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.     

Glow-worm larvae bioluminescence (Coleoptera: Lampyridae) operates as an aposematic signal upon toads (Bufo bufo)

It is an established fact that the spectacular bioluminescentdisplays of adult fireflies and glow-worms are used as courtshipsignals; however, the survival value of the glowing behaviorof their larvae remained the subject of speculation for manyyears. Our study is the first that demonstrates that lampyridlarvae use luminescence to signal unpalatability to nocturnal,visually guided predators. Wild-caught toads (Bufo bufo) weremore reluctant to attack luminescent artificial prey, and weshow that avoidance learning increased this reluctance. Afterbeing exposed to glow-worm larvae (Lampyris noctiluca), whichthe toads experienced as disagreeable, attack latencies to luminescentprey increased, but not those to nonglowing prey. Not all toadsshowed avoidance learning to the same extent, because of eitherdifferences in previous experience with glow-worms or differencesin memory.     

Flash behavior of femalePhoturis versicolor fireflies (Coleoptera: Lampyridae) in simulated courtship and predatory dialogues

FemalePhoturis versicolor fireflies attempt to capture males by responding to heterospecific flash patterns. A mating-dependent switch occurs which affects response timing and frequency of female flashes. We examined the switch using females of known age, mating status, and flash experience to assess how accurate mimicry is, what factors influence it, and what mechanism produces it. Presentations of simulated male flash patterns before and after mating revealed elements of an entrainment mechanism controlling female responsiveness. Unmated females preferentially answered conspecific patterns with variable latencies, averaging 1 s. Mating induced changes in both response frequency and response latency: Females answered heterospecific patterns more frequently, and latencies elicited by conspecific patterns shifted away from the unmated range. Heterogeneity in mean and variance of response latency among individuals indicates that females do not share a discrete reply to a given pattern. Little correspondence exists between latencies of sympatric species andP. versicolor females, suggesting that the flash response mechanism produces entriainment to any rhythmic pattern, not a one-to-one matching between prey and predator latencies. Different selective scenarios underlie strict mimicry versus entrainment mimicry.     

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.     

萤火虫的发光行为及其功能起源

发光生物因其活体能自发荧光而倍受关注,萤火虫(鞘翅目:萤科)是被研究最多的发光生物,因为成虫发光在其性选择以及求偶交配中起着重要作用.由于萤火虫的发光是一个消耗能量(ATP),的化学反应过程,因而其发光在成虫之外的虫态也具有重要意义.本文就萤火虫成虫和幼虫的发光行为、功能意义进行了综述,并结合萤火虫的饲养观察对其卵和蛹的发光行为进行了描述,探讨了卵和蛹的形态阶段发光的生物学功能以及生物荧光的起源进化.这将有助于理解萤火虫及其它生物自发荧光的本质和起源进化过程.     

The occurrence of synchrony in the North American fireflyPhotinus carolinus (Coleoptera: Lampyridae)

Synchronous flashing has been described for some Southeast Asian fireflies but has rarely been reported in North American fireflies. Our field observations indicated thatPhotinus carolinus flash synchronously. The flash pattern of individualP. carolinus was characterized by a burst of five to eight flashes over a period of approximately 4 s. These flash bursts were repeated about every 12 s. Groups of fireflies comprised individuals exhibiting this species-specific flash pattern. Remarkably, members of the group flashed synchronously: Flash bursts started and stopped at the same time and the flashes among individuals occurred at the same time as well. We used low-light level videography to examine this behavior in caged groups of (3 or 10) fireflies for synchronic flashing. The occurrence of concurrent rhythmic group flashing satisfies the criteria for synchrony as defined by Buck (1988). The intermittent nature of the bursts of flashes shown byP. carolinus makes this a discontinuous synchrony. The mechanisms underlying discontinuous synchrony are not known.     

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.     

Mechanisms of synchrony in the North American fireflyPhotinus carolinus (Coleoptera: Lampyridae)

Photometric recordings combined with computer stimulation, acquisition and analysis were used to study synchrony in the North American fireflyPhotinus carolinus. A computer-generated burst of simulatedP. carolinus flashes was used to trigger a firefly flash burst. We found that the first triggered firefly flashes occurred after the second or third flashes in the stimulus burst, that there were fewer flashes in a triggered burst than a spontaneous burst, and that extending the stimulus flashes into the firefly's interburst interval inhibited firefly flashing. When the stimulus flash interval (389–560 ms) was changed, no change was seen in the interflash interval. When the stimulus flash interval was changed, the average time between stimulus flash and firefly flash (flash delay) changed as if the firefly interflash interval was constant. Thus, interflash interval inP. carolinus does not change its length, making it similar to the Southeast Asian synchronizerPteroptyx cribellata and different fromPteroptyx malaccae, which can change its interval. We suspect that the time between bursts is functionally analogous to the time between flashes in Southeast Asian synchronizers.     

