Search

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."]}

Nipponoluciola owadai (Satô & Kimura, 1994) gen. et comb. nov. Figs 1–2, 5, 8–9 Luciola owadai Satô & Kimura, 1994: 159. Luciola owadai – Ohba et al. 1994: 13–26. — Ohba 1996: 17; 2004b: 10; 2005: 240. — Suzuki 1997: 19. — Jeng et al. 2003: 543, 550–551, 553–556. — Kawashima et al. 2003: 248. — Suzuki et al. 2004: 297. — Geisthardt & Satô 2007: 232. Diagnosis Distinguished from Nipponoluciola cruciata gen. et comb. nov. by the yellowish orange pronotum, (where the underlying fat body is yellowish), MS and MN (the pronotum of N. cruciata has underlying pink fat body and a median dark marking in most specimens, and the MS is black) (Figs 1–2). Material examined JAPAN ‒ Kume-jima Is., Okinawa Isls, Middle Ryukyus • 1 ♀; Mt Uegusuku-dake; 25 Apr. 1993; I. Kawashima leg.; CIK • 4 ♂♂, 1 ♀; Ôta-gawa Riv.; 25 Apr. 1993; I. Kawashima leg.; CIK • 2 larvae; Shirase-gawa Riv.; 27 Dec. 1993; Y. Goto leg.; CIK • 20 ♂♂, 8 ♀♀ (paratypes); same collection data as for preceding; 26 Apr. 1994; Y. Goto leg.; YCM • 7 ♂♂, 1 ♀ (paratypes); Shimajiri; 28 Apr. 1994; Y. Goto leg.; YCM. Notes Locality labels on the type specimen were incorrectly quoted in Satô & Kimura (1994) and are repeated here in their correct form (Fig. 2H) as follows (from top to bottom, a slash indicates a line break): Kumejima-Uegusuku-dake, Minami (in Japanese “Kanji” characters) / Japan, Ryukyus / Kumejima Is. / Mt. Uegusuku (south) / 90 m. 27. IV. 1993 / Mamoru Owada leg. (original white label, printed in black ink); HOLOTYPE / Luciola owadai / M. Satô et Kimura, 1994 / DET. M. SATÔ 1994 (red holotype label, printed in black ink). Description Male (Figs 1–2) BODY LENGTH. 9.5–15.8 mm. COLOUR (Figs 1–2). All of body including head, mouthparts and antennae, ventral thorax, legs and abdomen, black, except for yellowish orange pronotum (with underlying yellowish fat body), having no median darker markings, yellowish MN and MS, and white LO in V6, 7. Pronotal width/GHW 1.39–1.46. GHW/SIW: 4–4.8. ABDOMEN. As depicted in Jeng et al. (2003: fig. 8), with a short broad apically rounded MPP. AEDEAGAL SHEATH (Fig. 5). Sheath sternite widely emarginated on right side, and scarcely so on left (see also Jeng et al. 2003: fig. 28). AEDEAGUS (Fig. 5). BP well sclerotised; median anterior margin of BP with deep and well-defined notch (see also Jeng et al. 2003: fig. 34); lateral margins of ML subparallel-sided in basal 7/₁₀, gradually narrowing in apical 3/₁₀. LL/ML 2.81; LL apex width considerably wider than width of apex of ML; dorsal base of LL symmetrical; apex of ML depressed on both sides; the outline of apex rounded when viewed from either side. AEDEAGAL PATTERNS. Since the distribution of this species is limited to the small island Kume-jima Is., there is little individual variation in the shape of the aedeagus. Female (Fig. 7B) Macropterous and flight capable. BODY LENGTH. 13.1–15.7 mm. COLOUR (Fig. 7B). Colouration as in male except for whitish LO in V6 (Satô & Kimura 1994 describe this as the 5 th abdominal segment) occupying all but a narrow transparent posterior margin; V7 mostly black, not transparent as that of female of N. cruciata, and no underlying fat bodies visible; basal portion of V7 may have irregular and indistinct pale patches where the blackish pigmentation appears to be lost; V8 black; dorsal surface of all tergites (up to T8) and dorsally reflexed margins of ventrites (to V5, and V7, V8) dark brown to black; T6 black with outer margins of dorsally reflexed parts of V6 white and semitransparent. PRONOTUM. Pronotal width/GHW 1.74–1.86. HEAD. GHW/SIW 2.78–2.90. ABDOMEN. Posterior margin of V7 shallowly and narrowly emarginated as in Jeng et al. (2003: fig. 16). Lateral margins of V8 converge posteriorly; with the median posterior margin very shallowly and narrowly emarginated. Jeng et al. (2003: fig. 16) depict the posterior margin of V7 with a shallow and narrow emargination. REPRODUCTIVE SYSTEM. Well defined thin walled SDG; MOP trapezoidal, filling half of the median oviduct, with the anterior margin almost straight; lateral margins straight and convergent toward the posterior margin; posterior margin nearly straight but slightly irregular. Ovipositor elongate slender. Larva (Figs 8B, 9E‒H) Approx. body length 13 mm (specimens measured were severely shrunken due to ethanol fixation); maximum (median) length of protergum 2.7 mm; maximum width of protergum 2.3 mm. Penultimate instar larvae were examined. The larvae of this species closely resemble those of N. cruciata, and are not clearly distinguishable from each other in general appearance and morphological characteristics. However, the ground colour of the membranous body of N. owadai is much paler (Ohba et al. 1994), light to greyish brown, while that of N. cruciata is dark to blackish brown, and it differs from N. cruciata in the tendency of the pale markings on the lateral margins of pronotum to separate anteriorly and posteriorly into four independent spots. Ohba et al. (1994) recorded 12 abdominal pygopodia in what was possibly the last instar larva. Jeng et al. (2003) considered the basal row of pygopods was composed of four retractable filaments. Flash communication The flash pattern of both sexes of this species is very similar to that of N. cruciata, and the communication system is the same as that of N. cruciata. Synchronous flashing of the males is similar to that of east Japan populations of N. cruciata, but the flash interval gets shorter over time, from 4 sec to 3 and 2 sec. However, unlike the west Japan populations of N. cruciata, the synchronous flashing does not last for a long time but continues intermittently through the night. Thus, it has characteristics of both east and west Japan N. cruciata. Spawning behaviour Females gather and lay eggs on the moss near water as in the western Japanese form of N. cruciata. However, they gather all at once just before dawn, not one after another in the night. Remarks This species is designated both a natural monument of Okinawa Prefecture and an endangered species of wild fauna and flora under the Law for the Protection of Cultural Properties and the Act on Conservation of Endangered Species of Wild Fauna and Flora, respectively. It is now difficult if not impossible to obtain specimens of this endangered species in the locality, Kume-jima Is. The few specimens examined by IK are listed above. All other remarks about this species are referenced. Ohba et al. (1994) addressed life history, behaviour and morphology. Ohba (2004b) depicted various aspects of morphology of this species including flashing patterns (page 7), dorsal aspect of female (page 11) and male (pages 38, 45), SEMs of anterior head (page 62) and a species overview (in Japanese, page 109). Jeng et al. (2003) keyed aquatic Taiwanese and Japanese Luciolinae males.

