The higher-level taxonomy of the stingrays (Dasyatidae) has never been comprehensively reviewed. Recent phylogenetic studies, supported by morphological data, have provided evidence that the group is monophyletic and consists of four major subgroups, the subfamilies Dasyatinae, Neotrygoninae, Urogymninae and Hypolophinae. A morphologically based review of 89 currently recognised species, undertaken for a guide to the world's rays, indicated that most of the currently recognised dasyatid genera are not monophyletic groups. These findings were supported by molecular analyses using the NADH2 gene for about 77 of these species, and this topology is supported by preliminary analyses base on whole mitochondrial genome comparisons. These molecular analyses, based on data generated from the Chondrichthyan Tree of Life project, are the most taxon-rich data available for this family. Material from all of the presently recognised genera (Dasyatis, Pteroplatytrygon and Taeniurops [Dasyatinae]; Neotrygon and Taeniura [Neotrygoninae]; Himantura and Urogymnus [Urogymninae]; and Makararaja and Pastinachus [Hypolophinae]), are included and their validity largely supported. Urogymnus and the two most species rich genera, Dasyatis and Himantura, are not considered to be monophyletic and were redefined based on external morphology. Seven new genus-level taxa are erected (Megatrygon and Telatrygon [Dasyatinae]; Brevitrygon, Fluvitrygon, Fontitrygon, Maculabatis and Pateobatis [Urogymninae], and an additional three (Bathytoshia, Hemitrygon and Hypanus [Dasyatinae]) are resurrected from the synonymy of Dasyatis. The monotypic genus Megatrygon clustered with 'amphi-American Himantura' outside the Dasyatidae, and instead as the sister group of the Potamotrygonidae and Urotrygonidae. Megatrygon is provisionally retained in the Dasyatinae pending further investigation of its internal anatomy. The morphologically divergent groups, Bathytoshia and Pteroplatytrygon, possibly form a single monophyletic group so further investigation is needed to confirm the validity of Pteroplatytrygon. A reclassification of the family Dasyatidae is provided and the above taxa are defined based on new morphological data.
Lomanius annae sp. nov. is described from southern Vietnam. The species is characterized by the greatly developed dorso-basal process on cheliceral hand of males and by the partial effacement of all mesotergal grooves. The genus Lomanius contains four generic synonyms and currently comprises eight valid species distributed in China, Java, peninsular Malaysia, the Philippines, and Taiwan. The new species displays a general morphology similar to the former genus Paralomanius, with a combination of sexually dimorphic interocular mound (which is very large and strongly leaned back in males) and pedipalpus (which is extremely elongate in males). This morphological suite of features is herein called facies reclinobunoides. The replacement name Metibalonius triceratops nom. nov. is proposed for Trispinibunus abnormis Roewer, 1915, which is a junior secondary homonym of Ibalonius abnormis Strand, 1911. Finally, numerous morphological structures found in Podoctidae are recognized and named: (1) the cheliceral comb, present on cheliceral fingers, (2) the chained tubercular ridges, present on dorsal scutum and (3) several others related to the ocular region. The distribution of these two structures among podoctid species is not fully known, but both are absent in the former Ibaloniinae. We suggest that both structures may be useful to define supra-generic groups in the clade composed of the former Podoctinae and Erecananinae.
The synchronous firefly genus Pteroptyx Olivier is reassessed from morphological, molecular, and habitat perspectives in Malaysia, and includes some reliably associated females described from morphological features and internal female reproductive anatomy. Phylogenetic analyses using combined morphological and molecular data (where available) for 158 taxa supported all the major features of the existing taxonomic categories within the Indopacific Luciolinae. They revealed a distinct Pteroptyx clade as a morphologically variable genus with Poluninius selangoriensis Ballantyne being newly synonymised with Luciola testacea Motschulsky, the type species, which is redescribed from the type series. Pteroptyx gelasina Ballantyne was shown to be distinct and three of the four morphological subdivisions within Pteroptyx malaccae (Gorham) considered useful. A new species Pt. balingiana Jusoh sp. nov. is described from Sarawak. A second specimen of Pt. gombakia Ballantyne is described and figured. Some females were reliably associated with identified males by molecular data, but investigation of their morphology showed consistent features that were for the most part not useful for species delineation, which still relies on association with the males and colour patterns. All females investigated had bursa plates.Habitat details for most Pteroptyx revealed an association with a riparian environment likely to support mangroves but not necessarily an obligatory association with mangroves or any particular species. Pteroptyx galbina Jusoh was found up to 30 km from the sea, and Pt. bearni Olivier displays in a variety of flowering plants alongside rivers, including mangroves.Keys to species and diagnoses of all species with coloured plates are given.
