Terminal double bonds in the alkenes and acetates of defensive secretion from the thrips Suocerathrips linguis Mound & Marullo, 1994 (Thysanoptera: Phlaeothripidae)

Defensive secretions of adult males and females of the subsocial thrips Suocerathrips linguis were examined using gas chromatography–mass spectrometry (GC–MS). The samples contained six long-chain acetates and five aliphatic alkanes and alkenes. Whole-body extracts yielded six methyl esters of fatty acids as well as hexadecyl acetate. Structures of the two main components, (11Z)-icosa-11,19-dienyl acetate and octadecyl acetate, were confirmed. The third most abundant substance was identified as octadec-17-enyl acetate, which is a new natural product. In addition, (9Z)-octadec-9-enyl acetate was identified along with other acetates. Pure icosadienyl acetate was found to remain liquid down to −15°C, and is the solvent for the solid components of the secretion cocktail. A tentative biosynthetic pathway for all of the 12 acetates, alkanes and alkenes is discussed, considering especially the components with terminal double bonds that are uncommon in Thysanoptera and other insects.     

Social influence on the correlation between behaviours in young-of-the-year perch

The connection between risk-taking behaviour and exploratory behaviour in young-of-the-year perch (Perca fluviatilis) was studied in aquarium experiments to see whether individual behaviour patterns could be identified in this species and also to investigate how individual behaviour is influenced by their social environment. Risk-taking was defined as the time spent foraging in an open area vs hiding in the vegetation in the presence of a piscivore. Explorative behaviour was measured as latency to enter a passage leading to an unknown area. Groups of four fish were used for the observations, and both behaviours measured were positively correlated with the mean scores of these behaviours in the other group members. Risk-taking and explorative behaviours were correlated only when data was adjusted for the behaviour of the other group members. Individuals that spent more time in the open than their companions also tended to be faster than the others to enter the passage to the unknown area and vice versa. The results indicate that there are consistent individual differences in boldness in perch, but also that behaviour could be modified according to the behaviour of group members.     

Structure and function of Dufour gland pheromones from the crazy ant <Emphasis Type="Italic">Paratrechina longicornis</Emphasis>

Summary. Dufour glands of Paratrechina longicornis were analyzed chemically using GC-MS techniques. The glands contained twelve alkanes between C₉ and C₂₀ chain length with undecane and tridecane as the main components, six alkenes (1- and 4-undecene, 1-, 4-, and 6- tridecene, and heptadecadiene), two alkyl formates (C₁₁ and C₁₃), and saturated and polyunsaturated acids. Many of the alkanes and alkenes were behaviorally active causing a short lasting attraction of ants with different intensity. Detection of the major gland compound undecane in recruitment trails by in vivo SPME sampling provided evidence for its use as a recruitment pheromone. Both poison gland (formic acid) and Dufour gland (undecane) contents were detected on SPME fibers which had been attacked by the ants.     

Comparison of the flower scent of the sexually deceptive orchid <Emphasis Type="Italic">Ophrys iricolor</Emphasis> and the female sex pheromone of its pollinator <Emphasis Type="Italic">Andrena morio</Emphasis>

Summary. Ophrys flowers mimic the female produced sex pheromone of their pollinator species to attract males for pollination. The males try to copulate with the putative female and thereby pollinate the flower. Using electrophysiological and chemical analyses, floral volatiles released by O. iricolor as well as the female sex pheromone of its pollinator species, Andrena morio are investigated. Overall, 38 peaks comprising 41 chemical compounds, were found to release reactions in the antennae of male A. morio bees. Analyses using coupled gas chromatography-mass spectrometry revealed the presence of alkanes and alkenes with 20 to 29 carbon atoms, aldehydes (C9 to C24) and two esters. Almost all of those compounds were found in similar proportions in both, the floral extracts of O. iricolor and cuticle surface extracts of A. morio females. The pattern of biologically active volatiles described here is very similar to that used by other Ophrys species pollinated by Andrena males.     

