Travel duration,energetic expenditure,and patch exploitation in the parasitic wasp <Emphasis Type="Italic">Venturia canescens</Emphasis>

Foraging animals usually keep track of how costly it is to reach new resource patches and adjust patch residence time and exploitation rate accordingly. There are at least two potential factors, which are not necessarily closely linked, that animals could measure to estimate costs of traveling: the time the forager needs to reach the next patch and the amount of energy it has to invest until arrival. In the parasitoid wasp Venturia canescens, females forage for hosts from which their offspring can develop. Two different types of this parasitoid exist. The thelytokous type lives in anthropogenic habitats where flight is not necessarily linked with foraging. The arrhenotokous type lives under field conditions and shows frequent flight activity. We tested whether the wasps would use time or energy needed to assess patch availability, by either confining them into vials or letting them travel actively in a flight mill between patch visits. Our results show that in thelytokous lines, time is a sufficient cue influencing patch exploitation and an additional effect of the energy needed was not visible. In the arrhenotokous wasps, however, only the number of rounds flown in the mill influenced subsequent behavior, while mere time spent traveling did not. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Multiple feeding stimulants in <Emphasis Type="Italic">Sinapis alba</Emphasis> for the oligophagous leaf beetle <Emphasis Type="Italic">Phaedon cochleariae</Emphasis>

Summary. In earlier investigations on host plant discrimination of leaf beetles glucosinolates were described as feeding stimulants for the Brassicaceae specialist Phaedon cochleariae F. (Coleoptera: Chrysomelidae). However, since these findings could not be confirmed in later studies offering 2-propenylglucosinolate in concentrations corresponding to those detected in host plant leaf material, the identification of feeding stimulants of this leaf beetle species remained unclear. In order to investigate which compounds of the host plant Sinapis alba (Brassicaceae) are involved in feeding stimulation, leaf extracts of different polarities were tested in bioassays with adults of P. cochleariae. Number of feeding beetles and net consumption rates were highest on pea leaves painted with methanol extracts of S. alba, whereas weak feeding responses were also detectable for hexane extracts. In subsequent bioassay-guided fractionations of methanol extracts with semi-preparative high performance liquid chromatography, two distinct fractions, one containing glucosinolates and another containing flavonoids, were found to stimulate beetles to feed to variable degrees. Other collected fractions had zero activity. The combination of both active fractions evoked significantly higher consumption rates and stimulated more beetles to feed than fractions tested individually. At least one compound of each fraction, among these the main glucosinolate of S. alba, 4-hydroxybenzylglucosinolate, act additively. Effects of two different naturally-occurring ratios of glucosinolates and flavonoids on the strength of feeding responses were investigated by use of extracts of two sets of host plants differently exposed to radiation. One set was outdoors-exposed, whereas the second set was kept in the greenhouse. However, the feeding behaviour of P. cochleariae was not affected by the significantly different relative compositions of both compound classes in the host material. In conclusion, mustard leaf beetles need a combination of distinct plant metabolites acting in concert for feeding stimulation, whereby the mere presence of these stimulants, but probably not the ratio of involved compounds, determines their feeding response.     

Differential feeding of worker larvae affects caste characters in the Cape honeybee,<Emphasis Type="Italic"> Apis mellifera capensis</Emphasis>

Sections of brood from colonies of the Cape honeybee ( Apis mellifera capensis), the African honeybee ( A. m. scutellata), and hybrid bees of the two races were exchanged between colonies to study the effect of different brood-origin/nurse-bee combinations on development of caste characters. When Cape larvae were raised by African workers the amount of food provided almost doubled in comparison with Cape larvae reared by their own workers. In contrast, African larvae raised by Cape workers were provided with only half the amount they received from their own workers. After the bees emerged, we found a large degree of plasticity in characters related to caste differentiation, which corresponded closely to the amount of food provided. Super-fed Cape bees had enlarged spermathecae, were heavier than normal workers and developed more rapidly, and had reduced pollen combs, all typical for a more queen-like condition. Ovariole numbers did not appear to be enhanced by additional feeding. Cape bees that behave as social parasites in African bee colonies were most queen-like in the characters studied, albeit within the range that was found for Cape bees from normal colonies, suggesting within-colony selection for characters that enhance reproduction.Communicated by R. Page     

