Contributions of a small collection of terrestrial microsnails (Pupilloidea, Hypselostomatidae) from Myanmar with description of three new species.

Land snails were collected for the project 'Conserving Myanmar's Karst Biodiversity' from the limestone karsts in Mon, Kayin, and Shan states and in the regions of Tanintharyi and Mandalay between 2015 and 2017, through cooperation with Fauna and Flora International (FFI) and the Forestry Department of Myanmar. Here, we report on a portion of the collection, and list 17 species from seven genera of the Hypselostomatidae microsnails. Three new species from two genera are described as Bensonellataiyaiorum Tongkerd & Panha, sp. nov., B.lophiodera Tongkerd & Panha, sp. nov., and Gyliotrachelaaunglini Tongkerd & Panha, sp. nov. All new species are known only from the type locality in Shan State (Bensonella) and Kayin State (Gyliotrachela). A new combination of Acinolaemusdayanum and three newly recorded species, namely A.cryptidentatus, B.anguloobtusa and G.hungerfordiana are discussed. The low morphological variability of the widely distributed G.hungerfordiana is discussed, and two species are proposed for formal synonymisation. Constituting the first records for Myanmar, five species of Bensonella and two species of Acinolaemus were collected.

Molecular phylogeny and taxonomic position of Macrobrachiumlanchesteri (De Man, 1911), with descriptions of two new species from Thailand (Decapoda, Caridea, Palaemonidae).

Macrobrachiumlanchesteri (De Man, 1911), a translucent freshwater prawn has a wide distribution range throughout mainland Southeast Asia. A high morphological variation and genetic divergence between different geographical M.lanchesteri populations in Thailand have peculiarly extended the uncertainty of species boundaries and blended confusingly with several Macrobrachium species. To clarify these circumstances, broad sample examinations of the morphological variation, including topotype specimens, and phylogenetic reconstruction based on the concatenated mitochondrial dataset (16s rRNA and COI genes) were performed. Broad morphological examination of M.lanchesteri has shown congruency with phylogenetic analyses by revealing prominent lineages of M.lanchesteri sensu stricto and two new sibling lineages with interspecific variation between 6.48-8.76% for COI and 3.06-4.23% for 16S. Descriptions of two new species, named herein as M.panhai Chaowvieng & Siriwut, sp. nov. and M.rostrolevatus Chaowvieng & Siriwut, sp. nov. are provided. Morphological investigation of rostral form suggested plasticity in M.rostrolevatus populations showing the morphological trait associated with their habitat preferences. Furthermore, phylogenetic positions of the three taxa affirmed the hidden diversity of Thai freshwater Macrobrachium fauna correlated with the river network in the Mekong and Chao Phraya basins, Thailand. The genetic data and distribution records obtained in this study may also assist future river conservation plans as well as the sustainable management of freshwater prawn diversity.

A new jewel-like species of the pill-millipede genus Sphaerobelum Verhoeff, 1924 (Diplopoda, Sphaerotheriida, Zephroniidae) from Thailand.

A new species of the giant pill millipede genus Sphaerobelum is described: Sphaerobelumturcosasp. nov. from the northeastern part of Thailand. Species delimitation is based on morphological characters and COI sequence data. The new species can be clearly discriminated from congeners by its greenish-blue body color, the face mask-like appearance of the thoracic and anal shields jointly when rolled up, and the combination of the following four characters: (1) the coxa of the second leg laterally with a sharp and long process, (2) the tarsi of legs 4-21 with 6-7 ventral spines, (3) the anterior telopods consisting of four conspicuous telopoditomeres, and (4) the immovable, slender (not strongly humped) and distally curved finger of the posterior telopods without a membranous spot. The interspecific COI sequence divergence between the new species and other Sphaerobelum species ranges from 17% to 23% (mean 20%). The intergeneric COI sequence divergence between the new species and Zephronia species ranges from 18% to 21% (mean 20%). The relationships among Sphaerobelum and Zephronia species based on the COI sequence data were not resolved in this study. Sphaerobelumturcosasp. nov. is restricted to limestone habitat in Loei province and is probably endemic for the Thai fauna.

Review of the pill millipede genus Hyperglomeris Silvestri, 1917 (Diplopoda, Glomerida, Glomeridae) with description of two new species from Laos.

