The species of the genus Coeligetes Jacoby, 1884 distributed in Malaysia and Indonesia are revised, illustrated and keyed. New species, C. howardi sp. nov. from Borneo is described. New synonymy Coeligetes submetallica Jacoby, 1884 = C. wilcoxi Mohamedsaid, 1994 (syn. nov.) is proposed. New genus and species Coeligetoides trifurcatus gen. nov., sp. nov. (Malaysia, Brunei, Indonesia and Thailand) is described, illustrated and compared with related genera.
The areolate Oriental family Heteropterygidae Kirby, 1893 is critically reviewed and the results of the present study contradict the arrangement suggested by Zompro (2004), but in most aspects agree with a molecular study presented by Whiting et al (2003) and a phylogenetic study presented by Bradler (2009). The family is critically discussed and new hypotheses are presented for the phylogeny and intra-familiar relationships, placing the subfamily Dataminae Rehn & Rehn, 1939 as the basalmost clade of Heteropterygidae. The subfamilies Obriminae Brunner v. Wattenwyl, 1893 and Heteropteryginae Kirby, 1893 together represent the sister-group of Dataminae. Arguments and a tree are presented to support this hypothesis. New diagnoses and lists of genera are provided for all three subfamilies contained in Heteropterygidae, along with keys to distinguish between them. The subfamily Obriminae is critically reviewed and the distinction between the three tribes Obrimini Brunner v. Wattenwyl, 1893, Eubulidini Zompro, 2004 and Miroceramiini Zompro, 2004 introduced by Zompro (2004) is shown to be poorly supported. While Obrimini sensu Zompro, 2004 is generally accepted (but now also contains genera that were placed in Eubulidini or Miroceramiini by Zompro (2004)), the tribes Eubulidini and Miroceramiini are not supported. A new arrangement is introduced, which is based on morphological characters neglected or overlooked by Zompro (2004) but were partly discussed by Bradler (2009). The genus Mearnsiana Rehn & Rehn, 1939 is removed from Miroceramiini and transferred to Obrimini. The genera Eubulides Stål, 1877, Heterocopus Redtenbacher, 1906, Theramenes Stål, 1875 and Stenobrimus Redtenbacher, 1906 are removed from Eubulidini and also transferred to Obrimini. Consequently, Eubulidini is synonymised with Obrimini (n. syn.). Miroceramiini is a monotypical tribe and only includes the Wallacean genus Miroceramia Günther, 1934. The new tribe Tisamenini n. trib. is established for the three basal genera Tisamenus Stål, 1875, Ilocano Rehn & Rehn, 1939 and Hoploclonia Stål, 1875 all of which were placed in Eubulidini by Zompro (2004). The latter genus differs from the other two genera by the morphology of the female genitalia, which is unique amongst the entire family. Three generic groups are recognized within Obrimini, the Obrimus-group, Stenobrimus-group and Theramenes-group. Keys are presented to distinguish between the three tribes now contained in the Obriminae, i.e. Obrimini, Tisamenini n. trib. and Miroceramiini. The genus Hennobrimus Conle, 2006 is synonymised with Mearnsiana Rehn & Rehn, 1939, based on the fact that the type-species of both genera are conspecific (n. syn.). Hennobrimus hennemanni Conle, 2006, the type-species of Hennobrimus, and Trachyaretaon manobo Lit & Eusebio, 2005 are synonymised with Mearnsiana bullosa Rehn & Rehn, 1939, the type-species of Mearnsiana (n. syn.). Theramenes dromedarius Stål, 1877 from the Philippines is removed from synonymy with the Wallacean Theramenes olivaceus (Westwood, 1859) and re-established as a valid species (rev. stat.). The subfamily Heteropteryginae Kirby, 1896 is revised at the species-level and a new diagnosis is presented. Keys to the two genera and all 16 known species are provided along with new descriptions, differential diagnoses, lists of examined material, detailed information on the known distributions, measurements and illustrations of the insects and eggs. The intra-subfamiliar and intra-generic relationships are discussed and a cladogram is presented. Heteropteryginae contains two genera: Heteropteryx Gray, 1835 (Type-species: Phasma dilatatum Parkinson, 1798) and Haaniella Kirby, 1896 (Type-species: Phasma (Heteropteryx) muelleri de Haan, 1842). The distribution of this subfamily is restricted to Sundaland with the exception of a single species that is found in Vietnam. All other species are distributed in Borneo, Sumatra, the Mentawai Islands, Singapore, Peninsular Malaysia and Thailand. Heteropteryginae contains the largest and most striking members of the entire family Heteropteryginae, some of which are amongst the heaviest insects known. The subfamily is characterized by apomorphies such as the presence of wings, having a tympanal area (= stridulatory organ) in the basal portion of the alae, straight profemora, strongly shortened tarsi, lack of rough sensory-areas on the prosternum