution of shell morphology in endemic Greek land snails of the genus Codringtonia

Extensive variation in land snail shell morphology has been widely documented, although few studies have attempted to investigate the ecological and evolutionary drivers of this variation. Within a comparative phylogenetic framework, we investigated the temporal and spatial evolution of the shell morphology of the Greek endemic land snail genus Codringtonia. The contribution of both inter‐ and intraspecies shell differentiation in the overall shell variability is assessed. The effect of climate, space, and evolutionary history on the shell variability was inferred using a variance partitioning framework. For Codringtonia species, intraspecies divergence of shell traits contributes substantially to the overall shell variability. By decomposing this variability, it was shown that the overall shell size of Codringtonia clades is phylogenetically constrained, related to early speciation events, and strongly affected by large‐scale spatial variability (latitudinal gradient). The effect of climate on shell size cannot be disentangled from phylogeny and space. Shell and, to a larger extent, aperture shape are not phylogenetically constrained, and appear to be mostly related to conspecific populations divergence events. Shell shape is substantially explained by both climate and space that greatly overlap. Aperture shape is mainly interpreted by medium to small‐scale spatial variables. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 796–813.     

Evidence for a multi‐peak adaptive landscape in the evolution of trophic morphology in terapontid fishes

Using the Australian marine‐freshwater terapontid fishes as a model system, we examined the role of dietary phenotypic optima in an adaptive macro‐evolutionary landscape. Comparative modelling relying on both a priori and data‐driven identification of selective regimes suggested multi‐peak models as best describing much of the dietary phenotypic landscape of terapontids. Both approaches identified common phenotypic optima for different lineages of marine and freshwater herbivores, and minimal differentiation between carnivores and omnivores, irrespective of their phylogenetic relationships, as the model best describing morphological evolution. Significant correlations also existed between these phenotypic axes and proportions of non‐animal dietary items in species’ diets. While simulation results provided evidence for a multi‐peak adaptive landscape in the evolution of trophic morphology in terapontids, they could not rule out chance convergence in these adaptive peaks. However, they do provide scope for identifying areas for more detailed, functionally specific study of phenotypic convergence in herbivorous terapontid trophic habits. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 623–634.     

Comparative morphology of pond,stream and phytotelm‐dwelling tadpoles of the South American Redbelly Toads (Anura: Bufonidae: Melanophryniscus)

We present a comprehensive review of larval morphology in the Neotropical toad genus Melanophryniscus. The taxa studied included 23 species with representatives of recognized phenetic groups and different larval ecomorphological guilds: pond, stream, and phytotelm‐dwelling tadpoles. Their external morphology variation is congruent with current phenetic arrangement based on adult features, but also reflects the habitat where larvae develop. Lotic tadpoles (i.e. M. tumifrons group and M. krauczuki) in general exhibit a more depressed body, a longer tail with lower fins, and larger oral discs than lentic forms (i.e. M. stelzneri group, M. moreirae, M. sanmartini, and M. langonei). Despite their peculiar, confined microhabitat, phytotelm larvae do not diverge markedly from non‐arboreal species. The distinctive features of all species are the presence of a pineal end organ and the placement of the intestinal reversal point at the left of the abdomen in typical larval stages. The buccal cavity and musculoskeletal anatomy are quite conserved between species, yet some characteristics differ from those of other bufonids. The presence of one pair of subhyoid muscles is apparently an exclusive trait of Melanophryniscus among Bufonidae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 417–441.     

Molecular phylogenetics and morphological variation reveal recent speciation in freshwater mussels of the genera Arcidens and Arkansia (Bivalvia: Unionidae)

Intra‐ and interspecific morphological variation due to both phenotypic plasticity and evolutionary convergence hinder the work of taxonomists and lead to over‐ and underestimates of species richness. Nevertheless, most species on Earth are recognized solely based on morphological characters. We used molecular phylogenetic and morphometric techniques to examine two freshwater mussel species. One is common and widespread, while the other is imperiled and endemic to the Interior Highlands of the USA. Phylogenetic and molecular clock analyses revealed that divergence of Arcidens confragosus and Arkansia wheeleri is small and relatively recent. Divergence in these and other taxa is probably due to isolation of streams in the Interior Highlands. Morphometric analyses showed distinct shell shapes using traditional morphometrics, but not through geometric morphometrics. Outlined shell shapes are indistinguishable; geometric morphometrics could not capture a three‐dimensional component. Our analyses support the validity of these two species as congeners, with the nomen Arcidens (Simpson 1900) having priority. Because shell morphologies are both heritable and environmentally determined, our study emphasizes the importance of considering both molecular and morphometric analyses for identification of freshwater molluscs of conservation concern. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 535–545.     

