Wing pigmentation in territorial male damselflies, Calopteryx haemorrhoidalis: a possible relation to sexual selection

One striking characteristic in adult males of some odonate species is the presence of wing pigmentation. In Calopteryx species, males show a series of pre- and postcopulatory behavioural displays during which they face females while showing their pigmented wings. One hypothesis to explain the precopulatory flying displays and the associated wing pigmentation is that they may serve a sexual selection function. I investigated this in the territorial damselfly Calopteryx haemorrhoidalis. Males of this species defend aquatic substrates that females use for oviposition. Observational evidence indicated that males with a higher proportion of wing pigmentation were more likely to defend a territory, obtained more matings, had fewer gut parasites, survived in the study site and stayed in territories for longer. Experimental evidence suggested that the relationship mating success and wing pigmentation still held when controlling for the size of the substrate defended by territorial males. Similar to other studies in the Calopterygidae, these results suggest that wing pigmentation may be favoured by sexual selection. I discuss, however, whether an alternative function for male copulatory courtship displays and wing pigmentation, as sexual and/or species recognition, may also explain the evolution of these traits. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Size and mating success in males of the western harvester ant,Pogonomyrmex occidentalis (Hymenoptera: Formicidae)

Mating success in males of the lek mating ant species,Pogonomyrmex occidentalis, increases with increased body size. We estimated the magnitude of the selection coefficients on components of size by collecting males in copula and comparing their morphology to that of males that were collected at the lek but that were not mating. Four characters, body mass, head width, wing length, and leg length, were measured for a sample of 225 mating and 324 nonmating males and 225 females. Significant direct selection favors increased wing length and leg length. Multiple regression of transformed variables (principal components) indicated that the increased mating success of larger males is a function of all four characters. We found no evidence of positive assortative mating on the basis of any individual character or on the multivariate general size variable (the first principal component).     

Differences in size and energy content affect the territorial status and mating success of a neotropical dragonfly

In male odonates, both size and fat content are related to territory defence and mating success. Males that are larger and have higher energy reserves win relatively more disputes for territory and attract more females. Wing colour has also been regarded as a mechanism that influences agonistic behaviour between males, as wing pigmentation might be regarded as a sign of male quality. In this study, we analysed whether a set of male physical (body size and wing colour), physiological (body fat content) and behavioural (disputes between males) characteristics were involved in the territory defence and mating behaviour of the neotropical dragonfly Zenithoptera lanei Santos, 1941 (Anisoptera: Libellulidae). Males were characterised as territorial whenever they warded‐off other males and remained in the same place within the pond for two consecutive days. In general, these territorial males were larger and had more abdominal and thoracic fat, engaged in pursuits more frequently, spent more time on sexual behaviour and female guarding, and mated more in comparison to subordinate males. By evaluating whether the percentage of wing area covered by black ink influenced male behaviour, we found that territorial males tended to act aggressively towards other males whose wings were partially painted, and sexually towards females irrespective of wing area painted. In Z. lanei, both body size and fat content play a role in defining territoriality. By subduing competitors and dominating preferred locations within high‐quality sites, these males are likely to be visited by females and engage in mating.     

Interactions between a Candidate Gene for Migration (ADCYAP1), Morphology and Sex Predict Spring Arrival in Blackcap Populations

Avian research has begun to reveal associations between candidate genes and migratory behaviors of captive birds, yet few studies utilize genotypic, morphometric, and phenological data from wild individuals. Previous studies have identified an association between ADCYAP1 polymorphism and autumn migratory behavior (restlessness, or zugunruhe), but little is known about the relationship between ADCYAP1 and spring migratory behavior. The timing of spring migration and arrival to the breeding ground are phenological traits which could be particularly favorable for establishing territories and acquiring mates, thus important to fitness and reproductive success. Here, we investigated how individual genotypic ADCYAP1 variation and phenotypic variation (wing length and shape) of blackcaps (Sylvia atricapilla) affect spring arrival date across nine natural populations in Europe. We hypothesized that longer alleles should be associated with earlier spring arrival dates and expected the effect on arrival date to be stronger for males as they arrive earlier. However, we found that longer wings were associated with earlier spring arrival to the breeding grounds for females, but not for males. Another female-specific effect indicated an interaction between ADCYAP1 allele size and wing pointedness on the response of spring arrival: greater allele size had a positive effect on spring arrival date for females with rounder wings, while a negative effect was apparent for females with more pointed wings. Also, female heterozygotes with pointed wing tips arrived significantly earlier than both homozygotes with pointed wings and heterozygotes with round wings. Stable isotope ratios (δ ² H) of a subset of blackcaps captured in Freiburg in 2011 allowed us also to assign individuals to their main overwintering areas in northwest (NW) and southwest (SW) Europe. NW males arrived significantly earlier to the Freiburg breeding site than both SW males and females in 2011. NW females had more pointed wing tips compared to SW females, but no difference in ADCYAP1 allele size was found between the different migration routes.     

