Sabah, a Malaysian state at the north-eastern tip of Borneo, is situated in one of the Earth's biodiversity hotspots yet its freshwater gastropod diversity remains poorly known. An annotated checklist of the freshwater gastropods is presented, based on specimens deposited in the BORNEENSIS collection of the Institute for Tropical Biology and Conservation at Universiti Malaysia Sabah, Malaysia. A KMZ file is also provided, which acts as a repository of digital images and complete collection data of all examined material, so that it can be shared and adapted to facilitate future research.
Sabah, situated in one of the world's biodiversity hotspots, has the largest number of islands in Malaysia with more than 500 of various sizes and degrees of isolation. However, information on the islands' biodiversity is limited. This study provides an up-to-date checklist of land snail species found on 24 west coast islands in Sabah. A total of 67 species (nearly 20% of the total number of land snail species in the state) representing 37 genera and 19 families is enumerated based on systematic field surveys of 133 sampling plots, BORNEENSIS database records and species checklists published between 2000 and 2016. The number of species on the islands ranges from four to 29. Labuan Island has the highest number of species (29), followed by Tiga Island (25), Mantanani Besar Island (24) and Gaya Island (23). However, the populations of some land snail species may have declined as several previously recorded species on the islands were not found in a recent systematic field sampling. This checklist is provided as a baseline inventory for future island land snail studies and to better inform biodiversity conservation plans of marine parks and other islands on the Sabah west coast.
A total of 11 species and 1 subspecies of Alycaeus were recognised in Peninsular Malaysia prior to this study. However, these taxonomic descriptions of Alycaeus taxa were based on limited numbers of examined materials, where a whole spectrum of morphological variations were not accounted for and diagnoses were often provided without sufficient comparison between congeners from across the peninsula. We reviewed Peninsular Malaysian Alycaeus through the examination of 5137 specimens in 522 collection lots from all major museum collections and literature sources. Based on these examined materials, we utilised a more comprehensive revised set of 39 shell and operculum characters, as well as living animal colour to describe all Alycaeus species in this paper. We also noted their habitat and ecology, as well as updated the distribution of each species. Of the 12 previously described taxa, 10 are reconfirmed as present on Peninsular Malaysia (Alycaeus balingensis, Alycaeus carinata, Alycaeus conformis, Alycaeus gibbosulus, Alycaeus kapayanensis, Alycaeus kelantanensis, Alycaeus liratulus, Alycaeus perakensis perakensis, Alycaeus perakensis altispirus and Alycaeus thieroti) and 2 are confirmed as absent from the peninsula (Alycaeus jagori and Alycaeus pyramidalis). A new record of Alycaeus robeleni is reported for Peninsular Malaysia. One species, Chamalycaeus jousseaumei is confirmed as present on the peninsula and is reassigned to Alycaeus. The subspecies Alycaeus perakensis altispirus Möllendorff, 1902, is elevated to species. Examined Peninsular Malaysian materials that do not fit previously recognised species are described as new species. A total of 11 new species are proposed (Alycaeus selangoriensissp. n., Alycaeus costacrassasp. n., Alycaeus ikanensissp. n., Alycaeus alticolasp. n., Alycaeus charasensissp. n., Alycaeus kurauensissp. n., Alycaeus regalissp. n., Alycaeus virgogravidasp. n., Alycaeus senyumensissp. n., Alycaeus expansussp. n., Alycaeus clementsisp. n.). Overall, 23 species of Alycaeus are now recognised in Peninsular Malaysia.
Among the six species of Cyrtodactylus occurring in Sumatra, two species were described based on non-Sumatran type series, C.consobrinus and C.quadrivirgatus. The latter species was described originally from Thailand thus the wider distribution in Sumatra should be clarified taxonomically. Cyrtodactylusquadrivirgatus from Sumatra Barat was examined using both morphology and the Natrium Dehydrogenase Subunit 2 (ND2) gene to clarify its taxonomic status and phylogenetic placement. It was found that these specimens form a sister clade to all other species of the sworderi group from Peninsular Malaysia and the genetic distance ranges from 20-24.3%. This subset is herein described as a new species. The new species is readily distinguished from C.quadrivirgatus and other Sumatran species by a combination of characters: small size SVL 37.5-53.78 mm; longitudinal rows of dorsal tubercles 16-19; paravertebral tubercles 31-41; ventral scales 32-43; 24-49 enlarged precloacal and femoral scales; precloacal pores rarely present; no precloacal depression; two postcloacal tubercles on each side; 14-19 subdigital lamellae on forth toe; 9-15 supralabial scales; 9-12 infralabial scales; three or four internasal scales; and 3-6 gular scales that border the first pair of postmental scales. This work underscores the importance of clarifying widely distributed species for taxonomic validation.
