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 synonymised with P. bearni. Luciola semilimbata Olivier is transferred to Pyrophanes, and Luciola cowleyi Blackburn to Pygoluciola. The following species are treated as species incertae sedis: L. melancholica Olivier, L. ruficollis Guérin-Ménéville. The New Guinean records of Luciola tenuicornis Olivier, L. timida Olivier and Photinus cinctellus Motschulsky are suspect. Fifteen of the species treated here are recognised by flashing patterns. The functions of the terminal abdominal modifications, origins of the Australopacific firefly fauna, and use of female and larval characters in interpretations of relationships are considered.
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.
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.