Predator-prey interactions on the wing: aerobatics and body size among dance flies and midges

Male dance flies (Diptera: Empididae) are major predators of mating swarms of male nonbiting midges (Diptera: Chironomidae). Studies have indicated that chironomid midges are excellent fliers with extraordinarily high wing muscle power, acceleration and tortuosity for their size. We compared the relative flight performance and morphometrics of male empidsEmpis tesselatta Meigen and their prey, Chironomus plumosus L., in a series of laboratory and field studies. High-speed digital video camera footage revealed that there were no significant differences between predator and prey in terms of straight-line acceleration or maximum flight velocity, although the mean speed of empids was higher. In curved flight, measured as tortuosity, the smaller chironomids held a considerable advantage over their larger predators. Flight muscle ratio (e.g. flight muscle mass/body mass) of chironomids was also greater in chironomids than in empids. In field studies, small male chironomids suffered much lower levels of predation than large males. This result is consistent with the prediction of small midges being less likely to be captured because of superior aerobatics, although we are unable to exclude empirically that empids select large male midges. We discuss the possible mechanisms of prey capture by empids given the greater manoeuvrability of their prey, the coevolutionary dynamics of flight performance and the apparent selective advantage of small size in male chironomids. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Fitness effects of parasite-mediated spatial heterogeneity within a swarm

The fitness consequences associated with the position an individualadopts within a dynamic group are not well understood. I investigatedmate acquisition by male chironomid midges using a simple swarmingmodel and empirically collected data on midge aerobatic ability.Previous work has suggested that the aerobatic ability of amale is an important predictor of his reproductive success,although there is contrary (and counterintuitive) evidence thatinfection with ectoparasitic mites increases a male's chanceof mating, despite having negative effects on flight speed.The model used here suggests that a male's location within theswarm, brought about passively through interindividual differencesin flight speed, may explain these contradictory results. Specifically,slower flying males (including those burdened with mites) adoptedpositions nearer the center of the swarm, whereas faster malestended to occupy the periphery. This in turn affected theiraccess to females because any mechanism that brought femalesnearer the swarm's center before capture (including high femaleflight speed and selective pairing by either males or females)significantly increased the relative reproductive success ofboth larger and parasitized males, with the benefits of parasitismpeaking at around 4 mites per host. There may be selective pressureon hosts and parasites to maintain this optimal mite densitybecause both are likely to benefit from the relationship: hostsenjoy an increased reproductive success, whereas only throughhost copulation can mites transfer to a female midge and returnto water (their next life-history stage) during host oviposition.     

SEXUAL SELECTION AND BUTTERFLY DESIGN—A COMPARATIVE STUDY

Temperate butterflies of 44 species were examined to determine if their mating system (perching and patrolling) affected flight design. To control for spurious effects due to ancestry, 25 of these species were assigned to eight contrasts within which a change in mating system had occurred. In perching species sexual selection was predicted to favor traits associated with high acceleration ability and speed, while in patrolling species traits associated with flight endurance were predicted. In conformance with these expectations males of perching species had larger thorax/body mass ratios, higher wing loadings, and higher aspect ratios than patrolling species. The male mating system affected females in the same direction in the same variables as males. This could be explained by a genetic correlation with males. When removing the covariance between the sexes, only male design was explained by the mating system. The mating system was also associated with different degrees of sexual dimorphism in wing size. This supported the hypothesis that male design was affected by the mating system.     

Interactions Between Swarming Chironomus annularius (Diptera: Chironomidae) Males: Role of Acoustic Behavior

Males of swarming species of chironomids use their auditory system (Johnston's organs) to recognize a female within swarm and do not respond to male flight tones. However, in some cases the male–male interactions were observed at a high frequency. The role of acoustic behavior in this phenomenon in C. annularius was studied. The results showed that male Johnston's organs were sensitive to male flight tones from a distance of about 1–1.5 cm. The carrier frequencies of these sounds negatively correlated with male body size. Thus we would expect that male–male interactions will occur mainly between large males. Nonetheless, the analysis of caught pairs revealed that in both male–male and female–male interactions small males had an advantage. The ability of males to perceive the male flight tones is discussed with respect to swarming behavior and mating success.     