Nipponoluciola gen. nov. urn:lsid:zoobank.org:act: BD75E8CA-BC67-4597-A0A5-6BCEF5BA7B4F Figs 1‒3, 5‒12 Type species Luciola cruciata Motschulsky, 1854, designated here. Diagnosis Nipponoluciola gen. nov. is a genus of Japanese Luciolinae fireflies and consists of two species both of which have aquatic larvae with abdominal gills. While Nipponoluciola belongs to that group of 18 genera of Luciolinae where the aedeagus in the male has the LL widely visible at the side of the ML (Ballantyne et al. 2019), it is one of only two genera in this group of 18 (the other is Missimia) where the elytral epipleuron at its base is very narrow and when viewed from beneath in a horizontal specimen the humerus is widely visible in both males and females (Fig. 3A–C). In the other genera of Luciolinae with LL visible at the sides of the ML the base of the elytral epipleuron is expanded, and the humerus is not visible from below in a horizontal specimen (example in Fig. 3D). It is distinguished from Missimia by distribution (Missimia is a rare genus known only from four specimens taken in the New Guinean highlands), and the unique feature of Missimia, where the labrum and clypeus are immovably joined (Fig. 4E; Table 1; Ballantyne & Lambkin 2009: figs 27, 29–30, 32). The clypeolabral suture in Nipponoluciola is flexible. It is distinguished from the genus Aquatica, with which it shares similar aquatic larvae bearing abdominal gills, by the following features: Aquatica species adult males may have pale margined elytra (Fig. 4A), the elytral epipleuron at its base is wide (Fig. 3D), the LO in V7 is usually retracted from the posterior margin (Fig. 4B), the lateral margins of V7 are not angulated, both left and right margins of the posterior area of the sheath sternite have pointed projections, the aedeagal basal piece is not notched (Fig. 4C–D). Females have thin paired bursa plates (Fu et al. 2010). See Table 2. Males have black elytra and the pronotum may have median dark brownish to blackish markings and an underlying pink fat body (in N. cruciata) (Figs 1, 2G, 6), or have no dark markings and underlying yellowish orange fat body (in N. owadai) (Figs 1, 2H). Females are coloured like males, and are larger, macropterous, with spermatophore digesting gland, no obvious bursa plates and a small transverse median oviduct plate in N. cruciata (Fig. 7). Larvae (Figs 8–9) are aquatic, have forked gills along the sides of the abdominal segments 1–8 and apical sense organs on maxillary and labial palpomeres. Distinguished most obviously from Aquatica larvae by the differences in the markings of protergum (outlined subsequently); distinguished from A. lateralis by the unbranched setae on the dorsal surface of the mandibles, and the absence of microspines on pale areas of paired tergites of meso- and metathoraces, and abdominal segments 1–8 (Okada 1928: figs 2‒3, 2‒4 depicted but did not name these microspines; IK confirmed their presence in A. lateralis). Etymology The generic name seeks to honour the country of origin of these two species and their status as celebrated icons. ‘Nippon’ is the Japanese name for Japan, and Luciola represents the genus to which these species were formerly assigned. Gender: feminine. Description Male (Figs 1–3, 5–6, 10‒12) PRONOTUM (Figs 1–2). Punctation dense; anterior margin broadly rounded and projecting beyond rounded obtuse anterolateral corners, slightly so in owadai; lateral margins slightly divergent, the pronotum being wider across its posterior area than elsewhere (C>A, B), width (C) subequal to humeral width; posterolateral corners narrowed, apically rounded, slightly less than 90°, projecting posteriorly slightly beyond median posterior margin, which is not emarginated, and separated from it by welldefined emarginations. Pronotal colour variations depicted in Fig. 6 are discussed under Nipponoluciola cruciata gen. et comb. nov. HYPOMERA. Not well flattened; pronotal width/GHW 1.43–1.47. ELYTRON (Figs 1–3). Punctation dense, not linear, not as large as that of pronotum, nor widely and evenly spaced; apices not deflexed, nor emarginated; epipleuron and sutural ridge extend beyond mid-point, almost to apex but not extending around apex, neither thickened in apical half; with three interstitial lines of which the inner two interstitial lines are well defined; in horizontal specimen viewed from below epipleuron at elytral base narrow with humerus visible; viewed from above in horizontal specimen anterior margin of epipleuron arises well behind the posterior margin of MS (Fig. 3A, C); epipleuron developed as a lateral ridge along most of length; sutural margins approximate along most of length in closed elytra; lateral margins parallel-sided. HEAD (Figs 1–2). Gently depressed between eyes; at least anterior portion exposed in front of pronotum, not capable of complete retraction within prothoracic cavity at rest; eyes moderately separated beneath at level of posterior margin of mouthpart complex; frons-vertex junction not defined this area rounded, posterolateral eye excavation not developed; antennal sockets on head between eyes, close but not contiguous, ASD subequal to ASW; clypeolabral suture present, flexible, not in front of anterior eye margin when head viewed with labrum horizontal; outer edges of labrum reach inner edges of closed mandibles. Mouthparts: functional; apical labial palpomere flattened, shaped like a broad triangle, longer than wide, with inner edge dentate. Antennae (Fig. 1) 11 segmented; approximately 2× GHW; FS simple elongate; GHW/SIW 3.70–4.50. LEGS (Figs 1–2). Without MFC; no legs with curved or expanded femora or tibiae; no basitarsi expanded or excavated. ABDOMEN (Fig. 2F). No ventrites with curved posterior margins nor extending anteriorly into emarginated posterior margin of anterior segment. LO in V7 entire, LO present in V6, occupying almost all V6 (Jeng et al. 2003: figs 7, 9). V7 with elongate apically rounded MPP and angulate lateral areas which are not prolonged and correspond to where PLP would originate in other species. T7 without prolonged anterolateral corners; T8 not heavily sclerotised; no concealed anterolateral arms of T8 and the anterolateral corners scarcely produced (see Jeng et al. 2003: figs 21–22); lateral margins subparallel sided; posterior margin evenly rounded and entire; T8 ventral surface smooth even without any ridges or projections. AEDEAGAL SHEATH (Fig. 5). Approx. 1.94–2.12 times as long as wide; without bulbous paraprocts; symmetrical in anterior half where sheath sternite tapers evenly to a broadly rounded apex; lateral arms of sheath tergite widely visible from below on left, narrowly on right; posterior area of sternite (posterior to the articulation of the tergite arms) narrow, emarginated on both sides (depending on orientation it may appear to be more deeply emarginated on right side) with margin even, without projections, and sternite apex rounded; tergite broad with anterior margin slightly and evenly emarginated; no obvious subdivision into more than one segment observed; posterior margin entire and evenly rounded; tergite projecting about as far as apex of sternite. AEDEAGUS (Figs 5, 10–12). Subparallel sided; basal piece well defined and usually with distinctive, asymmetrical median notch; LL widely visible beside ML; LL of similar length, slightly shorter than ML; width across LL, (measured from beneath), much wider than width across widest part of median lobe (3‒3.3); ML slender gradually narrowing in apical 1/7; apices of LL with ventral surfaces broad, slightly curved and hairy. Differences in aedeagal patterns in Nipponoluciola cruciata gen. et comb. nov. are addressed under that species in Figs 10‒12. Female (Fig. 7) Macropterous and flight capable. BODY LENGTH. 14.0– 18.4 mm (Jeng et al. (2003) listed a range of 15.0– 18.6 mm long). COLOUR (Fig. 7A, C–D). As for male except white LO restricted to V6, pinkish fat bodies beneath V 7 in cruciata, or V7 completely black and underlying fat body not visible in owadai; black V8 with paired semitransparent areas; all tergites black except for T7 which has a median dark band and semitransparent lateral areas under which pink fat bodies are visible in cruciata. PRONOTUM (Fig. 7). Outline as for male; pronotal width/GHW 1.88–2.20. HEAD. GHW/SIW 2.67–2.90. ABDOMEN (Fig. 7C–D, I). Lateral margins of V8 taper posteriorly, V8 with shallow median posterior emargination; anterior apodeme of V8 well sclerotised and separated from posterior area of V8 by clear transparent cuticle (Fig. 7I). REPRODUCTIVE SYSTEM (Fig. 7E‒H, J). Spermatophore digesting gland well defined usually with particulate remnants inside; spermatheca with expanded non sclerotised base; no obvious bursa plates; median oviduct plate well defined (Fig. 7G–H), anterior margin curved and irregular, lateral margins straight and converging slightly posteriorly, posterior margin slightly and broadly emarginated; lying transversely in the median oviduct and curved possibly to accommodate to the outline of the duct. Ovipositor: elongate slender (Fig. 7J). Larva (Figs 8A–B, 9A‒H) (see also Fu et al. 2012b: figs 81–82) DORSAL SURFACE (Figs 8–9). With 3 thoracic and 9 obvious abdominal segments of which abdominal segments 1–8 bear lateral tracheal gills which are forked and compound, with a non-functional spiracle located at the end of the shorter branch. A narrow