The genus Kelawakaju Maddison & Ruiz, gen. nov., is described for a lineage of bark-dwelling Asian marpissine jumping spiders that represent a dispersal to Eurasia separate from that of the Marpissa-Mendoza lineage, according to the phylogeny recovered from analysis of four gene regions. All species of Kelawakaju are new to science except Kelawakajufrenata (Simon, 1901), comb. nov., which is transferred from Ocrisiona Simon, 1901. Kelawakajufrenata is known from Hong Kong, Guangdong, Guangxi, and likely Taiwan. The five new species are Kelawakajumulu Maddison & Ruiz, sp. nov. (type species of Kelawakaju, from Sarawak, Malaysia, ♂♀), K.intexta Maddison & Ruiz, sp. nov. (from Sarawak, ♂), K.leucomelas Maddison & Ng, sp. nov. (Singapore and Johor Bahru, ♂♀), K.sahyadri Vishnudas, Maddison, & Sudhikumar, sp. nov. (India, ♂♀), and K.singapura Maddison & Ng, sp. nov. (Singapore, ♂♀).
Examination of numerous specimens characterised by predorsal scute, long maxilla, indented preopercle and pelvic scute lacking a spine and previously identified as Stolephorusbengalensis (Dutt & Babu Rao, 1959) or Stolephorusinsularis Hardenberg, 1933, revealed four distinct species, true S.bengalensis (distributed from the Bay of Bengal to Pakistan) and three new species, viz., Stolephoruseldorado sp. nov. (Taiwan to Java, Indonesia), Stolephorusdiabolus sp. nov. (Strait of Malacca, from Penang , Malaysia, to Singapore) and Stolephoruseclipsis sp. nov. (Bintan Island, Riau Archipelago, Indonesia). Characters separating the four species include numbers of gill rakers on each gill arch and vertebrae and pelvic fin and dorsal-fin ray lengths. Two molecular markers (mitochondrial cytochrome b and cytochrome oxidase I genes) demonstrated the distinction of three of the species examined morphologically and enabled a reconstruction of their phylogenetic relationships. Each species was genetically divergent from the others by 3.5%-7.7% mean uncorrected distance in the mitochondrial cytochrome oxidase I gene.
Although Borneo has received more attention from biologists than most other islands in the Malay Archipelago, many questions regarding the systematic relationships of Bornean mammals remain. Using next-generation sequencing technology, we obtained mitochondrial DNA sequences from the holotype of Suncusater, the only known specimen of this shrew. Several shrews collected recently in Sarawak are closely aligned, both morphologically and mitochondrially, with the holotype of S.ater. Phylogenetic analyses of mitochondrial sequences indicate that the S.ater holotype and new Sarawak specimens do not belong to the genus Suncus, but instead are most closely related to Palawanosorexmuscorum. Until now Palawanosorex has been known only from the neighboring Philippine island of Palawan. Additional sequences from nuclear ultra-conserved elements from the new Sarawak specimens strongly support a sister relationship to P.muscorum. We therefore transfer ater to Palawanosorex. The new specimens demonstrate that P.ater is more widespread in northern Borneo than previously recorded. Continued sampling of Bornean mammal diversity and reexamination of type material are critical in understanding the evolutionary history of the biologically rich Malay Archipelago.
A new species of Spirobranchus (Annelida: Serpulidae) is described based on specimens collected at the coastal Shonan area of Sagami Bay and the adjacent areas of Honshu, Japan. Spirobranchusakitsushima sp. nov. forms large aggregations in the intertidal rocky zone of warm-temperate Japanese shores. This species was referred to as Pomatoleioskraussii (Baird, 1864) until the monotypic genus Pomatoleios was synonymized with Spirobranchus. This new species is formally described based on morphologically distinct Japanese specimens with supporting DNA sequence data. The calcareous opercular endplate of Spirobranchusakitsushima sp. nov. lacks a distinct talon, but some specimens have a slight rounded swelling on the endplate underside, while in other species of the S.kraussii complex a talon is present, usually extended, and with bulges. We examined sub-fossil tube aggregations of the new species and suggest that such aggregation stranded ashore is a good indicator of vertical land movements (uplift and subsidence) resulting from past events, such as earthquakes, in Honshu, Japan.
