Response of the predatorRhizophagus grandis to host plant derived chemicals inDendroctonus micans larval frass in wind tunnel experiments (Coleoptera: Rhizophagidae,Scolytidae)

Summary Adults of the predatory beetleRhizophagus grandis are strongly attracted to both adult and larval frass of its specific prey,Dendroctonus micans, in walking bioassays. Spruce bark and resin are relatively unattractive. Solvent extracts of larval frass that were attractive toR. grandis adults in a flight wind tunnel contained a mixture of monoterpenes of host plant origin. A synthetic mixture of these monoterpenes, (+)—-pinene, (–)—-pinene, -phellandrene,dl limonene and 3-carene, was responsible for 70–80% of the activity of the most attractive extract. We suggest that a blend of monoterpenes in frass acts as a kairomone forR. grandis but that attraction to monoterpenes is only the first of a sequence of behavioural responses that ensures successful host location and identification.     

A whole-animal bioassay for the determination of the food attractants of the lobster Homarus gammarus

A semi-quantitative assay is described for assessing the effectiveness of bait extracts for the lobster Homarus gammarus (L.), using an extract of the squid Loligo vulgaris Lamarck as a standard. Fractionation of this extract indicated that, while no single component was as attractive to lobsters as the whole, the recombination of the various components yielded a solution which was as attractive as the original extract. It appears likely that several different classes of chemosensory cell must be simultaneously stimulated for the initiation of feeding behaviour in the lobster.     

Mating expenditures reduced via female sex pheromone modulation in the primitively eusocial halictine bee, Lasioglossum (Evylaeus) malachurum (Hymenoptera: Halictidae)

The present paper reports on behavioral experiments and gas chromatographic analysis of chemical communication in the mating biology of the primitively eusocial sweat bee Lasioglossum (Evylaeus) malachurum. In a dual-choice experiment, a female made odorless was significantly less attractive than an untreated one. Attraction in L. (Evylaeus) malachurum is therefore mediated by a female-produced sex pheromone. Further bioassays showed that unmated gynes are more attractive to males than mated ones. Males are able to differentiate between the two groups of females as little as 3 h after mating. Biotests with different samples obtained from attractive gynes showed surface extracts to be most attractive. Behavioral tests with synthetic copies of the compounds identified as cuticular constituents of virgin gynes were highly attractive to males; the volatile bouquets consisting of n-alkanes, n-alkenes and iso- pentenyl esters of unsaturated fatty acids were the most attractive samples. Isopentenyl esters of unsaturated fatty acids were the key compounds in inducing male inspections as well as stimulating pounces and copulatory attempts. Virgin and nesting gynes differed clearly in the relative and absolute amounts of the volatiles on the cuticle. The total amount of volatiles was significantly higher in virgin gynes and decreased in breeding queens. Hydrocarbons were the dominant group of compounds in both groups of females. The relative amounts of the wax-type ester, hexadecyl oleate, iso- pentenyl esters, and a hitherto unidentified steroid were higher in attractive virgin gynes, while the relative proportions of hydrocarbons and lactones dominated in nesting queens. The site of sex pheromone production in attractive young L. (Evylaeus) malachurum gynes remains unknown. Head glands or Dufour's gland secretions may be involved. Another possible source of the `active principle' found among the cuticular lipids could be glandular cells of the epidermis. The significance of modulation of female sex pheromone composition is discussed in terms of a reduction in mating expenditures. Received: 30 April 1998 / Accepted after revision: 24 July 1998     

Identification of polyenic hydrocarbons from the northern winter moth, Operophtera fagata, and development of a species specific lure for pheromone traps

