Life cycle of the aquatic firefly Luciola ficta (Coleoptera: Lampyridae)

This research is the first to record the complete life history of the aquatic firefly Luciola ficta (Olivier) using a unique individual rearing method. Transparent containers (250 ml; height: 6 cm; bottom diameter: 8 cm; mouth diameter: 9.5 cm) were used to rear individuals from egg to adult in the laboratory, so that they could be observed throughout the whole life cycle. Larvae were fed on the meat of the water snail Cipangopaludina chinensis (Gray). Temperature ranged from 18 °C to 30 °C, relative humidity (RH) was 80 ± 5%, and the light:dark (L:D) ratio was 10:14. Of 80 eggs, 35 individuals completed their life cycle under these laboratory conditions in Jiji, Nantou County, Taiwan. The external morphological characteristics of each growing stage were described. Egg hatching rate was 95%. On average, one generation spanned 388.2 ± 25.7 days. The durations of egg, larva, climbing larva, cocoon, and adult stages were 19.1 ± 1.5 days, 328.9 ± 33.2 days, 10.9 ± 7.8 days, 14.7 ± 5.3 days, and 15.7 ± 5.2 days, respectively. The number of larval instars ranged from five to seven, with a modal value of six instars for males and seven instars for females. Female larval duration averaged 337.1 ± 31.2 days, which was higher than the 307.6 ± 34.1 days of the males. From January to December 2002, adult emergence peaked twice, with the main high peak appearing in April and the second peak occurring in August. The results of indoor rearing and of field investigations in Jiji, Nantou County, suggested that L. ficta is univoltine. Adult body length is negatively correlated with larval duration (P < 0.01). The life history traits of L. ficta show plasticity in adult occurrence, egg size, egg duration, larval duration, larval instars, and adult body length. Some variations were discussed in the context of survivorship in field habitats.     

Firefly luciferase genes from the subfamilies Psilocladinae and Ototretinae (Lampyridae, Coleoptera)

Firefly luciferase genes have been isolated from approximately 20 species of Lampyrinae, Luciolinae, and Photurinae. These are mostly nocturnal luminescent species that use light signals for sexual communication. In this study, we isolated three cDNAs for firefly luciferase from Psilocladinae (Cyphonocerus ruficollis) and Ototretinae (Drilaster axillaris and Stenocladius azumai), which are diurnal non-luminescent or weakly luminescent species that may use pheromones for communication. The amino acid sequences deduced from the three cDNAs showed 81-89% identities to each other and 60-81% identities with known firefly luciferases. The three purified recombinant proteins showed luminescence and fatty acyl-CoA synthetic activities, as observed in other firefly luciferases. The emission maxima by the three firefly luciferases (λmax, 545-546 nm) were shorter than those by known luciferases from the nocturnal fireflies (λmax, 550-568 nm). These results suggest that the primary structures and enzymatic properties of luciferases are conserved in Lampyridae, but the luminescence colors were red-shifted in nocturnal species compared to diurnal species.     

Bioluminescence in firefly larvae: A test of the aposematic display hypothesis (Coleoptera: Lampyridae)

We investigated the hypothesis that bioluminescence in firefly larvae (Coleoptera: Lampyridae) functions as an aposematic display. In two experiments, we confirmed the distastefulness of firefly larvae, and tested the hypothesis that a naive, nocturnal predator can learn to use light signals as aposematic cues for avoiding distasteful prey. Larvae were rejected as acceptable prey by 100% of the house mice (Mus musculus) tested. Mice learned to avoid bitter food associated with light cues significantly faster (P=0.003) than mice presented with food lacking light cues. We conclude that luminescent glowing in firefly larvae meets the requirements of an aposematic signal.     

Wolbachia infection in the Korean endemic firefly,Luciola unmunsana (Coleoptera: Lampyridae)

Wolbachia is one of the most prevalent endosymbiontic bacteria of arthropods. The bacterium induces sex ratio distortions in various host insects through processes such as cytoplasmic incompatibility, feminization, male killing, and parthenogenesis. We investigated if the Korean endemic firefly, Luciola unmunsana was infected with the bacterium because the species had an abnormal sex ratio in the field. The results show that some individuals are infected with the bacterium. Phylogenetic analyses showed that the bacterial strain infecting the firefly is closely related to strains that infect phylogenetically distant hosts.     

Systematic review of the firefly genus Scissicauda (Coleoptera,Lampyridae, Amydetinae) from Brazil

The Amydetinae genus Scissicauda McDermott, 1964 is reviewed and redescribed. We describe Scissicauda balena sp. n. from Brazil as new, and provide illustrations of the structural features and a key to species of both sexes.     

