Size-related sexual selection and yeast diet inDrosophila buzzatii (Diptera: Drosophilidae)

Size-related sexual selection (SRSS) was examined on four traits (thorax and wing length and head and face width) inDrosophila buzzatii, by scoring male copulatory status in two mass-mating experiments. Using axenic females, experiment 1 was carried out with axenic males, and experiment 2 with yeast-supplemented males. While there was no indication of SRSS in experiment 1, such selection was substantial in yeast-supplemented males, which transmitted yeasts to mating females. Multivariate analyses of selection indicated that face width is the measured trait on which directional SRSS essentially acted in yeast-supplemented males, resulting in indirect selection on body size. Because this selection was affected by yeast diet in males, its possible interaction with the yeast transmission from males to females during the courtship is discussed.     

Courtship success and multivariate analysis of sexual selection on morphometric traits inDrosophila buzzatii (Diptera: Drosophilidae)

Using wild-reared flies, we examined sexual selection on five phenotypic traits (thorax length, wing length, wing width, head width, and face width) inDrosophila buzzatii, by scoring copulatory status in nine mass mating cages. Only male face width was identified as a direct target of sexual selection in an analysis of selection gradient, while indirect selection was present on all other studied traits, as expected from their correlations with face width. In contrast to males, there was no indication of selection in females. Nor was there evidence of assortative mating. The suggested direct selection on face width seems to take place during licking behavior of the courtship and might be related to courtship feeding. This study suggests that courtship success gives rise to indirect selection on body size.     

Sexual dimorphism of guava weevil Conotrachelus psidii Marshall (Coleoptera: Curculionidae)

We investigate structural and behavioral differences between male and female adults of guava weevil, Conotrachelus psidii Marshall, 1922 to help in sex determination. Weevils collected from a commercial guava orchard were individually caged and maintained under laboratory conditions. Every day ten individuals were grouped and their behavior was observed. During mating the males and females were identified. Fifty weevils of each sex were killed and they were observed with stereoscopic microscope. The antenna insertion on the rostrum and hair on the thorax and abdomen surfaces were examined. Length and width of body, pronotum, last abdominal sternite and rostrum were also taken. Behaviorally, the male weevils produce audible sound by stridulation of abdomen while females do not. In the females, the body and pronotum width and body a rostrum length were higher than in the males, but the last abdominal sternite was smaller. Three other structural differences were visible with the naked eye: 1) the anterior region of pronotum have scarce hair or have not in the females and hair is dense in the males; 2) the last tergite of females is hidden by the penultimate tergite whereas in the males is visible; 3) the first abdominal sternite of females is more prominent, very convex and without or with little bristle. In the males it is slightly concave and with abundant hair. The male-female differences found in this work allow, without hurt or change insect behavior, an efficient sex determination of guava weevil.     

Sex‐specific selection and sexual size dimorphism in the waterstrider Aquarius remigis

We estimated selection on adult body size for two generations in two populations of Aquarius remigis, as part of a long‐term study of the adaptive significance of sexual size dimorphism (SSD). Net adult fitness was estimated from the following components: prereproductive survival, daily reproductive success (mating frequency or fecundity), and reproductive lifespan. Standardized selection gradients were estimated for total length and for thorax, abdomen, genital and mesofemur lengths. Although selection was generally weak and showed significant temporal and spatial heterogeneity, patterns were consistent with SSD. Prereproductive survival was strongly influenced by date of eclosion, but size (thorax and genital lengths in females; total and abdomen lengths in males) played a significant secondary role. Sexual selection favoured smaller males with longer external genitalia in one population. Net adult fitness was not significantly related to body size in females, but was negatively related to size (thorax and total length) in males.     

Female Mate Choice, Calling Song and Genetic Variance in the Cricket, Gryllodes sigillatus

Female preferences for song patterns of males of Gryllodes sigillatus and genetic variance of morphological traits correlated with them were analyzed. Females preferred short pulses associated with large males. The males’ thorax width, wing length and femur III length showed stronger relationship with the song pulse duration, whereas the relationship between pulse duration and wing width was not significant. Interestingly, this last trait was the only one that showed significant levels of genetic variance. Perhaps these results could be explained by the evolutionary response to sexual selection. Sexual selection could deplete the genetic variance in the male traits related to male‐mating success.     

