萤火虫(鞘翅目:萤科)两性交流中的闪光信号

对国内外萤火虫两性交流闪光信号的研究进行了综述,萤火虫发光器因种而异,多数发出黄绿色萤光,闪光信号的频率、光谱、强度及其时空分布的闪光模式包含着两性交流信息。萤火虫闪光交流系统有两种分类方法,其一是萤火虫具两个类型的闪光信号交流系统,及系统和系统,前者多在旧大陆,后者多在新大陆;其二是萤火虫具6个类型闪光信号交流系统,即HP,LL,LC,PR,CR和LB型,其中PR型与系统相对应,HP型与系统对应。萤火虫两性交流闪光信号常因时间和空间上的差异及外界物体的干扰使两性闪光交流的效率受到影响。萤火虫两性交流的闪光信号起源于鞘翅目的幼虫阶段,并起警戒天敌的作用,经过两性选择成为成虫两性交流的一种途径,进而成为新大陆的一些萤火虫间捕食猎物和逃避天敌的生存策略。     

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.     

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。     

Tuning of photoreceptor spectral sensitivity in fireflies (Coleoptera: Lampyridae)

Sexual communication between male and female fireflies involves the visual detection of species-specific bioluminescent signals. Firefly species vary spectrally in both their emitted light and in the sensitivity of the eye, depending on the time when each is active. Tuning of spectral sensitivity in three firefly species that occupy different photic niches was investigated using light and electron microscopy, microspectrophotometry, and intracellular recording to characterize the location and spectral absorption of the screening pigments that filter incoming light, the visual pigments that receive this filtered light, and the visual spectral sensitivity. Twilight-active species had similar pink screening pigments, but the visual pigment of Photinus pyralis peaked near 545 nm, while that of P. scintillans had a λ_max near 557 nm. The night-active Photuris versicolor had a yellow screening pigment that was uniquely localized, while its visual pigment was similar to that of P. pyralis. These results show that both screening and visual pigments vary among species. Modeling of spectral tuning indicates that the combination of screening and visual pigments found in the retina of each species provides the best possible match of sensitivity to bioluminescent emission. This combination also produced model sensitivity spectra that closely resemble sensitivities measured either with electroretinographic or intracellular techniques. Vision in both species of Photinus appears to be evolutionarily tuned for maximum discrimination of conspecific signals from spectrally broader backgrounds. Ph. versicolor, on the other hand, appears to have a visual system that offers a compromise between maximum sensitivity to, and maximum discrimination of, their signals. Accepted: 29 September 1999     

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.     

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。     

Contributions to molecular systematics of water scavenger beetles (Hydrophilidae,Coleoptera)

Phylogenetic relationships within Hydrophilidae were examined by analyses of separate and combined nuclear and mitochondrial markers (28S rRNA, 18S rRNA, 16S rRNA, 12S rRNA, COI and COII genes). The preferred (Bayesian) tree topology suggests a sister group relationship between Spercheidae and Hydrophilidae, supporting the ‘hydrophilid lineage’; Epimetopidae are placed on the base of the ‘helophorid branch’, the monophyly of Sphaeridiinae is highly supported, nested deeply within Hydrophilidae closest to Enochrus, making Hydrophilinae and Acidocerini paraphyletic; Hydrobius appears as sister taxon to (Hydrochara + Hydrophilus) without a closer relationship to Acidocerini; the hydrophiloid–histeroid sister group relationship is confirmed. The topology of several taxa remains contradictory, and requires further investigations with a larger taxon sampling and additional molecular markers.     

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.     

Vicariance of Pyrocoelia fireflies (Coleoptera: Lampyridae) in the Ryukyu islands,Japan

We performed molecular phylogenetic analyses based on the mitochondrial COI gene (687 bp) and the nuclear 28S rRNA gene (715 bp) and reconstructed phylogenetic trees of the Pyrocoelia fireflies in the Ryukyu Islands and eastern Asia. Age calibration was done using a robust geological constraint: the Okinawa trough and associated straits began to rift at 1.55 Ma, isolating the Ryukyu Islands from the Chinese continent, Japanese islands, Taiwan island and some of the islands from each other. We suggest that the physical isolation of these islands began to generate the allopatric speciation within these islands, so the timing of this isolation was assigned to an appropriate node. The topology is completely concordant among phylogenetic trees reconstructed using MEGA (maximum‐likelihood), raxmlGUI (maximum‐likelihood) and BEAST (Bayesian inference; including combined analysis of COI and 28S rRNA genes). Two lineages are recognized, related to their emergence time; spring to summer, and autumn. In each lineage, vicariance is inferred to have begun at 1.55 Ma from our phylogenetic and geological analyses. In lineage 1, P. oshimana (Amami), P. matsumurai (Okinawa), P. discicollis (W. Japan), P. fumosa (E. Japan) and P. abdominalis (Yaeyama) were differentiated. In lineage 2, P. rufa (Tsushima and Korea), P. miyako (Miyako‐jima), P. atripennis (Ishigaki‐jima) and P. praetexta (Taiwan and HongKong) were differentiated. Pyrocoelia analis (Taiwan and China) emerges throughout the year except for winter, and constitutes another lineage. We suggest that Pyrocoelia fireflies differentiated at 2 Ma to generate these three lineages. The base substitution rate for the COI gene is estimated as 4.48% Myr^–1 and that for the 28S rRNA gene is 0.394%  Myr^–1, and these rates were used in a combined BEAty analysis in BEAST.     

