Clarification of the status of the type series and of the holotype of Cyclophorus (Glossostylus) koboensis Godwin-Austen, 1915 (Mollusca, Caenogastropoda, Cyclophoridae) in Nantarat et al. (2014).

Here, the clarification of the "type" status for Cyclophorus (Glossostylus) koboensis Godwin-Austen, 1915 catalogued by Nantarat et al. (2014) is corrected and briefly discussed.

Notes on the sinistral helicoid snail Bertia cambojiensis (Reeve, 1860) from Vietnam (Eupulmonata, Dyakiidae).

Since the time of the original description there have been no precise locality records in Cambodia of Bertia cambojiensis (Reeve, 1860) and it was believed to be extinct. In 2012, a joint Natural History Museum survey with Vietnamese colleagues rediscovered living populations of this huge sinistral helicoid snail in a protected area of southern Vietnam. The genitalia and radula morphology are re-assessed and type specimens of all recognised congeners are figured herein. The unique morphological characters of this species are a small and simple penis, well-developed amatorial organ complex that incorporates four amatorial organ ducts, a short gametolytic organ complex and spiked papilla, and radula morphology with unicuspid teeth. The type locality of B. cambojiensis, which has been contentious, is determined here to be in the vicinity of 'Brelum', Vietnam, near the border with Cambodia. In addition, the nucleotide sequences of barcoding genes COI, 16SrRNA and 28S fragments were provided for further comparison.

Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species.

Limestone karsts can form terrestrial habitat islands for calcium-dependent organisms. In Vietnam, many karst habitats are threatened, while their rich biodiversity is still far from being thoroughly explored. Given that conservation of karst biota strongly relies on correct species identification, the presence of undetected cryptic species can pose severe problems. The present study focuses on cryptic diversity among karst-inhabiting land snails of the genus Cyclophorus in northern Vietnam, where specimens with a similar shell morphology have been reported from various regions. In order to examine the diversity and evolutionary history of this "widespread morphotype", we generated a Bayesian phylogeny based on DNA sequence data. Automatic Barcode Gap Discovery (ABGD) and the Bayesian implementation of the Poisson tree processes model (bPTP) contributed to species delimitation and analyses of shell shape and size aided the morphological characterisation of individual species. We found that the examined specimens of the widespread morphotype did not form a single monophyletic group in the phylogeny but clustered into several different clades. We delimited nine different species that develop the widespread morphotype and described four of them as new. Processes of convergent evolution were probably involved in the origin of the delimited species, while their generally allopatric distribution could result from interspecific competition. Our findings indicate ongoing processes of speciation and a potential case of morphological character displacement. The high degree of morphological overlap found among the species underlines the importance of DNA sequence data for species delimitation and description in the genus Cyclophorus. Given the findings of the present study and the high potential that as yet undiscovered cryptic taxa have also evolved in other groups of karst-inhabiting organisms, we argue for a systematic and efficient detection and description of Vietnam's karst biodiversity to provide a solid basis for future conservation planning.

Phylogenetics and species delimitations of the operculated land snail Cyclophorus volvulus (Gastropoda: Cyclophoridae) reveal cryptic diversity and new species in Thailand.

Recent conceptual and practical advances in phylogenetic species delimitation have enabled progressively robust biodiversity studies. Delimiting species in widespread taxa is an intriguing problem; the edible operculated land snail Cyclophorus volvulus (Müller, 1774) is a good example since it shows a high degree of shell and color variation along with a widespread distribution throughout Thailand. Taxonomic boundaries for C. volvulus were examined and clarified using a combined morphological and phylogenetic approach, the latter of which was based on both nuclear and mitochondrial gene sequences. Moreover, three species delimitation analyses were applied: Poisson tree processes (PTP), automatic barcode gap discovery (ABGD), and generalized mixed Yule-coalescent (GMYC). All phylogenetic trees revealed that C. volvulus was polyphyletic and comprised of three clades that coincided with their geographic distribution. The three species delimitation analyses concurred with the phylogenies and formed at least three groups. According to the results, C. volvulus s.l., as currently recognized, consists of three distinct species in Thailand: C. volvulus s.s., C. occultus sp. nov., and C. borealis sp. nov., which are described herein. Moreover, several of these highly distinct C. volvulus evolutionarily significant units (ESU) are likely to require urgent conservation attention.

