Male-Biased Capture in Light Traps in <Emphasis Type="Italic">Spodoptera exigua</Emphasis> (Lepidoptera: Noctuidae): Results from the Studies of Reproductive Activities

Nocturnal insects show phototactic behavior. Interestingly, males are caught by light traps more often than females in many species. In the present study, the male-biased capture in light traps in the field was evaluated to test the hypothesis that male-biased capture was associated with the reproductive activities of Spodoptera exigua. During a 16-wk. period in 2011, a total of 5075 moths caught by light traps in the field revealed a significant male-biased capture. However, the sex ratio of the natural population was not significantly different from 1:1 from July to October 2011. The ratio of moths caught per hour by light traps showed that females were captured in the first half of the night, whereas the peak capture by light traps for males occurred in the second half of the night. Among the captured adults, the percentage of unmated females was significantly lower than that of the mated females and males. Experiments in the laboratory indicated that the time of activities for both sexes was consistent with the active patterns of oviposition and copulation. The percentage of flight-to-light of the unmated females was significantly lower than that of the mated females and males. Thus, the lower capture of the unmated females, which is associated with reduced positive phototactic responses of S. exigua, probably is one of the reasons resulting in the male-biased sex ratio in light traps.     

Small‐plot studies comparing pheromone and juice baits for mass‐trapping invasive Synanthedon myopaeformis in Canada

Recent introduction of Synanthedon myopaeformis (Borkhausen) (Lepidoptera: Sesiidae) into organic apple‐growing areas of Canada has stimulated research on semiochemical‐based management of this European pest. Replicated, small‐plot (0.16 ha) experiments were conducted to compare sex pheromone, 3Z,13Z‐octadecadienyl acetate (10 mg), Concord grape juice (300 ml), or their combination, as mass‐trapping lures at trap densities equivalent to 12.5, 25, 50, and 100 traps ha^?1. Total numbers of male and female moths removed from test plots increased significantly with trap density in all juice‐based mass‐trapping experiments. In pheromone mass‐trapping experiments, however, total catches of males did not increase significantly as trap densities were increased and catches appeared to plateau with 25–50 traps ha^?1. With pheromone‐based mass‐trapping, significantly fewer males were caught in pheromone‐baited assessment traps at the centre of each mass‐trapping plot than in identical traps in untreated plots. This reduction is indicative of significant trap interference or trap ‘shut‐down’. Increasing the density of juice‐based mass‐trapping had no effect on catches of male or female moths in juice‐baited assessment traps, indicating a short range of attraction and lack of interference between juice traps. Pheromone‐ and juice‐based mass trapping removed similar numbers of males at each trap density tested, respectively, but summed catches of males and females were greatest with juice baits. Combining pheromone and juice into a single mass‐trapping treatment (50 traps ha^?1) did not significantly increase catches of males or females relative to either treatment alone. If a practical bisexual mass‐trapping system is going to be developed for S. myopaeformis, then identification of volatile kairomones in Concord grape juice may be useful.     

Monitoring oriental fruit moth (Lepidoptera: Tortricidae) with sticky traps baited with terpinyl acetate and sex pheromone

Studies in Argentina and Chile during 2010–2011 evaluated a new trap (Ajar) for monitoring the oriental fruit moth, Grapholita molesta (Busck). The Ajar trap was delta‐shaped with a jar filled with a terpinyl acetate plus brown sugar bait attached to the bottom centre of the trap. The screened lid of the jar was inserted inside the trap, and moths were caught on a sticky insert surrounding the lid. The Ajar trap was evaluated with and without the addition of a sex pheromone lure and compared with delta traps left unbaited or baited with a sex pheromone lure and a bucket trap filled with the same liquid bait. Studies were conducted in a sex pheromone‐treated orchard in Argentina and an untreated orchard in Chile. In Chile, the Ajar trap without the sex pheromone lure caught significantly fewer males, females and total moths than the bucket trap, and fewer males and more females than the sex pheromone‐baited delta trap. Total moth catch did not differ between the Ajar trap without a sex pheromone lure and the sex pheromone‐baited trap. Adding a sex pheromone lure to the Ajar trap significantly increased total moth catches to levels not different from those in the bucket trap. However, the Ajar trap with the sex pheromone lure caught significantly more males and fewer females than the bucket trap. In Argentina, the Ajar trap with or without the addition of a sex pheromone lure caught similar numbers of both sexes and total moths as the bucket trap. The sex pheromone‐baited delta trap caught <4% of the number of moths as these three traps. The bucket trap in both studies caught significantly more non‐targets than the delta and Ajar traps. Moth catches in the Ajar trap declined significantly after 2–3 weeks when the bait was not replaced.     

