A new genus of the tribe Alysiini (Hymenoptera, Braconidae, Alysiinae) is described with specimens from India, Indonesia, Malaysia, Singapore, Thailand, and Vietnam, and six new species are described: Anamalysiaidiastimorpha sp. nov. (type species), A.knekosoma sp. nov., A.mellipes sp. nov., A.transversator sp. nov., A.vandervechti sp. nov., and A.vanhengstumi sp. nov.. We transfer one species from Coelalysia to Anamalysia: A.urbana (Papp, 1967) comb. nov. from Singapore and one species from Alysiasta to Anamalysia: A.triangulum (Fischer, 2006) comb. nov. from Malaysia, Laos, Indonesia and Vietnam. A key to the genus of Anamalysia is included.
A new genus of onchidiid slugs, Wallaconchis Goulding & Dayrat, gen. n., is described, including ten species. Five species were previously described but known only from the type material: Wallaconchis ater (Lesson, 1830), W. graniferum (Semper, 1880), W. nangkauriense (Plate, 1893), W. buetschlii (Stantschinsky, 1907), and W. gracile (Stantschinsky, 1907), all of which were originally classified in Onchidium Buchannan, 1800. Many new records are provided for these five species, which greatly expand their known geographic distributions. Five species are new: Wallaconchis achleitneri Goulding, sp. n., W. comendadori Goulding & Dayrat, sp. n., W. melanesiensis Goulding & Dayrat, sp. n., W. sinanui Goulding & Dayrat, sp. n., and W. uncinus Goulding & Dayrat, sp. n. Nine of the ten Wallaconchis species are found in the Coral Triangle (eastern Indonesia and the Philippines). Sympatry is high, with up to six species found on the island of Bohol (Philippines) and eight species overlapping in northern Sulawesi (Indonesia). Wallaconchis is distinguished from other onchidiids by its bright dorsal colors (red, yellow, orange) but those are extremely variable and not useful for specific identification. Internally, the reproductive system can be used to identify all Wallaconchis species. The copulatory organs of Wallaconchis species are especially diverse compared to other onchidiid genera, and the possible role of reproductive incompatibility in species diversification is discussed. All specimens examined were freshly collected for the purpose of a worldwide revision of the Onchidiidae Rafinesque, 1815. The species are well delineated using DNA sequences and comparative anatomy. Mitochondrial DNA analysis yields thirteen molecular units separated by a large barcode gap, while nuclear DNA yields nine units. By integrating nuclear DNA and mitochondrial DNA with morphology, ten species are recognized. The natural history of each species (e.g., the microhabitat where they are found) is also documented. Nomenclature is addressed thoroughly (the types of all onchidiid species were examined, lectotypes were designated when needed, nomina dubia are discussed). Morphological characters, transitions to new microhabitats, and diversification processes are discussed in the context of a robust molecular phylogeny.
The sarawakensis species group of the termitophilous carabid genus Orthogonius MacLeay, 1825 is defined and reviewed. Members of this species group are distributed in Southeast Asia and represented by four species, including two new species: Orthogonius sabahicus sp. n. (Sabah, northern Borneo, Malaysia) and Orthogonius morvanianus sp. n. (southern Thailand). A key to all species of the species group is also 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.
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.
