Studies on the maturation-accelerating pheromone of the desert locustSchistocerca gregaria (Orthoptera: Acrididae)

Summary The accelerating effect of mature males of the desert locust,Schistocerca gregaria, on the maturation of immature male and female conspecifics was confirmed. The onset of sexual activity was found to correlate with yellowing of the male insects. Using the colour as an indicator of maturation, the maturation-hastening effect of an hexane extract from mature males was also confirmed. Likewise, volatiles from mature males placed in the upper storey of a two-chamber bioassay system (no visual or tactile contact possible) and charcoal-trapped volatiles from the mature males also induced accelerated maturation in recipient males placed in the lower storey. GC and GC-MS analysis of volatiles collected from 4-week-old mature males showed the presence of anisole, benzaldehyde, veratrole, phenylacetonitrile and 4-vinylveratrole, roughly at the ratio 4.8:7.0:3.3:79.8:5.0. In addition, guaiacol, phenol, benzoylnitrile, benzyl alcohol and 2-benzoyloxyphenylacetonitrile were present in smaller amounts. These compounds were either absent or found only in trace amounts in the female volatiles. Earlier, we had shown that essentially the same blend collected from younger males (10–20 d old, not yet demonstrating signs of maturation) induced strong aggregation of both sexes of adult desert locusts, suggesting a parsimonious role for the pheromone system: as an adult aggregation signal and as a maturation accelerant in young adults.     

Mate choice tactics and swarm size: a model and a test in a dance fly

In the dance flyEmpis borealis (L.) (Diptera: Empididae) females gather to swarm and males visit swarms for mating. A model was constructed, based on previously published data, simulating how males may choose among females of different sizes in swarms of different sizes. The focal question was, what influences the number of individuals in the swarm in this and possibly other swarming insects? The relationships between original swarm size and both the number of males arriving per minute and the proportion of males mating are both logarithmic. The model predicted that if these relationships were linear, or if males were able to judge absolute female size, the mean swarm size should increase and be at least four times as large as those found in the field. The only type of male mate choice strategy that gave rise to very large swarms (>25) was size-related choice (if males are able to assess the size of a female in relation to the entire population and not merely to the swarm). Furthermore, no swarming behaviour would occur if males mate independently of swarm size. Thus, the numbers of females attending a given swarm site are influenced by male arrival pattern, male preference for larger swarms, the inability of males to judge the absolute body size of females, and female polyandry. Males searching for mates seem to prefer larger swarms than females searching for a swarm to join, but the mean swarm size is primarily set by the swarm size preference of females. Optimal swarm size predicted from the model was 4.68±0.53 females. In order to test model predictions, 69 natural swarm sites were studied during one season. The mean swarm size was 4.85±4.54 females (median 4.03), and about 90% of swarms consisted of 11 females or fewer. Predicted and observed swarm size did not differ significantly.     

Mating behaviour of male Carcinus maenas in relation to a putative sex pheromone: behavioural changes in response to antennule restriction

Temporary suspension of the receptivity of the male antennules confirmed these organs as the site of putative pheromone detection in the crab Carcinus maenas. Comparative analysis of male performance in individual components of sexual behaviour failed to identify any difference between male crabs with antennules restricted and others without, when individuals from each group were placed in contact with receptive females. However, even though pheromone perception by males was not a prerequisite for the initiation of sexual behaviour, in its absence the control of normal sequence and duration of pairing behaviour was disrupted.     

