A well-supported and well-resolved phylogeny based on a concatenated data set from one mitochondrial and two nuclear genes, six morphological characters, and nine color pattern characters for 44 of the 50 species of the Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) is consistent with the previous taxonomy of Cnemaspis based solely on morphology and color pattern. Cnemaspis is partitioned into four major clades that collectively contain six species groups. The monophyly of all clades and species groups is strongly supported and they are parapatrically distributed across well-established, biogeographical regions ranging from southern Vietnam westward through southern Indochina, southward through the Thai-Malay Peninsula, then eastward to Borneo. Eight new species (Cnemaspis omari sp. nov. from the Thai-Malaysian border; C. temiah sp. nov. from Cameron Highlands, Pahang, Malaysia; C. stongensis sp. nov. from Gunung Stong, Kelantan, Malaysia; C. hangus sp. nov. from Bukit Hangus, Pahang, Malaysia; C. sundagekko sp. nov. from Pulau Siantan, Indonesia; C. peninsularis sp. nov. from southern Peninsular Malaysia and Singapore, and C. mumpuniae sp. nov. and C. sundainsula sp. nov. from Pulau Natuna Besar, Indonesia) are described based on morphology and color pattern and all but C. sundagekko sp. nov. are included in the phylogenetic analyses. Cnemaspis kendallii is polyphyletic and a composite of six species. An updated taxonomy consistent with the phylogeny is proposed for all 50 species and is based on 25 morphological and 53 color pattern characters scored across 594 specimens. Cladogenetic events and biogeographical relationships within Cnemaspis were likely influenced by this group's low vagility and the cyclical patterns of geographical and environmental changes in Sundaland over the last 25 million years and especially within the last 2.5 million years. The phylogeny indicates that nocturnality, diurnality, substrate preferences, and the presence of ocelli in the shoulder regions have evolved independently multiple times.
Species of the bark louse genus Trichadenotecnum Enderlein (Insecta: Psocodea) from Peninsular Malaysia and Singapore are revised with illustrations and identification keys. Twenty species are here recognised, with four new species and ten recorded for the first time from this region, together with an unnamed species represented by a single female. The previously described species T. marginatum New & Thornton is not included because its generic assignment is questionable. Females of T. cinnamonum Endang & New, T. imrum New & Thornton and T. sibolangitense Endang, Thornton & New, and the male of T. kerinciense Endang & New are described for the first time. A new species group is defined for T. krucilense Endang, Thornton & New.
The taxonomy of three genera of little-known Phaneropterinae from Malay Peninsula (Singapore and Peninsular Malaysia) is reviewed: Cesasundana Koçak & Kemal, 2009, Elbenia Stål, 1876 and Macedna Karsch, 1891. One new species each from the genera Cesasundana and Elbenia are described from Singapore and Peninsular Malaysia respectively: Cesasundana lorniensis sp. n. and Elbenia fraser sp. n. Keys to species of Cesasundana and Elbenia are also included. The taxonomic position of monotypic genus Macedna is validated, male is redescribed and the female is described for the first time.