ring of cuticle at the posterior end of segment 9 has been attributed to segment 10 (Ballantyne & Menayah 2002: figs 2–3; Nada et al. 2021) and may be completely covered and concealed by the posterior margin of segment 9 from above in aquatic species, but visible from beneath. Lawrence & Ślipiński (2013) consider that segment 10 has been transformed into the holdfast organ (pygopodia) at the end of the abdomen. We refer to abdominal terga 1‒9 only as these are clearly visible. Elongate, tapering at front and behind, membranous and very soft bodied, and somewhat flattened. The thorax and abdomen are almost membranous, with ventral sclerites degenerate and extent barely demarcated by wrinkles or folds, except for a single coloured dorsal plate on both the prothorax and abdominal segment 9, and paired well sclerotised and coloured tergites on the meso and metathorax, and abdominal segments 1–8. A wide membranous area/line extends from the anterior margin of the mesothorax to the posterior margin of abdominal segment 8. Without laterally explanate tergal margins thus laterotergites and gills visible from above. VENTRAL SURFACE (Fig. 9B, D, F). The extent of the following areas in very soft bodied aquatic larvae is often difficult to determine. A lateral pleural suture delimits laterotergites above in the thorax and abdomen (e.g., Ballantyne & Menayah 2002: fig. 1; Fu et al. 2012b: figs 81–82). In the thorax the median ventral surface of both meso and metathorax is subdivided into two areas by a sternocostal suture running anterior to the coxae, an anterior basisternum with laterotergites at the sides in the mesothorax; (in terrestrial larvae these mesothoracic laterotergites bear functional spiracles, but spiracles were not clearly visible in the larvae examined); a posterior median subrectangular sternellum bears the legs and is margined laterally by laterotergites; the episterna and epimera of both the meso and metathorax are visible above the coxae of the meso and metathoracic legs as thin dark sclerotised plates (see interpretation in Fu et al. 2012b: 5). This subdivision does not extend to the dorsal surface. In the abdominal segments 1–8 a median subrectangular area is attributed to the sternum; this is margined laterally by elongate narrow paired laterosternites, which are delimited by folds from the median sternal plate below, and the laterotergites above; in this aquatic genus the absence of any additional colour on the ventral surface makes interpretation of these areas difficult, and they may be incompletely distinguished only by folds in the membrane; laterotergites of segments 1–8 bear branched gills. Eversible branched defensive organs arise at the sides of the meso and metathorax, and abdominal segments 1–8, in the membrane at the sides of the terga of those segments and above the laterotergites of the abdomen (see below). HEAD (Fig. 8C) (Kanda 1935: fig. 13; Hara 1962: figs 1, 5; Fu et al. 2012b: figs 25–30, 69). Subparallel sided, dorsoventrally flattened, prognathous, well sclerotised, with an ocellus at each side; not visible when retracted into the prothoracic cavity; extensible neck membrane forming two layered envelope around retracted head; capable of considerable extension beyond the anterior protergal margin; head capsule divisible into median dorsal frontoclypeus, bounded laterally by the U shaped frontal arms of the ecdysial line (= epicranial suture); lateral parietal plates at the sides of the frontoclypeus are separated behind by the epicranial stem and are reflexed ventrally but not meeting; maxillae and labium fused forming a maxillolabial complex covering most of ventral head area. ANTENNAE (Fu et al. 2012b: figs 27–29). Slender, cylindrical, 3 segmented, with elongated scape and pedicel, and apical very short (3 rd) segment (the flagellum), with apical sensilla, subequal in length to adjacent sense cone; elongate ‘articulating membrane’ forms two layered envelope around retracted antenna. MOUTHPARTS (Fu et al. 2012b: figs 25, 29, 69). Well developed; mandibles falcate, strongly sclerotised; densely covered in fine hair along outer margins; densely pubescent along basal half of inner margins; perforated along length by a fine canal that opens on the outer margin just behind the apex; without a basal retinaculum; fine reticulate sculpture on the dorsal surface; with a row of elongate curved setae arising in a line just inside the outer margin. Maxillae with short, squat, four segmented palp bearing sense organs at the apex of segment 4; basal segment (which may be interpreted as the palpifer) large and well defined, segments 2 and 3 very short and diminishing in width towards apex, apical segment longer and narrower than palpomere 3, with apical sense organs; palp may obscure galea, which is long, thin, two segmented, bearing long and short setae at apex, and on its inner margin an elongate, flattened, dense profusion of anteromedially directed hairs (lacinia); cardo well defined, articulating with broad elongate stipes; fused along its outer edges with the reflexed head margins, and along its inner margins with the median labium. Labium with two segmented labial palpi, bearing sense organs at the apex of segment 2, and arising at anterolateral corners of small prementum which lacks a ligula; apical palpomere elongate conical with two or three peg-like sensilla on the sides; anterior area of prementum densely covered by minute spines on both sides of dorsal surface; postmentum elongate, not well sclerotised and colourless, and joined along sides by membrane to the cardines. THORAX (Figs 8–9) (Fu et al. 2012b: figs 81–82). Prothorax longer than wide, and containing retracted head within; with ventral surface little differentiated. Protergum elongate or elliptical, with lateral margins arcuate and shallowly expanded laterad, divergent posteriorly; anterior and posterior margins rounded and widely arcuate, connecting directly to lateral margins without forming corners. Each tergite of meso- and metathorax as are those of abdominal segments 1–8 separated from each other as a pair of plates, subequal in size; the shape more or less round pentagonal; any corners more or less rounded; the margins lightly pale margined, broadened at the lateral margins to form pale spots; lateral margins arcuate, expanded to laterad; hind margins also arcuate; inner margins straight and parallel-sided to each other. Legs are four segmented – short cylindrical coxae with bases widely separated; elongate trochanters joining femora obliquely; tibiotarsus terminated by a single claw. In contrast with previous treatments the leg segments are attributed to coxa, trochanter, femur and tibiotarsus. ABDOMEN (Figs 8–9) (Fu et al. 2012b: figs 81–82). Each tergite of abdominal segments 1–8, as are those of the meso- and metathorax, is separated as a pair of plates, similar form to those of meso- and metathoracic terga, more or less round pentagonal, diminishing slightly in both length and width toward posterior ones, not forming any corners; the margins lightly pale margined, broadened at the lateral margins to form pale spots; segment 9 has a single round tergal plate, with the margin also lightly pale margined as in segments 1–8; pale posterior margin often widens slightly to form a narrow pale spot; abdominal tergal width decreasing a little towards posterior end, ventral surface of all but terminal abdominal segment (segment 9) with paired laterotergites bearing gills at the sides (visible along sides of body as lateral projections); median sternal element in each of abdominal segments 1–8 margined by elongate slender laterosternites but these areas difficult to discern in soft bodied larvae; (laterosternites appear membranous on segment 8 which houses the light organs). DEFENSIVE ORGANS. With 10 pairs of similar pale white, forked eversible defensive organs (Okada 1928: fig. 1a; Kanda 1935: 57–58, pl. 5-5; Trice et al. 2004: 2; Fu et al. 2009: fig. 4; 2012b: figs 7–12) located laterally on the meso- and metathorax, and above the tracheal gills on each of the 8 abdominal segments; these organs arise in the thorax in folds of membrane on the upper side of the laterotergites and just below the lateral margins of the tergal plates, and in the abdomen above the abdominal laterotergites and below the lateral margins of the tergal plates; organs have protuberances on their external surface, each composed of a well-developed globular body attached to a long thick stalk; globular bodies consist of secretory cells. PYGOPODIA (Fig. 8, partly visible in Fig. 9A). On the external surfaces of the tubular larval eversible holdfast organs, numerous hook-shaped protuberances grow in circular rows on each tube. List of species of Nipponoluciola gen. nov. Nipponoluciola cruciata (Motschulsky, 1854) gen. et comb. nov. Nipponoluciola owadai (Satô & Kimura, 1994) gen. et comb. nov. Key to species of Nipponoluciola gen. nov. using males 1. Widely distributed in Japan; pronotum usually with pinkish fat body visible through cuticle, and variable median blackish marking which may be expanded in the median area to assume a cross like formation; MS and elytra black (Figs 1–2).................................................................................................................. N. cruciata (Motschulsky, 1854) (Honshu, Shikoku, Kyushu and their sub-islands) – A rare and protected species known only from the Okinawa Islands group; pronotum yellowish orange, without m