The Role of Wing Pigmentation, UV and Fluorescence as Signals in a Neotropical Damselfly

Pigmentation patterns, ultraviolet reflection and fluorescent emission are often involved in mate recognition and mate quality functions in many animal taxa. We investigated the role of wing ultra-violet reflection, fluorescence emission, and pigmentation on age and sexual signals in the damselfly Mnesarete pudica. In this species, wings are sexually dimorphic in colour and exhibit age dependency: males and females show a smoky black colouration when young, turning red in mature males while it turns brown in females. First, we investigated wing UV patterns through reflectance and emission spectra. Second, behavioural experiments were undertaken to show male and female responses to manipulated wing pigmentation and experimentally reduced UV (UV-). Reflectance spectra of the wings of juvenile and mature males and females were used to show the differences between controls and individuals with manipulated colouration used in the behavioural experiment. UV-reduced, females with wings painted red, and control males and females were tethered and presented to conspecific males and females, and their behavioral responses were recorded. The male red wing pigmentation and females with red wings elicited an aggressive response in territorial males and a sexual response in females. Both males and females showed neutral responses towards individuals with reduced UV. Wing signals of juvenile individuals also provoked neutral responses. These results suggest that UV, together with pigmentation, plays a role during mate recognition in males and females. Other than butterflies and spiders, it seems that fluorescence signals and UV reflectance can also be part of communication in odonates.     

Effect of Drosophila melanogaster Female Size on Male Mating Success

In this study, we examined the influence of female size on mating success in Drosophila melanogaster. The results that were obtained from experiments performed in mating chambers allowed us to confirm the results of previous studies, demonstrating higher mating success of larger D. melanogaster males, and to conclude that female size also affects mating success, either when considering a single male or two competing males. We observed that the advantage for larger males depends on their size relative to that of the female, demonstrating a previously unknown role for female size in mating behavior studies. This effect of female size on mating success depends on various factors: males take longer to initiate courtship toward larger females, large females receive more wing vibrations from males prior to mating, and large females tend to keep moving for longer periods during male courtship. The importance of this finding is discussed in the context of recent reports on sexual conflict in D. melanogaster, in which males were observed to depress fitness in females as a result of intercourse.     

Recapture of male and female dragonflies in relation to parasitism by mites, time of season, wing length and wing cell symmetry

For aquatic mites parasitic on dragonflies, completion of their life cycle depends on their being returned to appropriate water bodies by their hosts, after completion of engorgement. We examined whether differences among hosts in timing of emergence or phenotypic attributes might affect their probability of return to an emergence pond, and hence success of mites. Parasitized males and females of the dragonfly Sympetrum obtrusum (Hagen) did not differ in overall recapture rates. Females that had wing cell symmetry and emerged early were more likely to be recaptured than females that emerged later or had wing cell asymmetry, but there were no consistent relations between these variables and parasitism by mites. No such relations between wing cell asymmetry, emergence date, and recapture likelihood were found for males. Using randomization tests, we found that mean intensities of Arrenurus planus (Marshall) mites at host emergence were the same for recaptured females and females not recaptured; however, males that were recaptured had lower mean intensities of mites at emergence than males not recaptured. Further, mature females carried more mites than mature males, and the latter had fewer mites than newly emerged males not recaptured. Biases in detachment of engorging mites do not explain the differences in parasitism between mature males and females, nor the differences in mite numbers between mature males and newly emerged males that were not recaptured. Rather, heavily parasitized males appear to disperse or die and are not recaptured, which should have implications for dispersal of mites and fitness of male hosts.     

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.     

Energetic cost of sexual attractiveness: ultrasonic advertisement in wax moths

Pair formation in the lesser wax moth,Achroia grisella(Lepidoptera: Pyralidae), is initiated by male ultrasonic signals that attract receptive females. Individual males vary in attractiveness to females, and the most attractive males are distinguished by exaggeration of three signal characters: pulse rate, peak amplitude and asynchrony interval (temporal separation between pulses generated by movements of the left and right wings during a given wing upstroke or downstroke). Using flow-through respirometry, we measured the resting and signalling metabolic rates of males whose relative attractiveness was known. Acoustic recordings and metabolic measurements were made simultaneously, and we calculated net metabolic rates and factorial metabolic scopes as measures for the energetic cost of signalling. On average, attractive males had higher net metabolic rates and factorial metabolic scopes than unattractive ones, but many unattractive males also had high values. Thus, high expenditure of energy on signalling is necessary but not sufficient for attractiveness. This may result because only one of the three signal characters critical for female preference, pulse rate, is correlated with energy expenditure. Although the results are consistent with the good genes model of sexual selection, they do not conflict with other indirect or direct mechanisms of female choice.     

