Swarming and mating behaviour ofChironomus flaviplumus (Diptera: Chironomidae), compared with a sympatric congeneric species,C. yoshimatsui

The swarming and mating behaviour ofChironomus flaviplumus was observed and compared with a sympatric congeneric species,C. yoshimatsui. C. flaviplumus males swarmed around sunset near foliage or angles of buildings near the emergence site and copulated with females entering the swarm. Swarming and mating occurred under conditions of higher light intensity in cooler seasons than in warmer ones. Results suggested that temperature had an effect on the timing of flight to the swarming site in both sexes. TheC. flaviplumus swarm marker and swarming behaviour seemed very similar to that toC. yoshimatsui, and their respective daily swarming time zone greatly overlapped. No mixed swarm, however, was observed in the study area. This is probably due to the distance between the species' larval habitats. Possible premating isolation mechanisms between these 2 species are discussed.     

Age-associated male mating success in three swarming caddis fly species (Trichoptera: Leptoceridae)

Abstract. 1. In the three caddis fly species, Athripsodes albifrons (L.), A. cinereus (Curtis) and Mystacides azurea (L.) (Leptoceridae; Trichoptera), males swarm above the water surface of lakes or rivers. Receptive females fly to swarms and are chased and/or courted by males. After one of the swarming males has grasped an approaching female, the pair flies in tandem to the shore where they copulate.
2. In males, wing wear indices were negatively correlated with the ratio of fat/dry weight. In the only species in which comparisons were possible between newly emerged and swarming males (M. azurea), the former had significantly lower indices. Unmated females on average had lower wing wear indices than spent females. These facts suggest that wing wear reflects relative age.
3. The tandem males had significantly less wing wear than those in swarms, and are probably therefore younger. Age is therefore likely to be significant in relation to mating success.
4. Among males of the same relative age, tandem males had higher fat ratio than swarming ones, indicating that male mating success was also influenced by traits other than age. It is suggested that the shortest possible duration of the period of adult prematurity is adaptive, especially in insects with marginal adult food intake.     

Temporal dynamics of mating and predation in mosquito swarms

We determined the numbers of copulations and predatory attacks in swarms ofAnopheles freeborni (Diptera: Culicidae), and the distribution of these events throughout the duration of the swarming period each day. On 19 evenings of observation, we recorded 2724 copulating pairs leaving swarms and 1351 dragonfly (Pantala hymenaea andErythemis collocata) attacks. Mating activity partially coincided with predator activity. Most copulations occurred between 10 and 20 min after the swarms formed, while predation events were most frequent during the initial 15 min of the swarm. We calculated the ratio of copulations to predatory attacks during the swarming period. This ratio was significantly higher in an area sheltered by trees than it was in the open. We suggest that physiological and ecological constraints other than predation operate on the mating system of this anopheline to affect the timing of swarm initiation and swarm site selection.     

Mating system ofTokunagayusurika akamusi (Diptera: Chironomidae): I. Copulation in the air by swarming and on the ground by searching

Observations on the mating system of the midge,Tokunagayusurika akamusi, revealed mating to occur both in the air by swarming and on the ground by searching. At the shores of Lake Biwa, midges appeared from November to early December. Newly emerged adults arrived at the resting place, lakeside vegetation, in the morning, during which time a number of males also walked about in search of mates. Many copulating pairs were observed at the resting place. Huge swarms occurred chiefly before sunset but the frequency of copulation observed in the swarm was extremely low. It is likely that, in the Lake Biwa population, the proportion of females inseminated by searching males at the resting place was much larger than that by swarming males in the air. Furthermore, by searching, males copulated with younger females than by swarming. The differences between the searching and swarming tactics are discussed.     

Variation in energy reserves and role of body size in the mating system of Anopheles gambiae

Anopheles gambiae mates in flight. Males gather at stationary places at sunset and compete for incoming females. Factors that account for male mating success are not known but are critical for the future of any genetic control strategy. The current study explored variations in nutritional reserves (sugars, glycogen, lipids, and proteins) in wild‐caught swarming and resting males and evaluated the effect of body size and wing symmetry on male mating success. Our results showed that glycogen and sugar reserves are mobilized for flight. Males consume proportionally 5.9‐fold as much energy derived from sugars in swarming activities than when they are at rest. Mated males were on average bigger than unmated ones (P<0.0001). A strong correlation between the left and right wings in both mated and unmated males was found and additional analysis on fluctuating asymmetry did not show any indication of mated males being more symmetrical than unmated ones. The distribution of wing size of mated males was focused around a central value, suggesting that intermediate size of males is advantageous in the An. gambiae mating system. The results are discussed in the context of sexual selection.     