The lowland region of Sumatra Barat has received little attention in previous biodiversity studies. Past studies have mainly focused on highland habitat and conservation areas. However, many populations of Cyrtodactylus in the lowland habitats of Sumatra Barat were not correctly identified. A phylogenetic tree based on the NADH dehydrogenase subunit 2 (ND2) gene showed that the lowland Sumatran population is the sister group of the Malaysian lowland species, C.semenanjungensis, together nesting within the agamensis group. The genetic divergence within the Sumatra Barat population is 0-4.2% and 18.3-20% to C.semenanjungensis. Further examination of morphological characters revealed that they differed from the sister clade and other Sumatran Cyrtodactylus members by a unique combination of characters such as absence of tubercle on brachium, presence of tubercle on ventrolateral fold, 32-41 paravertebral tubercles, 38-46 ventral scales, enlarged femoral scales, presence of precloacofemoral pores and 22-23 subdigital lamellae under fourth toe. Based on the morphological and molecular evidence, the lowland Sumatran population is herein described as a new species, increasing the number of species in Sumatra to seven. More comprehensive and intensive sampling efforts would most likely yield further discoveries in the group of Sumatran Cyrtodactylus in the near future.
The southern lesser pomfret (Pampus minor) is an economically important fish, and its numbers are declining because of overfishing and environmental pollution. In addition, owing to the similarities of its external morphological characteristics to other species in the genus Pampus, it is often mistaken for grey pomfret (P. cinereus) or silver pomfret (P. argenteus) juveniles. In this study, the genetic diversity and structure of 264 P. minor individuals from 11 populations in China and Malaysia coastal waters were evaluated for the first time, to the best of our knowledge, using mitochondrial cytochrome b fragments. The results showed that P. minor had moderate haplotype diversity and low nucleotide diversity. Furthermore, two divergent lineages were detected within the populations, but the phylogenetic structure corresponded imperfectly with geographical location; thus, the populations may have diverged in different glacial refugia during the Pleistocene low sea levels. Analysis of molecular variation (AMOVA) showed that genetic variation originated primarily from individuals within the population. Pairwise FST results showed significant differentiation between the Chinese and Malaysian populations. Except for the Xiamen population, which was classified as a marginal population, the genetic differentiation among the other Chinese populations was not significant. During the Late Pleistocene, P. minor experienced a population expansion event starting from the South China Sea refugium that expanded outward, and derivative populations quickly occupied and adapted to the new habitat. The results of this study will provide genetic information for the scientific conservation and management of P. minor resources.
A background study is important for the conservation and stock management of a species. Terapon jarbua is a coastal Indo-Pacific species, sourced for human consumption. This study examined 134 samples from the central west and east coasts of Peninsular (West) Malaysia and East Malaysia. A 1446-bp concatenated dataset of mtDNA COI and Cyt b sequences was used in this study and 83 haplotypes were identified, of which 79 are unique haplotypes and four are shared haplotypes. Populations of T. jarbua in Malaysia are genetically heterogenous as shown by the high level of haplotype diversity ranging from 0.9167-0.9952, low nucleotide diversity ranging from 0.0288-0.3434, and high FST values (within population genetic variation). Population genetic structuring is not distinct as shown by the shared haplotypes between geographic populations and mixtures of haplotypes from different populations within the same genetic cluster. The gene flow patterns and population structuring observed among these regions are likely attributed to geographical distance, past historical events, allopatric speciation, dispersal ability and water currents. For instance, the mixture of haplotypes revealed an extraordinary migration ability of T. jarbua (>1200 km) via ancient river connectivity. The negative overall value of the neutrality test and a non-significant mismatch distribution are consistent with demographic expansion(s) in the past. The median-joining network concurred with the maximum likelihood haplotype tree with three major clades resolved. The scarcity of information on this species is an obstacle for future management and conservation purposes. Hence, this study aims to contribute information on the population structure, genetic diversity, and historical demography of T. jarbua in Malaysia.
For the first time in 21 years, a new genus of cardiochiline braconid wasp, Orientocardiochiles Kang & Long, gen. nov. (type species Orientocardiochiles joeburrowi Kang, sp. nov.), is discovered and described. This genus represents the ninth genus in the Oriental region. Two new species (O. joeburrowi Kang, sp. nov. and O. nigrofasciatus Long, sp. nov.) are described and illustrated, and a key to species of the genus, with detailed images, is added. Diagnostic characters of the new genus are analyzed and compared with several other cardiochiline genera to allow the genus to key out properly using an existing generic treatment. The scientific names validated by this paper and morphological data obtained from this project will be utilized and tested in the upcoming genus-level revision of the subfamily based on combined morphological and molecular data.