Raorchestes is a speciose genus of bush frogs with high diversity occurring in the Western Ghats of India. Relatively fewer species have been recorded across India, through Bangladesh, southern China, into Vietnam and Peninsular Malaysia. Many bush frogs are morphologically cryptic and therefore remain undescribed. Here, a new species, Raorchestes rezakhani sp. nov., is described from northeastern Bangladesh based on morphological characters, genetics, and bioacoustics. The 16S rRNA gene distinguished this species from 48 known species of this genus. Bayesian Inference and Maximum Likelihood analyses indicated that the new species was most similar to R. tuberohumerus, a species found in the Western Ghats, and to R. gryllus, a species found in Vietnam. Bioacoustics indicated that their calls were similar in pattern to most Raorchestes species, although number of pulses, duration of pulses, pulse intervals and amplitude differentiated it from a few other species. It is suggested that northeastern India, Bangladesh, northern Myanmar, and southern China represent important, relatively unexplored areas that could yield additional species of Raorchestes. Since many remaining habitat patches in Bangladesh are under severe threat from deforestation, efforts should be made to protect these last patches from further degradation.
The genus Passiena is recorded for the first time from China with Passienaduanisp. nov. (♂♀) from Guangxi described here. In addition, the male of P.spinicrus Thorell, 1890 is described for the first time based on a specimen from Malaysia and colour photographs of freshly collected material are also presented. Detailed morphological descriptions, photographs, genital illustrations, and a distribution map for the two species are provided.
Sabah, Malaysia, is well known for its extensive and diverse coral reefs. It is located on the northwestern edge of the Coral Triangle, the region with the highest marine biodiversity. Much of the marine fauna here is still unknown, especially inconspicuous animals, such as small stoloniferous octocorals, which are common on coral reefs. Here, we describe two new monospecific genera of the family Arulidae found off the coast of Kota Kinabalu, Sabah, East Malaysia; Bunga payung gen. nov. et sp. nov. and Laeta waheedae gen. nov. et sp. nov. As well, the stoloniferan genus Phenganax Alderslade & McFadden, 2011 belonging to the family Clavulariidae is expanded with three new species, P. marumi sp. nov., P. subtilis sp. nov., and P. stokvisi sp. nov., which are all sclerite-free. Additionally, we report a possibly undescribed species, closely related to the clavulariid genera Azoriella Lopez-Gonzalez & Gili, 2001 and Cervera Lopez-Gonzalez et al., 1995. As this and other recent studies have shown, discoveries of small stoloniferous octocorals are helping to fill gaps in our knowledge of the overall systematics of Octocorallia.
Amphibians of Sekayu lowland forest have been studied more than a decade, with discoveries of new records of species showing no sign of abating between the years 2003 to 2020, indicating the remarkably rich diversity of anurans in this forest. Despite ceaseless anthropogenic activities in this area, this study successfully recorded 52 species of amphibians from 32 genera in the lowland forest of Sekayu. The species composition consisted of a single species from the family Ichthyophiidae and 51 species of anurans of 31 genera and six families. The number of species recorded has steadily increased especially during more recent surveys from 2015 to 2020. This study augments the total number of amphibian species recorded from Hulu Terengganu by ten additional species, increasing the total to 70 species for the district.
Euborellia (Anisolabididae: Anisolabidinae) is one of the most speciose genera of earwigs (Dermaptera), and its species-level classification is difficult. To settle the classification of brachypterous species with abbreviated tegmina recorded from East and Southeast Asia, we examined the morphology and reproductive isolation of three tentative Euborellia species, and analyzed the DNA barcoding region of the mitochondrial cytochrome oxidase subunit I (COI) gene. The observed complete reproductive isolation among the three Euborellia taxa and considerable differentiation in the COI sequences clearly show that each should be treated as a separate species. Based on morphology, distribution and the DNA sequence, we identify Euborellia sp. 1 of Malaysia as E.annulata (Fabricius), a circumtropical cosmopolitan with no records of a fully winged form. Samples from Ioto Island (= Iwo-jima Island: Ogasawara Islands, southern Japan) were also identified as this species. Euborellia sp. 3, from the main islands of Japan, was generally larger and lacked a Y-shaped pigmented area on the penis lobe, which is characteristic of Euborellia sp. 1. We propose reinstating E.pallipes (Shiraki) as the oldest name for this taxon. Euborellia sp. 2, even the brachypterous form, can be distinguished from these two species by its paler coloration (particularly the femora), ecarinate post-abdomen, and the shape of the male genitalia (parameres). We tentatively identify this species as E.philippinensis Srivastava based on the morphology of the brachypterous form, although the macropterous form cannot be distinguished from E.femoralis (Dohrn).