Host-plant green-leaf volatiles synergize the synthetic sex pheromones of the corn earworm and codling moth (Lepidoptera)

Summary The capture of adult male moths in female sex pheromone traps of two key agricultural pests, the corn earworm (Helicoverpa zea) and the codling moth (Cydia pomonella), is enhanced or synergized by a certain group of host-plant volatiles, the green-leaf volatiles (GLVs). Since female adults of both species call and release their sex pheromones while perched upon the leaves of their host-plants, the volatile constituents from the leaves of a number of host-plants were compared. Sex pheromone traps containing one of the prominent leaf volatiles of certainH. zea hosts, (Z)-3-hexenyl acetate, not only significantly increased the capture ofH. zea males but were preferred over traps baited only with sex pheromone. Similarly, traps baited with synthetic sex pheromome ofC. pomonella plus a blend of GLVs captured significantly more males than traps baited only with sex pheromone. Since male moths are not captured in traps baited only with these GLVs, it appears that these GLVs act as pheromone synergists which increase or enhance the attraction or arrestment of male moths in pheromone traps.     

Sexually differentiated functions of female-produced pheromone of the black chafer <Emphasis Type="Italic">Holotrichia loochooana loochooana</Emphasis> (Sawada) (Coleoptera: Scarabaeidae)

Summary. Both male and female Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) were attracted with female-produced pheromone, anthranilic acid (2-aminobenzoic acid), in the field. Male chafers were observed to apparently directly locate cotton balls impregnated with 1 to 10 mg of pheromone. In contrast, females never directly oriented to the treated balls but landed 0.2-1.5 m away and exposed their abdominal glands in a calling posture, which occasionally resulted in aggregation of both females and males. This suggested the mating aggregation of this species could be primarily induced by pheromone released by females. A hypothesis for adaptability of female aggregation is proposed and discussed.     

Indirect evidence indicates female semiochemicals release male precopulatory behaviour in the snow crab,Chionoecetes opilio (Brachyura: Majidae)

Summary Experiments were conducted in aquaria to test the hypothesis that females of the snow crab,Chionoecetes opilio, release a sex pheromone to attract mates. Males exhibited significantly increased activity to water from a source aquarium containing recently-moulted pubescent females, egg-stripped multiparous females and recently-moulted immature females than to water from an aquarium containing berried multiparous females, eggs alone, adolescent males or an empty aquarium. Males without their cephalic chemoreceptors maintained low activity levels in the presence of recently-moulted pubescent females, whereas maxilla-ablated males reacted as strongly as intact males. These findings are discussed within the context of the ecdysteroid-sex pheromone hypothesis, proposed by Kittredgeet al. (1971) and Kittredge & Takahashi (1972), but subsequently rejected. We propose that ecdysteroids from both pubescent and multiparous females may elicit male search-and-clasp behaviour inC. opilio and that the reproductive biology of species used to refute the hypothesis was inappropriate to test the role of ecdysteroids as a cue in the mating process.     

The effect of male sodium diet and mating history on female reproduction in the puddling squinting bush brown<Emphasis Type="Italic"> Bicyclus anynana</Emphasis> (Lepidoptera)

The males of butterflies transfer a spermatophore to the female during mating that can contain nutrients enhancing the reproductive potential of their partners. The nutrients transferred by males can be derived from both larval and adult feeding. These nutrients may be depleted by multiple matings. An apparent difference in adult feeding behaviour between the sexes is puddling on mud, dung and carrion, which in most butterfly species is exclusively a male behaviour. A possible explanation for this division in feeding behaviour is that nutrients derived from puddling by males are transferred to the female during mating. Here, we test this hypothesis in the African fruit-feeding butterfly Bicyclus anynana. We varied the male nuptial gift by (1) feeding males either a diet with or without sodium, and (2) varying the number of previous successful copulations by remating males up to five times on consecutive days. The results show both a strong effect of order of mating on the mating duration, and an individual effect with some males typically copulating for a shorter time than others. The effects on female reproduction were, however, minimal. The total number of eggs per female and the sodium content of the eggs did not differ significantly between diets, nor were they affected by the mating histories of the males. Eggs showed a non-significant lower hatching for females partnered by a male who had already mated several times. There was an indication of an interaction with male diet: the sodium treatment showing a decline in egg hatchability with order number of male mating, whilst the control treatment showed a constant hatchability. The results are discussed in relation to determinants of male gift-giving strategy and to other potential explanations for a restriction of puddling to males in butterflies.Communicated by M.A. Elgar     