Acceptability and digestion of diets fed to larval stages ofHomarus gammarus and the role of dietary conditioning behaviour

Whilst the role of chemoreception in juvenile crustacean feeding behaviour has received considerable attention, the chemosensory responses of larval planktonic crustacean stages have been neglected. The present study investigated the acceptability of processed, microparticulate and microencapsulated diets and the possible role of chemoattractants for larval and juvenile stages ofHomarus gammarus (Linnaeus) and juvenile stages ofPalaemon elegans (Rathke). Larval and juvenile lobster were obtained from MAFF (Ministry of Agriculture, Fisheries and Food, Conwy, UK), and juvenileP. elegans were collected from the Menai Straits (Gwynedd, UK) during the summer months of 1985/1986. During the larval stages, food capture results from chance encounter, and the role of low molecular-weight chemoattractants appears to be in the determination of the edible from the inedible. Dietary conditioning behaviour in decapod larvae, which has until now not been investigated, enables the larvae to discriminate more rapidly between particles of different digestibilities. The digestibility and assimilation efficiency displayed on artificial diets in the present study suggest that lobster larvae possess limited enzymatic capabilities, although later stages are more efficient and display longer food-retention time. Growth (0.114 mm d^–1) and survival (80.0%) of lobster larvae fed natural diets were significantly better (F=14.8;P=0.001, andF=54.0,P=0.001, respectively) than of individuals fed artificial diets. Although pre-digested artificial diets (ingredients pre-treated with pancreatin) resulted in higher survival (74 and 63%) compared to microencapsulated diets (37%), there was no significant growth difference between the artificial diets tested.     

Is the alkaloid in 2spot ladybirds (Adalia bipunctata) a defence against ant predation?

Garden black ants,Lasius niger L., in a laboratory colony, attacked three species of live ladybirds found near their nest, killing the smaller two species. A second colony was offered artificial diets containing crushed ladybirds of two species, and the ants' choice of feeding site noted. Both the diets were aversive compared to control, but that containing 7spot,Coccinella septempunctata L., was more aversive than the diet containing 2spot,Adalia bipunctata L. The implications of this lesser protection for 2spots in terms of the chemical defence of the species are discussed.     

Oviposition site guarding by male walnut flies and its possible consequences for mating success

Field studies showed that male Rhagoletis juglandis and R. boycei flies guard egg-laying punctures (and the eggs within) on host walnut (Juglans major) fruit and defend those sites from conspecific and heterospecific males. In field experiments with artificially punctured fruit,as well as field observations on unmanipulated fruit, males were consistently more likely to be sighted and stayed longer on damaged fruit than on undamaged fruit. On artificially punctured fruit, they consistently spent more time in the vicinity of a puncture than expected by chance alone. Males together on damaged fruit were more likely to engage in contests over those fruit than males together on undamaged fruit. Copulations were consistently more frequent for either species on damaged than undamaged fruit, both in observations of unmanipulated fruit and in artificial puncture experiments. Analyses which controlled for the longer male residence time on damaged fruit suggested strongly that copulations were consistently achieved at higher per capita rates on damaged than on undamaged fruit, indicating that puncture-guarding functions to increase access to females. An exception to the pattern in male mating success was noted at a site where both species used host fruit on the same trees. In this case, R. juglandis males were only slightly more common on punctured fruit than on control fruit and male success in copulation did not differ significantly between the two types of fruit. This anomalous result was apparently due to an almost absolute advantage enjoyed by R. boycei males in on-fruit contests with R. juglandis males. A likely basis for improvements in mating success associated with puncture guarding was a propensity for females to deposit eggs into existing punctures. Both in observations of unmanipulated fruit and in artificial puncture experiments, females consistently attempted oviposition more often in damaged than undamaged fruit. In artificial puncture experiments, both species at both sites deposited most clutches in damaged fruit. Mating generally took place as females initiated oviposition. The possible functions of puncture use by females as well as alternative functions of puncture guarding by males are discussed.     

