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RESUMO Desde seu lançamento, Lucíola (1862) tem sido objeto de análises que abordam especialmente o paralelo com a A dama das camélias (Alexandre Dumas Filho) e Paulo e Virgínia (Bernardin de St-Pierre). Porém, raramente as questões ali presentes foram tratadas em seu conjunto, de modo a constituírem uma narrativa única e coerente. Pois, a despeito da linguagem erótica dos capítulos iniciais, as imagens idílicas ou “analogia da inocência” (Frye) acabam por sobrepor-se gradualmente na construção de cenas, diálogos e situações, assim como nas remissões à obra de St. Pierre, a Atala (Chateaubriand) e aos idílios de Teócrito. Assim, Lucíola sofre uma modificação crucial em seu mythos, que passa a orbitar em torno do conceito de idílio, não apenas enquanto locus amoenus (Curtius) mas sobretudo no sentido moderno atribuído a ele por Schiller (1800), segundo o qual trata-se de um vir-a-ser que só se realiza plenamente na chave da transcendência. Tal reorientação, da imanência à transcendência, aproxima-o de uma das chamadas “formas simples” (Jolles), que é a legenda cristã. Ao objetificar bem e mal, a legenda culmina na personagem qualitativamente superior do santo, cujos mythos se dão através do “trajeto que conduz à santidade” (Jolles). Discutindo tais premissas teóricas, o artigo propõe que a crítica a Lucíola revelou profunda incompreensão quanto ao papel discursivo de G.M., a digna senhora responsável por publicar as cartas a ela entregues por Paulo, o amante da heroína. G.M. compreende a construção do romance enquanto vita e exemplum - algo que o narrador não chega a entender. ABSTRACT Since its release, Lucíola (1862) has been the subject of analyses that especially address the parallel with La dame aux camélias (Alexandre Dumas, fils) and Paul et Virginie (Bernardin de St-Pierre). However, rarely have the issues therein been treated as a whole, so as to constitute a single coherent narrative. For, despite the erotic language of the initial chapters, the idyllic images or “analogy of innocence” (Frye) end up gradually overlapping in the construction of scenes, dialogues and situations, as well as in the references to the work of St. Pierre, to Atala (Chateaubriand) and to the idylls of Theocritus. Thus, Lucíola undergoes a crucial modification in its mythos, which comes to orbit around the concept of the idyll, not only as locus amoenus (Curtius) but above all in the modern sense attributed to it by Schiller (1800), according to which it is a coming-to-be which is only fully realised in the key of transcendence. Such a reorientation from immanence to transcendence brings it closer to one of the “simple forms” (Jolles), which is the Christian legend. By objectifying good and evil, the legend culminates in the superior character of the saint, whose mythos takes place through the “path that leads to sanctity” (Jolles). Discussing such premises, the article proposes that the criticism of Lucíola revealed a profound misunderstanding of the discursive role of G.M., the worthy lady responsible for publishing the letters delivered to her by Paulo, the heroine’s lover. G.M. understands the construction of the novel as vita and exemplum - something the narrator fails to grasp.

Systematics of Luciola Laporte, 1833 – quo vadis ? Luciola s. str. Ballantyne and colleagues identified very early on that the grouping of 276 species under Luciola in McDermott (1966) was at best arbitrary, and morphological phylogenetic analyses (see Ballantyne et al. 2019 for a review) identified within McDermott’s Luciola many new genera, many including species which were transferred from what McDermott had placed under Luciola. It became clear that ‘ Luciola ’ was a heterogeneous assemblage of species and that it was necessary to be able to define what constituted Luciola s. str. This was achieved by comparison of morphological features of the type species Luciola italica with the other species. Fu et al. (2010) first identified two species, Luciola italica and the Japanese species Luciola parvula Kiesenwetter, 1874 as Luciola s. str., and Fu et al. (2012a) an additional species but without further definition. Ballantyne et al. (2013) first formally addressed and redescribed Luciola s. str. Subsequently Ballantyne et al. (2015: fig. 1; 2016: figs 1–2; 2019: fig. s1), and Jusoh et al. (2021: 1) further refined the concept and composition of Luciola s. str. Our exhaustive literature reviews and taxonomic revisions reveal two possible options to resolve Luciola taxonomy insofar as it relates to a definition of Luciola s. str. Such a definition is essential to our argument (and the status of Hotaria) as well as the resolution of a position for Luciola cruciata and L. owadai. Here we follow Jusoh et al. (2021: 8, fig. 3). The options are: 1. Subsume all three current members of Luciola subgenus Hotaria Yuasa, 1937 (unmunsana, papariensis and tsushimana) into one genus, i.e., Luciola s. str. (Jusoh et al. 2021: fig. 3 Clade G), and retain the remaining five species in Luciola s. str. Luciola s. str. thus comprises the 7 species from Jusoh et al. (2021: fig. 3) including the type species L. italica, and is clearly distinct from the clade E which includes L. cruciata / owadai and species of Aquatica. 2. Subsume all current members of Luciola subgenus Hotaria into one genus, i.e., Luciola s. str. (Jusoh et al. 2021: Clade G) which subsequently restricts Luciola s. str. to four species within the same clade as Luciola italica (Clade G Part II), and erect a new genus to accommodate Luciola pallidipes + niah Jusoh, 2019 (Clade G Part I) Option 2 while extreme, may indicate a future path, but is untenable at present without evidence to support the split between niah/ pallidipes and Luciola s. str., the extensive and presently uninvestigated ‘ Luciola ’ of Africa (about 100 species are assigned to Luciola in McDermott 1966), and species of ‘ Luciola ’ in Europe such as L. lusitanica Charpentier, 1825, L. italica, and L. mingrelica Motschulsky, 1854. There is no present agreement on just which populations comprise the aforementioned three species. Option 1 permits a wider definition of Luciola s. str., where all the species in Jusoh’s Clade G are assigned, and is supported by morphological considerations (including males females and larvae). This definition of Luciola s. str. will accommodate possible future directions as well as existing problems, while still defining a narrow group of species forming a clade with the type species. Neither of these options address specifically what course of action should be taken with three of the four species of ‘ Hotaria ’ which the analysis indicates may well be the same species, distinct from L. parvula. However, as each option recommends that Hotaria be subsumed under Luciola s. str. (as already undertaken by Kawashima et al. 2003) we advise this recommendation (Supp. file 1: 3). All comments above relating to suggested placements of species still listed as Luciola sp. relate specifically to the wide, and defined study area of Ballantyne et al. (2019: 5) in SE Asia and the Australopacific area. The remaining Luciola The situation with regard to the remaining species still standing under Luciola by McDermott (1966) can also be addressed. Ballantyne et al. (2019) addressed Luciola s. str. from 17 species, including the type species L. italica. However, they did not regard their treatment of all the species listed by McDermott (1966) under Luciola as finished. Clearly they did not attempt to address the Luciola of Africa (ca 100 species), nor the Luciola of Europe. Their specified area of coverage extended from India in the west through Asia (excluding Russia), the Philippines to the Australopacific region (see listing in Ballantyne et al. 2019: 5), where in the Australopacific region the firefly fauna is exclusively Luciolinae. In attempting to address the range of species, Ballantyne et al. (2019) were confronted with many of the problems we have outlined here. Two solutions were proposed specifically for the remaining species still standing under Luciola, occurring in the study area as defined above: 1. Species incertae sedis. Thirty-five species of ‘ Luciola ’ were assigned to species incertae sedis (Ballantyne et al. 2019: 151, table 28). While each species was individually addressed, each had certain attributes which the authors felt would preclude any further attempt to conclusively identify them. These included absence of types, poor condition of types (if represented at all what body parts that remained were not useful in diagnosis), and types that were females. This sex is presently of little use in what is still a male based Luciolinae taxonomy. 2. Luciola s. lat. The second solution addressed the remaining species from the defined study area still assigned to Luciola in McDermott (1966). Although a ‘heterogeneous assemblage of species’ (Ballantyne et al. 2019: 102), they were considered sufficiently distinctive (usually in colour, and in many cases because of the existence of a type specimen in some sort of condition that might permit re-examination), that the very real possibility existed for future assignment of males and elucidation of generic affinities. All are in MNHN. Of the thirty-three species listed in Ballantyne et al. (2019: 103–104, table 18) seven have female types but distinctive colouration, 18 have male types in various condition, and types were not found for 11. It was anticipated that a visit to the MNHN in Paris and work on the collection would eventuate, as this museum will no longer loan types. However Covid intruded, as did the very real financial situation the retired senior author finds herself in, that further precludes any possibilities of additional investigation in the immediate future., Published as part of Ballantyne, Lesley, Kawashima, Itsuro, Jusoh, Wan F. A. & Suzuki, Hirobumi, 2022, A new genus for two species of Japanese fireflies having aquatic larvae (Coleoptera, Lampyridae) and a definition of Luciola s. str., pp. 1-54 in European Journal of Taxonomy 855 on pages 42-44, DOI: 10.5852/ejt.2022.855.2023, http://zenodo.org/record/7501294, {"references":["McDermott F. A. 1966. Lampyridae. In: Steel W. O. (ed.) Coleopterorum Catalogus Supplementa, Pars 9: 1 - 149. Uitgeverij Dr. W. Junk, The Hague.","Ballantyne L. A., Lambkin C. L., Ho J-Z., Jusoh W. F. A., Nada B., Nak-eiam S., Thancharoen A., Wattanachaiyingcharoen W. & Yiu V. 2019. Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species. Zootaxa 4687 (1): 1 - 174. https: // doi. org / 10.11646 / zootaxa. 4687.1.1","Fu X. H., Ballantyne L. A. & Lambkin C. L. 2010. Aquatica gen. nov. from mainland China with a description of Aquatica wuhana sp. nov. (Coleoptera: Lampyridae: Luciolinae). Zootaxa 2530 (1): 1 - 18. https: // doi. org / 10.11646 / zootaxa. 2530.1.1","Kiesenwetter E. A. H. v. 1874. Die Malacodermen Japans nach dem Ergebnisse der Sammlungen des Herrn G. Lewis wahrend der Jahre 1869 - 1871. Berliner entomologische Zeitschrift 18: 241 - 288.","Fu X. H., Ballantyne L. A. & Lambkin C. L. 2012 a. Emeia gen. nov., a new genus of Luciolinae fireflies from China (Coleoptera: Lampyridae) with an unusual trilobite-like larva, and a redescription of the genus Curtos Motschulsky. Zootaxa 3403 (1): 1 - 53. https: // doi. org / 10.11646 / zootaxa. 3403.1.1","Ballantyne L. A., Fu X-H., Lambkin C., Jeng M-L., Faust L., Wijekoon W. M. C. D., Li D-Q. & Zhu T. F. 2013. 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). Zootaxa 3721 (1): 1 - 48. https: // doi. org / 10.11646 / zootaxa. 3721.1.1","Ballantyne L. A., Lambkin C. L., Boontop Y. & Jusoh W. F. A. 2015. Revisional studies on the Luciolinae fireflies of Asia (Coleoptera: Lampyridae): 1. The genus Pyrophane s 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. Zootaxa 3959 (1): 1 - 84. https: // doi. org / 10.11646 / zootaxa. 3959.1.1","Jusoh W. F. A., Ballantyne L., Chan S. H., Wong T. W., Yeo D. & Nada B. 2021. Molecular systematics of the firefly genus Luciola (Coleoptera: Lampyridae: Luciolinae) with the description of a new species from Singapore. Animals 2021 11: 687. https: // doi. org / 10.3390 / ani 11030687","Yuasa H. 1937. Description of a new genus of the family Lampyridae (Coleoptera). Kontyu, Tokyo 11 (1 - 2): 107 - 109.","Motschulsky V. 1854. Lampyrides. Etudes Entomologiques 3: 47 - 62.","Kawashima I., Suzuki H. & Sato M. 2003. A check-list of Japanese fireflies (Coleoptera, Lampyridae and Rhagophthalmidae). Japanese Journal of Systematic Entomology 9 (2): 241 - 261."]}

The firefly subgenus Hotaria sensu lato of the genus Luciola currently includes four morphospecies: L. (H.) parvula, L. (H.) unmunsana, L (H.) papariensis, and L. (H.) tsushimana. The latter three are taxonomically controversial based on both morphological and molecular data. We examined the phylogenetic relationships and evolutionary history of the species and related congeners using partial COI gene sequences (DNA barcoding). Our phylogenetic analyses consistently supported the monophyly of Hotaria sensu lato, but did not resolve the generic rank. The two types of L. (H.) parvula in Japan can be considered distinct species that arose by pseudocryptic speciation during the Miocene, with substantial genetic divergence (15.41%). Three morphospecies, L. (H.) unmunsana, L (H.) papariensis, and L. (H.) tsushimana, split into several polyphyletic or paraphyletic groups, forming entangled species groups. They are considered an incipient group that is distinguishable genetically but not morphologically, with evidence for recent allopatric speciation events corresponding to geologic events and sea-level changes during the Pliocene and Pleistocene. Group III of L. (H.) unmunsana collected from the Jeolla region is a new taxon.