The first cytogenetic report of fireflies (Coleoptera, Lampyridae) from Brazilian fauna

The first cytogenetic analysis of fireflies from Brazilian fauna was carried out in this work. The investigation of two species of the subfamily Lampyrinae, Aspisoma maculatum and Photinus sp. (aff. pyralis), showed the diploid number 2n = 19 and an X0 sex determination system in males. These observations are similar to those already described for all the Lampyrinae species previously studied. In contrast, Bicellonycha lividipennis (Photurinae) revealed the karyotype 2n = 16 + neoXY, which has not yet been registered for any firefly species. The neoXY sex determination system encountered in this species probably arose through fusion between an ancestral X sex chromosome, belonging to the X0 system, and an autosomal element. This event also reduced the diploid number from 2n = 19, which is more frequent in the family Lampyridae, to 2n = 18 in B. lividipennis. The analysis of meiotic cells showed that the neoXY sexual bivalent of B. lividipennis exhibited a prominent terminal chiasma, indicating that the sex chromosomes are not wholly differentiated and still retain a region of homology. A review of the cytogenetic data known for the family Lampyridae was also documented in this work, as well as a discussion on the main trends of chromosomal evolution that seem to have occurred in this group.     

The nearly complete mitogenome of the Southeast Asian firefly Pteroptyx tener (Coleoptera: Lampyridae)

Molecular Biology Reports - The estuarine firefly, Pteroptyx tener, aggregates in the thousands in mangrove trees lining tidal rivers in Southeast Asia where they engage one another in a nocturnal,...     

Variation in the flash pattern of the firefly,Photuris versicolor quadrifulgens (Coleoptera: Lampyridae)

We sampled a population of signalingPhoturis versicolor quadrifulgens fireflies to quantify the variation in flash patterns emitted by males. Males produced five distinct flash patterns during their mate-searching flights. Four of the patterns consisted of two to five equal-intensity pulses and the fifth pattern type was a flicker, a group of rapid modulations in intensity. We found that the proportions of each pattern remained relatively constant from night to night throughout the season. The different flash patterns produced varied significantly with time of night; patterns having fewer pulses occurred earlier in the evening. Local density, an estimate of competition, did not significantly correlate with flash pattern type. On consecutive emissions, individuals changed their flash types with a mean probability of 0.12 (over all males), and they usually switched between patterns differing by a single pulse (from a two- to a three-pulse pattern, from a three- to a two-pulse pattern, etc.). The nightly temporal changes in flash patterns may be related to tradeoffs between female availability and energetic costs of signaling or the changes may be related to increased predation risk from visual predators. Photuris versicolor quadrifulgens was originally described by Barber (1951) as a subspecies ofP. versicolor. The genusPhoturis is currently under revision by Dr. James E. Lloyd, and this firefly will be given species status.     

条背萤的形态和生物学研究

首次在中国大陆发现水栖萤火虫条背萤Luciola substriata。形态观察发现,条背萤成虫橙黄色, 鞘翅末端灰黑色;发光器均为白色,雄虫发光器位于第5、6腹节,位于第5腹节的发光器呈带状,第6腹节的发光器呈“V”字形;雌虫的发光器呈带状,位于第5腹节;卵椭圆形,橙黄色。幼虫有两种形态,1～2龄具有7对呼吸鳃,3～6龄幼虫无呼吸鳃。幼虫具有一对发光器,位于第7腹节腹面;初蛹期仍保留幼虫形态的发光器,后呈现成虫的发光器,两种形态的发光器并存直至羽化。对条背萤生活史及习性调查发现,条背萤生活在水草较多的池塘、湖泊和流速缓慢的河流中。该虫1年发生1代,以幼虫在水中越冬,5月初老熟幼虫开始上岸化蛹。在25℃下,条背萤预蛹期平均为6.17天,蛹期平均为4.43天。成虫5月上旬至9月中旬发生。日落后的1 h内是条背萤成虫闪光求偶的最盛期。卵期平均12.5天。幼虫的猎物为静水椎实螺Lymnaea stagnalis,凸旋螺Gyraulus conwexiusculus等,天敌为克氏原螯虾Procambarus clarkii、中华绒毛蟹Eriocheir sinensis、草鱼Ctenopharyngodonidellus等。利用光谱仪对条背萤的发光光谱进行测定发现,条背萤的萤光光谱为425～603 nm,峰值为504 nm,颜色为黄绿混合色。     