Chemical characterization of contact semiochemicals for host-recognition and host-acceptance by the specialist parasitoid <Emphasis Type="Italic">Cotesia plutellae</Emphasis> (Kurdjumov)

Summary. Cotesia plutellae is a specialist parasitoid of Plutella xylostella. This specificity is potentially under the control of several factors before and after oviposition. Thereby, the stimuli that lead female parasitoids to host locations and to oviposition, might be at the basis of the specificity. We explore here the response of C. plutellae females exposed to host cuticular lipids. A total cuticular lipid extract of host caterpillars was fractionated into a hydrocarbon fraction and a non-hydrocarbon fraction. Neither fraction alone had any effect on oviposition behaviour in C. plutellae but the hydrocarbon fraction alone did seem to have a positive effect on the rate of antennal contact by the females. To induce oviposition behaviour, both fractions were necessary and reflect cooperation between at least one compound in each fraction. Identification of cuticular lipids shows that hydrocarbons were dominant (77%). Non-hydrocarbon compounds were mainly represented by 15-nonacosanone (18% of the total lipid extract). This ketone is rare in insect cuticle lipids and is thought to originate from the cabbage epicuticle where it is dominant with n-C₂₉ and 14- and 15-nonacosanol also found among the cuticular lipids of the host caterpillar.     

Experimental evidence of chemokinesis in newly hatched cod larvae (Gadus morhua L.)

The swimming behaviour of laboratory-reared newly hatched cod larvae (Gadus morhua L.) was observed in a control solution of artificial seawater and in seven solutions, each with a different concentration of arginine (10⁹ to 10^-3 M). The behaviour of 20 larvae was analysed in each of the eight solutions; the individual observation time was 1 min. Individual movements were recorded on video and analyzed using a computer-assisted program. The larvae swam in straight lines (a trajectory), rested, moved and started swimming again. For the parameters analyzed, i.e., number of movements, angle between successive trajectories and straightness index, there was no significant difference between the behaviour of the larvae in the different solutions. However, for the larvae in 10^-5, 10^-4 and 10^-3 M arginine solutions, the analyzed parameters, i.e., time active, frequency of trajectories (number of movements exceeding body length), distance swum min^-1, length of individual trajectories and trajectory velocity, were all significantly lower than for the larvae in the control solution of artificial seawater and for larvae in the solutions of 10^-9, 10^-8, 10^-7 and 10^-6 M arginine. The results show that the mean distance swum by cod larvae min^-1 was two to five times longer in artificial seawater without arginine and in the four lower concentrations of arginine than in the three higher concentrations. Scanning micrographs show that newly hatched (pre-feeding) cod larvae possess olfactory organs. It seems reasonalbe to assume that the observed changes in swimming behaviour are mediated by the olfactory sense and are important in the feeding strategy of cod larvae. We suggest that the observed behaviour increases the probability of the larvae localizing patches of prey organisms and remaining in the patch once they have found it. The results show that chemokinesis is a mechanism by which the spatial distribution of fish larvae will be correlated with their prey.     

Is boldness affected by group composition in young-of-the-year perch (Perca fluviatilis)?

In young-of-the-year perch (Perca fluviatilis), individuals within groups differed in the degree of boldness, estimated by habitat utilisation and feeding activity in visual contact with a potential predator. We looked at changes in individual behaviour in connection with change of group composition. During the first period, perch were randomly assigned to groups, and time spent in open habitat versus in vegetation and number of prey attacks were registered. The perch were then categorised into personality types (shy, bold, intermediate) according to their behaviour. During the second period, fish were observed when sorted into new groups, each containing only one personality type. Shy individuals showed the largest changes in behaviour, and increased both the time spent in the open and the number of prey attacks when placed into the new groups. Feeding activity in shy fish during the second period was affected by group composition during the first period. After regrouping, bold individuals decreased their time in the open, whereas intermediate individuals did not change behaviour. Time in the open habitat was, to some extent, influenced by the behaviour of the other members of the group, but number of prey attacks was not. The behaviour of fish of the different personality types we have defined in this study seemed to be based on innate traits, but also modified by the influence of other group members and by habituation to the environment.Communicated by J.Krause     

