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.     

Sexual dimorphism inAnastrepha suspensa (Loew) and other tephritid fruit flies (Diptera: Tephritidae): Possible roles of developmental rate,fecundity, and dispersal

Larger male Caribbean fruit flies are more likely to be chosen as mates and defeat rivals in territorial contests. Yet males are smaller than females. Adaptive explanations for relatively small male size include (1) acceleration of male development to maximize female encounter rates, (2) selection for greater female size to increase fecundity, and (3) selection for body sizes most suitable for sexually dimorphic degrees of mobility, speed, and distance flight. None of these unambiguously accounts for the degree of sexual dimorphism. Male development is not accelerated relative to that of females. On average, males remain inside fruit longer than females and those males with extended development periods are smaller than more rapidly developing individuals. There is no evidence that female enlargement alone, presumably for greater fecundity, has generated the degree of dimorphism in the Caribbean fruit fly or other fruit flies. The relationship between dimorphism and mean female body size in 27 species of Tephritidae is the opposite of what would be predicted if differences in dimorphism were due to differences in unilateral female enlargement. Larger size in a species or in one sex of a species may be an adaptation for extensive flight. In general, among 32 species of fruit flies, as body size increases, wing shape becomes progressively more suited for distance flight. However, there are important exceptions to this correlation. Both sexual selection and nonadaptive allometries may contribute to the range of dimorphisms within the family.     

A comparative study on the evolution of reversed size dimorphism in monogamous waders

Sexual dimorphism in size is common in birds. Males are usually larger than females, although in some taxa reversed size dimorphism (RSD) predominates. Whilst direct dimorphism is attributed to sexual selection in males giving greater reproductive access to females, the evolutionary causes of RSD are still unclear. Four different hypotheses could explain the evolution of RSD in monogamous birds: (1) The ‘energy storing’ hypothesis suggests that larger females could accumulate more reserves at wintering or refuelling areas to enable an earlier start to egg laying. (2) According to the ‘incubation ability’ hypothesis, RSD has evolved because large females can incubate more efficiently than small ones. (3) The ‘parental role division’ hypothesis suggests that RSD in monogamous waders has evolved in species with parental role division and uniparental male care of the chicks. It is based on the assumption that small male size facilitates food acquisition in terrestrial habitats where chick rearing takes place and that larger females can accumulate more reserves for egg laying in coastal sites. (3) The ‘display agility’ hypothesis suggests that small males perform better in acrobatic displays presumably involved in mate choice and so RSD may have evolved due to female preference for agile males. I tested these hypotheses in monogamous waders using several comparative methods. Given the current knowledge of the phylogeny of this group, the evolutionary history of waders seems only compatible with the hypothesis that RSD has evolved as an adaptation for increasing display performance in males. In addition, the analysis of wing shape showed that males of species with acrobatic flight displays had wings with higher aspect ratio (wing span/²wing area) than non-acrobatic species, which probably increases flight manoeuvrability during acrobatic displays. In species with acrobatic displays males also had a higher aspect ratio than females although no sexual difference was found in non-acrobatic species. These results suggest that acrobatic flight displays could have produced changes in the morphology of some species and suggest the existence of selection favouring higher manoeuvrability in species with acrobatic flight displays. This supports the validity of the mechanisms proposed by the ‘display agility’ hypothesis to explain the evolution of RSD in waders.     

Butterfly contests and flight physiology: why do older males fight harder?