Luciola lambkinae Keller and Ballantyne, nomen novum Luciola roseicollis Pic 1935: 6. McDermott 1966: 112 partim; Fu et al. 2010: 15 partim (nec Luciola roseicollis Pic 1933: 109). Type. Male. CHINA as Kina S Kansu. MNHN. Labelled: 1. Handwritten roseicollis n sp; 2. handwritten voisin de calceata; 3. Red card printed TYPE; 4. Printed Sven Hedins Exp. Ctr. Asien Dr Hummel; 5. Printed Kina S. Kansu. Etymology. The species is named for Christine Lambkin (Queensland Museum, South Brisbane, Australia). Dr. Lambkin has been collaborating with LB since 2000 trying to make sense of the Australopacific and Asian lucioline fauna. This is a small honor for 21 years of shared interest and collaboration. Diagnosis. Rosy pink pronotum; narrow anterior dark marking may reflect retraction of fat body in this area and the dark color is probably the dark head showing through the transparent cuticle; mesoscutellum dingy yellow; elytra black and subparallel-sided; most of ventral surface (and head) black except for brown femora of hind legs (interpretation of other legs not possible), and white light organ occupying all of the terminal two abdominal ventrites except for a narrow median posterior yellowish portion; lateral margins of light organ in ventrite 7 converge posteriorly, and posterior margin truncate, with very slight broad emargination. Remarks. This specimen was not dissected and thus reliable assignment to any genus is not possible. We have assigned it to the genus Luciola sensu lato, a category Ballantyne et al. (2019) used for Luciolinae species still not investigated nor assigned to any genus. This category is one of convenience and has no taxonomic standing. Pic (1935) highlighted the very slight emargination of the posterior margin of the ventrite 7 light organ (���ultimo truncato aut paulo emarginato���) and considered the species close to the similarly colored Luciola cerata Olivier differing in the pinkish pronotal tint with an anterior dark marking. Luciola cerata was redescribed from a lectotype designation by Fu and Ballantyne (2008) and subsequently assigned to the genus Abscondita by Ballantyne et al. (2013). It is only known from Taiwan., Published as part of Keller, Oliver & Ballantyne, Lesley, 2021, Luciola lambkinae Keller and Ballantyne, new replacement name for Luciola roseicollis Pic, 1935 (Coleoptera: Lampyridae), pp. 1-3 in Insecta Mundi 2021 (874) on pages 1-2, DOI: 10.5281/zenodo.5113782, {"references":["Pic M. 1935. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sii Ping-chang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927 - 1930. 16. Coleoptera. 2. Helmidae, Dermestidae, Anobiidae, Cleridae, Malacodermata, Dascillidae, Heteromera (ex. p.), Bruchidae, Cerambycidae, Phytophaga (ex. p.). Arkiv for Zoologi 27 A (2): 1 - 14.","McDermott FA. 1966. Lampyridae. p. 1 - 149. In: Steel WO (ed.). Coleopterorum Catalogus Supplementa, pars 9. Uitgeverij Dr. W. Junk; Gravenhage, Netherlands. 149 p.","Fu X-H, Ballantyne LA, Lambkin CL. 2010. Aquatica gen. nov. from mainland China with a description of Aquatica wuhana sp. nov. (Coleoptera: Lampyridae: Luciolinae). Zootaxa 2530 (1): 1 - 18.","Pic M. 1933. Malacodermes exotiques. L'Echange. Revue Linneenne 49 (451): 109 - 112.","Ballantyne LA, Lambkin CL, Ho JZ, Jusoh WFAW, Nada B, Nak-Eiam S, Thancharoen A, Wattanachaiyingcharoen W, Yiu V. 2019. The Luciolinae of S. E. Asia and the Australopacific region: A revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species. Zootaxa 4687 (1): 1 - 174.","Fu X-H, Ballantyne LA. 2008. Taxonomy and behaviour of lucioline fireflies (Coleoptera: Lampyridae: Luciolinae) with redefinition and new species of Pygoluciola Wittmer from mainland China and review of Luciola LaPorte. Zootaxa 1733 (1): 1 - 44.","Ballantyne LA, Fu X-H, Lambkin CL, Jeng M-L, Faust LF, Wijekoon WMCD, Li D, Zhu T. 2013. 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). Zootaxa 3721 (1): 1 - 48."]}

Luciola roseicollis Pic, 1933 Luciola roseicollis Pic 1933: 109. McDermott 1966: 112 partim. Fu et al. 2010: 15 partim. Type. Not located. Diagnosis. In the absence of any reliably identified specimens this diagnosis is based on Pic���s original description. 7 mm long. With thorax rosy red, mesoscutellum yellow; elytra black; having a distinct costa (���in disco distincte costatis���). Remarks. Pic���s (1933) description relied on color patterns but he did highlight the presence of a (?single) elytral costa or ridge which is a feature of the genus Curtos Motschulsky (see Ballantyne et al. 2019) and indicated the similarity of this species to Luciola impolita Olivier, which has subsequently been assigned to Curtos. Curtos is also known to have distinctive dorsal punctation but Pic described the elytral punctation as ���minute et dense punctatis���. We are unable to interpret what Pic intended in his description of the elytral apices as ���depressis���. Assignment to the genus Luciola sensu lato is made under the same provisions as outlined above. Fu et al. (2010) confused the issue in referring to Luciola roseicollis Pic, 1933. While their description of color patterns followed the 1933 format, they incorrectly referenced Pic���s (1933) description of the abdomen as possibly being that of a female. Pic (1935) had described the abdomen as truncated and slightly emarginate which suggested a female., Published as part of Keller, Oliver & Ballantyne, Lesley, 2021, Luciola lambkinae Keller and Ballantyne, new replacement name for Luciola roseicollis Pic, 1935 (Coleoptera: Lampyridae), pp. 1-3 in Insecta Mundi 2021 (874) on page 2, DOI: 10.5281/zenodo.5113782, {"references":["Pic M. 1933. Malacodermes exotiques. L'Echange. Revue Linneenne 49 (451): 109 - 112.","McDermott FA. 1966. Lampyridae. p. 1 - 149. In: Steel WO (ed.). Coleopterorum Catalogus Supplementa, pars 9. Uitgeverij Dr. W. Junk; Gravenhage, Netherlands. 149 p.","Fu X-H, Ballantyne LA, Lambkin CL. 2010. Aquatica gen. nov. from mainland China with a description of Aquatica wuhana sp. nov. (Coleoptera: Lampyridae: Luciolinae). Zootaxa 2530 (1): 1 - 18.","Ballantyne LA, Lambkin CL, Ho JZ, Jusoh WFAW, Nada B, Nak-Eiam S, Thancharoen A, Wattanachaiyingcharoen W, Yiu V. 2019. The Luciolinae of S. E. Asia and the Australopacific region: A revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species. Zootaxa 4687 (1): 1 - 174.","Pic M. 1935. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sii Ping-chang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927 - 1930. 16. Coleoptera. 2. Helmidae, Dermestidae, Anobiidae, Cleridae, Malacodermata, Dascillidae, Heteromera (ex. p.), Bruchidae, Cerambycidae, Phytophaga (ex. p.). Arkiv for Zoologi 27 A (2): 1 - 14."]}

In this study, we announce the complete mitochondrial genome (mitogenome) of the Korean endemic firefly, Luciola unmunsana Doi, 1931. The full-length circular genome was 15,858 bp, with 77.94% A/T content. It contained the typical set of 37 metazoan genes: 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, and 2 ribosomal RNA (rRNA) genes, as well as an A + T-rich region. The gene arrangement of the species is identical to that of the ancestral arrangement found in the majority of insects. The maximum-likelihood tree, built using all PCGs and two rRNAs via randomized accelerated maximum likelihood (RAxML) showed that L. unmunsana was grouped as a sister to L. curtithorax with the highest nodal support. However, another Luciola species clustered with the Aquatica species such that the genus Luciola was a non-monophyletic group. Therefore, more sampling is required to clarify the phylogeny of Luciola.

Several studies have been carried out on flashing of fireflies till now. From patterns of its flashes in different ambient conditions, different conclusions have been drawn and hypotheses put forward. Here we observe flashes emitted by two Indian species of firefly Luciola praeusta and Asymmetricata circumdata. For the species L. praeusta, as the temperature is lowered below about 21 °C, simple pulses become compound or combination ones. Males begin to emit bi-modal flashes while females emit both bi- and tri-modal flashes, along with the simple ones, in a regular manner. For the species A. circumdata, this feature is evident even at their normal flashing temperatures. Though rare, three-peaked flashes do appear from male specimens of both these species, and decay times of all the three peaks in a male- or a female-flash come out to be a few tens of, or a hundred-odd, milliseconds - in contrast to the nanosecond lifetimes determined by different workers for different analogs of the light emitter molecule oxyluciferin. We propose that the tri-modal feature in a flash represents three luminescent forms of the excited state of oxyluciferin, which in the normal flashing state in the living firefly decays via a pathway followed by molecules exhibiting phosphorescence.

To date, two Iranian luciferase genes from the Lampyris turkestanicus and Lampyroidea maculata have been carefully studied. Here, we report the cloning and characterization of the gene and protein of luciferase enzyme from the beetle of an Iranian lampyrid species, Luciola sp. (Coleoptera-Lampyridae). In this study, a Luciola sp. firefly was collected from the Yasouj area of Iran and its luciferase gene sequence was cloned and characterized. The genomic DNA length for this luciferase was the 1950 bp that combined of seven exons and separated by six introns. The results of multiple sequence alignment show that this gene has the most similarity with DNA gene luciferase from the Hotaria unmunsana species. Further analysis determined accurately the location of these introns in the luciferase gene. However, the deduced amino acid sequences of the luciferase gene (548 residues) showed that this luciferase had 97.8% resemblance to luciferase from Lampyroidea maculata species. By in silico modeling of firefly luciferase in an I-TASSER server, the 3D structure of this enzyme was evaluated. The results of phylogenetic tree analysis display the close evolutionary relationship of this luciferase gene and luciferase gene from the Lampyroidea maculata and Hotaria unmunsana.

Simple Summary Fireflies have a scattered distribution in Singapore but are not as uncommon as many would generally assume. A nationwide survey of fireflies in 2009 across Singapore documented 11 species, including “Luciola sp. 2”, which is particularly noteworthy because the specimens were collected from a freshwater swamp forest in the central catchment area of Singapore and did not fit the descriptions of any known Luciola species. Ten years later, we revisited the same locality to collect new specimens and genetic material of Luciola sp. 2. Subsequently, the mitochondrial genome of that population was sequenced and specimens were subjected to rigorous morphological examinations. We then collated published mitogenomes and shorter mitochondrial markers from closely related taxa to infer a phylogeny for the subfamily Luciolinae. Our results reveal that Luciola sp. 2 is both genetically and morphologically distinct from other congeners and is thus described herein as a new species Luciola singapura sp. nov. This marks 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 that can still harbor undescribed biodiversity. Abstract 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.

Luciola italica (Linnaeus, 1767) La luciole italienne est répartie de l’Italie jusqu’à la Grèce en passant par la péninsule balkanique selon Geisthardt & Satô (2007), mais sa distribution devrait être précisée suite aux résultats d’études génétiques obtenus par Day et al. (2014) (voir Discussion plus bas). Une population de L. italica forte de plusieurs centaines d’individus vient d’être découverte en France dans un petit village du département du Doubs à Blussangeaux, en juin 2018. Elle résulte d’une introduction accidentelle dans les années 1980 à partir d’individus provenant de Varese en Italie, vraisemblablement avec des plantes ornementales (Le Tallec & Cotte 2020). En Suisse, elle est indigène au Sud des Alpes (Tessin et Val Mesolcina aux Grisons), dans la continuité des populations d’Italie du Nord. Au Nord des Alpes par contre, toutes les populations recensées résultent d’introductions (Fig. 4a). Certaines sont très bien documentées, notamment celle de la région lausannoise. Faes (1941) explique ainsi s’être fait envoyer en 1938 des lucioles du Tessin qu’il introduisit près de Lausanne, «dans le parc Bourget, non loin de la rivière Chamberonne». Ce premier essai d’introduction fut néanmoins un échec et aucun individu ne put être trouvé en 1939 («peut‐être les insectes envoyés l’année précédente étaient‐ils exclusivement des mâles?»). En 1940, son correspondant tessinois lui renvoya de nouveaux spécimens et notamment des femelles, que Faes relâcha auprès de l’étang du parc Bourget. Des individus furent ensuite régulièrement observés sur ce site dès 1941. Cette population, issue de seulement quelques individus, était toujours présente en 2019 dans le même site. Actuellement forte de milliers d’individus (V. Cosandey, comm. pers.), elle a pourtant connu d’importantes fluctuations au fil du temps.Allenspach& Wittmer écrivaient ainsi en1979:«Toumayeff H. [Georges?], le 20.6.197 6 n’a pu trouver que quatre exemplaires. Il faut donc compter sur l’extinction prochaine de cette population au Nord des Alpes». Cela ne s’est toutefois pas produit. En parallèle à cette introduction dans la région lausannoise, Faes a également tenté une introduction de l’espèce en Valais central, qui est restée sans succès, comme il le constatait lui-même en 1941: «l’essai d’acclimatation tenté en 1938 seulement, auprès du petit lac de Géronde, sur Sierre en Valais, semble n’avoir pas eu de succès, les bords du lac (vignes et rochers) étant peut‐être trop arides». Un exemplaire de L. italica étiqueté « Pfynwald [Finges, près de Sierre], 18.– 27.7.1966, leg. Anonymous » existe pourtant dans les collections du Naturhistorisches Museum der Burgergemeinde Bern. Une petite population s’est donc peut-être tout de même maintenue dans la région durant deux décennies, ou cette capture résulte d’une nouvelle importation. Dans tous les cas, l’espèce n’a plus été signalée en Valais depuis plus de cinquante ans. Mis à part ces deux introductions locales attestées, il existe également plusieurs autres implantations de l’espèce au Nord des Alpes, vraisemblablement plus récentes et généralement non documentées. L. italica existe ainsi dans les environs d’Yvonand (VD). Plusieurs spécimens capturés en 2019 (29.6.2019, leg. A. Maibach) nous ont permis de confirmer l’identification spécifique. Suite à des discussions avec M. Antoniazza, il apparaît qu’une population existe dans cette localité depuis au moins 1975. Cette population a toujours été très localisée et n’a jamais colonisé des sites similaires situés le long de la Menthue, à quelque 400 m de distance, respectivement 600 à 700 m en amont (A. Maibach, comm. pers.). Une autre population, aujourd’hui éteinte, existait également à proximité dans la région de Châble-Perron dans les années 1970. Dans la région zurichoise, les premières données datent d’une trentaine d’années (Zürich, Kreuzkirche, 1989, leg. C. Meier; plusieurs centaines d’individus, Zürich-Hottingen, 2.– 12.7.1991, leg. A. Ion Plitzco & R. Neumeyer), mais cette population existe depuis au moins 1952 (S. Hostettler, comm. pers., in Neumeyer 1991). Dans cette région, l’espèce colonise principalement une prairie de fauche de basse altitude d’environ 0,25 ha située sur une pente exposée au sud et partiellement ombragée par les arbres d’un parc. Ce biotope est entouré de zones défavorables pour les larves et l’espèce s’y trouve confinée. La capture d’un spécimen en 2019 (leg. R. Neumeyer) permit de confirmer l’identification spécifique. Les mâles (Fig. 4d et 5a) et les femelles (Fig. 5b) de L. italica se ressemblent beaucoup du point de vue morphologique, les femelles étant toutefois plus trapues. Les mâles émettent en vol différents signaux lumineux jaune‐vert sous forme d’éclairs répétés et les femelles, au sol, clignotent pour attirer les mâles. Selon Mikšić & Mikšić (1965), les femelles possèdent des élytres, mais leurs ailes, incomplètement formées, ne leur permettent pas de voler. L’espèce dépasse rarement les 800 m d’altitude en Suisse (Fig. 4b) et les adultes s’observent principalement en juin (Fig. 4c)., Published as part of Gurcel, Kevin, Chittaro, Yannick, Sanchez, Andreas & Rieger, Ingo, 2020, Contribution à la connaissance des lucioles et lampyres de Suisse et observation de Luciola lusitanica Charpentier, 1825 à Genève (Coleoptera, Lampyridae), pp. 81-96 in Entomo Helvetica 13 on pages 86-88, DOI: 10.5169/seals-985886, http://zenodo.org/record/8110753, {"references":["Geisthardt M. & Sato M. 2007. Lampyridae. In: Lobl I. & Smetana A. (Eds), Catalogue of Palaearctic Coleoptera, Vol. 4. Elateroidea - Derodontoidea - Bostrichoidea - Lymexyloidea - Cleroidea - Cucujoidea, pp. 225 - 234. Apollo Books, Stenstrup.","Day J. C., Bonaduce A., Sabelli B. & de Cock R. 2014. Phylogeography of European Fireflies: Five Species in One. Firefly Symposium in Florida. Poster, non publie.","Le Tallec Q. & Cotte B. 2020. Decouverte de Luciola italica (Linnaeus, 1767) dans le Doubs, nouvelle espece pour la faune de France (Coleoptera Lampyridae). L'Entomologiste. In print.","Faes H. 1941. L'introduction de la luciole (Luciola italica) dans le canton de Vaud. Bulletin de la Societe Vaudoise des Sciences Naturelles 61 (256): 451 - 452.","Neumeyer R. 1991. Ein neuer Fundort von Luciola italica (Col., Lampyridae) nordlich der Alpen. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 64 (3 - 4): 349 - 350."]}