Identification of trail pheromone compounds from the labial glands of the stingless bee Geotrigona mombuca

Foragers of several species of stingless bees deposit pheromone spots in the vegetation to guide recruited nestmates to a rich food source. Recent studies have shown that Trigona and Scaptotrigona workers secrete these pheromones from their labial glands. An earlier report stated that species within the genus Geotrigona use citral from their mandibular glands for scent marking. Since convincing experimental proof for this conjecture is lacking, we studied the glandular origin of the trail pheromone of Geotrigona mombuca. In field bioassays, newly recruited bees were diverted by artificial scent trails that branched off from the natural scent trail deposited by their nestmates only when they were baited with extracts from the foragers’ labial glands. Compounds extracted from the mandibular glands, however, did not release trail following behavior. This demonstrates that the trail pheromone of G. mombuca is produced in the labial glands, as in Trigona and Scaptotrigona. Furthermore, in chemical analyses citral was identified exclusively in the foragers’ mandibular glands, which disproves its supposed role as a trail pheromone. The labial glands contained a series of terpene- and wax type esters, with farnesyl butanoate as major constituent. We, therefore, postulate that the trail pheromone of G. mombuca is composed of a blend of esters.     

Cleptoparasitism increases the host finding ability of a polyphagous parasitoid species,<Emphasis Type="Italic"> Rhopalicus tutela</Emphasis> (Hymenoptera: Pteromalidae)

Host finding abilities were investigated in the braconid Coeloides bostrichorum and the pteromalid Rhopalicus tutela, two ectoparasitoid wasps attacking the late instar of the bark beetle Ips typographus. Under laboratory conditions, important differences in the host-searching behaviour of these species were found, R. tutela being highly mobile as compared to C. bostrichorum. In the presence of C. bostrichorum, R. tutela behaved as a cleptoparasitoid, displacing the females from their oviposition sites and stealing the hosts previously located by the braconid. This behaviour facilitated host finding in R. tutela and could partly explain the relative success of the polyphagous R. tutela when occurring with the more specialised C. bostrichorum.Communicated by R.F.A. Moritz     

Sex, intimidation and severed limbs: the effect of simulated predator attack and limb autotomy on calling and emergence behaviour in the field cricket Gryllus bimaculatus

Crickets can autotomize a limb in order to evade predation; however, this autotomy compromises their escape speed and, therefore, their ability to avoid subsequent predation events. We recorded two measures of predator-avoidance behaviour in field crickets (Gryllus bimaculatus) exposed to varying levels of predation threat, with the most extreme level leading to autotomy of a limb. Our first measure of caution was time to emergence from cover, which was affected by perceived predation treatment, with both autotomized males and females being significantly more cautious than intact individuals. For males (but not females), the presence of a calling conspecific encouraged earlier emergence. Our second measure of caution was alteration of male calling behaviour. Autotomized males remained silent for significantly longer time after disturbance than intact males, but there was no difference in call rate once they had resumed calling, suggesting that behavioural changes were not merely a result of injury response but a controlled modification of behaviour as a result of autotomy. These data suggest that autotomy in field crickets results in altered calling behaviour, which was not significantly altered with different acoustic environments.     