and typically X-shaped micropylar plate of the eggs. The sister-group of Heteropteryginae is represented by the Obriminae, with which it shares a beak-like secondary ovipositor in the females and presence of a medio-apical spine on the area apicalis. Both features are synapomorphies of Heteropteryginae + Obriminae. The genus Haaniella Kirby, 1904 contains 16 known species, five of which are newly described herein. The genus Miniopteryx Zompro, 2004 (Type-species: Haaniella parva Günther, 1944) is synonymised with Haaniella on the basis that the distinguishing feature mentioned in the original description is a character that is frequently found throughout the genus (n. syn.). The type-species H. parva Günther, 1944 is automatically retransferred to Haaniella (rev. stat.). Haaniella aculeata n. sp. from western Sumatra is described from the male. Haaniella macroptera n. sp. from Singapore and the Johor state in southern Peninsular Malaysia is described from both sexes and the eggs. Haaniella gintingi n. sp. from Central Sumatra is described from both sexes and the eggs and Haaniella kerincia n. sp. from Western Sumatra is described from the insects only, the eggs being still unknown. One new species, Haaniella gorochovi n. sp., is the only representative of the genus and subfamily Heteropteryginae known from Vietnam and both sexes as well as the eggs are described. Haaniella erringtoniae (Redtenbacher, 1906) is endemic in Peninsular Malaysia, here removed from synonymy with H. muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.). The Sumatran Haaniella glaber (Redtenbacher, 1906) is removed from synonymy with H. muelleri (Haan, 1842) and re-established as a valid species (rev. stat.). Leocrates glaber Redtenbacher, 1906 and Haaniella muelleri simplex Günther, 1944 are removed from synonymy with H. muelleri (Haan, 1842) (rev. stat.) and synonymised with H. glaber. Haaniella mecheli (Redtenbacher, 1906) and H. rosenbergii (Kaup, 1871) are removed from synonymy with H. muelleri (Haan, 1842) and re-established as valid species (rev. stat.). Haaniella erringtoniae novaeguineae Günther, 1934 and Haaniella muelleri var. b. (Haan, 1842) are synonymized with H. rosenbergii (Kaup, 1871) (n. syn.). The type-species Haaniella muelleri (Haan, 1842) is shown to be a fairly rare species that is restricted to Sumatra. All subsequent records of H. muelleri from outside Sumatra and references to captive breeding of stock originating from Peninsular Malaysia in Europe relate to H. erringtoniae (Redtenbacher, 1906). The previously unknown males and eggs of H. rosenbergii (Kaup, 1871) as well as the previously unknown females and eggs of H. parva Günther, 1944 are described and illustrated for the first time. Based on morphological characters of the insects and eggs three distinct species-groups are recognized within Haaniella. The muelleri species-group contains nine species that are distributed throughout Sumatra, the Mentawei Islands, Singapore and Peninsular Malaysia. These are characterized by the smooth ventral surface of the meso- and metafemora and lemon-shaped eggs which entirely lack the setae seen in the two other species-groups. The grayii species-group comprises four species, two of which are endemic in Borneo, one endemic in Sumatra and the fourth species being the only known representative of the subfamily in Vietnam. These species are characteristic for the prominent pair of spines on the abdominal tergites II-IV of males and long apically multidentate epiproct of females. The echinata species-group contains three exceptionally Bornean species, which are characterized by the long and apically pointed subgenital plate of females, which clearly projects beyond the epiproct, as well as the sub-basal lateral tooth of the anal segment of males. The muelleri species-group is sister to the remainder two species-groups. Heteropteryx Gray, 1853 is a monotypical genus and only contains the type-species H. dilatata (Parkinson, 1798), which is found throughout Peninsular Malaysia, Thailand, Sumatra and Northeastern Borneo. This genus differs from Haaniella by the strongly conically elevated head, which posteriorly projects over the anterior margin of the pronotum, females being bright green or yellow in colour with plain and translucent pink alae and having distinct spines on the abdominal tergites, and males having a strongly shortened mesothorax and dull pink alae. Lectotypes are designated for Haaniella parva Günther, 1944, Heteropteryx echinata Redtenbacher, 1906, Heteropteryx saussurei Redtenbacher, 1906 and Heteropteryx scabra Redtenbacher, 1906 to guarantee stability of these names. Information on the habitats, host-plants, biology, life cycle, parasitism and captive breeding of the species of Heteropteryginae is presented and a list summarising all taxonomic changes presented herein.