The relative performance of pheromone and light traps in monitoring the seasonal activity of both sexes of the eastern hemlock looper, Lambdina fiscellaria fiscellaria

The seasonal flight activity of both sexes of the eastern hemlock looper, Lambdina fiscellaria fiscellaria Guenée (Lepidoptera: Geometridae) was studied during two consecutive years in Quebec and Newfoundland, using light (L), pheromone (P) and combined light and pheromone (LP) traps. Moth density significantly affected the performance of the different traps, with P traps being more effective at low than high density. However, P trap catches decreased just prior to the onset of female captures, probably as a result of competition between traps and virgin calling females. Nearly all females caught in L and LP traps were already mated and even the first females caught had laid at least half of their egg complement. In Quebec under warm nights, the pattern of male activity occurred at different times, with peak P catches being later in the scotophase than those of L traps, but overall similar numbers of males were caught in both traps. In contrast, under cool nights, males were caught early in the night in both P and L traps, suggesting a strong competition effect between traps, although more males were caught in P than L traps overall. In Newfoundland, the pattern of male captures in L and P traps was similar at both high and low temperatures, so competition between trap types would always be high. Under these conditions P traps were more effective than L traps. Irrespective of the region, year or temperature, significantly more males were captured in LP, with the effect of L and P being additive. In both regions, females responded similarly to L and LP traps with peak activity occurring early in the night. Captures of females were lower than those of males under cool temperatures, suggesting that the temperature threshold for flight is higher for females. The use of L and P traps simultaneously and/or in combination is discussed in relation to integrated pest management programs and ecological considerations.     

Reversed sexual size dimorphism in microtines: Are females larger than males or are males smaller than females?

Summary We analysed sexual size dimorphism for 21 populations of microtine rodents. Female to male size ratio varied considerably among populations from females significantly larger than males (ratio=1.18) to males larger than females (ratio=0.78). In a multiple regression analysis female to male home range size ratio explained 94% of the total variation in body size dimorphism and was the only one of eight independent variables that was selected in a stepwise regression procedure. When females are the larger sex, males have home range sizes much larger than females. We suggest that the relationship between home range size ratio and body weight size dimorphism reflects different selection pressures on males and females in competition for resources and mates.     

Using water bottles for trapping of Indianmeal moths in stored peanuts

Sticky traps baited with sex pheromone are the most common trapping devices used in monitoring of moth pests in food warehouses and food processing. However, these traps only capture males, and it is debatable whether captures of male moths can be used as spatio‐temporal indicators of hot spots of conspecific larvae (only larvae are responsible for damage to food products). Water has been documented as highly attractive to stored product moths, and here we present the first performance data on water bottles as monitoring devices. On average, water bottles caught 15 times more Indianmeal moths [Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae)] than unbaited sticky traps and 74 times more moths than probe traps. We showed that hole size in water bottles had negligible effect on their trapping performance in a naturally infested peanut warehouse. Experimental evaluation of water loss over time showed that smaller holes dramatically reduced water evaporation (less frequent service required), and detergent can be added to the water to reduce moth decomposition without adversely affecting water attractiveness (trap performance). Trap captures of males and females were linearly correlated, and based on quantitative statistical analysis [Spatial Analysis by Distance IndicEs (SADIE)], we showed that weekly captures of the two sexes were spatially correlated. The applied implications of using water bottles in improved IPM of moths in food facilities are discussed.     

Xestia communities (Lepidoptera, Noctuidae) in eastern Finnish Forest Lapland as indicated by light trap sampling

Xestia moths (Lepidoptera, Noctuidae, subgenera Anomogyna and Pachnobia ) were collected with eleven light traps installed in various virgin habitats in the Värriötunturi fell area, E Finnish Forest Lapland (67° 44'N, 29° 37'E). Seven Xestia species were caught during the years 1978–84. The most abundant species, X. tecta and X. alpicola inhabited the widest range of habitats, from spruce-covered ravines up to the treeless summit. The other species were almost entirely caught in the sprucecovered ravine. The largest catches were obtained in odd years, although some moths were also caught in the even years. There was a striking male dominance. The trap catches were separated into three groups, corresponding to the original classification of the trap habitats. Of the habitats, the open pine forest was nearest to the summit and the spruce forest farthest away. X. alpicola and X. tecta appear to be ecologically related species, while X. gelida and X. laetabilis are located at the other end of this ecological scale.     