Evolutionary genetics of Drosophila ananassae: evidence for trade-offs among several fitness traits

Correlated responses to bi‐directional selection on thorax length, examined on several life‐history traits and chromosome inversion polymorphisms, have revealed apparent novel trade‐offs in Drosophila ananassae. We provide evidence of trade‐offs between hatching time and pupal period, pupal period and egg‐pupa development time, and pupal period and larval development time (LDT). Body size shows positive correlations with ovariole number, LDT and DT (egg–fly). We provide evidence of sexual dimorphism for trade‐offs between longevity and body size and starvation and longevity in females only. Trade‐offs between wing/thorax (W/T) ratio and longevity, W/T ratio and starvation, and DT (egg‐ fly) and longevity are evident in males only. Sexual dimorphism is also evident for inversion polymorphism with body size and longevity. A longevity assay suggests that low line females outlived high line females whereas high line males outlived low line males. The mean longevity in males is negatively correlated with the 2L‐ST and 3R‐ST arrangement frequencies whereas the 3L‐ST arrangement frequency is positively correlated with the mean longevity in males but opposite arrangements are found in females. Absolute starvation resistance is negatively correlated with 2L‐ST and 3R‐ST chromosome arrangements and results in a trade‐off between longevity and absolute starvation resistance in females. Analyses of fecundity, hatchability, and viabilities based on age intervals in both G₁₀ and G₁₃ suggest that the early reproduction is favoured in D. ananassae. The productivity percentage is highest in the high line and there is no effect of late reproduction on it. Overall, we provide some unravelled trade‐offs and striking sex differences, which may help in understanding the life‐history evolution of the species. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 669–685.     

Morphological and histological examination of short‐wing formation in the winter moth Protalcis concinnata (Insecta: Lepidoptera,Geometridae)

Winter geometrid moths exhibit sexual dimorphism in wing length and female‐specific flightlessness. Female‐specific flightlessness in insects is an interesting phenomenon in terms of sexual dimorphism and reproductive biology. In the winter geometrid moth, Protalcis concinnata (Wileman), adult females have short wings and adult males have fully developed wings. Although the developmental process for wing reduction in Lepidoptera is well studied, little is known about the morphology and the developmental pattern of short‐winged flightless morphs in Lepidoptera. To clarify the precise mechanisms and developmental processes that produce short‐winged morphs, we performed morphological and histological investigations of adult and pupal wing development in the winter geometrid moth P. concinnata. Our findings showed that (a) wing development in both sexes is similar until larval‐pupal metamorphosis, (b) the shape of the sexually dimorphic wings is determined by the position of the bordering lacuna (BL), (c) the BL is positioned farther inward in females than in males, and (d) after the short pupal diapause period, the female pupal wing epithelium degenerates to approximately two‐thirds its original size due to cell death. We propose that this developmental pattern is a previously unrecognized process among flightless Lepidoptera.     

Reproductive behavior of the dragonfly,Orthetrum japonicum (Odonata: Libellulidae)

The reproductive behavior of the dragonfly,Orthetrum japonicum, is described. Behavioral processes of turnover of territorial males, simultaneous guarding of 2 females by a male, and copulation by non-territorial males are described. The males with longer hind wings won the territorial conflicts more frequently. The total duration of territorial residence of a given male was correlated with the number of his matings, but not correlated with the length of his abdomen or hind wings. The territorial site with the lower degree of vegetation cover was occupied by males more consistently. Males in more consistently occupied territorial sites did not have longer abdomen and hind wings than males in less consistently occupied sites. The territorial site where the larger number of copulations was observed was not occupied more consistently. Selection episode analysis using the method of Arnold & Wade (1984a, b) showed that direct selection on the hind wing length favored the short wing and that direct selection on the abdomen length favored the long abdomen during mating.     

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).     

Behavioural evidence for a female sex pheromone in Cotesia flavipes (Hymenoptera: Braconidae)

Abstract.  The mating behaviour of Cotesia flavipes Cameron, 1891 (Hymenoptera: Braconidae) was observed under laboratory conditions. The results demonstrate that chemical stimuli play an important role in bringing males and females together for mating. Females from this species release chemical compounds to attract males, which exhibit a distinct courtship behaviour divided into three main steps: (i) antennation, (ii) lateral bouncing and (iii) wing fluttering. Ethological tests using hexane extracts from two different parts of the female's body (head plus thorax and abdomen) demonstrate that males are more attracted and also display a courtship behaviour when a filter paper containing hexane extracts from the abdomen is presented to them, indicating that the abdomen is the source of the sex pheromone of this species. Naive males of this species respond to hexane extracts of this gland by displaying courtship behaviour. Furthermore, additional tests show that C. flavipes females mate only once, whereas virgin males do not show any preference for virgin or mated females, suggesting that mated females of this species continue to produce sex attractants after mating.     