The Oriental, Australasian and Oceanian genus Caiusa Surcouf, 1920 is revised, species concepts being based on male and female genitalia. A key to males for all known species, and a key to females for all except one are given. All relevant types still in existence have been studied, complete synonymies given and the geographical distribution reconsidered. The eight species included in the genus are: Caiusa borneoensis sp. nov. (Malaysia, Thailand, Vietnam); Caiusa coomani Séguy, 1948 (China, Malaysia, Singapore, Thailand, Vietnam); Caiusa indica Surcouf, 1920 (Australia, Cambodia, India, Indonesia, Malaysia, Papua New Guinea, Philippines, Singapore, Solomon Islands, Sri Lanka, Thailand, Vietnam); Caiusa karrakerae sp. nov. (Malaysia, Thailand); Caiusa kurahashii sp. nov. (Indonesia, Japan, Philippines); Caiusa pooae sp. nov. (Thailand); Caiusa testacea Senior-White, 1923 (India, Nepal, Sri Lanka) and Caiusa violacea Séguy, 1925, stat. rev. (Cambodia, China, Laos, Malaysia, Taiwan, Thailand, Vietnam). A lectotype is designated for Caiusa indica to fix the interpretation of the name. Caiusa nigronitens Senior-White, 1923, syn. nov. and Caiusa surcoufi Bezzi, 1927, syn. nov. are established as junior synonyms of Caiusa indica. Caiusa violacea is correctly diagnosed and errors in the original description of the female holotype are pointed out. Caiusa dubiosa Villeneuve, 1927 is established as a junior synonym of C. violacea, syn. nov. Seven Caiusa species have been reared from the egg mass of various species of frogs. The reproductive mode of the eighth species, i.e., C. indica, is unknown. Five species, i.e., C. borneoensis, C. coomani, C. karrakerae, C. kurahashii and C. violacea have been reared from one or more of the foam nesting frog species Chiromantis nongkhorensis (Cochran, 1927), Polypedates leucomystax (Gravenhorst, 1927), Polypedates megacephalus Hallowell, 1861, Rhacophorus annamensis Smith, 1924, Rhacophorus dulitensis Boulenger, 1892, Rhacophorus kio Ohler & Delorme, 2005 and Rhacophorus owstoni (Stejneger, 1907) all belonging in the family Rhacophoridae in Anura. These five Caiusa species all have a specialised ovipositor tip, with small spine-like setae on the ST8 and the hypoproct, probably enabling the flies to oviposit on a foam nest with a hardened outer surface. They form a monophyletic group on account of these features of the ovipositor, unique in the Oestroidea. The sixth species, C. testacea, has been reared from a frog egg mass, the frog species being unknown. Its ovipositor structure is also unknown. The seventh species, C. pooae, has been reared once from the jelly-like egg mass of Feihyla hansenae (Cochran, 1927), also in Rhacophoridae. Caiusa pooae females do not have spine-like setae on the ovipositor, a fact correlated with the soft outer surface of the jelly-like egg mass on which a C. pooae female had oviposited. The extreme rarity of C. pooae oviposition on Feihyla hansenae egg masses may indicate that this fly perhaps has another, unknown, regular oviposition substrate. Caiusa pooae and C. indica make up a second monophyletic group within Caiusa. Caiusa indica, the most common and most widespread species of the genus, has an ovipositor structure similar to C. pooae. Its breeding substrate is unknown and it occurs both within and outside the distributional area of Rhacophoridae. Possibly both C. indica and C. pooae share a regular oviposition substrate that has still to be discovered. The holotype female of Plinthomyia emimelania Rondani, 1875 from Sarawak is established as a member of the genus Bengalia Robineau-Desvoidy, 1830, thus Plinthomyia Rondani, 1875 becomes a junior synonym of Bengalia Robineau-Desvoidy, 1830, syn. nov. It is removed from the synonymy of Phumosia Robineau-Desvoidy, 1830.
Horaeomorphus bicornis sp. n., a remarkable species with strongly modified head from Yunnan, Southwest China is described. A distinctly different female from the same locality is also recorded, its identity remains unconfirmed until associated males become available. The previously unknown female of H. hujiayaoi Zhou & Zhang, 2016, is discovered, with its spermatheca and female terminalia illustrated. Horaeomorphus punctatissimus Franz, 1992 is newly recorded from Mount Trus Madi, Malaysia. Two females of H. eumicroides Schaufuss, 1889 were discovered from Singapore, with female terminalia illustrated.
Here, taxonomic notes on species from three genera of bark crickets are presented. A new locality is recorded for Duolandrevus (Bejorama) parvulus Gorochov, 2016: Singapore. Two new species of Landrevinae are described: one species of Endodrelanva Gorochov, 1999 from Singapore: Endodrelanva jimini sp. nov.; one species of Odontogryllodes Chopard, 1969 from Fraser's Hill, Peninsular Malaysia: Odontogryllodes gratus sp. nov. The male of Odontogryllodes latus Chopard, 1969 is also described for the first time and female is redescribed. Keys to all known species of Endodrelanva and Odontogryllodes is provided.