Effects of kinship on territorial conflicts among groups of lions,<Emphasis Type="Italic"> Panthera leo</Emphasis>

Inclusive fitness theory predicts that cost of tolerant behaviour during competitive interactions is lower for relatives than for nonrelatives. Many studies have examined the effect of relatedness on behaviour within social groups. In contrast, kin selection acting among groups has received less attention. The genetic structure of African lion (Panthera leo) populations creates a strong possibility that kin selection among groups modifies behaviour during group conflicts. We used playback experiments and genetic data to investigate the importance of relatedness during simulated territorial disputes in lions. However, we found no effect of relatedness on territorial behaviour. Degree of relatedness did not affect the decision to approach simulated intrusions, nor did it affect the behaviour during approaches. The decision to approach was instead affected by position within the territory and consecutive playback number (a measure of habituation). For playbacks that did elicit an approach, the speed of response was not detectably affected by relatedness, but was affected by odds (the ratio of residents to intruders), number of intruders, number of bouts, presence of cubs, position within the territory, temperature and playback number. Although responses were unaffected by relatedness, it remains possible that other aspects of behaviour during natural encounters among prides are affected by kin selection.Communicated by L. Sterck     

Defence effectiveness of easy bleeding sawfly larvae towards invertebrate and avian predators

Summary. Easy bleeding is a phenomenon discovered in some tenthredinid insects which possess a particularly low mechanical resistance of the integument, leading under mechanical stress to haemolymph exudation. It has a defensive effect against ants and wasps through harmful plant compounds which are sequestered in the haemolymph. Here we describe etho-ecological and some chemical aspects of the defence of easy bleeders and specify the range of predators to which easy bleeding might be effective. Beside a high haemolymph deterrence associated with low integument resistance across sawfly species, we also detected toxicity of the haemolymph of some species to workers of the ant Myrmica rubra. The behaviour of easy bleeders is to move slowly and, once disturbed, to become motionless, thereby probably impeding the tendency of a predator to attack. This behaviour had no beneficial effect for easy bleeders when attacked by the predatory bug Podisus maculiventris. Bugs could successfully and without harm prey on sawfly larvae without evoking easy bleeding. For the easy bleeder Athalia rosae, host plants with different secondary metabolite profiles, and, consequently, changes in haemolymph chemistry only slightly affected the feeding behaviour of the bugs. To test the effectiveness of easy bleeding towards a vertebrate predator, easy bleeders were offered to birds, Sturnus vulgaris. The body colouration of the sawfly larvae was of prime importance in determining the predators response when testing birds in a group. It is likely that easy bleeding is a defence strategy directed primarily towards foraging insects with biting-chewing mandibles and that it is much less active towards predatory insects with piercing-sucking mandibles as well as birds. The involvement of chemical and/or physical cues in the strategy is discussed with respect to these types of predators.     

Complex chemical communication in the crazy ant <Emphasis Type="Italic">Paratrechina longicornis</Emphasis> Latreille (Hymenoptera: Formicidae)

Summary. The formicine ant Paratrechina longicornis is known for its extremely opportunistic foraging behaviour. Only a single trail pheromone source, the rectum, was previously described from this ant. Our detailed examination of this ant’s chemical communication system revealed the presence of at least four sources of pheromones. Rectum, poison sac, and Dufour gland contain orientation components with decreasing effectiveness and persistence (in the sequence mentioned) as well as attractants with increasing effectiveness. Furthermore, the mandibular gland contains repellents, and a releaser of defensive behaviour. This set of various signals of different strength and persistence allows an extraordinary degree of flexibility and efficiency in the collective behaviour of P. longicornis, especially food exploitation, and thus may contribute to this insect’s overall ecological success.     

Higher removal rate of eggs laid by anarchistic queens—a cost of anarchy?

Anarchy, where honeybee workers produce males in the presence of a queen, is extremely rare in natural honeybee populations, suggesting that there are colony-level costs associated with being anarchistic. Yet, no significant costs have yet been identified. A possible reason for this may be that researchers have only focused on the behaviour of anarchistic workers, which have been shown to perform worker-tasks as do wild-type workers. Possibly, therefore, costs associated with anarchy should be sought in anarchistic queens and not workers. A potential cost could be a lower survival rate of eggs laid by anarchistic queens perhaps because their egg-marking signal is not as clear as that of a wild-type queen. In this paper, we determined the removal rate of eggs laid by anarchistic queens in standard worker-policing bioassays. Our results show that eggs laid by anarchistic queens are removed at a higher rate than eggs laid by wild-type queens. This does not seem to be due to differences in hydrocarbons found on the surface of eggs, as both egg types showed the same alkanes and alkenes in similar proportions. We postulate that higher removal rates of queen-laid eggs due to recognition errors may be one reason that anarchy is rare in natural honeybee populations.     