Summary. In order to elucidate the composition of the female sex pheromone of the northern (beech) winter moth, Operophtera fagata Scharf. (Lepidoptera: Geometridae), ovipositor extracts of unmated, calling females were analysed by gas chromatography with simultaneous electroantennographic and flame ionization detection (GC-EAD/FID). Male antennal responses indicated three active components, two of which had distinct matching peaks in the FID trace. Using coupled gas chromatography- mass spectrometry (GC-MS), these two compounds were identified as (9Z)-nonadecene (9Z-19:Hy), and (6Z,9Z)-nonadecadiene (6Z9Z-19:Hy), respectively. The third component, present in very small amounts only, was identified as (1,3Z,6Z,9Z)-nonadecatetraene (1,3Z6Z9Z-19:Hy), known as the sex pheromone of the common winter moth, O. brumata. Field tests revealed that traps baited with 6Z9Z-19:Hy and 1,3Z6Z9Z-19:Hy caugth large numbers of male O. fagata. Both compounds were found to be essential for attraction of O. fagata. In addition, the diene prevented captures of co-occurring O. brumata. In contrast, 9Z-19:Hy neither influenced the attractiveness of the two-component mixture towards O. fagata nor contributed to bait specificity. A binary mixture of 6Z9Z-19:Hy and 1,3Z6Z9Z-19:Hy in a ratio of 10:1, applied to pieces of rubber tubing, constituted a highly attractive and species-specific bait for O. fagata, which can be used for monitoring of the flight of this defoliator pest of deciduous forests.     

Male-biassed response of garden chafer, Phyllopertha horticola L., to leaf alcohol and attraction of both sexes to floral plant volatiles

Summary. Field tests were performed to evaluate the response of the garden chafer, Phyllopertha horticola (Coleoptera: Scarabaeidae) to known scarab attractants and to investigate the influence of trap colour and height on the number of captured beetles. Both sexes were attracted by a lure mixture composed of geraniol, eugenol, and 2-phenylethyl propionate (PEP) (ratio 3:7:3). When testing floral volatiles individually, only geraniol, eugenol and methyl anthranilate but not PEP was attractive. Response of garden chafer to (Z)- 3-hexen-1-ol was strongly male-biassed. Both the response to floral volatiles and (Z)-3-hexen-1-ol was increased by using funnel traps with yellow instead of grey vanes. Traps positioned at 50 and 125 cm above ground captured significantly more garden chafers than those at 200 cm. The strongly male-biassed response to (Z)-3-hexen-1-ol suggests that in P. horticola (subfamily Rutelinae) orientation towards plant volatiles emitted upon mechanical damage of plants is part of the male mate finding strategy as recently demonstrated for cockchafers of the genus Melolontha (subfamily Melolonthinae). Possible application of plant volatiles for control of P. horticola is discussed.     

Pheromone of the elm bark beetle Scolytus laevis (Coleoptera: Scolytidae): stereoisomers of 4-methyl-3-heptanol reduce interspecific competition

Stereoisomers of 4-methyl-3-heptanol (MH) are pheromone components of several Scolytus bark beetles. The elm bark beetle Scolytus laevis (Coleoptera: Scolytidae) has in previous studies been caught in traps baited with commercial MH containing all four stereoisomers, but the lure has been considered a weak attractant. In this study, we addressed the question whether stereospecific responses by S. laevis to stereoisomers of MH might contribute to its niche separation from other sympatric Scolytus species. Using GC–MS, we analyzed extracts of hindguts and abdomens from male and female S. laevis and the sympatric S. triarmatus. We also tested all four MH-stereoisomers individually and in combinations in the field to determine their role for S. laevis. All four stereoisomers were synthesized via a boronic ester method with 1,2-dicyclohexylethanediol as chiral director. In addition, the (3S,4R)-stereoisomer of MH was prepared through enantioselective, lipase-mediated transesterification of a mixture of the four stereoisomers of MH. Females of both species contained small amounts of syn-MH, and males contained trace amounts of anti-MH. The anti stereoisomer (3R,4S)-MH was attractive to male and female S. laevis, whereas the syn stereoisomer (3S,4S)-MH acted as an inhibitor or deterrent and reduced the catch when added to the attractive isomer. The syn isomer is the main aggregation pheromone component of the larger and sympatric S. scolytus and possibly also of S. triarmatus. The avoidance response of S. laevis to the (3S,4S)-stereoisomer may reduce interspecific competition for host trees.     