The genus Doryscus Jacoby, 1887 is revised. Twelve new species are described: D. indochinensis sp. nov. from China (Yunnan), India, Laos, Thailand, Vietnam; D. krausi sp. nov. from Laos and Thailand; D. kubani sp. nov., from China (Yunnan); D. geiseri sp. nov. from Singapore and Thailand; D. nepalensis sp. nov. from Bhutan, north India, and Nepal; D. luzonensis sp. nov., and D. mindanaoensis sp. nov. from Philippines; D. barclayi sp. nov., D. boreri sp. nov., D. javanensis sp. nov., and D. sumatrensis sp. nov. from Indonesia; D. wangi sp. nov. from East Malaysia (Sabah). Doryscus chujoi Takizawa, 1978 and D. varians (Gressitt & Kimoto, 1963) are removed from synonymy with D. testaceus Jacoby, 1887. Doryscus nigricollis Jiang, 1992 and D. marginicollis Jiang, 1992 are regarded as junior synonyms of D. varians (Gressitt & Kimoto, 1963). A lectotype is designated for Doryscus testaceus Jacoby, 1887.
Five species of the genus Polyonyx Stimpson, 1858 are reported from southern Japan, the Philippines, Singapore, and Peninsular Malaysia. All of them belong to the P. sinensis group, which is morphologically defined by having the dactyli of the ambulatory legs each with the dorsal claw being much smaller than the ventral claw. Two species are described as new to science, one each from Peninsular Malaysia and the Ryukyu Islands in Japan, and their affinities are discussed. Polyonyx boucheti Osawa, 2007, P. heok Osawa Ng, 2016, and P. thai Werding, 2001, are also newly reported from southern Japan, the Philippines, and Singapore. A key to the Indo-West Pacific species of the P. sinensis group is provided.
The Sundaic swamp clade of the genus Cyrtodactylus contains nine species that collectively range through Peninsular Malaysia and its associated land bridge islands, Singapore, Sumatra, Java, and Pulau Natuna Besar. Ancestral range reconstruction analyses using BioGeoBEARS based on an updated molecular phylogeny of the nine Sundaic swamp clade species of Cyrtodactylus demonstrated that this lineage evolved in Peninsular Malaysia, dispersed independently to Sumatra and Pulau Natuna Besar, Indonesia and most likely back into Peninsular Malaysia from Sumatra. This scenario is consistent with climate-driven, cyclical, ephemeral, geographic reconfigurations of Sundaic landmasses from at least the mid-Miocene to present.
A new species of the porcellanid genus Raphidopus Stimpson, 1858, is described on the basis of specimens from Penang and Kuala Selangor, Peninsular Malaysia. Raphidopus brevipes n. sp. is distinguished from its allied congeners, R. indicus Henderson, 1893 and R. johnsoni Ng Nakasone, 1994, by the proportionally narrower carapace and stouter ambulatory legs, as well as the different dentition on the cutting edge of the dactylus of the larger cheliped. Raphidopus johnsoni, previously known only by the holotype from Singapore, is now also recorded from Phuket (Thailand) and Java (Indonesia). Raphidopus indicus and R. johnsoni are rediagnosed on the basis of the respective type material and other specimens. A key to species of the genus Raphidopus is provided.
Twenty new species of Agrilus jewel beetles from the Oriental region are described and illustrated: Agrilus cicadelloides sp. nov. (Malaysia: Sarawak); A. draco sp. nov. (Malaysia: Sabah); A. hergovitsi sp. nov. (Malaysia: Johor); A. hik sp. nov.(Cambodia); A. ika sp. nov. (Solomon Islands); A. jankae sp. nov. (Indonesia: Sumatra, Singapore); A. jum sp. nov. (Laos); A. kon sp. nov. (Cambodia); A. mimicus sp. nov. (Laos); A. qom sp. nov. (Laos); A. titi sp. nov. (Malaysia: Perak); A. uxo sp. nov. (Vietnam); A. wos sp. nov. (Laos); A. xen sp. nov. (Laos); A. xia sp. nov. (Laos); A. xis sp. nov. (Laos); A. yoa sp. nov. (Laos, Vietnam); A. yuk sp. nov. (Laos); A. zao sp. nov. (Indonesia: Java and Sumba Islands); A. zim sp. nov. (Malaysia: Pahang).