The males of many butterfly species compete for territoriesvia conspicuous aerial wars of attrition, in which the determinantsof persistence ability are largely unclear. Flight performancefeatures, such as stamina, acceleration, and maneuverability,are often assumed to be important in this context, yet thereis no direct evidence by which to evaluate these possibilities.Recent research has indicated that competitive ability increaseswith age in a notably territorial species, Hypolimnas bolina,which could arise from lifetime morphological or physiologicalchanges that directly affect flight performance. I evaluatedthis hypothesis by investigating how size-independent variancein body composition, energy stores, flight muscle ratio (FMR),and wing condition change with age in this species. Males infive age categories (spanning the functional life span of territorialindividuals) were sampled from encounter sites in tropicalAustralia. Analysis of body composition with respect to anestimate of eclosion mass (forewing length) indicated thattotal body mass, abdomen mass, and wing area decrease throughoutan individual's lifetime, but thorax mass remains unchanged.Wing loading (the ratio of wing area to body mass) is lowestin intermediately aged individuals, but FMR and energetic statusremain largely similar regardless of age. On average, therefore,the energetic cost of sustained flight should first decrease,then increase, with age in a male H. bolina (of standardizedbody size), while available energy reserves decline slightly. Acceleration and maneuverability should remain relatively constant.These results, coupled with the fact that body size is unrelatedto contest success in this territorial butterfly, fail to supportthe idea that age-related competitive ability is mediated simplyby energetics or flight performance.     

The Role of Wing Coloration in Sex Recognition and Competitor Recognition in Rubyspot Damselflies (Hetaerina spp.)

The decision rules that animals use for distinguishing between conspecifics of different age and sex classes are relevant for understanding how closely related species interact in sympatry. In rubyspot damselflies (Hetaerina spp.), the red wing coloration of mature males is hypothesized to be a key trait for sex recognition and competitor recognition within species and the proximate trigger for interspecific male–male aggression. We tested this hypothesis by manipulating the wing coloration of tethered conspecific intruders and measuring the responses of territory holders of three species in the field. As predicted, covering the red spots of mature males with black ink nearly eliminated territorial responses, and in some cases, territorial holders clasped the blackened males as if they were females. Adding red spots to female wings triggered territorial responses and nearly eliminated sexual responses. Immature males with artificial red spots were attacked at the same rate as mature male intruders, and much more frequently than were immature male controls. The results varied somewhat by species. In H. titia, the only species of Hetaerina with substantial black wing pigmentation, the effects of blackening the red spots of intruders varied both geographically and seasonally. But even when blackening the red spots of male intruders did not reduce the aggressive response of H. titia territory holders, adding artificial red spots to female wings elicited aggressive responses and nearly eliminated sexual responses. The results of this study further strengthen the evidence that interspecific aggression in Hetaerina results from overlap in territorial signals and that the derived black wing pigmentation of H. titia reduces interspecific aggression.     

Is natural selection promoting sexual dimorphism in the Green-backed Firecrown Hummingbird ( Sephanoides Sephaniodes )?

In many hummingbird species there is an opposite pattern of sexual dimorphism in bill length and other morphometric measures of body size. These differences seem to be closely related with differences in foraging ecology directly associated with a different resource exploitation strategy. The aim of this study was to assess if natural selection is acting on wing length and bill size in hummingbird males and females with different resource exploitation strategies (i.e., territorial males and non-territorial females). If competition for resources promotes sexual dimorphism as a selective pressure, males should be subjected to negative directional selection pressure for wing length and no selection pressure over bill size, while females should undergo positive directional selection pressure for both bill size and wing length. The morphometric data we collected suggests that there is no selection for wing length and bill size in male hummingbirds. In contrast, our females exhibited positive directional selection for both wing length and bill size. Although we cannot reject sexual selection acting on sexually dimorphic traits, this study suggests that natural selection may promote sexual dimorphism in traits that are closely related with hummingbird foraging ecology and resource exploitation strategies.     

Success in territorial defence by male tarantula hawk wasps Hemipepsis ustulata: the role of residency

1. Some male tarantula hawk wasps Hemipepsis ustulata defend landmark shrubs growing on mountain tops, while others only visit a succession of territory sites, generally fleeing from territorial males. To determine whether residency asymmetries contribute to the short duration of most interactions between territorial males and visiting intruders, established residents were held captive for varying periods while other males replaced them at their territories.
2. Replacement males immediately assumed resident status and began to defend the shrubs and trees with success against non-resident intruders, many of which were larger. Interactions between replacements and non-resident intruders rarely escalated into pursuits leading to ascending flights.
3. In contrast, when the original resident was released, its subsequent encounter with the replacement male almost always resulted in one or more ascending flights, even when the replacement had been on territory for less than 10 min. The longer the period of territory occupation by replacements, the longer on average the interaction and the more ascending flights, showing that the duration of time on a territory affects male motivation to defend the site. Despite strong resistance from long-term replacements, however, returning residents almost always regained their territories.
4. The strong residency effect in H. ustulata may be an adaptive proximate control mechanism for male behaviour that arises because established residents typically have high resource holding power, which makes them difficult to displace.     