The pill millipede genus Hyperglomeris Silvestri, 1917 is reported from Laos for the first time. Two new species, namely H.bicaudata Likhitrakarn, sp. nov. and H.inkhavilayi Likhitrakarn, sp. nov., from Houaphanh and Khammouane provinces, northern Laos, are described and illustrated based on morphological characters and molecular analyses. Sequences of COI gene were used as DNA barcoding markers, and successfully supported the accurate identification of other Glomeridae species. Interspecific divergence of the COI uncorrected p-distance between these new species and other Hyperglomeris species ranged from 7.84-13.07%, while the intraspecific divergence was 0.45% in H.inkhavilayisp. nov. and 5.3% in H.bicaudatasp. nov. The updated status of Hyperglomeris, a map of its distribution, and identification keys for all species are given.

The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z.siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae).

Material of the giant pill-millipede genus Zephronia Gray, 1832 recently collected from Thailand contains three new species: Zephroniaenghoffi sp. nov., Zephroniagolovatchi sp. nov., and Zephroniapanhai sp. nov. The first Zephronia species recorded for Thailand, Z.siamensis Hirst, 1907, is also redescribed based on new specimens collected both from the type locality in Chonburi Province and from neighboring areas. Morphological characters of all new species, Z.phrain Likhitrakarn & Golovatch, 2021, and Z.siamensis are illustrated, and a distribution map of the confirmed Zephronia species occurring in Thailand is also provided.

Two new species of the millipede genus Tylopus Jeekel, 1968 from Shan State, Myanmar (Diplopoda, Polydesmida, Paradoxosomatidae).

The predominantly Indochinese to southern Chinese millipede genus Tylopus presently comprises 76 described species, including two new, T. monticola sp. nov. and T. sutchariti sp. nov., both described and illustrated based on material from a limestone mountain in Taunggyi District, Shan State, Myanmar. Both new species have been found to occur syntopically near limestone caves and are assumed to be narrowly endemic to the Taunggyi Mountains, southwestern Shan State, Myanmar. A key to all six Tylopus species known to occur in Myanmar is provided, and their distributions are also mapped.

A new micropolydesmoid millipede of the genus Eutrichodesmus Silvestri, 1910 from Cambodia, with a key to species in mainland Southeast Asia (Diplopoda, Polydesmida, Haplodesmidae).

The micropolydesmoid millipede family Haplodesmidae is here recorded from Cambodia for the first time through the discovery of the first, new species of the genus Eutrichodesmus Silvestri, 1910: E. cambodiensis sp. nov. This new species is described from two limestone habitats in Kampot Province, based on abundant material. It is easily distinguished from all related congeners by the following combination of characters: body greyish-brown; limbus roundly lobulate; solenomere partially divided from acropodite by a digitiform lobe, but without hairpad. Brief remarks on the previously-proposed "pecularis-group" are provided and a second group, the "demangei-group", is established and discussed on the basis of morphological evidence, updating the number of recognised species groups of Eutrichodesmus to two. Detailed morphological illustrations, photographs and a distribution map, as well as remarks on its habitat and mating behaviour of the new species are presented. Furthermore, the current distributions of all 55 presently-known species of Eutrichodesmus are provided and a key to all 23 species that occur in mainland Southeast Asia is given.

Annotated checklist of freshwater molluscs from the largest freshwater lake in Southeast Asia.

The Tonle Sap Lake in Cambodia is a crucial freshwater biodiversity hotspot and supports one of the world's largest inland fisheries. Within the Tonle Sap basin, freshwater molluscs provide vital ecosystem services and are among the fauna targetted for commercial harvesting. Despite their importance, freshwater molluscs of the Tonle Sap basin remain poorly studied. The historical literature was reviewed and at least 153 species of freshwater molluscs have been previously recorded from throughout Cambodia, including 33 from the Tonle Sap basin. Surveys of the Tonle Sap Lake and surrounding watershed were also conducted and found 31 species, 15 bivalves (five families) and 16 gastropods (eight families), in the Tonle Sap basin, including three new records for Cambodia (Scaphula minuta, Novaculina siamensis, Wattebledia siamensis), the presence of globally invasive Pomacea maculata and potential pest species like Limnoperna fortunei. This study represents the most comprehensive documentation of freshwater molluscs of the Tonle Sap basin, and voucher specimens deposited at the Inland Fisheries Research and Development Institute, Cambodia, represent the first known reference collection of freshwater molluscs in the country. In order to combat the combined anthropogenic pressures, including invasive species, climate change and dams along the Mekong River, a multi-pronged approach is urgently required to study the biodiversity, ecology, ecosystem functioning of freshwater molluscs and other aquatic fauna in the Tonle Sap basin.