Depreissia is a little known genus comprising two hymenopteran-mimicking species, one found in Central Africa and one in the north of Borneo. The male of Depreissia decipiens is redescribed, the female is described for the first time. The carapace is elongated, dorsally flattened and rhombus-shaped, the rear of the thorax laterally depressed and transformed, with a pair of deep pits; the pedicel is almost as long as the abdomen. The male palp is unusual, characterized by the transverse deeply split membranous tegulum separating a ventral part which bears a sclerotized tegular apophysis and a large dagger-like retrodirected median apophysis. The female epigyne consists of one pair of large adjacent spermathecae and very long copulatory ducts arising posteriorly and rising laterally alongside the spermathecae continuing in several vertical and horizontal coils over the anterior surface. Relationships within the Salticidae are discussed and an affinity with the Cocalodinae is suggested. Arguments are provided for a hypothesis that Depreissia decipiens is not ant-mimicking as was previously believed, but is a mimic of polistinine wasps. The species was found in the canopy in the Kinabalu area only, in primary and old secondary rainforest at 200-700 m.a.s.l. Overlap of canopy-dwelling spider species with those in the understorey are discussed and examples of species richness and endemism in the canopy are highlighted. Canopy fogging is a very efficient method of collecting for most arthropods. The canopy fauna adds an extra dimension to the known biodiversity of the tropical rainforest. In southeast Asia, canopy research has been neglected, inhibiting evaluation of comparative results of this canopy project with that from other regions. More use of fogging as a collecting method would greatly improve insight into the actual species richness and species distribution in general.
Five sea cucumber species including one new species of the genus Stichopus are reported from the shallow coral reefs of Straits of Malacca. The new species Stichopus fusiformiossa has unusual fusiform spicules in the tentacles, which are not found in the other species of the genus. Pseudo-tables and large perforated plates are newly recorded for Stichopus hermanni Semper, 1868 and Stichopus vastus Sluiter, 1887, respectively.
Octomeris is a chthamalid intertidal barnacle with eight shell plates. There are currently two species of such barnacles: O. brunnea Darwin, 1854 (type locality in the Philippines), common in the Indo-Pacific region, and O. angulosa Sowerby, 1825, only recorded in South Africa. Octomeris intermedia Nilsson-Cantell, 1921, identified from the Mergui Archipelago in Myanmar, was considered to be conspecific with O. brunnea by Hiro (1939) based on samples collected in Taiwan. The morphological differences in shell and opercular plates between O. brunnea and O. intermedia are believed to be intra-specific variations due to different degrees of shell erosion. In the present study, the genetic and morphological differentiations of Octomeris in the Indo-Pacific region were examined. This study found two molecular clades (with inter-specific differences) based on the divergence in the COI genes, and the species also have distinct geographical distributions. The Octomeris brunnea clade covers samples collected from the Philippines and Taiwan waters and the other clade, which we argue is O. intermedia, is distributed in Phuket and Krabi, Thailand and Langkawi, Malaysia. Phuket and Krabi are located approximately 300 km south of the Mergui Archipelago, the type locality of O. intermedia. The morphology of samples collected from Thailand fits the type description of O. intermedia in Nilsson-Cantell (1921). Our study concludes that O. intermedia is a valid species based on morphological and molecular evidence.
The genus Dichaetophora Duda comprises 61 described species classified into four species groups: agbo, tenuicauda, acutissima and sinensis. This genus is distributed exclusively in the Old World, and is rich in species in the tropical and subtropical areas of the Oriental, Australasian, and Afrotropical regions. In this paper, a new species group, the trilobita group, is established for six new species discovered from the Oriental region. The delimitation of these species is firstly performed in light of morphology and further with the aid of DNA sequences of the mitochondrial COI and COII (cytochrome c oxydase, subunits I and II, respectively) genes, considering also their respective geographical origins. Then, the new species (trilobita Yang & Gao, sp. n., heterochroma Yang & Gao, sp. n., flatosternata Yang & Gao, sp. n., borneoensis Yang & Gao, sp. n., javaensis Yang & Gao, sp. n., and sumatraensis Yang & Gao, sp. n.) are described, and a key, based on not only morphological but also molecular information, is provided.
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.