Y-maze experiments on the chemotactic behaviour of the tetrabranchiate cephalopod<Emphasis Type="Italic"> Nautilus pompilius</Emphasis> (Mollusca)

In contrast to the modern cephalopods, the nautiloids use primarily the chemosensory sense to explore their environment. So far there have been few studies on sexual-selection processes in solitary-living nautiloid cephalopods, but it can be posited that conspecifics are also discovered by odour. In order to determine whether a special area of the rectum, the rectal gland, plays a role in the intraspecific communication processes of Nautilus pompilius, Y-maze experiments were performed. We tested the reaction of juvenile, early-adolescent male and female, and adult male N. pompilius to homogenates of the rectum of male and female conspecifics. As negative controls, homogenates of gills or mantle, or seawater were used. To check the set-up of the experiment, carrion was presented as a positive attractant. We demonstrated that the adult (mature) males significantly preferred the rectum homogenate containing the stimulus of females, whereas the homogenates from males and/or females had no influence on the behaviour of immature animals of both sexes. Our behavioural studies provide evidence that sexually mature male N. pompilius are attracted to females by excretions of the rectum.Communicated by O. Kinne, Oldendorf/Luhe     

The aggregation pheromone ofIps duplicatus and its role in competitive interactions withI. typographus (Coleoptera: Scolytidae)

Summary Ips duplicatus withI. typographus co-inhabiting Norway spruce (Picea abies (L.) Karst.) would benefit from a pheromone blend distinct from that of the larger competitorI. typographus. GC-MS analysis showed thatI. duplicatus males feeding in the host produced ipsdienol (Id),cis-verbenol (cV),trans-verbenol (tV), myrtenol (Mt), andE-myrcenol (EM) and traces of 2-methyl-3-buten-2-ol (MB).I. duplicatus produced Id in approximately racemic form (48.9-54.5% (+)-(S)-isomer). The amounts of Id and EM released over a 9 day period had a maximum of 250 and 5 ng/h/male, respectively, on day 2. Exposure ofI. duplicatus males to myrcene and -pinene resulted in the production of small amounts of Id, cV, tV, Mt, andtrans-pinocarveol, but not of EM. In laboratory bioassays with walking beetles, the pheromone component Id alone was weakly attractive while EM was inactive, but in binary combination with Id strongly synergized attraction. A combination of EM and Id at a release rate equivalent to 100–200 males was more attractive in the field than 70 unmated males in a spruce log. The addition of myrcene ( a suggested pheromone precursor of Id) to Id did not enhance trap caches, while addition of EM increased catches > 10-fold. Subtracting EM from a blend of Id, EM, cV and MB drastically reduced trap catches while subtraction of cV or MB or both had no significant effect. Addition of EM over a wide concentration range to the synthetic pheromone ofI. typographus did not reduce the attraction of females of this species in the laboratory. A two-species pheromone interaction field test releasingI. typographus pheromone components (MB + cV) at 10–1000 male equivalents (ME) andI. duplicatus pheromone (Id + EM) at 0, 10–1000 ME in all possible combinations showed both positive intraspecific dose-response effects and an interspecific inhibition. Higher release rates of EM appeared to inhibitI. typographus, especially males. In a tree colonization model, the response of the two competing species to their respective pheromones show a good separation during the mass-attack with a small initial cross-attraction. It remains to be shown whether either of the two pheromone systems have in fact evolved in the present sympatry, or if they are an incidental effect of ancestry of these phylogenetically distantIps.     

Effect of colour on male orientation to female pheromone in the black chafer Holotrichia loochooana loochooana

Summary. Under field conditions significantly more black chafer, Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) males landed on black and gray sources than white ones containing 10 mg of female pheromone, anthranilic acid. When a broader spectrum of colours was tested the frequency of male landing was intensively proportional to as the lightness of the colour of the lures. These findings demonstrated that mate location by H. l. loochooana males is dependent on both olfactory and visual stimuli of the source. In contrast, female aggregation was not affected by colour.     