Swarming Behavior, Sexual Dimorphism, and Female Reproductive Status in the Sex Role-Reversed Dance Fly Species Rhamphomyia marginata

Dance flies are predaceous insects which often form male mating swarms. In many species males prior to swarming catch an insect prey, which is presented to the female at mating. In Rhamphomyia marginata, females in contrast to males gather to swarm, while males carrying a prey visit swarms for mating. Here I describe the swarming and courtship behavior in R. marginata and provide data on sexual dimorphism and swarming female reproductive status. Females swarm in small clearings in the forests. There was no specific swarm-maker. The swarming period lasted for 2–3 h and peaked around sunset. Identical swarm sites were used each evening and for several years. The mean number of females in swarms (swarm sites with at least one female) was 9.9 ± 9.1 (range, 1–40; n = 107) in 1993 and 7.1 ± 7.0 (range, 1–35; n = 68) in 1994. No obvious competition between females in swarms was observed. The operational sex ratio in swarms was extremely female biased (all swarms, 0.04). Less than one-third of male visits to swarms resulted in mating and males were found more often in larger swarms. Nuptial prey consisted of male midges. Females seem to mate more than once. Swarming females had undeveloped eggs, whereas mated females in swarms had further developed eggs than unmated females. Amount of sperm in the spermatheca was correlated with egg size. Amount of sperm and egg size did not correlate with wet weight, wing length, or wing load, except for egg size and weight. The wing coloration pattern and shape in R. marginata females are unique among dance flies, being greatly enlarged (1.6 times larger than that of males) and bicolored (gray part, 60% of wing area). When females, instead of males, possess extravagant secondary sexual characters, it is predicted from sexual selection theory that females should compete for males and that males should be selective in their choice of partner. A sex-role reversal will evolve when assess to males limit female reproductive success. The dance fly species R. marginata, like Empis borealis, another dance fly species studied earlier and discussed here, seems to fit these predictions.     

Wing length and asymmetry of male Tokunagayusurika akamusi chironomid midges using alternative mating tactics

Male Tokunagayusurika akamusi chironomids have alternative mating tactics.One is to search for females on vegetation (ground mating),and the other is to wait for females in an aerial swarm (swarmmating). Simultaneous sampling of ground-unpaired and ground-pairedmales and of swarm-unpaired and swarm-paired males were performed.The average wing length and right-left wing length difference(wing asymmetry) were compared between males from the four differentcategories. Swarm-unpaired males were larger than ground-unpaired ones,swarm-paired males were larger than swarm-unpaired ones, and ground-pairedmales were not larger than ground-unpaired ones. Thus, large malestended to aggregate in swarms, and larger swarming males matedmore successfully. On the other hand, small males probably enjoyedmating on the ground, especially when large males swarmed. Thewing asymmetry was not significantly different between unpairedand paired males both within and between tactics. There wasa flat or U-shaped relationship between wing length and asymmetry,underpinning the lack of a symmetrical advantage of swarmingto large males. The right-left difference was not normally distributedin four of six samples of unpaired males but, in contrast, wasnot normally distributed in only one of six samples of pairedmales. The non-normal distributions were leptokurtic and includedoutliers. Removal of the outliers improved normality, suggestingthat males with extremely asymmetric wings were not successfulin mating.     

Acoustic orientation of male Aedes diantaeus during mating

The reaction of males of Aedes diantaeus to different auditory stimuli was studied in natural environment during swarming and under laboratory conditions. The males were attracted by 250-420 Hz that corresponds to the main frequencies of flight tones of sympatric females and the range of sensitivity of male Johnston's organs. Behavioural data show that in natural environment swarming males are attracted by flying conspecific and heterospecific females rather than by dead or immobilized females. Contact with heterospecific females always ended in parting of mosquitoes while contact with cospecific females resulted in pairing. Thus, swarming males of Aedes diantaeus localize females by their flight tones but recognize the females of their own species in contact.     