Material collected between 2010 and 2014 on the Indonesian islands of Sumatra, Bali, Sumba, Sumbawa, Sulawesi, and Seram unveiled the enormous diversity of Labiobaetis Novikova & Kluge in this country. Five species were reported from Indonesia previously (L. fulmeki (Ulmer), L. obscurum (Ulmer), L. necopinatum (Müller-Liebenau), L. ulmeri (Müller-Liebenau), and L. boettgeri (Ulmer)); all were described from adults only and no species were previously known at larval stage. We identified 18 new species by integrative taxonomy using genetic distance (COI, Kimura-2-parameter) and morphology, and they are described and illustrated based on their larvae. Another species, L. multus (Müller-Liebenau) from Malaysia, was also found in Indonesia, increasing the total number of species in Indonesia to 24. Seven morpho-groups of species are proposed based on morphological characters and a key to the larvae of all species from Indonesia and adjacent countries is provided. The total number of Labiobaetis species worldwide is augmented to 123. The examination of the new species allowed us to slightly modify the generic attributes of the larvae. The interspecific K2P distances are usually between 11% and 24%, the intraspecific distances are usually between 0% and 3%. The remarkable richness of the genus in Indonesia is discussed.
Material collected between 2000 and 2014 on the island Borneo, including the Indonesian province of Kalimantan, the Malaysian province of Sabah and Brunei Darussalam, substantially increased our knowledge of Labiobaetis on this island. The total number of Labiobaetis species in Borneo increased to five, as only one species, L. borneoensis (Müller-Liebenau, 1984), was previously reported. Three new species were identified by morphology and partly by using genetic distance (COI, Kimura 2-parameter). They are described and illustrated based on their larvae (Labiobaetis bakerae sp. nov., L. penan sp. nov. and L. dayakorum sp. nov.); in one case, the imago is described as well. New reports of L. borneoensis are presented and the imago of this species is described for the first time. Labiobaetis moriharai (Müller-Liebenau, 1984), originally described from mainland Malaysia (Province Selangor), is reported from Borneo for the first time. The interspecific K2P distances in Borneo are between 19% and 25%, the intraspecific distances are usually between 0% and 1%. The total number of Labiobaetis species worldwide is augmented to 126.
A reinvestigation of type and other material of Baetisjavanicus Ulmer, 1913 and Baetissabahensis Müller-Liebenau, 1984, together with new material from Southeast Asia revealed a new genus, Branchiobaetis gen. nov. The above species are formally assigned to the new genus Branchiobaetis gen. nov. It is characterized by the presence of accessory gills ventrally near fore coxa and at the base of maxillae, a peculiar folding of the gonostyli developing under the cuticle of last instar male larvae, together with a unique combination of other larval characters. Besides the two formerly described species, five new species are identified using a combination of morphology and molecular characters (COI, Kimura 2-parameter distances), four species from Sumatra and one from the Philippines. They are described and illustrated at the larval stage. Additionally, a complementary description of larva and adult stages of the generic type species B.javanicus comb. nov. as well as the first description of the eggs are provided. Furthermore, new reports of B.javanicus comb. nov. and B.sabahensis comb. nov. are indicated. The distribution of Branchiobaetis gen. nov. includes the Indonesian Sunda Islands, Borneo, and the Philippines. A key to the larval stage of all species is provided.
Species identities of Goniobranchus nudibranchs with white bodies and various marginal bands have long been problematic. In this study, specimens of these Goniobranchus nudibranchs from the Philippines, Peninsular Malaysia, Japan, Papua New Guinea, and Madagascar were analyzed and molecular data were obtained in order to re-examine the relationships between species within this "white Goniobranchus with marginal bands" group. The analyses clearly recovered six species groups corresponding to the described species Goniobranchusalbonares, G.preciosus, G.rubrocornutus, G.sinensis, and G.verrieri as well as one new species, G.fabulus Soong & Gosliner, sp. nov. Notably, G.preciosus, G.sinensis, G.rubrocornutus, G.verrieri, and G.fabulus Soong & Gosliner, sp. nov. exhibit color variation and polymorphism, suggesting that some aspects of color patterns (e.g., presence or absence of dorsal spots) may not always be useful in the identification of species in the "white Goniobranchus with marginal bands" group, whereas other features such as gill and rhinophore colors and the arrangement and colors of the mantle marginal bands are more diagnostic for each species.