Luciola lusitanica (Charpentier, 1825) Citée de toute l’Europe méridionale, du Portugal à l’Ukraine (Geisthardt & Satô 2007), cette espèce présente une large distribution qui doit cependant être précisée. Si elle est bien connue d’Italie et n’est confirmée que de quelques localités en France, principalement des Alpes-Maritimes et plus rarement du Var, du Gard et de Corse (Constantin 2014), Day et al. (2014) indiquent sur la base de résultats génétiques que les spécimens de L. lusitanica du Portugal et de Grèce pourraient appartenir à deux autres espèces distinctes. Tout comme L. italica, les femelles de L. lusitanica (Fig. 4f et 5d) sont plus trapues que les mâles (Fig. 4e et 5c) et possèdent des élytres mais pas d’ailes complètement formées, si bien qu’elles se déplacent au sol. Selon Papi (1969), les mâles de L. lusitanica émettent un flash en vol toutes les 1,08 secondes. Les femelles prêtes à s’accoupler répondent par trois flashs courts à la suite, avec un intervalle de 70 à 250 ms selon la température. En Suisse, une donnèe historique est rapportèe du Tessin par Allenspach & Wittmer (1979), mais l’espèce s’avère absente du travail de synthèse de Geisthardt & Satô (2007). L’identification spécifique a pu être confirmée récemment: il existe en effet un spécimen de cette espèce étiqueté « Locarno, 1886, leg. O. Schneider-Orelli » au Naturhistorisches Museum Basel dans la collection de V. Allenspach et deux spécimens identiques sont déposés à l’Eidgenössische Technische Hochschule, Zürich. S’agissant de l’unique mention de l’espèce pour le Tessin, alors que de très nombreuses L. italica tessinoises existent dans les collections suisses, cette donnée doit être considérée comme douteuse (erreur d’étiquetage des spécimens?), ou résulte peut-être d’une importation ponctuelle au Tessin. En 2017, l’un de nous (KG) a été intrigué par l’apparence des lucioles genevoises (Fig. 4e–f) qui avaient été identifiées comme L. italica par la communauté naturaliste locale. Des comparaisons plus poussées sur photos indiquaient en effet qu’il pouvait s’agir de L. lusitanica. Grâce à l’appui de M. Geisthardt et la collecte d’échantillons supplémentaires en 2019, il nous fut en effet possible de confirmer l’identification spécifique de L. lusitanica. La première donnée genevoise authentifiée date de 2005 (Bernex, 12.7.2005, leg. G. Carron, un spécimen étant conservé au Musée d’histoire naturelle de Neuchâtel). Bien que portant une étiquette d’identification de M. Geisthardt l’attribuant à L. italica, nos identifications mènent également à L. lusitanica. Mais la prèsence de l’espèce dans la région genevoise semble antérieure à 2005, en particulier sur la commune de Satigny. La principale station est une pépinière et les lucioles y sont observées depuis 1998 au moins (G. Bousquet, comm. pers.). Plus ponctuellement, une autre observation fut rapportée de la commune de Collonge-Bellerive, où trois mâles en vol furent repérés le soir du 13.6.2017 dans le jardin d’une propriété privée située aux Crêts-de-la-Capite (B. Guibert, comm. pers.). Bien que l’espèce ne soit pas originellement indigène en Suisse et que toutes les observations résultent probablement d’importations, L. lusitanica forme actuellement des populations pérennes sur notre territoire (Fig. 4a) et est donc considérée comme rèsidente dans le pays., Published as part of Gurcel, Kevin, Chittaro, Yannick, Sanchez, Andreas & Rieger, Ingo, 2020, Contribution à la connaissance des lucioles et lampyres de Suisse et observation de Luciola lusitanica Charpentier, 1825 à Genève (Coleoptera, Lampyridae), pp. 81-96 in Entomo Helvetica 13 on pages 88-90, DOI: 10.5169/seals-985886, http://zenodo.org/record/8110753, {"references":["Geisthardt M. & Sato M. 2007. Lampyridae. In: Lobl I. & Smetana A. (Eds), Catalogue of Palaearctic Coleoptera, Vol. 4. Elateroidea - Derodontoidea - Bostrichoidea - Lymexyloidea - Cleroidea - Cucujoidea, pp. 225 - 234. Apollo Books, Stenstrup.","Constantin R. 2014. Contribution a l'etude des Lampyridae de France, actualisation de leur distribution et observations en France de Lampyris iberica Geisthardt, Figueira, Oay & Oe Cock, 2008 (Coleoptera, Elateroidea). Le Coleopteriste 17 (1): 34 - 44.","Day J. C., Bonaduce A., Sabelli B. & de Cock R. 2014. Phylogeography of European Fireflies: Five Species in One. Firefly Symposium in Florida. Poster, non publie.","Papi F. 1969. Light emission, sex attraction and male flash dialogues in a firefly, Luciola lusitanica (Charp.) Monitore Zoologico Italiano - Italian Journal of Zoology 3: 135 - 184.","Allenspach V. & Wittmer W. 1979. Cantharoidea, Cleroidea, Lymexylonoidea. Insecta Helvetica 4. Schweizerischen Entomologischen Gesellschaft, Zurich, 139 pp."]}

Luciola sp. (spécimens non identifiés à l’espèce) Mis à part les populations attribuèes à Luciola italica ou L. lusitanica, pour lesquelles nous avons généralement pu vérifier des spécimens collectés, il existe encore quelques populations de lucioles du Nord des Alpes pour lesquelles les preuves permettant une attribution spécifique sont actuellement insuffisantes (photos peu nettes, pas de spécimens collectés…). Ces populations sont pour l’heure et par prudence attribuées à Luciola sp. Une population située à Uster (ZH) dans un jardin privé a ainsi été signalée en 2016, bien qu’elle existe depuis au moins 2006 (observateur souhaitant rester anonyme, comm. pers. à I. Rieger en 2016). L’espèce a vraisemblablement été introduite avec des plantes dans ce périmètre restreint. Une autre population est connue à Niederhasli (ZH) depuis 2015 (leg. H. Brunner). La population se trouvant à côté d’une gare industrielle, une introduction liée à des wagons de fret est supposèe. Les larves se dèveloppent dans environ 1 ha de champs entourés de zones industrielles, dont l’extension actuelle pourrait menacer directement la population. Dans cette population, l’un de nous (IR) a pu étudier la communication lumineuse entre mâles et femelles et a constaté que lorsque les mâles reçoivent des réponses (flashs) de la femelle, ils volent plus bas et produisent une lumière continue. À Widnau (SG), une population de plusieurs centaines d’individus a été découverte en 2017 (leg. R. Naef) à côté de l’autoroute. L’espèce semble cantonnée à une petite parcelle d’environ 200 m de long, bien que des milieux apparemment favorables existent à proximité. La contiguité de l’autoroute laisse supposer une introduction via un véhicule (transport de plantes?)., Published as part of Gurcel, Kevin, Chittaro, Yannick, Sanchez, Andreas & Rieger, Ingo, 2020, Contribution à la connaissance des lucioles et lampyres de Suisse et observation de Luciola lusitanica Charpentier, 1825 à Genève (Coleoptera, Lampyridae), pp. 81-96 in Entomo Helvetica 13 on pages 90-91, DOI: 10.5169/seals-985886, http://zenodo.org/record/8110753

Numerous studies have been carried out on different aspects of the light from summer-active fireflies. Characteristics of this light have led to very interesting conclusions on the chemiluminescence reaction as well as on the nature of the light from live fireflies. Here we present a first report on bioluminescence emissions from a newly found winter-active Indian species of firefly Diaphanes sp. The steady-state emission spectrum from this species comes out to be apparently similar to those from the other two Indian summer species, Luciola praeusta and Asymmetricata circumdata: asymmetric in nature with a little bit of change in the position of the peak wavelength and in the width of the full width at half maximum. An increase in temperature to approximately 28°C causes a red-shift in the peak wavelength, which probably indicates denaturation of the enzyme luciferase in the live, flashing condition. Emissions in the time domain reveal that the light is never completely off - it decreases in intensity to a low value, sometimes very close to zero, and then increases - a characteristic unheard-of till date. Flash durations are considerably longer than those from the two Indian summer species; those become shorter at about 28°C and increase to noticeably larger values at higher temperatures.

There is an urgent need to preserve the ever-decreasing number of different species of fireflies all over the world. We sought to develop a vitrification procedure to cryopreserve the firefly embryos. The late stages of Luciola praeusta Kiesenwetter embryos were collected. Several impermeable and permeable protectants with various concentrations in different mediums (TNM-FH insect medium, Grace's medium, Dulbecco's Modification of Eagle's Medium (DMEM) and Dulbecco's Phosphate-Buffered Saline (DPBS)) were used. Embryos culturing in TNM-FH medium yielded the highest survival rate of 75.3 ± 3.6%. One-step, two-step and three-step methods were used in equilibrium procedure respectively. The highest survival rate (73.7% ±3.2%) occurred in embryos treated by three-step method ((1.5 M ethylene glycol (EG) + 2 M EG plus 8% polyvinylpyrrolidone (PVP) + 3 M EG, 8% PVP and 15% trehalose). Additionally, embryos exposed to 0.5 M trehalose presented a significantly higher survival rate (71.8 ± 2.7%) than embryos preserved in 0.5 M sucrose.

The lampyrids (Coleoptera: Lampyridae) belong to the few insects that have been named and observed accurately for thousands of years. The most common names in European languages are formed as diminutives of light and related terms or they are composed of a term like light, fire and glow in combination with bug, fly or worm. There is a huge variety of idiomatic names - in Italian dialects e.g. about 500 different expressions denominating species like Lampyris noctiluca or Luciola italica are reported. Beyond that, lampyrids are interpreted as signs or used as metaphors in a wide range of the semantic field. Thus, the notion of lampyrids leads to a vast network of associations including such distinct and even contradictory significances as childhood, crop, doom, elves, fear, habitat change, idyll, love, luck, mortality, prostitution, solstice, stars and fleetingness of words and cognition. All these connotations are evoked by six features of the observed lampyrids: light in the darkness, season of appearance, mating, direction of flying in the case of species like Lamprohiza splendidula and Luciola sp., flashing (of Luciola-fireflies) and finally disappearance of lampyrids in landscape and perception. The examples of the presence of lampyrids in literature, visual arts, music and evidence in traditional popular culture collected in past years show that the accent of the connotations is shifted by the change of cultural context: in early times the seasonal indication was relevant for the agricultural calendar, the romantic connotations had their golden age during Belle Epoque while in recent decades the associations circle around the disappearance of glowworms and fireflies. Although, the observer is not normally aware of the full richness of significances, the extraordinary complexity of connotations is an important factor for the fascination produced by lightning bugs during summer nights.