The Oriental pselaphine genus Horniella Raffray, 1905 (tribe Tyrini: subtribe Somatipionina) is redefined and revised. Twenty-five new species are described: H. centralis Yin & Li, sp. n., H. confragosa Yin & Li, sp. n., H. dao Yin & Li, sp. n., H. hongkongensis Yin & Li, sp. n., H. nakhi Yin & Li, sp. n., H. schuelkei Yin & Li, sp. n., H. sichuanica Yin & Li, sp. n., H. simplaria Yin & Li, sp. n., and H. tianmuensis Yin & Li, sp. n. from China, H. himalayica Yin & Li, sp. n. from Nepal and North India, H. asymmetrica Yin & Li, sp. n., H. burckhardti Yin & Li, sp. n., H. intricata Yin & Li, sp. n., H. kaengkrachan Yin & Li, sp. n., H. khaosabap Yin & Li, sp. n., H. loebli Yin & Li, sp. n., H. phuphaman Yin & Li, sp. n., H. prolixo Yin & Li, sp. n., and H. schwendingeri Yin & Li, sp. n. from Thailand, H. philippina Yin & Li, sp. n. from the Philippines, H. awana Yin & Li, sp. n., H. gigas Yin & Li, sp. n., H. pilosa Yin & Li, sp. n., and H. smetanai Yin & Li, sp. n. from Malaysia, and H. cibodas Yin & Li, sp. n. from Indonesia. The two previously described species, H. hirtella Raffray, 1901 (type species) from Sri Lanka and H. falcis Yin & Li, 2010 from China are redescribed, and a lectotype is designated for H. hirtella. Illustrations of habitus and important diagnostic features, an identification key, and distributional maps for all species are provided. Eleven unidentified species represented only by females are left unnamed. Illustrations of the habitus and the genital complex, and label data of these species are given to facilitate future study. All available data indicates that species of Horniella typically inhabit leaf litter of various kinds of forests, and can be most efficiently collected by sifting and use of Winkler-Moczarski extractors.