Spawning behaviour and sex hormone levels in adult and precocious brown trout (Salmo trutta L.) males and the effect of anosmia

Summary. In the present experiment the behaviour and endocrine status of males of the brown trout, Salmo trutta L., (Salmoniformes: Salmonidae) were studied when males were kept in a stream tank with a nest digging female. Groups of mature adult males and precocious intact or anosmic male parr were placed with the nesting female so that the group resembled a natural spawning situation with big anadromous fish acting as dominant males and precocious parr acting as “sneakers”. A control experiment was also run with only males without a female present. In intact parr there were significant positive correlations between the per cent of the total observation time spent with a female, milt volume, and plasma concentration of 17α, 20β-dihydroxy-4-pregnen-3-one. Anosmic parr had significantly lower volumes of strippable milt and gonadal steroid hormone levels compared with intact parr. However, no differences were found in the control experiment. Significantly fewer anosmic parr attended and courted the nesting female and those anosmic fish that attended the female had significantly lower plasma levels of gonadal hormones. Intact parr also displayed a greater number of agonistic acts against other parr without any difference in fighting ability. No differences in aggression occured in the control experiment. In adult males together with a female, post-experimental gonadal steroid hormone levels were higher than pre-experimental levels. Positive correlations between aggression and androgen hormone levels were observed in adult males. No differences in plasma hormone levels were observed between adult males and intact precocious males. The results show that olfactory occlusion results in low steroid hormone levels and milt volumes in precocious males placed in a spawning situation. The courting behaviour was also affected by anosmia. Odours from the nesting female may have caused the enhanced plasma hormone levels and stimulated the males to attend the female. Received 15 May 1997; accepted 29 June 1997.     

Male endogenous pheromonal component of Bactrocera carambolae (Diptera: Tephritidae) deterred gecko predation

Summary. Geckos consumed immature Bactrocera carambolae males readily, but their consumption was reduced significantly when offered sexually mature males. They showed a similar aversion for sexually mature males previously fed with methyl eugenol. When male flies at different ages were offered to starved geckos, geckos’ consumption showed an age-related anti-feeding effect, which was concomitant with the production trend of 6-oxo-1-nonanol, a major endogenous component synthesized in the male rectal gland. The production of 6-oxo-1-nonanol was also age-related and corresponded with sexual maturity as reported previously. Topical application of the authentic chemical of 6-oxo-1-nonanol onto the thorax of female fruit flies, at the amount normally detected in the rectal gland of a sexually mature male, had the same deterrent effect against geckos.     

Differential activity of non-fluorinated and fluorinated analogues of the European corn borer pheromone

Summary.  The differing antagonist activity of (Z)-13-hexadecen-2-one (Z11 – 14 :MK, 1) and its 1,1,1-trifluoro derivative (Z11 –14 :TFMK, 2), two closely related analogues of the European corn borer pheromone Ostrinia nubilalis (Z strain), and their rationale is reported. Both chemicals exhibited some electrophysiological activity, and topical application of 10 pg of pheromone analogue on male antennae was sufficient to induce significantly lower depolarization responses to the pheromone versus untreated insects. In a wind tunnel, the number of European corn borer males attracted to sources containing mixtures of 1 + pheromone in ratios ≥ 1 :1 was significantly lower than the number attracted to a source containing pheromone alone. Source contact behaviour was dramatically impaired when the 1 + pheromone blend reached a ratio of 10 :1, in which only 2% of males displayed source contact in the presence of antagonist. When compound 1 was present at the source, males usually flew upwind with occasional downwind reversals; when compound 2 was present at the lure, males performed wider crosswind reversals, with little progress toward the source. In the field, traps baited with mixtures of both compounds with the pheromone in ratios of 5 :1 and 10 :1 elicited a significantly decreased number of male catches. In esterase inhibition assays, compound 2 was a potent inhibitor (IC₅₀ = 70 nM), whereas the non-fluorinated compound 1 was not. The different activity of both compounds is presumed to be due to different mechanisms of action; considerations for using methyl ketone analogues as new behavioural antagonists of the pheromone are outlined.     