In this paper, four species of the genus Diduga Moore, [1887] from China (Chongqing and Guangdong) and Malaysia (Borneo, Sabah) are described as new to science, namely D. simianshanasp. nov., D. chebalingasp. nov., D. chewisp. nov., and D. hollowayisp. nov. Adults of these species are illustrated in color, and images of the male and female genitalia are provided. A distribution map of the new species is provided, together with an updated checklist of all species of Diduga.
Three myrmecophilous genera of Aleocharinae (Staphylinidae) associated with Leptogenys distinguenda (Emery, 1887) and Leptogenys mutabilis (Smith, 1861) are reviewed with descriptions of new taxa: Maschwitzia Kistner, 1989, Togpelenys Kistner, 1989 and Witteia Maruyama & von Beeren, gen. n. (type species: Witteia dentilabrumMaruyama & von Beeren, sp. n.). The following new combinations are proposed: Zyras (s. lat.) aenictophila (Kistner, 1997),comb. n. (exMaschwitzia), Zyras (s. lat.) dichthadiaphila (Kistner in Kistner et al., 2003), comb. n. (exMaschwitzia), Maschwitzia derougemonti (Pace, 1984), comb. n. (exWroughtonilla Wasmann, 1899), Maschwitzia watanabei (Maruyama, 2004), comb. n. (exWroughtonilla), Maschwitzia dilatata (Pace, 2005), comb. n. (exWroughtonilla), Witteia borneensis (Pace, 1986), comb. n. (ex Wroughtonilla). These genera belong to the Wroughtonilla genus group of the tribe Lomechusini.
Rompindessus jenisi Balke, Bergsten & Hendrich, gen. n. et sp. n. is described from near Rompin village in West Malaysia. The new genus is characterized by the presence of an occipital line and basal pronotal striae, the presence of a thick anterior bead on clypeus and two-segmented parameres as well as by the absence of basal elytral striae, the absence of sutural line on elytron, the absence of basal epipleural transverse carina, and the absence of longitudinal elytral carina. Moreover, male pro- and mesotarsus appear stout, and distinctly dilated laterally; the pronotum is comparably long and parallel-sided and the colour of beetle conspicuous dark orange. Leiodytes kualalipis Balke, Wang, Bergsten & Hendrich, sp. n. is described from West Malaysia (Pahang) and South Vietnam (Cat Tien). It is well characterized by its large size, elongate body and the form of the median lobe. Limbodessus fijiensis (J. Balfour-Browne, 1944), comb. n. described from Fiji is a new synonym of Limbodessus curviplicatus (Zimmermann, 1927) described from Samoa.
A new silvanid genus Borneophanusgen. n. is described based on specimens collected from Malaysian Borneo. A new species, B.spinosussp. n., is described herein. Digitiform sensilla on the apical maxillary palpomere is reported in Silvanidae for the first time.
Layang-Layang is a small island part of an oceanic atoll in the Spratly Islands off Sabah, Malaysia. As the reef coral fauna in this part of the South China Sea is poorly known, a survey was carried out in 2013 to study the species composition of the scleractinian coral families Fungiidae, Agariciidae and Euphylliidae. A total of 56 species was recorded. The addition of three previously reported coral species brings the total to 59, consisting of 32 Fungiidae, 22 Agariciidae, and five Euphylliidae. Of these, 32 species are new records for Layang-Layang, which include five rarely reported species, i.e., the fungiids Lithophyllonranjithi, Podabaciasinai, Sandalolithaboucheti, and the agariciids Leptoseriskalayaanensis and Leptoseristroglodyta. The coral fauna of Layang-Layang is poor compared to other areas in Sabah, which may be related to its recovery from a crown-of-thorns seastar outbreak in 2010, and its low habitat diversity, which is dominated by reef slopes consisting of steep outer walls. Based on integrative molecular and morphological analyses, a Pavona variety with small and extremely thin coralla was revealed as Pavonamaldivensis. Since specimens from Sabah previously identified as Pavonamaldivensis were found to belong to Pavonaexplanulata, the affinities and distinctions of Pavonamaldivensis and Pavonaexplanulata are discussed.