Aggressive behavior of the male parent predicts brood sex ratio in a songbird

Brood sex ratio is often affected by parental or environmental quality, presumably in an adaptive manner that is the sex that confers higher fitness benefits to the mother is overproduced. So far, studies on the role of parental quality have focused on parental morphology and attractiveness. However, another aspect, the partner’s behavioral characteristics, may also be expected to play a role in brood sex ratio adjustment. To test this hypothesis, we investigated whether the proportion of sons in the brood is predicted by the level of territorial aggression displayed by the father, in the collared flycatcher (Ficedula albicollis). The proportion of sons in the brood was higher in early broods and increased with paternal tarsus length. When controlling for breeding date and body size, we found a higher proportion of sons in the brood of less aggressive fathers. Male nestlings are more sensitive to the rearing environment, and the behavior of courting males may often be used by females to assess their future parental activity. Therefore, adjusting brood sex ratio to the level of male aggression could be adaptive. Our results indicate that the behavior of the partner could indeed be a significant determinant in brood sex ratio adjustment, which should not be overlooked in future studies.     

Differential timing and latitudinal variation in sex ratio of Aquatic Warblers during the autumn migration

Differential migration has been extensively reported in spring, but less so in autumn, particularly in relation to sex in monomorphic bird species. Here, we analysed the autumn passage of a monomorphic, globally threatened passerine, the Aquatic Warbler Acrocephalus paludicola throughout Western Europe, with regard to age and sex. We showed that, overall, adults migrated earlier than first-year birds, and males migrated earlier than females during the autumn migration. This may be caused by an overall social dominance of adults over immatures, and differentiated migration strategy of males and females. In addition, we found male-skewed sex proportions, with a tendency to an equalised ratio in more southern stopover sites. This may indicate a male bias in the global population or different migration strategies of the sexes. Differential migration may cause the age and sex classes to be exposed differently to various threats affecting demographic structure of the species.     

Cognitive ability correlates positively with son birth and predicts cross-cultural variation of the offspring sex ratio

Human populations show remarkable variation in the sex ratio at birth which is believed to be related to the parental condition. In the present study, the global variation of sex ratio at birth (SRB, proportion of male offspring born) was analyzed with respect to indirect measure of condition, the intelligence quotient (IQ). IQ correlates strongly with lifespan across nations, which makes it a good indicator of health of the large populations. Relation between three standard measures of average national IQ and SRB was studied using multiple linear regression models. Average national IQ was positively correlated with SRB (r?=?0.54 to 0.57, p?<?0.001). Further, IQ emerged as a powerful predictor of SRB after controlling for the effects of all the known covariates like fertility, maternal age, polygyny prevalence, wealth, son preference, latitude, low birth weight, and neonatal mortality in the regression models. These results suggest that the striking variation of offspring sex ratio across nations could be caused in part by the difference in general condition of populations.     

Sex pheromone component ratios and mating isolation among three Lygus plant bug species of North America

The plant bugs Lygus hesperus, Lygus lineolaris, and Lygus elisus (Hemiptera: Miridae) are major pests of many agricultural crops in North America. Previous studies suggested that females release a sex pheromone attractive to males. Other studies showed that males and females contain microgram amounts of (E)-4-oxo-2-hexenal, hexyl butyrate, and (E)-2-hexenyl butyrate that are emitted as a defense against predators. Using gas chromatography–mass spectrometry, we found that female L. lineolaris and L. elisus have a 4:10 ratio of hexyl butyrate to (E)-2-hexenyl butyrate that is reversed from the 10:1 ratio in female L. hesperus (males of the three species have ~10:1 ratio). These reversed ratios among females of the species suggest a behavioral role. Because both sexes have nearly equal amounts of the major volatiles, females should release more to attract males. This expectation was supported because L. hesperus females released more hexyl butyrate (mean of 86 ng/h) during the night (1800–0700 hours) than did males (<1 ng/h). We used slow-rotating pairs of traps to test the attraction of species to blends of the volatiles with a subtractive method to detect synergism. Each species’ major butyrate ester was released at 3 μg/h, the minor butyrate according to its ratio, and (E)-4-oxo-2-hexenal at 2 μg/h. The resulting catches of only Lygus males suggest that (E)-4-oxo-2-hexenal is an essential sex pheromone component for all three species, (E)-2-hexenyl butyrate is essential for L. elisus and L. lineolaris, and hexyl butyrate is essential for L. hesperus. However, all three components are recognized by each species since ratios of the butyrate esters are critical for conspecific attraction and heterospecific avoidance by males and thus play a role in reproductive isolation among the three species. Because L. hesperus males and females are known to emit these major volatiles for repelling ant predators, our study links defensive allomones in Lygus bugs with an additional use as sex pheromones.     