Annotated checklist of the land snail fauna from southern Cambodia (Mollusca, Gastropoda).

Prior to this study, few collections and records were made of the land snails in Cambodia and the historical taxa had never been reviewed. Herein a report on the land snail diversity based on specimens collected recently from karstic and non-karstic areas in southern Cambodia is provided. This checklist presents 36 species of land snails (two Neritimorpha, six Caenogastropoda, and 28 Heterobranchia). Illustrations and brief taxonomic notes/remarks are provided for every species. We also described Georrisa carinata Sutcharit & Jirapatrasilp, sp. nov. based on some distinct shell morphological characters. Since the first descriptions during the colonial period in the nineteenth century, some land snail species (e.g., Trichochloritis norodomiana, Durgella russeola, Anceyoconcha siamensis obesula comb. nov., Anceyoconcha chaudoensis comb. nov., and Succinea tenuis) have not been reported subsequently. This probably reflects a lack of knowledge concerning land snail biodiversity in this country. To our knowledge, this is the first comprehensive survey of land snails in southern Cambodia. A need for more field research and systematic revision of the land snails in this interesting region is also highlighted and demonstrated.

Two new species of the millipede genus Plusioglyphiulus Silvestri, 1923 from Cambodia (Diplopoda, Spirostreptida).

Two new species of Plusioglyphiulus are described from southern Cambodia. Plusioglyphiulus biserratus sp. nov. is clearly distinguished from all congeners by the shape of the telopodites of the posterior gonopods which are distinctly serrate laterally and by the anterior gonopods showing only a pair of single, smooth and curved coxosternal processes. Plusioglyphiulus khmer sp. nov. is distinguished by having most crests on the collum being complete and male legs 1 showing long, prominent, one-segmented telopodites, coupled with the oblong-subtrapeziform, membranous, posterior gonopods with a small bifid process at about a third of the telopodite length. Notes on the variation of Plusioglyphiulus boutini Mauriès, 1970 are also given, including a colour photograph of fresh, live material. A key to all four species of Plusioglyphiulus currently known to occur in Cambodia is also presented.

A remarkable new species of the millipede genus Trachyjulus Peters, 1864 (Diplopoda, Spirostreptida, Cambalopsidae) from Thailand, based both on morphological and molecular evidence.

A new, giant species of Trachyjulus from a cave in southern Thailand is described, illustrated, and compared to morphologically closely related taxa. This new species, T. magnus sp. nov., is much larger than all other congeners and looks especially similar to the grossly sympatric T. unciger Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2012, which is widespread in southern Thailand. Phylogenetic trees, both rooted and unrooted, based on a concatenated dataset of the COI and 28S genes of nine species of Cambalopsidae (Trachyjulus, Glyphiulus, and Plusioglyphiulus), strongly support the monophyly of Trachyjulus and a clear-cut divergence between T. magnus sp. nov. and T. unciger in revealing very high average p-distances of the COI gene (20.80-23.62%).

A new species of the millipede genus Cryptocorypha Attems, 1907, from northern Thailand (Polydesmida, Pyrgodesmidae).

The millipede family Pyrgodesmidae and the genus Cryptocorypha are recorded from Thailand for the first time, being represented there by C.enghoffi sp. n. The new species is distinguished by the evident apicodorsal trichostele on the last tibia of both sexes and the gonopodal telopodite being particularly complex, quadripartite, consisting of the longest, mesal, suberect solenomere branch; a slightly shorter, similarly slender, acuminate endomere branch tightly appressed to the solenomere; a somewhat shorter, caudal, strongly curved, armed exomere process; and a very distinct, low, lateral, sac-shaped velum at their base. This situation strongly resembles the one observed in the geographically closest C.perplexa Golovatch & VandenSpiegel, 2015, from Myanmar, but the shapes and armament of all outgrowths of the gonopodal telopodite are clearly different. A key to all three Cryptocorypha pecies known from Indochina or Myanmar and an updated checklist of all 21 species of the genus are provided.

Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda).

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.

A revision of dragon millipedes IV: the new genus Spinaxytes, with the description of nine new species (Diplopoda, Polydesmida, Paradoxosomatidae).