Is bigger better? Male body size affects wing‐borne courtship signals and mating success in the olive fruit fly,Bactrocera oleae (Diptera: Tephritidae)

Variations in male body size are known to affect inter‐ and intrasexual selection outcomes in a wide range of animals. In mating systems involving sexual signaling before mating, body size often acts as a key factor affecting signal strength and mate choice. We evaluated the effect of male size on courtship displays and mating success of the olive fruit fly, Bactrocera oleae (Diptera: Tephritidae). Wing vibrations performed during successful and unsuccessful courtships by large and small males were recorded by high‐speed videos and analyzed through frame‐by‐frame analysis. Mating success of large and small males was investigated. The effect of male–male competition on mating success was evaluated. Male body size affected both male courtship signals and mating outcomes. Successful males showed wing‐borne signals with high frequencies and short interpulse intervals. Wing vibrations displayed by successful large males during copulation attempt had higher frequencies over smaller males and unsuccessful large males. In no‐competition conditions, large males achieved higher mating success with respect to smaller ones. Allowing large and small males to compete for a female, large males achieve more mating success over smaller ones. Mate choice by females may be based on selection of the larger males, able to produce high‐frequency wing vibrations. Such traits may be indicative of “good genes,” which under sexual selection could means good social‐interaction genes, or a good competitive manipulator of conspecifics.     

Optimal progeny body size in a solitary bee,Osmia bicornis (Apoidea: Megachilidae)

1. Females of solitary, nest‐constructing bees determine both sex (haplo‐diploidy) as well as body size (by amount of provision) of each single offspring. 2. According to the Optimal Allocation Theory, females should allocate resources in portions that maximise fitness returns. A key fitness component in bees is body size that is determined solely by the provisions supplied by the mother. 3. The optimal progeny body size relies on different factors in both sexes. In females, provision efficiency is crucial for reproductive success. In males, however, fitness depends primarily on mating success. 4. Provision efficiency of Osmia bicornis L. females depends on the capacity to stow pollen (scopa) and their ability to carry the packed loads. Scopa capacity increases isometrically with body size whereas indices of flight performance (EPI, free lift) decrease. These complementary effects substantially contribute to the adjustment of optimal body size in daughters. 5. The impact of body mass on fitness of males is determined by the mating system. Owing to the opportunistic polygyny in O. bicornis, there was no detectable correlation between body size and male mating success. Consequently, mother bees distribute their provisions to many, but small sons to increase the number of descendant competitors in the race for matings. Optimal body size in sons is a trade‐off between a large male advantage in rare scrambles over receptive females and small‐size‐related disadvantages in viability.     

Calling,Courtship, and Condition in the Fall Field Cricket,Gryllus pennsylvanicus

Theoretically, sexual signals should provide honest information about mating benefits and many sexually reproducing species use honest signals when signalling to potential mates. Male crickets produce two types of acoustic mating signals: a long-distance mate attraction call and a short-range courtship call. We tested whether wild-caught fall field cricket (Gryllus pennsylvanicus) males in high condition (high residual mass or large body size) produce higher effort calls (in support of the honest signalling hypothesis). We also tested an alternative hypothesis, whether low condition males produce higher effort calls (in support of the terminal investment hypothesis). Several components of long-distance mate attraction calls honestly reflected male body size, with larger males producing louder mate attraction calls at lower carrier frequencies. Long-distance mate attraction chirp rate dishonestly signalled body size, with small males producing faster chirp rates. Short-range courtship calls dishonestly reflected male residual mass, as chirp rate and pulse rate were best explained by a curvilinear function of residual mass. By producing long-distance mate attraction calls and courtship calls with similar or higher effort compared to high condition males, low condition males (low residual mass or small body size) may increase their effort in current reproductive success at the expense of their future reproductive success, suggesting that not all sexual signals are honest.     

No Evidence for Significant Effect of Body Size and Age on Male Mating Success in the Spot-legged Treefrog

In anurans, body size and age of individuals generally affect male mating success. To test whether body size and age have effects on male mating success in the foam-nesting treefrog Polypedates megacephalus, a species widely distributed in China, we analyzed differences in body size and age between mated and unmated males for three populations using a Generalized Linear Mixed Model(GLMM). The results showed that mated males did not exhibit larger body size and older age than unmated males, suggesting that large and/or old male individuals did not have greater mating success than small and/or young males. Moreover, we also found a non-significant size-assortative mating pattern for all populations. Our findings suggest that body size and age of the foam-nesting treefrog do not affect male mating success.     