Luciola angusticollis Olivier 1886 Luciola angusticollis Olivier 1886: 184; 1902: 74. Schultze 1916: 43. McDermott 1966: 99. Ballantyne et al. 2016 Tables 6, 7. Type. Male. PHILIPPINES. Not located. Remarks. Olivier (1886) described a small (5.5 mm long) species with yellowish dorsum having black elytral apices, abdominal segments anterior to the LO black, and the terminal ventrite rounded. He highlighted the rectangular outline of the pronotum which was considerably narrower than the elytra. While the dorsal colour pattern and outline of V7 suggests A bscondita, we do not know of any Abscondita species as small as this., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 153, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187.","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","Schultze, W. (1916) A catalogue of Philippine Coleoptera. The Philippine Journal of Science, XI (1), 43 - 44.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241"]}

Luciola japonica (Thunberg 1784) Lampyris japonica Thunberg 1784: 79. Luciola japonica (Thunberg). Motschulsky 1854: 48. McDermott 1966: 107 (partim). Kawashima Suzuki et Sâtô 2003: 256. Sâtô 1974: 133. Type. Female. JAPAN. Upsala University Upsala Sweden. Other specimen examined. JAPAN. Labelled: 1. Handwritten ‘ Luciola Gorhami Ritsema affinis Gorh’; 2 handwritten ‘ Luciola praeusta’; handwritten ‘Yokohama’; 4. Printed ‘Japan G. Lewis 1910-320’. (NHML). Remarks. Kawashima et al. (2003) ably overviewed the confusion relating to the identification and misidentifications of this species and presented a colour picture of the holotype female. Despite the name they considered it was not from Japan and probably had never been. It is impossible to identify the type female as it belongs to a wide range of S. E. Asian species which have a pale yellowish dorsum with black tipped elytra. The single male from Japan in NHML was not dissected but appears to be an Abscondita sp. and has pale yellow abdominal tergites, and pale yellow ventrites anterior to the LOs. This would suggest Abscondita perplexa. However it presents a second problem. We are assuming that the Thunberg specimen was mislabelled and actually is not from Japan but from some other S. E. Asian country where Thunberg travelled. Short of suggesting that the species once existed in Japan and has since become extinct, the more likely explanation is that these two specimens were actually mislabelled. Japan reveres its firefly population and it is almost impossible to conceive that the species has gone unnoticed there., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 157-158, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Thunberg, C. P. (1784) Insectorum Novaes Species IV. In: Dissertatio Entomologica Novas Insectorum Species, Sistens, Cujus Partem Qvartum. Cons. Exper. Facult. Med. Upsal. Publice Ventilandam Exhibent, Upsaliae, pp. 69 - 84.","Motschulsky, V. de (1854) Lampyrides. Etudes Entomologiques, III, 47 - 62.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Kawashima, I., Suzuki, H. & Sato, M. (2003) A Check-List of Japanese Fireflies (Coleoptera, Lampyridae and Rhagophthalmidae). Japanese Journal of Systematic Entomology, 9 (2), 241 - 261."]}

Luciola intricata (Walker 1858) Colophotia intricata Walker 1858: 282. Luciola intricata (Walker). Olivier 1902: 81. McDermott 1966: 106. Ballantyne et al. 2016 Tables 6, 7. Type. Male (abdomen missing). SRI LANKA(as Ceylon). Labelled: 1. Male symbol; 2. Blue syntype; 3. Red Type; 4. Ceylon on one side, 59/106 on obverse (handwritten); 5. Intricata (handwritten); 6. Colophotia intricata Walker A. N. H. (type) (handwritten). (NHML) Diagnosis. In the absence of the abdomen it is possible to give certain diagnostic features of this species as follows: 6.1 mm long; dorsally yellowish brown with dark tipped elytral apices; head dingy yellowish brown with darker brown median line; labrum brown. Antennal sockets very close, ASD twice GHW; apical left labial palpomere broadly expanded in the form of a broad triangle with inner margin dentate (6 irregularly shaped teeth); no MFC. Luciola intricata differs most obviously from the very similarly coloured Serratia sibuyania gen. et sp. nov. by the lateral expansion of FS 2–8 and the broad apical labial palpomere the inner margin of which is dentate (in the latter only FS1–4 are clearly serrate, with the remaining FS elongate slender, and the apical labial palpomere is not much wider than long, with inner margin entire. Remarks. Including Serratia sibuyania gen. et sp. nov. we know of four Luciolinae species having serrate or flattened antennae. Luciola intricata (Walker) has FS 2–8 serrate (the type is without an abdomen). Olivier (1886) described as Luciola exstincta a single female from Alitaptap (Philippines) which is of similar size but different colouration to Serratia sibuyania gen. et sp. nov., and in which the antennal ‘articulis 4–8’ were expanded. It is unclear if Olivier was referring to the flagellar segments only (as we do here), or more likely to the antennal segments counting the scape as number 1. If the latter then his reference is to flagellar segments 2–6 which is not inconsistent with what we describe for Serratia sibuyania gen. et sp. nov. We cannot locate the type of this species and it is difficult to assign males to any species described from a female only. Antennal modifications in the Luciolinae are more usually seen in males (Ballantyne & Lambkin 2013). Bourgeois (1905) described as L. antennalis a single female from Sri Lanka as dorsally yellow with antennal segments 4−7 depressed. The type has not been located but the description of the antennal segments appears consistent with what we describe for Serratia sibuyania gen. et sp. nov. Bourgeois considered, and we agree, “elle mériterait peut-être de constituer un genre nouveau’. Wijekoon et al. (2012) recorded the species from various locations in Sri Lanka but their method involved a catch and release and it is not possible to confirm their identifications. Wijekoon (pers. com.) indicated his identifications were tentative only., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 157, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Walker, F. (1858) XXX. Characters of some apparently undescribed Ceylon Insects. Annals and Magazine of natural history, Series 3, 2 (No. 10), 280 - 286. https: // doi. org / 10.1080 / 00222935808697026","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241","Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187.","Ballantyne, L. A. & Lambkin, C. L. (2013) Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa, 3653 (1), 1 - 162. https: // doi. org / 10.11646 / zootaxa. 3653.1.1","Bourgeois, J. (1905) Voyage du D. W. Horn a Ceylan. Malacodermes et Lymexylonides. II. Lampyridini. Annales de la Societe entomologique de France, 74, 127.","Wijekoon, W. M. C. D., Wegiriya, H. C. E. & Bogahawatha, C. N. L. (2012) Regional diversity of fireflies of the subfamily Luciolinae (Coleoptera: Lampyridae) in Sri Lanka. Lampyrid, 2, 138 - 141."]}

Luciola spectralis Gorham 1880 Luciola spectralis Gorham 1880: 104. McDermott 1966: 113. Type. NEW ZEALAND. Male. (MNHN). Remarks. Based on a single male having pale dorsum and black tipped elytral apices, this species was incorrectly recorded from New Zealand and the single specimen was not dissected for any further examination., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 161, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Gorham, H. S. (1880) Materials for a revision of the Lampyridae. Transactions of the Entomological Society of London, 1880, 83 - 112.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149."]}

Luciola fukiensis Pic 1955 Luciola fukiensis Pic 1955: 29. McDermott 1966: 104. Ballantyne et al. 2016 Table 4. Type. CHINA. Not located. Remarks. Described as 6−7 mm long with yellow pronotum and black elytra; the colour pattern is suggestive of Aq. ficta as there may be a faint clear area on the front portion of the suture., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 156, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1955) Coleopteres nouveaux de Chine (Suite). Bulletin de la Societe Entomologique de Mulhouse, 1955, 21 - 23 + 25 - 26 + 29 - 32.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241"]}

Luciola pallidipes Pic 1928 Figs 292–308 Luciola pallidipes Pic 1928: 58. McDermott 1966: 111. Type. MALAYSIA. Malacca. Not located. Luciola fletcheri Pic 1935: 10. McDermott 1966: 104. New Synonymy. Type. Syntype male (no abdomen). INDIA. Sikkim. (NHML) 27.5– 20.9 N, 87.59E to 88.56E approximately. Type labelled: 1. Handwritten “ Luciola Fletcheri n. sp.” (Ballantyne interprets this as being in Pic’s handwriting); 2. Red printed “ Type ”; 3. Blue printed “ syntype ”; 4. Handwritten “ desiré”; 5. Sikkim 5000ft Kurseong 7.20 June 22 Fletcher coll. (NHML) (Figs 292, 293, 300, 301, 302). Other specimens examined. INDONESIA. Province of Maluku, island of Seram, 12 kms S. E. Wahai, 17.i−6. ii.1997 S. Bilý, male (Prague) (Figs 296–299). 0.03S 113.59E, Kalimantan, Tengah Busang / Rekut confl. MV light, August 2001, Brendell / Mendel, 2 males, female (NHML). BRUNEI. Temburong Prov. 14–15.ii.2010 Kuala Belalong F. stn., at light, 1 male (NHML) (Fig. 295). MALAYSIA. Nada’s collection in various forest reserves has compartment numbers and transect number following in []. # = killed fixed and preserved in 70% ethanol. Peninsular. Pahang: District, King George V Nat. Park: Kuala Tahang 7−14.xii.1958, J. L. Gressitt, 2 males (BPBM) (Figs 303, 304). Perak: Banjaran Bintang Bukit Berapit (Tai ping) 11– 12.3.1997, Ivo Jeniš, male (Prague) (Fig. 294). Pahang: Fraser’s Hill, Bishop Trail, 13.iv.2016, B. Nada, ♀ (B 1 I) (FRIM); 27.iv.2015, B. Nada, ♀ (FH 27 Out) (FRIM). Selangor: Gunung Nuang, 250 m a.s.l., 6.v.2015, B. Nada, ♂, 3 ♀ (GN 250) (FRIM); FRIM, Canopy Trail, 20.iv.2015, B. Nada, 2 ♀; 23.iv.2015, B. Nada, 2 ♂, ♀ (Canopy Trail 23) (FRIM); Mersawa Trail, 22.iv.2015, B. Nada, ♂, ♀ (Mersawa) (FRIM); Rover Track, 171 m a.s.l., 14.vi.2011, O. Khirul Faizal, male (ADULT 11) (FRIM); Non-dipterocarp Arboretum, 8.viii.2011, W.M. Wan Shaina, male (ADULTNOND3) (FRIM). Sabah: Mt Tambuyukon, 2350−2400m, 02.05.2011, Ultrabasic Team, 3 males (ANIC). Diagnosis. A small species (about 6 mm long) dorsally very dark brown to black, except for paler brown lateral surfaces of the metathorax and light brown legs in the L. fletcheri type, and paler legs in Perak and Maluku males; other specimens have dark margins to the metathorax and dark brown legs; white light organs in V6, 7, with pale T7, 8; eyes without a posterolateral emargination, inner eye margins very close above antennal sockets, and antennal sockets very close but not contiguous; median anterior margin of aedeagal sheath tergite broadly produced and apically squarely truncate (Figs 305, 306); posterior margin of sheath sternite rounded, entire. In the Brunei male the elytra are widely separated along their sutural margins and it is unclear if this is the natural state or due to dehydration. Female not reliably associated but coloured as for male and macropterous. Remarks. Four species approaching L. pallidipes in both size and colouration have been described. Luciola flebilis Olivier (1909a) from Sumatra was characterised from a female. Luciola pallidipes Pic 1928 from Malacca was described as 5 mm long and similar to L. picea but with legs and underbody of thorax yellow. Gorham (1882) described a small black species from Sumatra as L. picea. We are unable to locate types of either L. picea or L. pallidipes. It is possible that flebilis is simply a female of L. picea both having been recorded from Sumatra. The type male of L. fletcheri is without an abdomen, but the very close antennal sockets and small interocular distance is shared by the other specimens we assign here. While L. fletcheri was recorded from a high altitude in Sikkim, only some of the other specimens here are from similar altitudes. The Pahang males have the posterior margin of the light organ in ventrite 7 retracted. Both the Sabah and Maluku males have the posterior margin of the pronotum bisinuate and paler legs, while the Sabah males have antennae exceeding GHW in length. Specimens from Mt Kinabalu, Maluku and Perak have pale legs and underbody of thorax paler than rest. Preliminary observations on a collection of Luciolinae from peninsular Malaysia indicates L. pallidipes as defined by the dark dorsal colouration and paler brown legs is widely distributed. (B. Nada pers. comm.). The handwritten label ‘desiré’ on the type indicates that Pic hoped to keep the specimen., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 97-100, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1928) Malacodermes exptiques. L'Echange, Revue Linneenne, XLIV, 58 - 63.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Pic, M. (1935) Nouveautes diverses. Melanges exotico-entomologique Moulins, 66, 1 - 36.","Olivier, E. (1909 a) Descriptions de Lampyrides nouveaux du Musee Civique de Genes. Annali del Museo Civico di Storia Naturale di Genova, IV (3 a), 316 - 319.","Gorham, H. S. (1882) New species of Lycidae, Lampyridae and Telephoridae from Sumatra. Notes of the Leyden Museum, 4, 93 - 109."]}

Luciola antennalis Bourgeois 1905 Luciola antennalis Bourgeois 1905: 128. Wijekoon et al. 2012: 138; 2016: 67. Type. Female. SRI LANKA (as Ceylon). Not located. Remarks. Bourgeois (1905) described as L. antennalis a single female from Sri Lanka as dorsally yellow with antennal segments 4−7 depressed. We know of three other Luciolinae species described as having serrate or flattened antennae (see discussion following description of Serratia sibuyania gen. et sp. nov.) Luciola intricata (Walker) has FS 2–8 serrate and is also treated here as species incertae. Serratia sibuyania gen. et sp. nov. has four FS laterally expanded. The single female of Luciola exstincta Olivier (1886) has FS 4−8 expanded but is a different colour to Serratia sibuyania gen. et sp. nov. Wijekoon et al. (2012) recorded the species from various locations in Sri Lanka but their method involved a catch and release and it is not possible to confirm their identifications. Wijekoon (pers. com.) indicated his identifications were tentative only., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 153, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Bourgeois, J. (1905) Voyage du D. W. Horn a Ceylan. Malacodermes et Lymexylonides. II. Lampyridini. Annales de la Societe entomologique de France, 74, 127.","Wijekoon, W. M. C. D., Wegiriya, H. C. E. & Bogahawatha, C. N. L. (2012) Regional diversity of fireflies of the subfamily Luciolinae (Coleoptera: Lampyridae) in Sri Lanka. Lampyrid, 2, 138 - 141.","Wijekoon, W. M. C. D., Wegiriya, H. C. E. & Bogahawatha, C. N. L. (2016) Systematic revision of the repository collection of Canthoroidea (sic) in the Department of National Museums, Colombo, Sri Lanka (Coleoptera: Cantharidae, Lampyridae, Lycidae, Rhagophthalmidae). Ceylon Journal of Science, 45 (1), 67 - 74. https: // doi. org / 10.4038 / cjs. v 45 i 1.7365","Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187."]}