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.     

Spectral sensitivity of the compound eyes in two day-active fireflies (Coleoptera: Lampyridae: Lucidota)

Summary The electroretinographic visual spectral sensitivity functions in day-active fireflies Lucidota luteicollis and Lucidota atra show a broad green sensitivity and a shoulder in the near-ultraviolet region of the spectrum (Figs. 1, 2) as is commonly found among day-active insects. The nomogram for P530 visual pigment matches the spectral sensitivity curves in the green. The adult L. luteicollis retains its larval bioluminescent light organ which has a peak emission at 562 nm. The _max of the ERG spectral sensitivity does not match the bioluminescent peak (Fig. 1B) as it does in twilight- and dark-active fireflies. Some relevant behavioural observations with respect to mating are presented.     

Phylogeny of North American fireflies (Coleoptera: Lampyridae): implications for the evolution of light signals

Representatives of the beetle family Lampyridae ("fireflies", "lightningbugs") are well known for their use of light signals for species recognition during mate search. However, not all species in this family use light for mate attraction, but use chemical signals instead. The lampyrids have a worldwide distribution with more than 2000 described species, but very little is known about their phylogenetic relationships. Within North America, some lampyrids use pheromones as the major mating signal whereas others use visual signals such as extended glows or short light flashes. Here, we use a phylogenetic approach to illuminate the relationships of North American lampyrids and the evolution of their mating signals. Specifically, to establish the first phylogeny of all North American lampyrid genera, we sequenced nuclear (18S) and mitochondrial (16S and COI) genes to investigate the phylogenetic relationships of 26 species from 16 North American (NA) genera and one species from the genus Pterotus that was removed recently from the Lampyridae. To test the monophyly of the NA firefly fauna we sequenced the same genes from three European lampyrids and three Asian lampyrids, and included all available Genbank data (27 additional Asian lampyrids and a former lampyrid from Asia, Rhagophthalmus). Our results show that the North American lampyrids are not monophyletic. Different subgroups are closely related to species from Europe, Asia and tropical America, respectively. The present classification of fireflies into subfamilies and tribes is not, for the most part, supported by our phylogenetic analysis. Two former lampyrid genera, Pterotus and Rhagophthalmus, which have recently been removed from this family, are in fact nested within the Lampyridae. Further, we found that the use of light as a sexual signal may have originated one or four times among lampyrids, followed by nine or four losses, respectively. Short flashes originated at least twice and possibly three times independently among our study taxa. The use of short flashes as a mating signal was replaced at least once by the use of long glows, and light signals as mating signals were lost at least three times in our study group and replaced by pheromones as the main signal mode.     

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.     

Correlated evolution of female neoteny and flightlessness with male spermatophore production in fireflies (Coleoptera: Lampyridae)

The beetle family Lampyridae (fireflies) encompasses ～100 genera worldwide with considerable diversity in life histories and signaling modes. Some lampyrid males use reproductive accessory glands to produce spermatophores, which have been shown to increase female lifetime fecundity. Sexual dimorphism in the form of neotenic and flightless females is also common in this family. A major goal of this study was to test a hypothesized link between female flight ability and male spermatophore production. We examined macroevolutionary patterns to test for correlated evolution among different levels of female neoteny (and associated loss of flight ability), male accessory gland number (and associated spermatophore production), and sexual signaling mode. Trait reconstruction on a molecular phylogeny indicated that flying females and spermatophores were ancestral traits and that female neoteny increased monotonically and led to flightlessness within multiple lineages. In addition, male spermatophore production was lost multiple times. Our evolutionary trait analysis revealed significant correlations between increased female neoteny and male accessory gland number, as well as between flightlessness and spermatophore loss. In addition, female flightlessness was positively correlated with the use of glows as female sexual signal. Transition probability analysis supported an evolutionary sequence of female flightlessness evolving first, followed by loss of male spermatophores. These results contribute to understanding how spermatophores have evolved and how this important class of seminal nuptial gifts is linked to other traits, providing new insights into sexual selection and life-history evolution.     