Male size,mating potential and lifetime reproductive success in the field cricket,Gryllus bimaculatus (De Geer)

Male size is known to influence short-term, competitive mating success in the field cricket, Gryllus bimaculatus. Short-term measures of mating success are, however, potentially misleading when considering the influence of sexual selection on male morphology and behaviour. The lifetime mating success of males was investigated in the absence of competition to determine the effects of male size per se on reproductive potential and any effects of female behaviour on male lifetime mating success. The cost of spermatophore production was relatively greater for small males who consequently had a longer refractory period between matings, thus limiting their reproductive potential. Large males had a mating advantage resulting from a differential response of females to the courtship attempts of large and small males. Furthermore, small males were ‘sexually incompetent’ in terms of their ability to attach spermatophores once mounted. There was, therefore, an overall positive relationship between male size and lifetime reproductive success.     

Aerial Contests, Sexual Selection and Flight Morphology in Solitary Pompilid Wasps

Aerial contest competition has proven to be a challenging phenomenon to interpret in many territorial insects. Because the duels often consist of elaborate and/or high speed ascending maneuvers, the hypothesis that they are settled due to asymmetries in flight performance is intuitively appealing. We evaluated this hypothesis by contrasting differences in known morphological determinants of flight performance between (1) residents vs. non-residents of the territorial wasp, Hemipepsis ustulata and between (2) H. ustulata vs. a non-territorial relative, Pepsis thisbe . In the first contrast, resident male H. ustulata were seen to be larger, and had a tendency for reduced wing loading, but they did not possess greater flight musculature or wing aspect ratios (i.e., more elongated wings) than their non-resident counterparts. In the second contrast, male H. ustulata exhibited clearly greater flight musculature and greater sexual dimorphism in this parameter (males more muscular), and also exhibited a slight tendency for greater wing loading and smaller aspect ratios than males of the patrolling species P. thisbe . Interestingly, although size is linked with territorial success in H. ustulata , males of this species were not larger than male P. thisbe , nor did the former species exhibit greater sexual size dimorphism. These results do not support the hypothesis that the repeated ascending contests of H. ustulata require, and select for, a high acceleration design. However, the observed intraspecific patterns of flight musculature suggest that high acceleration is favored in males of the perching species, perhaps for the ability to intercept passing receptive females.     

Virgin and recently mated males are equally able to achieve new matings in Neoleucinodes elegantalis

Lepidopteran male mating success is recognized to be directly related to physical and behavioural traits such as ability, vigour, activity and persistence in courtship. In the tomato fruit borer Neoleucinodes elegantalis Guenée, the mating system is known to be monogamic and therefore males' sexual investment is apparently low. The hypothesis that recently mated males have a remating probability equal to that of virgin males is tested. The impact of body size in remating success and the cost of remating are also analyzed. Mated males show as much propensity to remate as naïve ones. Copula duration and the time taken to copulate are similar in mated and virgin males. However, spermatophore size is not related to male size. The results suggest the ability of N. elegantalis males to remate within 24 h between mating events and their propensity to remate are not affected by copula investment. © 2013 The Royal Entomological Society     

Male size, sperm transfer, and colony fitness in the western harvester ant, Pogonomyrmex occidentalis

Mating success in the western harvester ant, Pogonomyrmex occidentalis, increases with male size. We tested the hypothesis that increased mating success increases male fitness and the fitness of colonies that make large males by comparing the sperm content of males prior to and at the conclusion of the mating swarm. The number of sperm a male initially possesses is a function of male size, and large males transfer a greater proportion of their sperm than do small males. For colonies, the payoff per unit of investment is an increasing function of male size, and investment in large males is not equivalent to investing in a larger number of small males. Allocation ratios in species that show size variation in reproductives may need to be modified by the individual fitness functions.     