Sex pheromone composition and reproductive isolation in two Busseola species (Lepidoptera: Noctuidae) in Kenya

Closely related species provide an interesting and useful model to understand mate communication diversification. Pre-mating isolation is the result of selection processes and is achieved by utilization of different sex pheromones, but also by different ecological and life traits such as geographic distribution, host-plant specialization and allochrony of sexual behavior. Here, we report sex pheromone identifications and pre-mating isolation of two closely related species of stemborers: Busseola segeta and the yet undescribed Busseola nov. sp., developing on different host-plants and in different geographical locations in Kenya. GC–MS analyses showed that the two species produced the same pheromone components, but in slight different ratios. The respective synthetic pheromone mixtures tested in the field were attractive for males of each species. The low inter-specific variation in the pheromone mixture would allow possible interbreeding in case of accidental introduction of both species in the same environment. Under laboratory conditions, allochrony of the mating period is total and might prevent interbreeding if confirmed in natural conditions. This point would ensure the reproductive isolation between the two species in case of modifications of ecological factors. This study and the chemical structures of the identified pheromone components clearly demonstrate that these Busseola species are closely related although they differ in morphology.     

Permanence of Euscorpius carpathicus (L.) larvae on the mother's back (Scorpiones,Chatidae)

Summary The tendency of Euscorpius carpathicus larvae to remain on immobilized live or dried scorpions of the same or different species (Euscorpius flavicaudis) has been tested. Dried scorpions are less attractive than live ones, which are equally attractive regardless of species, sex or reproductive phase. The larvae find old (one year) dried specimens less attractive than freshly or recently (15 days) killed females, which are equally as attractive as live mothers (Table 1). A substratum treated with a chloroform extract of the mother's cuticle is consistently preferred by the larvae. The maternal behaviour (tolerating the larvae on the back) usually disappears about a week after the moult of the larvae, but it can be protracted if they are continually replaced by younger larvae (Table 2). The survival rate of larvae on live mothers is higher than on dried specimens.     

Responses of Metaphycussp. nr. flavus to semiochemicals released from a scale host, Coccus hesperidum

Summary. Metaphycus sp. nr. flavus (Encyrtidae: Hymenoptera) is a parasitoid species collected from the Mediterranean region which lays its eggs in the immature stages of several economically important soft scale insects (Hemiptera: Coccidae), including brown soft scale, Coccus hesperidum L. (= host insect). Preliminary tests suggested that the parasitoid is most successful in producing offspring when it oviposits in the younger stages of brown soft scale. In Y-olfactometer bioassays measuring wasp choices and residence times, naïve parasitoids were significantly more attracted to yucca leaves infested with 26, 27, or 28 d-old scale than to uninfested leaves, whereas leaves with older (29-30 d-old) scale were no more attractive than uninfested leaves. Parasitoids also spent significantly more time in the arm with yucca leaves infested with 26 d-old scale than in the arm with uninfested leaves. These results are consistent with observations of the parasitoids reproductive success on scale of different ages, whereby older scale are more likely to encapsulate the developing eggs of M. sp. nr. flavusfemales than are younger scale. Further bioassays determined that yucca leaves that had been infested with 26 d-old scale but from which the scale had been removed were as attractive as infested leaves. In contrast, infested yucca leaves from which scale had been removed and the leaves subsequently washed with distilled water were less attractive than infested leaves. Furthermore, the wash water containing scale residues was attractive to female wasps. In total, these results suggest that Metaphycussp. nr. flavus females utilize volatile, water soluble compounds produced by brown soft scale as cues to locate suitable hosts.     