Differential aggressiveness between fire salamander (<Emphasis Type="Italic">Salamandra infraimmaculata</Emphasis>) larvae covaries with their genetic similarity

Responding differentially to kin and non-kin is known to be adaptive in many species. One example is the inclusive fitness benefits of reducing aggression toward closer relatives. Little is known, however, about the ability of animals to assess differential degrees of genetic relatedness and to respond accordingly with differential levels of aggression. In the present study, we tested whether aggressiveness between body mass-matched pairs of fire salamander (Salamandra infraimmaculata) larvae covaried with the genetic similarity between them. We quantified aggressiveness at three levels of genetic similarity by selecting pairs within and across pools from recently genotyped populations. We also assessed aggression between pairs of siblings. Aggression and associated injuries decreased as genetic similarity increased across the groups. These findings suggest that cannibalistic salamanders can assess their degree of genetic relatedness to conspecifics and vary their behavioral responses depending on the degree of similarity between them along a genetic relatedness continuum.     

Leaf surface compounds of the potato (<Emphasis Type="Italic">Solanum tuberosum</Emphasis>) and their influence on Colorado potato beetle (<Emphasis Type="Italic">Leptinotarsa decemlineata</Emphasis>) feeding

Summary.  The sampling behavior of the Colorado potato beetle Leptinotarsa decemlineata Say (CPB) involves examination of the surface of potato leaves. It has been suggested that leaf surface compounds (volatiles and cuticular waxes) may be involved in host-plant recognition, acceptance or discrimination. Here we report on the effect of leaf surface extracts of six Polish commercial potato varieties on CPB feeding. We tested the influence of potato leaf surface extracts on CPB adult and larval feeding, then separated the extracts with HPLC, and finally tested the effect of the HPLC-separated fractions on CPB feeding. The bioassays were performed using potato leaf discs deprived of their original surface compounds. Applied to test discs at concentrations ten times higher than natural (10 leaf area equivalent), the extracts deterred CPB adults and larvae from feeding. HPLC-separated fractions composed of alkanes, sesquiterpene hydrocarbons, wax esters, benzoic acid esters, fatty acid methyl, ethyl, isopropyl and phenylethyl esters, aldehydes, ketones, methyl ketones, fatty acids, primary alcohols, β-amyrin and sterols did not affect adult CPB feeding. Similarly, alkanes, sesquiterpene hydrocarbons, wax esters, methyl ketones, sesquiterpene alcohols and secondary alcohols had no effect on larval CPB feeding. The sterol fraction (cholesterol, β-sitosterol and stigmasterol) acted as a phagostimulant to CPB larvae. We isolated a fraction demonstrating a phagodeterrent effect on CPB adults and larvae. The qualitative composition of the deterrent fraction was quite similar in all potato extracts, but there were quantitative differences between the varieties. Much further work is needed to identify the compounds that can produce the deterrent effect.     

Differential reproductive success among subfamilies in queenless honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.) colonies

With very rare exceptions, queenright worker honeybees (Apis mellifera L.) forego personal reproduction and suppress reproduction by other workers, preferring to rear the queens sons. This is in stark contrast to colonies that have lost their queen and have failed to rear a replacement. Under these conditions workers activate their ovaries and lay many eggs that develop parthenogenetically into a final brood of males (drones) before the colony perishes. Interestingly, not all workers contribute equally to this final generation of drones in queenless colonies. Some subfamilies (workers that share the same father) contribute a disproportionately greater number of offspring than other subfamilies. Here we explore some of the mechanisms behind this reproductive competition among subfamilies. We determined the relative contribution of different subfamilies present in colonies to laying workers, eggs, larvae and pupae by genotyping samples of all life stages using a total of eight microsatellite loci. Our colonies were headed by free-mated queens and comprised 8–17 subfamilies and therefore differed significantly from colonies used in an earlier study investigating the same phenomena where colonies comprised an artificially low number of subfamilies. We show that, first, subfamilies vary in the speed with which they activate their ovaries after queen-loss and, second, that the survival of eggs to the larval stage is unequal among subfamilies suggesting that some subfamilies lay eggs that are more acceptable than others. However, there is no statistically significant difference among subfamilies in the survival of larvae to pupae, indicating that ovary activation and egg survival are the critical components to reproductive competition among subfamilies of queenless honeybee workers.Communicated by R. Page     