Swarm Site Fidelity in the Sex Role-Reversed Dance Fly Empis borealis

In the dance fly species Empis borealis (L.), females (1–40) gather to swarm at landmarks (swarm markers, like trees and bushes), and males carrying an insect prey visit these swarms for mating. We noticed earlier that some swarm sites were used for several years and that they appeared to be frequented by a similar number of swarming females in each year, although the numbers of females varied greatly among swarm sites and certain sites attracted more swarming individuals than others. To explore swarm site fidelity in this mating system, in 1993 we monitored the same swarm sites that we studied in 1989, addressing the questions, Would the same swarm sites still attract the same number of females and males after 4 years? and Why do some swarm sites attract more displaying females than others? The number of females swarming at the different markers in 1993 was approximately the same as 4 years earlier. Some of these swarm sites are known to have been used for 18 years. The swarm sites with the largest number of flies had a high sun exposure during the day and were found at coniferous swarm marker trees and in a mixed forest habitat. A swarm site with few females attending and with a low amount of insolation during the day can be predicted to be abandoned as a swarming site soon. Empis borealis swarm sites thus persist over many years and are attended by a similar number of individuals each year. To our knowledge, such site fidelity has not been demonstrated for any swarming insect species earlier.     

Swarm behaviour and mate competition in mayflies (Ephemeroptera)

Although mayfly swarms are frequently cited as an example of lekking by insects, little is known about the behaviour of individuals within a swarm, or how mate-selection takes place. A study of five species of mayfly over a period of 10 consecutive years has revealed species-specific differences in the flight pattern of swarming males and in the ability of males to recognize swarms of their own species. Males of four of the five species jostle other males in the swarm at all times except when mating: mating pairs are not jostled. The pattern of jostling varies with the species. Measurements of the sperm content of the vesicula seminalis and of the wing length of members of individual swarms show that larger wing size is positively correlated with the presence of less sperm. The vesicula seminalis is always filled with sperm at the beginning of the imaginal stage and the testes regress before the beginning of the imaginal stage. If the volume of sperm in the vesicula seminalis is a valid index of mating success then males with larger wings have the highest success. Large wings may bestow an advantage during jostling. The males of Ephemera danica , which do not jostle, glide with outspread wings; these outspread wings may attract females, the largest wings being the most attractive. Females of all five species enter the swarm a few at a time, although many females may be resting beneath the swarm. This phased entry may decrease the attraction of the swarm for predators. The number of females in a swarm is not correlated with swarm size, and the factors which enable females to regulate their entry into a swarm remain obscure.     

Body size and mating success inDrosophila willistoni are uncorrelated under laboratory conditions

Mating activity and wing length were investigated in the F₁ progeny ofDrosophila willistoni females collected in the field to examine any possible relationship between body size and mating success. The flies were observed in a mating chamber under laboratory conditions. No significant differences in wing length were observed between copulating and noncopulating flies, and there was no significant correlation between wing length and copulation latency for both males and females. These results therefore suggest that the commonly accepted view that large body size is positively correlated with mating success inDrosophila does not always hold true. The results support the view that the extent of environmentally induced variation in body size may be an important factor in determining whether an association between body size and mating success is observed inDrosophila species.     

Effect of body size on swarming behavior and mating success of maleAnopheles freeborni (Diptera: Culicidae)

We examined whether body size affects the swarming behavior and mating success of male Anopheles freeborninear California rice fields. Swarms formed after dusk and persisted for approximately 30 min. The proportion of males in 33 swarms sampled n=6028 ranged from 100 to 92% but decreased over time (r=0.73, t=6.03, P<0.001).On average, swarming males (n=1058) were larger than males sampled from the resting population (n=735, H=35.6, P<0.0001),indicating that some males never swarm at all. Males swarming early were significantly smaller than those swarming during the peak (H=6.71, P=0.009)or final minutes of the swarm (H=4.86, P=0.002). Mated males returned to the swarm after mating, and larger males enjoyed greater mating success than did smaller ones (n=398, H=16.1, P=0.0005).     