Predator inspection behaviour in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): body size,local predation pressure and cooperation

The apparently maladaptive tendency of fish to approach and inspect potential predators has been explained in terms of useful information gathering or as a signal to the predator that it has been seen. We examined this behaviour in 16 populations of wild-caught stickleback (Gasterosteus aculeatus) from ponds with and without predatory perch (Perca fluviatilis). Three large and three small individuals per population were each exposed to three model predators differing in realism. A final cooperative treatment entailed pairing subjects with a second individual from the same population, but of the alternative size class, during predator presentation. As might be expected, predator inspection behaviour was much greater in the predator-sympatric populations, and only these fish increased their level of inspection as the models became incrementally more realistic. This suggests that reductions occur in the level of costly inspection behaviour in populations without predators. Subject body size had no effect on inspection effort, which suggests a limited role for experience (we assumed larger fish to be older than smaller fish), at least over the relative age differences utilized. However, small predator-sympatric fish were the only subjects to increase inspection significantly when in a cooperative context, perhaps reflecting the inherent value of a relatively larger partner in this context. These results confirm that levels of predator inspection are both population- and situation-dependent, suggesting a trade-off in the potential costs and benefits of this behaviour.Communicated by C. St. Mary     

Emission of oilseed rape volatiles after pollen beetle infestation; behavioural and electrophysiological responses in the parasitoid <Emphasis Type="Italic">Phradis morionellus</Emphasis>

Summary. The pollen beetle, Meligethes aeneus, is an important pest of oilseed rape, Brassica napus. Larvae of this species feed only in the buds and flowers of Brassicaceae. One important natural enemy of this beetle is the parasitoid Phradis morionellus that attacks larvae in buds and flowers and also feeds on the flowers. The preferences for odours of non-infested and infested rape were tested for both starved and fed parasitoids in Y-tube olfactometer experiments. The volatile blend released from pollen beetle-infested and non-infested flowering rape and from pollen beetle larvae was identified and quantified. Gas chromatography-electroantennodetection analyses were performed with female P. morionellus. Parasitoids in both treatment groups preferred infested rape, but the proportion of responding female P. morionellus was significantly lower for the group that was starved. Six of the 20 volatiles identified were released at higher rates from infested rape than from non-infested. None of these compounds was found in pollen beetle larvae headspace. P. morionellus antennae detected both major and minor components in the volatile blend. The volatiles released at a significantly higher rate from infested rape and detected by P. morionellus antennae were (Z)-3-hexenylacetate, (Z)-3-hexenol, 3-butenyl isothiocyanate and (E,E)-α-farnesene.     

Reduced nest defence intensity and improved breeding success in terns as responses to removal of non-native American mink

Lack of parental experience or differences in reproductive effort may lead to variation in nest defence behaviour among individuals in a prey population. In this experimental study, we analysed nest defence behaviour using a model of an American mink, Mustela vison, a non-native predator, at colonies of arctic terns, Sterna paradisaea, in two large areas where mink had been removed and two comparable control areas with mink in the south-western archipelago of Finland, Baltic Sea, in June 2000. Furthermore, we recorded breeding success of arctic terns in the same four areas during 1998–2001. Arctic terns took higher risks in nest defence in control areas and in a short-term (mink-free for 2 years) removal area than in the long-term (mink-free for 8 years) removal area. Thus, colonies with recent experience of mink were more active in defending their offspring. The breeding success of arctic terns was significantly higher in mink-removal areas than in control areas. We conclude that arctic terns modify their nest defence behaviour in the presence of mink. However, they cannot defend their nests sufficiently against this mainly nocturnal predator, since their breeding success is reduced in areas where mink are present.Communicated by J. Graves