Lampyridae (Coleoptera): A plethora of mollicute associations

Beetles (Coleoptera) harbor many species ofAcholeplasma andSpiroplasma (division Tenericutes, class Mollicutes). Mollicutes were isolated from guts and/or hemocoels of firefly beetles (Lampyridae) from the United States (Maryland and West Virginia), Ecuador, and Tobago. Firefly beetles were frequent hosts for the group XIV spiroplasma, isolated from Ellychnia corrusca, and the group XIX spiroplasma, isolated fromPhoturis spp. The most unusual feature of the firefly-mollicute association is the carriage of four Mycoplasma species. Recent phylogenetic studies indicate that these species are members of a clade that includes a vertebrate pathogen,Mycoplasma mycoides. The high rate of occurrence ofMycoplasma species (which are, otherwise, infrequent in insects) in lampyrid beetles suggests that the association is significant. The unusual light-producing physiology of lampyrids (which is dependent on large pools of energy) and the production of large amounts of cardenolides from cholesterol (a critical growth factor for many mollicutes) may favor colonization by mollicutes. Offprint requests to: K. J. Hackett.     

条背萤的闪光求偶行为

水栖萤火虫条背萤Luciola substriata (Gorh.)发出单脉冲周期性特异闪光信号进行求偶。室外观测发现,在23℃、86% RH时,雄萤飞行求偶闪光信号脉冲闪光持续时间为0.52 s,间隔时间为0.28 s,闪光信号的最大亮度为0.6 lx; 雌萤求偶信号周期为 0.67 s,最大亮度为0.4 lx。雄萤发出求偶信号0.22 s后,雌萤发出两个连续的回应信号。第一个回应信号为0.49 s,第二个为0.41 s, 两个回应信号的间隔时间为0.11 s。雄萤发现雌萤回应信号后,降落至离雌萤5～10 cm处,继续发出闪光信号,但闪光脉冲频率减小,闪光脉冲时间为1.23 s,间隔时间为0.50 s。条背萤交配时呈“V”或“一”字形交配姿势。     

胸窗萤的防卫行为：反射性出血及翻缩腺反应

为了探讨胸窗萤Pyrocoelia pectoralis Oliver, 1883的防卫行为, 对成虫的反射性出血行为及幼虫的翻缩腺体形态进行了研究。结果表明： 胸窗萤具有复杂的多重防卫行为策略。成虫利用闪烁、 假死、 反射性流血进行防卫, 幼虫则利用闪烁、 假死和微小的翻缩腺体进行防卫。反射性流血发生在雄成虫的前胸背板和鞘翅边缘, 而雌成虫仅在前胸背板边缘观察到有反射性流血行为。扫描电镜观察到直径大约32 μm的圆坑状结构环绕前胸背板或鞘翅边缘一圈。雌虫鞘翅牙也具有相似的圆坑状结构。幼虫具有9对乳头状的翻缩腺体, 位于中胸、 后胸和第1～7节侧板上。超薄切片发现, 腺体表面的棒状结构内部中空, 棒状结构连接一个发达的分泌细胞。细胞内连接棒状结构的部位着生致密的管状内质网。行为学实验发现, 胸窗萤血液对蚂蚁具有非常有效的拒避作用。     

Gene diversity and geographic differentiation in mitochondrial DNA of the Genji firefly, Luciola cruciata (Coleoptera: Lampyridae)

The Genji firefly, Luciola cruciata, is divided into two ecological types, the fast-flash and slow-flash types, on the basis of the interflash interval of mate-seeking males. To evaluate the evolutionary origin of the two types, 62 populations were examined by restriction fragment length polymorphism analysis of the mitochondrial cytochrome oxidase (CO) II gene. As a result, 19 haplotypes were detected, and their distributions were indigenous to local areas. Phylogenetic trees constructed from sequence comparison of the haplotypes revealed three major clades (I, II, and III). The boundary of haplotypes between clades I and II is approximately concordant with the geological structure of the Japanese Islands, which is a great rupture zone called the Fossa Magna, and the distribution of haplotypes in clades III and I-II corresponds to the Kyushu and Honshu-Shikoku Islands, respectively. The results suggest a vicariant scenario in which current L. cruciata diversity would have arisen from phylogenetic separations subsequent to the formation process of the Japanese Islands based on the molecular clock. The CO II gene trees also suggested that the fast-flash type should be considered an ancestral form, while the slow-flash type would be a derived one. The divergence time between the slow- and the fast-flash types is estimated to be about 4.6 to 2.0 mya (the Pliocene epoch).     