Sexual dimorphism in relation to swarming and pair formation patterns in leptocerid caddisflies (Trichoptera: Leptoceridae)

North European Leptoceridae (Trichoptera) perform three types of swarming flight patterns: (1) swarming males of Athripsodesand Ceracleafly in horizontal zigzag patterns over the water surface, (2) the Mystacidesspp. perform vertical zigzag movements, and (3) the flight of males of Triaenodes unanimisMcLach. is a mixture of the horizontal and vertical zigzagging. Also three groups of pair formation behavior can be distinguished. In the first group, of Athripsodesand Ceraclea,the females fly into the male swarms, where they are grasped and carried to the riparian vegetation by the flying males with the females hanging upside-down in genitalia coupling. In the second group, a Mystacidesfemale is caught by a male, when approaching a swarm and both use their wings to fly in tandem to the shore where they copulate. In the third group, of Triaenodes bicolor(Curt.) and Oecetis lacustris(Curt.), the males fly searching for females sitting on aquatic plants and when a female is found the male lands and they copulate immediately while clinging to the plant. The different swarming and mating behaviors might have favored selection for three types of sexual dimorphism: (1) longer forewings in males than females in species which fly in copula, (2) larger eyes in males of the vertically zigzagging species, and (3) much smaller males in the group where males search for females sitting on aquatic plants. In the second group approaching females are detected by males before reaching the swarm and in the third group the female almost always mates with the male which is the first to find her. In conclusion, we suggest that females of Athripsodesand Ceracleahave a greater choice among swarming males than do females of Mystacides, T. bicolor,and O. Lacustris.     

Swarming and mating of Aedes communis mosquitoes in nature

The date of the beginning of mating behaviour in males and females, the rate of insemination and the increasing of bloodsucking activity of females were studied in natural environments. Over 80% of females mated on the 3-4th day after emergence; after fertilization their behaviour changed from looking for males for coupling to looking for ones for a prey. The male swarming began on the 5th day after emergence and simultaneously the appearance of inseminated females was observed. The places of mating of males and females were investigated. It was established that coupling took place in swarms with swarming males and out of swarms with freely flying males.     

Dietary constraints on sexual activity, mating success, and survivorship of male Delia antiqua

The effects of protein-deprivation on the sexual activity and reproductive fitness of male onion flies, Delia antiqua (Meigen) (Diptera: Anthomyiidae), were investigated under laboratory conditions. The percentage of males inseminating gravid females, the magnitude of ovipositional response, and the total numbers of eggs deposited in 1:1 or 1:10 male:female matchings over two days was unaffected by deprivation of dietary protein. The LT₅₀'s (median survival time) for solitary males provided proteinaceous, sucrose, or water diets were 38.0, 25.8, and 6.0 days, respectively. Yet independent of diet effects, males lost 50% of their wing tissue by fragmentation after 26 days, suggesting that wing condition is more important in determining male reproductive fitness than longevity. Male mating frequency in single pairings with previtellogenic females deprived of proteinaceous diet for ten days was similar to that of gravid, protein-fed females. In no-choice and choice mating bioassays at a 10:1 female:male ratio, however, males inseminated significantly fewer previtellogenic than gravid females over 24 h. Despite evidence for male autogeny, removal of exogenous protein resources in the Allium agroecosystem may have important effects on the reproductive competency and fecundity of D. antiqua.     

Widespread 'hilltopping' in Acraea butterflies and the origin of sex-role-reversed swarming in Acraea encedon and A. encedana

In some populations of the butterflies Acraea encedon and A. encedana, most females are infected with a bacterium that kills their sons. The resulting shortage of males is associated with females adopting a sex‐role‐reversed mating system, in which females swarm at landmarks such as hilltops and compete for males. We have observed the mating behaviour of Acraea species that are not known to be infected with the male‐killer. In over half of these species, males were found to aggregate on hilltops. It is likely that this behaviour was ancestral to the sex‐role‐reversed swarms of Acraea encedon and A. encedana, and we discuss how the spread of the male‐killing infection may have converted this mating system into sex‐role‐reversed swarming.     