Call matching in the quacking frog (Crinia georgiana)

Males of the quacking frog Crinia georgiana produce calls consisting of 1–11 notes. Playback experiments using synthetic calls showed that males tend to match the number of notes in 2-note and 4-note stimuli; however, males tended to produce more than 1 note in response to a 1-note stimulus and fewer than 8 notes in response to an 8-note stimulus. Successive playbacks of two, 4-note calls from separate speakers indicate that males are likely to match the combined number of notes in the calls of two neighbors, even if they are not equidistant from the focal male. The results are compared with the few other studies of matching in anurans, and interpreted in terms of hypotheses developed to explain matching in songbirds. One attractive and testable hypothesis for call matching in C. georgiana is that males are attempting to produce calls that are at least as attractive to females as those of rivals, without wasting energy. Received: 14 February 2000 / Received in revised form: 11 April 2000 / Accepted: 3 May 2000     

Yolk androgen deposition as a compensatory strategy

The deposition of androgens into the yolks of eggs can have long-lasting effects on the growth and development of young birds. It has been proposed that female birds and reptiles deposit yolk androgens according to the differential allocation hypothesis (DAH), which posits the allocation of more resources to offspring sired by more attractive, higher-quality males. We examined deposition patterns of yolk androgens in relation to mate attractiveness in the house finch Carpodacus mexicanus. Contrary to the predictions of the DAH, female house finches deposited significantly more androgens into eggs sired by less attractive males. We propose that, rather than serving as resources, androgens are used as mediators in a compensatory distribution strategy, enabling females to improve the quality of young produced with less attractive males.     

Effects of three phamaceuticals on the responses of tadpoles to predator alarm cues

The aim of this study is to measure the effects of two selective serotonin reuptake inhibitors, fluoxetine and sertraline, and one selective serotonin and norepinephrine reuptake inhibitor, venlafaxine, on the swimming and lateralization behaviour of Sclerophrys arabica tadpoles exposed to predator alarm cues. Tadpoles were exposed to the three pharmaceuticals either at 0.5 µg/L or 2.0 µg/L, either individually or as a mixture for 14 days. Control tadpoles and those exposed to high concentration of drugs individually or as a mixture, and those exposed to the low-concertation mixture reduced their swimming speed. Tadpoles exposed to 0.5 µg/L fluoxetine or venlafaxine did not respond to the alarm cues. Tadpoles exposed to the mixture had similar responses to that of the control. Results indicate that these drugs have an additive mode of action. Tadpoles exposed to the low-concentration mixture increased lateralization of movement.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Efficacy of created and restored nesting sites for the conservation of colonial Laridae in the South of France

By rapidly modifying key habitat components, habitat restoration is at risk of producing attractive cues for animals without providing habitats of sufficient quality. As such, individual fitness components, such as reproduction, could be reduced and restored habitats could become ecological traps. This risk notably appears by using artificial constructions in restoration projects, yet few studies have evaluated their efficacy in a robust way. We investigated this by analyzing 154 islets that were created or restored to improve the conservation status of 7 colonial Laridae species in the South of France. From 2007 to 2016, we compared occupancy dynamics and breeding parameters of these species between the restored sites and 846 unmanaged nesting sites. We also explored species’ preference for different nesting site characteristics and their respective effect on breeding parameters. Restored nesting sites were 2–9 times as attractive as unmanaged sites for all species except the Black-headed Gull (Chroicocephalus ridibundus). Colonization probability was up to 100 times higher in sites already used by other species the previous year and increased with distance to the shore until >0.2 when distance was over 250 m. Abandonment probability was 29–70% lower when breeding was successful the previous year in all species except the Sandwich Tern (Thalasseus sandvicensis). Productivity and breeding success probability were 2 times higher on managed sites. Distance from the shore was an important attractive characteristic of artificial nesting sites in all species. Other nesting site characteristics had species-specific effects on colonization, abandonment, and breeding success. Our results indicate that managed nesting sites are successful conservation tools for colonial Laridae in the Mediterranean and do not act as ecological traps. Our study showed that testing the ecological trap hypothesis is a robust way to evaluate the success of restoration projects of breeding habitats.     