Modification of spectral sensitivities by screening pigments in the compound eyes of twilight-active fireflies (Coleoptera: Lampyridae)

1. ERG S(lambda) were determined in dark-adapted intact preparations of 6 North American firefly species (Photinus collustrans, marginellus, pyralis, macdermotti, scintillans and Bicellonycha wickershamorum) which restrict their flashing activity to twilight hours. The curves possess narrow (1/2 bandwidth = 50-60 nm) peaks in the yellow (560-580 nm) and a shoulder in the violet (370-420 nm), with a marked attenuation (1.4-2.2 log units) of sensitivity in the green (480-530 nm) region of the spectrum (Fig. 1). Two additional species (Photuris potomaca and frontalis) which initiate flashing at twilight and continue on late into the night (twi-night) possess broad sensitivity maxima around 560 nm (Fig. 3). 2. Selective adaptation experiments isolated near-UV and yellow in P. scintillans (Fig. 2). In the dorsal frontal region of the compound eyes in P. frontalis, high sensitivity existed only in the short wavelength region (near-UV and blue) with a maximum in the blue (lambda max 435 nm) (Fig. 4). 3. The in situ MSP absorption spectrum of the screening pigments was determined in preparations of firefly retina. a) Two kinds of dark brown granules were found in the clear zone region. These granules absorb all across the spectrum with a gradual increase in optical density in the shorter wavelength region in P. pyralis (Fig. 5). b) Besides dark granules, pink-to-red colored screening pigments were present in the vicinity of the rhabdoms. The absorption spectra of these pigments determined in five species were narrow (1/2 bandwidth = 50-80 nm) with species-specific differences in their peak absorption in the green at 525 nm, 510 nm, 512 nm and 517 nm in P. scintillans, macdermotti, collustrans and pyralis, respectively (Fig. 6). A similar pigment was found in P. marginellus with a lambda max at 512 nm (Fig. 7). In all cases, transmission increased both at long and short wavelengths, but more sharply in the long wavelength region (Figs. 6 and 7). Hence each twilight-restricted species has its own unique colored screening pigment. A yellow pigment whose absorption spectrum differed from those found in genus Photinus was found in twi-night active Photuris potomaca (lambda max 461 nm) and night-active P. versicolor (lambda max 456 nm). The transmission of the Photuris pigment increased sharply only in the long wave-length region (Fig. 8).(ABSTRACT TRUNCATED AT 400 WORDS)     

Ulva flexuosa (Ulvaceae,Chlorophyta) inhabiting inland aquatic ecosystems: molecular,morphological and ecological discrimination of subspecies

Species of Ulva (Ulvaceae, Chlorophyta) are relatively common macroalgae in most coastal areas. In this paper, we examine Ulva in inland aquatic ecosystems to determine relationships between the chemical parameters of the water and the Ulva taxa present. Two groups of samples of freshwater Ulva were selected from: (i) habitats with high salinity levels, and (ii) habitats with fresh or slightly saline waters. We showed that these two groups of Ulva, which occupy distinct habitats, represent two separate taxa. The first group represents Ulva flexuosa subsp. pilifera (Kützing) M.J. Wynne, which is very common and regularly encountered in fresh waters, but is equally well represented in slightly saline waters. The second is U. flexuosa subsp. paradoxa (C. Agardh) M.J. Wynne, which is less common, preferring mainly salt-wedge saline waters. We did not find both Ulva flexuosa subspecies together at any of the research sites.

There were significant morphological differences between the Ulva flexuosa subspecies. Ulva flexuosa subsp. pilifera had longer and wider thalli and wider cells than those of subsp. paradoxa. Ulva flexuosa subsp. paradoxa cells were longer and of greater circumference than those of subsp. pilifera. Analyses of the rRNA ITS region and partial rbcL gene sequences demonstrated minor sequence variation within the two subspecies. The results of this study provide detailed information on the characteristics of limnic Ulva habitats and indicate the niches where such populations are likely to occur.     

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.     

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.     

Comparative morphology study of the male genitalia in the tribe Astathini from China (Coleoptera: Cerambycidae)

The male genitalia of 13 species from four genera of Astathini were described and analyzed. The result showed that five genital characters, such as shape of the apex of 8th abdominal tergaum and sternum, ratio of the length of lateral lobes to tegmen, can be used to identify genera of Astathini; six characters, such as ratio of the length of lateral lobes to tegmen, ratio of the length of roof to lateral lobes, shape of the apex of ventral plate of median lobe, can be used to identify species in Bacchisa.