Competition matters: Determining the drivers of land snail community assembly among limestone karst areas in northern Vietnam.

The insular limestone karsts of northern Vietnam harbor a very rich biodiversity. Many taxa are strongly associated with these environments, and individual species communities can differ considerably among karst areas. The exact processes that have shaped the biotic composition of these habitats, however, remain largely unknown. In this study, the role of two major processes for the assembly of snail communities on limestone karsts was investigated, interspecific competition and filtering of taxa due to geographical factors. Communities of operculate land snails of the genus Cyclophorus were studied using the dry and fluid-preserved specimen collections of the Natural History Museum, London. Phylogenetic distances (based on a Bayesian analysis using DNA sequence data) and shell characters (based on 200 semilandmarks) were used as proxies for ecological similarity and were analyzed to reveal patterns of overdispersion (indicating competition) or clustering (indicating filtering) in observed communities compared to random communities. Among the seven studied karst areas, a total of 15 Cyclophorus lineages were found. Unique communities were present in each area. The analyses revealed phylogenetic overdispersion in six and morphological overdispersion in four of seven karst areas. The pattern of frequent phylogenetic overdispersion indicated that competition among lineages is the major process shaping the Cyclophorus communities studied. The Coastal Area, which was phylogenetically overdispersed, showed a clear morphological clustering, which could have been caused by similar ecological adaptations among taxa in this environment. Only the community in the Cuc Phuong Area showed a pattern of phylogenetic clustering, which was partly caused by an absence of a certain, phylogenetically very distinct group in this region. Filtering due to geographical factors could have been involved here. This study shows how museum collections can be used to examine community assembly and contributes to the understanding of the processes that have shaped karst communities in Vietnam.

Revision of the Alycaeidae of China, Laos and Vietnam (Gastropoda: Cyclophoroidea) I: The genera Dicharax and Metalycaeus.

This is the first part of the revision of the Alycaeidae of China, Laos and Vietnam. Here we revise the species hitherto classified in Chamalycaeus Möllendorff, 1897 (genera Dicharax and Metalycaeus in this study). We show that no Chamalycaeus live in the region, and the alycaeid species of the investigated region can be classified into five genera: Alycaeus Baird, 1850, Dicharax Kobelt & Möllendorff, 1900, Dioryx Benson, 1859, Metalycaeus Pilsbry, 1900 and Pincerna Preston, 1907. We propose Dioryx dautzenbergi Páll-Gergely nom. nov. as a replacement name for Alycaeus (Dioryx) major Bavay & Dautzenberg, 1900, non Alycæus (Dioryx) granum var. major Godwin-Austen, 1893. Alycaeus anceyi Mabille, 1887, A. eydouxi Venmans, 1956, A. mouhoti L. Pfeiffer, 1862, A. rolfbrandti Maassen, 2006 and A. vanbuensis Bavay & Dautzenberg, 1900 are retained in Alycaeus. Alycaeus costulosa Bavay & Dautzenberg, 1912 and A. maolanensis Luo, Zhang & Zhuo 2009 were transferred to Pincerna. We examined newly collected specimens and the type specimens of all species with the exception of the following: A. elevatus Heude, 1886, A. expansus Heude, 1890, A. neglectus Heude, 1885, Chamalycaeus libonensis Chen, Li & Luo 2003, C. panshiensis Chen, 1989, C. tangmaiensis Chen & Zhang, 2001, C. zayuensis Zhang, Chen & Zhou, 2008. Fifteen new species are described as follows: Dicharax (?) abdoui Páll-Gergely, n. sp., Dicharax (?) alticola Páll-Gergely & Hunyadi n. sp., Dicharax (?) ellipticus Páll-Gergely n. sp., Dicharax (?) immaculatus Páll-Gergely n. sp., Dicharax bison Páll-Gergely & Hunyadi n. sp., Dicharax draco Páll-Gergely & Hunyadi n. sp., Dicharax imitator Páll-Gergely & Hunyadi n. sp., Dicharax microcostatus Páll-Gergely n. sp., Dicharax micropolitus Páll-Gergely & Hunyadi n. sp., Dicharax robustus Páll-Gergely & Hunyadi, n. sp., Metalycaeus (?) awalycaeoides Páll-Gergely & Hunyadi n. sp., Metalycaeus (?) ibex Páll-Gergely & Hunyadi n. sp., Metalycaeus laosensis Páll-Gergely n. sp., Metalycaeus oharai Páll-Gergely & Hunyadi n. sp., Metalycaeus (?) okuboi Páll-Gergely & Hunyadi, n. sp. A total of twelve names are synonymised with valid species. In this revision we payed special attention to conchological characters hitherto neglected, such as the fine sculpture of the embryonic whorls and the fine structure of the microtunnels along the breathing tube. We described the radula and penis morphology of five Dicharax species.