Sex pheromone components of pyralid moths Terastia subjectalis and Agathodes ostentalis feeding on the coral tree, Erithrina variegata: Two sympatric species share common components in different ratios

Summary. Two common components, identified as (E)-11-hexadecenal (E11-16:Ald) and (E,E)-10,12-hexadecadienal (E10E12-16:Ald), were found in the extract of virgin females of two sympatric pyralid moths, Terastia subjectalis Lederer, and Agathodes ostentalis (Geyer). The amount of E11-16:Ald and E10E12-16:Ald was 0.12 and 2.8 ng/female for T. subjectalis and 2.0 and 1.8 ng/female for A. ostentalis, respectively. Hexadecanal (16:Ald) was also found at 0.7 ng/female in the extract of A. ostentalis, but EAG activity was unclear. In the field, T. subjectalis and A. ostentalis males were captured with species specific blends in the proportions of 5:95 and 50:50 blends. Blends attractive to one species did not attract the opposite one. Rubber septa loaded with 1 mg each of 5:95 and 50:50 blends of E11-16:Ald and E10E12-16:Ald were as attractive as two virgin females of T. subjectalis and A. ostentalis , respectively. This demonstrated that different ratios of shared pheromone components could provide species specific cues for critical mate location in the two sympatric pyralid moths. It is suggested that directional selection probably occurs in the pheromone blends in opposite directions in both species.     

Response of Asterias vulgaris to chemical stimuli

Groups of the sea star Asterias vulgaris approached a variety of chemical stimuli in a flow tank. Urea, lactic acid, succinic acid, and handsweat were attractive to some animals; some simple acids and amino-acid mixtures were weakly attractive; pH changes and sea-water salts were not attractive. Although the groups were similarly maintained and responded with high internal consistency, they differed in responsiveness to different substances. These variations are not easily accounted for as contamination or effects of prior animal experience. A. vulgaris detects many simple organic substances, but the behavioral response is not constant and species-wide; it depends on nonchemical factors. This sensory/behavioral mode presumably suits a nonspecialist predator, and may be widespread among similar asteroids. Individuals attracted to lactic acid could be induced to cling to and evert their stomachs over a sponge soaked in lactic-acid solutions. The chemicals found atrractive in this study do not account for most of the attractiveness of shellfish tissue extracts studied previously.Contribution No. 2854 of the Woods Hole Oceanographic Institution.     

Mating in the red-sided garter snake,Thamnophis sirtalis parietalis: differential effects on male and female sexual behavior

Summary Female red-sided garter snakes, Thamnophis sirtalis parietalis, become unattractive to most males after mating in the field and in the laboratory. Male red-sided garter snakes vary in their latencies to court attractive females following copulation, with courtship resuming in minutes to hours. Unsuccessful males in mating balls disperse from mating pairs, but are not residually inhibited from courting attractive females. These patterns of behavior indicate that males have evolved mechanisms to maximize opportunities for copulation with several females, while females mate only once per season.     

有机磷农药对蛋白核小球藻的毒性相互作用研究

水体中农药复合污染产生的毒性效应具有潜在风险。为系统考察有机磷农药(OPs)混合物对淡水生态系统中绿藻的联合毒性效应,以马拉硫磷(MIT)、敌敌畏(DDVP)、敌百虫(TRC)、乐果(DIT)和氧乐果(OMT)等5种OPs作为混合物组分,运用直接均分射线法设计9组二元混合物体系共45条混合物射线。利用96孔微板测定5种OPs及其二元混合物对蛋白核小球藻(C. pyrenoidosa)的生长抑制毒性,通过基于置信区间的组合指数法分析混合物的联合毒性及毒性相互作用。结果表明,以p EC50为毒性指标,5种OPs对C. pyrenoidosa的毒性大小顺序为:TRCMITDDVPOMTDIT,OPs对C. pyrenoidosa的毒性大小受其中心磷原子的电正性影响;因混合组分的不同,部分OPs混合物对C. pyrenoidosa的联合毒性依赖于组分浓度比; OPs混合物对C. pyrenoidosa的毒性相互作用以加和为主,部分发生拮抗作用,发生拮抗作用的混合体系具有低效应区域呈加和作用,高效应区域呈拮抗作用的规律;与MIT混合的体系均有发生拮抗作用,且依赖于MIT浓度,MIT浓度比例越高,拮抗作用越强,OPs混合物的毒性相互作用与组分浓度比相关; OPs混合物的毒性相互作用组分浓度比依赖性与其联合毒性的组分浓度比依赖性规律不相关。     