Take-off flight performance in the butterfly Pararge aegeria relative to sex and morphology: a quantitative genetic assessment

Adult fitness components may strongly depend on variation in locomotory performance such as flight; this variation can be sex specific. Fast take-off to intercept females and competing males is an essential behavioral component of the territorial perching behavior in male speckled wood butterflies (Pararge aegeria L.). Females on the other hand avoid frequent take-offs particularly under suboptimal temperatures, typically showing fewer but longer flights than males. We estimated the heritability of take-off acceleration performance under suboptimal body temperatures by a restricted maximum-likelihood model. We calculated genetic correlations between this performance and a selection of morphological traits: size (body mass), flight muscle investment (relative thorax mass), and wing shape (forewing aspect ratio). Our results show significant additive genetic variation for mean acceleration performance and a similar but nonsignificant trend (P= 0.08) for maximal acceleration performance during take-off in males (h(2)= 0.15). In females, heritability was not significantly different from zero for either of the acceleration performance measures. Morphological traits and take-off performance were genetically linked in a sex-specific way. In males, relative thorax mass and forewing aspect ratio were positively genetically correlated with acceleration performance. In females, there was a negative genetic correlation between acceleration performance and abdomen mass, but not with residual abdomen mass (i.e., regressed on total body mass). To fully understand the evolution of sexual differences in flight performances and morphology, several other flight performances will have to be included. This multifunctional nature of flight and its consequences for the evolutionary study of flight has not yet been fully appreciated in the literature.     

Temperature shapes the costs,benefits and geographic diversification of sexual coloration in a dragonfly

The environment shapes the evolution of secondary sexual traits by determining how their costs and benefits vary across the landscape. Given the thermal properties of dark coloration generally, temperature should crucially influence the costs, benefits and geographic diversification of many secondary sexual colour patterns. We tested this hypothesis using sexually selected wing coloration in a dragonfly. We find that greater wing coloration heats males – the magnitude of which improves flight performance under cool conditions but dramatically reduces it under warm conditions. In a colder region of the species’ range, behavioural observations of a wild population show that these thermal effects translate into greater territorial acquisition on thermally variable days. Finally, geo‐referenced photographs taken by citizen scientists reveal that this sexually selected wing coloration is dramatically reduced in the hottest portions of the species’ range. Collectively, our results underscore temperature's capacity to promote and constrain the evolution of sexual coloration.     

Flight behaviour of seabirds in relation to wind direction and wing morphology

We studied flight direction relative to wind direction (hereafter referred to as "flight direction"), the relation between wing morphology and flight behaviour and interspecies relationships in flight behaviour among all major seabird taxa. We calculated wing loading and aspect ratios for 98 species from 1029 specimens. Species were sorted into 13 groups on the basis of similarity in patterns of flight direction. The primary flight direction of Pelecaniformes and Charadriiformes was into and across headwinds. The most common flight direction of Procellariiformes was across wind. Seabirds avoided flying with tailwinds. Wing loading and aspect ratios were positively correlated in Procellariiformes, Pelecaniformes and alcids but negatively correlated in larids. In Procellariiformes, incidence of headwind flight and that of tailwind flight were significantly correlated with wing loading and aspect ratio; species with higher wing loading and aspect ratios flew more often into headwinds and less often with tailwinds. In contrast, the proportion of Pelecaniformes and Charadriiformes flying with tailwinds increased significantly with increased wing loading. Our results demonstrate a close link in seabirds between flight behaviour, wing morphology and natural history patterns in terms of distribution, colony location, dispersal and foraging behaviour.     