Layers of contingency shroud pervasive ecological divergence in a local radiation of land snails

The predictability of evolution depends on the roles that selection and historical contingency play in determining its outcomes, but the relative importance of these evolutionary mechanisms has attracted considerable debate. One view is that historical events have such a profound impact on the genetic structure of populations that patterns of phenotypic evolution are essentially unpredictable. The opposing view is that selection is so powerful that evolutionary change is primarily deterministic, and thus highly predictable. By controlling for the effects of phylogeny, geographic location and habitat, this study examined the relative roles of contingency and determinism in a local radiation of land snails, genus Rhagada, in a continental archipelago. Informed by previous studies on a single island, which revealed a strong association between low‐spired shells and rocky habitats, 28 population pairs were sampled in directly adjoining rocky and spinifex plain habitats. When considered in their respective pairs, the effect of habitat was remarkably consistent, with lower‐spired shells observed in the rocky habitat in 24 of the comparisons. However, when analyzed outside the context of those pairs, the association was obscured by broad variation in shell shape within habitat types and among lineages. These results reveal the complex nature of a morphological radiation; while the pattern of ecological divergence is highly predictable at the scale that selection acts, deterministic evolution is largely obscured by phylogenetic and population history.     

The specialisation of the third metacarpal and hand in arboreal frogs: Adaptation for arboreal habitat?

Skeletal morphology is directly associated with habitat characteristics. To investigate the arboreal adaptation, we studied the osteological morphology of the forelimbs and the third metacarpals of arboreal frogs (Rhacophoridae and Hylidae) and non‐arboreal frogs (Bombinatoridae, Bufonidae, Megophryidae, Ranidae, and Microhylidae) using transparent specimens or X‐rays of skeletons. Our results revealed that the bony knob on the third metacarpal, which formed by a dilated and elongated lateral articular cartilage (AL) through endochondral ossification, occurred only in species of Rhacophorinae. The results of the phylogenetic comparative methods and correlation analysis strongly supported the conclusion that the bony knob is a phylogenetic independent evolution trait and had a significant correlation with the arboreal habitat. Furthermore, anatomical observation showed that a muscle adhered to the bony knob. Therefore, we speculated that the bony knob might act as an enlarged attachment point for larger or more musculatures to help with grasping. In addition, the relative length of the hand showed a significant difference between arboreal and non‐arboreal species (p = .007), suggesting that longer hands might be an arboreal adaptive trait. Overall, this study leads to a deeper understanding of the arboreal adaptation.     

Systematic revision of the arboreal snail Satsuma albida species complex (Mollusca: Camaenidae) with descriptions of 14 new species from Taiwan

The taxonomy of the endemic arboreal snail Satsuma albida species complex from Taiwan was unclear due to the animals' highly similar morphology, and their nocturnal and strict arboreal behaviour, leading to difficulties in collecting living specimens. This article is the first comprehensive comparative study on the systematics and taxonomy of this species complex using external morphology, anatomy of the reproductive system and molecular phylogeny. Consequently, two subspecies of S. albida are raised to species status, namely S. insignis and S. mollicula. Fourteen new species are also described. Fourteen of the 17 species showed polymorphism in banding pattern amongst populations and other species retained the whitish unity as seen in S. albida. Distributions of almost all taxa are geographically limited, with the exception of S. polymorpha sp. nov . The phylogeny of these species was reconstructed using 20 morphological characters and molecular data from the partial sequences of mtDNA CO1 and 16S rRNA genes, and the complete ITS2 sequence. The molecular phylogeny revealed three subclades (west, east and polymorpha clade) and revealed that these snails are monophyletic, originating from a ground‐dwelling ancestor. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 437–493.     