Accurate identification of mosquitoes is crucial for the management and control of vector species. Although Singapore is a tropical country with high mosquito diversity, there are no identification keys specific to its mosquito fauna. An illustrated identification key to the genera of the adult female mosquitoes of Singapore is presented. Twenty genera are characterized and two genera found in Malaysia, Culiseta and Topomyia, are also included. The key will aid non-taxonomically trained operational officers in adult mosquito identification. The key to the genera provides the first crucial step in the process to species identification.
Since the book-formatted, monographic treatment of the lizards of Peninsular Malaysia, Singapore, and their adjacent islands by Grismer (2011), 47 additional species have been added to that fauna bringing the total to 174. One species, Cyrtodactylus stresemanni, was removed from the lizard fauna based on its likely mislabeled type locality; 44 species were described as new-29 gekkonids, 11 scincids, five agamids, and one dibamid; and three others, Cnemaspis narathiwatensis, Cyrtodactylus brevipalmatus, and Scincella melanosticta were recently discovered in northern Peninsular Malaysia. Taxonomic changes for seven previously described species occurring after 2011 are noted and significant, new locality records are documented here for additional species. The intent of this review is to provide an up to date peer-reviewed checklist for the lizard species of this region prior to a pending updated book monograph.
The intertidal serpulid polychaete Spirobranchus kraussii was originally described from South Africa and has since been reported in numerous sub (tropical) localities around the world. Recently, however, S. kraussii was uncovered as a complex of morphologically similar and geographically restricted species, raising the need to revise S. cf. kraussii populations. We formally describe S. cf. kraussii from Singapore mangroves as Spirobranchus bakau sp. nov. based on morphological and molecular data. Despite their morphological similarities, Maximum Likelihood and Bayesian Inference analyses of 18S and Cyt b DNA sequence data confirm that S. bakau sp. nov. is genetically distinct from S. kraussii and other known species in the complex. Both analyses recovered S. bakau sp. nov. as part of a strongly supported clade (96% bootstrap, 1 posterior probability), comprising S. sinuspersicus, S. kraussii and S. cf. kraussii from Australia and Hawaii. Additionally, paratypes of S. kraussii var. manilensis, described from Manila Bay in the Philippines, were examined and elevated to the full species S. manilensis. Finally, we tested the hypothesis that fertilisation and embryonic development of S. bakau sp. nov. can occur under the wide range of salinities (19.630.9 psu) and temperatures (2531C) reported in the Johor Strait. Fertilisation success of ≥70% was achieved across a temperature range of 2532C and a salinity range of 2032 psu. Embryonic development, however, had a narrower salinity tolerance range of 2732 psu. Clarifying the taxonomic status of S. cf. kraussii populations reported from localities elsewhere in Singapore and Southeast Asia will be useful in establishing the geographical distribution of S. bakau sp. nov. and other members of the S. kraussii-complex.
Six Oriental species of Evaza Walker, 1856 were described by J.C.H. de Meijere in the period between 1911 and 1924. The type specimens are now deposited in the Museum Naturalis (Leiden). All species were redescribed, photographed and their diagnostic characters were defined. Male terminalia of five species with known males were illustrated in detail. The revised species were compared with recently described new species from Oriental China. Additional material belonging to the revised species was recorded for the first time from the following countries: Evaza demeijerei Brunetti, 1923 (= E. pallipes de Meijere, 1916a) and E. discolor de Meijere, 1916a were found in Malaysia, E. javanensis de Meijere 1911 in Singapore, E. kerteszi de Meijere, 1914 in Laos and Malaysia and E. maculifera de Meijere, 1914 in Malaysia, Philippines, Thailand and Vietnam.
A species of scaly cricket is described here: Ornebius lupus sp. nov. from the mangrove forests in Singapore. Ornebius pullus Ingrisch, 2006 is recorded in eastern Sabah for the first time. The calling songs of Cycloptiloides bimaculatus Tan et al., 2021 and Ornebius pullus from Sabah are described. We also revise the diagnosis of Ectatoderus nigrofasciatus Tan et al., 2021 from Brunei Darussalam.