Sexual size dimorphism and assortative mating in Carolina Wrens

ABSTRACT.   Sexual size dimorphism (SSD) may be due to sexual and natural selection, but identifying specific mechanisms that generate such dimorphism in a species is difficult. I examined SSD in Carolina Wrens ( Thryothorus ludovicianus ) by examining (1) the degree of SSD in the population and between pairs using five morphometrics, (2) assortative mating patterns based on size and age, and (3) relationships between size and longevity. Analysis revealed that males were significantly larger than females in all body measurements. For example, mass, bill, and wing measurements yielded a canonical variable that permitted separation of the sexes and linear classification functions correctly determined the sex of 95% (238/250) of all wrens measured. No evidence was found to suggest that SSD was related to resource partitioning. However, assortative mating trends based on morphometrics (e.g., wing length), positive associations between longevity and morphometrics (e.g., wing length in females and body size in males), and intense male-male contests for territorial resources year-round provide evidence that sexual selection may contribute to SSD in Carolina Wrens.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

A small badge of longevity: opposing survival selection on the size of white and black wing markings

According to handicap principle, exaggerated ornamental traits are supposed to exert costs on their bearers. However, there is much less theoretical and practical consensus about whether and under which conditions ornament expression should positively correlate with survival. We measured age‐related variation and survival selection on the size of white wing patches and black wing tips in a long‐lived monogamous seabird, the common gull Larus canus. Males had larger white patches than females but patch size showed concave relationship with age irrespective of sex, suggesting that white patch size was prone to senescence in both sexes. Extent of wing tip abrasion correlated negatively with the size of white patch, suggesting, in agreement with the Zahavian handicap hypothesis that only individuals with largest ornaments are able of maintaining them and not paying cost of displaying them. Areas of white wing patches and black wing tips correlated negatively. Irrespective of sex, survival selection favored birds with larger white wing patches and smaller black wing tips, which suggests that white and black wing markings may have coevolved as reverse components of a single ornament. Altogether, our results provide an evidence for the case where survival selection on ornamental traits in females is not weaker than in males. Absence of sex differences with respect to most of observed patterns is consistent with a prediction that among monogamous long‐lived species with biparental care, mutual mate choice leads to evolution of elaborate ornamental traits in both sexes.     

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.     

Sex differences in age structure, growth rate and body size of common frogs Rana temporaria in the subarctic

The thermal environment and length of the activity season are important factors in shaping life-history trait variation in ectotherms. Many ectothermic vertebrates living at high latitudes or altitudes tend to be larger and older than their conspecifics living at lower latitudes or altitudes. However, detailed data on age, body size and growth variation—and how they may differ between males and females—are still scarce, especially from extreme high-latitude environments. We studied growth (body length increment), age and size structure of common frogs (Rana temporaria) in subarctic Finland (69°04′N) by applying skeletochronological methods to individually marked adults (n?=?169) captured and recaptured between 1999 and 2003. We found that breeding males were on average younger (mean?=?8.5?years) than females (11.9?years) and that males started reproducing earlier (≥3–4?years of age) than females (>4–5?years). The oldest encountered individual was an 18-year-old female, which to our knowledge is the oldest wild common frog ever reported. Females were on average larger (mean body length?=?76.6?mm) than males (70.7?mm), and this appeared to be mainly due to their older age as compared to males. While body length increased and growth rate decreased with age in both sexes, growth rate declined significantly faster with age in males than in females. The latter finding provides a proximate explanation for the observation that even after accounting for age differences among sexes (females?>?males), females were longer than males.     

Avoidance of maladaptive,precocious copulation in the cabbage white butterfly,Pieris rapae crucivora

In the cabbage white butterfly, Pieris rapae crucivora, copulation hinders normal expansion and hardening of the wings of newly emerged females. The resulting permanent wing deformation makes it impossible for females to fly and therefore, to find an oviposition site and nectar sources. An attempt was made to clarify whether the newly emerged female butterfly avoids copulation. Observation of wing expansion and hardening reveals that the wings are fully expanded and hard by 20–30 min after emergence. In the field, presentation of females with soft wings to males shows that males will attempt to copulate with these females. However, newly emerged females prevent successful completion of copulation by assuming the mate refusal posture, and thereby avoid a potentially maladaptive copulation. The discussion focusses on the question as to why females and not males avoid early copulation.     