Sex pheromone communication of tentiform leaf-miners <Emphasis Type="Italic">Phyllonorycter insignitella</Emphasis> and <Emphasis Type="Italic">Ph. nigrescentella</Emphasis> from two related species groups

Summary.  Females of both species start their pheromone-releasing activity on the second day after emergence at the beginning of the photophase. During the present work, a peak of calling activity with close to 100% of active Ph. nigrescentella females was registered 1.5 hour after the light had been put on. The high pheromone release behaviour with 50% active females lasted for 3 hours. The calling activity of the group of females was about 6 h/day. The beginning of a photophase under laboratory conditions or an early morning in nature is a common period for sex pheromone release in the genus Phyllonorycter. (8Z,10E)-tetradecadien-1-yl acetate (8Z,10E-14:Ac), (8Z,10E)-tetradecadien-1-ol (8Z,10E-14:OH) and (8E,10Z)-tetradecadien-1-yl acetate (8E,10Z-14:Ac) in the ratio 96:4:traces as well as 8Z,10E-14:Ac and 8Z,10E-14:OH in the ratio 88:12 collected by Solid Phase Micro Extraction (SPME) were found to be specific for the calling periods of virgin Phyllonorycter insignitella and Ph. nigrescentella females respectively. Field trapping experiments demonstrated that all three compounds are important for the attraction of Ph. insignitella males while only 8Z,10E-14:Ac is the essential sex pheromone component for Ph. nigrescentella. The pheromone activity of all three compounds is reported for the first time. Addition of either 8Z,10E-14:OH or 8E,10Z-14:Ac to 8Z,10E-14:Ac did not have a significant effect on the attraction of Ph. nigrescentella males, while the efficiency of the three component blend was 5 times lower as compared to that of 8Z,10E-14:Ac. Our data demonstrate that 8Z,10E-14:OH and 8E,10Z-14:Ac play a dual function, they are minor sex pheromone components of Ph. insignitella essential for attraction of conspecific males and show an allelochemical, antagonistic effect on Ph. nigrescentella males and, thus, ensuring specificity of the mate location signal in two related Phyllonorycter species.     

Antioxidant enzyme activities of <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis> in response to nonylphenols and degradation of nonylphenols by <Emphasis Type="Italic">M. aeruginosa</Emphasis>

The aim of this study was to examine the effects of chemical nonylphenols (NPs) on the antioxidant system of Microcystis aeruginosa strains. The degradation and sorption of NPs by M. aeruginosa were also evaluated. High concentrations of NPs (1 and 2 mg/l) were found to cause increases in superoxidase dismutase (SOD) and glutathione-S-transferase (GST) activities and in glutathione (GSH) levels. These results suggest that toxic stress manifested by elevated SOD and GST levels and GSH contents may be responsible for the toxicity of NPs to M. aeruginosa and that the algal cells could improve their antioxidant and detoxification ability through the enhancement of enzymatic and nonenzymatic prevention substances. The observed elevations in GSH levels and GST activities were relatively higher than those in SOD activities, indicating that GSH and GST contributed more in eliminating toxic effects than SOD. Low concentrations of NPs (0.05–0.2 mg/l) enhanced cell growth and decreased GST activity in algal cells of M. aeruginosa, suggesting that NPs may have acted as a protecting factor, such as an antioxidant. The larger portion of the NPs (>60%) disappeared after 12 days of incubation, indicating the strong ability of M. aeruginosa to degrade the moderate persistent NP compounds. The sorption ratio of M. aeruginosa after a 12-day exposure to low nominal concentrations of NPs (0.02–0.5 mg/l) was relatively high (>30%). The fact that M. aeruginosa effectively resisted the toxic effects of NPs and strongly degraded these pollutants indicate that M. aeruginosa cells have a strong ability to adapt to variations in environmental conditions and that low and moderate concentrations of organic compounds may favor its survival. Further studies are needed to provide detailed information on the fate of persistent organic pollutants and the survival of algae and to determine the possible role of organic pollutants in the occurrence of water blooms in eutrophic lakes.     