Nine new species constituting the 'spiny' group of dragon millipedes are assigned to the new genus Spinaxytes Srisonchai, Enghoff & Panha, gen. n. Seven new species are described from Thailand: S.biloba Srisonchai, Enghoff & Panha, sp. n. and S.palmata Srisonchai, Enghoff & Panha, sp. n. from Surat Thani Province, S.hasta Srisonchai, Enghoff & Panha, sp. n. from Chumphon Province, S.krabiensis Srisonchai, Enghoff & Panha, sp. n. (type species) and S.sutchariti Srisonchai, Enghoff & Panha, sp. n. from Krabi Province, S.uncus Srisonchai, Enghoff & Panha, sp. n., and S.macaca Srisonchai, Enghoff & Panha, sp. n. from Phang Nga Province; as well as one from Malaysia, S.tortioverpa Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar, S.efefi Srisonchai, Enghoff & Panha, sp. n. The new genus is endemic to South Myanmar, South Thailand, and Malaysia, and all new species are restricted to limestone habitats. All were exclusively found living on humid rock walls and/or inside small caves. Complete illustrations of external morphological characters, an identification key, and a distribution map are provided.

A revision of dragon millipedes I: genus Desmoxytes Chamberlin, 1923, with the description of eight new species (Diplopoda, Polydesmida, Paradoxosomatidae).

The dragon millipede genus Desmoxytes s.l. is split into five genera, based on morphological characters and preliminary molecular phylogenetic analyses. The present article includes a review of Desmoxytes s.s., while future articles will deal with Hylomus Cook and Loomis, 1924 and three new genera which preliminarily are referred to as the 'acantherpestes', 'gigas', and 'spiny' groups. Diagnostic morphological characters of each group are discussed. Hylomus is resurrected as a valid genus and the following 33 species are assigned to it: H. asper (Attems, 1937), comb. n., H. cattienensis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. cervarius (Attems, 1953), comb. n., H. cornutus (Zhang & Li, 1982), comb. n., H. draco Cook & Loomis, 1924, stat. rev., H. enghoffi (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. eupterygotus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. getuhensis (Liu, Golovatch & Tian, 2014), comb. n., H. grandis (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. hostilis (Golovatch & Enghoff, 1994), comb. n., H. jeekeli (Golovatch & Enghoff, 1994), comb. n., H. lingulatus (Liu, Golovatch & Tian, 2014), comb. n., H. laticollis (Liu, Golovatch & Tian, 2016), comb. n., H. longispinus (Loksa, 1960), comb. n., H. lui (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. minutuberculus (Zhang, 1986), comb. n., H. nodulosus (Liu, Golovatch & Tian, 2014), comb. n., H. parvulus (Liu, Golovatch & Tian, 2014), comb. n., H. phasmoides (Liu, Golovatch & Tian, 2016), comb. n., H. pilosus (Attems, 1937), comb. n., H. proximus (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. rhinoceros (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. rhinoparvus (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. scolopendroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. scutigeroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. similis (Liu, Golovatch & Tian, 2016), comb. n., H. simplex (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. simplipodus (Liu, Golovatch & Tian, 2016), comb. n., H. specialis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. spectabilis (Attems, 1937), comb. n., H. spinitergus (Liu, Golovatch & Tian, 2016), comb. n., H. spinissimus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n. and H. variabilis (Liu, Golovatch & Tian, 2016), comb. n.Desmoxytes s.s. includes the following species: D. breviverpa Srisonchai, Enghoff & Panha, 2016; D. cervina (Pocock,1895); D. delfae (Jeekel, 1964); D. des Srisonchai, Enghoff & Panha, 2016; D. pinnasquali Srisonchai, Enghoff & Panha, 2016; D. planata (Pocock, 1895); D. purpurosea Enghoff, Sutcharit & Panha, 2007; D. takensis Srisonchai, Enghoff & Panha, 2016; D. taurina (Pocock, 1895); D. terae (Jeekel, 1964), all of which are re-described based mainly on type material. Two new synonyms are proposed: Desmoxytes pterygota Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes cervina (Pocock, 1895)), Desmoxytes rubra Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes delfae (Jeekel, 1964)). Six new species are described from Thailand: D. aurata Srisonchai, Enghoff & Panha, sp. n., D. corythosaurus Srisonchai, Enghoff & Panha, sp. n., D. euros Srisonchai, Enghoff & Panha, sp. n., D. flabella Srisonchai, Enghoff & Panha, sp. n., D. golovatchi Srisonchai, Enghoff & Panha, sp. n., D. octoconigera Srisonchai, Enghoff & Panha, sp. n., as well as one from Malaysia: D. perakensis Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar: D. waepyanensis Srisonchai, Enghoff & Panha, sp. n. The species can mostly be easily distinguished by gonopod structure in combination with other external characters; some cases of particularly similar congeners are discussed. All species of Desmoxytes s.s. seem to be endemic to continental Southeast Asia (except the 'tramp' species D. planata). Some biological observations (relationship with mites, moulting) are recorded for the first time. Complete illustrations of external morphological characters, an identification key, and distribution maps of all species are provided.