Ontogeny of sexual dimorphism in the Long-tailed Manakin Chiroxiphia linearis: long maturation of display trait morphology

Males of dimorphic species often show ornaments that are thought to have evolved through female choice or/and male–male competition. The sexual differentiation of similar morphologies occurs during ontogeny, resulting in differential sex and age-specific selection. The Long-tailed Manakin is a dimorphic species with a highly skewed mating system, the males of which delay plumage maturation over 3 to 4 years. We describe ontogenetic changes in feather morphology in this species through sexual maturity. Males showed a significant increase in length of the central rectrices with age, hence their degree of sexual dimorphism increased from zero in 1-year-old males to 189.5% in adults. In contrast, male tail length decreased with age. Wing length did not vary significantly with age, but females had relatively longer wings than males. Wing loading was greater in females and decreased with age in males. In adults, rectrix length was positively correlated with testis volume, supporting the hypothesis that secondary sexual characters can signal the condition of primary sexual characters. Rectrix length showed positive allometry with body size in males less than 4 years old, whereas older males showed negative allometry and females showed isometry. Wing area and wing loading shifted from negative to positive allometry in males of 2 to 3 years of age. Changes in male morphology during ontogeny in the Long-tailed Manakin appeared to be associated with their specific display behaviours. Age-related changes in allometric growth of rectrices in the Long-tailed Manakin suggested that young males invest disproportionately more in the length of this trait relative to their body size. This investment could act as a signal of competitive ability to move status position in their orderly queue.     

On the sex-specific mechanisms of butterfly flight: flight performance relative to flight morphology, wing kinematics, and sex in Pararge aegeria

Many evolutionary ecological studies have documented sexual dimorphism in morphology or behaviour. However, to what extent a sex-specific morphology is used differently to realize a certain level of behavioural performance is only rarely tested. We experimentally quantified flight performance and wing kinematics (wing beat frequency and wing stroke amplitude) and flight morphology (thorax mass, body mass, forewing aspect ratio, and distance to centre of forewing area) in the butterfly Pararge aegeria (L.) using a tethered tarsal reflex induced flight set-up under laboratory conditions. On average, females showed higher flight performance than males, but frequency and amplitude did not differ. In both sexes, higher flight performance was partly determined by wing beat frequency but not by wing stroke amplitude. Dry body mass, thorax mass, and distance to centre of forewing area were negatively related to wing beat frequency. The relationship between aspect ratio and wing stroke amplitude was sex-specific: females with narrower wings produced higher amplitude whereas males show the opposite pattern. The results are discussed in relation to sexual differences in flight behaviour.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 675–687.     

Positive Relationship between Signalling Time and Flight Capability in the Texas Field Cricket, Gryllus texensis

A trade‐off between dispersal ability and reproduction is generally thought to explain the persistence of wing dimorphism in insects, although this trade‐off has received minimal attention in male insects. Research on male sand cricket, Gryllus firmus, supports the trade‐off hypothesis insofar as flight capable cricket’s spend significantly less time signalling for potential mates than their flightless counterparts. By contrast, here I show that this expected trade‐off between signalling time and wing dimorphism does not exist in a male congener, the Texas field cricket (Gryllus texensis). In G. texensis, flight capable males signal twice as often as flightless males. Thus, unless male G. texensis express trade‐offs between dispersal ability and other, presently unmeasured components of reproduction, the trade‐off hypothesis may not explain the persistence of wing dimorphism in all male insects.     

Territoriality and male-biased sexual size dimorphism in Argia reclusa (Odonata: Zygoptera)

In Odonata, many species present sexual size dimorphism (SSD), which can be associated with male territoriality in Zygoptera. We hypothesized that in the territorial damselfly Argia reclusa, male–male competition can favor large males, and consequently, drive selection pressures to generate male-biased SSD. The study was performed at a small stream in southeastern Brazil. Males were marked, and we measured body size and assessed the quality of territories. We tested if larger territorial males (a) defended the best territories (those with more male intrusions and visiting females), (b) won more fights, and (c) mated more. Couples were collected and measured to show the occurrence of sexual size dimorphism. Results indicated that males are larger than females, and that territorial males were larger than non-territorial males. Larger territorial males won more fights and defended the best territories. There was no difference between the mating success of large territorial and small non-territorial males. Although our findings suggest that male territoriality may play a significant role on the evolution of sexual size dimorphism in A. reclusa, we suggest that other factors should also be considered to explain the evolution of SSD in damselflies, since non-territorial males are also capable of acquiring mates.     