Swarming and mating activity of Anopheles gambiae mosquitoes in semi‐field enclosures

Anopheles gambiae Giles sensu stricto (Diptera: Culicidae) is the major Afro‐tropical vector of malaria. Novel strategies proposed for the elimination and eradication of this mosquito vector are based on the use of genetic approaches, such as the sterile insect technique (SIT). These approaches rely on the ability of released males to mate with wild females, and depend on the application of effective protocols to assess the swarming and mating behaviours of laboratory‐reared insects prior to their release. The present study evaluated whether large semi‐field enclosures can be utilized to study the ability of males from a laboratory colony to respond to natural environmental stimuli and initiate normal mating behaviour. Laboratory‐reared males exhibited spatiotemporally consistent swarming behaviour within the study enclosures. Swarm initiation, peak and termination time closely tracked sunset. Comparable insemination rates were observed in females captured in copula in the semi‐field cages relative to females in small laboratory cages. Oviposition rates after blood feeding were also similar to those observed in laboratory settings. The data suggest that outdoor enclosures are suitable for studying swarming and mating in laboratory‐bred males in field‐like settings, providing an important reference for future studies aimed at assessing the comparative mating ability of strains for SIT and other vector control strategies.     

Genetic structure in a swarming brown long-eared bat (<Emphasis Type="Italic">Plecotus auritus</Emphasis>) population: evidence for mating at swarming sites

Plecotus auritus, a small, gleaning bat species, lives in small, isolated summer colonies in which both males and females show a high degree of natal philopatry. Despite this, colonies have high gene diversities and low inbreeding coefficients. It has been suggested that inbreeding is avoided because mating occurs during autumnal and spring swarming at hibernation sites. We tested this hypothesis by comparing microsatellite profiles, based on eight loci, of bats from six summer colonies and two swarming sites they were known to visit from radiotelemetry studies. We found high gene diversities (H _s = 0.77) at both swarming sites and summer colonies which were not statistically different. There was no detectable isolation by distance and F_ST was low (0.001). Together, these results suggest high gene flow between sites. Despite this, there was small but significant genetic differentiation amongst summer colonies and between summer colonies and the primary swarming site. We suggest that swarming is important for gene flow and for maintaining genetic diversity in this highly philopatric species and discuss possible reasons for the genetic differentiation observed. The identification and protection of swarming sites should be a major conservation priority for this and other temperate bat species.     

Effects of supraoptimal temperatures on the myxomycetePhysarum polycephalum

Vibrio parahaemolyticus, the flagellated nonswarming marine bacteria were induced to swarm on solid media under three different conditions: growth at 20–26°C on medium containing 1% NaCl, growth on a medium in a sealed Petridish and growth on H₂O₂-treated medium. The morphological transformations observed in cells during swarming of V. parahaemolyticus are similar to those found jor the naturally swarming Vibrio alginolyticus. The mechanism of swarming in both species involves massive formation of peritrichous flagella and a negative chemotactive response to metabolic byproducts.     

Swarming mechanisms in the yellow fever mosquito: aggregation pheromones are involved in the mating behavior of Aedes aegypti

Mosquitoes of various species mate in swarms comprised of tens of thousands of flying males. In this study, we examined Aedes aegypti swarming behavior and identified associated chemical cues. Novel evidence is provided that Ae. aegypti females aggregate by means of olfactory cues, such as aggregation pheromones. Isolation of Ae. aegypti aggregation pheromones was achieved by aeration of confined mosquitoes and collection of associated volatiles by glass filters. The collected volatiles were identified through gas chromatography mass spectrometry (GCMS). Three aggregation pheromones were collected and identified as 2,6,6‐trimethylcyclohex‐2‐ene‐1,4‐dione (ketoisophorone) (CAS# 1125–21–9, t_R = 18.75), 2,2,6‐trimethylcyclohexane‐1,4‐dione (the saturated analog of ketoisophorone) (CAS# 20547–99–3, t_R = 20.05), and 1‐(4‐ethylphenyl) ethanone (CAS# 937–30–4, t_R = 24.22). Our biological studies revealed that the identified compounds stimulated mosquito behavior under laboratory conditions. The mechanism of mosquito swarm formation is discussed in light of our behavioral study findings. A preliminary field trial demonstrated the potential application of the isolated aggregation pheromones in controlling Ae. aegypti.