A taxonomic revision and phylogenetic analysis of the spider genus Glenognatha Simon, 1887 is presented. This analysis is based on a data set including 24 Glenognatha species plus eight outgroups representing three related tetragnathine genera and one metaine as the root. These taxa were scored for 78 morphological characters. Parsimony was used as the optimality criterion and a sensitivity analysis was performed using different character weighting concavities. Seven unambiguous synapomorphies support the monophyly of Glenognatha. Some internal clades within the genus are well-supported and its relationships are discussed. Glenognatha as recovered includes 27 species, four of them only known from males. A species identification key and distribution maps are provided for all. New morphological data are also presented for thirteen previously described species. Glenognatha has a broad distribution occupying the Neartic, Afrotropic, Indo-Malaya, Oceania and Paleartic regions, but is more diverse in the Neotropics. The following eleven new species are described: G. vivianae n. sp., G. caaguara n. sp., G. boraceia n. sp. and G. timbira n. sp. from southeast Brazil, G. caparu n. sp., G. januari n. sp. and G. camisea n. sp. from the Amazonian region, G. mendezi n. sp., G. florezi n. sp. and G. patriceae n. sp. from northern Andes and G. gouldi n. sp. from Southern United States and central Mexico. Females of G. minuta Banks, 1898, G. gaujoni Simon, 1895 and G. gloriae (Petrunkevitch, 1930) and males of G. globosa (Petrunkevitch, 1925) and G. hirsutissima (Berland, 1935) are described for the first time. Three new combinations are proposed in congruence with the phylogenetic results: G. argyrostilba (O. P.-Cambridge, 1876) n. comb., G. dentata (Zhu & Wen, 1978) n. comb. and G. tangi (Zhu, Song & Zhang, 2003) n. comb., all previously included in Dyschiriognatha Simon, 1893. The following taxa are newly synonymized: Dyschiriognatha montana Simon, 1897, Glenognatha mira Bryant, 1945 and Glenognatha maelfaiti Baert, 1987 with Glenognatha argyrostilba (Pickard-Cambridge, 1876) and Glenognatha centralis Chamberlin, 1925 with Glenognatha minuta Banks, 1898.
Species of Signoretia Stål from the Oriental region are reviewed and types of five species described by Baker, two species described by Distant and one species described by Schmidt are illustrated. A checklist of 20 species of the genus from the Oriental region including 9 new species is given. The new species described and illustrated are Signoretia dulitensis sp. nov. (Malaysia: Mt Dulit), S. lunglei sp. nov. (India: Mizoram), S. mishmiensis sp. nov. (Myanmar: Mishmi Hills), S. quoinensis sp. nov. (Malaysia: Quoin Hill), S. rubra sp. nov. (Thailand: Chiang Mai), S. sahyadrica sp. nov. (India: Kerala), S. similaris sp. nov. (Vietnam: Fyan), S. sinuata sp. nov. (India: West Bengal) and S. takiyae sp. nov. (India: Andaman Is.). Both S. aureola Distant and S. maculata Baker are redescribed and illustrated. Lectotypes are designated for S. greeni Distant and S. aureola Distant.
The bee fly genus Euchariomyia Bigot is reviewed and new records from the Oriental Region are given. Five names (for four species-level taxa) have been associated with species in the genus. Examinations of types, as well as homotypic and topotypic specimens, shows all five names to belong to a highly variable single species, Euchariomyia dives Bigot. The following species are here shown to be the same as Euchariomyia dives Bigot: Bombylius pulchellus Wulp, 1880, Bombylius scintillans Brunetti, 1909, and Bombylius brunettii Senior-White, 1922, n. syn. The genus is known primarily from the southern and eastern Oriental Region and ranges into the Palaearctic in eastern China. We extend the distribution of the genus with new records in the southeastern Oriental Region [Indonesia (Sumatra), Laos, Peninsular Malaysia, and Vietnam].
The apparently rare chelonine wasp genus Wushenia Zettel was previously known only from a single species Wushenia nana Zettel, collected by Townes at 1150 m from Wushe, Taiwan in 1983. Here we describe a second species, Wushenia australiensis sp. nov. from coastal New South Wales, Australia. This second species extends the known distribution of the genus from the Oriental into the Australasian region, indicating either an extreme disjunct distribution or that Wushenia may also occur on the landmasses inbetween, e.g. the Philippines, Malaysia, Indonesia and/or Papua New Guinea. In addition to a detailed description of the new species, a re-diagnosis of the genus and type species, and a key to species are presented.
A new species of Narrow-mouthed frog of the genus Kaloula is described from northern Peninsular Malaysia. Kaloula latidisca sp. nov. is genetically and morphologically most similar to K. baleata and K. indochinensis but differs from those and other congeners by the unique combination of the following characters: (1) adult males SVL 49.2-56.2 mm (x̅=53.5 ± 3.0; N=4); (2) finger tips expanded into large, transversely expanded discs (disc width 2.8-3.1 mm, x̅=3.0 ± 0.1); (3) inner metatarsal tubercle large, oval, distinctly raised, slightly shorter than first toe; (4) three subarticular tubercles on fourth toe; (5) toe webbing formula: I 1-2 II 1-3 III 2-3.5 IV 4-2 V; and (6) yellow to orange irregularly shaped patch on the axillary, inguinal and posterior region of thigh.