Offspring sex in a TSD gecko correlates with an interaction between incubation temperature and yolk steroid hormones

We incubated eggs of the Japanese gecko Gekko japonicus at three temperatures, and measured yolk testosterone (T) and 17??-estradiol (E2) levels at three time points in embryonic development (oviposition, 1/3 of incubation, and 2/3 of incubation), to examine whether maternal influence on offspring sex via yolk steroid hormone deposition is significant in the species. Eggs incubated at 24?°C and 32?°C produced mostly females, and eggs incubated at 28?°C almost a 50:50 sex ratio of hatchlings. Female-producing eggs were larger than male-producing eggs. Clutches in which eggs were incubated at the same temperature produced mostly same-sex siblings. Yolk T level at laying was negatively related to eggs mass, and yolk E2/T ratio was positively related to egg mass. Results of two-way ANOVA with incubation temperature and stage as the factors show that: yolk E2 level was higher at 32?°C than at 24?°C; yolk T level was higher, whereas yolk E2/T ratio was smaller, at 28?°C than at 24?°C; yolk E2 and T levels were higher at 2/3 than at 1/3 of incubation. Our data in G. japonucus show that: (1) maternal influence on offspring sex via yolk steroid hormone deposition is significant; (2) incubation temperature affects the dynamics of developmental changes in yolk steroid hormones; (3) influences of yolk steroid hormones on offspring sex are secondary relative to incubation temperature effects; and (4) offspring sex correlates with an interaction between incubation temperature and yolk steroid hormones.     

Selective aggressiveness in European free-tailed bats (Tadarida teniotis): influence of familiarity,age and sex

Bats are highly social mammals that often form large groups and represent good models to test the role played by individual status in shaping social relationships. Social cohesion relies on the ability of group and individual recognition, which is mediated by a range of sensorial cues. In this study, we selected the European free-tailed bat Tadarida teniotis as a model species to test the effects of familiarity, sex and age on aggressiveness and mutual tolerance. We hypothesize that T. teniotis is able to recognize group members and exhibit selective aggressiveness, and thus we predict fewer aggressive events and more amicable encounters between colony mates than between strangers. As female bats are generally more sociable and perform prolonged parental care to juveniles even after weaning, we hypothesize that sex and age of bats have significant influences on aggressive behaviours and thus predict that females will perform more amicable behaviours than males and that adults of both sexes will be less aggressive towards juveniles. Our results confirm that T. teniotis is able to discriminate between familiar and stranger individuals, showing higher rates of aggressive behaviours towards the latter. Females are more prone to exhibit amicable behaviours, particularly during same-sex interactions, while males show higher level of aggressiveness. Juveniles are subjected to fewer aggressive behaviours by adults of both sexes. Familiarity appears crucial for T. teniotis in determining the degree of aggressiveness during social interactions but the rate of aggressive events is also influenced by intrinsic individual factors such as sex and age.     

Differential autumn migration of the aquatic warbler Acrocephalus paludicola

Relatively little attention has been paid to sex differences in the migration of birds in autumn. We studied the autumn migration strategy of molecularly sexed males and females in the globally threatened aquatic warbler Acrocephalus paludicola. We captured 176 birds at a stopover site in the Loire estuary at Donges, France. The median date for the passage of adults was 8 days earlier in males than females, although the timing of migration in first-year males and females was similar. This indicates that males, who are without parental duties, can start their migration earlier than females and first-year birds. Adults were significantly heavier than immature birds but did not have higher fat scores. In both age categories, more males (two to three times more) were captured. However, various factors (including tape-luring) can affect observed sex ratio.     

Forest clearing and sex ratio in forest-dwelling wood ant <Emphasis Type="Italic">Formica aquilonia</Emphasis>

Sex ratios of ants have been shown to vary with food resource levels in several studies, but it is not known whether forest clear-cutting has any effect on sex ratio of aphid-tending forest-dwelling ants. We investigated whether the offspring sex ratio of the forest dwelling ant Formica aquilonia varied as a response to clear-cutting. We found that the proportion of males was smaller in clear-cuts than in adjacent forests. Our results are among the first showing that anthropogenic changes in forest structures may have a potential to modify sex ratios of social insects and other forest-dwelling animals.     