Effects of predation by the copepod Mesocyclops ogunnus on the sex ratios of mosquito Aedes albopictus

Cyclopoid copepods are important predators in many aquatic ecosystems and have been used as biological agents in successful programs to control mosquito larvae. However, the impacts of this predation on adult mosquito populations are still poorly understood. The present study compared the sex ratios and body sizes (measured as wing length) of Aedes albopictus mosquitoes emerging from recipients containing the copepod predator Mesocyclops ogunnus with control situations without this predator. We found that copepod predation significantly biased mosquito sex ratios toward females, and that both the males and females emerging from copepod-containing recipients were significantly larger than control insects. The ecological and epidemiological consequences of the changes induced by copepod predation on mosquito populations are discussed.     

Male Age,Copulation Duration and Insemination Success in Mystacides azurea (Leptoceridae,Trichoptera)

Time in copula in the swarming caddis fly Mystacides azurea L. (Trichoptera: Leptoceridae) ranged from 0.33 to 44.5 min. It increased with male age (wing wear) and male dry weight, but was independent of male and female size (forewing length), female wing wear and number of eggs in the females. Older males failed to transfer sperm during copulation more frequently than did younger ones. It is suggested that females benefit by interrupting prolonged copulations if sperm is not transferred rapidly, since being in copula might increase the risk of predation. Alternatively, young and old males follow different mating tactics; old males have a lower chance to acquire new mates and they do better by monopolizing a female, once they get one, by prolonging the copulation.     

Love is in the air: olfaction-based mate-odour identification by jumping spiders from the genus Cyrba

Jumping spiders (Salticidae) are known for having good eyesight, but the extent to which they rely on olfaction is poorly understood. Here we demonstrate for the first time that olfactory pheromones are used by two species from the salticid genus Cyrba (C. algerina and C. ocellata). Using a Y-shape olfactometer, we investigated the ability of adult males and females of both species to discriminate between mate and non-mate odour. A hidden spider or a spider’s draglines (no spider present) were used as odour sources. There was no evident response by females of either Cyrba species to any tested odour. Males of both species chose odour from conspecific females, or their draglines, significantly more often than the no-odour control, but there was no evident response by males to any of the other odours (conspecific male and heterospecific female). Our findings demonstrate that C. algerina and C. ocellata males can make sex- and species-specific discriminations even when restricted to using olfaction alone. Also, by showing that draglines can be a source of olfactory pheromones, our findings illustrate the difficulty of ruling out olfaction when attempting to test for chemotactile cues.     

Sexual Dimorphism in Sturnira lilium (Chiroptera,Phyllostomidae): Can Pregnancy and Pup Carrying Be Responsible for Differences in Wing Shape?

Competition is one of the most cited mechanisms to explain secondary sexual dimorphism in animals. Nonetheless, it has been proposed that sexual dimorphism in bat wings is also a result of adaptive pressures to compensate additional weight caused by fetus or pup carrying during the reproductive period of females. The main objective of this study is to verify the existence of sexual dimorphism in Sturnira lilium wings. We employed geometric morphometrics techniques using anatomical landmarks superimposition to obtain size (Centroid Size) and shape variables of wings, which were reduced by Linear Discriminant Analysis (LDA). We also employed classical morphometrics using wing length measurements to compare efficiency between these two morphometric approaches and make comparisons using wing area measurements. LDA indicated significant differences between wing shapes of males and females, with 91% (stepwise classification) and 80% (leave-one-out cross validation) of correct classification. However, the size variable obtained did not contribute to such classifications. We have observed larger areas in female wings, but we found no differences in wing length measurements and no allometric effects in wing length, shape and area measurements. Interestingly, our study has provided evidences of morphological differences where classical morphometrics have failed. LDA and area measurements analyses revealed that females have a different area distribution in distinct portions of the wing, with wider dactylopatagia and plagiopatagia, and wingtips more triangular than males. No differences in body length or relative wing length were observed between the sexes, but pregnant females have more body weight than non-pregnant females and males. Our findings suggest that sexual dimorphism in the wing shape of S. lilium is probably related to the increase in flight efficiency of females during reproductive period. It decreases wing loading in specific portions of the wing and reduces energy cost to maintain a faster and maneuverable flight.