Nipponoluciola cruciata (Motschulsky, 1854) gen. et comb. nov. Figs 1‒3, 5‒12 Luciola cruciata Motschulsky, 1854: 53. Type specimen represented by label only, ZMMU. Luciola picticollis Kiesenwetter, 1874: 262. Series of syntype males. NHMUK. Syn. nov. Luciola cruciata towadensis Nakane, 1987: 173. Luciola cruciata PARTIM – Lacordaire 1857: 338. — Heyden 1879: 350. — Gorham 1880: 102. — Olivier 1902a: 77; 1902b: 188; 1907: 51; 1910: 41. — McDermott 1962: 24; 1966: 102. — Takakura 1977: 7. — Jeng et al. 2003: 541. — Kawashima et al. 2003: 247. Luciola cruciata – Motschulsky 1866: 167. — von Harold 1877: 357. — Lewis 1879: 17. — Matsumura 1918: 86, 87. — Okada 1928: 102; 1931: 134, 146. — Kanda 1935:31. — Kishida 1936: 12, 20, 21. — Hasama 1942a: 366; 1942b: 378; 1943: 23; 1944: 155. — Nakane et al. 1959: 170. — Nakane 1960: 36, 118; 1968: 5; 1970: 285; 1983: 111; 1987: 173. — Minami 1961: 21‒106, 152‒238. — Bertrand 1972: 599; 1973: 107. — Satô 1974: 133; 1978: 17; 1985: 121; 1989: 352. — Ohba 1978: 25; 1983: 24; 1984: 23; 2001: 45; 2004a: 228; 2005: 240; 2012: 13. — Yuma 1981: 57. — Ohba et al. 1994: 13. — Suzuki et al. 1996a: 191; 1996b: 682; 2004: 297. — Suzuki 1997: 1; 2001: 99. — Sawada 2000: 93. — Branham & Wenzel 2003: figs 1–2, 4–5, 8. — Takeda et al. 2006: 177. — Ballantyne & Lambkin 2009: 21; 2013: 9. — Oba et al. 2011: 771. — Ballantyne et al. 2015: 8; 2016: 204; 2019: 163. — Fu & Ballantyne 2009: 243. — Fu et al. 2009: 155; 2010: 3; 2012a: 6; 2012b: 14. — Jusoh et al. 2018: 14; 2021: 1. — Kato et al. 2020: 1. Luciola picticollis – Von Harold 1877: 357. — Lewis 1879: 17. — Olivier 1902a: 85; 1902b: 189; 1907: 54; 1910:45. — Matsumura 1918: 87. — Okada 1931: 130. — McDermott 1966: 111 (partim). — Nakane 1983: 111. Luciola vitticollis – Sensu Gorham 1883: 409. — Okada 1931: 146. Luciola cruciata vitticollis – Okamoto 1924: 182–183, 226 (incorrect record from Korea). — Matsumura 1928: 59 (misidentification). Luciola cruciata towadensis – Kawashima et al. 2003: 247 (synonymy). Luciola cruciata cruciata – Geisthardt & Satô 2007: 232. non Luciola picticollis – Gorham 1883: 409 (= lateralis). non Luciola vitticollis – Sensu Gorham 1883: 409. — Okada 1931: 146 (= lateralis). non Luciola cruciata var. vitticollis – Olivier 1902a: 71; 1902b: 188; 1907: 54; 1910: 42. — McDermott 1966: 111 (= lateralis). non Lueiola erciata – Matsumura 1918: 86 (misspelling, typographical error). non Luciola Cxuca’a var. vitticollis – Matsumura 1918: 82 (misspelling, typographical error). non Luciola cruciata vitticollis – Matsumura 1928: 59, pl. 5 fig. 15, pl. 6 figs 5–6, pl. 7 fig. 14. non Luciola cruciata – Thapa 2000: 115 (incorrect record). Diagnosis One of the most famous fireflies in Japan, distinguished most obviously from the only other species in the genus, N. owadai gen. et comb. nov., by the pinkish pronotum with median blackish markings (that of owadai is yellowish orange without dark markings), and black MS and elytra (owadai has yellow MS and black elytra). Widely distributed throughout the main islands of Japan in contrast with owadai, which is restricted to a part of the Ryukyu islands, Kume-jima Island, and presently considered endangered in the locality. Macropterous females coloured as for males, except for white LO in V6, semitransparent anterior ⅔ of V7, with underlying pink fat body, and median posterior area black; V8 largely black. Type material Neotype of Luciola cruciata Motschulsky, 1854 (here designated) JAPAN • ♂; Kanagawa Pref., Yokosuka-shi, Nobi (Aza-Yatonota); 10 Jun. 2000; Itsuro Kawashima leg.; registration number: KPM-NK 80923. Remarks This designation fulfils the requirements of the ICZN neotype designation as follows: 75.3: in the absence of a type specimen there is a need to designate a neotype to preserve the existing taxonomy; 75.3.1: having established that the type locality given in Motschulsky (1854) as Java is incorrect, we designate a specimen from mainland Japan; 75.3.2: characters differentiating the neotype from other genera in the Luciolinae, and the other species in this new genus, owadai, are given in the generic description and key to species; 75.3.3: the specimen is fully labelled (outlined below) and given an identifying number in the type depository below; 75.3.4: introductory sections of this paper outline the steps taken to establish that no type specimen could be found; 75.3.5: this paper outlines all the references to this species including the original description by Motschulsky (1854), and we believe the morphological features of this specimen are consistent with all; 75.3.6: we established that the original type specimen could not have come from Java, but Japan; the locality chosen for this neotype designation is an area of high density of the species; 75.3.7: upon publication of this paper the neotype specimen, already lodged in the Kanagawa Prefectural Museum of Natural History, Odawara, Kanagawa, will be permanently lodged there and become the property of that museum. Notes Labels attached to the neotype are as follows (from top to bottom, a slash indicates a line break): (Nobi, Yatonota (in Japanese “Kanji” characters)) / Yokosuka-shi/ Kanagawa Pref. / VI ‒10, 2000 / I. Kawashima leg. (original white label, hand written in black ink); Luciola cruciata / Motschulsky, 1854 / det. I. Kawashima, 2004 (white determination label, printed in black ink); NEOTYPE / Luciola cruciata / Motschulsky, 1854 / = Nipponoluciola cruciata / (Motschulsky, 1854) / Ballantyne, Kawashima, Jusoh & Suzuki, 2021 (designated) (pink neotype label, printed in black ink). Other material examined JAPAN ‒ Akita Prefecture • 1 ♂; Higashi, Kawabe-machi; 39°39′ N, 140°12′ E; 25 Jun. 1994; K. Umetsu leg.; AKPM • 1 ♂; Sunakobuchi, Kawabe-machi; 39°44′ N, 140°33′ E; 25 Jun. 1994; K. Umetsu leg.; AKPM • 1 ♂; Yunosato, Akita-shi; 39°48′ N, 140°13′ E; Jul. 1995; K. Umetsu leg.; AKPM • 1 ♂; Noda, Taihei, Akita-shi; 39°43′ N, 140°20′ E; 25 Jun. 1994; K. Umetsu leg.; AKPM. • 1 ♂; Mt Ôtaki-yama, Akita-shi; 39°43′ N, 140°16′ E; 12 Jul. 1988; F. Satô leg.; AKPM • 1 ♀; Kawabemachi; 13 Jun. 2002; collector unknown; CIK. ‒ Niigata Prefecture • 1 ♂; Yomogihira, Nagaoka-shi; 37°36′ N, 138°53′ E; Jul. 1994; N. Harayama leg.; CIK. ‒ Ishikawa Prefecture • 2 ♂♂; Kamitokuyama, Nomi-shi; 36°43′ N, 136°54′ E; 3 Jul. 2012; H. Fukutomi leg.; CIK. ‒ Fukushima Prefecture • 1 ♀; Arakawa; 37°42′ N, 140°35′ E; Jun. 1993; K. Matsumoto leg.; CIK • 2 ♂♂; Jikiri-onsen Spa., Nagasaki, Iwaki-shi; 36°96′ N, 140°93′ E; 3 Jun. 1998; I. Kawashima leg.; CIK. ‒ Ibaraki Prefecture • 2 ♀♀; Ishizuka, Jôhoku-machi, Higashi-ibaraki-gun; 36°47′ N, 140°38′ E; Jun. 1988; K. Nakayama leg.; CIK • 4 ♂♂; same collection data as for preceding; 30 May 1990; CIK • 1 ♂; same collection data as for preceding; 1 Jun. 1992; CIK • 1 ♂; same collection data as for preceding; 20 Jun. 1995; CIK • 2 ♂♂; same collection data as for preceding; 25 Jun. 1998; CIK • 2 ♂♂; same collection data as for preceding; 8 Jun. 1999; CIK • 3 ♂♂; same collection data as for preceding; 10 Jun. 1999; CIK • 4 ♂♂, 2 ♀♀; same collection data as for preceding; 6 Jun. 2000; CIK • 1 ♂, 1 ♀; same collection data as for preceding; 6–7 Jun. 2002; CIK • 1 ♂, 1 ♀; same collection data as for preceding; 3 Jun. 2009; CIK. ‒ Saitama Prefecture • 2 ♂♂; Shôgun-sawa Riv., Ranzan-machi; 36°01′ N, 139°33′ E; 5 Jun. 1997; S. Arai leg.; CIK. ‒ Chiba Prefecture • 13 ♂♂, 1 ♀; Hirasawa-gawa Riv., Ôtaki-machi; 35°23′ N, 140°22′ E; Jun. 2019; S. Nishihara and I. Kawashima leg.; CIK • 1 ♂; Tsutsumori, Ôtaki-machi; 35°22′ N, 140°14′ E; 13 Jun. 2018; I. Kawashima leg.; CIK. ‒ Tokyo Metropolitan • 10 larvae; Hamura-shi; 35°45′ N, 139°31′ E; 2001; T. Sano leg.; CIK. ‒ Kanagawa Prefecture • 3 ♂♂; Kurokawa, Asao-ku, Kawasakishi; 35°36′ N, 139°45′ E; 9 Jun. 1990; Y. Goto leg.; CIK • 6 ♂♂; Inukura, Miyamae-ku, Kawasaki-shi; 35°58′ N, 139°56′ E; 9 Jun. 1991; T. Arikawa leg.; CIK • 3 ♂♂; Nobi (Aza-Yatonota), Yokosukashi; 35°21′ N, 139°41′ E; 10 Jun. 2000; I. Kawashima leg.; KPM • 3 ♂♂; same collection data as for preceding; CIK • 18 ♂♂; same collection data as for preceding; 11 Jun. 2001; CIK • 1 ♂; same collection data as for preceding; 26 Jun. 2021; CIK • 5 ♂♂; Morito-gawa Riv., Sakurayama-Ôyama Pass, Hayamamachi; 35°28′ N, 139°59′ E; 15 Jun. 2000; I. Kawashima leg.; CIK. ‒ Shizuoka Prefecture • 1 ♂; Katasumata-gawa Riv., Kanaya, Haibara-gun; 34°49′ N, 138°13′ E; 15 Jun. 1995; Y. Sato leg.; CIK • 3 ♂♂; Shizuoka-shi; 34°58′ N, 138°38′ E; 6 Jun. 1998; Y. Sato leg.; CIK. ‒ Yamanashi Prefecture • 1 ♂, 1 ♀; Yamanashi; 35°41′ N, 138°41′ E; 16 Jun. 1978; unknown leg.; (id N. Ohba); ANIC. ‒ Nagano Prefecture • 2 ♂♂; Sanozaka, Hakuba-mura; 36°37′ N, 137°50′ E; 13 Jul. 1986; M. Takakuwa leg.; CIK • 5 ♂♂, 1 ♀; Matsumoto-shi; 36°23′ N, 137°58′ E; 5 Jul. 1990; Y. Sato leg.; CIK. ‒ Mie Prefecture • 1 ♂; Osugi; 34°21′ N, 136°15′ E; 6 Jun. 1932; N. Tamu leg.; (id T. Nakane); ANIC. ‒ Gifu Prefecture • 4 ♂♂; Gujôhachiman, Gujo-shi; 35°44′ N, 136°58′ E; 5 Jun. 1988; H. Kuwano leg.; CIK • 4 ♂♂, 1 ♀; Kitaogi, Tajimi-shi; 35°33′ N, 137°11′ E; 18 Jun. 1994; H. Suzuki leg.; CIK. ‒ Hyogo Prefecture • 1 ♂; Kawanishi-shi; 34°49′ N, 135°41′ E; 1 Jul. 1992; K. Matsuda leg.; CIK • 3 ♂♂, 3 ♀♀; same locality data as for preceding; 6 Jul. 1996; T. Ochi leg.; CIK. ‒ Shimane