The genus Hemiplecta is a group of large-sized land snails which have long been used as a food resource by Indochinese people. There are five dextral and four sinistral species currently recognized from Thailand. The dextral group is comprised of two previously recorded species (H. humphreysiana and H. distincta), two newly recorded species (H. funerea and H. esculenta), and one new species (H. nemorosa sp. nov.) from northern Thailand is being proposed. We reassessed the diagnostic characters of the genitalia, mantle edge, and radula. Specimens were classified into the genus Hemiplecta on the basis of the penial verge and shell lobe, and on the characters of a bulbous gametolytic sac without a gametolytic duct. A complete species list, together with photographs of the name-bearing types or authenticated specimens and the taxonomic status of Hemiplecta s.l. that are known from Indochina including Peninsular Malaysia and Myanmar, is provided for the first time. In total, this species list contains 39 available nominal species names described from this area. Type or authentic specimens can be located for 37 nominal species names, of which 25 are illustrated herein and the other 12 were recently illustrated. However, two available species-level names could not be traced to any type specimens. In addition, lectotypes of H. funerea and H. pluto are designated herein to stabilize the names.
The large genus Orthomorpha is rediagnosed and is shown to currently comprise 51 identifiable species ranging from northern Myanmar and Thailand in the Northwest to Lombok Island, Indonesia in the Southeast. Of them, 20 species have been revised and/or abundantly illustrated, based on a restudy of mostly type material; further 12 species are described as new: Orthomorpha atypicasp. n., Orthomorpha communissp. n., Orthomorpha isarankuraisp. n., Orthomorpha picturatasp. n., Orthomorpha similanensissp. n., Orthomorpha suberectasp. n., Orthomorpha tuberculiferasp. n.,Orthomorpha subtuberculiferasp. n. and Orthomorpha latitergasp. n., all from Thailand, as well as Orthomorpha elevatasp. n.,Orthomorpha spiniformissp. n. and Orthomorpha subelevatasp. n., from northern Malaysia. The type-species Orthomorpha beaumontii (Le Guillou, 1841) is redescribed in due detail from male material as well, actually being a senior subjective synonym of Orthomorpha spinala (Attems, 1932), syn. n. Two additional new synonymies are proposed: Orthomorpha rotundicollis (Attems, 1937) = Orthomorpha tuberculata (Attems, 1937), syn. n., and Orthomorpha butteli Carl, 1922 = Orthomorpha consocius Chamberlin, 1945, syn. n., the valid names to the left. All species have been keyed and all new and some especially widespread species have been mapped. Further six species, including two revised from type material, are still to be considered dubious, mostly because their paraterga appear to be too narrow to represent Orthomorpha species. A new genus, Orthomorphoidesgen. n., diagnosed versus Orthomorpha through only moderately well developed paraterga, coupled with a poorly bi- or trifid gonopod tip, with at least some of its apical prongs being short spines, is erected for two species: Orthomorpha setosus (Attems, 1937), the type-species, which is also revised from type material, and Orthomorpha exaratus (Attems, 1953), both comb. n. ex Orthomorpha.
We examined allozyme variation in two camaenid tree snails, Amphidromus atricallosus and A. inversus, across two principal regions of Thailand and from Singapore, plus for A. inversus, one site in peninsular Malaysia. Using horizontal starch gel electrophoresis, 13 allozyme loci (11 polymorphic) were screened for A. atricallosus and 18 (5 polymorphic) for A. inversus. Heterozygosity was higher in A. atricallosus (Hexp=0.018-0.201, mean=0.085) than in A. inversus (Hexp=0-0.023, mean= 0.002). Genetic heterogeneity among samples was higher in A. inversus (Fst=0.965) than in A. atricallosus (Fst=0.781). Within A. atricallosus, populations were more differentiated in southern Thailand (Fst=0.551) than in eastern Thailand (Fst=0.144). The high Fst and low Hexp in populations of A. inversus suggest that this species is likely to have experienced a series of strong bottlenecks, perhaps occurring chiefly on offshore continental-shelf islands. The low Fst values of A. atricallosus in eastern Thailand suggest frequent gene flows among populations in this region. The southern and eastern samples of A. atricallosus exhibited fixed allele differences at four loci and great genetic distance (Nei's D=0.485-0.946), suggesting that these two samples may actually represent, or else be evolving into, separate species.