Extra DNA replication and cell proliferation in wing imaginal discs of aDrosophila species hybrid

Summary Species hybrids obtained from corsses betweenD. azteca andD. athabasca display characteristic growth anomalies. These are restricted to hybrid males while hybrid females resemble the pure species. The ratio of the wing length to body length is 1 in pure species, 1.3 in hybrid males from the crossD. azteca x D. athabasca and 0.65 in hybrid males from the reciprocal cross. This indicates a disproportionate enlargement and diminuation, respectively, restricted to some tissues. Whereas the number of ommatidia in the compound eye is normal, the number of wing cells is approximately twice as high in hybrid giant males as in hybrid females and pure species. Microdeterminations of the DNA content of wing discs show that an extra DNA replication step occurs during the early pupal stage of hybrid giant males; the pupal stage of these males lasts about 58 h longer than that of the pure species. In eye-antenna discs no extra DNA synthesis has been found. Incorporation experiments reveal a delayed uptake of radioactively labeled thymidine into the DNA of wing discs in late third instar larvae and early pupae of hybrid giant males. The results of this study are interpreted to indicate that a disturbed balance between sex-linked and autosomal genes in hybrid males is responsible for the modified rhythm of DNA synthesis and of cell proliferation in certain cell types.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

Relative growth and morphological sexual maturity of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) (Decapoda,Brachyura, Sesarmidae) on the southern coast of the state of São Paulo,Brazil

Growth in crustaceans is characterized by ontogenetic differentiation during the development of their body structures, so studies on relative growth are widely applied in this group. In this study, the growth pattern of the body structures was verified through the analysis of relative growth, and then, morphological sexual maturity of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) was estimated. The carapace width (CW), cheliped propodus length (PL), cheliped propodus height (PH), propodus width (PW), abdomen width (AW), and first pleopod length (FPL) of the crabs were measured. The relationships that best showed changes in the allometric coefficient among demographic categories were FPL vs. CW for males and AW vs. CW for females. This study verified the increased size of the cheliped in terms of length, width, and height, which occurred mainly in adult males. This increase reflects the importance of this structure in the reproductive processes of A. pisonii. For females, the increase in abdomen growth reflects their reproductive potential, since it is a structure that provides protection for eggs.     

Quantitative morphometrical analysis of a North African population of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: sexual dimorphism,and comparison with European populations

Genetic variability of quantitative traits was investigated in aMoroccan population of Drosophila melanogaster, with an isofemale line design. Results were compared with data previously obtained from French populations. Although the environmental and thermal conditions are very different in France and Morocco, only two significant differences were observed: a shorter wing and a lighter abdomen pigmentation in Morocco. It is, therefore, concluded that Moroccan D. melanogaster are quite typical temperate flies, belonging to the Palaearctic region, and very different from the ancestral Afrotropical populations. Almost all traits were genetically variable, as shown by significant intraclass correlations among lines. Genetic correlations were highly significant among three size-related traits, while much lower between size and bristle numbers. Fluctuating asymmetry was greater for abdominal bristles than for sternopleural bristles. Sex dimorphism, analysed as a female/male ratio, was identical in French and Moroccan populations. Examination of the thorax length/thorax width ratio showed that the thorax is more elongated in females. Sexual dimorphism of wing length was significantly more correlated to thorax width than to thorax length. The results illustrate the value of measuring numerous quantitative traits on the same flies for characterizing the genetic architecture of a natural population. In several cases, and especially for genetic correlations, some interesting suggestions could be made, which should be confirmed, or invalidated, by more extensive investigations.     

Extreme Eye Projection in the Male Weevil <Emphasis Type="Italic">Exechesops leucopis</Emphasis> (Coleoptera: Anthribidae): Its Effect on Intrasexual Behavioral Interferences

Fungus weevils, Exechesops leucopis (Anthribidae), are sexually dimorphic in the degree of eye protrusion and antenna elongation. I examined the allometric relationships of eye span, eyestalk length, antenna length, elytra width, and wing length against body size (pronotum width), and their effects on the outcome of male-male combat in the laboratory. Male eye span, eyestalk length, and antenna length indicated positive allometry, while elytra width showed isometry, and wing length showed negative allometry. In male-male combat, males with a larger eye span, antenna length, and body size defeated those with smaller attributes. However, when males fought experimentally males of similar body size, only eye span affected the outcome of combat, independent of body size and antenna length. In the female-female contests, the prior residency was an important determinant of victory the other than any morphological traits.     