Females decide whether size matters: plastic mate preferences tuned to the intensity of male-male competition

Recent studies have indicated that mating success of large malesmay improve under increasing levels of mating competition. Thisoutcome is explained 1) if male mating competition is overridingfemale preferences for male traits that are unrelated to, ornegatively correlated with, male size and dominance and, inso doing, dictates the distribution of matings or 2) if femalesalter their preferences with respect to large males when male–malecompetition is intense. Under both hypotheses, one could expectlarge, dominant males to be more successful under intense competitionthan under weak competition. However, only the first explanationpredicts that male mating success under intense competitionshould be determined by dominance; traits that are unrelatedto male dominance should be uncorrelated to mating success.In contrast, if females change their preferences (explanation2), males with traits beneficial to females independent of thecompetitive environment can maintain a high mating success underall levels of male–male competition. We tested these alternativesusing a small marine fish, the sand goby, Pomatoschistus minutus.The mating success of large males increased under conditionsallowing intense male competition, whereas females showed apreference for good nest building independent of the level ofcompetition. These findings suggest that females are in controlof their choice by altering their preference for male size inresponse to the intensity of male–male competition ratherthan female preference being overshadowed by male dominance.This plasticity of preferences implies that the strength ofsexual selection is not constant at the population level.     

Sexual size dimorphism,prey morphology and catch success in relation to flight mechanics in the peregrine falcon: a simulation study

In common with many other raptors, female peregrine falcons Falco peregrinus are about 50% heavier than males. Their sexual dimorphism is thought to allow breeding pairs to exploit a wider range of prey through a division of labor: the male being able to catch more maneuverable prey species; the female capable of carrying larger ones. Given the difficulty of assessing the catch success and load carrying capacity of both sexes of falcon in the field, we here adopt a novel approach to test the division‐of‐labor theory by using a detailed physics‐based flight simulator of birds. We study attacks by male and female peregrines on prey species ranging from small passerines to large ducks, testing how catch success relates to the flight performance of predator and prey. Males prove to be better than females at catching highly maneuverable prey in level flight, but the catch success of both sexes improves and becomes more similar when diving, because of the higher aerodynamic forces that are available to both sexes for maneuvering in high‐speed flight. The higher maximum roll acceleration of the male peregrine explains its edge over the female in catching maneuverable prey in level flight. Overall, catch success is more strongly influenced by the differences in maneuverability that exist between different species of prey than between the different sexes of falcon. On the other hand, the female can carry up to 50% greater loads than the male. More generally, our detailed simulation approach highlights the importance of several previously overlooked features of attack and escape. In particular, we find that it is not the prey's instantaneous maximum centripetal acceleration but the prey's ability to sustain a high centripetal acceleration for an extended period of time that is the primary driver of the variation in catch success across species.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

Time and energy constraints and the evolution of sexual size dimorphism — to eat or to mate?

Summary We present an empirical test of the Ghiselin—Reiss small-male hypothesis for the evolution of sexual size dimorphism (SSD). In mating systems dominated by scramble competition, where male reproductive success is a function of encounter rate with females, small males may be favoured when food is limiting because they require lower absolute amounts of food. Given a trade-off between time and energy devoted to foraging and to mate acquisition, small males should be able to devote more time to the latter. If at the same time larger females are favoured, this mechanism will contribute to the evolution of SSD and may be the major determinant of the female-biased SSDs that characterize most animal taxa. We tested this hypothesis using the water strider,Aquarius remigis (Heteroptera: Gerridae), a scramble competitor which mates many times over a prolonged mating season and which shows female-biased SSD. Laboratory experiments demonstrated that foraging success and giving up times (GUTs) are lower for males than for females during the reproductive season and that male water striders flexibly alter their time budgets under conditions of energy limitation. Controlled feeding experiments showed that male and female longevity, female fecundity and male mating success are positively related to food availability. As predicted, male body size is negatively correlated with several indices of male fitness (longevity, number of mating attempts and mating success), while female body size is positively correlated with longevity. These results are consistent with the hypothesis that scramble competition for mates favours small males in this species and provides empirical support for the Ghiselin—Reiss small-male hypothesis.     