The predominantly tropical ophionine genus Enicospilus Stephens, 1835 is one of the largest genera of Darwin wasps (Hymenoptera, Ichneumonidae), with more than 700 extant species worldwide that are usually crepuscular or nocturnal and are parasitoids of Lepidoptera larvae. In the present study, the Japanese species of Enicospilus are revised using an integrative approach (combined morphology and DNA barcoding). On the basis of 3,110 specimens, 47 Enicospilus species are recognised in Japan, eight of which are new species (E. acutus Shimizu, sp. nov., E. kunigamiensis Shimizu, sp. nov., E. limnophilus Shimizu, sp. nov., E. matsumurai Shimizu, sp. nov., E. pseudopuncticulatus Shimizu, sp. nov., E. sharkeyi Shimizu, sp. nov., E. takakuwai Shimizu, sp. nov., and E. unctus Shimizu, sp. nov.), seven are new records from Japan (E. jilinensis Tang, 1990, E. laqueatus (Enderlein, 1921), E. multidens Chiu, 1954, stat. rev., E. puncticulatus Tang, 1990, E. stenophleps Cushman, 1937, E. vestigator (Smith, 1858), and E. zeugos Chiu, 1954, stat. rev.), 32 had already been recorded in Japan; three (E. biharensis Townes, Townes & Gupta, 1961, E. flavicaput (Morley, 1912), and E. merdarius (Gravenhorst, 1829)) have been erroneously recorded from Japan based on misidentifications, and four names that were previously on the Japanese list are deleted through synonymy. The following taxonomic changes are proposed: E. vacuus Gauld & Mitchell, 1981, syn. nov. (= E. formosensis (Uchida, 1928)); E. multidensstat. rev.; E. striatus Cameron, 1899, syn. nov. = E. lineolatus (Roman, 1913), syn. nov. = E. uniformis Chiu, 1954, syn. nov. = E. flatus Chiu, 1954, syn. nov. = E. gussakovskii Viktorov, 1957, syn. nov. = E. striolatus Townes, Townes & Gupta, 1961, syn. nov. = E. unicornis Rao & Nikam, 1969, syn. nov. = E. unicornis Rao & Nikam, 1970, syn. nov. (= E. pungens (Smith, 1874)); E. iracundus Chiu, 1954, syn. nov. (= E. sakaguchii (Matsumura & Uchida, 1926)); E. sigmatoides Chiu, 1954, syn. nov. (= E. shikokuensis (Uchida, 1928)); E. yamanakai (Uchida, 1930), syn. nov. (= E. shinkanus (Uchida, 1928)); E. ranunculus Chiu, 1954, syn. nov. (= E. yezoensis (Uchida, 1928)); and E. zeugosstat. rev. = E. henrytownesi Chao & Tang, 1991, syn. nov. In addition, the following new regional and country records are also provided: E. flavocephalus (Kirby, 1900), E. puncticulatus, and E. vestigator from the Eastern Palaearctic region, E. laqueatus from the Eastern Palaearctic and Oceanic regions, and E. maruyamanus (Uchida, 1928) from the Oriental region; E. abdominalis (Szépligeti, 1906) from Nepal, E. flavocephalus from Laos, E. formosensis from Laos and Malaysia, E. insinuator (Smith, 1860) from Taiwan, E. maruyamanus from India and Philippines, E. nigronotatus Cameron, 1903, E. riukiuensis (Matsumura & Uchida, 1926), and E. sakaguchii from Indonesia, E. pungens from 14 countries (Australia, Bhutan, Brunei, Indonesia, Laos, Malaysia, Nepal, New Caledonia, Papua New Guinea, Philippines, Solomon Islands, Sri Lanka, Tajikistan, and Taiwan), and E. yezoensis from South Korea. An identification key to all Japanese species of Enicospilus is proposed. Although 47 species are recognised in the present study, approximately 55 species could potentially be found in Japan based on ACE and Chao 1 estimators. The latitudinal diversity gradient of Enicospilus species richness is also tested in the Japanese archipelago based on the constructed robust taxonomic framework and extensive samples. Enicospilus species richness significantly increases towards the south, contrary to the 'anomalous' pattern of some other ichneumonid subfamilies.
