The land area of Laos is composed of a large variety of undisturbed habitats, such as high mountainous areas, huge limestone karsts and the lower Mekong Basin. Therefore, Laos is expected to have a high species diversity, especially for the land snails. However, with respect to research on malacology, Laos is probably the least well-researched area for land snail diversity in Indochina (including Laos) over the past few centuries. The handful of species lists have never been systematically revised from the colonial period to the present, so these classifications are outdated. Herein we present the first comprehensive annotated checklist with an up-to-date systematic framework of the land snail fauna in Laos based on both field investigations and literature surveys. This annotated checklist is collectively composed of 231 nominal species (62 ‘prosobranch’ and 169 heterobranches), of which 221 nominal species are illustrated. The type specimens of 143 species from several museum collections and/or 144 species of newly collected specimens are illustrated. There are 58 species recorded as new to the malacofauna of the country, and two new replacement names are proposed as Hemiplectalanxangnica Inkhavilay and Panha, nomen novum (Ariophantidae) and Chloritiskhammouanensis Inkhavilay and Panha, nomen novum (Camaenidae). Four recently described species of the genus Amphidromus from Laos, “thakhekensis”, “richgoldbergi”, “attapeuensis” and “phuonglinhae” are synonymized with previously described species. In addition, thirteen nominal species are listed as uncertain records that may or may not occur in Laos. This annotated checklist may inspire malacologists to carry on systematic research in this region.
Diverse animals exhibit left–right asymmetry in development. However, no example of dimorphism for the left–right polarity of development (whole‐body enantiomorphy) is known to persist within natural populations. In snails, whole‐body enantiomorphs have repeatedly evolved as separate species. Within populations, however, snails are not expected to exhibit enantiomorphy, because of selection against the less common morph resulting from mating disadvantage. Here we present a unique example of evolutionarily stable whole‐body enantiomorphy in snails. Our molecular phylogeny of South‐east Asian tree snails in the genus Amphidromus indicates that enantiomorphy has likely persisted as the ancestral state over a million generations. Enantiomorphs have continuously coexisted in every population surveyed spanning a period of 10 years. Our results indicate that whole‐body enantiomorphy is maintained within populations opposing the rule of directional asymmetry in animals. This study implicates the need for explicit approaches to disclosure of a maintenance mechanism and conservation of the genus.
Seven Scolopendra species from the Southeast Asian mainland delimited based on standard external morphological characters represent monophyletic groups in phylogenetic trees inferred from concatenated sequences of three gene fragments (cytochrome c oxidase subunit 1, 16S rRNA and 28S rRNA) using Maximum likelihood and Bayesian inference. Geometric morphometric description of shape variation in the cephalic plate, forcipular coxosternite, and tergite of the ultimate leg-bearing segment provides additional criteria for distinguishing species. Colouration patterns in some Scolopendra species show a high degree of fit to phylogenetic trees at the population level. The most densely sampled species, Scolopendra dehaani Brandt, 1840, has three subclades with allopatric distributions in mainland SE Asia. The molecular phylogeny of S. pinguis Pocock, 1891, indicated ontogenetic colour variation among its populations. The taxonomic validation of S. dawydoffi Kronmüller, 2012, S. japonica Koch, 1878, and S. dehaani Brandt, 1840, each a former subspecies of S. subspinipes Leach, 1814 sensu Lewis, 2010, as full species was supported by molecular information and additional morphological data. Species delimitation in these taxonomically challenging animals is facilitated by an integrative approach that draws on both morphology and molecular phylogeny.
A high degree of intraspecific variation, both genetic and in shell morphology, of the operculate land snail Cyclophorus fulguratus (Pfeiffer, 1854) suggests that its classification as a single species warrants reconsideration. We sequenced two nuclear (18S and 28S) and two mitochondrial (16S and COI) genes of 46 C. fulguratus specimens and used them to estimate the phylogeny and to determine the validity of species boundaries. Molecular phylogenetic analyses revealed the presence of three lineages corresponding to three geographically disjunctive populations of C. fulguratus in Thailand. Likelihood tests of topologies significantly supported the non-monophyly of the C. fulguratus–complex and Bayesian species delimitation analysis significantly supported the potential representation as distinct species of these three lineages. Discriminant function analysis based on geometric-morphometrics of shell shape allowed for significant distinction of these three candidate species, although they revealed a considerable degree of overlap of shell shape reflecting their crypsis morphologically. The diagnostic characters are provided by color pattern, pattern of protoconch and pattern of jaw. In conclusion, the results support that the C. fulguratus s.l., as currently recognized, consists of three distinct species in Thailand: C. fulguratus s.s., C. rangunensis and C. abditus sp.nov., which are described herein.
Sophina is a poorly known and neglected genus due to the inaccessibility of the Salween Basin, Southern Myanmar. Taxonomic status, distribution, and phylogeny are being revised based on an integrative analysis of genitalia, radula, and molecular data. Morphological variation in shells and genitalia, together with a phylogenetic tree from concatenated data of mitochondrial and nuclear genes, revealed the existence of ten species/subspecies. Penial morphology and genetic divergence were generally consistent and useful in delimiting species, while shell characters showed little overall taxonomic utility in some species. Taxonomic placement of the previous subspecies “bensoni” shows clear distinction in both genitalia and molecular evidence, and is currently recognized as a distinct species. The nominal species “S. schistostelis” and “S. calias” possess similar genitalia and shell morphology, and molecular evidence suggested that they are sister taxa representing geographically isolated populations. Four new species are additionally described herein as S. furfuracea n. sp., S. pisinna n. sp., S. salweenica n. sp., and S. tonywhitteni n. sp. based on both morphology and molecular evidence. Phylogenetic analyses supported monophyly of Sophina, and further a split into two principal clades. These two clades showed little difference in genitalia, but more clear differences in the umbilical area and allopatric distribution in upper and lower reaches of the Gyaing River. High genetic divergence was revealed and this coincided with remarkable degree of endemism and localization with a pattern of one outcrop for one lineage. These data highlight the importance of the Salween Basin's karst ecosystems as an evolutionary and endemic biodiversity hotspot, and indicate that a focus on conservation and management in this area is urgently required.
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