The firefly genus Luciola sensu McDermott contains 282 species that are distributed across major parts of Asia, Europe, Africa, Australia, and the Pacific islands. Due to phenotypic similarities, species identification using external morphological characters can be unreliable for this group. Consequently, decades of piecemeal taxonomic treatments have resulted in numerous erroneous and contentious classifications. Furthermore, our understanding of the group’s evolutionary history is limited due to the lack of a robust phylogenetic framework that has also impeded efforts to stabilize its taxonomy. Here, we constructed molecular phylogenies of Luciola and its allies based on combined mitogenomes and Cytochrome c oxidase subunit 1 (COX1) sequences including a newly sequenced mitogenome of an unidentified taxon from Singapore. Our results showed that this taxon represents a distinct and hitherto undescribed evolutionary lineage that forms a clade with L. filiformis from Japan and L. curtithorax from China. Additionally, the Singaporean lineage can be differentiated from other congeners through several external and internal diagnostic morphological characters, and is thus described herein as a new species. Our phylogeny also strongly supported the paraphyly of Luciola with regard to L. cruciata and L. owadai, which were inferred to be more closely related to the genus Aquatica as opposed to other members of Luciola sensu stricto. The genus Hotaria was inferred as a derived clade within Luciola (sister to L. italica), supporting its status as a subgenus of Luciola instead of a distinct genus. This is the first time since 1909 that a new species of luminous firefly has been discovered in Singapore, highlighting the need for continued biodiversity research, even in small, well-studied and highly developed countries, such as Singapore.

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...

This revision completes a taxonomic survey of fireflies (Coleoptera: Lampyridae) in the area encompassed by Australia, the Republic of Palau, Federated States of Micronesia, Papua New Guinea, Indonesia (West Irian/Papua), Solomon Islands, New Caledonia, Vanuatu and Fiji. It finalises the taxonomic issues arising from the 1969–70 voyage of the scientific vessel Alpha Helix to New Guinea. The firefly fauna of this area is exclusively Luciolinae. The scope of the revision was extended to include all known Luciolinae genera and certain species from SE Asia, and a phylogenetic analysis of 436 morphological characters of males, females, and associated larvae includes 142 Luciolinae species (Ballantyne & Lambkin 2009, and Fu et al. 2012a). The phylogenetic analyses infer four major groups within the Luciolinae. The monotypic Missimia Ballantyne is sister to all remaining Luciolinae and forms a grade to Aquatica Fu etBallantyne. The large clade of Curtos Motschulsky, Photuroluciola Pic, Colophotia Motschulsky, Poluninius gen. nov., Pyrophanes Olivier, Pteroptyx s. str. Olivier, Medeopteryx gen. nov., Trisinuata gen. nov., and Australoluciola gen. nov.forms a grade to the clade of Luciola s. str. Laporte (including Bourgeoisia Olivier). The monotypic Emeia Fu et al.forms a grade with a clade of Luciola and Pygoluciola Wittmer, sister to a large clade of Convexa Ballantyne, Pacifica gen. nov., Magnalata Ballantyne, Lloydiella Ballantyne, Asymmetricata Ballantyne, Pygatyphella s. str. Ballantyne, Atyphella Olliff, Aquilonia Ballantyne, and Gilvainsula Ballantyne. Luciola is paraphyletic, found in up to six clades across the tree. Together with Luciola, Magnalata, Aquilonia, and Gilvainsula render Atyphella paraphyletic. The new genera described here are all monophyletic and supported in the phylogenetic analyses that also provide evidence for the inclusion of taxa within them. Twenty-three genera including five new ones, and ten new species, are recognised and keys are presented for the males and females. Certain females are characterised by the nature of their bursa plates.Australoluciola gen. nov. is proposed for ten species from Australia and New Guinea, seven transferred from Luciola and three new, with species keyed from males, all of which have an entire light organ in ventrite 7. Aus. anthracina (Olivier), Aus. aspera (Olivier), Aus. australis (F.), Aus. flavicollis (MacLeay), Aus. foveicollis (Olivier), Aus. nigra (Olivier) and Aus. orapallida (Ballantyne) are transferred from Luciola with males assigned to Aus. aspera(Olivier), and a lectotype designated for Luciola foveicollis Olivier; Aus. baduria sp. nov., Aus. fuscamagna sp. nov.,Aus. fuscaparva sp. nov., Aus. japenensis sp. nov. and Aus. pharusaurea sp. nov. are described. Females of Aus. australis and Aus. flavicollis have two pairs of wide bursa plates.The bent-winged fireflies of New Guinea and Australia are removed from Pteroptyx Olivier and assigned to Medeopteryx gen. nov. and Trisinuata gen. nov. Medeopteryx gen. nov. is erected for 17 species including two new; all have ventrite 7 with an entire light organ, trisinuate posterior margin and short posterolateral projections; the following 14 species in which males have deflexed elytral apices are transferred from Pteroptyx Olivier: M. amilae (Satô), M. antennata (Olivier), M. corusca (Ballantyne), M. cribellata (Olivier), M. effulgens (Ballantyne), M. elucens (Ballantyne), M. flagrans (Ballantyne), M. fulminea (Ballantyne), M. hanedai (Ballantyne), M. platygaster (Lea), M. similisantennata(Ballantyne), M. sublustris (Ballantyne), M. tarsalis (Olivier), and M. torricelliensis (Ballantyne). M. clipeata sp. nov. is described. Two species without deflexed elytral apices include M. pupilla (Olivier) which is transferred from Luciola, and M. similispupillae sp. nov. A Lectotype is designated for Luciola pupilla (Olivier). Females of M. corusca(Ballantyne), M. cribellata (Olivier), M. effulgens (Ballantyne), and M. similispupillae sp. nov. have two pairs of wide bursa plates. The second genus including species in which the males have deflexed elytral apices is Trisinuata gen. nov., where all males have light organ in ventrite 7 bipartite and posterolateral projections expanded; it is proposed for eight New Guinean species: T. microthorax (Olivier), T. minor (Ballantyne), T. papuae (McDermott) and T. similispapuae(Ballantyne) are transferred from Pteroptyx Olivier, T. papuana (Olivier) previously known only from a female, has males associated and is transferred from Luciola, and T. caudabifurca sp. nov., T. dimidiata sp. nov. and T. apicula sp. nov. are described. Females of T. similispapuae (Ballantyne) have two pairs of wide bursa plates.Luciola s. str. is defined by scoring the type species L. italica (L), Bourgeoisia Olivier and Lampyroidea (based on its type species syriaca Costa) both of which are submerged into Luciola; Luciola s. str is addressed here from four Pacific Island species: L. hypocrita Olivier, L. antipodum Bourgeois both transferred from Bourgeoisia; L. aquilaclarasp. nov. and L. oculofissa sp. nov. are described. L. oculofissa sp. nov. is the only Luciolinae male known to lack light organs. Females of L. italica and L. hypocrita lack bursa plates.Pacifica gen. nov. is proposed for five species from the Solomon Islands transferred from Pygatyphella(Ballantyne), and which the phylogenetic analysis shows to be distinctive viz. P. limbatifusca (Ballantyne), P. limbatipennis (Pic), P. plagiata (Blanchard), P. russellia (Ballantyne), and P. salomonis (Olivier).A monotypic genus Poluninius gen. nov. is proposed for Pol. selangoriensis sp. nov. from Selangor, Malaysia. The genera Colophotia, Pteroptyx, Pyrophanes, and Pygoluciola are treated in an abbreviated fashion with generic diagnoses, lists of, and keys to, species. Pteroptyx bearni Olivier and P. tener Olivier are characterised from type specimens and female bursae and P. similis Ballantyne is...