Mating success in the spittlebug Cercopis sanguinolenta (Scopoli, 1763) (Homoptera, Cercopidae): the role of body size and mobility

In insects, a sexual size dimorphism commonly occurs, with larger females. However, as a deviation from this general rule, larger males are found in some species. In these species often sexual selection for large males has been presumed. The spittlebug Cercopis sanguinolenta exhibits a distinct sexual size dimorphism with larger males. Mating behaviour was studied in a field population in respect to mating success of males and females. The aim of this study was to examine the mechanisms that lead to the observed non-random mating pattern. The results showed a mating pattern without size-assortative mating. A correlation was found between mating success and body size in males. In females no such correlation was found. The mobility of males depends on their body size and mobility is high only when females are present. However, in an analysis of covariance it was found that male mating success is not correlated with mobility, when controlled for body size. The mating system of the spittlebug was classified as scramble competition polygyny. Electronic Publication     

Is size assortative mating in Paracalliope fluviatilis (Crustacea: Amphipoda) explained by male–male competition or female choice?

Field and laboratory studies were used to assess: (1) whether size assortative mating occurred in the New Zealand amphipod Paracalliope fluviatilis and (2) hypotheses developed to explain size assortative mating. We found that assortative mating occurred and that larger females carried more eggs, suggesting they may be more valuable as mates. Laboratory experiments were then used to determine whether: (1) male size influenced the size of the female selected (mechanical constraints hypothesis); (2) male size influenced pairing success in the presence of competition (intrasexual selection hypothesis); (3) take‐overs of females occurred and whether large males were more successful (intrasexual selection hypothesis); (4) guard duration varied relative to male and female size (guard duration hypothesis); and (5) females had control over pairing success and guard duration (intersexual selection hypothesis). Although there was evidence to suggest the existence of intrasexual competition for mates (i.e. both small and large males preferred large females), there was no evidence of overt competition (i.e. takeovers of paired females). There was also no difference with respect to how long small and large males guarded females, but large females were guarded longer by both male size classes. Females handicapped by having their mobility reduced were guarded for the same duration as control females but males were more likely to pair with handicapped females, suggesting that they were easier to amplex. Given the lack of evidence for direct male–male competition or female choice, we suggest that assortative mating may be the result of: (1) indirect competition (e.g. in situ large males may be better able to access and amplex the largest females) or (2) female resistance to small males in combination with higher costs that small males may incur in securing large females. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 173–181.     

Sexual Competition,Coercive Mating and Mate Assessment in the One‐Sided Livebearer,Jenynsia multidentata: Are They Predictive of Sexual Dimorphism?

We investigated the mechanisms of sexual selection operating on body size in the one‐sided livebearer (Jenynsia multidentata), a small fish characterized by male dwarfism. Mating in the one‐sided livebearer is coercive: males approach females from behind and try to thrust their copulatory organ at the female genital pore. Females counter males' mating attempts by either swimming away or attacking them. We tested the hypothesis that the components of sexual selection favouring small size in males (sexual coercion) were more effective than those favouring a large size (male competition and mate choice). When alone, small males had a significantly higher success in their mating attempts than large males. The proportion of successful attempts was also positively correlated with female size. When two males competed for the same female, the large male had a significant mating advantage over the small one. With a 1 : 1 sex ratio, the large‐male mating advantage vanished because each male tended to follow a different female. Large males, however, preferentially defended large females, thus compelling small males to engage with smaller, less fecund females. Males did not discriminate between gravid and non‐gravid females, but preferred mating with larger females. This preference disappeared when males were much smaller than the female, probably in relation to the risk for the male of being eaten or injured by the female. In a choice chamber, male‐deprived females that had their sperm storage depleted remained close to males and showed a preference for large individuals, a behaviour not observed in non‐deprived females. Nonetheless, when placed with males in the same aquarium, all females showed avoidance and aggression. Struggling may represent a way by which the female assesses the skill and endurance of males.     

A Life History Trade-Off Between Flight Ability and Reproductive Behavior in Male Field Crickets (<Emphasis Type="Italic">Gryllus texensis</Emphasis>)

Male field crickets (Gryllus texensis) that differ in flight ability incur a life history trade-off between flight ability and reproduction, where flight ability comes with a male fitness cost. In courtship trials, flight-capable males produced courtship song, a necessary signal for mating success, with a significantly lower probability than flight-incapable males. The trade-off was evident in young males, and a similar trend occurred in older males. Males that lost the ability to fly through histolysis of flight muscles produced courtship song with a similar probability as males incapable of flight for their entire lives. Time of day did not affect the expression of the trade-off. Neither male morph nor time of day influenced female mating behavior.