In a genomic study by Chan and colleagues, pit-vipers of the Trimeresuruserythrurus-purpureomaculatus complex from the Ayeyarwady and Yangon regions in Myanmar were demonstrated to be a distinct species based on robust population genetic and species delimitation analyses. Here, we provide morphological characterizations and a formal description of those populations as a new species. The new species, Trimeresurusayeyarwadyensissp. nov., is most closely related to T.erythrurus and T.purpureomaculatus and shares morphological characteristics with both of those species. Some specimens of T.ayeyarwadyensissp. nov. have green dorsal coloration and no distinct dorsal blotches (a trait shared with T.erythrurus but not T.purpureomaculatus), while others have dark dorsal blotches (a trait shared with T.purpureomaculatus but not T.erythrurus). The distinct evolutionary trajectory of the new species, coupled with the lack of obvious morphological differentiation, represents a classic example of the cryptic nature of species commonly found in the Trimeresurus group of Asian pit-vipers and underscores the need for data-rich analyses to verify species' boundaries more broadly within this genus.
Lobonychiumpalpiplus Roewer, 1938, originally reported from Indonesian Borneo, is redescribed based on the specimens from Malaysia. The genitalia of this species are described for the first time and a new genital terminology is proposed. The rediscovery expands the known distribution of the species to Malaysian Borneo.
The south-east Asian genus Eupoa is redescribed and diagnosed. Seven new species are diagnosed, described and illustrated: E. daklak sp. n. (♀) from Viet-Nam; E. lehtineni sp. n. (♂♀) from India, Thailand and Viet-Nam; E. lobli sp. n. (♂) from Malaysia; E. pappi sp. n. (♂) from Thailand; E. pulchella sp. n.(♂) from Thailand; E. schwendingeri sp. n. (♂♀) from Thailand; and E. thailandica sp. n. (♂♀) from Thailand. Eupoa prima Żabka, 1985 and E. yunnanensis Peng & Kim, 1997 are redescribed and illustrated on the basis of type and/or newly collected materials. The female of E. yunnanensis Peng & Kim, 1997 is found and described for the first time.
Tetrasticta gnatha sp. n., collected under the bark of a rotten fallen tree in Peninsular Malaysia, is described. A habitus photograph, line drawings of diagnostic characters, and a diagnosis are provided. The new species is readily distinguished from all known congeners by having long mandibles, and long, curved maxillary palpi.
Coptotermocola clavicornisgen. & sp. n. and Neotermitosocius bolivianusgen. & sp. n. of the termite inquilinous tribe Termitohospitini are described from peninsular Malaysia and Bolivia, respectively. The Termitohospitini are most readily diagnosable by the distally migrated anterior tentorial pits that are no longer associated with the antennal fossae, and by the enlarged vertex which obscures the antennal fossae dorsally. Additionally, the Termitohospitini are hypothesized to share a recent common ancestor with the Masuriini and Myllaenini due to shared derived morphologies of the lacinia distal teeth with lateral cuticular processes, presence of a unique maxillary palpomere III sensilla, and anterolateral angles of mentum produced. Habitus photographs and illustrations of diagnostic features are provided for the two new genera in order to facilitate future work.
Cheleionwatanabeisp. n. is described from Pahang, Peninsular Malaysia and represents the third species of the genus Cheleion Vårdal & Forshage, 2010 (Coleoptera, Scarabaeidae, Aphodiinae, Stereomerini). A description of the previously unknown male of C.jendeki Král & Hájek, 2015, and a key to the species of the genus Cheleion are also provided.
Blood meal analysis (BMA) from ticks allows for the identification of natural hosts of ticks (Acari: Ixodidae). The aim of this study is to identify the blood meal sources of field collected on-host ticks using PCR analysis. DNA of four genera of ticks was isolated and their cytochrome b (Cyt b) gene was amplified to identify host blood meals. A phylogenetic tree was constructed based on data of Cyt b sequences using Neighbor Joining (NJ) and Maximum Parsimony (MP) analysis using MEGA 5.05 for the clustering of hosts of tick species. Twenty out of 27 samples showed maximum similarity (99%) with GenBank sequences through a Basic Local Alignment Search Tool (BLAST) while 7 samples only showed a similarity range of between 91-98%. The phylogenetic trees showed that the blood meal samples were derived from small rodents (Leopoldamyssabanus, Rattustiomanicus and Sundamysmuelleri), shrews (Tupaiaglis) and mammals (Tapirusindicus and Prionailurusbengalensis), supported by 82-88% bootstrap values. In this study, Cyt b gene as a molecular target produced reliable results and was very significant for the effective identification of ticks' blood meal. The assay can be used as a tool for identifying unknown blood meals of field collected on-host ticks.