A new species of Argiope Audouin 1826, A. hoiseni new species is described from Perak and Selangor, Peninsular Malaysia based on morphology and DNA information of the mitochondrial (16S rRNA, COI and COII) and nuclear-encoded (H3A, 18S rRNA) molecular markers. Epigynal structure suggested Argiope hoiseni to be similar to A. jinghongensis Yin, Peng Wang 1994, A. luzona (Walckenaer 1841), A. pulchella Thorell 1881 and A. taprobanica Thorell 1887. Molecular sequence data including the new species inferred that it is monophyletic with an intraspecific variation of 0.87-3.59 % based on the 16S+COI+COII+H3A dataset. Phylogenetic analyses also revealed insights into the evolutionary lineages of Argiope species in Southeast Asia as well as corroborated recent taxonomic changes and species synonymies associated with Argiope. Two new distribution records were also reported for A. chloreis Thorell,1877 and A. doleschalli Thorell, 1873 in Peninsular Malaysia.
Up to three nominal species of the cyprinid fish genus Poropuntius (i.e. P. deauratus [Valenciennes in Cuvier Valenciennes 1842], P. normani [Smith 1931], and P. smedleyi [de Beaufort 1933]) have been reported to occur in Peninsular Malaysian freshwater ecosystems. However, low morphological differentiation among species of Poropuntius causes confusion and it is still unknown how many valid species of Poropuntius occur in this region. The goal of this study is to review the taxonomic status of Poropuntius in Peninsular Malaysia by using morphological and molecular characters. Principal Component Analysis (PCA) on a morphometric dataset including 281 specimens of Poropuntius from Peninsular Malaysia and P. normani from Thailand (type locality) failed to identify non-overlapping clusters within sampled specimens. A phylogenetic tree based on cytochrome oxidase subunit I (COI) showed intraspecific levels of genetic differentiation within Poropuntius of Peninsular Malaysia and the specimens of P. normani from Thailand form a monophyletic group. Our results strongly support the presence of only one species of Poropuntius in Peninsular Malaysia, P. normani. We demonstrate that P. smedleyi described from Johor, southern Peninsular Malaysia, is a junior synonym of P. normani. The previous reports of the presence of P. deauratus in Peninsular Malaysia are doubtful because this species was described from Vietnam where, in all evidence, it is endemic.
The Sundaic swamp clade of the genus Cyrtodactylus contains nine species that collectively range through Peninsular Malaysia and its associated land bridge islands, Singapore, Sumatra, Java, and Pulau Natuna Besar. Ancestral range reconstruction analyses using BioGeoBEARS based on an updated molecular phylogeny of the nine Sundaic swamp clade species of Cyrtodactylus demonstrated that this lineage evolved in Peninsular Malaysia, dispersed independently to Sumatra and Pulau Natuna Besar, Indonesia and most likely back into Peninsular Malaysia from Sumatra. This scenario is consistent with climate-driven, cyclical, ephemeral, geographic reconfigurations of Sundaic landmasses from at least the mid-Miocene to present.
The taxonomic position of the rare Selangor Mud Snake (Raclitia indica) Gray to other species of homalopsids has remained uncertain due to the scarcity of specimens in collections and the lack of genetic material for comparison. Here we report the first molecular phylogenetic examination of this species based on recently acquired material. The study recovered R. indica nested within the clade of advanced, fanged homalopsids and the sister species to Erpeton tentaculatus Lácèpede. We also present notes on variation observed in the new specimens as well as range extensions for the species.