Conchological differentiation and genital anatomy of Nepalese Glessulinae (Gastropoda, Stylommatophora, Subulinidae), with descriptions of six new species.

Eleven species of Glessulinae belonging to the genera Glessula Martens, 1860 (three species) and Rishetia Godwin-Austen, 1920 (eight species) are reported from Nepal, six of which are new to science and are described here, viz., G. tamakoshi Budha & Backeljau, sp. n., R. kathmandica Budha & Backeljau, sp. n., R. nagarjunensis Budha & Naggs, sp. n., R. rishikeshi Budha & Naggs, sp. n., R. subulata Budha & Naggs and R. tribhuvana Budha, sp. n. and two are new records for Nepal viz. G. cf. hebetata and R. cf. mastersi. The relation between the shell height-width ratio and the structure of the proximal part of the male reproductive organs in Glessulinae is explored. Illustrations and a key for the identification of the Nepalese Glessulinae are provided, including the first record of a spermatophore in Rishetia.

Molecular phylogeny of the Achatinoidea (Mollusca: Gastropoda).

This study presents a multi-gene phylogenetic analysis of the Achatinoidea and provides an initial basis for a taxonomic re-evaluation of family level groups within the superfamily. A total of 5028 nucleotides from the nuclear rRNA, actin and histone 3 genes and the 1st and 2nd codon positions of the mitochondrial cytochrome c oxidase subunit I gene were sequenced from 24 species, representing six currently recognised families. Results from maximum likelihood, neighbour joining, maximum parsimony and Bayesian inference trees revealed that, of currently recognised families, only the Achatinidae are monophyletic. For the Ferussaciidae, Ferussacia folliculus fell separately to Cecilioides gokweanus and formed a sister taxon to the rest of the Achatinoidea. For the Coeliaxidae, Coeliaxis blandii and Pyrgina umbilicata did not group together. The Subulinidae was not resolved, with some subulinids clustering with the Coeliaxidae and Thyrophorellidae. Three subfamilies currently included within the Subulinidae based on current taxonomy likewise did not form monophyletic groups.

Novel shell device for gas exchange in an operculate land snail.