Phylogenetic relationships among Malaysia's long-tailed macaques have yet to be established, despite abundant genetic studies of the species worldwide. The aims of this study are to examine the phylogenetic relationships of Macaca fascicularis in Malaysia and to test its classification as a morphological subspecies. A total of 25 genetic samples of M. fascicularis yielding 383 bp of Cytochrome b (Cyt b) sequences were used in phylogenetic analysis along with one sample each of M. nemestrina and M. arctoides used as outgroups. Sequence character analysis reveals that Cyt b locus is a highly conserved region with only 23% parsimony informative character detected among ingroups. Further analysis indicates a clear separation between populations originating from different regions; the Malay Peninsula versus Borneo Insular, the East Coast versus West Coast of the Malay Peninsula, and the island versus mainland Malay Peninsula populations. Phylogenetic trees (NJ, MP and Bayesian) portray a consistent clustering paradigm as Borneo's population was distinguished from Peninsula's population (99% and 100% bootstrap value in NJ and MP respectively and 1.00 posterior probability in Bayesian trees). The East coast population was separated from other Peninsula populations (64% in NJ, 66% in MP and 0.53 posterior probability in Bayesian). West coast populations were divided into 2 clades: the North-South (47%/54% in NJ, 26/26% in MP and 1.00/0.80 posterior probability in Bayesian) and Island-Mainland (93% in NJ, 90% in MP and 1.00 posterior probability in Bayesian). The results confirm the previous morphological assignment of 2 subspecies, M. f. fascicularis and M. f. argentimembris, in the Malay Peninsula. These populations should be treated as separate genetic entities in order to conserve the genetic diversity of Malaysia's M. fascicularis. These findings are crucial in aiding the conservation management and translocation process of M. fascicularis populations in Malaysia.
Two new species of urothoid amphipods from Pulau Sibu and Pulau Tinggi, Johor are described and illustrated. The specimens of Urothoe sibuensis new species were collected by vertical haul plankton net and is distinctively different from other existing Urothoe species by these combination of special characters; similar gnathopods 1-2 with short and stout propodus expanded into poorly defined palms; large eyes and epimeron 3 smooth. Urothoe tinggiensis new species as collected using an airlift suction sampler at seagrass area is characterized by its different gnathopodal configuration with setose dactylus of 5th pereopod; eyes minute; carpus is wider than merus in the 5th pereopod; subquadrate coxa 4; merus and carpus of pereopods 6-7 are linear.
The Bornean representatives of the genus Georissa (Hydrocenidae) have small, dextral, conical, calcareous shells consisting of ca. three teleoconch whorls. Our recent study on the Georissa of Malaysian Borneo has revealed high intra- and inter-specific variation in the "scaly" group (a group of species with striking scale-like surface sculpture). The present study on the "non-scaly" Georissa is the continuation of the species revision for the genus. The "non-scaly" species are also diverse in shell sculptures. This informal group comprises Georissa with subtle spiral and/or radial sculpture. The combination of detailed conchological assessment and molecular analyses provides clear distinctions for each of the species. Conchological, molecular, and biogeographic details are presented for 16 species of "non-scaly" Georissa. Three of these are new to science, namely Georissacorrugatasp. n., Georissainsulaesp. n., and Georissatrusmadisp. n.
The family Symphytognathidae is reported from Thailand for the first time. Three new species: Anapistula choojaiae sp. nov., Crassignatha seeliam sp. nov., and Crassignatha seedam sp. nov. are described and illustrated. Distribution is expanded and additional morphological data are reported for Patu shiluensis Lin & Li, 2009. Specimens were collected in Thailand between July and August 2018. The newly described species were found in the north mountainous region of Chiang Mai, and Patu shiluensis was collected in the coastal region of Phuket. DNA sequences are provided for all the species here studied. The relations of these symphytognathid species were tested using previously published phylogenetic analyses on micro orb-weavers. Also, we used micro CT analysis to build 3D models of the male genitalia and somatic characters of two species of Crassignatha Wunderlich, 1995. The molecular phylogeny and 3D models were used to discuss the taxonomy and circumscription of the currently valid symphytognathid genera, with focus on Crassignatha and Patu Marples, 1951. Based on this, three new combinations are suggested: Crassignatha bicorniventris (Lin & Li, 2009), comb. nov., Crassignatha quadriventris (Lin & Li, 2009), comb. nov., and Crassignatha spinathoraxi (Lin & Li, 2009), comb. nov. A new record of Crassignatha danaugirangensisMiller et al. 2014 is reported from Brunei.