This paper reports a new weevil genus Devernodes gen. n. established for five new species from Southeast Asia: D. alkippe sp. n. (China: Mt. Emei), D. asteria sp. n. (Vietnam: Tam Dao), D. chthonia sp. n. (Vietnam: Tam Dao; the type species), D. drimo sp. n. (Malaysia: Pasoh Forest Reserve) and D. methone sp. n. (Malaysia: Tanah Rata). All Devernodes are wingless and inhabit the forest leaf litter. Adult Devernodes share a combination of two head characters unique among weevils in Asia: antenna with apparently unsegmented club and 6-segmented antennal funicle, as well as strong constriction separating the eye-bearing rostrum from the head capsule. To test monophyly and investigate phylogenetic relationships of Devernodes, Maximum Likelihood phylogenetic analysis was undertaken using parts of mitochondrial (COI) and nuclear ribosomal (28S) genes, as well as the nuclear internal transcribed spacer (ITS2) from 14 Devernodes and 55 outgroup Curculionidae specimens. Results strongly corroborated monophyly of Devernodes and did not suggest its realistic sister-group. The new genus is assigned to the molytine tribe Lymantini (not represented in the DNA analysis) based on two potential synapomorphies: head markedly constricted behind eyes and presence of undivided female hemisternites IX (= "merged coxite and stylus"). Thus interpreted, Devernodes is the twelfth nominal genus of Lymantini and the first record of the tribe outside of the Americas. All original data (localities, DNA sequences, specimen images) are available online in public datasets dx.doi.org/10.5883/DS-DEVERNO1 and dx.doi.org/10.5883/DS-DEVERNO2.
An integrative taxonomic analysis of the Ptychozoon lionotum group across its range in Indochina and Sundaland recovers P. lionotum sensu lato Annandale, 1905 as paraphyletic with respect to P. popaense Grismer, Wood, Thura, Grismer, Brown, Stuart, 2018a and composed of four allopatric, genetically divergent, ND2 mitochondrial lineages. Multivariate and univariate analyses of continuous and discrete morphological and color pattern characters statistically and discretely diagnose each lineage from one another and together, with maximum likelihood and Bayesian inference analyses, provide the foundation for the recognition of each lineage as a new species-hypotheses corroborated with a Generalized Mixed Yule Coalescent species delimitation analysis. Ptychozoon cicakterbang sp. nov. ranges throughout Peninsular Malaysia to Pulau Natuna Besar, Indonesia; P. kabkaebin sp. nov. is endemic to northern and central Laos; and P. tokehos sp. nov. ranges from southern Thailand south of the Isthmus of Kra northward to Chiang Mai, fringing the Chao Phraya Basin and ranging southward through Cambodia to southern Vietnam. Ptychozoon lionotum sensu stricto ranges from northwestern Laos through southern Myanmar to eastern India. The phylogeographic structure within each species varies considerably with P. lionotum s.s. showing no genetic divergence across its 1,100 km range compared to P. cicakterbang sp. nov. showing upwards of 8.2% sequence divergence between syntopic individuals. Significant phylogeographic structure exists within P. tokehos sp. nov. and increased sampling throughout Thailand may require additional taxonomic changes within this species.
We collected a specimen of a scincid lizard of Larutia Böhme, 1981 from the edge of a primary forest on Gunung Penrissen, Kuching Division, Sarawak, Malaysian Borneo. The single specimen of the new species differs from all other known congeners by the molecular divergence in the mitochondrial ND1 gene and morphological characters including small adult body size (SVL 84 mm); 22 longitudinal scale rows around midbody; first pair of chinshields contacting second infralabial; second pair of chinshields separated from infralabials by an elongated scale; two subdigital lamellae on second toe; and body without yellow or pale bands or spots. It is the ninth species described in the genus and the second species of Larutia in Borneo.