Nest-building by crested tit Parus cristatus males: an analysis of costs and benefits

In a population of crested tits in Belgium, one out of three breeding males participated in nest-building. Since variation in the amount of nest-building among 11 building males was low, analysis of male nest-building allowed a logistic regression design. The probability of building increased with male condition and for late nests. The effect of male condition decreased with date, and date was the only factor predicting nest-building for repeat broods. The effect of male condition on nest-building suggests that building entails important costs for males. Based on correlative data, we argue that building by males shortens the interval between the start of nest-building and the onset of egg-laying by about 5 days. Since clutch size, incubation and nestling time did not differ between early and late broods, male nest-building apparently affects fledging date. Since early fledgers (early dispersers) are more likely to settle in future winter flocks, building behaviour by crested tit males should be favoured by natural selection. So far, male nest-building in hole-breeding birds might have been overlooked due to the widespread use of artificial nest-boxes in behavioural studies.     

Population differentiation in female sex pheromone and male preferences in a solitary bee

Population differentiation in female mating signals and associated male preferences can drive reproductive isolation among segregated populations. We tested this assumption by investigating intraspecific variation in female sex pheromone and associated male odour preferences among distant populations in the solitary bee Colletes cunicularius (L.) by using quantitative gas chromatography and by performing field bioassays with synthetic blends of key sex pheromone compounds. We found significant differences in sex pheromone blends among the bee populations, and the divergence in odour blends correlated positively with geographic distance, suggesting that genetic divergence among distant populations can affect sex pheromone chemistry. Our behavioural experiments, however, demonstrate that synthetic copies of allopatric female sex pheromones were cross-attractive to patrolling males from distant populations, making reproductive isolation by non-recognition of mating signals among populations unlikely. Our data also show that patrolling male bees from different populations preferred odour types from allopatric populations at the two sites of bioassays. These male preferences are not expected to select for changes in the female sex pheromone, but may influence the evolution of floral odour in sexually deceptive orchids of the genus Ophrys that are pollinated by C. cunicularius males.     

Moult in relation to some aspects of reproduction and growth in swarms of Antarctic krill,Euphausia superba

In a 14-d period 38 swarms of Antarctic krill,Euphausia superba Dana, were sampled in an area 55.5 x 55.5 km to the southwest of Elephant Island, at the north of the South Shetland Islands. Moult stage, maturity stage, and size of ca. 100 krill from each swarm were measured. Each of the characteristics varied greatly between swarms. Moulting krill were found in most swarms, but in one swarm all of the krill were just about to moult. Ways in which moulting may act as a possible sorting mechanism are discussed. Data for all the analysed krill (ca. 3000 specimens) were used to investigate the interdependence of moult rate, sexual maturation and growth. While all immature krill moulted at approximately the saine rate in the study, there were significant differences in the moulting rates of mature male and female krill. Gravid female krill continued to moult, although less frequently than mature males. As a consequence males had to attach spermatophores to females after each moult. It is likely that variation in moult rate of females and males was related to the energy expenditure required for ovary development in females and spermatophore production and searching behaviour in males. Spawning and moulting were only partly coupled in gravid females. Spawning appeared to take place predominantly during Moult Stage D₂. There was no evidence of intermoult growth by intersegmental dilation.     