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.
Previously, the genus Merizocera Fage, 1912 comprised only seven species from Indonesia, Malaysia, Sri Lanka, and Thailand. In this study, 28 new species are described from South and Southeast Asia: M. baoshan Li, sp. nov. (♂♀), M. betong Li, sp. nov. (♂♀), M. colombo Li, sp. nov. (♂♀), M. galle Li, sp. nov. (♂♀), M. hponkanrazi Li, sp. nov. (♂), M. kachin Li, sp. nov. (♂♀), M. kandy Li, sp. nov. (♂♀), M. mandai Li, sp. nov. (♂♀), M. krabi Li, sp. nov. (♂♀), M. kurunegala Li, sp. nov. (♂♀), M. lincang Li, sp. nov. (♀), M. mainling Li, sp. nov. (♂♀), M. nyingchi Li, sp. nov. (♀), M. peraderiya Li, sp. nov. (♂♀), M. phuket Li, sp. nov. (♂♀), M. putao Li, sp. nov. (♂♀), M. ranong Li, sp. nov. (♂♀), M. ratnapura Li, sp. nov. (♂♀), M. salawa Li, sp. nov. (♂), M. tak Li, sp. nov. (♀), M. tanintharyi Li, sp. nov. (♂♀), M. tengchong Li, sp. nov. (♂), M. thenna Li, sp. nov. (♂♀), M. uva Li, sp. nov. (♀), M. wenshan Li, sp. nov. (♂♀), M. wui Li, sp. nov. (♂♀), M. yala Li, sp. nov. (♀), and M. yuxi Li, sp. nov. (♂♀). Among them the genus Merizocera is reported for the first time from China, Myanmar, and Singapore.
An estuarine species of Nereididae (Annelida), Neanthesglandicincta (Southern, 1921) has been newly recorded on the eastern coast of Peninsular Malaysia located in the South China Sea based on 23 specimens collected from three estuaries (Tumpat, Kelantan Delta, Kelantan; Setiu Lagoon, Terengganu; Kuala Ibai, Terengganu). The morphological characteristics of the Malaysian specimens agree well with those of the previous original description and the redescription of N.glandicincta based on Indian, Myanmar and Singapore specimens. The number of paragnaths in all groups on the proboscis of our Malaysian specimens is within the range of the intraspecific variation of N.glandicincta as shown in the previous descriptions. An identification key to species of the Neanthesglandicincta species complex, which includes two morphologically similar species, is provided.
A taxonomic description of all castes of Colobopsis explodens Laciny & Zettel, sp. n. from Borneo, Thailand, and Malaysia is provided, which serves as a model species for biological studies on "exploding ants" in Southeast Asia. The new species is a member of the Colobopsis cylindrica (COCY) group and falls into a species complex that has been repeatedly summarized under the name Colobopsis saundersi (Emery, 1889) (formerly Camponotus saundersi). The COCY species group is known under its vernacular name "exploding ants" for a unique behaviour: during territorial combat, workers of some species sacrifice themselves by rupturing their gaster and releasing sticky and irritant contents of their hypertrophied mandibular gland reservoirs to kill or repel rivals. This study includes first illustrations and morphometric characterizations of males of the COCY group: Colobopsis explodens Laciny & Zettel, sp. n. and Colobopsis badia (Smith, 1857). Characters of male genitalia and external morphology are compared with other selected taxa of Camponotini. Preliminary notes on the biology of C. explodens Laciny & Zettel, sp. n. are provided. To fix the species identity of the closely related C. badia, a lectotype from Singapore is designated. The following taxonomic changes within the C. saundersi complex are proposed: Colobopsis solenobia (Menozzi, 1926), syn. n. and Colobopsis trieterica (Menozzi, 1926), syn. n. are synonymized with Colobopsis corallina Roger, 1863, a common endemic species of the Philippines. Colobopsis saginata Stitz, 1925, stat. n., hitherto a subspecies of C. badia, is raised to species level.