The genus Antheromorpha is redefined and shown to comprise 11 valid species: Antheromorpha miranda (Pocock, 1895), Antheromorpha bistriata (Pocock, 1895), Antheromorpha comotti (Pocock, 1895), Antheromorpha festiva (Brölemann, 1896), Antheromorpha harpaga (Attems, 1937), Antheromorpha mediovirgata (Carl, 1941), Antheromorpha minlana (Pocock, 1895), Antheromorpha pardalis (Pocock, 1895), Antheromorpha paviei (Brölemann, 1896), comb. n., Antheromorpha rosea Golovatch, 2013 and Antheromorpha uncinata (Attems, 1931). Three new synonymies are proposed: Antheromorpha bivittata (Pocock, 1895) and Antheromorpha melanopleuris (Pocock, 1895) are synonymized under Antheromorpha miranda (Pocock, 1895), and Antheromorpha orophila (Carl, 1941) under Antheromorpha comotti (Pocock, 1895). Detailed descriptions and illustrations of fresh material from Thailand and Malaysia are given, especially regarding colour patterns which appear to be crucial for accurate species identifications. Two Antheromorpha species proposed by Attems are redescribed, based on type material. The genus is rediagnosed and a key and a distribution map are also provided. At least in Thailand, adult Antheromorpha rosea have been found to occur every year only for one or two weeks in September or October, disappearing thereafter.
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.
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.
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.
A new species of the megascolecid earthworm genus Pontodrilus Perrier, 1874, Pontodrilus longissimus sp. n., is described from seashores of Thailand and Peninsular Malaysia. The new species differs from congeners, especially the cosmopolitan P. litoralis (Grube, 1855) in the size of the body, number of segments and the shape of the spermathecae. P. litoralis is redescribed, based on specimens collected from the same region and the same type of habitat. DNA fragments of the mitochondrial cytochrome oxidase subunit I of both species were sequenced. Morphological as well as DNA sequence-based comparisons confirm that P. longissimus sp. n. is a lineage distinct from P. litoralis and in fact a new species. The illustrated descriptions are accompanied by a key to species of Pontodrilus.
Earthworm specimens collected from various parts of Thailand were found to contain seven new species of the genus Metaphire Sims & Easton, 1972. These are M. songkhlaensis sp. n. in the octothecal pulauensis species group, M. trangensis sp. n. in the octothecal ignobilis species group, M. khaoluangensis sp. n. and M. khaochamao sp. n. in the sexthecal houlleti species group, M. doiphamon sp. n. in the sexthecal peguana species group, M. saxicalcis sp. n. in the quadrithecal planata species group, and the bithecal M. surinensis sp. n. Type material of some established species from Thailand or northern Malaysia was reinvestigated and illustrated to confirm the status of the new species and to facilitate species comparisons: M. pulauensis (Beddard, 1900), M. baruana (Stephenson, 1932), both with newly designated lectotypes, and M. planata (Gates, 1936), illustrated and redescribed.
The cosmopolitan littoral earthworm Pontodrilus litoralis is distributed in tropical and sub-tropical coastal habitats, whereas P. longissimus is reported only in the Thai-Malay coastal line. In the present study, we examined the difference in salinity effect on the survival rate, wet weight (hereafter weight) change, behaviour, and osmolality of these two Pontodrilus species. A 28 d exposure to varying salinity concentration (0-50 ppt) revealed that P. litoralis is able to survive over a wide salinity range than P. longissimus, with the latter species exhibiting a low survival rate over the same salinity range. During short-term exposure (0-96 h) to a salinity of less than 30 ppt, P. litoralis exhibited weight gain and this was significant in the first 12 h of exposure. However, P. longissimus gained weight when exposed to salinity at under 10 ppt in the first 72 h of exposure. The two species of Pontodrilus behaved differently when exposed to different salinities. The coelomic fluid osmolarity of Pontodrilus was related to the exposure medium and was mostly maintained as hyperosmotic to the external medium over the range of salinities tested. This study shows how two different species of the littoral earthworm genus Pontodrilus respond to a change in salinity, which may explain their dispersal pattern and shape their distribution pattern throughout Southeast Asia.
The status of the indigenous Southeast Asian apple snails belonging to the genus Pila is of concern due to their fast rate of population decline, possibly as a result of multiple factors including habitat loss or disturbance and the introduction of globally-invasive apple snails, Pomacea spp. Conservation actions, including captive breeding of the native Pila species, have been suggested as urgent remedial practices, but the lack of knowledge regarding the fundamental reproductive biology of indigenous Pila spp. makes such practices difficult. In the present study, observations on the mating and egg-laying behaviour of an economic valuable apple snail native to Southeast Asia, P.virescens, were conducted using video recording to examine and describe their reproductive behaviour under a laboratory condition. A total of 15 types of mating and seven egg-laying behaviour were recorded. The mating sequence which subsequently resulted in egg laying was comprised of seven types of major sequential behaviour: mate probing, mounting, shell circling, positioning, insemination posture, sheath withdrawal and dismounting. Rejection of mating attempts by the female was frequently observed. Egg laying occurred during either day or night. A sequence of seven distinct types of behaviour were performed during oviposition: climbing, positioning, forming a temporary tube, mucous secreting, egg depositing, leaving and resting. Overall, these results provide an understanding of the egg-laying behavioural process and highlight its complexity in P.virescens. In addition, detailed ethograms of mating and egg-laying behaviour were derived. These will promote further systematic comparative studies of the reproductive behaviour of apple snails.