Genetic and morphological divergence between Littorina fabalis ecotypes in Northern Europe

Low dispersal marine intertidal species facing strong divergent selective pressures associated with steep environmental gradients have a great potential to inform us about local adaptation and reproductive isolation. Among these, gastropods of the genus Littorina offer a unique system to study parallel phenotypic divergence resulting from adaptation to different habitats related with wave exposure. In this study, we focused on two Littorina fabalis ecotypes from Northern European shores and compared patterns of habitat‐related phenotypic and genetic divergence across three different geographic levels (local, regional and global). Geometric morphometric analyses revealed that individuals from habitats moderately exposed to waves usually present a larger shell size with a wider aperture than those from sheltered habitats. The phenotypic clustering of L. fabalis by habitat across most locations (mainly in terms of shell size) support an important role of ecology in morphological divergence. A genome scan based on amplified fragment length polymorphisms (AFLPs) revealed a heterogeneous pattern of differentiation across the genome between populations from the two different habitats, suggesting ecotype divergence in the presence of gene flow. The contrasting patterns of genetic structure between nonoutlier and outlier loci, and the decreased sharing of outlier loci with geographic distance among locations are compatible with parallel evolution of phenotypic divergence, with an important contribution of gene flow and/or ancestral variation. In the future, model‐based inference studies based on sequence data across the entire genome will help unravelling these evolutionary hypotheses, improving our knowledge about adaptation and its influence on diversification within the marine realm.     

Under pressure: morphological and ecological correlates of bite force in the rock‐dwelling lizards Ouroborus cataphractus and Karusasaurus polyzonus (Squamata: Cordylidae)

Rock‐dwelling lizards are hypothesized to be highly constrained in the evolution of head morphology and, consequently, bite force. Because the ability to generate a high bite force might be advantageous for a species' dietary ecology, morphological changes in head configuration that allow individuals to maintain or improve their bite force under the constraint of crevice‐dwelling behaviour are to be expected. The present study addressed this issue by examining head morphology, bite force, and a number of dietary traits in the rock‐dwelling cordylid lizards Ouroborus cataphractus and Karusasaurus polyzonus. The results obtained show that O. cataphractus has a larger head and higher bite force than K. polyzonus. In K. polyzonus, head width, lower jaw length, and jaw closing‐in lever are the best predictors of bite force, whereas head height is the main determinant of bite force in O. cataphractus. Although the observed difference in bite force between the species does not appear to be related to dietary patterns or prey handling, the prey spectrum available for intake was greater in O. cataphractus compared to K. polyzonus. We discuss the influence of interspecific differences in anti‐predator morphology on head morphology and bite force in these rock‐dwelling species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 823–833.     

Interspecific interference competition alters habitat use patterns in two species of land snails

Focus has been placed on the relative importance of environmental heterogeneity and biological interactions on community structure. For land snails, abiotic factors have commonly been assumed to be the most important factors for shaping communities because resources for land snails are commonly not limiting, and because interspecific resource competition would not be strong enough to promote ecological divergence. However, clear divergence in habitat use is often observed among sympatric land snail species. Such an ecological divergence would be promoted not by resource competition but by other interactions such as aggression. To test this hypothesis, laboratory experiments were designed to explore aggressive behaviour in the land snails Euhadra quaesita and E. peliomphala and field surveys were conducted to examine their habitat use. In the laboratory experiments, we examined how the presence of one species affects the growth of the other species by supplying sufficient amounts of food and calcium for both species. Experimental trials showed that adult E. quaesita decreased the growth of E. peliomphala under constant diet conditions. In contrast, E. peliomphala did not affect the growth of E. quaesita. Because E. peliomphala was often attacked by E. quaesita and its shell was often eroded by E. quaesita gnawing, aggression by E. quaesita appears to be the primary factor reducing the growth of E. peliomphala. When the two species coexist in nature, adults and juveniles of E. quaesita are terrestrial, while those of E. peliomphala are arboreal. When these species occurred alone, E. quaesita were still terrestrial, but E. peliomphala were more terrestrial than when they were in sympatry. Our results suggest that habitat use of these species in the natural communities is affected by interspecific interference. These findings further suggest that the divergence of habitat use between the species can occur by aggression even in environments with unlimited resources.     