The operculum of terrestrial snails tightly seals the shell aperture providing protection from predators and body-water loss. To allow respiration with a closed operculum, operculate land snails repeatedly evolved shell devices such as tubes or channels that open to the air. In all Asian members of the Alycaeidae, an externally closed tube lies along the suture behind the aperture that possesses a small internal opening into the last whorl at the tube's anterior end. However, this structure presents a paradox: how is gas exchanged through an externally closed tube? Here we show that many microtunnels open into the tube and run beneath radial ribs along the growth line of the last whorl in Alycaeus conformis These tunnels open to the outside of the shell surface near the umbilicus. Examination under high magnification revealed that the outermost shell layer forms these tunnels only in the whorl range beneath the sutural tube. Each tunnel (ca 16 µm diameter) is far narrower than any known metazoan parasite. These findings support our hypothesis that the externally closed sutural tube functions with microtunnels as a specialized apparatus for predator-free gas exchange with minimal water loss when the operculum seals the aperture.

Review of the genus Endothyrella Zilch, 1960 with description of five new species (Gastropoda, Pulmonata, Plectopylidae).

All known taxa of the genus Endothyrella Zilch, 1960 (family Plectopylidae) are reviewed. Altogether 23 Endothyrella species are recognized. All species are illustrated and whenever possible, photographs of the available type specimens are provided. Five new species are described: Endothyrella angulata Budha & Páll-Gergely, sp. n., Endothyrella dolakhaensis Budha & Páll-Gergely, sp. n. and Endothyrella nepalica Budha & Páll-Gergely, sp. n. from Nepal, Endothyrella robustistriata Páll-Gergely, sp. n. from the Naga Hills, India, and Endothyrella inexpectata Páll-Gergely, sp. n. from Sichuan, China. Helix (Plectopylis) munipurensis Godwin-Austen, 1875 is synonymized with Helix (Plectopylis) serica Godwin-Austen, 1875, and Plectopylis (Endothyra) gregorsoni Gude, 1915 is synonymized with Helix (Plectopylis) macromphalus W. Blanford, 1870. Plectopylis plectostoma var. exerta Gude, 1901 is a synonym of Plectopylis plectostoma var. tricarinata Gude, 1896, which is a species in its own right. Five species of the genus Chersaecia viz. Plectopylis (Chersaecia) bedfordi Gude, 1915, Helix (Plectopylis) brahma Godwin-Austen, 1879, Helix (Plectopylis) Oglei Godwin-Austen, 1879, Helix (Plectopylis) serica Godwin-Austen, 1875, and Plectopylis (Endothyra) williamsoni Gude, 1915 are moved to Endothyrella. The holotype of Plectopylis hanleyi Godwin-Austen, 1879 seems to be lost; therefore, Plectopylis hanleyi is considered to be a nomen dubium.

Annotated checklist of the terrestrial gastropods of Nepal.

This is the very first checklist of the terrestrial gastropods of Nepal. It includes 138 species and six subspecies, of which 22 species are endemic and four are introduced. It highlights 34 species recorded for the first time in Nepal and provides new distribution records for another 30 species.

Illustrated type catalogue of Amphidromus Albers, 1850 in the Natural History Museum, London, and descriptions of two new species.

The collection of the Southeast Asian tree snail genus Amphidromus Albers, 1850 at the Natural History Museum, London includes more than 100 lots of type specimens representing 85 name-bearing types, 9 paratypes and 6 paralectotypes, and one nomen nudum. Lectotypes are here designated for Amphidromuscambojiensis, Amphidromusperakensisglobosus, Amphidromuscolumellarisgloriosa, Amphidromusmaculiferusinflata, Amphidromuslepidus, Amphidromussinistralislutea, Amphidromusmoniliferus, Amphidromusmaculiferusobscura, Amphidromussinistralisrosea and Amphidromussinensivicaria. In addition, the missing types of A.A. Gould were discovered and their type status is discussed. A complete catalogue of these types, including colour photographs is provided for the first time. After examining these type specimens, two new Amphidromus species, Amphidromus (Syndromus) globonevilli Sutcharit & Panha, sp. n. and Amphidromus (Syndromus) principalis Sutcharit & Panha, sp. n. were recognized and are described herein.