Key to species of Luciola s. str. as defined here, using males and associated females 1. Male............................................................................................... 2 Female............................................................................................ 14 2. Light organs absent; very dark brown to black species with pronounced posterolateral eye emargination; known only from New Caledonia (Ballantyne & Lambkin 2013 figs 120 121, 124, 125)............................... oculofissa Ballantyne Light organs present in at least one abdominal ventrite; eye emargination present or not............................. 3 3. Eyes with strong posterolateral emargination clearly visible in resting head (Ballantyne & Lambkin 2013 fig. 110)....... 4 Eyes without a visible posterolateral emargination, or with a weak emargination usually covered by the pronotum........ 5 4. No light organs in V7; light organs in V6 reduced to anterolateral plaques; known only from Fiji......... hypocrita Olivier Light organs in both V6 and 7 entire, may be retracted from lateral margins (Ballantyne & Lambkin 2013 figs 114, 119)....................................................................................... antipodum Bourgeois 5. Dorsal surface completely black or very dark brown.......................................................... 6 Dorsal surface never completely dark brown; pronotum usually pale coloured sometimes with a median dark marking..... 7 6. Known only from New Caledonia; aedeagal LL widely separated along their inner dorsal margins with a rounded projection along inner dorsal margins approximately half their length from their apices (Ballantyne & Lambkin 2013 figs 111, 112)...................................................................................... aquilaclara Ballantyne A S. E. Asian species sometimes from high altitudes; aedeagal LL not widely separated along their inner dorsal margins and without rounded projection as described above (Fig. 307)........................................... pallidipes Pic 7. Dorsal surface pale yellowish brown with dark elytral apices....................................................8 Dorsal surface not as above; pronotum may be black; elytra usually dark brown to black............................. 9 8. Ventral abdomen with median brown markings on V4–5 with lateral margins converging anteriorly (Figs 271, 275)........................................................................................... jengai Nada sp. nov. Ventral abdomen not coloured as above; ventrites anterior to the LO yellowish except for dark posterior margin of V5 (Fig. 285)................................................................................. niah Jusoh sp. nov. 9. Pronotum MN and MS black; elytra light brown with extensive darker brown markings at base and apex (Figs 309, 310)................................................................................ tiomana Ballantyne sp. nov. Dorsal surface with elytra black and pronotum pale coloured sometimes with a median dark marking.................. 10 10. Pronotum uniformly pale coloured with no darker markings.................................................. 11 Pronotum always with median darker markings.............................................................12 11. Large thick bodied, 10.5− 15 mm long (Chen 2003: 168)........................................ kagiana Matsumura Smaller, 6.5− 10 mm long (Chen 2003: 169 as Luciola sp. 3)........................................ satoi Jeng et al. 12. Pronotum orange with darker median markings; pronounced tubercles along posterior margin (Yiu 2017 fig. 22f).................................................................................................. tuberculata Yiu If pronotum with darker median markings then no tubercles across posterior margin............................... 13 13. Elytra dark brown to black with dark apices; 6 mm long species from Sri Lanka with dark brown elytra and orange pronotum having a wide median dark band (Figs 258, 266, 267)............................................ horni Bourgeois Elytra pale brown with white apices; 10 mm long; known from Vietnam; pronotum pale yellowish, may have pale brown median marking (Figs 246−247)........................................................... chapaensis comb. nov. 14. Female fully winged and flight capable (Chen: 172)............................................ curtithorax Olivier Female flightless..................................................................................... 15 15. Female with elytra usually covering abdomen and hind wings absent............................ parvula Kiesenwetter Female with short elytra up to twice as long as median pronotal length (Chen: 170, 171)................ filiformis Olivier

Luciola tuberculata Yiu 2017 Luciola tuberculata Yiu 2017: 96. Type. CHINA. Hong Kong. Holotype male (TLES). Diagnosis. Yiu (2017) distinguished this species from Hong Kong by the tubercles along the posterodorsal surface of the pronotum., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 102, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Yiu, V. (2017) A Study of Rhagophthalmidae and Lampyridae in Hong Kong with descriptions of new species (Coleoptera): Part 2. Lampyrid, 4, 67 - 119."]}

Luciola niah Jusoh sp. nov. Figs 284–291 Type. Holotype male. MALAYSIA. Sarawak Lambir Hill 24.ix.2010, 19.24, Nazeri Abghani (SK 0017 | MYFI 018-14) (FRIM) Figs 284, 285, 288–291. Paratypes. MALAYSIA. Sarawak: Kapit, I Polunin, no date 2 males (ANIC). Niah National Park forest track, M. Musbah and colleagues, 2.vii.2010, 2 males (SK 0023 | GenBank: KY 572917.1, MF 948239.1; SK 0064). Long Aton, Ulu Baram, 2.ix.2011, Musa Musbah 2 males (SK 0067 | GenBank: MF 063274.1; SK 0069 | GenBank: MF 063275.1) (FRIM). Diagnosis. 4.8–6.1 mm long; one of only two Luciola s. str. having dorsal colouration of yellowish brown with black elytral apices; other Luciola s. str. species with pale yellowish or pale brown pronota have dark brown to black elytra. Distinguished from the other similarly coloured species, L. jengai sp. nov., most obviously by the abdominal colour in jengai, and which also does not have leaf like lobes on the inner margins of the LL. Description of male. 4.8–6.1 mm long. Colour (Figs 284, 285): (Kapit males preserved in 70% ethanol differ in colour to remainder and are described here in brackets); pronotum yellowish with diffuse median darker markings (retraction of underlying fat body gives impression of darker markings); MN, MS and elytra yellowish brown with elytral apices black (elytra pale yellow with apices quite pale brown); elytra semitransparent and underlying hind wings may confuse interpretation of colour; head black, antennae and palpi dark brown (head reddish brown, antennae and palpi light brown); venter of thorax, basal abdominal ventrites and all abdominal tergites yellowish brown except for narrow dark posterior margin of V5 (all areas yellow except for thin dark marking on posterior margin of V5); legs yellowish brown with tibiae and tarsi dark brown (legs yellow, tarsi light brown); LO creamy white. Pronotum: slightly wider across middle (B> A, C); width a little less than humeral width; anterolateral corners angulate obtuse, posterolateral corners angulate. Elytron: subparallel sided. Head: minimal depression between eyes; GHW 11 X SIW; ASD much less than ASW (sockets are almost contiguous). Antennae: longer than GHW but not as long as 2 X GHW; segments elongate slender, scape longer than FS. Mouthparts: apical labial palpomere laterally flattened with inner edge dentate. Abdomen (Fig. 285): LO occupying entire area of V6, 7; V7 a little longer than wide, posterior margin rounded and no MPP; T8 with anterolateral arms elongate slender slightly longer than entire posterior portion which is rounded and without median emargination. Aedeagal sheath (Figs 288, 289): apical margin of sheath sternite narrowly prolonged and medianly slightly emarginated and hairy; median anterior margin of sheath tergite strongly prolonged and apically pointed.Aedeagus (Figs 290, 291): apices of LL narrow; LL longer than ML and widely divergent. Female: Jusoh associated the female by DNA barcode (Process ID: MYFI179-14 | GenBank: KY572915.1). Macropterous and capable of flight. Coloured as for male except for white LO restricted to V6 (Figs 286, 287). Bursa with two wide plates on each side. Etymology. The locality name Niah is used as a noun in apposition. Remarks. Ballantyne et al. (2016 Table 6) listed 30 species with this colour pattern (dorsally pale yellowish or pale brown with or without dark elytral apices) that is common among S. Asian Luciolinae. Many of these species have been assigned to species incertae here. Of the remainder the only species recorded from Sarawak having similar colouration is L. maculipennis Olivier, where the description of the terminal abdominal ventrite suggests a prolongation that could be typical of a Pygoluciola sp. We have presently little information about shape and numbers of bursa plates in this genus. Paired plates as seen here are characteristic of other non-related genera like Pteroptyx and Colophotia. Luciola italica does not have bursa plates. Jusoh et al. (2018) scored this species as “ Luciola WFA”., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 96-97, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241","Jusoh, W. F. A., Ballantyne, L., Lambkin, C. L., Hashim, N. R. & Wahlberg, N. (2018) The firefly genus Pteroptyx Olivier revisited (Coleoptera: Lampyridae: Luciolinae). Zootaxa, 4456 (1), 1 - 71. https: // doi. org / 10.11646 / zootaxa. 4456.1.1"]}

Luciola lata Olivier 1883 Figs 484−494 Luciola lata Olivier 1883: 79. Nec Luciola pallescens Gorham 1880: 102. Olivier 1890: xxx (synonymy). McDermott 1966: 111. Stat. rev. Type. BORNEO (no indication of specific area). Not located. Specimens examined. MALAYSIA. Sabah Danum Valley 4.28N 117.47E June 1999 H. Mendel, male (Figs 484−494) (NHML). Sarawak 4 th division Gn. Mulu NP. V–viii.1978 PM Hammond, JE Marshall female (NHML). Sarawak Quop, iii.1914, GE Bryant 2 females (NHML). Sarawak Nov. 1895 female, handwritten label C. miranda ♀ (MNHN). BRUNEI. Female labelled 1. Handwritten ‘ Colophotia miranda E. Ol. ♀ ’; 2. Handwritten ‘ Atyphella n. sp.’; 3. Handwritten ‘ Brunei N Borneo’ (MNHN). Male. For reasons outlined below it was considered appropriate to describe the single male listed above. 14.0 mm long. Colour: Pronotum and MN yellowish, semitransparent, with white patches of fat body irregularly scattered beneath cuticle; MS and elytra very pale brown, thin lateral margin of elytra paler; head antennae and palpi very dark brown; venter of thorax mid brown; legs yellowish with dark brown tibiae and tarsi; dorsal and ventral aspects of abdomen yellowish, except for white LO in V6, 7; cuticle semitransparent and underlying fat body is visible especially beneath tergites 7, 8. Pronotum: width subequal to humeral width; dorsal surface very convex, only depressed areas are across posterior margin. Elytra slightly convex sided; inner two interstitial lines visible. Head: GHW 5 X SIW; ASD ASW; apical labial palpomere laterally flattened with inner margin dentate. Antennae slightly longer than GHW, segments simple elongate. Abdomen: LO material retracted from posterior margin of V7; MPP well defined, about as long as wide and apically rounded. Aedeagal sheath: asymmetrical in posterior half only as sheath sternite is emarginated on both sides; short even edged emargination on left side of sternite posterior to tergite articulation; longer, deeper and wider emargination on right side, having a pointed projection at base near tergite articulation; anterior margin broadly rounded; anterior margin of sheath tergite evenly emarginated; posterior margin squarely truncate. Aedeagus: LL visible at sides of ML, broader at apices than apex of ML, separated along their mid dorsal length and with anterior margin produced and rounded; basal piece narrow; ML expanding a little along its length, then narrowed near apex; ML moderately strongly arched with apex inclined slightly ventrally and rounded. Remarks. It is appropriate to address these few specimens separately. This single male may represent Luciola lata. It may also be a new genus (the outline of the aedeagal ML is suggestive of Luciola s. str. but the aedeagal sheath is unlike that genus). The females are associated by similarity of size, colour pattern and nature of the pronotum only. In MNHN they stand under the label of Colophotia miranda. However doubts about the definitive identification of both males and females suggest further investigation is necessary. Whether this male represents Luciola lata is difficult to determine. Olivier (1883) described the species as 12 mm long, with an oblong shape with convex sided elytra which are wider at their base than the pronotum, and dorsally pale brown. However Olivier also indicated a median carina on the last abdominal ventrite (‘Dernier segment de l’abdomen ….charge sur son milieus d’une forte carène longitudinale obtuse’). This V7 median carina would indicate the species was most probably a Colophotia. The species was synonymised with L. pallescens Gorham by Olivier (1890), who indicated that he had seen Gorham’s type. However the apparent anomaly of a median carina on V7 cannot be explained (specimens of pallescens described here do not have a median carina on V7), and the synonymy of lata with pallescens is rejected. Chey (2008 fig. 3) illustrated as Luciola sp. 4 a large (14 mm long) male with pale dorsal colouration, entire LO in V7 and no median carina. This presently unidentified specimen approaches L. lata in size and colour only and may be conspecific with the single male we address here. Chey saw several males which he indicated were solitary fliers. The females are very likely conspecific with the single male. They also possess the somewhat convex dorsal surface of the pronotum, and the posterior margin of V7 is scarcely emarginated, unlike that of Abs. pallescens comb. nov. (which is strongly emarginated). We record here 22 females of Abs. pallescens from Quop of which six have yellowish pronotum; none have the convex dorsal surface we describe here. The confusion surrounding the identity of Colophotia miranda is addressed under that species and some of that discussion is relevant here. Olivier’s identification of specimens he assigned to C. miranda is confusing and contradictory. In 1886 he described a large male from the Philippines (Bohol) as totally yellow dorsally and with a single recurved appendage of the terminal abdomen. Subsequently (1913b) he indicated the occurrence of the species from Borneo, but this time the abdomen having “two long elevated appendicles curved upwards”. It is possible the first specimen may have been a species of Pygoluciola, or even a female, while the second specimen could have been a Colophotia sp. The issue is further confused as several Luciolinae have large females which approach Olivier’s (1886) description of C. miranda. Abscondita anceyi is a large pale specimen but with black elytral apices. Abscondita pallescens comb. nov. female is entirely pale brownish yellow dorsally except for variously sized darker brown marks on the pronotum. Luciola candezei and L. ochracea are both pale coloured dorsally but with distinctive black abdomens and are probably the same species. Luciola concreta Olivier from Java is a little smaller at 10 mms. See further discussion under Abscondita pallescens (Gorham) comb. nov. This male possesses several unique morphological features that could suggest a new genus is necessary: the convex pronotal outline, the emarginations of the posterior area of the aedeagal sheath sternite where there is a pointed projection. Further investigation when more specimens become available will be necessary., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 158-160, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Olivier, E. (1883) Lampyrides nouveaux ou peu connus. Revue D'Entomologie, 3, 73 - 80, 326 - 332.","Gorham, H. S. (1880) Materials for a revision of the Lampyridae. Transactions of the Entomological Society of London, 1880, 83 - 112.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Chey, V. K. (2008) Fireflies of Kionsom. Sepilok Bulletin, 8, 15 - 22.","Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187."]}