Studies on South-east Asian fireflies: Abscondita , a new genus with details of life history, flashing patterns and behaviour of Abs. chinensis (L.) and Abs. terminalis (Olivier) (Coleoptera: Lampyridae: Luciolinae)

Revisional studies on the Luciolinae fireflies of Asia (Coleoptera: Lampyridae): 1. The genus Pyrophanes Olivier with two new species. 2. Four new species of Pteroptyx Olivier and 3. A new genus Inflata Boontop, with redescription of Luciola indica (Motsch.) as Inflata indica comb. nov.

Sclerotia Ballantyne gen. nov. is established for seven species. Sclerotia aquatilis (Thancharoen, 2007), Scl. brahmina (Bourgeois, 1890), Scl. carinata (Gorham, 1880), Scl. flavida (Hope, 1845), Scl. seriata (Olivier, 1891), and Scl. substriata (Gorham, 1880) are transferred from Luciola Laporte. Sclerotia fui sp. nov. from China is described as new. Luciola cingulata Olivier, 1885 is synonymised with Luciola substriata Gorham. Luciola formosana Pic, 1916 is removed from synonymy with Luciola substriata and newly synonymised with Luciola flavida (Hope). Males are characterised by linear elytral punctation, a set of three sclerites surrounding the aedeagal sheath and a light organ in ventrite 7 with emarginated anterior margin. Associated females lack bursa plates. Larvae associated for three species are back swimmers, having heavily sclerotised exoskeleton and a metapneustic respiratory system. Males and associated females are keyed. Analysis of molecular evidence for Chinese populations of three species indicates monophyly of Sclerotia Ballantyne gen. nov. and distinct separation from other genera.Triangulara Pimpasalee gen. nov. is described from Tri. frontoflava Pimpasalee gen. et sp. nov. from Thailand. It is superficially similar to Sclerotia gen. nov. but larger, and specimens in collections appear to have been confused with the similarly coloured Scl. substriata (Gorham) comb. nov. Males differ in having an elongate triangular light organ in ventrite 7 and are without sclerites around the aedeagal sheath. Larvae are not reliably associated but may be aquatic.