Species limits in molecular phylogenies: a cautionary tale from Australian land snails (Camaenidae: Amplirhagada Iredale, 1933)

By complementing two independent systematic studies published recently on the Western Australian land snail Amplirhagada, we compare levels of morphological variation in shells and genitalia with those in the mitochondrial markers cytochrome c oxidase (COI) and 16S to evaluate the utility of mtDNA markers for delimiting species. We found that penial morphology and mitochondrial divergence are generally highly consistent in delimiting species, while shells have little overall taxonomic utility in these snails. In addition to this qualitative correspondence, there is almost no overlap between intraspecific and interspecific genetic distances in COI, with the highest intraspecific and lowest interspecific distance being 6%. This value is twice the general level suggested as a DNA barcode threshold by some authors and higher than the best average found in stylommatophoran land snails. Although in Amplirhagada land snails DNA barcoding may provide meaningful information as a first‐pass approach towards species delimitation, we argue that this is due only to specific evolutionary circumstances that facilitated a long‐termed separate evolution of mitochondrial lineages along spatial patterns. However, because in general the amounts of morphological and mitochondrial differentiation of species depend on their evolutionary history and age, the mode of speciation, distributional patterns and ecological adaptations, and absence or presence of mechanisms that prevent gene flow across species limits, the applicability of DNA barcoding has to be confirmed by morphological studies for each single group anew. Based on evidence from both molecular and morphological markers, we describe six new species from the Bonaparte Archipelago and revise the taxonomy of a further two. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 337–362.     

Effects of habitat history and extinction selectivity on species-richness patterns of an island land snail fauna

Aim Local‐scale diversity patterns are not necessarily regulated by contemporary processes, but may be the result of historical events such as habitat changes and selective extinctions that occurred in the past. We test this hypothesis by examining species‐richness patterns of the land snail fauna on an oceanic island where forest was once destroyed but subsequently recovered. Location Hahajima Island of the Ogasawara Islands in the western Pacific. Methods Species richness of land snails was examined in 217 0.25 × 0.25 km squares during 1990–91 and 2005–07. Associations of species richness with elevation, current habitat quality (proportion of habitat composed of indigenous trees and uncultivated areas), number of alien snail species, and proportion of forest loss before 1945 in each area were examined using a randomization test and simultaneous autoregressive (SAR) models. Extinctions in each area and on the entire island were detected by comparing 2005–07 records with 1990–91 records and previously published records from surveys in 1987–91 and 1901–07. The association of species extinction with snail ecotype and the above environmental factors was examined using a spatial generalized linear mixed model (GLMM). Results The level of habitat loss before 1945 explained the greatest proportion of variation in the geographical patterns of species richness. Current species richness was positively correlated with elevation in the arboreal species, whereas it was negatively correlated with elevation in the ground‐dwelling species. However, no or a positive correlation was found between elevation and richness of the ground‐dwelling species in 1987–91. The change of the association with elevation in the ground‐dwelling species was caused by greater recent extinction at higher elevation, possibly as a result of predation by malacophagous flatworms. In contrast, very minor extinction levels have occurred in arboreal species since 1987–91, and their original patterns have remained unaltered, mainly because flatworms do not climb trees. Main conclusions The species‐richness patterns of the land snails on Hahajima Island are mosaics shaped by extinction resulting from habitat loss more than 60 years ago, recent selective extinction, and original faunal patterns. The effects of habitat destruction have remained long after habitat recovery. Different factors have operated during different periods and at different time‐scales. These findings suggest that historical processes should be taken into account when considering local‐scale diversity patterns.     

Does sympatry predict life history and morphological diversification in the Mexican livebearing fish Poeciliopsis baenschi?