Prefecture, Oki Islands (off W Honshu) • 1 ♂; Mt Akahage-yama, Chiburi Is.; 36°01′ N, 133°03′ E; 26 Jul. 2003; T. Shimada leg.; CIK • 7 ♂♂; Araki-gawa Riv., Saigo-chô, Dogo Is.; 36°20′ N, 133°32′ E; 13 Jun. 2003; T. Shimada leg.; CIK. ‒ Ehime Prefecture • 1 ♂; Nakayama, Omogo-mura, Kami-ukena-gun (Kumakôgen-chô at present); 33°39′ N, 133°06′ E; 3 Jul. 1999; Y. Goto leg.; CIK. ‒ Kochi Prefecture • 1 ♂; Amikawa, Tosayamamura; 33°39′ N, 133°50′ E; 19–20 Jun. 1999; Y. Goto leg.; CIK • 11 ♂♂; Shimotsui, Shimanto-chô; 33°29′ N, 132°55′ E; 17 Jun. 2006; I. Kawashima leg.; CIK. ‒ Fukuoka Prefecture • 1 ♂, 2 ♀♀; Doubaru, Kitakyushu-shi; 33°46′ N, 130 ° 49′ E; Jun. 1978; M. Nakamura leg.; (id by N. Ohba); ANIC. ‒ Nagasaki Prefecture • 6 ♂♂, 1 ♀; Tsushima Is. (off N. Kyushu), Kyôzuka, Izuhara-chô (Tsushimashi at present); 33°46′ N, 129°55′ E; 18 Jun. 1990; Y. Goto leg.; CIK; 9 ♂♂; Urakami-gawa Riv., Azebettô-chô, Nagasaki-shi; 32°48′ N, 129°55′ E; 20 May 2018; K. Tanaka leg.; CIK. ‒ Kumamoto Prefecture • 3 ♂♂, 1 ♀; Yubune, Kyokushi-mura (Kikuchi-shi at present); 32°55′ N, 130°52′ E; 29 May 1991; Y. Goto leg.; CIK. ‒ Miyazaki Prefecture • 10 ♂♂; Kita-gawa Riv., Kitagawa-chô; 32°43′ N, 131°39′ E; 3 Jun. 1993; Y. Goto leg.; CIK. ‒ Kagoshima Prefecture • 7 ♂♂, 4 ♀♀; Yaku-shima Is. (off S. Kyushu), Miyanoura to Ryûjinsugi-tozanguchi, Yakushima-chô; 30°37′ N, 130°52′ E; 29 May 2021; A. Ueno leg.; CIK. – Without locality • 7 ♀♀, pupa; reared by Y. Haneda; ANIC. Description Male (Figs 1–3, 5–6, 10–12) BODY LENGTH. 10.5–15.8 mm (Jeng et al. (2003) listed a range of 10.5‒16.5 mm long). COLOUR (Figs 1–2, 6). Entirely black except for pale pronotum with underlying pink fat body, and usually with a series of black markings extending narrowly across the anterior margin, continuous with a narrow median band that expands just anterior to midsection into short arms (the median cross) which may be wide or narrow, and continuing to posterior area where there may be a wide expansion across the posterior margin (Fig. 6). Aged pinned and ethanol preserved specimens may fade and the femoral base may appear brownish. A few specimens have very reduced pronotal colour patterns, and even fewer have no dark pronotal markings at all. IK observed many individual variations in pronotal colour among specimens from the same locality (see Pronotal colouration variations below). ABDOMEN (Fig. 2). Jeng et al. (2003: fig.7) illustrated a narrow MPP on V7 with well-defined posterolateral corners which were not prolonged. AEDEAGAL SHEATH (Fig. 5). Anterior margin of sheath tergite entire and slightly and broadly emarginated; sternite emarginated strongly on right side. Jeng et al. (2003: fig. 27) depict the same stronger emargination of the sheath sternite on its right side as we do here. AEDEAGUS (Figs 5, 10–12). L/W 3/1; either subparallel-sided or with lateral margins converging slightly; maximum width across LL/maximum width of ML 2.57–3.57; LL separated along the dorsal surface longitudinally by7/₉ of their length; LL apex width considerably wider than width of apex of ML; dorsal base of LL symmetrical, somewhat irregularly rounded; ML symmetrical, expanding slightly along its length to a maximum width around the ejaculatory orifice then narrowing in apical 1/6 or less where it is approximately 2/7‒2/9 the width of the more anterior portion; with rounded apex; BP well sclerotised, not hooded, emargination along anterior margin, may be absent. Jeng et al. (2003: figs 33–34) show an anterior median notch in the BP and the ML narrowing towards the apex. AEDEAGAL PATTERNS (Figs 10–12). For the first time we are able to demonstrate intraspecific variation in the aedeagal patterns of N. cruciata. Aedeagal patterns corresponded with the areas of east Honshu, west Honshu and the island of Kyushu. In E Honshu the lateral margins of the LL converge slightly; the apices of the LL are slightly narrowed; the ML is broader (LL/ML range: 2.57‒2.78), becoming widest around ⅓ of its length from the apex; when viewed from the side, the apex of the ML is not flattened vertically, but becomes thin and stick-like; the emargination of the anterior margin of the BP is more pronounced (Figs 10‒12, A‒E). In W Honshu and Kyushu the lateral margins of the LL are subparallel-sided; the apices of the LL are broader and more rounded; the ML is narrower (LL/ML range: 2.66‒3.57); when viewed laterally, the tip of the ML is depressed along both sides and becomes slightly wider vertically in lateral view; the emargination of the anterior margin of the BP is less pronounced (Figs 10‒12, F‒P). In populations in E Honshu, from the north to the Pacific coast (Fig. 12A‒E), aedeagus is generally upturned towards the ventral side when viewed from the side. In contrast, populations from Hokuriku to western Honshu, Shikoku and Kyushu (Fig. 12F‒P) are generally straight. Depictions of the aedeagus from line figures only are sometimes difficult to interpret. The figures in Jeng et al. (2003) are the exception. Assuming McDermott (1962: 24, fig. 20c) is left lateral then despite the absence of a basal piece, the ML can be interpreted as having a thin apex and curving strongly dorsally, similar to what we depict here for the E. Honshu pattern. Ohba’s (2004b: 91, fig. 3) depiction of the aedeagus shows a basal piece with median notch. Takakura’s (1977: fig. 2l) interpretation of the aedeagus may show a dorsal view but does show narrowing of the apex of the ML. Female (Fig. 7) BODY LENGTH. 14.0– 18.4 mm (Jeng et al. (2003) listed a range of 15.0– 18.6 mm long). COLOUR (Fig. 7). Coloured as for male except for abdomen; abdominal ventrites black except for white LO in V6 occupying all but a narrow transparent posterior margin; V7 (Fig. 7D) semitransparent with pink fat body granules visible beneath cuticle and clustered around anterior ⅔; median posterior area of V7 is largely devoid of fat bodies and appears black mainly due to the underlying black V8; V8 (observed when removed from intact specimen; Fig. 7I) black with lateral areas light brown; dorsal surface of basal tergites (up to T5), and dorsally reflexed margins of V2–V5 black; T6 (Fig. 7C) black with dorsally reflexed margins of V6 white, semitransparent; T7 (Fig. 7C) semitransparent with median dark stripe, dorsally reflexed margins of V6 appearing pink due to underlying fat body granules; T8 black (Fig. 7C). ABDOMEN (Fig. 7C–D, I). V8: lateral margins converge posteriorly; median posterior margin shallowly and narrowly emarginated; anterior apodeme and narrow anterior margin of V8 well sclerotised, pale coloured, appearing separate to posterior area of V8 because of intervening transparent area (Jeng et al. 2003: fig. 15, depict the posterior margin of V7 with a broad shallow emargination). REPRODUCTIVE SYSTEM (Fig. 7E–H). No intact spermatophores observed but spermatophore digesting gland may contain white particulate material which may represent digested spermatophore; median oviduct plate filling half of the median oviduct, anterior margin slightly irregular, lateral margins straight, posterior margin evenly shallowly emarginated. Ovipositor elongate slender (Fig. 7J). Larva (Figs 8–9) Final instar larvae were examined. Approx. body length 25–29 mm; maximum (median) length of protergum 3.2–3.4 mm; maximum width of protergum 2.6–2.9 mm. The following is modified and expanded from Fu et al, 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 28-39, DOI: 10.5852/ejt.2022.855.2023, http://zenodo.org/record/7501294, {"references": ["Motschulsky V. 1854. Lampyrides. Etudes Entomologiques 3: 47 - 62.", "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.", "Nakane T. 1987. Notes on some little-known beetles (Coleoptera) in Japan. Kita-Kyushu no Konchu, Fukuoka 34 (3): 171 - 176. [In Japanese with English title and descriptions.]", "Lacordaire J. T. 1857. Tribu II. Lampyrides. In: Lacordaire J. T. (ed.) Histoire Naturelle des Insectes. Genera des Coleopteres ou Expose Methodique et Critique de tous les Genres Proposes jusqu'ici dans cet Ordre d'Insectes. Volume 4: 304 - 341. Librairie encyclopedique de Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 8864", "Heyden L. v. 1879. Die coleopterologishce Ausbeute des Prof. Dr. Rein in Japan 1874 - 1875. Deutsche entomologische Zeitschrift 23: 321 - 365. https: // doi. org / 10.1002 / mmnd. 48018790221", "Gorham H. S. 1880. VIII. Materials for a revision of the Lampyridae. The Transactions of the Entomological Society of London 1880: 83 - 112. https: // doi. org / 10.1111 / j. 1365 - 2311.1880. tb 02023. x", "Olivier E. 1902 a. 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. 