Restricted genetic variation in populations of Achatina (Lissachatina) fulica outside of East Africa and the Indian Ocean Islands points to the Indian Ocean Islands as the earliest known common source.

The Giant African Land Snail, Achatina ( =  Lissachatina) fulica Bowdich, 1822, is a tropical crop pest species with a widespread distribution across East Africa, the Indian subcontinent, Southeast Asia, the Pacific, the Caribbean, and North and South America. Its current distribution is attributed primarily to the introduction of the snail to new areas by Man within the last 200 years. This study determined the extent of genetic diversity in global A. fulica populations using the mitochondrial 16S ribosomal RNA gene. A total of 560 individuals were evaluated from 39 global populations obtained from 26 territories. Results reveal 18 distinct A. fulica haplotypes; 14 are found in East Africa and the Indian Ocean islands, but only two haplotypes from the Indian Ocean islands emerged from this region, the C haplotype, now distributed across the tropics, and the D haplotype in Ecuador and Bolivia. Haplotype E from the Philippines, F from New Caledonia and Barbados, O from India and Q from Ecuador are variants of the emergent C haplotype. For the non-native populations, the lack of genetic variation points to founder effects due to the lack of multiple introductions from the native range. Our current data could only point with certainty to the Indian Ocean islands as the earliest known common source of A. fulica across the globe, which necessitates further sampling in East Africa to determine the source populations of the emergent haplotypes.

An annotated catalogue of type specimens of the land snail genus Cyclophorus Monfort, 1810 (Caenogastropoda, Cyclophoridae) in the Natural History Museum, London.

The collection of land caenogastropod snails in the genus Cyclophorus Monfort, 1810 housed in the Natural History Museum, London (NHM), includes 52 type lots. Lectotypes have been designated for 43 available species-level names to stabilize existing nomenclature, two previously designated lectotype, two holotypes, one paratype, one syntype, one possible syntype and two paralectotypes are also listed. A complete catalogue of the Cyclophorus types in NHM, London is provided for the first time.

Revision of the carnivorous snail genus Discartemon Pfeiffer, 1856, with description of twelve new species (Pulmonata, Streptaxidae).

Twelve new species of the streptaxid snail genus Discartemon Pfeiffer, 1856 are described from southern Thailand and western Malaysia, D. afthonodontia sp. n., D. circulus sp. n., D. deprima sp. n., D. discadentus sp. n., D. discamaximus sp. n., D. expandus sp. n., D. flavacandida sp. n., D. kotanensis sp. n., and D. megalostraka sp. n. from southern Thailand, as well as D. conicus sp. n., D. epipedis sp. n. and D. triancus sp. n. from western Malaysia. All 15 previously described species are revised and commented upon based on examined material. Streptaxis paradiscus Möllendorff, 1900 is considered a junior subjective synonym of the type species D. discus (Pfeiffer, 1853). Details of the genital anatomy of twelve species, and the radula and pallial system, are provided for the first time. An identification key is provided.

The mitochondrial 16 s rRNA reveals high anthropogenic influence on land snail diversity in a preliminary island survey.

A total of 30 specimens belonging to five species, namely; Cryptozona siamensis, Sarika resplendens and Sarika sp. from the family Ariophantidae as well as Quantula striata and Quantula sp. from the family Dyakiidae were collected from the Langkawi Island in Northern Peninsular Malaysia. All specimens were identified through comparisons of shell morphology and amplification of a 500 bp segment of the 16S rRNA mtDNA gene. To assess phylogenetic insights, the sequences were aligned using ClustalW and phylogenetic trees were constructed. The analyses showed two major lineages in both Maximum Parsimony and Neighbour Joining phylogenetic trees. Each putative taxonomic group formed a monophyletic cluster. Our study revealed low species and intraspecies genetic diversities based on the 16S rRNA gene sequences. Thus, this study has provided an insight of land snail diversity in populations of an island highly influenced by anthropogenic activities through complementary use of shell morphological and molecular data.