The island of Borneo lies within one of the most biodiverse regions in the world. Despite this, its documented gekkonid diversity is not commensurate with other areas of Southeast Asia. The megadiverse genus Cyrtodactylus is especially underrepresented. Limestone-karst ecosystems, in particular, harbor many endemic Cyrtodactylus species, but only one karst-dwelling species is currently recognized from Borneo. This paper adds two additional karst-dwelling Cyrtodactylus species-C. muluensis sp. nov. and C. limajalur sp. nov.-from Sarawak, Malaysia. Cyrtodactylus muluensis sp. nov. is endemic to Gunung Mulu and is distinguished from its congeners by having a precloacal groove, 31-38 ventral scales, a maximum SVL of at least 88 mm, enlarged subcaudals, 19-20 subdigital lamellae, and a banded dorsal body pattern. Cyrtodactylus limajalur sp. nov. is endemic to the Serian region and is distinguished from its congeners by having 33-42 ventral scales, enlarged subcaudals, a precloacal pit, a maximum SVL of at least 94 mm, 5-6 enlarged femoral scales, 19-22 subdigital lamellae, and five distinct bands on the dorsum. Both species are phylogenetically distinct and deeply divergent from all other congeners. The description of two new karst-dwelling species highlights the need to conserve karst habitats and the endemic species they harbor.
The genus Rhitymna Simon, 1897 is revised by means of new material. Four new species are described: R. gerdmangel spec. nov. (Thailand, Malaysia; male, female), R. merianae spec. nov. (Indonesia: Bali; male), R. flores spec. nov. (Indonesia: Flores; male, female), R. senckenbergi spec. nov. (Philippines; male). The male of R. plana Jäger, 2003 and the female of R. tangi Quan Liu, 2012 are described for the first time. Rhitymna simoni Jäger, 2003 is recognised as junior synonym of R. cursor (Thorell, 1894) comb. nov., the latter transferred from the genus Olios Walckenaer, 1837. New records are given for further Rhitymna species, among them new country or island records for R. verruca (Wang, 1991) (Thailand), R. tangi Quan Liu, 2012 (Laos), R. plana Jäger, 2003 (Cambodia), R. pinangensis (Thorell, 1891) (Thailand), R. deelemanae Jäger, 2003 (Bali). The number of cheliceral bristles close to the fang base is recognised as size dependent, therefore without true phylogenetic signal. Two main types of copulatory organs within the genus are recognised and discussed. R. gerdmangel spec. nov. has a special biology as it lives exclusively in bamboo. Holes made by beetles or woodpeckers are used to enter the bamboo stem. Spiders hide during the day and lay their eggs in bamboo internodes.
A new species of limestone karst-adapted gecko of the Cyrtodactylus pulchellus complex, C. dayangbuntingensis sp. nov., is described from Dayang Bunting Island of the Langkawi Archipelago off the northwest coast of Peninsular Malaysia. It is the third species of the group to be described from the archipelago after C. langkawiensis and C. macrotuberculatus. The new species can be distinguished from all other species of Cyrtodactylus based on molecular evidence from the mitochondrial gene ND2 and its flanking tRNAs as well as having unique combinations of morphological and color pattern characteristics. This discovery underscores the need for continued surveys of the many islands in the archipelago to properly ascertain its true herpetological diversity.
Micryletta inornata (Boulenger 1890), the type species of the genus Micryletta, was originally described from the island of Sumatra in Indonesia. Subsequently, this species has been widely reported from Sundaland (Sumatra and Malay Peninsula), Indo-China, Northeast India and South Andaman, up to southern China and Taiwan. However, since the original description there has been no further report of this species from the type locality or the island. During a herpetofaunal survey in Sumatra, several specimens that are morphologically concordant with the original description and the syntypes of M. inornata were found, and thus the species was rediscovered after 125 years. Here, we provide a redescription of the species based on the freshly collected specimens, along with a detailed morphological and molecular comparison with known congeners. Further, using molecular data from the mitochondrial 16S rRNA gene, our study recovered the Sumatran M. inornata as a phylogenetically distinct lineage from all other populations previously referred to this species. This confirms that all known Micryletta 'inornata' populations from regions outside Sumatra constitute several other lineages representing either new species or previously available names currently considered as synonyms, consequently requiring taxonomic validation in the future.