1902 b. Coleopteres Lampyrides receueillis aux environs de Tokio (Japon) par M. le Dr Harmand. Bulletin du Museum d'histoire naturelle 8 (2): 188 - 190. Available from https: // www. biodiversitylibrary. org / page / 42896473 [accessed 17 Nov. 2022].", "Olivier E. 1907. Coleoptera. Fam. Lampyridae. In: Wytsman P. A. (ed.) Genera Insectorum 54: 1 - 74, pls 1 - 3. Verteneuil & Desmet, Bruxelles. Available from https: // www. biodiversitylibrary. org / page / 33991802 [accessed 17 Nov. 2022].", "Olivier E. 1910. Lampyridae. In: Schenkling S. (ed.) Coleopterorum Catalogus pars 9: 1 - 68. W. Junk, Berlin, Germany. https: // doi. org / 10.1007 / 978 - 94 - 011 - 9697 - 0 _ 1", "McDermott F. A. 1962. Illustrations of the aedeagi of the Lampyridae (Coleoptera). The Coleopterists Bulletin 16 (1): 21 - 27.", "McDermott F. A. 1966. Lampyridae. In: Steel W. O. (ed.) Coleopterorum Catalogus Supplementa, Pars 9: 1 - 149. Uitgeverij Dr. W. Junk, The Hague.", "Takakura Y. 1977. Cantharoidea of Fukuoka Prefecture (Coleoptera). Biologia Fukuokana 17: 6 - 13. [In Japanese with English title.]", "Jeng M. - L., Lai J. & Yang P. - S. 2003. Lampyridae: a synopsis of aquatic fireflies with description of a new species. In: Jach M. A. & Ji L. (eds) Water Beetles of China Vol. 3: 539 - 562. Zoologisch-Botanische Gesellschaft in Osterreich und Wiener Coleopterologenverein, Vienna, Austria.", "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.", "Available from https: // www. biodiversitylibrary. org / page / 51901305 [accessed 16 Nov. 2022]. Motschulsky V. 1866. Catalogue des insectes recus du Japon. Bulletin de la Societe imperiale des naturalistes de Moscou 39 (1): 163 - 200. Available from https: // www. biodiversitylibrary. org / page / 44324258 [accessed 16 Nov. 2022].", "Harold E. 1877. Beitr age zur K aferfauna von Japan (Zweites Stuck). Deutsche Entomologische Zeitschrift 21: 337 - 367. https: // doi. org / 10.1002 / mmnd. 4800210220", "Lewis G. 1879. A Catalogue of Coleoptera from the Japanese Archipelago. Taylor and Francis, London, UK.", "Matsumura S. 1918. Nihon no Hotaru (The fireflies of Japan). Kyoiku-Gaho 6 (3): 82 - 89. [In Japanese.]", "Okada Y. K. 1928. Two Japanese aquatic glowworms. Transactions of the Royal Entomological Society of London 76 (1): 101 - 108. https: // doi. org / 10.1111 / j. 1365 - 2311.1928. tb 01193. x", "Okada Y. K. 1931. Honposan Hotaru-rui Gakumei no Kousatsu (Notes on the scientific names of the Japanese fireflies). Zoological Magazine Tokyo 43: 130 - 149. [In Japanese.]", "Kanda S. 1935. Hotaru (The Fireflies). Association of Luminous Organisms of Japan, Tokyo, Japan. [In Japanese.]", "Kishida K. 1936. Family Cantharidae, Insects of Jehol (VI) - Order Coleoptera (I). Report of the First Scientific Expedition Manchoukuo V (X) Art. No. 46: 1 - 21, pls 1 - 11. [In Japanese and English.]", "Hasama B. 1942 a. Uber die Biolumineszenz der Luciola lateralis im zytologischen Bild sowie im Potentialbild ihres Leuchtorgans. Cytologia 12: 366 - 377. https: // doi. org / 10.1508 / cytologia. 12.366", "Hasama B. 1942 b. Uber die Biolumineszenz der Larve von Luciola cruciata sowie von Pyrocoelia rufa im Aktionsstrombild und im histologischen Bild ihres Leuchtorgans. Cytologia 12: 378 - 388. https: // doi. org / 10.1508 / cytologia. 12.378", "Hasama B. 1943. Lebensgeschichte der Luciola cruciata und der Luciola lateralis. Annotationes Zoologicae Japonenses 22 (1): 23 - 47.", "Hasama B. 1944. Entwicklung des imaginalen Leuchtorgans der Luciola cruciata in histologischer sowie bioelektrischer Hinsicht. Cytologia 13 (2): 155 - 161. https: // doi. org / 10.1508 / cytologia. 13.155", "Nakane T., Ohbayashi K., Nomura S. & Kurosawa Y. 1959. Iconographia Insectorum Japonicorum Colore Naturali Edita. Vol. II (Coleoptera). Lampyridae: 169 - 170, pl. 85. Hokuryu-kan Publishing Co., Tokyo. [In Japanese with Latin title.]", "Nakane T. 1960. Lampyridae. In: Nakane T. & The Japan Coleopterological Society (eds) Coloured Illustrations of the Insects of Japan. Coleoptera: 118 - 119, pl. 36. Kinki Coleopterological Society, Hoikusha Publishing, Tokyo. [In Japanese.]", "Nakane T. 1968. Hotaru no bunrui (Classification of lampyrid beetles). The Nature and Insects 3 (6): 3 - 6. [In Japanese.]", "Nakane T. 1970. On the cantharoid Coleoptera found in the islands of Tsushima (Insecta). Memoirs of the National Science Museum, Tokyo 3: 287 - 288. [In Japanese with English summary.]", "Nakane T. 1983. In: Hotaru no Kansatsu to Shiiku (Observation and Breeding of Fireflies): 81 - 113. New Science Co.", "Minami K. 1961. Hotaru no Kenkyu (A study of fireflies). Ohta-shoten, Hikone. [In Japanese.]", "Bertrand H. P. I. 1972. Lampyridae. In: Bertrand H. P. I. (ed.) Larves et Nymphes des Coleopteres aquatiques du Globe: avec Tableaux de Determination des Genres: 598 - 603. Impr. F. Paillart, Abbeville.", "Bertrand H. P. I. 1973. Part XI: larvae and pupae of water beetles collected from the island of Ceylon. Bulletin of the Fisheries Research Station, Sri Lanka (Ceylon) 24 (1 & 2): 95 - 112.", "Sato M. 1974. Luminous insects. In: Konchu no Sekai (The World of Insects): 130 - 137. Hoikusha Publishing Co., Ltd., Osaka. [In Japanese.]", "Sato M. 1978. Lampyrid-beetles of Japan. The Insectarium, Tokyo 15: 168 - 173. [In Japanese.]", "Sato M. 1985. Lampyridae. In: Kurosawa Y., Hisamatsu S. & Sasaji H. (eds) Colored Illustrations of the Coleoptera of Japan Vol. III: 121 - 124, pl. 20. Hoikusha Publishing Co., Ltd., Osaka, Japan. [In Japanese.]", "Sato M. 1989. A Check List of Japanese Insects. I. Lampyridae: 351 - 353. Kyushu University, Fukuoka. [In Japanese with English title.]", "Ohba N. 1978. Morphology and behavior of the Lampyridae (Coleoptera, Insecta). Science Report of the Yokosuka City Museum 25: 15 - 28, pls 2 - 3. [In Japanese with English abstract.]", "Ohba N. 1983. Studies on the communication system of Japanese fireflies. Science Report of the Yokosuka City Museum 30: 1 - 62, pls 1 - 6.", "Ohba N. 1984. Synchronous flashing in the Japanese firefly, Luciola cruciata (Coleoptera: Lampyridae). Science Report of the Yokosuka City Museum 32: 23 - 32, pl. 8.", "Ohba N. 2001. Geographical variation, morphology and flash pattern of the firefly, Luciola cruciata (Coleoptera: Lampyridae). Science Report of the Yokosuka City Museum 48: 45 - 89. [In Japanese with English abstract.]", "Ohba N. 2004 a. Flash communication systems of Japanese fireflies. Integrative Comparative Biology 44: 225 - 233. https: // doi. org / 10.1093 / icb / 44.3.225", "Ohba N. 2005. Lampyridae, Rhagophthalmidae. In: Ishiwata S. et al. (eds) Insect Larvae of Japan: 237 - 243. Gakken, Tokyo. [In Japanese.]", "Ohba N. 2012. The restoration and maintenance of firefly habitats in Japan - case studies from Yokosuka City. Lampyrid 2: 13 - 23.", "Yuma M. 1981. The body size variations of the climbing larvae of the firefly, Luciola cruciata (Coleoptera; Lampyridae). Japanese Journal of Ecology 31 (1): 57 - 66.", "Ohba N., Azuma S., Nishiyama K., Goto Y., Suzuki H., Sato Y. & Kawashima I. 1994. Morphology, life history and behaviour of the firefly Luciola owadai (Coleoptera: Lampyridae). Science Report of the Yokosuka City Museum 42: 13 - 26. [In Japanese with English abstract.]", "Suzuki H., Sato Y., Fujiyama S. & Ohba N. 1996 a. Allozymic differentiation between two ecological types of flashing behavior in the Japanese firefly, Luciola cruciata. Japanese Journal of Entomology 64 (3): 682 - 691.", "Suzuki H., Sato Y., Fujiyama S. & Ohba N. 1996 b. Biochemical systematics of Japanese fireflies of the subfamily Luciolinae and their flash communication systems. Biochemical Genetics 34 (5 / 6): 191 - 200. https: // doi. org / 10.1007 / BF 02407019", "Suzuki H., Sato Y., Ohba N., Bae J-S., Jin B-R., Sohn H-D. & Kim S-E. 2004. Phylogeographic analysis of the firefly, Luciola lateralis, in Japan and Korea based on mitochondrial cytochrome oxidase II gene sequences (Coleoptera: Lampyridae). Biochemical Genetics 42 (9 / 10): 287 - 300. https: // doi. org / 10.1023 / B: BIGI. 0000039805.75118.8 f", "Suzuki H. 1997. Molecular phylogenetic studies of Japanese fireflies and their mating systems (Coleoptera: Cantharoidea). Tokyo Metropolitan University Bulletin of Natural History 3: 1 - 53.", "Suzuki H. 2001. Studies on biological diversity of firefly in Japan. International Journal of Industrial Entomology 2 (2): 91 - 105.", "Sawada Y. 2000. A List of Japanese Insect Collection by P. F. von Siebold and H. Burger preserved in Nationaal Natuurhistorisch Museum, Leiden, The Netherlands Part 2. Coleoptera. Bulletin Kitakyushu Museum Natural History 19: 77 - 104", "Branham M. A. & Wenzel J. W. 2003. The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). Cladistics 19 (1): 1 - 22. https: // doi. org / 10.1111 / j. 1096 - 0031.2003. tb 00404. x", "Takeda M., Amano T., Katoh K. & Higuchi H. 2006. The habitat requirement of the Genji-firefly Luciola cruciata (Coleoptera: Lampyridae), a representative endemic species of Japanese rural landscapes. Biodiversity and Conservation 15: 177 - 189. https: // doi. org / 10.1007 / s 10531 - 004 - 6903 - y", "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): 1 - 188. https: // doi. org / 10.11646 / zootaxa. 1997.1.1", "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", "Oba Y., Branham M. A. & Fukatsu T. 2011. The terrestrial bioluminescent animals of Japan. Zoological Science 28 (11): 771 - 789. https: // doi. org / 10.2108 / zsj. 28.771", "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", "Ballantyne L. A., Lambkin C. L., Luan X., Boontop Y., Nak-Eiam S., Pimpasalee S., Silalom S. & Thancharoen A. 2016. Further studies on south eastern Asian Luciolinae: 1. Sclerotia Ballantyne, a new genus of fireflies with back swimming larvae 2. Triangulara Pimpasalee, a new genus from Thailand (Coleoptera: Lampyridae). Zootaxa 4170 (2): 201 - 249. https: // doi. org / 10.11646 / zootaxa. 4170.2.1", "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. 