Phylogenetic relationships of the operculate land snail genus Cyclophorus Montfort, 1810 in Thailand.

Operculate land snails of the genus Cyclophorus are distributed widely in sub-tropical and tropical Asia. Shell morphology is traditionally used for species identification in Cyclophorus but their shells exhibit considerable variation both within and between populations; species limits have been extremely difficult to determine and are poorly understood. Many currently recognized species have discontinuous distributions over large ranges but geographical barriers and low mobility of snails are likely to have led to long periods of isolation resulting in cryptic speciation of allopatric populations. As a contribution towards solving these problems, we reconstructed the molecular phylogeny of 87 Cyclophorus specimens, representing 29 nominal species (of which one was represented by four subspecies), plus three related out-group species. Molecular phylogenetic analyses were used to investigate geographic limits and speciation scenarios. The analyses of COI, 16S rRNA and 28S rRNA gene fragments were performed using neighbour-joining (NJ), maximum likelihood (ML), and Bayesian inference (BI) methods. All the obtained phylogenetic trees were congruent with each other and in most cases confirmed the species level classification. However, at least three nominate species were polyphyletic. Both C. fulguratus and C. volvulus appear to be species complexes, suggesting that populations of these species from different geographical areas of Thailand are cryptic species. C. aurantiacus pernobilis is distinct and likely to be a different species from the other members of the C. aurantiacus species complex.

Taxonomic revision of the Elephant Pupinid snail genus Pollicaria Gould, 1856 (Prosobranchia, Pupinidae).

The status of species currently assigned to the Southeast Asian Elephant Pupinid snail genus Pollicaria Gould, 1856 is reassessed. Shell, radular and reproductive morphology are investigated and analysed with reference to karyotype patterns previously reported and to distribution patterns among the species. Six previously described species are recognised: Pollicaria gravida (Benson, 1856), Pollicaria myersii (Haines, 1855), Pollicaria mouhoti (Pfeiffer, 1862), Pollicaria elephas (Morgan, 1885), Pollicaria crossei (Dautzenberg & d'Hamonville, 1887) and Pollicaria rochebruni (Mabille, 1887). A new subspecies, Pollicaria mouhoti monochroma ssp. n.,is proposed and a dichotomous key to species is provided.

Three new species of the carnivorous snail genus Perrottetia Kobelt, 1905 from Thailand (Pulmonata, Streptaxidae).

Three new species of the streptaxid snail genus Perrottetia are described from north and northeastern Thailand, Perrottetia aquilonaria sp. n., Perrottetia dermapyrrhosa sp. n. and Perrottetia phuphamanensis sp. n. Each species is endemic to a single or a few limestone mountain ranges. The species are characterized by the morphology of their genital organs, as well as by shell characters. Perrottetia aquilonaria sp. n. has a club shaped distal penis and large penial hooks are present and penial papillae cover almost the entire penial hook portion; adjacent areas possess low reticulated folds. Perrottetia dermapyrrhosa sp. n. has a long genital atrium and the penial sheath is about two-thirds of the penis length. Penial hooks are long, scattered and sunken into deep ovate hollows; vaginal hooks are present. Perrottetia phuphamanensis sp. n. has a rounded and protruded shell periphery. The aperture is subcircular, peristome is thick and the second parietal lamella is adjacent to the first parietal lamella; a basal lamella is the smaller than in the other Thai species.

Darwininitium - a new fully pseudosigmurethrous orthurethran genus from Nepal (Gastropoda, Pulmonata, Cerastidae).

A new genus and species of pseudosigmurethrous orthurethran pulmonate of the family Cerastidae, Darwininitium shiwalikianumgen. n. and sp. n. is described from the Lesser Himalaya of Nepal. It represents the first record of an orthurethran with a fully developed pseudosigmurethrous pallial system, having a completely closed secondary ureteric system. Biogeographically this new taxon provides a significant range extension for the family north of the previously known distribution range.