Luciola tiomana Ballantyne sp. nov. Figs 309–314 Types. Holotype male, paratype male. MALAYSIA. Johor: Tioman Isl. Jungle track 10–600 masl 15– 29.3.2009 V Hula (Prague). Terengganu: Pasir Raja Selatan Forest Reserve, Compt. 52, 29.vi. 2016, B. Nada, ♂ (PS 52 [4]) (FRIM). Nada’s collection in forest reserve has compartment number and transect number following in (). # = killed fixed and preserved in 70% ethanol. Diagnosis. A small species (4.2–4.3 mm long) with distinctive dorsal colouration of black head, pronotum, dark brown to black MN and MS, and brownish yellow elytra which have diffuse darker brown markings in apical ¾, with paler basal markings restricted to basal area near suture, and an accumulation of whitish fat body along apex of suture and round elytral apex. Description of Male. 4.2–4.3 mm long. Colour (Figs 309–313): pronotum black with narrow brownish orange band across posterior margin; MN and MS black in type, MS with narrow irregular paler brown lateral margins; MN and MS brownish orange in paratypes; elytra of holotype with basal ¼ and suture and lateral margin narrowly brownish orange; rest of elytron in type is diffuse dark brown with apices dark; elytra of paratype with basal paler colour occupying slightly less than basal ¼ and brownish orange colour of suture and lateral margin expanding around apex; head antennae palpi black, scape and pedicel paler on underside; venter of thorax and all ventrites anterior to the LO very dark brown to black; Legs brownish orange; LO in V6, 7 pale creamy white; T2–5 semitransparent grey, T6, 7 black, T8 semitransparent, pale. Pronotum: slightly narrower than humeral width; sides subparallel (A = B = C); posterolateral corners acute. Elytron: subparallel sided; interstitial lines not well defined. Head: GHW 10 x SIW; ASD Etymology. The type locality is latinised thus Tioman to tiomana, a noun in apposition. Remarks. The similarly coloured Luciola notaticollis Pic 1914 from Madura India is 7.0 mm long, with black pronotum, MN, and MS, and the basal area of the elytra (slightly less than half the length) orange, with the apical ½ of the elytron black. The type in MNHN was examined but not dissected as it is glued face down on a card. Ballantyne & Jusoh (2016) suggested a closer examination was necessary to determine if L. notaticollis should be assigned to Curtos., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 100-102, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1914) Nouveau genre, especes et varietes nouvelles de diverses familles. Melanges exotico-Entomologiques, 10, 3.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241"]}

Luciola truncata Olivier 1886 Luciola truncata Olivier 1886: 183; 1902: 87. Ballantyne & Lambkin 2009: 64. Colophotia truncata (Olivier). McDermott 1966: 116. Type. PHILIPPINES. Female. Not located by Ballantyne MNHN 2013. Remarks. Olivier (1886) described a 10 mm long female having a pale dorsum with black tipped elytra from the Philippines (Mindoro). He highlighted the pronotal outline and the paired impressions across the posterior margin. McDermott (1966) erroneously assigned the species to Colophotia. External female morphology does not permit generic assignments unless there are distinctive colour patterns and similarities of geographical locations., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 162, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187.","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","Ballantyne, L. A. & Lambkin, C. L. (2009) Systematics of Indo-Pacific fireflies with a redefinition of Australasian Atyphella Olliff, Madagascan Photuroluciola Pic, and description of seven new genera from the Luciolinae (Coleoptera: Lampyridae). Zootaxa, 1997, 1 - 188.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149."]}

Luciola ruficollis (Boisduval 1835) Lampyris ruficollis Boisduval 1835: 128. Guérin-Méneville 1838: 75. Motschulsky 1854: 53. Luciola ruficollis (Guérin-Méneville). Lacordaire 1857: 338. Olivier 1902: 85; 1907: 54; 1910a: 45. Nec Girard 1873, plate xxxv, figs 9, 9a, 9b. Luciola (Luciola) ruficollis (Guérin-Méneville). McDermott 1966: 112. Ballantyne, 1987a: 131. Ballantyne & Lambkin 2013: 123. Luciola ruficollis (Boisduval). Ballantyne & Lambkin 2013: 123. Type. NEW GUINEA. Not located. Remarks. This species has not been reliably identified in collections. It is 6 mm long with orange pronotum, black elytra and ventrite 7 trisinuate. Its possible identity was discussed by Ballantyne & Lambkin (2013)., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 161, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Boisduval, J. B. A. D. (1835) Voyage de decouvertes de l' Astrolabe, execute par ordre du Roi, pendant les annees 1826 - 1829, sous le commandement de M. J. Dumont- D'Urville. Faune entomologique de l'ocean Pacifique avec l'illustration des insecte nouveaux recueillis pendant le voyage. Deuxieme partie. Coleopteres et autres ordres. Tastu, Paris, 716 pp.","Motschulsky, V. de (1854) Lampyrides. Etudes Entomologiques, III, 47 - 62.","Lacordaire, T. (1857) Histoire naturelle des insectes. Genera des Coleopteres. Tome Quatrieme. Roret, Paris, 579 pp.","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","Olivier, E. (1907) Coleoptera, Lampyridae. Genera Insectorum, 53, 1 - 74. [Luciolinae, 48 - 56].","Olivier, E. (1910 a) Pars 9. Lampyridae. In: Coleopterorum Catalogus (Junk, Schenkling). Junk, Berlin, pp. 1 - 68. [Luciolinae, 40 - 48]","Girard, M. (1873) Les Insectes. Traite elementaire d'entomologie. Introduction. Coleopteres. Tome 3. J. - B. Bailliere et Fils, Paris, 1110 pp.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A. (1987 a) Further Revisional Studies on the firefly genus Pteroptyx Olivier (Coleoptera: Lampyridae: Luciolinae). Transactions of the American Entomological Society, 113, 117 - 170.","Ballantyne, L. A. & Lambkin, C. L. (2013) Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa, 3653 (1), 1 - 162. https: // doi. org / 10.11646 / zootaxa. 3653.1.1"]}

Luciola binhana Pic 1927 Luciola binhana Pic 1927: 37. McDermott 1966: 100. Ballantyne et al. 2016 Table 4. Type. VIETNAM (as Tonkin). Presumed male, fragmentary (part of head, antennae and two legs only). Labelled: 1 handwritten ‘ Hoa Binh’; 2 handwritten ‘type’; 3 printed on red label ‘type’; 4. Handwritten ‘ binhana s. sp.’ (MNHN). Remarks. Pic described the species with yellowish pronotum and brown elytra with pale margins. He thought it resembled L. limbatipennis Pic. Nothing sufficiently distinctive about this colour pattern, nor the locality data, would permit an attempt to associate males., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 154, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1927) Malacodermes exotiques. L'Echange, Revue Linneenne, XLIII, 37 - 52.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241"]}

Luciola marginipennis (Boisduval 1835) Lampyris marginipennis Boisduval 1835: 126 Luciola Marginipennis Laporte 1833: 151. Nomen nudum. Colophotia marginipennis Dejean 1837: 103. Nomen nudum. Germar 1848: 184. (Incorrect record from Adelaide). Luciola marginipennis Guérin-Méneville. Motschulsky 1854: 53. Lacordaire 1857: 338. Olivier 1902: 83. Lea 1909: 46, 106. McDermott 1966: 109. Luciola marginipennis Boisduval. Olivier 1913a: 417. Incorrect attribution of author to Guérin-Méneville. Ballantyne & Lambkin 2009: 106. Type. Ile Waigiou (may be Waigeo off NW tip of Irian Jaya 130E, 0S). Location of holotype unknown. Remarks. A large species (ca. 12 mm) with orange pronotum, black elytra with orange margins and red mesoscutellum. Ballantyne & Lambkin (2009: 106) discussed this species at length, concentrating on the original description of the MS as red to contrast it with specimens they assigned to Luciola limbata., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 160-161, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Boisduval, J. B. A. D. (1835) Voyage de decouvertes de l' Astrolabe, execute par ordre du Roi, pendant les annees 1826 - 1829, sous le commandement de M. J. Dumont- D'Urville. Faune entomologique de l'ocean Pacifique avec l'illustration des insecte nouveaux recueillis pendant le voyage. Deuxieme partie. Coleopteres et autres ordres. Tastu, Paris, 716 pp.","Laporte, F. L. de (1833) Essai d'une revision du genre Lampyre. Annales de la Societe Entomologique de France, 1833, 122 - 153.","Dejean, P. F. M. A. (1837) Catalogue des Coleopteres de la collection de M. Le Comte Dejean 4 th edition. Mequignon-Marvis, Paris, 176 pp.","Germar, E. F (1848) Beitrag zur insectenfauna von Adelaide. Linnaea Entomologica, III, 153 - 247.","Motschulsky, V. de (1854) Lampyrides. Etudes Entomologiques, III, 47 - 62.","Lacordaire, T. (1857) Histoire naturelle des insectes. Genera des Coleopteres. Tome Quatrieme. Roret, Paris, 579 pp.","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","Lea, A. M. (1909) Revision of the Australian and Tasmanian Malacodermidae. Transactions of the Entomological Society of London, 1909, 45 - 251. [Lampyridae, 106 - 111]","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Olivier, E. (1913 a) Lampyridae. Resultats de L'Expedition Scientifique Neerlandaise a la Nouvelle-Guinee en 1907 et 1909 sous les auspices de Dr H. R. Lorentz. Zoologie Livraison III. Nova Guinea, 9, 417 - 422.","Ballantyne, L. A. & Lambkin, C. L. (2009) Systematics of Indo-Pacific fireflies with a redefinition of Australasian Atyphella Olliff, Madagascan Photuroluciola Pic, and description of seven new genera from the Luciolinae (Coleoptera: Lampyridae). Zootaxa, 1997, 1 - 188."]}