2009. Larval respiration system and evolution in aquatic fireflies (Coleoptera: Lampyridae: Luciolinae). In: Meyer-Rochow V. B. (ed.) Bioluminescence in Focus - A Collection of Illuminating Essays: 243 - 253. Research Signpost, Kerala.", "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", "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", "Fu X. H., Ballantyne L. A. & Lambkin C. L. 2012 b. The external larval morphology of aquatic and terrestrial Luciolinae fireflies (Coleoptera: Lampyridae). Zootaxa 3405 (1): 1 - 34. https: // doi. org / 10.11646 / zootaxa. 3405.1.1", "Jusoh W. F. A., Ballantyne L. A., 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", "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", "Kato D., Suzuki H., Tsuruta A., Maeda J., Hayashi Y., Arima K., Ito Y. & Nagano Y. 2020. Evaluation of the population structure and phylogeography of the Japanese Genji firefly, Luciola cruciata, at the nuclear DNA level using RAD-Seq analysis. Scientific Reports 2020 (10): 1533. https: // doi. org / 10.1038 / s 41598 - 020 - 58324 - 9", "Gorham H. S. 1883. Revision of the genera and species of Malacoderm Coleoptera of the Japanese fauna. The Transactions of the Entomological Society of London 1883: 393 - 411. https: // doi. org / 10.1111 / j. 1365 - 2311.1883. tb 02955. x", "Okamoto H. 1924. The insect fauna of Quelpart island (Saishu-to). Bulletin of the Agricultural Experiment Station, Government-General of Chosen 1 (2): IV + 47 - 233, pls VII - X.", "Matsumura S. 1928. II. Hotaru (Chapter II The fireflies). In: Matsumura S. (ed.) Omoshiroki Chukai no Kyozai (Interesting Insects for Educational Materials). Tokyo-do Publishing Company, Tokyo. [In Japanese.]", "Geisthardt M. & Sato M. 2007. Lampyridae. In: Lobl I. & Smetana A. (eds) Catalogue of Palaearctic Coleoptera, Vol. 4: 225 - 234. Apollo Books, Stenstrup, Denmark.", "Thapa V. K. 2000. An Inventory of Nepal's Insects. Vol. III. IUCN Nepal, Kathmandu.", "Ohba N. (ed.) 2004 b. Hotaru Tenmetsu no Fushigi, Chikyu no Kiseki (Mystery of Fireflies). Yokosuka City Museum, Yokosuka. [In Japanese with English title.]", "Hara S. 1962. Larvae of Luciola cruciata and L. lateralis (Coleoptera). Kontyu 30 (4): 230 - 235. [In Japanese with English title.]", "Kanda S. 1933. Hotaru no kenkyu (1) Genji-botaru no seikatushi (Study of firefly (1) Life history of Genji firefly). Kontyu, Tokyo 7 (5 - 6): 219 - 239. [In Japanese.]", "Kuribayashi S. 1979. Genji-botaru (Luciola cruciata). Nature Books, Tokyo. [In Japanese with English explanation.]", "Ohba N. 1988. Genjibotaru (Genji firefly). Bun-ichi Publishing Co. Ltd., Tokyo. [In Japanese.]", "Watase S. 1902. Hotaru no Hanashi (The story of fireflies). Kaiseikan Publishing, Tokyo. [In Japanese.]", "Iguchi Y. 2010. Temperature-dependent geographic variation in the flashes of the firefly Luciola cruciata (Coleoptera: Lampyridae). Journal of Natural History 44 (13 - 14): 861 - 867. https: // doi. org / 10.1080 / 00222930903528206", "Ohba S-y., Numata K. & Kawano K. 2020. Variation in flash speed of Japanese firefly, Luciola cruciata (Coleoptera: Lampyridae), identifies distinct southern \" quick-flash \" population on Goto Islands, Japan. Entomological Science 23 (2): 117 - 230. https: // doi. org / 10.1111 / ens. 12403", "Suzuki H., Sato Y. & Ohba N. 2002. Gene diversity and geographic differentiation in mitochondrial DNA of the Genji firefly, Luciola cruciata (Coleoptera: Lampyridae). Molecular Phylogenetics and Evolution 22 (2): 193 - 205. https: // doi. org / 10.1006 / mpev. 2001.1046", "Yuma M. & Hori M. 1981. Gregarious oviposition of Luciola cruciata Motschulsky (Coleoptera: Lampyridae). Physiology and Ecology Japan 18: 93 - 112.", "Kawano K. 2013. Studies on Mating Behavior of Luciola cruciata Motschulsky (Coleoptera: Lampyridae). Personal Publication Yamaguchi. [In Japanese with English abstract.]", "Tamura M., Yokoyama J., Ohba N. & Kawata M. 2005. Geographic differences in flash intervals and pre-mating isolation between populations of the Genji firefly Luciola cruciata. Ecological Entomology 30: 241 - 245. https: // doi. org / 10.1111 / j. 0307 - 6946.2005.00683. x", "Miyashita M. 2011. Effects of LED lighting on the oviposition of firefly, Luciola cruciata and Luciola lateralis. Journal of Japan Society of Civil Engineers Ser G (Environmental Research) 67 (1): 21 - 29. [In Japanese with English abstract.] https: // doi. org / 10.2208 / jscejer. 67.21", "Moriya S., Yamanichi T. & Nagaghoshi N. 2009. Sex ratios in the Japanese firefly, Luciola cruciata (Coleoptera: Lampyridae) at emergence. Japanese Journal of Limnology 69 (3): 255 - 258. https: // doi. org / 10.3739 / rikusui. 69.255", "Yuma M. 1984. Egg size and viability of the firefly, Luciola cruciata (Coleoptera, Lampyridae). Kontyu, Tokyo 52 (4): 615 - 629.", "Yuma M. 1986. Growth and size variations in the larvae of Luciola cruciata (Coleoptera: Lampyridae) in relation to the egg size. Physiology and Ecology Japan 23: 45 - 78.", "Yuma M. 2001. Effects of illuminance and tree coverage on the density of adult Genji-firefly, Luciola cruciata (Coleoptera). Ecology and Civil Engineering 4 (1): 59 - 63. https: // doi. org / 10.3825 / ece. 4.59 [In Japanese with English abstract.]", "Yuma M. 2007. Effect of rainfall on the long-term population dynamics of the aquatic firefly Luciola cruciata. Entomological Science 10 (3): 237 - 244. https: // doi. org / 10.1111 / j. 1479 - 8298.2007.00219. x", "Yuma M. & Ono K. 1985. Seasonal changes and population estimate of the adult firefly, Luciola cruciata (Coleoptera: Lampyridae), along the Biwako Canal, Kyoto City. Science Report of the Yokosuka City Museum 33: 1 - 11. [In Japanese with English abstract.]", "Yuma M. & Hori M. 1990. Seasonal and age-related changes in the behaviour of the Genji firefly, Luciola cruciata (Coleoptera, Lampyridae). Japanese Journal of Entomology 58 (4): 863 - 870.", "Iguchi Y. 2001. Seasonal variation in the adult body size of the Genji-firefly Luciola cruciata (Coleoptera: Lampyridae). Entomological Review of Japan 56 (1): 35 - 38.", "Iguchi Y. 2009 a. The ecological impact of an introduced population on a native population in the firefly Luciola cruciata (Coleoptera: Lampyridae). Biodiversity and Conservation 18: 2119 - 2126. https: // doi. org / 10.1007 / s 10531 - 009 - 9576 - 8", "Iguchi Y. 2009 b. Genji firefly import problem. Journal of the National Firefly Research Association 42: 35 - 38. [In Japane

Colophotia miranda Olivier in Baer, 1886 Colophotia miranda Olivier in Baer 1886: 186; Olivier 1902: 76; 1913: 59. McDermott 1966: 116. Ballantyne and Lambkin 2013: 64. Ballantyne et al. 2019: fig. 474���483. Type. Not located in MNHN during LB���s visit in 2013. Specimen examined. PHILIPPINES as Philippine islands, no collection date nor collector. Male determined Colophotia miranda Olivier by KG Blair (NHMUK). Summary of diagnostic features. Dorsal colour pale brown; 15.8 mm long; light organs in ventrite 7 bipartite and widely separated; median posterior area of ventrite 7 terminated by a single elongate slender apically acute ���appendage��� which is considered to conform to Olivier���s (1886) description. Male. 15.8 mm long; 6.8 mm wide measured at region of pin through right elytron. Colour (Fig. 1A, B): Dorsal surface very pale brown; pronotum semitransparent with fat body showing irregularly through cuticle; head between eyes light brown, semitransparent, with fat body visible; left antenna with pale brown scape and remaining segments darker brown; right antenna colour ventral surface scape pale brown, pedicel and apical two segments dark brown, dorsal surface as for left; ventral surface of thorax and legs pale brown; abdomen in original state before immersion in any fluids with dorsum pale brown semitransparent and fat bodies visible beneath cuticle in all but terminal tergite; ventral surface with V 2���4 pale brown semitransparent, V 5 slightly paler, V 6 with LO plaques adhering to lateral areas of ventrite appearing mid brown and intervening area semitransparent with irregular distribution of fat body beneath; anterior area of V 7 in dry specimen with lateral brown LO plaques and an apparent posterior margin separating this from the posterior area which is encircled by paler brown PLP incurving, and was covered in light brown material of unknown origin; after soaking in KOH all of the ventral surface of the MPP and PLP, and a narrow area anterior to the emarginations appears to be clumped fat body beneath. Pronotum (Fig. 1A, B): 3.0 mm long, 4.7 mm wide; anterior margin broadly rounded scarcely projecting beyond the rounded obtuse anterolateral corners; lateral margins diverging in anterior half and pronotum widest across middle; lateral margins converge posteriorly with slightly angulate posterior corners not behind anterior corners but to the sides; posterior corners slightly obtuse; median posterior margin shallowly emarginated and projecting a little beyond the posterior corners; most of dorsal surface smooth with flattened and widely explanate lateral margins; median area of disc a little elevated and shallow depressions just behind anterior margin behind head, and across and in front of posterior margin, not extending to laterally explanate region; pronotal width slightly less than humeral width. Elytra (Fig. 1A): 12.8 mm long; no elevated interstitial lines; lateral margins slightly convex sided; with wide epipleuron extending laterally just behind anterior margin of hypomeron and diminishing in width towards elytral apex. Head (Fig. 1A): GHW 