Two new species of the millipede family Cambalopsidae from Myanmar (Diplopoda, Spirostreptida).

Two new species of cave-dwelling millipedes are described from Myanmar, one each in the genera Plusioglyphiulus Silvestri, 1923 and Trachyjulus Peters, 1864. Plusioglyphiulus digitiformissp. n. joins the small peculiar group of congeners from Thailand which is characterized by such plesiomorphies as the tergal crests on the collum and following metaterga being transversely divided into two, not three parts, as in species of the genus Glyphiulus Gervais, 1847. However, this new species differs by the 3-segmented telopodites of ♂ legs 1, the anterior gonopodal coxosternum showing higher and nearly straight apicomesal processes and very evident apicolateral teeth, as well as the higher and acuminate paramedian coxal processes of the posterior gonopods, the latter's telopodites demonstrating an apical fovea bearing a group of microsetae at the bottom. Trachyjulus bifidussp. n. is primarily distinguished by the telopodites of their anterior gonopods being strikingly and deeply bifid. A key to the five species of Cambalopsidae currently known to occur in Myanmar is presented, and a map showing their distributions given.

Two new species of the millipede genus Glyphiulus Gervais, 1847 from Laos (Diplopoda, Spirostreptida, Cambalopsidae).

Two new species of Glyphiulus are described and illustrated from northern Laos. The epigean Glyphiulus subbedosae Likhitrakarn, Golovatch & Panha, sp. n. is the second member of the granulatus-group to be found in that country and it seems to be especially similar to G. bedosae Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007. However, it differs from the latter species by a row of several strong setae near the median marginal ridge on the paraprocts, combined with the gnathochilarium being considerably less densely setose on the caudal face, and the anterior gonopods showing a pair of smaller, apical, but larger lateral teeth on the coxosternal plate. Glyphiulus semicostulifer Likhitrakarn, Golovatch & Panha, sp. n. is the fourth member of the javanicus-group to be discovered in Laos, taken from a cave. It seems to be particularly similar to G. costulifer Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007, but is distinguished by the more sparsely alveolate background fine structure of the metazonae, coupled with the gnathochilarium being considerably less densely setose on the caudal face, much stronger paramedian prongs and 4-segmented telopodites on ♂ coxae 1, the slightly longer and more slender apicoparamedian sternal projections on the anterior gonopods, and the much longer flagella of the posterior gonopods. An identification key to and a distribution map of Glyphiulus species in Laos are also presented.

A new species of Trichopeltis Pocock, 1894 from southern China, with a checklist and a distribution map of Trichopeltis species (Diplopoda, Polydesmida, Cryptodesmidae).

The millipede genus Trichopeltis Pocock, 1894 contains 12 described species including a new species from southern China described here. Trichopeltis sutcharitisp. n. can be distinguished from congeners by its gonopods that are strongly caudolaterally curved and have a prominent, high, curved, densely setose process on each coxa. An updated checklist and a distribution map are provided for all species of the genus.

Four colorful new species of dragon millipedes, genus Desmoxytes Chamberlin, 1923, from northern Thailand (Diplopoda: Polydesmida: Paradoxosomatidae).

Four new dragon millipede species of the genus Desmoxytes from northern Thailand are described and illustrated: D. des sp. n. from Chiang Mai Province, D. breviverpa sp. n. from Phrae Province, D. takensis sp. n. from Tak Province and      D. pinnasquali sp. n. from Phitsanulok Province. The new species were compared with type specimens of closely related species and were all found to exhibit explicit morphological differences from these. They can be distinguished from other members of Desmoxytes by the shape of the gonopods, the sternal process between male coxae 4, the pattern of spines or tubercles on the metaterga, and the shape of the paraterga. The coloration of all new species is clearly aposematic: ranging from purple-pink to red. The new species are discussed in relation to their congeners, and a distribution map is provided.