Biased sex ratio and low population density increase male mating success in the bug Nysius huttoni (Heteroptera: Lygaeidae)

Demographic factors such as operational sex ratio (OSR) and local population density (LPD) are temporally and spatially dynamic in the natural environment but the influence of these variables on male mating success and the mechanisms behind it are still poorly understood and highly controversial. Here, we manipulated the OSR and LPD of a seed bug, Nysius huttoni, and carried out a series of mating trials to test how these variables affected male mating success. The two demographic factors had no significant interactions, suggesting that they affect male mating success independently in N. huttoni. In this species male mating success was significantly higher in both male- and female-biased OSR than in even OSR. It is suggested that, in male-biased OSR, the increased intensity of competition and interference does not result in lower male mating success; rather, males may make more effort in courting and females may have more chance to encounter better males, resulting in higher male mating success. In female-biased OSR, females may become less choosy and less likely to reject male mating attempt, leading to the higher male mating success. Lower male mating success in N. huttoni in high LPD may be due to increased interference between males and/or delayed female receptiveness for mating. OSR had a stronger effect on male mating success than LPD in N. huttoni, suggesting that OSR and LPD affect mating success in different ways and intensities.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Reversed sex change by widowed males in polygynous and protogynous fishes: female removal experiments in the field

Sex change, either protogyny (female to male) or protandry (male to female), is well known among fishes, but evidence of bidirectional sex change or reversed sex change in natural populations is still very limited. This is the first report on female removal experiments for polygnous and protogynous fish species to induce reversed sex change in the widowed males in the field. We removed all of the females and juveniles from the territories of dominant males in the cleaner wrasse Labroides dimidiatus (Labridae) and the rusty angelfish Centropyge ferrugata (Pomacanthidae) on the coral reefs of Okinawa. In both species, if new females or juveniles did not immigrate into the territories of the widowed males, some of them emigrated to form male–male pairs. When a male–male pair formed, the smaller, subordinate partner began to perform female sexual behaviours (n = 4 in L. dimidiatus; n = 2 in C. ferrugata) and, finally, released eggs (n = 1, respectively). Thus, the reversed sex change occurred in the widowed males according to the change of their social status. These results suggest that such female removal experiments will contribute to the discovery of reversed sex change in the field also in other polygnous and protogynous species.     

Diploid males, diploid sperm production, and triploid females in the ant Tapinoma erraticum

Under complementary sex determination (CSD), females of Hymenoptera arise from diploid, fertilized eggs and males from haploid, unfertilized eggs. Incidentally, fertilized eggs that inherit two identical alleles at the CSD locus will develop into diploid males. Diploid males are usually unviable or sterile. In a few species, however, they produce diploid sperm and father a triploid female progeny. Diploid males have been reported in a number of social Hymenoptera, but the occurrence of triploid females has hardly ever been documented. Here, we report the presence of triploid females, diploid males, and diploid sperm (produced by diploid males and stored in queen spermathecae) in the ant Tapinoma erraticum. Moreover, we show variations in the frequency of triploids among female castes: Triploid females are more frequent among workers than virgin queens; they are absent among mated, reproductive queens. The frequency of triploid workers also varies between populations and between nests within populations.     

(<Emphasis Type="Italic">Z</Emphasis>)-9-tricosene identified in rectal gland extracts of <Emphasis Type="Italic">Bactrocera oleae</Emphasis> males: first evidence of a male-produced female attractant in olive fruit fly

It is well-known that Bactrocera oleae (olive fruit fly) females attract conspecific males by using 1,7-dioxaspiro[5,5]undecane (1) as the main component of their sex pheromone, and that 1 is produced in the female rectal gland. Although some authors have claimed that B. oleae males also attract females, to date no male-produced female attractants have been found in this species. In this paper, we report the first identification of a substance unique to males and able to attract females. The findings of the study include the following: (1) females responded in a bioassay to hexane extracts obtained from rectal glands of 15-day-old B. oleae males, (2) the presence of (Z)-9-tricosene (2) was consistently and unambiguously identified in these extracts using gas chromatography (GC) and GC-mass spectrometry methods, (3) in preliminary bioactivity tests, low doses (equivalent to a few males) of chemically and stereoisomerically pure synthetic (Z)-9-tricosene (2) attracted olive fruit fly females. Interestingly, compound 2, commonly called muscalure, is also a well-known component of the house fly (Musca domestica) sex pheromone.     