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 exploration of unsurveyed areas in southern Thailand discovered two new karst-adapted species, Cyrtodactylussungaiupesp. nov. and Cyrtodactyluswangkhramensissp. nov., from Thung Wa and La-ngu Districts, Satun Province, respectively. These new species are members of the C.pulchellus group that occur along the Thai-Malay Peninsula. The new species can be distinguished from all other congeners by their key morphological characters and genetic divergence. Morphologically, Cyrtodactylussungaiupesp. nov. and Cyrtodactyluswangkhramensissp. nov. can be diagnosed from other members by having a combination of differences in body size; degree of dorsal tuberculation; absence of tubercles on ventral surfaces; number of ventral scales, paravertebral tubercles and femoroprecloacal pores in males only; deep precloacal groove only in males; absence of a scattered pattern of white dorsal tubercles; number of dark body bands; and the extent of caudal tubercles on an original tail. Although the two species are sister taxa and have nearly identical morphologies, they are considered to be different species, based on a relatively high uncorrected pairwise genetic divergence of the mitochondrial ND2 gene (6.59-6.89%), statistically significant univariate and multivariate morphological differences (PERMANOVA and ANOVA) and diagnostic characteristics of caudal tuberculation on the original tail. Moreover, Cyrtodactylussungaiupesp. nov. and Cyrtodactyluswangkhramensissp. nov. are currently restricted to their karstic type localities which may serve as a geographic barrier to dispersal and gene flow.
The subgenus Dudaica Strand of the genus Drosophila Fallén has been known to comprise only two species: Drosophila (Dudaica) senilis Duda, 1926 (recorded from Indonesia, Philippines, Vietnam, Bhutan, and India) and D.malayana (Takada, 1976) (recorded from Malaysia). In the present study, this subgenus is revised, with D.malayana redescribed and six new species discovered and described from China, Malaysia, and Indonesia: gracilipalpis Katoh & Gao, sp. n., puberula Katoh & Gao, sp. n., albipalpis Katoh, Toda & Gao, sp. n., qiongzhouensis Katoh & Gao, sp. n., orthophallata Katoh, Toda & Gao, sp. n., and dissimilis Katoh & Gao, sp. n. Both morphological and molecular data (DNA barcodes) are used to distinguish the above species. A key to species of this subgenus is provided.
Species of the Oriental subtropical and tropical genus Phylladothrips of fungus-feeding thrips exhibit some diagnostic character states, usually with abdominal tergite VIII bearing two pairs of wing-retaining setae and male tergite IX setae S2 about as long as S1. These species are quite small, and the maxillary stylets unusually broad for Phlaeothripinae. Phylladothripstrisetaesp. nov. from Xizang, China and P.selangorsp. nov. from Selangor, Malaysia are described, and P.fasciae is newly recorded from China. All 11 species in this genus are revised with an illustrated key.
The Torrent frogs of the genus Amolops are widely distributed in Nepal and northern India eastwards to southern China and southwards to Malaysia. The genus currently contains 84 species. Previous studies indicated underestimated species diversity in the genus. In the context, a new species occurring from the mountains in the northwestern Guizhou Province, China is found and described based on morphological comparisons and molecular phylogenetic analyses, Amolopsdafangensissp. nov. Phylogenetic analyses based on DNA sequences of the mitochondrial 16S rRNA and COI genes supported the new species as an independent lineage. The uncorrected genetic distances between the 16S rRNA and COI genes in the new species and its closest congener were 0.7% and 2.6%, respectively, which are higher than or at the same level as those among many pairs of congeners. Morphologically, the new species can be distinguished from its congeners by a combination of the following characters: body size moderate (SVL 43.2-46.8 mm in males); head length larger than head width slightly; tympanum distinct, oval; vocal sacs absent; vomerine teeth present; dorsolateral folds weak formed by series of glands; nuptial pads present on the base of finger I; heels overlapping when thighs are positioned at right angles to the body; tibiotarsal articulation reaching the level far beyond the tip of the snout when leg stretched forward.
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.
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.
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.
This study documents information on the composition, diversity, richness, and temporal occurrence of snakes at Sekayu's lowland forest (SLF), Terengganu, Peninsular Malaysia for the first time. The snakes recorded within the SLF were sampled opportunistically from 2013 to 2019, employing the Visual Encounter Survey method (VES) and L-shape pitfall traps with drift fences. Forty-six snake species from 37 genera belonging to the nine families were recorded, of which 11 were new records to Terengganu. Individual-based rarefaction and extrapolation curves were not reaching asymptote, indicating that additional species can be recorded at the study area. Non-parametric species richness estimators estimated and produced a range between 51 and 57 species. ACE was the best estimator based on the quantitative evaluation. All species showed some variations of occurrence patterns across months. Fourteen species were only encountered once across the sampling years, and interestingly 11 of them were only detected during the rainy season (late October to January). In general, the number of species richness, abundance, and rare species were high during this season. Species richness of snakes is high at SLF but sampling effort should be intensified, especially during these rainy months, to obtain a robust estimated snake species richness in SLF. Terengganu harbor considerably high species richness of snakes with a total of 71 species to date (excluding marine snakes), but snake diversity is still underestimated as only a few localities were surveyed in the past years, primarily at the northern part. Future surveys should be commenced at the central and southern parts of Terengganu to complement the current investigation.