Understanding why some species coexist and others do not remains one of the fundamental challenges of ecology. Although there is evidence to suggest that closely‐related species are unlikely to occupy the same habitat because of competitive exclusion, there are many cases where closely‐related species do co‐occur. Research comparing sympatric and allopatric populations of co‐occurring species provides a framework for understanding the role of phenotypic diversification in species coexistence. In the present study, we compare phenotypic divergence between sympatric and allopatric populations of the livebearing fish, Poeciliopsis baenschi. We focus on life‐history traits and body shape, comprising two sets of integrated traits likely to diverge in response to varying selective pressures. Given that males and females can express different phenotypic traits, we also test for patterns of divergence among sexes by comparing size at maturity and sexual dimorphism in body shape between males and females in each population type. We take advantage of a natural experiment in western Mexico where, in some locations, P. baenschi co‐occur with a closely‐related species, Poeciliopsis turneri (sympatric populations) and, in other locations, they occur in isolation (allopatric populations). The results obtained in the present study show that sympatric populations of P. baenschi differed significantly in life‐history traits and in body shape compared to their allopatric counterparts. Additionally, males and females showed different responses for size at maturity in sympatric conditions versus allopatric conditions. However, the amount of sexual dimorphism did not differ between sympatric and allopatric populations of P. baenschi. Hence, we conclude that not all traits show similar levels of phenotypic divergence in response to sympatric conditions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 608–618.     

Morphology and aestivation behaviour in some Madagascan acavid land snails

Nine species of Madagascan acavid land snails were compared in a phylogenetic context. The two most plesiomorphic, Clauator johnsoni and C. moreleti, differ from the others by their high-spired shells, short tentacles, short tails, long necks, and crawling mode of hitching the shell along the ground. In the seven more apomorphic species, the crawling mode is smooth, with the shell resting on the tail, and the relative lengths of tail and shell correlate significantly. Among these seven species, three pairs of closest relatives (Helicophanta petiti and H. uesicalis, H. farafanga and H. souuerbiana, Ampeltta decaryi and A. julii) show evidence of phylogenetic constraints on ranked shell size. Aestivation site (as tentatively inferred from rare data) does not correlate with shell shape or size: burrowers have H/D = 2.7 to 0.6 and D = 70 to 25 mm; arboreals have H/D = 0.8 to 0.5 and D = 70 to 30 mm; the species with both the highest spire and the smallest diameter (C. moreleti) is neither a burrower or an arboreal, but stays on the ground surface. Inferred aestivation sites are randomly distributed phylogenetically. Climate shows no correlation, except that the arborcals are only from humid to wet regimes. Uniform shell colouration occurs only in burrowers (C. johnsoni, H. petiti, H. uestcalis), but disruptive shell colouration occurs in all others, including burrowers (H. farafanga, A. decaryt), ground-surface aestivators (C. moreleti), arboreals (H. souuerbiana, A. julii), and semi-arboreals (Ampefita subfunebris). Among all nine species, burrowers have significantly thicker shells (than their close relatives of similar size), wider bodies, and longer snouts than non-burrowers (H. souuerbiana is exceptional in being arboreal despite its huge size and in having the broad foot and snout of a burrower). Thus, although there is some evidence for phylogenetic constraints, natural selection for aestivation and crawling behaviours seems to have dominated the evolution of external body morphology and of shell thickness (but not shell size and shape) in these snails.     

Microgeographic differentiation among closely related species of Biomphalaria (Gastropoda: Planorbidae) from the Andean Altiplano

Direct development and water dependence entail limited vagility in freshwater fauna. In these organisms, the population structure is probably linked to restrictions imposed by the habitat. In this study we investigate the relative contribution of processes stimulating the divergence of populations of Biomphalaria costata (Biese, 1951) and Biomphalaria crequii (Courty, 1907), two freshwater snails occurring in two contiguous and fragmented closed basins from the Andean Altiplano using mitochondrial DNA (cytochrome c oxidase subunit I) sequences, shell morphometric and radular morphology. In order to clarify the species boundaries, a third allopatric species was included: Biomphalaria aymara Valdovinos & Stuardo, 1991. Molecular analyses recovered two distinct clades: one composed of B. aymara from the Isluga swamps and B. costata from Spring 1 in Salar de Carcote, the single spring occupied by this species, and another integrated by snails from 12 springs spread across the Salar de Carcote and the Salar de Ascotán assigned to B. crequii, originally described from the Salar de Ascotán. Unlike shell morphometrics, radular morphology was informative for distinguishing these species. The division of the lineages occurred in the Late Pleistocene. A subclade that includes snails from the southernmost springs in Salar de Ascotán suggests fragmentation of the distribution of B. crequii associated with landscape discontinuities. In addition to microvicariance signals, the private haplotypes scattered around both salt spans show that close‐range dispersal is a common biogeographic process in this species. As evolutionary units, the single isolated and restricted population of B. costata has a high priority for conservation. © 2013 The Linnean Society of London     