Composite,haustellate mouthparts in netwinged beetle and firefly larvae (Coleoptera,Cantharoidea: Lycidae,Lampyridae)

During lycid embryogenesis, labral and mandibular anlagen develop separately. Each anterolateral corner of the labrum elongates into a stylet, while the whole of each mandible involutes into a flute that is open to the lateral aspects. The stylets and flutes interlock in such a way as to leave canals, fashioning left and right haustellate apparatuses. During lampyrid embryogenesis, only one pair of anlagen forms instead of two. The pair matures into sharp, arcuate appendages that appear to be typical mandibles, but nevertheless a canal forms internally. The canal is not formed by coadaptation of two appendages; instead, cells within the pair create the canal autonomously. Indications are that the canal is homologous between these two families. It is suggested that in lampyrids, labral and mandibular cell fields develop together in the same anlage but behave according to their own respective programs, forming the canal with morphogenetic movements that are comparable to those of lycids. © 1994 Wiley-Liss, Inc.     

Aposematism and Bioluminescence: Experimental evidence from Glow-worm Larvae(Coleoptera: Lampyridae)

Bioluminescence most likely evolved under selection from the visually guided behaviours of co-occurring organisms, in particular that of predators. Many possible functions of light signals have been proposed and some are supported, but whatever their function may be, they make an easy target of the emitter unless it is defended. Therefore, we want to emphasise that in many cases bioluminescence can only have evolved through a defensive function. If this were the case, one would expect multimodal adaptiveness of luminescence with at least some evidence for a defensive function. Light signals could be used in many ways to reduce predation, but for spontaneous glowing species in particular, aposematism seems the only functional strategy. In a preliminary experiment with glowing and non-glowing dummy prey, we found that wild-caught toads discriminated against glowing prey. They showed significantly lower attack responses and higher latencies towards glowing prey dummies. However, some of the toads were less reluctant because they did not distinguish initially between prey with or without the light stimulus. Since the toads were collected in areas abundant with lampyrid glow-worms, which is the only luminous organism at this locality, and our results concur with the general evidence that they may have had previous experiences with this prey, we attribute the result to luminescent aposematism. From the literature, and from our own experiments, we know that toads and many other potential predators experience lampyrids as disagreeable prey. In future experiments we will test whether glow-worms are defended by luminescent aposematism or not. This revised version was published online in July 2006 with corrections to the Cover Date.     

The ecological impact of an introduced population on a native population in the firefly Luciola cruciata (Coleoptera: Lampyridae)

In Japan, Tatsuno Town has been famous for many Luciola cruciata fireflies emerging every summer at least since the 1920s. However, in the 1960s, L. cruciata fireflies were intentionally introduced from the Lake Biwa area into Matsuo-kyo, the most famous habitat of fireflies in that town. In this study, I examined ambient temperatures and flash rates of L. cruciata at four sites including Matsuo-kyo in the Tatsuno area and two sites in the Lake Biwa area. The linear regression of flash rates on temperatures indicated that the Matsuo-kyo population was distinct from the other three populations native to the Tatsuno area, but similar to the two populations native to the Lake Biwa area in terms of flash rates. These results were also supported by a recent molecular biological study, suggesting that the introduced fireflies had a strong ecological impact on the native ones at Matsuo-kyo. The present study emphasizes that we should not transport and release L. cruciata fireflies without careful consideration.     

The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae)

Through a phylogenetic analysis using adult morphological characters, we show that the origin of bioluminescence in cantharoid beetles appears to predate the origin of the family Lampyridae. The ability to produce and emit photic signals was first gained by larvae and appears to function as an aposematic warning display; it was subsequently gained in adults and is used as a sexual signal. Our analysis also suggests that while pheromonal sexual signals are used basally in the family, they are used in conjunction with and then subsequently replaced by photic signals in some lampyrid lineages. Both photic signals and the photic organs used to produce them have become greatly elaborated in the fireflies that no longer employ pheromonal sexual signals. In addition, the ability to produce a flashed sexual signal appears to have arisen at least three times in the family Lampyridae. Convergent evolution is also evident in a number of adult male photic organ morphologies. Further, we recommend that individual signal system components be compared rather than overall signal system complexity. The use of this strategy may allow one to recognize and better interpret adaptive correlations despite convergence or loss. We demonstrate that phylogenetic analysis is a powerful tool even for rapidly evolving traits.