Cuticular volatiles,attractivity of worker larvae and invasion of brood cells by Varroa mites. A comparison of Africanized and European honey bees

Summary. Africanized honey bees (AHBs) of Brazil and Mexico have proven to be tolerant to Varroa destructor mites. In contrast, European honey bees (EHBs: Apis mellifera carnica) at the same tropical study site are highly intolerant to these ectoparasites. A lower attractiveness of Varroa-tolerant AHB larvae has been hypothesised to be an important trait in reducing the susceptibitlity of AHBs to these mites. Thus, selection for EHB brood that is less attractive to mites is thought to be one possibility for limiting mite population growth and thus increase the tolerance of EHBs to the mite.?In Ribeir?o Preto, Brazil, European A. m. carnica bees and AHBs were tested with respect to their rate of brood infestation and brood attractiveness to Varroa mites. For the comparison of brood infestation rates, we introduced combs with pieces of EHB and AHB brood into honey bee colonies (18 repetitions). The relative infestation rate of EHB brood was significantly higher compared to AHB brood.?The preference behaviour of single Varroa mites was tested in a laboratory bioassay where either living host stages were offered or host extracts were presented on dummies. By these tests we could confirm the preference of Varroa females for certain developmental host stages and for their corresponding extracts. In contrast to the within-colony results, Varroa mites in the laboratory bioassay showed a slight preference for AHB compared to EHB larvae.?The gas chromatographic analysis revealed differences in the chemical spectrum of extracts obtained from different larvae. In accord with the results of the bioassays, we could detect stage-specific odour differences in larval cuticular compounds, including methyl esters and hydrocarbons that have been described as kairomones. None of these substances, however, revealed significant race-specific differences. Therefore, the quantity and composition of certain cuticular compounds seem to be responsible only for the recognition of a suitable host stage by Varroa females. The different infestation rates in the colonies, however, seem to be caused neither by race-specific differences in attractiveness of bee larvae nor by an extended attractive period of EHB larvae: both AHB and EHB larvae become attractive approximately 21 h before capping of the brood cell, and thus have the same window of time when they can be parasitised.?Therefore differential Varroa-infestation rates are not related to larval attraction but probably are determined by other race-specific and colony-related factors. Received 11 June 2001; accepted 19 November 2001.     

A fungus infecting domestic flies manipulates sexual behaviour of its host

The Enomopathogenic fungus Entomophthora muscae infects and kills domestic flies Musca domestica, and rates of transmission are higher at high population densities. I tested whether features of parasitized hosts increased the likelihood of transmission of the fungus in a series of experiments. Individual flies contacting an infected conspecific had a much higher probability of becoming infected than had flies contacting a dead fly without signs of infection. Males especially were attracted to and behaved sexually towards dead, infected flies even when a choice was available between a dead, infected fly and a dead, uninfected individual. The abdomen of infected flies swelled considerably as a consequence of infection, and uninfected female flies with larger abdomens are more fecund than the average female and maybe more attractive. Experiments in which abdomens of infected and uninfected flies were cut off and glued back on other flies demonstrated that the abdomens of infected flies were highly attractive to males. When the size of the abdomen of infected and uninfected individual flies was held constant, infected dead flies were still more attractive than corpses of uninfected flies. This result suggests that features other than the size of the swollen abdomen enhanced the attractivity of infected flies. Features of infected hosts therefore increased the likelihood of transmission of the parasitic fungus.