To date, 26 species of Theloderma have been described and all are distributed throughout Southeast Asia from Assam in northeastern India to Myanmar, Indochina, the Malay Peninsula, and the islands of the Greater Sundas: Sumatra and Borneo (Frost 2019). The tadpoles of only 12 species have been described and published: T. asperum (Boulenger); T. auratum Poyarkov, Kropachev, Gogoleva Orlov; T. bicolor (Bourret); T. corticale (Boulenger); T. gordoni Taylor; T. horridum (Boulenger); T. leave (Smith); T. moloch (Annandale); T. nebulosum Rowley, Le, Hoang, Dau Cao; T. palliatum Rowley, Le, Hoang, Dau Cao; T. stellatum Taylor; T. vietnamense Poyarkov, Orlov, Moiseeva, Pawangkhanant, Ruangsuwan, Vassilieva, Galoyan, Nguyen Gogoleva (Boulenger 1903; Annandale 1912; Wassersug et al. 1981; Inger et al. 1999; Leong Lim 2003; Inthara et al. 2005; Rowley et al. 2011; Gawor et al. 2012; Orlov et al. 2012; Poyarkov et al. 2015; Kropachev et al. 2018).
Molecular phylogenetic analyses of the sister species Sphenomorphus stellatus and S. praesignis based on the mitochondrial genes 12S and 16S rRNA recover the former as paraphyletic with respect to the latter in that a specimen of S. stellatus from the type locality in Peninsular Malaysia is more closely related to S. praesignis than to Indochinese populations of S. stellatus. Furthermore, the phylogeny indicates that the Indochinese populations represent two species, thus resulting in four major lineages within this clade. These relationships are consistent with multivariate and univariate analyses of morphological and discrete color pattern data which statistically define and diagnose the four lineages and together with the molecular data, provide the foundation for robust, testable, species-level hypotheses. As such, S. stellatus is herein restricted to Peninsular Malaysia; S. annamiticus is resurrected for the circum-continental populations ranging through southeastern Thailand, southern Cambodia, and southern Vietnam; a new species-S. preylangensis sp. nov.-is described from an isolated mountain, Phnom Chi, from the Prey Lang Wildlife Sanctuary in central Cambodia; and the taxonomy of S. praesignis remains unchanged. The description of S. preylangensis sp. nov. underscores the necessity to conserve this remnant of lowland evergreen rainforest in the Prey Lang Wildlife Sanctuary.
A new species of Ansonia is described from the Shan Plateau of Myanmar based on an integrative taxonomic analysis that differentiates it from all other congeners. Molecular phylogenetic analyses based on the mitochondrial genes 12S and 16S rRNA and tRNA-val recover A. kyaiktiyoensis sp. nov. as the sister species to A. inthanon from Thailand but differs from it and other congeners by at least a 5.0% sequence divergence. It is further differentiated by the following combination of morphological characters: (1) maximum SVL 24 mm in males and females; (2) first finger shorter than second; (3) absence of interorbital and tarsal ridges; (4) presence of light-coloured interscapular spot; (5) presence of yellow rictal tubercle; (6) absence of wide, light-coloured patch below eye; (7) presence of large, discrete, bright-yellow submandibular spots along the underside of lower jaw; (8) iris yellow-gold; (9) presence of markings on the snout consisting of streaks below the eye to the lip, and on the canthus rostralis to the nostril; (10) dorsum grey-brown with orange-beige spots, a dark-brown X-shaped marking on the back surrounding the interscapular spot, and dark-coloured markings on rump; (11) fore- and hind limbs with orange-beige cross-bars; and (12) venter light-gray with yellow spotting, especially near flanks and underside of hind limbs. Ansonia kyaiktiyoensis sp. nov. is the westernmost known record for the genus and the only species west of the Salween Basin. Its discovery echoes the increasing number of herpetological discoveries being made in upland regions fringing the Ayeyarwady and Salween Basins.