Effect of trap design, trap size and pheromone dose on male capture of two pine bast scales species (Hemiptera: Matsucoccidae): implications for monitoring and mass-trapping

Abstract 1 The effects of pheromone dose, trap size and trap design on the capture of male Matsucoccus feytaudi were studied in maritime pine forests of Portugal, France and Italy, and Matsucoccus josephi in Aleppo pine stands of Israel. 2 Males of both species respond to racemic mixtures of the respective sex pheromone at a dosage as low as 25 µg. However, for low population densities of M. feytaudi, a 50 µg dosage was needed to guarantee male catches significantly different from the control trap. 3 Male capture increased with increasing dosage up to a threshold level for both species. Overdose repellence was not observed even with baits containing 1600 and 2200 µg of the pheromone of M. josephi and M. feytaudi, respectively. 4 For M. feytaudi, a higher dose–response was observed at medium population densities, whereas lesser captures were registered at low and high population densities, suggesting female competition in the latter case. 5 Catches of M. feytaudi males were not affected by trap design, whereas M. josephi males were caught in significantly greater numbers in delta traps. Large traps caught significantly more males of both species. 6 The relative higher male catches in the marginal zone of the sticky traps is probably related to males landing behaviour in the vicinity of the pheromone source. 7 The shape and size of the trap did not affect the bias of the estimates of male catches. However, the plate traps provided higher precision. Both bias and precision improved with increasing dose.     

Female‐targeted attractant containing pear ester for Synanthedon myopaeformis

When testing pear ester (ethyl‐2,4‐decadienoate) + acetic acid (PEAA) lures to catch codling moths, Cydia pomonella (L.) (Lepidoptera: Tortricidae), in Hungary, significant catches of the apple clearwing moth, Synanthedon myopaeformis (Borkhausen) (Lepidoptera: Sesiidae), were also recorded. This sesiid is one of the most important pests of apple in Europe. Pear ester plus acetic acid lures were attractive to S. myopaeformis no matter whether the two compounds were provided in separate dispensers or mixed together in a single dispenser, and a large percentage (40–80%) of the clearwing moths caught were females. In all cases, traps baited with binary combinations of PEAA caught far more than traps baited with either of the compounds presented alone. Traps with PEAA lures in some tests caught (females and males together) up to ca. 20% of the catch in traps baited with the synthetic apple clearwing moth sex attractant (all males). Consequently, the PEAA lure shows potential for future practical applications as a female‐targeted lure. To our knowledge, this is the first report of attractiveness of a lure containing pear ester for non‐tortricid Lepidoptera. Our finding suggests that the compound may be exploited as a host location stimulus by a wider array of insects than was indicated previously.     

小木蠹蛾性行为和性信息素产生与释放的时辰节律

观察了小木蠹蛾Holcocerus insularis的性行为反应,并采用腺体提取、空气收集 、触角电位和田间试验等方法对雌蛾产生和释放性信息素的时辰节律进行了研究。结果表明： (1) 该虫羽化24 h后性成熟,婚飞和交配活动主要在1：00～4：00,交配历时15～45 min;(2) 大部分雌蛾一生交配1～3次,雄蛾多数一生只交配1次,雌雄比为1∶0.89; (3) 雌蛾腺体提取物中性信息素含量同蛾龄有关,2日龄雌蛾腺体性信息素含量最高;(4) 雌蛾腺体中性信息素含量在1：00时最高,而性信息素释放高峰在2：30。     

Resurrecting the differential mortality model of sexual size dimorphism

In some animal groups males may be several times smaller than females. One of the hypotheses proposed to explain the evolution of this extreme sexual size dimorphism (SSD) is the differential mortality model (DMM), which is based on the assumption that when males are the searching sex, higher male mortality relaxes male–male contest competition, leading to the adaptive evolution of early‐maturing, small males that are favoured by viability selection. Evidence for the main prediction of this model, i.e. that there is a negative relationship between differential mortality and SSD, has remained elusive. Using sex differences in pitfall trap catches – a proxy of sex differences in mobility and mortality – across 40 spider species, and using the evolutionary comparative method, we found significant negative relationships between differential mortality and SSD for three size traits. Thus, the DMM can still explain part of the observed variance in SSD.     