2.3 mm; SIW 0.9 mm; ASD subequal to ASW; vertex barely depressed between eyes. Antennae (Fig. 1A): 11 segmented, three times as long as GHW, with elongate scape, small pedicel and all flagellar segments longer than wide; FS 1���7 four times as long as wide; FS 8 �� as long as FS 7, and FS 9 �� as long as FS8. Mouthparts (Fig. 1A): well developed and male can probably feed; apical labial palpomere in the form of an elongate slender triangle (L 4���5 �� W) with inner margin entire. Legs (Fig. 1A): no tibiae swollen or curved; hind femora stout slightly wider than femora 1 and 2. Abdomen (Fig. 1D, 2): V 6: appears to have no light organs in the median area and they may be emarginated in this segment, or the median area devoid of light organ is a consequence of some aspect of dehydration and the dissection, as there are no obvious muscle attachments in the median area of V 6; V 7: with broadly and evenly emarginated anterior margin (not obvious in intact specimen); lateral margins converging posteriorly; posterior margin trisinuate (Fig. 2A, B) with oblique PLP which are half the length of the MPP and curve inwards partly embracing the MPP in dried specimen (Fig. 1D); deep slightly irregular emarginations separate MPP and PLP; MPP broad and deep with concave dorsal surface; apically truncated with a single elongate very slender apically acute appendage arising from the left side of the apex, and remaining margins densely hairy; dorsal surface V 7 with brown facing cuticle covering sides of MPP and around emarginations to the PLP. Muscle attachments (Fig. 2A, B): a large block of longitudinal muscles attach on the median area of V 7 between light organ halves and narrow to a median band along the dorsal median surface of the MPP; in the anterior portion these muscles split into two wide bands attaching to the anterior margin on either side of the anterior emargination of V 7; small lateral bands reach the sides anterior to the PLP; T 8 (Fig. 2 D���F): lateral margins straight converging slightly posteriorly; median posterior margin with a deep and narrow emargination, margined by acute corners, and further shallow lateral emarginations between median area and rounded posterolateral corners; dorsal surface with deep median longitudinal emargination which houses the aedeagus and sheath in repose and distinct acute margins separating this median channel from flat side areas which have no spines; there is no muscle attachment posterior to the flanges in T 8; strongly sclerotized and dark flanges with obliquely truncated apices each projecting posteriorly arise at the anterior end of the acute margins of the median trough; anterolateral paired prolongations of T 8 rounded and slightly shorter than the median length of the entire posterior portion. Muscle attachments (Fig. 2 E, F): strong bands of longitudinal muscles attach onto the basal portion of the flanges. Aedeagal sheath Fig. 3 A���C): elongate slender about 7 �� as long as wide; tergite of sheath in two sections; anterior margin evenly emarginated. Aedeagus (Fig. 3 D���F): elongate slender about 6 �� as long as wide; appearing slightly asymmetrical (basal piece is turned to one side); lateral lobes very short scarcely separated., Published as part of Ballantyne, Lesley & Jusoh, Wan F. A., 2021, The strange case of Colophotia miranda Olivier, 1886 (Coleoptera: Lampyridae: Luciolinae), pp. 1-9 in Insecta Mundi 2021 (857) on page 3, DOI: 10.5281/zenodo.5041739, {"references":["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. 1913. The Lampyridae of Borneo. Sarawak Museum Journal 3: 55 - 60.","McDermott FA. 1966. Luciolini. p. 98 - 118. In: Steel WO (ed.). Coleopterorum catalogus supplementa. Pars 9 (Editio secunda). Lampyridae. W. Junk; The Hague. 149 p.","Ballantyne LA, Lambkin CL. 2013. Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa 3653: 1 - 162.","Ballantyne LA, Lambkin CL, Ho JZ, Jusoh WFA, 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 - 174.","Olivier E. 1886. Colophotia miranda. p. 186 - 187. In: Baer G-A. Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France 1886: 97 - 200."]}

Ballantyne, Lesley, Jusoh, Wan F. A. (2021): The strange case of Colophotia miranda Olivier, 1886 (Coleoptera: Lampyridae: Luciolinae). Insecta Mundi 2021 (857): 1-9, DOI: http://doi.org/10.5281/zenodo.5041739, {"references":["Ballantyne LA. 1987. Lucioline morphology, taxonomy and behaviour: A reappraisal (Coleoptera, Lampyridae). Transactions of the American Entomological Society 113(2): 171-188.","Ballantyne LA, Lambkin CL. 2013. Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa 3653: 1-162.","Ballantyne LA, Lambkin CL, Ho JZ, Jusoh WFA, 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: 1-174.","Gorham HS. 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 34: 293-307.","McDermott FA. 1966. Luciolini. p. 98-118. In: Steel WO (ed.). Coleopterorum catalogus supplementa. Pars 9 (Editio secunda). Lampyridae. W. Junk; The Hague. 149 p.","Olivier E. 1883a. Lampyrides nouveaux ou peu connus. Revue D'Entomologie 3: 73-80,","Olivier E. 1883b. Lampyrides nouveaux ou peu connus. Revue D'Entomologie 3: 326-332.","Olivier E. 1886. Colophotia miranda. p. 186-187. In: Baer G-A. Catalogue des Coleopteres des Iles Philippines. Annales de la Societe Entomologique de France 1886: 97-200.","Olivier E. 1890. Lampyrides rapportes de Borneo par M. Platteeuw. Annales de la Societe Entomologique de Belgique 34: xxx-xxxi.","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. 1903. Lampyrides nouveaux du Musee de Bruxelles. Annales de la Societe Entomologique de Belgique 47: 11-12.","Olivier E. 1907. Luciolinae. p. 48-56. In: Olivier E. Coleoptera, Lampyridae. Genera Insectorum 53: 1-74.","Olivier E. 1913. The Lampyridae of Borneo. Sarawak Museum Journal 3: 55-60.","Pic M. 1924. Lampyrides et Telephorides des Iles Philippines. The Philippine Journal of Science 25: 711-731.","Pic M. 1927. Notes diverses, descriptions et diagnoses. L'Echange, Revue Linneenne XLIII: 37.","Received January 27, 2021; accepted February 27, 2021."]}

Pygoluciola dunguna Nada, 2018 was described from Peninsular Malaysia, using males and reliably associated females. This paper details description of the larva which has been conclusively identified as Pygoluciola dunguna based on DNA barcoding technique and uses morphology, brief habitat and behavioural data. A total of 70 larval specimens were measured and their main features described. The larvae exhibit a riparian or semi-aquatic behaviour, observed crawling on the sandy edge of shallow streams. The stake-like projections along the length of the body suggest a form of defensive mechanism from falling prey to aquatic predators.

Fireflies are charismatic beetles with attractive bioluminescent courtship displays that have recently been swept onto the global stage of nature tourism. Here, we provide the first comprehensive review of the geographic scope, magnitude, focal species, and other attributes of the major firefly tourism sites worldwide. Through targeted interviews and surveys, we estimate that in recent years over one million tourists travelled annually to sites located in at least 12 countries for firefly‐watching. Rapid proliferation of firefly tourism provides a timely opportunity to examine how such activities may impact local firefly populations, and to highlight the biological factors that make certain species especially vulnerable to tourism‐associated threats. We offer science‐based best practices for firefly ecotourism that can be tailored to fit local circumstances, including recommendations to: (1) Conserve the habitats required for all life stages to thrive, (2) involve local communities as key stakeholders, and (3) provide training programs for guides and interpretive materials for visitors. Finally, we provide suggestions for transforming tourist behavior to minimize impact on firefly populations. By developing management plans that incorporate these recommendations, firefly tourist sites can enhance the visitor experience, protect natural resources, benefit local communities, and help promote the conservation of invertebrate biodiversity.

Identification and revision of firefly species are largely based on morphological characteristics. Subsequently, female firefly specimens have been neglected in taxonomic inventories because morphological descriptions relied only on male firefly specimens. In light of successful DNA barcoding in many invertebrates, this study explored the potential of DNA barcoding to complement morphological identification of fireflies in Malaysia. Firefly specimens were collected from riparian forests (including mangroves) in Peninsular Malaysia and Sarawak. Specimens were morphologically examined, and their DNA was extracted for molecular analysis. The morphological data of the male specimens was found to be congruent with molecular data, i.e., both indicated 12 species. However, inclusion of females and larvae in the DNA analysis increased the species count to 14 species. We also found distinct geographically structured variations in the DNA barcodes of two species: Pteroptyx tener Olivier and Pteroptyx bearni Olivier. The study indicates that firefly species delimitation using traditional taxonomic knowledge and new technology like DNA barcoding are compatible, therefore increasing the reliability of species identification.