An integrative taxonomic analysis of Subdoluseps herberti from southern Thailand and Peninsular Malaysia and S. samajaya from Sarawak, East Malaysia (Borneo) recovers the former as paraphyletic with respect to the latter. The analyses recover the three southernmost populations of S. herberti in Peninsular Malaysia as conspecific and the sister lineage of S. samajaya, whereas S. herberti from Thailand and northern Peninsular Malaysia constitute the sister species to S. samajaya plus the southern three Peninsular Malaysian populations. As such, the southern populations are described herein as S. malayana sp. nov. and all three species are referred to as the S. herberti group. Clade boundaries and breaks within this group on the Thai-Malay Peninsula occurring at the Isthmus of Kra, across the Kangar-Pattani line, and between the Thai-Malay Peninsula and Borneo are consistent with phylogeographic patterns of other Sundaic taxa. The discovery of S. malayana sp. nov. continues to underscore the fact that, despite the well-studied nature of the lizard fauna of Peninsular Malaysia, much of it still remains unrealized and for conservation efforts to move forward, field research followed by expeditiously revised taxonomies must continue.
This overview of the Luciolinae addresses the fauna of S. E. Asia including India, Sri Lanka, China, Japan, Malaysia, Thailand, Laos, Cambodia, Vietnam, Indonesia, the Philippines, the Republic of Palau, Federated States of Micronesia, and the Australopacific area of Australia, Papua New Guinea, Solomon Islands, New Caledonia, Vanuatu and Fiji.Of the 28 genera now recognised in the Luciolinae we address 27 genera from the study area as defined above, including three new genera which are described herein, and 222 species including 13 species newly described herein. Photuroluciola Pic from Madagascar is the only Luciolinae genus not addressed here. A key to genera is presented. Keys to species are either included here or referenced in existing literature. Twelve genera have had no new taxonomic decisions made nor are any new species records listed, and are addressed in an abbreviated fashion, with short diagnoses and plates of features of life stages: Aquatica Fu et al. 2010, Australoluciola Ballantyne 2013, Convexa Ballantyne 2009, Emeia Fu et al. 2012a, Inflata Boontop 2015, Lloydiella Ballantyne 2009, Missimia Ballantyne 2009, Pteroptyx Olivier 1902, Pyrophanes Olivier 1885, Sclerotia Ballantyne 2016, Triangulara Pimpasalee 2016, and Trisinuata Ballantyne 2013. Abscondita Ballantyne 2013 contains 8 species, and includes new records for Abs. anceyi (Olivier 1883), Abs. chinensis (L.) (which is newly synonymised with Luciola succincta Bourgeois), Abs. terminalis (Olivier 1883) including a first record from both Laos and Thailand, and Abs. perplexa (Walker 1858). Luciola pallescens Gorham 1880 is transferred to Abscondita and the pronotal colour range is addressed from a wide range of localities. Abs. berembun Nada sp. nov. and Abs. jerangau Nada sp. nov. are described from Malaysia. Hooked bursa plates are described for pallescens and berembun. Aquilonia Ballantyne 2009 is expanded to include 3 species. Gilvainsula Ballantyne 2009, represented by two species from the south eastern coast of New Guinea is synonymised under Aquilonia Ballantyne 2009, which is briefly redescribed and keyed from: Aquil. costata (Lea) from northern Australia, including many new records, Aquil. messoria (Ballantyne) comb. nov. and Aquil. similismessoria (Ballantyne) comb. nov. Asymmetricata Ballantyne 2009 now includes 4 species. As. bicoloripes (Pic 1927) comb. nov. and As. humeralis (Walker 1858) comb. nov. are transferred from Luciola, with L. doriae Olivier 1885, L. impressa Olivier 1910b and L. notatipennis Olivier 1909a newly synonymised with As. humeralis. Luciola aemula Olivier 1891 is synonymised with As. ovalis (Hope 1831). The variation in the extent of the anterior median emargination of the light organ in ventrite 7, and the possibility of a bipartite light organ in males of As. circumdata (Motsch. 