Main nutrient compounds in food bodies of Mexican <Emphasis Type="Italic">Acacia</Emphasis> ant-plants

Summary. Myrmecophytic plants use obligate ant mutualists as a constitutive indirect defence mechanism. These plants often produce cellular food bodies (FBs) to nourish their resident ants. Lipids, proteins, and even highly specialised compounds such as glycogen have been reported from FBs, but detailed chemical analyses of FB composition have so far been presented only for Southeast Asian Macaranga and Central American Piper myrmecophytes. Here we report the chemical composition of FBs of five myrmecophytic Acacia (Fabaceae) species from Mexico using HPLC (carbohydrates and proteins) and GC-MS (lipids). Feeding experiments revealed no hints on any use of external food sources by the inhabiting Pseudomyrmex ants. These ants obviously rely completely on FBs and extrafloral nectar provided by their hosts. The total content of nutrients in Acacia FBs was 15-25% of FB dry mass, being much lower than in Macaranga or Piper FBs. Proteins were dominating (8-14 % dm) in Acacia FBs and thus were present in higher amounts than in Macaranga FBs, yet in lower amounts than in Piper. Lipids contributed 1-9 % of dry mass, showing a lower proportion than in FBs of Macaranga or Piper. Carbohydrates made up 3-11 % dm, reaching in most Acacia species the same range as observed in Macaranga and in Piper FBs. Water content was 18-24 % of FB fresh mass, and structural tissue obviously made up a much higher proportion in Acacia FBs than in Macaranga or Piper FBs. Both characters might represent an adaptation to producing FBs unprotected at the leaf tips under dry conditions. Acacia FBs contain all amino acids and all fatty acids that are considered essential for insects, and their contents of lipids and proteins are higher than in the leaves from which they are ontogenetically derived. This indicates a putatively adaptive enrichment of nutritionally valuable compounds in structures functioning as ant-food.     

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     

The trail pheromone of the ant <Emphasis Type="Italic">Crematogaster castanea</Emphasis>

Summary. The major component of the trail pheromone of the myrmicine ant Crematogaster castanea has been identified as (R)-2-dodecanol from the tibial glands of the hind legs. The substance gave activity comparable to the contents of 8 tibial glands at a concentration of 1 pg per 32 cm trail.     

Different policing rates of eggs laid by queenright and queenless anarchistic honey-bee workers (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Worker-reproduction is rare in queenright honey-bee colonies. When workers do lay eggs, their eggs are normally eaten by other workers presumably because they lack the queen's egg-marking signal. Workers use the absence of this queen signal to enforce the queen's reproductive monopoly by policing any worker-laid eggs. In contrast, in anarchistic colonies, the majority of the males arise from worker-laid eggs. Anarchistic worker-laid eggs escape policing because workers perceive anarchistic eggs as queen-laid. However, in this study, we show that eggs laid by queenless anarchistic workers do not escape policing and have very similar removal rates to worker-laid eggs from queenless wild-type (i.e. non-anarchistic) colonies. This suggests that, under queenless conditions, eggs laid by anarchistic workers lose their chemical protection and are therefore no longer perceived as queen-laid. Hence, the egg-marking signal seems to be only applied to eggs when queen and brood are present. This suggests that in the absence of queen and brood, the biosynthetic pathway that produces the egg-marking signal is switched off.Communicated by L. Keller     

Factors affecting vigilance within wild mixed-species troops of saddleback (<Emphasis Type="Italic">Saguinus fuscicollis</Emphasis>) and moustached tamarins (<Emphasis Type="Italic">S. mystax</Emphasis>)

We examine vigilance within a mixed-species troop of saddleback (Saguinus fuscicollis) and moustached (S. mystax) tamarins over a complete year. Saddleback tamarins were consistently more vigilant than moustached tamarins. This may be linked to their preference for lower strata. In accordance with previous studies of other primates, vigilant tamarins of both species were significantly further away from their nearest neighbours, and were also at lower heights in the forest than non-vigilant individuals. There was no observed sex difference in the amount of time spent vigilant. In terms of modes of scanning, the saddleback tamarins looked up significantly more frequently than the moustached tamarins, whereas there was no difference between the species in the frequency of side sweeps. There were no differences between the sexes in the frequencies of either type of vigilant behaviour. The proportion of time spent vigilant was higher than average immediately prior to entering a sleeping site for saddleback tamarins, but not for moustached tamarins. Both species were more vigilant immediately after exiting a sleeping site than at other times of the day. There was significant variation in the amount of time devoted to vigilance over the course of the year. These findings are discussed with respect to the social structure, ecology and main predator threats facing these species.Communicated by D. Watts     