Luciola curtithorax Pic 1928 Luciola curtithorax Pic 1928: 49. McDermott 1966: 102. Jeng et al. 2003b: 255. Chen 2003: 172. Fu et al. 2010: 15. Fu 2014: 26. Yiu 2017: 92. Type. Holotype male. VIETNAM. (MNHN). Other specimens examined. THAILAND. S.N. = Sorasak Nak-eiam. 18°14’ N, 98°32’ E, Chiang Mai province, Hot district, 286 msl., 31.v.2011, mixed deciduous forest, 2 males (SN ˗ CMB ˗003) S.N. (NUNHM). 18°13’ N, 98°29’ E, Chiang Mai province, Hot district, 364 msl., 31.v.2011, mixed deciduous forest, 8 males, 2 females (SN ˗ CMB ˗005) S.N. (NUNHM). 18°29’ N, 99°18’ E, Lamphun province, Mea Ta district, 1,214 msl., 28.v.2010, lower montane forest, 6 males, 2 females (SN ˗ LNB ˗001) S.N. (NUNHM). 18°50’ N, 99°28’ E, Lampang province, Muang Pan district, 484 msl., 10.v.2011, forest park, 17 males, 2 females (SN ˗ LPA ˗002) S.N. (NUNHM). 18°50’ N, 99°29’ E, Lampang province, Muang Pan district, 408 msl., 10.v.2011, paddy field area, 2 males (SN ˗ LPA ˗007) S.N. (NUNHM). 18°09’ N, 97°55’ E, Mae Hong Son province, Mae Sariang district, 213 msl., 1.vi. 2011, paddy field area, 2 males (SN ˗ MSB ˗008) S.N. (NUNHM). 18°09’ N, 97°54’ E, Mae Hong Son province, Mae Sariang district, 255 msl., 1.vi.2011, mixed deciduous forest, 5 males (SN ˗ MSB ˗011) S.N. (NUNHM). 18°09’ N, 97°53’ E, Mae Hong Son province, Mae Sariang district, 290 msl., 1.vi.2011, dry evergreen forest, 1 male (SN ˗ MSB ˗015) S.N. (NUNHM). 19°21’ N, 98°26’ E, Mae Hong Son province, Pai district, 514 msl., 3.vi.2011, freshwater area, 1 male (SN ˗ MSB ˗032) S.N. (NUNHM). 19°13’ N, 99°50’ E, Phayao province, Muang district, 401 msl., 11.v.2011, orchard, 1 male (SN ˗ PYA ˗009) S.N. (NUNHM). 19°07’ N, 99°50’ E, Phayao province, Muang district, 417 msl., 11.v.2011, paddy field area, 2 males, 2 females (SN ˗ PYA ˗011) S.N. (NUNHM). 18°55’ N, 100°13’ E, Phayao province, Chi- ang Muan district, 357 msl., 12.v.2011, mixed deciduous forest, 1 male (SN ˗ PYA ˗012) S.N. (NUNHM). 17°56’ N, 99°35’ E, Phrae province, Wang Chin district, 147 msl., 9.v.2011, freshwater area, 6 males (SN ˗ PRA ˗005) S.N. (NUNHM). 18°40’ N, 100°13’ E, Phrae province, Wang Chin district, 246 msl., 12.v.2012, freshwater area, 4 males (SN ˗ PRA ˗009) S.N. (NUNHM). 17°14’ N, 99°00’ E, Tak province, Sam Ngao district, 150 msl., 8.vii.2007, forest park, 1 male (SN ˗ TKB ˗011) S.N. (NUNHM). 15°28 ‘N, 99°17’ E, Uthai Thani province, Lan Sak district, 474 msl., 12˗16.v.2008, dry evergreen forest, 4 males, 1 female (SN ˗ UTA ˗002) S.N. (NUNHM). 15°36’ N, 99°19’ E, Uthai Thani province, Lan Sak district, 195 msl., 6.vi.2008, mixed deciduous forest, 8 males, 1 female (SN ˗ UTA ˗006) S.N. (NUNHM). 15°11’ N, 99°16’ E, Uthai Thani province, Ban Rai district, 325 msl., 7.vi.2008, dry evergreen forest, 1 male, 2 females (SN ˗ UTA ˗011) S.N. (NUNHM). 18°13’ N, 101°02’ E, Uttaradit province, Ban Khok district, 487 msl., 9.v.2009, forest park, 4 males (SN ˗ UDB ˗003) S.N. (NUNHM). Diagnosis. One of many Luciolinae with dark brown elytra; pronotum often pinkish, with a median dark brown area which reaches the anterior margin but not the posterior margin; most of ventral surface except for light organs very dark brown to black; the type male has lateral and most of the sutural elytral margin very narrowly orange; aedeagal sheath with posterior margin of sheath sternite rounded entire; median anterior margin of sheath tergite produced and rounded; aedeagus with basal 3/4 wide, narrowing in apical ¼ which is ¼ the width of the wider basal portion; leaf like lobes of LL elongate slender; LL closely approach along mid dorsal line and are expanded at their apices; anterolateral prolongations of T8 narrow, less than half the length of the posterior entire portion. Jeng et al. (2003b: 255) characterised this species as close to L. filiformis Olivier but having a more transverse pronotum and larger head. Fu (2014) depicted males with a narrow reddish sutural and lateral margin of the elytra. Females of filiformis are brachelytral while those of curtithorax are fully winged (Chen 2003: 172). Remarks. The redescription and the aedeagal diagrams of Jeng et al. (2003), Fu (2014: 27) and Yiu (2017: 92) indicate this species falls under Luciola s. str. We have followed Jeng’s characterisation of this species where they described specimens from Taiwan, but the identifications are further discussed below. The type has not been dissected. L. curtithorax was characterised and named from what appeared to Pic to be a short pronotum. Describing the shape as short may be a little misleading as the median anterior margin in the type is strongly prolonged anteriorly, and it may be that the somewhat short lateral margins give the impression of a short pronotum. The pronotum of the type is twice as wide as long and the ratio of the length of the lateral margins to median pronotal length is 2: 3. There are inconsistencies with the characterisation of several Asian species which we are here assigning to Luciola s. str. and there is need for further investigation of some. Fu et al. (2010) discussed the similarities between the descriptions of L. curtithorax and L. roseicollis Pic 1933 when they were justifying the erection of a new species Aquatica wuhana Fu et al. Pic indicated he thought roseicollis near L. impolita, which was subsequently assigned to Curtos (Jeng et al. 1998). Jeng et al. (2003b) distinguished L. satoi Jeng et al. from L. roseicollis by its smaller size (6.5−8 vs 8−9 mm long) and being without ‘rugulose’ sculpture on the elytra. However Ballantyne’s observations of the type of L. roseicollis (in MNHN, 2013) appear to conflict with those of Jeng and revealed a species with a uniformly orange pronotum, and elytra without any pronounced sculpturing. It is very similar to Luciola cerata which was transferred to Abscondita by Ballantyne et al. (2013). Pic indicated on a handwritten label with this specimen ‘voisin de calceata’ and L. calceata is transferred to Pygoluciola here. Yiu (2017) described specimens as ‘near curtithorax ’ which appear to conform with what we describe here. However his confusion with the similarity of this and other similarly coloured species (including laticollis Gorham and robusticeps Pic) was indicated in his Table 1 on page 94 but was not resolved. Yiu did not consider L. roseicollis in his discussion., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 90-91, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1928) Malacodermes exptiques. L'Echange, Revue Linneenne, XLIV, 58 - 63.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Jeng, M. L., Yang, P. S. & Lai, J. (2003 b) Notes on the genus Luciola (Coleoptera, Lampyridae, Luciolinae) of Taiwan. Special Bulletin of the Japanese Society of Coleopterolog y, 6, 247 - 262.","Chen, T. R. (2003) The fireflies of Taiwan. Field Image Publisher, Taipei, 256 pp. [in Chinese]","Fu, X. H., Ballantyne, L. & Lambkin, C. (2010) Aquatica gen. nov. from mainland China with a description of Aquatica wuhana sp. nov. (Coleoptera: Lampyridae: Luciolinae). Zootaxa, 2530 (1), 1 - 18. https: // doi. org / 10.11646 / zootaxa. 2530.1.1","Fu, X. H. (2014) An illustrated handbook of Chinese fireflies. The Commercial Press, Beijing, 167 pp. [in Chinese]","Yiu, V. (2017) A Study of Rhagophthalmidae and Lampyridae in Hong Kong with descriptions of new species (Coleoptera): Part 2. Lampyrid, 4, 67 - 119.","Jeng, M. L., Lai, J. & Yang, P. S. (2003 a) Lampyridae: A synopsis of aquatic fireflies with description of a new species. In: Jach, M. A. & Ji, L. (Eds.), Water Beetles of China III. Zoologisch-Botanische Gesellschaft in Osterreich, and Wiener Coleopterologenverein, Wien, pp. 1 - 572.","Pic, M. (1933) Notes diverses, descriptions et diagnoses. L'Echange, Revue Linneenne, XLIX, 109.","Jeng, M. L., Yang, P. S., Sato, M., Lai, J. & Chang, J. C. (1998) The genus Curtos (Coleoptera, Lampyridae, Luciolinae) of Taiwan and Japan. Japanese Journal of Systematic Entomology, 4, 331 - 347.","Ballantyne, L., Fu, X. - H., Lambkin, C., Jeng, M. - L., Faust, L., Wijekoon, W. M. C. D., Li, D. - Q. & Zhu, T. - F. (2013) 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). Zootaxa, 3721 (1), 1 - 48. https: // doi. org / 10.11646 / zootaxa. 3721.1.1"]}

Luciola trivandrensis Raj 1947 Luciola trivandrensis Raj 1947: 188. McDermott 1966: 114. Fu et al. 2012a: Table 1, 3. Type. INDIA. Not located. Raj (1947) reported that the three larvae he collected and tried to rear to adults, all died. Remarks. There are few clues to aid in the possible identification of this species as Raj presented figures of the dorsal surface of the larval head and antenna only. The projections he described along the posterior margins of most dorsal segments are suggestive of described larvae of Pygoluciola qingyu Fu et al. The three larvae were collected in marshy area but not recorded with gills., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 162, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Raj, J. S. (1947) Two species of undescribed Lampyrid larvae from S. India. Proceedings of the Indian Academy of Science, 25, 188 - 194.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Fu, X. H., Ballantyne, L. & Lambkin, C. (2012 a) Emeia gen. nov., a new genus of Luciolinae fireflies from China (Coleoptera: Lampyridae) with an unusual trilobite-like larva, and a redescription of the genus Curtos Motschulsky. Zootaxa, 3403 (1), 1 - 53. https: // doi. org / 10.11646 / zootaxa. 3403.1.1"]}

Luciola jengai Nada sp. nov. Figs 270−283 Types. Holotype male, paratype male. MALAYSIA. Pahang: Fraser’s Hill, Bishop Trail, 19.ix.2016, B. Nada (B 1 III) (FRIM). Terengganu: Jengai Forest Reserve, Compt.52, 30.vi.2016, B. Nada (JI 52 [5]) (FRIM). Diagnosis. 8.7 mm long. Very pale semitransparent yellow with small dark brown markings at elytral apex (Figs 270, 273). Most obviously distinguished from other Luciola by the colour of abdominal ventrites V4 and 5 (Figs 271, 272, 275), where the median area is dark brown with margins which narrow anteriorly, and the very close interocular distance which occurs above the insertions of the antennae. Macropterous females tentatively associated by similarity of colour pattern only and described separately below. These two specimens are tentatively included in Luciola s. str. (see comments below). Male. 8.7 mm long. Colour (Figs 270−275): pronotum creamy yellow semitransparent, with main colour due to underlying white fat body, which is retracted along anterior and posterior margins, (anterior margin may appear black because of underlying dark head); fat body retracted in four other patches in median area of disc corresponding to underlying muscle attachments and these areas appear slightly darker; MN creamy white; MS and elytra semitransparent and pale yellow, with elytral apex black; lateral and sutural elytral margins are very narrowly white due to underlying fat body which is scattered near apex and near inner margin in apical third; head, antennae and palpi black; venter of thorax orange-yellow, legs same with tibiae and tarsi and apices of femora black; basal abdominal ventrites orange yellow, V4, 5 with median dark brown markings: in B1 V5 has extensive dark markings with sloping sides, not reaching anterior margin, and V4 has darker markings restricted to two spots along posterior margin; J1 the dark markings in V5 reach the anterior margin of this ventrite, and V4 has similarly shaped markings which are narrower; all abdominal tergites except T8 are dark brown with T6, 7 almost black; T8 is pale semitransparent. Pronotum: width slightly less than humeral width. Elytron: interstitial lines not well defined; subparallel-sided. Head: barely depressed between eyes; GHW 10 X SIW which occurs above the antennal insertions; ASD Female. (Figs 280–283). 8.3 mm (GL350)–11.1 mm (GL 1400) long. Macropterous. Not reliably associated but the similarity of abdominal colour pattern in five females suggests they could belong to this species. Colour: Pronotum yellowish; elytra yellowish semitransparent with small black area at tip; head antennae palpi very dark almost black; venter of thorax orange yellow; legs orange yellow with dark brown tibiae and tarsi; basal abdominal ventrites yellow; V4, 5 with yellow margins and median dark brown markings which are wider across posterior margin and narrow anteriorly, with that of V4 narrower than V5; LO in V6 white occupying all of the visible area; V7, 8 pale yellow semitransparent, with underlying fat body visible through cuticle with most of posterior margin clear; basal abdominal tergites brown (usually visible through the semitransparent elytra), with T8 pale yellow and semitransparent (T7 and 8 usually protruding beyond the elytral apices). Abdomen: median posterior margin of V7 widely and shallowly emarginated, with rounded corners; V8 without median posterior emargination. Female specimens examined. All specimens are preserved in 70% ethanol. MALAYSIA. Perak: Gunung Liang 350 m a.s.l., 21.vi.2015, B. Nada, female (GL 350); 18.vi.2015, 1400 m a.s.l., B. Nada, female (GL 1400); 21.vi. 2015, 345 m a.s.l., B. Nada, female (GL 345) (FRIM). Negeri Sembilan: Gunung Besar Hantu, 9.vi. 2015, 350 m a.s.l., B. Nada, female (GBH 350) (FRIM). Terengganu: Pasir Raja Barat Forest Reserve, Compt. 87, 2.viii. 2016, B. Nada, female (PB 87 [4]) (FRIM). Etymology. The locality name is used as a noun in apposition with the generic name. Remarks. The two males described here are tentatively assigned to Luciola s. str. as they differ from the conventional description in certain aspects. The ML of the aedeagus is strongly curved but not inclined ventrally at its apex; the LL are not expanded at their apices and there are no elongated leaf like lobes on the inner ventral margin. Females are addressed separately and are not reliably associated, however their colour patterns, size, and especially the pattern on the ventral abdomen, strongly suggest this species., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 93-96, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098

Luciola melancholica Olivier 1913a Luciola melancholica Olivier 1913a: 417. McDermott 1966: 109. Ballantyne & McLean, 1970: 267. Ballantyne & Lambkin 2013: 122. Holotype. Female. NEW GUINEA: Iles Bertrand (MCSN). Remarks. The holotype female was described with dark brown elytra and an orange pronotum with dark marginal markings. Ballantyne & Lambkin (2013) and Ballantyne & McLean (1970) addressed various species using both males and females having marginal dark markings on the pronotum, but were unable to resolve its identity. No males from the type locality of the Bertrand Islands have been located., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 161, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Olivier, E. (1913 a) Lampyridae. Resultats de L'Expedition Scientifique Neerlandaise a la Nouvelle-Guinee en 1907 et 1909 sous les auspices de Dr H. R. Lorentz. Zoologie Livraison III. Nova Guinea, 9, 417 - 422.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A. & McLean, M. R. (1970) Revisional studies on the firefly genus Pteroptyx Olivier (Coleoptera: Lampyridae: Luciolinae: Luciolini). Transactions of the American Entomological Society, 96, 223 - 305.","Ballantyne, L. A. & Lambkin, C. L. (2013) Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa, 3653 (1), 1 - 162. https: // doi. org / 10.11646 / zootaxa. 3653.1.1"]}

Luciola klapperichi Pic 1955 Luciola Klapperichi Pic 1955: 26. Luciola klapperichi Pic. McDermott 1966: 107. Type. CHINA. Not located. Remarks. Pic described a species 7−8 mm long with orange pronotum and black elytra having reddish markings on the humeral angle. He felt it could be distinguished from other similarly coloured species by the convex sided (Ballantyne’s words) elytra. It may be synonymous with Luciola dilatata., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 158, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1955) Coleopteres nouveaux de Chine (Suite). Bulletin de la Societe Entomologique de Mulhouse, 1955, 21 - 23 + 25 - 26 + 29 - 32.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149."]}

Luciola satoi Jeng et Yang 2003 Luciola satoi Jeng et Yang 2003: 251. Fu 2014: 32. Type. Holotype male. TAIWAN. (NMNS). Diagnosis. A Taiwanese species with pinkish red pronotum having central apex dark brown and margins yellowish brown; elytra dark brown, margins yellowish brown (Jeng did not specify extent or width of the margins); posterior margin of V7 rounded; posterior margin of aedeagal sheath sternite two lobed; ML wide at base, narrowing gradually just posterior to the origins of the lobes on the LL; narrow apical portion is ¼ as wide as basal portion; anteromedian margin of sheath tergite broadly and narrowly produced and truncate. Jeng et al. (2003b: 251) distinguished this species from L. roseicollis Pic by its much smaller size, and from the similarly coloured L. kagiana Matsumura by the much larger size of kagiana. Inconsistencies with the identifications of various of these similarly coloured Luciola are discussed under the description of L. curtithorax above, and requires further investigation. Chen (2003: 169) figures a ‘ Luciola sp. 3’ which is L. satoi (Jeng pers. comm.)., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 100, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Jeng, M. L., Yang, P. S. & Lai, J. (2003 b) Notes on the genus Luciola (Coleoptera, Lampyridae, Luciolinae) of Taiwan. Special Bulletin of the Japanese Society of Coleopterolog y, 6, 247 - 262.","Fu, X. H. (2014) An illustrated handbook of Chinese fireflies. The Commercial Press, Beijing, 167 pp. [in Chinese]","Chen, T. R. (2003) The fireflies of Taiwan. Field Image Publisher, Taipei, 256 pp. [in Chinese]"]}