1854) is explored. Females of both As. circumdata and As. ovalis (Hope 1831) are without bursa plates and the distinctively shaped median oviduct plate in each is described. Records from Thailand are recorded for both As. circumdata and As. ovalis. Atyphella Olliff 1890 now contains 28 species with 4 transferred from other genera, and one new species: Aty. abdominalis (Olivier 1886) comb. nov. and Aty. striata (Fabricius 1801) comb. nov. are transferred from Luciola, with Aty. carolinae Olivier 1911b and Aty. rennellia (Ballantyne 2009) comb. nov. transferred from Magnalata Ballantyne 2009. Atyphella telokdalam Ballantyne sp. nov. from Indonesia is described herein. Atyphella is now known from records in the Philippines and Indonesia as well as Australia and New Guinea. Colophotia Motschulsky 1853 is considered here from seven species for which intact types can be located for three. An abbreviated revision based on the United States National Museum collection only is presented, with specimens of C. bakeri Pic 1924, C. brevis Olivier 1903a, C. plagiata (Erichson 1834) and C. praeusta (Eschscholtz 1822) redescribed, using where possible features of males, females and larvae. Colophotia particulariventris Pic 1938 is newly synonymised with C. praeusta. Colophotia miranda Olivier 1886 and L. truncata Olivier 1886 are treated as species incertae sedis. Curtos Motschulsky 1845 includes 19 species with suggestions made, but not yet formalised, for the possible transfer of the following seven species from Luciola: Luciola complanata Gorham 1895, L. costata Pic 1929, L. delauneyi Bourgeois 1890, L. deplanata Pic 1929, L. extricans Walker 1858, L. multicostulata Pic 1927 and L. nigripes Gorham 1903. Curtos is not revised here. Emarginata Ballantyne gen nov. is described for E. trilucida (Jeng et al. 2003b) comb. nov., transferred from Luciola and characterised by the emarginated elytral apex. An extended range of specimens from Thailand is listed. Kuantana Ballantyne gen. nov. from Selangor, Malaysia is described from K. menayah gen. et sp. nov. having bipartite light organs in ventrite 7 and an asymmetrical tergite 8 which is not emarginated on its left side. Female has no bursa plates. Luciola Laporte 1833 s. stricto as defined by a population of the type species Luciola italica (L. 1767) from Pisa, Italy, is further expanded and considered to comprise the following19 species: L. antipodum (Bourgeois 1884), L. aquilaclara Ballantyne 2013, L. chapaensis Pic 1923 which is synonymised with L. atripes Pic 1929, L. curtithorax Pic 1928, L. filiformis Olivier 1913c, L. horni Bourgeois 1905, L. hypocrita Olivier 1888, L. italica (L. 1767), L. kagiana Matsumura 1928, L. oculofissa Ballantyne 2013, L. pallidipes Pic 1928 which is synonymised with L. fletcheri Pic 1935, L. parvula Kiesenwetter 1874, L. satoi Jeng Yang 2003, L. tuberculata Yiu 2017, and two species treated as near L. laticollis Gorham 1883, and near L. nicollieri Bugnion 1922. The following are described as new: L. niah Jusoh sp. nov., L. jengai Nada sp. nov. and L. tiomana Ballantyne sp. nov. Luciola niah sp. nov. female has two wide bursa plates on each side of the bursa. Luciola s. lato (as defined here) consists of 36 species. Twenty-seven species formerly standing under Luciola have been assigned to other genera or synonymised. Seven species are recommended for transfer to Curtos, and 32 species now stand under species incertae sedis. Magnalata Ballantyne is reduced to the type species M. limbata and redescribed. Medeopteryx Ballantyne 2013 is expanded to 20 species with the addition of two new combinations, Med. semimarginata (Olivier 1883) comb. nov. and Med. timida (Olivier 1883) comb. nov., both transferred from Luciola, and one new species, Med. fraseri Nada sp. nov. from Malaysia. The range of this genus now extends from Australia and the island of New Guinea to SE Asia. Medeopteryx semimarginata females have wide paired bursa plates. Pygoluciola Wittmer 1939 now includes 19 species with 5 new species: P. bangladeshi Ballantyne sp. nov., P. dunguna Nada 2018, P. matalangao Ballantyne