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

Pheromone-based aggregation behaviour of the firebrat, <Emphasis Type="Italic">Thermobia domestica</Emphasis> (Packard) (Thysanura: Lepismatidae)

Summary. We tested the hypothesis that aggregation behaviour of the firebrat, Thermobia domestica (Packard) (Thysanura: Lepismatidae), an inhabitant of enclosed microhabitats, is mediated, at least in part, by a pheromone. Individual insects were released into the central chamber of a 3-chambered olfactometer and test stimuli were placed in lateral chambers. Paper discs previously exposed for 3 days to 10 female, male, or juvenile T. domestica were all preferred by female, male, or juvenile T. domestica over unexposed paper discs, indicating the presence of an aggregation/arrestment pheromone. In additional experiments, frass and scales from female T. domestica, tested singly and in combination, proved not to be the source of the pheromone. Physical contact was required for pheromone recognition, indicating that the pheromone arrests rather than attracts conspecifics. Arrestment by the long-tailed silverfish, Ctenolepisma longicaudata Escherich (Thysanura: Lepismatidae), but not by the common silverfish, Lepisma saccharina L. (Thysanura: Lepismatidae), to T. domestica exposed paper discs suggests closer phylogenetic relatedness between C. longicaudata and T. domestica, than between C. longicaudata and L. saccharina. Whether C. longicaudata or L. saccharina produce an aggregation signal, and whether T. domestica respond to this signal is unknown. Received 10 June 2002; accepted 30 September 2002.     

Precopulatory assessment of male quality in an arctiid moth (Utetheisa ornatrix): hydroxydanaidal is the only criterion of choice

Females of the moth Utetheisa ornatrix (Lepidoptera: Arctiidae) mate preferentially with males that excel in three quantitatively correlated attributes: body mass, systemic content of defensive pyrrolizidine alkaloid (derived from the larval diet), and glandular content of the courtship pheromone hydroxydanaidal (derived from the alkaloid). By so choosing, the females obtain direct phenotypic benefits (alkaloid and nutrient received with the spermatophore), and indirect genetic benefits (genes for large size, a heritable trait). We asked whether the female appraises the courting male on the basis of all three attributes, or whether, as had been postulated, she does so on the basis of the intensity of the pheromonal scent alone. We present data indicating that male possession of hydroxydanaidal is indeed the sole criterion of choice. Females fail to differentiate between males that differ in body mass or alkaloid content if the males lack hydroxydanaidal, but choose between males that are size-matched and alkaloid-free if one of the males has been experimentally endowed with hydroxydanaidal. We show moreover that females are able to differentiate between males that contain unequal quantities of hydroxydanaidal. Females abide by these criteria whether or not they themselves contain alkaloid. Their choice was also unaffected by whether they were confined singly with 2 males in small mating chambers, or were in groups of 10 with 20 males in large flight cages.     

Liver and renal lesions in mercury-contaminated narwhals (Monodon monoceros) from North West Greenland

Narwhals (Monodon monoceros) from North West Greenland are known to bioaccumulate mercury (Hg) in tissues and internal organs. This is postulated to be a health concern and therefore studies were undertaken to conduct a screening study of Hg concentrations and histopathology of liver and renal tissues in a total of 12 specimens. The sample consisted of two sub-adults (one male, one female) and 10 adult (six males, four females) collected in Qaanaaq (Thule) 2010. In liver, Hg mean ± SD was 11.88 ± 10.47 μg/g ww (range: 0.39?31.8 μg/g ww) while the concentrations in kidneys were 1.85 ± 1.20 μg/g ww (range: 0.41?4.03 μg/g ww). There was no marked difference in Hg concentrations between males and females while sub-adults had significantly lower concentrations. The histological examinations of renal tissue showed glomerular capillary dilatation and basement membrane thickening, dilatation, and hyalinization of Bowman's space/capsule and tubular lesions with hyaline casts accompanied by interstitial fibrosis in the kidneys. In liver tissue, portal cell infiltrates and fibrosis, bile duct proliferation, lipid-filled Ito cells, and steatosis were found. There was no marked difference in histological prevalence between males and females and in Hg concentrations in individuals with or without lesions. Four of seven renal lesions and one liver lesion were found in the two sub-adult whales. Based upon these findings, as well as the nature of the lesions, evidence suggests that histopathological alterations were a result of age but that Hg might be an aggravating co-factor in development of renal lesions in particular.