Chemical ecology involved in invasion success of the cuckoo bumblebee <Emphasis Type="Italic">Psithyrus vestalis</Emphasis> and in survival of workers of its host <Emphasis Type="Italic">Bombus terrestris</Emphasis>

In bumblebees all species of the subgenus Psithyrus are social parasites in the nests of their Bombus hosts. In the bumblebee B. terrestris we investigated how colony size influences survival rates of nest entering females of the social parasite Psithyrus vestalis. Furthermore, we studied whether the host worker’s dominance status and age are reflected in its individual scent and whether Psithyrus females use volatiles to selectively kill host workers. The survival rate of Psithyrus vestalis females drops from 100%, when entering colonies with five workers, to 0% for colonies containing 50 host workers. Older host workers, born before the nest invasion, were selectively killed when Psithyrus females entered the nest. In contrast, all workers born after the nest invasion survived. The host workers’ dominance status and age are reflected by their individual odours: newly emerged workers produced a significantly lower total amount of secretions than 4-day-old workers. In chemical analyses of female groups we identified saturated and unsaturated hydrocarbons, aldehydes, and unsaturated wax-type esters of fatty acids. In a discriminant function analysis different worker groups were mainly separated by their bouquets of hydrocarbons. Killed workers release significantly more scent and of a different chemical composition, than survivors. Survivors alter scent production and increase it beyond the level of the killed workers within 1 day of the invasion. The Psithyrus female clearly maintains reproductive dominance utilizing these differences in the odour bouquets as criteria for killing workers that compete for reproduction.     

New repellent semiochemicals for three species of <Emphasis Type="Italic">Dendroctonus</Emphasis> (Coleoptera: Scolytidae)

Summary. Nine compounds identified from captured volatiles of the Douglas-fir beetle, Dendroctonus pseudotsugae, the mountain pine beetle, D. ponderosae, and the spruce beetle, D. rufipennis, that elicited antennal responses in males and females of one or more of these species were tested in the field to determine behavioural activity. 1-Octen-3-ol, found in the volatiles of females of all three species decreased the response of male and female coastal and male interior D. pseudotsugae and both sexes of D. ponderosae to their aggregation pheromones. Acetophenone, identified in the volatiles of females of all three species, significantly decreased the response of interior female D. pseudotsugae. trans-Verbenol, a potent aggregation pheromone of D. ponderosae, decreased the response of both sexes of D. pseudotsugae, while 3-methyl-2-cyclohexen-1- one (MCH), the antiaggregation pheromone of D. pseudotsugae and D. rufipennis decreased the response of both sexes of D. ponderosae. While it has been demonstrated that semiochemical mediated interspecific communication occurs among bark beetles infesting the same host, this study demonstrates that beetles can perceive signals emitted by heterospecifics attacking nonhosts and can potentially use them to avoid attacking the wrong species of conifer.     

Patterns of gamete incompatibility between the blue mussels<Emphasis Type="Italic"> Mytilus edulis</Emphasis> and<Emphasis Type="Italic"> M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis>

Previous research on gametic incompatibility in marine invertebrates suggests that for highly dispersive marine invertebrate species, barriers to fertilization among closely related taxa are often incomplete and sometimes asymmetric. The nature of these barriers can dramatically affect the patterns of gene flow and genetic differentiation between species, and thus speciation. Blue mussels, in the genus Mytilus, are genetically distinct in allopatry yet hybrids are present wherever any two species within the group co-occur. The present study sampled M. edulis (L.) and M. trossulus (Gould) in May and June 2001 from the East Bay section of Cobscook Bay, Maine, USA (latitude 44°56′30″N; longitude 67°07′50″W), where the two species are sympatric. Gamete incompatibility was investigated in a series of laboratory fertilizations carried out in July 2001. The proportion of fertilized eggs typically exceeded 80% at sperm concentrations of 10³–10⁴ ml^?1 among intraspecific matings (n=18), but was <30% even at sperm concentrations in excess of 10⁵–10⁶ ml^?1 for interspecific matings (n=13). Further analysis indicated that approximately 100- to 700-fold higher sperm concentrations were required to achieve 20% fertilization in interspecific matings relative to intraspecific matings, indicating strong barriers to interspecific fertilization. The proportion of fertilized eggs did not follow this general pattern in all matings, however. The eggs from two (out of five) M. edulis females were almost as receptive to M. trossulus sperm as they were to M. edulis sperm. In contrast, the eggs from all M. trossulus females (n=3) were unreceptive to M. edulis sperm, suggesting that fertilization barriers between these species may be asymmetric. Given the experimental design employed in this study, the results are also consistent with a strong maternal or egg effect on the level of interspecific gamete compatibility in M. edulis.     