Sex-dependent obesogenic effect of tetracycline on Drosophila melanogaster deteriorated by dysrhythmia

Antibiotics have been identified as obesogens contributing to the prevalence of obesity. Moreover, their environmental toxicity shows sex dependence, which might also explain the sex-dependent obesity observed. Yet, the direct evidence for such a connection and the underlying mechanisms remain to be explored. In this study, the effects of tetracycline, which is a representative antibiotic found in both environmental and food samples, on Drosophila melanogaster were studied with consideration of both sex and circadian rhythms (represented by the eclosion rhythm). Results showed that in morning-eclosed adults, tetracycline significantly stimulated the body weight of females (AM females) at 0.1, 1.0, 10.0 and 100.0 µg/L, while tetracycline only stimulated the body weight of males (AM males) at 1.0 µg/L. In the afternoon-eclosed adults, tetracycline significantly stimulated the body weight of females (PM females) at 0.1, 1.0 and 100.0 µg/L, while it showed more significant stimulation in males (PM males) at all concentrations. Notably, the stimulation levels were the greatest in PM males among all the adults. The results showed the clear sex dependence of the obesogenic effects, which was diminished by dysrhythmia. Further biochemical assays and clustering analysis suggested that the sex- and rhythm-dependent obesogenic effects resulted from the bias toward lipogenesis against lipolysis. Moreover, they were closely related to the preference for the energy storage forms of lactate and glucose and also to the presence of excessive insulin, with the involvement of glucolipid metabolism. Such relationships indicated potential bridges between the obesogenic effects of pollutants and other diseases, e.g., cancer and diabetes.     

Paternal inheritance in mealybugs (Hemiptera: Coccoidea: Pseudococcidae)

Mealybugs have a haplodiploid reproduction system, with paternal genome elimination (PGE); the males are diploid soon after fertilization, but during embryogenesis, the male paternal set of chromosomes becomes heterochromatic (HC) and therefore inactive. Previous studies have suggested that paternal genes can be passed on from mealybug males to their sons, but not necessarily by any son, to the next generation. We employed crosses between two mealybug species—Planococcus ficus (Signoret) and Planococcus citri (Risso)—and between two populations of P. ficus, which differ in their mode of pheromone attraction, in order to demonstrate paternal inheritance from males to F2 through F1 male hybrids. Two traits were monitored through three generations: mode of male pheromone attraction (pherotype) and sequences of the internal transcribed spacer 2 (ITS2) gene segment (genotype). Our results demonstrate that paternal inheritance in mealybugs can occur from males to their F2 offspring, through F1 males (paternal line). F2 backcrossed hybrid males expressed paternal pherotypes and ITS2 genotypes although their mother originated through a maternal population. Further results revealed other, hitherto unknown, aspects of inheritance in mealybugs, such as that hybridization between the two species caused absence of paternal traits in F2 hybrid females produced by F1 hybrid females. Furthermore, hybridization between the two species raised the question of whether unattracted males have any role in the interactions between P. ficus and P. citri.     

A new class of mealybug pheromones: a hemiterpene ester in the sex pheromone of Crisicoccus matsumotoi

Mealybugs, which include several agricultural pests, are small sap feeders covered with a powdery wax. They exhibit clear sexual dimorphism; males are winged but fragile and short lived, whereas females are windless and less mobile. Thus, sex pheromones emitted by females facilitate copulation and reproduction by serving as a key navigation tool for males. Although the structures of the hitherto known mealybug pheromones vary among species, they have a common structural motif; they are carboxylic esters of monoterpene alcohols with irregular non-head-to-tail linkages. However, in the present study, we isolated from the Matsumoto mealybug, Crisicoccus matsumotoi (Siraiwa), a pheromone with a completely different structure. Using gas chromatography–mass spectrometry and nuclear magnetic resonance spectroscopy, we identified the pheromone as 3-methyl-3-butenyl 5-methylhexanoate. Its attractiveness to males was confirmed in a series of field trapping experiments involving comparison between the isolated natural product and a synthetic sample. This is the first report of a hemiterpene mealybug pheromone. In addition, the acid moiety (5-methylhexanoate) appears to be rare in insect pheromones.     

Geographical variation in sexual behavior and body traits in a sex role reversed wolf spider

Mating partners need to recognize, assess each other, and exchange information through behavioral events that occur before, during, and after mating. Sexual signals, as well as life history traits, are influenced by selective pressures and environmental factors that can vary across distant geographical areas. Allocosa senex is a sand-dwelling wolf spider which constructs burrows along the sandy coasts of Argentina, Brazil, and Uruguay. Females are the mobile sex that searches for males and initiates courtship. They prefer males which construct longer burrows, and males prefer virgin females in good body condition. The objective of this study was to compare sexual behavior patterns, as well as body characteristics and burrow dimensions, between two geographically distant locations of A. senex, one in Uruguay (Uruguayan location) and the other from central Argentina (Argentinean location). We found differences in the number of male abdominal vibrations, male and female touches during mating, and number of erections of male leg spines, which all were higher in matings of Argentinean pairs. On the other hand, male body mass and female body condition were higher in Uruguayan individuals. The wide distribution of A. senex could be determining variations in the biotic and abiotic features that affect the species, generating differences in the strength of selective forces acting on individuals from the two studied locations.     