Sexual size dimorphism in shorebirds, gulls, and alcids: the influence of sexual and natural selection

Abstract. Charadrii (shorebirds, gulls, and alcids) have an unusual diversity in their sexual size dimorphism, ranging from monomorphism to either male-biased or female-biased dimorphism. We use comparative analyses to investigate whether this variation relates to sexual selection through competition for mates or natural selection through different use of resources by males and females. As predicted by sexual selection theory, we found that in taxa with socially polygynous mating systems, males were relatively larger than females compared with less polygynous species. Furthermore, evolution toward socially polyandrous mating systems was correlated with decreases in relative male size. These patterns depend on the kinds of courtship displays performed by males. In taxa with acrobatic flight displays, males are relatively smaller than in taxa in which courtship involves simple flights or displays from the ground. This result remains significant when the relationship with mating system is controlled statistically, thereby explaining the enigma of why males are often smaller than females in socially monogamous species. We did not find evidence that evolutionary changes in sexual dimorphism relate to niche division on the breeding grounds. In particular, biparental species did not have greater dimorphism in bill lengths than uniparental species, contrary to the hypothesis that selection for ecological divergence on the breeding grounds has been important as a general explanation for patterns of bill dimorphism. Taken together, these results strongly suggest that sexual selection has had a major influence on sexual size dimorphism in Charadrii, whereas divergence in the use of feeding resources while breeding was not supported by our analyses.     

Coevolution between flight morphology,vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight‐related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit‐feeding butterflies sampled in seven sites across 3341 ha. We used wing–thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing–thorax ratio in both sexes. Stouter fast‐flying species preferred the canopy, whereas more slender slow‐flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait–habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms.     

Sexual selection for male mobility in a giant insect with female-biased size dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.     

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.     

Seasonal sexual segregation in two Thalassarche albatross species: competitive exclusion,reproductive role specialization or foraging niche divergence?

Sexual segregation by micro- or macrohabitat is common in birds, and usually attributed to size-mediated dominance and exclusion of females by larger males, trophic niche divergence or reproductive role specialization. Our study of black-browed albatrosses, Thalassarche melanophrys, and grey-headed albatrosses, T. chrysostoma, revealed an exceptional degree of sexual segregation during incubation, with largely mutually exclusive core foraging ranges for each sex in both species. Spatial segregation was not apparent during brood-guard or post-guard chick rearing, when adults are constrained to feed close to colonies, providing no evidence for dominance-related competitive exclusion at the macrohabitat level. A comprehensive morphometric comparison indicated considerable species and sexual dimorphism in wing area and wing loading that corresponded, both within and between species, to broad-scale habitat preferences relating to wind strength. We suggest that seasonal sexual segregation in these two species is attributable to niche divergence mediated by differences in flight performance. Such sexual segregation may also have implications for conservation in relation to sex-specific overlap with commercial fisheries.     

Causes of sex‐biased adult survival in ungulates: sexual size dimorphism,mating tactic or environment harshness?

Using both a conventional and a phylogenetic approach, we tested whether sexual size dimorphism, mating tactic and environmental conditions influenced the between-sex differences in adult survival among 26 populations of polygynous ungulates. As a general rule, male survival was both lower and more variable among species than female survival. Whatever the method we used, sexual size dimorphism had no direct influence on male-biased mortality. In food-limited environments, the survival of males relative to that of females was lower than in good environments, suggesting a cost of large size for males facing harsh conditions. On the other hand, the survival of males relative to that of females tended to increase with sexual size dimorphism in good environments, indicating that large size may be profitable for males facing favourable conditions. Lastly, we found that the between-sex differences in adult survival did not vary with sexual size dimorphism in harem-holding or tending species, but tended to increase with sexual size dimorphism in territorial species. Our analyses indicate that sexual size dimorphism does not lead directly to a decrease in male survival compared to that of females. Thus, environmental conditions rather than the species considered could shape between-sex differences in adult survival observed in ungulate populations.