Allopatry and overlap in a clade of snails from mangroves and mud flats in the Indo‐West Pacific and Mediterranean (Gastropoda: Potamididae: Cerithideopsilla)

Cerithideopsilla is a genus of potamidid snails found in high abundance on sedimentary intertidal flats and beneath mangrove trees on continental shores in the tropical and subtropical Indo‐West Pacific region and Mediterranean Sea. Taxonomic revisions have recognized four species, but recent molecular studies have hinted at a higher diversity. Here, we analyse 377 individuals sampled from across the known range and use a combination of molecular phylogenetic (mitochondrial COI and 16S rRNA, and nuclear 28S rRNA genes), statistical (generalized mixed Yule‐coalescent GMYC method) and morphological (shell form) criteria to delimit 16 species. These form four species groups, corresponding with the traditionally recognized species C. alata, C. ‘djadjariensis’ (for which the valid name is C. incisa), C. cingulata and C. conica. Distribution maps were compiled using museum specimens identified by diagnostic shell characters. In combination with the molecular phylogenetic trees, these suggest an allopatric speciation mode, with diversification centred on the East Asian coastline and northern Australia, and a pronounced gap in the ‘eastern Indonesian corridor’, an area of low oceanic productivity. There is, however, frequently geographical overlap between sister species and we suggest from several sources of evidence (e.g. presence of C. conica in isolated saline lakes 900 km from the sea) that post‐speciation transport by migratory birds has occurred. Nine of the 16 species occur between the Gulf of Tonkin and Hong Kong, so southern China is significant for both the evolution and conservation of Cerithideopsilla species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 212–228.     

Extreme,continuous variation in an island snail: local diversification and association of shell form with the current environment

On Rosemary Island, a small continental island (11 km²) in the Dampier Archipelago, Western Australia, snails of the genus Rhagada have extremely diverse morphologies. Their shells vary remarkably in size and shape, with the latter ranging from globose to keeled‐flat, spanning the range of variation in the entire genus. Based primarily on variation in shell morphology, five distinct species are currently recognized. However, a study of 103 populations has revealed continuity of shell form within a very closely‐related group. A phylogenetic analysis of specimens from Rosemary Island, and other islands in the Dampier Archipelago, indicates that much of the morphological variation has evolved on the island, from within a monophyletic group. Within the island, snails with distinct shell morphologies could not be distinguished based on variation in mitochondrial DNA or their reproductive anatomy. The shell variation is geographically structured over a very fine scale, with clines linking the extreme forms over distances less than 200 m. Although there is no evidence that the different forms have evolved in isolation or as a consequence of drift, there is a strong association between keeled‐flat shells and rocky habitats, suggesting that shell shape may be of adaptive significance. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 756–769.     

Evidence of introgressive hybridization between the morphologically divergent land snails Ainohelix and Ezohelix

Hybridization between different taxa is likely to take place when adaptive morphological differences evolve more rapidly than reproductive isolation. When studying the phylogenetic relationship between two land snails of different nominal genera, Ainohelix editha and Ezohelix gainesi, from Hokkaido, Japan, using nuclear internal transcribed spacer and mitochondrial 16S ribosomal DNA, we found a marked incongruence in the topology between nuclear and mitochondrial phylogenies. Furthermore, no clear association was found between shell morphology (which defines the taxonomy) and nuclear or mitochondrial trees and morphology of reproductive system. These patterns are most likely explained by historical introgressive hybridization between A. editha and E. gainesi. Because the shell morphologies of the two species are quite distinct, even when they coexist, the implication is that natural selection is able to maintain (or has recreated) distinct morphologies in the face of gene flow. Future studies may be able to reveal the regions of the genome that maintain the morphological differences between these species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 77–95.