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.     

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.     

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,...     

Flash communication between the sexes of the firefly, Photuris lucicrescens

ABSTRACT. The courtship signal of the male firefly, Photuris lucicrescens Barber (Coleoptera, Lampyridae), is a brilliant crescendo flash which grows in intensity, reaches a peak and abruptly terminates. It was found to be triggered by a long neural burst from the brain. Males and females produce weak, twinkling flashes which induce male crescendo flashes. Female responses were triggered by a slowly rising intensity, and female response latency is therefore variable. Male and female P.lucicrescens fireflies produce two different types of flashes and both these flashes play an integral part in their courtship communication.     

Proximate Mechanisms of Sexual Selection in the Firefly Photinus pyralis (Coleoptera: Lampyridae)

Male competition in courtships of the firefly Photinus pyralis is intense. Morphometric analysis revealed that male size is variable, possibly exhibiting alternative body plans. We examined the outcomes of courtships to determine the proximate mechanisms of selection that may have led to male allometry. Two distinct selective events exist within mate competition: (1) searching and (2) scramble competition. Although many courtships failed, those without rivals were shorter and successful males spent less time in the mate-guarding position than when several males were in competition. Winners outflashed rivals, but flash dialogs occurred at similar frequencies across rival densities, indicating that females require some quantum of information before mating. Female flash preference was examined in choice tests to evaluate female discrimination ability. Females preferred flashes of greater intensity and precedence. This suggests that flash synchronization is a competitive display rather than a cooperative behavior to maintain female responsiveness. Selection for male size was density dependent: females preferred large males in searching, while small males were favored in scrambles. Simulated scrambles showed that small males have superior locomotor abilities, allowing them to reach the female rapidly. Comparison with a species lacking scrambles, P. macdermotti, revealed that selection in P. pyralis is disruptive, targeting traits alternately, and may have led to the evolution of higher allometric ratios in a species with direct male competition. Both male competition and female choice are important determinants of the outcome of P. pyralis courtships.     

Bioluminescence emissions of the firefly <Emphasis Type="Italic">Luciola praeusta</Emphasis> Kiesenwetter 1874 (Coleoptera: Lampyridae: Luciolinae)

We recorded the in vivo emission and time-resolved spectra of the firefly Luciola praeusta Kiesenwetter 1874 (Coleoptera: Lampyridae: Luciolinae). The emission spectrum shows that the full width at half maximum (FWHM) value for this particular species is 55 nm, which is significantly narrower than the in vivo half-widths reported till now. The time-resolved spectrum reveals that a flash of about 100 ms duration is, in fact, composed of a number of microsecond pulses. This suggests that the speed of the enzyme-catalysed chemiluminescence reaction in the firefly for the emission of light is much faster than was previously believed. A version of this article has already appeared in an archive Nature Precedings; see     

Determinants of reproductive success across sequential episodes of sexual selection in a firefly

Because females often mate with multiple males, it is critical to expand our view of sexual selection to encompass pre-, peri- and post-copulatory episodes to understand how selection drives trait evolution. In Photinus fireflies, females preferentially respond to males based on their bioluminescent courtship signals, but previous work has shown that male paternity success is negatively correlated with flash attractiveness. Here, we experimentally manipulated both the attractiveness of the courtship signal visible to female Photinus greeni fireflies before mating and male nuptial gift size to determine how these traits might each influence mate acceptance and paternity share. We also measured pericopulatory behaviours to examine their influence on male reproductive success. Firefly males with larger spermatophores experienced dual benefits in terms of both higher mate acceptance and increased paternity share. We found no effect of courtship signal attractiveness or pericopulatory behaviour on male reproductive success. Taken together with previous results, this suggests a possible trade-off for males between producing an attractive courtship signal and investing in nuptial gifts. By integrating multiple episodes of sexual selection, this study extends our understanding of sexual selection in Photinus fireflies and provides insight into the evolution of male traits in other polyandrous species.     

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.