The role of fecundity and sexual selection in the evolution of size and sexual size dimorphism in New World and Old World voles (Rodentia: Arvicolinae)

Evolutionary ecologists dating back to Darwin (1871) have sought to understand why males are larger than females in some species, and why females are the larger sex in others. Although the former is widespread in mammals, rodents and other small mammals usually exhibit low levels of sexual size dimorphism (SSD). Here, we investigate patterns of sexual dimorphism in 34 vole species belonging to the subfamily Arvicolinae in a phylogenetic comparative framework. We address the potential role of sexual selection and fecundity selection in creating sex differences in body size. No support was found for hyperallometric scaling of male body size to female body size. We observed a marginally significant relationship between SSD and the ratio of male to female home range size, with the latter being positively related to the level of intrasexual competition for mates. This suggests that sexual selection favours larger males. Interestingly, we also found that habitat type, but not mating system, constitutes a strong predictor of SSD. Species inhabiting open habitats – where males have extensive home ranges in order to gain access to as many females as possible – exhibit a higher mean dimorphism than species inhabiting closed habitats, where females show strong territoriality and an uniform distribution preventing males to adopt a territorial strategy for gaining copulations. Nonetheless, variation in the strength of sexual selection is not the only selective force shaping SSD in voles; we also found a positive association between female size and litter size across lineages. Assuming this relationship also exists within lineages (i.e. fecundity selection on female size), this suggests an additional role for variation in the strength of fecundity selection shaping interspecific differences in female size, and indirectly in SSD. Therefore our results suggest that different selective processes act on the sizes of males and females, but because larger size is favoured in both sexes, SSD is on average relatively small.     

Sexual dimorphism in leg length among orb-weaving spiders: a possible role for sexual cannibalism

The degree and direction of sexual dimorphism across different species is commonly attributed to differences in the selection pressures acting on males and females. The extent of these differences is especially apparent in species that practise sexual cannibalism, where the female attempts to capture and eat a courting male. Here, we investigate the relationship between sexual dimorphism in size and leg length, sexual cannibalism and courtship behaviour in three taxonomic groups of orb-weaving spiders, using morphological data from 249 species in 36 genera. Females are larger than males in all three taxonomic groups, and males have relatively longer legs than females in both the Araneinae and Tetragnathidae. Across genera within each taxonomic group, male body size is positively correlated with both female body size and male leg length, and female body size is positively correlated with female leg length. Sexual size dimorphism is negatively correlated with relative male leg length within the Araneinae, but not within either the Tetragnathidae or the Gasteracanthinae. There was no negative correlation between sexual size dimorphism and relative female leg length in any taxonomic group. We argue that the relationship between sexual size dimorphism and relative male leg length within the Araneinae may be the result of selection imposed by sexual cannibalism by females.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

Sexual dimorphism, extrapair fertilizations, and operational sex ratio in great frigatebirds (Fregata minor)

Across taxa, the presence of sexual ornaments in one sex isusually correlated with disproportionately great parental effortby the other. Frigatebirds (Fregatidae) are sexually dimorphic,with males exhibiting morphological and behavioral ornaments,but males and females share in all aspects of parental effort.All other taxa in a clade of 237 species exhibit biparentalcare, but only frigatebirds exhibit pronounced sexual dimorphism. We tested for the presence of two factors that could contributeto the evolution of male ornaments in great frigatebirds: ahigh frequency of extrapair fertilizations and a male-biasedoperational sex ratio. In 92 families sampled over two breedingseasons, there was only one extrapair fertilization. However,in both seasons, there were more males than females availablefor mating, and the sex ratio among individuals actively engagedin mate-acquisition behavior was strongly male biased, withtypically five or six males available per female. Our resultssuggest that extrapair fertilizations are not responsible forthe exaggeration of sexual ornaments in male frigatebirds,and that operational sex ratio may be related to sexual dimorphismin this species. Further work is needed to determine whetherthe male-biased operational sex ratio creates the variancein male reproductive success that would be needed to drivethe evolution of male ornaments.     

Sex-specific spawning behavior and its consequences in an external fertilizer

Identifying the target of sexual selection in externally fertilizing taxa has been problematic because species in these taxa often lack sexual dimorphism. However, these species often show sex differences in spawning behavior; males spawn before females. I investigated the consequences of spawning order and time intervals between male and female spawning in two field experiments. The first involved releasing one female sea urchin's eggs and one or two males' sperm in discrete puffs from syringes; the second involved inducing males to spawn at different intervals in situ within a population of spawning females. In both, fertilization success was measured as the fraction of eggs fertilized and the paternity share of each male. The results indicate that spawning after females imposes a cost on males but only during sperm competition. Further, the optimal interval between the initiations of male and female spawning depends on degree of sperm competition, distance between males and females, and water velocity. The results show that sex differences in spawning timing of marine invertebrates can be explained on the basis of the differential costs and benefits of spawning out of synchrony with the other sex and that the result of sexual selection on external fertilizers may be behavioral rather than morphological differentiation of the sexes.     