Brood sex ratio adjustment in collared flycatchers (<Emphasis Type="Italic">Ficedula albicollis</Emphasis>): results differ between populations

Recently, a number of studies have found adaptive brood sex ratio (BSR) manipulation in birds. The reason for such manipulations is thought to be the different reproductive value of male and female nestlings. Several studies have found that parental quality and food supply can affect BSR, however results are sometimes inconsistent between species and populations. We investigated BSR patterns in a Hungarian population of Collared Flycatchers (Ficedula albicollis) and compared the results with those obtained in a previous study of the same species in Sweden. We found two significant differences. First, the male forehead patch size, a heritable, sexually selected trait, affected the brood sex ratio in the Swedish population, but not in our Hungarian study population. This difference might be a consequence of the different information content of the forehead patch size in the two populations. Second, a seasonal shift in BSR (more sons late in the season) was observed in the Hungarian, but not in the Swedish population.Communicated by J. Graves     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

A mark-recapture study of male<Emphasis Type="Italic"> Colletes cunicularius</Emphasis> bees: implications for pollination by sexual deception

An unusual pollination strategy is pollination by sexual deception in which orchids sexually attract male insects as pollinators. One gap in knowledge concerns the pattern and extent of pollinator movement among these sexually deceptive flowers and how this translates to pollen and gene flow. Our aim was to use mark and recapture techniques to investigate the behavior and movement of male Colletes cunicularius, an important bee pollinator of Ophrys. Our study site was located in northern Switzerland where a large population of the bees was nesting. Within two plots, (10×40 m), we marked bees with different colors and numbered tags. Seventeen percent of the 577 marked bees were recaptured over a period of 1 to a maximum of 11 days. However, the number of recaptures dropped dramatically after 3–5 days, suggesting an average lifetime of less than 10 days. Mark-recapture distances varied from 0 to 50 m, with a mean of 5 m. Our findings show that individual male bees patrol a specific and restricted region of the nesting area in search of mates. This mark-recapture study provides the first clues about the potential movement of pollen within populations of Ophrys orchids. We predict that orchid-pollen movements mediated by bees will be similar to the mark-recapture distances in this study. Parallel studies within orchid populations, including direct studies of pollen movement, are now required to better understand how pollinator mate-searching behavior translates to pollination success and pollen movement within sexually deceptive orchid populations.Communicated by R.F.A. Moritz     

Correlates of boldness in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)

Behavioural variation is known to occur between individuals of the same population competing for resources. Individuals also vary with respect to their boldness or shyness. An individuals position along the shy-bold axis may be defined as the extent to which it is willing to trade off potentially increased predation risks for possible gains in resources. Similarly, group living may be interpreted as a trade-off between anti-predatory tactics and foraging efficiency. The responses of three-spined sticklebacks (Gasterosteus aculeatus) were tested across four social contexts to assess relative boldness or shyness and to further examine whether their behaviour would be consistent within and between contexts. Individuals displayed consistent responses within and between the first two contexts: those individuals which resumed foraging rapidly after a simulated aerial predator attack also displayed low shoaling tendencies. Such fish were deemed to be bold, whilst those which displayed the converse behaviour, slow resumption of foraging and a high shoaling tendency, were deemed to be shy. In a third context, bold individuals out-competed shy conspecifics for food. Boldness was also positively correlated with growth over a 6-week period. The position adopted by an individual within a group is usually interpreted as a trade-off between predation risk and foraging efficiency—both are greater at the front of a mobile group. Bold individuals showed significantly stronger tendencies towards front positions than shy conspecifics. The results suggest that, contrary to some previous studies on other animals, bold or shy behaviour in sticklebacks is consistent between contexts.Communicated by T. Czeschlik