Identification and electrophysiological studies of (4S,5S)-5-hydroxy-4-methyl-3-heptanone and 4-methyl-3,5-heptanedione in male lucerne weevils

An investigation to identify a sex or aggregation pheromone of Sitona discoideus Gyllenhål (Coleoptera: Curculionidae) is presented. Antenna flicking and attraction behaviors evoked by conspecifics of both sexes were recorded in arena bioassays, where attraction of females to males was observed. Air entrainment of both males and females was conducted in separate chambers. Gas chromatographic–mass spectrometric analysis of headspace volatiles revealed that two male-specific compounds, 4-methyl-3,5-heptanedione (major) and (4S,5S)-5-hydroxy-4-methyl-3-heptanone (minor), were emitted during the autumnal post-aestivatory flight period. The stereoisomers of the minor component were separated by enantioselective gas chromatography and their absolute configurations assigned by NMR (diastereomers) and the known preference of enantioselective transesterification reactions catalyzed by Candida antarctica lipase B. Electroantennogram and single sensillum recording studies indicate that 4-methyl-3,5-heptanedione as well as all individual stereoisomers of 5-hydroxy-4-methyl-3-heptanone are detected by the antennae of male and female S. discoideus. Further, single sensillum recordings suggest that both sexes of S. discoideus have specialized olfactory receptor neurons (ORNs) for detecting 4-methyl-3,5-heptanedione and different populations of stereoselective ORNs for detecting the stereoisomers of 5-hydroxy-4-methyl-3-heptanone. Some of these stereoselective ORNs appear to be sex-specific in S. discoideus.     

Male behaviors reveal multiple pherotypes within vine mealybug Planococcus ficus (Signoret) (Hemiptera; Pseudococcidae) populations

The vine mealybug (VM) females collected in Israel produce two sex pheromone compounds: lavandulyl senecioate (LS) and (S)-lavandulyl isovalerate (LI). The males display ambiguous behavior to LI: repulsion in the vineyard and attraction of laboratory-reared males. We addressed the question of individual male behavior, i.e., do males respond to both LS and LI, or might they display a distinct response to each of the two pheromone compounds. We compared male pherotype frequencies between wild-caught and laboratory-reared populations. Then, we examined the relationship between pherotype composition and male capture rates in pheromone traps. Finally, we addressed the heredity of the pherotypes. The Israeli VM populations contain nine different male pherotypes, as defined according to the male behavior to pheromone compounds. The studied Portuguese populations included five of the nine pherotypes; none of the Portuguese males were attracted to LI. It seems that the high frequency of males that were attracted to LI is related to dense VM populations. It is hypothesized that selection for the male pherotypes, I males, those that respond to LI, occur under high-density rearing conditions. This may result from shorter development times of males and females that produce more I male pherotypes. The lower relative frequency of trapping of males in LI-baited traps than expected from the percentage determined in a Petri dish arena suggests that males that respond solely to LS (S males) are better fliers. The results also suggest that the pherotype trait is inherited by both sexes of the VM.     

Male antenna morphology and its effect on scramble competition in false garden mantids

Well-developed antennae are crucial for many insects, but especially for scramble competitors, who race to find their mates using female sex cues. In these systems, the ability of males to locate females quickly is thought to be under strong selection. A rarely tested assumption is that males with more sensory structures are able to locate females faster. In the present study, we used the false garden mantid Pseudomantis albofimbriata to investigate male antennal morphology and its effect on male efficiency in finding a mate. We used scanning electron microscopy to describe the major sensilla types and their arrangement along the length of male antennae. We also conducted field enclosure trials relating male antennal morphology to scramble competition in this system. We identified six different types of antennal sensilla (cheatic, trichoid, basiconic, grooved peg, ceolocapitular and campaniform) on male P. albofimbriata antennae. As expected, males who had more trichoid sensilla located females quicker than did males with fewer sensilla. Results of the current study suggest that antenna morphology plays a significant role in mate location and hence scramble competition in the P. albofimbriata mating system.