The ground beetle genus Hexachaetus Chaudoir, 1871 is re-defined and reviewed. Bearing six setae on ligula is no more considered as a crucial characteristic for Hexachaetus. Members of Hexachaetus share the following combination of morphological features: body polish, smooth, and impunctate, ligula more or less dilated at apex, bearing 4, 6, or even 12 setae apically, prosternal process unbordered at apex, elytra distinctly and obliquely truncated at apex, with the apical inner angles very sharp in most species (except for H. mulan n. sp.), and interval 3 with anterior and posterior setiferous pores (median one lacking). The members of Hexachaetus are about 20 species which could be divided into six species groups. All except angulatus species group are dealt with in this paper, with descriptions of four new species: H. kirschenhoferi n. sp. (Indonesia: Kalimantan), H. brunki n. sp. (Malaysia: N. Borneo), H. vietnamensis n. sp. (Vietnam: Annam) and H. mulan n. sp. (Malaysia: Perak and Pahang). H. maindroni Tian & Deuve, 2006 is proposed as a subspecies of H. lateralis Guérin, 1843, n. stat. A key to species groups and species of the genus is also provided.
Prodasineura yulan is described from a male from Maludam National Park, Betong Division, Sarawak, Malaysian Borneo. It is allied to Prodasineura interrupta.
Tree snails of the subgenus Amphidromus s. str. are unusual because of the chiral dimorphism that exists in many species, with clockwise (dextrally) and counter-clockwise (sinistrally) coiled individuals co-occurring in the same population. Given that mating in snails is normally impeded when the two partners have opposite coil, positive frequency-dependent selection should prevent such dimorphism from persisting. We test the hypothesis that a strong population structure with little movement between tree-based demes may result in the fixation of coiling morphs at a very small spatial scale, but apparent dimorphism at all larger scales. To do so, we describe the spatial structure in a Malaysian population of A. inversus (Müller, 1774) with 36% dextrals. We marked almost 700 juvenile and adult snails in a piece of forest consisting of 92 separate trees, and recorded dispersal and the proportions of dextrals and sinistrals in all trees over a 7-day period. We observed frequent movement between trees (155 events), and found that no trees had snail populations with proportions of dextrals and sinistrals that were significantly different from random. Upon recapture 1 year later, almost two-thirds of the snails had moved away from their original tree. We conclude that population structure alone cannot stabilise the coil dimorphism in Amphidromus.
Phaenandrogomphus safei is described from a male from the Kalabakan Forest Reserve, Sabah, Malaysian Borneo. It is the first species of Phaenandrogomphus to be recorded from Borneo. Onychogomphus treadawayi, known from Busuanga Island in the Palawan region of the Philippines, is transferred to Phaenandrogomphus.
A new species of gecarcinucid freshwater crab of the genus Parathelphusa H. Milne Edwards, 1853, is described from freshwater swamp habitats in Pekanbaru, Riau Province, in central-eastern Sumatra, Indonesia. Parathelphusa pardus sp. nov., has a very distinctive colour pattern, and in this respect, resembles P. maindroni (Rathbun, 1902) from Sumatra and Peninsular Malaysia; P. batamensis Ng, 1992, from Batam Island, Indonesia; P. reticulata Ng, 1990, from Singapore; and P. oxygona Nobili, 1901, from western Sarawak. It can be distinguished from these species and congeners by a suite of carapace, ambulatory leg, thoracic sternal and most importantly, male first gonopod characters.