Phenotypic variation in adult morphology and pupal colour within and among families of the speckled wood butterfly Pararge aegeria

Abstract .1. Larvae from eggs of fifteen wild-caught speckled wood females were reared individually under common conditions (LD 18:6 h and 17 °C) in the laboratory.
2. Pupal colour (green or brown) and the following adult morphological variables that are known or assumed to be related to behavioural variation (thermoregulation, flight, crypsis) were studied: size, relative thorax mass, area of yellow patches on the dorsal forewing, size of submarginal eyespots on the dorsal hindwing, colour of the dorsal thorax, dorsal basal wing, and dorsal distal wing.
3. The results of the breeding experiment indicate significant differences in adult morphological traits among families, sexes, and pupal colour types.
4. All adult morphological variables (except spot size and thorax colouration) differed significantly among families, suggesting genetic variation underlying the phenotypic variation. Heritabilities for these features were intermediate (0.38) to high (> 1).
5. Apart from known aspects (e.g. size), novel aspects of the sexual dimorphism were found: females had paler thoraxes than males, which relates to higher abundance of fur on the thorax and hence to thermoregulatory differences.
6. Green pupae produced larger individuals with a larger relative thorax mass than brown pupae. Green pupae produced adults with a paler basal wing colour in females, but not in males. These relationships are novel and suggest a trade-off between juvenile and adult investment.     

Causes of variation in wing loading among Drosophila species

We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.     

Population genetics of Drosophila ananassae: inversion polymorphism and body size in Indian geographical populations

Twelve Indian natural populations of Drosophila ananassae, a cosmopolitan and domestic species, were sampled and laboratory populations (mass cultures) were established from naturally impregnated females. These populations were maintained in the laboratory for some generations and were analysed chromosomally to know the frequency of different inversions. The chromosomal analysis revealed the presence of three cosmopolitan inversions. The data on the whole show that there are significant differences in the frequencies of different chromosome arrangements in these populations. Body size (wing length and thorax length) was measured in both sexes (50 females and 50 males), in all the 12 geographical populations of D. ananassae. There are statistically significant differences in wing length as well as in thorax length of both sexes among different geographical populations. Five geographical strains were crossed reciprocally and body size (wing length and thorax length) was measured in F₁ and F₂ progeny. The comparison of body size (both traits) between mid‐parent, F₁ and F₂ shows that there is an increase in body size in F₁ and F₂ progeny as compared with parents. Thus, there is no break down of heterosis in F₂, which suggests absence of coadaptation in geographical populations of D. ananassae. Scaling test statistical analysis showed additive, dominance and epistatic effects in certain crosses involving geographical strains of D. ananassae. Correlation between chromosome arrangement frequency and body size has also been tested and significant negative correlation has been found between 2L – ST chromosome arrangement and male thorax.     

Divergence of reaction norms of size characters between tropical and temperate populations of Drosophila melanogaster and D. simulans

Reaction norms to growth temperature of two size-related traits, wing and thorax length, were compared in tropical (West Indies) and temperate (France) populations of the two sibling species, Drosophila melanogaster and D. simulans. A major body size difference was found in D. melanogaster, with much smaller Caribbean flies, while D. simulans exhibited little size variation between geographical populations. The concave norms of reaction were adjusted to second- or third-degree polynomials, and characteristic points calculated i.e. maximum value (MV) and temperature of maximum value (TMV). TMVs were confirmed to be higher for thorax than for wing length, higher in D. melanogaster than in D. simulans, and higher in females than in males. For both traits Caribbean populations exhibited higher TMVs in the two species, strongly suggesting an adaptive shift of the reaction norms toward higher temperature in warm-adapted populations. The wing/thorax ratio was also analysed, and found to be significantly lower in tropical populations of both species. This ratio, which is related to wing loading and flight capacity, might evolve independently of body weight itself.     

Cryptically dimorphic caste differences in a Neotropical, swarm-founding paper wasp genus, Parachartergus (Hymenoptera: Vespidae)

Morphological differences in body shape between females of different reproductive conditions (in terms of insemination and ovarian development) were examined in two species of the Neotropical polistine genus Parachartergus: P. smithii and P. fraternus. The present study shows, for the first time, that non‐size‐based morphological divergence between queens and workers occurs in Parachartergus, an epiponine genus once believed to have little or no morphological caste differences. In the P. smithii colony examined, queens were significantly larger than workers in five of the eight body parts measured (head width, eye width, genal width, mesosomal length, wing length, first metasomal tergum width, and width and length of the second tergum), but the mean values of wing length and first and second tergum widths were not significantly different between them. The queen : worker size ratios tended to be greater anteriorly and smaller posteriorly, although the size ratio was greatest in second tergum length. Analysis of covariance (ancova ) with mesosomal length as covariate showed that queens had proportionally wider heads and narrower first terga than did workers. In the P. fraternus colony, size differences between queens and workers were not significant, and there was little or no difference in shape, but queens had significantly proportionally wider first terga than did workers.