The Ontogeny of Sexual Size Dimorphism of a Moth: When Do Males and Females Grow Apart?

Sexual dimorphism in body size (sexual size dimorphism) is common in many species. The sources of selection that generate the independent evolution of adult male and female size have been investigated extensively by evolutionary biologists, but how and when females and males grow apart during ontogeny is poorly understood. Here we use the hawkmoth, Manduca sexta, to examine when sexual size dimorphism arises by measuring body mass every day during development. We further investigated whether environmental variables influence the ontogeny of sexual size dimorphism by raising moths on three different diet qualities (poor, medium and high). We found that size dimorphism arose during early larval development on the highest quality food treatment but it arose late in larval development when raised on the medium quality food. This female-biased dimorphism (females larger) increased substantially from the pupal-to-adult stage in both treatments, a pattern that appears to be common in Lepidopterans. Although dimorphism appeared in a few stages when individuals were raised on the poorest quality diet, it did not persist such that male and female adults were the same size. This demonstrates that the environmental conditions that insects are raised in can affect the growth trajectories of males and females differently and thus when dimorphism arises or disappears during development. We conclude that the development of sexual size dimorphism in M. sexta occurs during larval development and continues to accumulate during the pupal/adult stages, and that environmental variables such as diet quality can influence patterns of dimorphism in adults.     

Female moths of cotton bollworm (Lepidoptera: Noctuidae) captured by waterbasin traps baited with synthetic female sex pheromone

Cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), is one of the most important pest insects in cotton fields in China. Female moths were captured by waterbasin traps with a synthetic female sex pheromone blend in cotton fields over three years. The blend contained (Z)‐11‐hexadecenal and (Z)‐9‐hexadecenal with a ratio of 97:3. Each pheromone dispenser was impregnated with 2.0 mg of pheromone blend and 0.2 mg of antioxidant dissolved with 0.1 mL of hexane, and there was a control dispenser with a similar amount of antioxidant and solvent only. Waterbasin traps were deployed in three configurations in the fields. ‘A’ was pheromone traps only, ‘B’ was both pheromone and control traps, ‘C’ was control traps only. (i) In four plots of ‘A’, the average weekly female catch was 1.5, and more females were captured by centrally located pheromone traps, (ii) In three plots of ‘Brsquo;, control traps also captured female as well as male moths, but average weekly female catches of control traps was significantly lower than that in pheromone‐baited traps. (iii) There were significant linear relationships between the average weekly female catch and the corresponding layer in pheromone‐baited traps in both ‘A’ and ‘B’ plots, and in quadratic equations in control in ‘B’ plots. (iv) With the increase of the interval of traps, average weekly female catches per trap increased but average weekly female catches per hectare decreased. (v) Among the female moths captured by pheromone traps, 88.3% were mated female moths which each containing 1.46 spermatophores, while in control traps 86.9% of the mated female moths had 0.90 spermatophores. There was a significant difference between the average numbers of spermatophores of mated females in pheromone traps and in controls.     

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.     

Communication interference in sympatrically occurring moth species

In moth species, females emit a species‐specific sex pheromone that is perceived over long distance by conspecific males. The species‐specificity in the chemical communication channel is achieved by a combination of unique components in specific ratios and sometimes also by interspecific behavioural antagonists to deter sympatrically occurring heterospecific males. In this study, we determined possible antagonistic effects in Helicoverpa gelotopoeon Dyar (Lepidoptera: Noctuidae) males to the major sex pheromone component of sympatrically occurring heliothine moths, Z11‐16:Ald, as well as to the sex pheromone of the sympatrically occurring Heliothis virescens (Fabricius) (Lepidoptera: Noctuidae) (Z11‐16:Ald and Z9‐14:Ald). We also explored whether other co‐occurring species are attracted to these pheromone blends. Our field experiments showed that the addition of Z11‐16:Ald alone or in combination with Z9‐14:Ald inhibited trap catches of H. gelotopoeon males and that this inhibition depended on the concentration of these compounds. In addition, other moth species were attracted to the blends. Together, our results confirm the antagonistic effect of heterospecific sex pheromone compounds of H. virescens to H. gelotopoeon.