Three species of the genus Amphiesma Duméril, Bibron & Duméril, 1854 have long been confused in the literature, with each other and with other species of the genus. Amphiesma khasiense (Boulenger, 1890) has been considered to inhabit a large geographical region, extending from north-eastern India, east to Vietnam and southern Thailand. Amphiesma boulengeri (Gressitt, 1937) has been regarded as a species endemic to south-eastern China. Amphiesma inas (Laidlaw, 1901) has been recorded from West Malaysia, Thailand and Indonesia (Sumatra). A multivariate analysis of morphometric and meristic characters shows that these three species can be separated by combinations of characters in the scalation and pattern, the most obvious being the structure of the postocular streak. On the basis of our analysis and after comparison with name-bearing type specimens, Amphiesma khasiense is restricted to north-eastern India, Myanmar, western Yunnan Province of China, northern Laos and northern and western Thailand. Other populations from south-eastern China, Vietnam, other parts of Laos, Cambodia and central Thailand, which have been recorded in the literature as A. khasiense, A.johannis or Amphiesma modestum (Günther, 1875), should be referred to Amphiesma boulengeri. Amphiesma inas (Laidlaw, 1901) is a valid species endemic to mountain ranges of southern Peninsular Thailand and West Malaysia. The mention of Amphiesma inas in Sumatra is erroneous, being based on the second known specimen of Amphiesma kerinciense David & Das, 2003, which is here redescribed. A key to species of the Amphiesma khasiense group and other species sharing a greyish-brown background without conspicuous dark and pale stripes, is provided.
The previously known distribution area of the genus Grossander Slater, 1976 (Hemiptera, Heteroptera, Rhyparochromidae, Drymini) is broadened with the description of two new species: Grossander papuanus sp. nov. (New Guinea) and Grossander eylesi sp. nov. (Burma, Laos, Thailand, Malaysia and Indonesia). Grossander (Oculoander) subgen. nov. is created for these new taxa. Drawings of habitus and male genitalia are presented. Keys to the subgenera of Grossander, and to the species of the new subgenus are provided.
Four new species of black flies are described, and three others are reported as newly recorded, based on adults reared from pupae, pupae and larvae collected in and near Tam Dao National Park, Vinh Phuc Province, Vietnam. New species include Simulium (Gomphostilbia) hongthaii sp. nov., S. (G.) tamdaoense sp. nov. (both species placed in the asakoae species-group), S. (Simulium) taythienense sp. nov. and S. (S.) xuandai sp. nov. (the two latter species placed in the striatum species-group). Newly recorded species are S. (G.) brinchangense Takaoka, Sofian-Azirun & Hashim, S. (Nevermannia) aureohirtum Brunetti and S. (S.) brevipar Takaoka & Davies. These discoveries increase the number of species of black flies known in Vietnam from 21 to 28.
A new species, Megatyrus femoralis sp. n., is described from the Koshi Zone, East Nepal, with major diagnostic features illustrated. Megatyrus masumotoi Nomura, Sakchoowong & Chanpaisaeng, originally described from southwestern Thailand, is recorded from the Noring Timur Mountain, West Malaysia. The above data extends the known range of Megatyrus about 1,200 km to the west, and 870 km to the south.
Mycomya Rondani specimens from the islands of South-East Asia, i.e. Malaysia, Indonesia and the Philippines, are revised. The paper includes a key to the Mycomya species of the South-East Asian islands. The following six new species are described: M. shimai sp. n. from Java, Indonesia, M. pongo sp. n. from Sabah, Malaysia, and M. apoensis sp. n., M. nakanishii sp. n., M. paraklossi sp. n. and M. yatai sp. n. from Mindanao, the Philippines. The holotypes of M. klossi Edwards from Borneo, Malaysia, and M. minutata Edwards from Sumatra, Indonesia, were examined and their genitalia are described. M. occultans (Winnertz) is recorded from Java, Indonesia.
Three new species of Dasyrhicnoessa Hendel, 1934 and one of Pseudorhicnoessa Malloch, 1914 from the Indo-Pacific area are described and the male terminalia illustrated. Among these new species, Dasyrhicnoessa paradoxa sp. nov. and Pseudorhicnoessa longicerca sp. nov. are especially noteworthy for the morphological peculiarities of the male terminalia.