sp. nov. (scored by the code name 'Jeng Matalanga' in Ballantyne Lambkin 2013), P. phupan Ballantyne sp. nov. and P. tamarat Jusoh sp. nov. Six species are transferred from Luciola: P. abscondita (Olivier 1891) comb. nov., P. ambita (Olivier 1896) comb. nov., P. calceata (Olivier 1905) comb. nov., P. insularis (Olivier 1883) comb. nov., P. nitescens (Olivier 1903b) comb. nov. and P. vitalisi (Pic 1934) comb. nov., and redescribed from males, and includes female reproductive anatomy for P. nitescens comb. nov. and P. dunguna, both of which have hooked bursa plates. Serratia Ballantyne gen. nov. is erected for S. subuyania gen. et sp. nov. and characterised by the serrate nature of certain antennal flagellar segments in the male. The following 37 species listed under species incertae sedis are further explored: Colophotia miranda Olivier 1886, Lampyris serraticornis Boisduval 1835, Luciola angusticollis Olivier 1886, L. antennalis Bourgeois 1905, L. antica (Boisduval 1835), L. apicalis (Eschscholtz 1822), L. aurantiaca Pic 1927, L. bicoloriceps Pic 1924, L. binhana Pic 1927, L. bourgeoisi Olivier 1895, L. dilatata Pic 1929, L. exigua (Gyllenhall 1817), L. exstincta Olivier 1886, L. fissicollis Fairmaire 1891, L. flava Pic 1929, L. flavescens (Boisduval 1835), L. fukiensis Pic 1955, L. immarginata Bourgeois 1890, L. incerta (Boisduval 1835), L. infuscata (Erichson 1834), L. intricata (Walker 1858), L. japonica (Thunberg 1784), L. klapperichi Pic 1955, L. lata Olivier 1883, L. limbalis Fairmaire 1889, L. marginipennis (Boisduval 1835), L. melancholica Olivier 1913a, L. robusticeps Pic 1928, L. ruficollis (Boisduval 1835), L. spectralis Gorham 1880, L. stigmaticollis Fairmaire 1887, L. tincticollis Gorham 1895, L. trivandrensis Raj 1947, L. truncata Olivier 1886, L. vittata (Laporte 1833) Pteroptyx atripennis Pic 1923 and P. curticollis Pic 1923. While phylogenetic analyses indicate their distinctiveness, no further taxonomic action is taken with Luciola cruciata Motschulsky 1854 and L. owadai Sâtô et Kimura 1994 from Japan given the importance of the former as a national icon. Analyses also indicate that Lampyroidea syriaca Costa 1875 belongs in Luciola s. str. A much wider taxonomic analysis of this genus including all the species is necessary before any further action can be taken.
The intertidal serpulid polychaete Spirobranchus kraussii was originally described from South Africa and has since been reported in numerous sub (tropical) localities around the world. Recently, however, S. kraussii was uncovered as a complex of morphologically similar and geographically restricted species, raising the need to revise S. cf. kraussii populations. We formally describe S. cf. kraussii from Singapore mangroves as Spirobranchus bakau sp. nov. based on morphological and molecular data. Despite their morphological similarities, Maximum Likelihood and Bayesian Inference analyses of 18S and Cyt b DNA sequence data confirm that S. bakau sp. nov. is genetically distinct from S. kraussii and other known species in the complex. Both analyses recovered S. bakau sp. nov. as part of a strongly supported clade (96% bootstrap, 1 posterior probability), comprising S. sinuspersicus, S. kraussii and S. cf. kraussii from Australia and Hawaii. Additionally, paratypes of S. kraussii var. manilensis, described from Manila Bay in the Philippines, were examined and elevated to the full species S. manilensis. Finally, we tested the hypothesis that fertilisation and embryonic development of S. bakau sp. nov. can occur under the wide range of salinities (19.630.9 psu) and temperatures (2531C) reported in the Johor Strait. Fertilisation success of ≥70% was achieved across a temperature range of 2532C and a salinity range of 2032 psu. Embryonic development, however, had a narrower salinity tolerance range of 2732 psu. Clarifying the taxonomic status of S. cf. kraussii populations reported from localities elsewhere in Singapore and Southeast Asia will be useful in establishing the geographical distribution of S. bakau sp. nov. and other members of the S. kraussii-complex.