Luciola kagiana Matsumura 1928 Luciola kagiana Matsumura 1928: 39. Jeng et al. 2003b: 260. Chen 2003: 168. Fu 2014: 30. Yiu 2017: 98. Type. Female. TAIWAN (as Formosa). Chiayi (as Kagi). Hokkaido University Museum. Diagnosis. Most obviously distinguished from other similarly coloured Luciola by its large size and uniformly coloured pronotum; 10.5−15.0 mm long; pronotum pinkish red; MS yellowish brown, elytra black. Assigned to Luciola s. str. based on diagrams of aedeagus in Fu (2014) and Yiu (2017). Aedeagal sheath with anterior margin of sheath tergite not produced; posterior area of sternite produced and apically emarginated. Aedeagus slender, with ML swollen at base and narrowing in apical 1/5; narrow apical portion 1/6 s wide as wider basal portion; leaf like lobes slender elongate; LL closely approach along median dorsal line; apices of LL expanded. Possible larvae have laterally explanate tergal margins., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 96, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Matsumura, M. (1928) The Fireflies. In: Using interesting insects as teaching tool. Tokyodo Shoten, Tokyo, pp. 39 - 70. [in Japanese]","Jeng, M. L., Yang, P. S. & Lai, J. (2003 b) Notes on the genus Luciola (Coleoptera, Lampyridae, Luciolinae) of Taiwan. Special Bulletin of the Japanese Society of Coleopterolog y, 6, 247 - 262.","Chen, T. R. (2003) The fireflies of Taiwan. Field Image Publisher, Taipei, 256 pp. [in Chinese]","Fu, X. H. (2014) An illustrated handbook of Chinese fireflies. The Commercial Press, Beijing, 167 pp. [in Chinese]","Yiu, V. (2017) A Study of Rhagophthalmidae and Lampyridae in Hong Kong with descriptions of new species (Coleoptera): Part 2. Lampyrid, 4, 67 - 119."]}

Luciola aurantiaca Pic 1927 Luciola aurantiaca Pic 1927: 37. McDermott 1966: 99. Type. INDIA. Female. (MNHN). Remarks. The type female is dorsally pale yellowish and in colour is close to L. candezei and L. ochracea. The ventral colouration was not examined (the specimen is stuck ventral side down on a card). Nothing sufficiently distinctive about this colour pattern, nor the locality data, would permit an attempt to associate males., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 154, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Pic, M. (1927) Malacodermes exotiques. L'Echange, Revue Linneenne, XLIII, 37 - 52.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149."]}

Luciola antica (Boisduval 1835) Lampyris antica Boisduval 1835: 138. Luciola antica Boisduval. McDermott 1966: 99. Type. Locality unclear. Type not located. Remarks. This species was described from “des Îles de la mer du Sud”, with a yellowish dorsum and the pronotum having an anterior dark mark, the underside yellowish except for the black rings of the abdomen. The locality is unclear., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on page 153, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Boisduval, J. B. A. D. (1835) Voyage de decouvertes de l' Astrolabe, execute par ordre du Roi, pendant les annees 1826 - 1829, sous le commandement de M. J. Dumont- D'Urville. Faune entomologique de l'ocean Pacifique avec l'illustration des insecte nouveaux recueillis pendant le voyage. Deuxieme partie. Coleopteres et autres ordres. Tastu, Paris, 716 pp.","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149."]}

Luciola s. lato sensu Ballantyne et al. 2015 As a heterogeneous assemblage of species, Luciola s. lato cannot be distinguished further here and its subdivision is being progressively addressed. Species still listed here are those assigned to Luciola in McDermott (1966) for which we currently cannot assign a genus. We believe they are sufficiently distinctive that the real possibility still exists for reliable association of males and determination of a generic placement. Many have types although in rather poor condition. All are in MNHN. McDermott’s (1966) catalogue of the Luciolinae distinguished several genera by male abdominal distinctiveness (Colophotia, Pteroptyx, Pygoluciola and Pyrophanes), and one genus by the very distinctive elytral punctation (Curtos). A subgenus of Luciola distinguished by a pronounced eye emargination in the males (Bourgeoisia) has since been submerged under Luciola s. str. Presence of flightless females distinguished Luciola s.g. Hotaria and Lampyroidea. However for various reasons, not entirely unrelated to the same problems we have encountered here, 279 species were assigned to Luciola s.g. Luciola. This arrangement in McDermott (1966) of 279 species under Luciola Laporte was the largest stumbling block to identification and understanding of the fauna of this area. However at least 100 species of African Luciolinae assigned in McDermott to Luciola are not investigated. Ballantyne et al. (2015) outlined their approach to Luciolinae taxonomy and the problems posed with this heterogeneous assortment of species. They outlined the historical nature of their approach, which has been to develop a working taxonomic framework for the Luciolinae while attempting to identify all the species listed under Luciola in McDermott (1966). Their success thus far is exemplified in the results presented here. They listed in Appendix 3 96 species from the defined study area which still stood under Luciola s.g. Luciola. Of the 45 species for which types could be located, or which were not based on females, they suggested possible placements for 24, many of which have been effected here. We list here 34 species still standing under Luciola s. lato. Of these eight have female types, only one of which, L. obscura, has sufficiently distinctive colouration that assignment of similarly coloured males might be possible. We have not located types for twelve species. Of the others where types can be located the somewhat fragile nature of the types has precluded any attempt thus far at dissection. Dissections must address the terminal abdomen for generic characters, and this is often the first part of the specimen to become dislodged. Ballantyne et al. (2016) have short diagnoses and pictures of many species (see Table 18), and preliminary comments have been made already, or can be made about many of the species included below. Luciola auritula although known only from a female has distinctive brown lateral markings on an otherwise yellowish pronotum. Luciola clara and L. curticollis are probably the same species (Ballantyne et al. 2016 figs 47–50). Although both known only from females L. candezei and L. ochracea are very distinctively coloured large species, and are probably the same (Ballantyne et al. 2016 figs 152–157). Both L. davidis and L. maculipennis may be species of Pygoluciola. Luciola nicollieri may be represented by specimens illustrated by Fu (2014: 47), and Yiu (2017: 98). In the absence of a type of laticollis and a definitive identification we retain laticollis here (see Yiu 2017: 94). ......continued on the next page Species PeninsulaMalay- Locationoftypes Myanmar Vietnam Cambodia Indochina SriLanka Indonesia Ambonisland Philippines Females Referencesfor illustrationsof * Borneo China Celebes India Japan sia Krea Larvae species roseicollis Pic 1933 * MNHN rugiceps Olivier 1886 * sordida Olivier 1905 * MNHN sudra Gorham 1903 * * NHML tenuicornis Olivier 1885 * MNHN 1 tincticollis Gorham 1895 * * unmunsana Doi 1931 * = tsushimana Nakane 1970 CTN = papariensis Doi 1932 * * varia Olivier 1908 * MNHN xanthura Gorham 1880 * IRNSB Luciola picea and L. pallidipes (which is described here) bear similar dorsal colouration and are a similar size. Luciola xanthura has similar aedeagal pattern to that illustrated by Fu (2014: 43, 52) and is very similar to L. sudra. Preliminary indications are that L. varia should be assigned to Sclerotia Ballantyne. Others are sufficiently distinctive either in dorsal colouration or in size, that it should be possible to obtain fresher specimens to allow dissection that has not been attempted on many of the fragile types. These species include L. gigas, L. obscura and L. limbatipennis., Published as part of Ho, - Z., 2019, The Luciolinae of S. E. Asia and the Australopacific region: a revisionary checklist (Coleoptera: Lampyridae) including description of three new genera and 13 new species, pp. 1-174 in Zootaxa 4687 (1) on pages 102-105, DOI: 10.11646/zootaxa.4687.1.1, http://zenodo.org/record/3508098, {"references":["Ballantyne, L., Lambkin, C. L., Boontop, Y. & Jusoh, W. F. A. (2015) Revisional studies on the Luciolinae (Coleoptera: Lampyridae): 1. The firefly genus Pyrophanes Olivier with two new species. 2. Four new species of Pteroptyx Olivier. 3. Inflata gen. nov. Boontop, with redescription of Luciola indica (Motsch.) as Inflata indica comb. nov. Zootaxa, 3959 (1), 1 - 84. https: // doi. org / 10.11646 / zootaxa. 3959.1.1","McDermott, F. A. (1966) Lampyridae. In: Steel, W. O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S'Gravenhage, pp. 1 - 149.","Ballantyne, L. A & Jusoh, W. F. A. (2016) List of genera and species in the Luciolinae from SE Asia and the Australopacific Version 2. Checklist of IndoPacific Luciolinae, 2016, 1 - 7. https: // doi. org / 10.13140 / RG. 2.1.4635.2241","Fu, X. H. (2014) An illustrated handbook of Chinese fireflies. The Commercial Press, Beijing, 167 pp. [in Chinese]","Yiu, V. (2017) A Study of Rhagophthalmidae and Lampyridae in Hong Kong with descriptions of new species (Coleoptera): Part 2. Lampyrid, 4, 67 - 119.","Olivier, E. (1910 b) XX. Descriptions of two new species of Luciola in the collection of Mr H. E. Andrewes. Transactions of the Entomological Society of London, 1910, 532 - 533. https: // doi. org / 10.1111 / j. 1365 - 2311.1910. tb 01183. x","Olivier, E. (1902) Catalogue des especes de ' Luciola' et genres voisins decrits jusqu'a ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69 - 88.","Olivier, E. (1907) Coleoptera, Lampyridae. Genera Insectorum, 53, 1 - 74. [Luciolinae, 48 - 56].","Pic, M. (1927) Malacodermes exotiques. L'Echange, Revue Linneenne, XLIII, 37 - 52.","Olivier, E. (1895) Deux especes nouvelles de Lucioles. Bulletin de la Societe Entomologique de France, 1895, 148 - 149.","Olivier, E. (1909 a) Descriptions de Lampyrides nouveaux du Musee Civique de Genes. Annali del Museo Civico di Storia Naturale di Genova, IV (3 a), 316 - 319.","Olivier, E. (1888) Etudes sur les Lampyrides IV. Descriptions d'especes nouvelles. Annales de la Societe Entomologique de France, 1888, 47 - 62.","Gorham, H. S. (1883) Review of the genera and species of Malacoderm Coleoptera of the Japanese fauna. Transactions of the Entomological Society of London, 1883, 393 - 411. https: // doi. org / 10.1111 / j. 1365 - 2311.1883. tb 02955. x","Pic, M. (1923) Deuxieme addenda. Faune entomologique de l'Indo-Chine Francaise, 6, 58 - 63.","Olivier, E. (1913 b) The Lampyridae of Borneo. Sarawak Museum Journal, 3, 55 - 60.","Bugnion, E. (1922) Luciola nicollieri sp. n. de Ceylan. Spolia Zeylandica, XII, 1 - 14.","Pic, M. (1914) Nouveau genre, especes et varietes nouvelles de diverses familles. Melanges exotico-Entomologiques, 10, 3.","Pic, M. (1928) Malacodermes exptiques. L'Echange, Revue Linneenne, XLIV, 58 - 63.","Gorham, H. S. (1895) List of the Coleoptera in the collection of H. E. Andrewes Esq. from India and Burma, with descriptions of new species and notes. Annales de la Societe Entomologique de Belgique, XXXIX, 293 - 307.","Pic, M. (1939) Notes diverses, nouveautes. L'Echange, Revue Linneenne, 55, 17 - 19 + 21 - 22 + 25 - 26 + 370.","Gorham, H. S. (1882) New species of Lycidae, Lampyridae and Telephoridae from Sumatra. Notes of the Leyden Museum, 4, 93 - 109.","Olivier, E. (1900) Contribution a l'etude de la faune entomologique de Sumatra. XI. Lampyrides determines et decrits. Societe royale d'entomologie de Belgique Annales, 44, 234 - 238.","Pic, M. (1933) Notes diverses, descriptions et diagnoses. L'Echange, Revue Linneenne, XLIX, 109.","Olivier, E. (1886) In: Baer, G. - A., Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France, 1886, 183 - 187.","Olivier, E. (1905) Descriptions de Lampyrides nouveaux. Annales de la Societe Entomologique de Belgique, XLIX, 206 - 209.","Gorham, H. S. (1903) On Coleoptera collected in India by MM. H. E. and H. L. Andrewes. Annales de la Societe Entomologique de Belgique, 47, 323 - 330.","Olivier, E. (1885) Catalogue des Lampyrides faisant partie des collections du Musee Civique de Genes. Annali del Museo Civico di Storia Naturale di Genova, 2 (2 a), 333 - 374.","Doi, H. (1931) A new species of Luciola from Korea. Journal Chosen Natural History Society, 12, 54 - 55. [in Japanese with English subtitle]","Nakane, T. (1970) On the cantharoid Coleoptera found in the Islands of Tsushima (Insecta). Memoirs of the Natural Science Museum, Tokyo, 3, 287 - 288. [in Japanese with English descriptions and summary]","Doi, H. (1932) A new species of Luciola from Korea. Journal Chosen Natural History Society, 14, 64. [in Japanese with English subtitle]","Olivier, E. (1908) Neue Lampyriden des Deutschen Entomol. National-Museums. (Col.) Deutsche Entomologische Zeitschrift, 1908, 491 - 495.","Gorham, H. S. (1880) Materials for a revision of the Lampyridae. Transactions of the Entomological Society of London, 1880, 83 - 112."]}