Sequestration of pyrrolizidine alkaloids by larvae ofTellervo zoilus (Lepidoptera: Ithomiinae) and their role in the chemical protection of adults against the spiderNephila maculata (Araneidae)

Summary Life stages of the primitive Australian ithomiine butterflyTellervo zoilus and its larval hostplant, the apocynaceous vineParsonsia straminea, were quantitatively assayed for pyrrolizidine alkaloids (PAs). PAs were found in all stages, mainly as N-oxides, being most concentrated in larvae and freshly-emerged adults. Although adults feed at various confirmed PA sources this probably does not compensate for losses, as wild-caught adults had considerably lower concentrations of PAs. The main alkaloid present in both freshly-emerged adults and in leaves of the host-plant was lycopsamine (1b), stored by butterflies in the N-oxide form. Its presence in higher proportion, in relation to intermedine (1a), in larvae, pupae and adults ofTellervo in relation to the host-plants suggests the inversion of intermedine to lycopsamine by the insects. No 14-member ring macrocyclic PAs were detected in either food-plant or butterflies. Several other PAs were found in wild-caught adults reflecting visits to other PA sources. PAs were also found in high concentrations in freshly-emerged individuals of the danaineEuploea core bred onParsonsia straminea. Wild-caughtDanaus affinis had high PA levels acquired from adult feeding. Freshly emergedEuploea raised onIschnocarpus frutescens andDanaus raised onIschnostemma carnosum (both PA-free) were preyed on by the orb weaving spiderNephila maculata, and showed no PAs. In all cases where PAs were present, most butterflies were liberated, usually cut out of the web unharmed, byNephila. The spider's response was not closely linked to PA concentration, however, and may also depend on hunger levels and previous experience with PA-containing butterflies. All control and other non-PA containing butterflies were consumed although rejection of some body parts of freshly-emergedDanaus affinis suggests that compounds other than PAs may be involved.     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Detoxification of pyrrolizidine alkaloids by the harvestman <Emphasis Type="Italic">Mitopus morio</Emphasis> (Phalangidae) a predator of alkaloid defended leaf beetles

Summary. The harvestman Mitopus morio (Phalangidae) is a generalist predator. It is known to prey on larvae of the chrysomelid leaf beetle Oreina cacaliae defended by plant acquired pyrrolizidine alkaloids (PAs). Tracer feeding experiments were performed to determine how harvestmen tolerate protoxic PAs. Minced meat containing either [¹⁴C]senecionine or [¹⁴C]senecionine N-oxide was fed to M. morio and subsequently feces and bodies were analyzed. Labeled alkaloid N-oxide remained stable and was eliminated almost unaltered with the feces; only 10% was recovered as tertiary PA. In contrast, approximately 80% of labeled tertiary alkaloid (senecionine) ingested with the diet was N-oxidized and eliminated; the remaining 20% consisted of unchanged senecionine and a polar metabolite of unknown structure. Harvestmen process their diet by excreting digestive juice, indicated by bleaching of the meat color. Analysis of the processed diet revealed some N-oxidation of [¹⁴C]senecionine, suggesting the gut as the site of Noxidation. Analysis of the bodies of harvestmen 80 hours after the tracer feeding pulse revealed only trace amounts of the polar metabolite. Neither senecionine nor its N-oxide could be detected in the body extracts. The results are discussed in relation to the strategies of PA adapted insects to avoid accumulation of tertiary PAs in living tissues.     

Variability in host plant chemistry: behavioural responses and life-history parameters of the Colorado potato beetle (Leptinotarsa decemlineata)

Summary. Many studies investigating effects of plant chemicals on herbivore performance have reported contradictory results, perhaps because of possible interaction between different chemicals. Also, a herbivore’s performance is not necessarily consistent with its food or oviposition preference. Our aim was to investigate simultaneously antibiosis (larval growth and survival) and antixenosis (oviposition and feeding preferences) responses in herbivore to three plant chemicals, of which one is expected to have positive and two are expected to have negative effects. Antibiosis was measured by correlating the nitrogen and glycoalkaloid levels in host plants to the survival and adult size of Leptinotarsa decemlineata, by rearing larvae on whole plants of three potato varieties. Although host plants differed in their glycoalkaloid levels, survival rate and adult body size did not differ among beetles reared on different potato varieties. This suggests that beetles are quite robust for differences in both foliar α-chaconine and foliar α-solanine content. However, differences in antixenosis were found although they could not be directly predicted from the leaf chemistry. Females preferred to lay their eggs on the variety with high α-solanine content (Nevsky) towards which males showed a tendency to feeding preference. Overall, our results confirm that beetles are well adapted to the chemical defences of potato plants as potato varieties did not significantly affect beetle performance, but differences in oviposition preference may still result in major differences in the amount of damage inflicted on plants in the fields.     

Offensive and defensive sperm competition roles in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

Sperm competition is predicted to generate opposing selection pressures on males. On one hand, selection should favour ‘defensive’ adaptations that protect a male’s ejaculate from displacement, while, on the other hand, selection should favour ‘offensive’ adaptations that overcome paternity assurance mechanisms of rivals. Here, we use the sterile male technique to assess sperm precedence when a male dung beetle Onthophagus taurus mates in both a defensive (first male) and an offensive (second male) role. Significant variation in a male’s sperm precedence (both P ₁ and P ₂) was detected, and an individual’s defensive (P ₁) and offensive (P ₂) abilities were positively correlated. Thus, it appears that sexual selection simultaneously selects for ‘defensive’ and ‘offensive’ adaptations in O. taurus. We discuss a variety of male traits in O. taurus that potentially contribute to a male’s ability to be successful when mating in an ‘offensive’ and a ‘defensive’ role.     

Determinants of vigilance behavior in the ring-tailed coati (Nasua nasua): the importance of within-group spatial position

Individuals living in social groups are predicted to live under unequal predation risk due to their spatial location within the group. Previous work has indicated that individuals located at the edge of groups have higher “domains of danger”, thus are more likely to engage in vigilance or antipredator behavior. We studied the determinants of vigilance behavior in two groups of ring-tailed coatis in Iguazu National Park, Argentina. In addition to the expected pattern that coatis were more vigilant at the edge of the group, we found that individuals were particularly vigilant at the front edge of the group. This pattern conforms to predictions of differing predation risk caused by sit-and-wait predators with respect to mobile animal groups. In addition, coatis exhibited less vigilance when the number of neighbors within 5 m and group size increased. Of the three spatial variables tested, within-group spatial position was the most important predictor variable determining vigilance levels. These results confirm that spatial position has major effects on vigilance behavior, and that group directionality is an important factor which should be taken into account when measuring vigilance behavior. Coatis were more vigilant when juveniles less than 6 months old were in the groups. The presence of these young juveniles also affected the relationship between alarm response and vigilance levels. Coatis were more vigilant after strong alarm reactions, but only when young juveniles were not present in the groups. This may indicate that coatis give differential responses to alarm calls depending on the age of the caller. A comparison of antipredator vigilance between coatis and sympatric capuchin monkeys is consistent with the hypothesis that terrestriality leads to higher perceive predation risk for coatis.     

Quantitative composition of the defensive secretion ofBledius species (Coleoptera: Staphylinidae: Oxytelinae) is adapted to naturally occurring predators

Summary The adaptation of defensive secretions to their target organisms was examined for the abdominal gland secretions ofBledius furcatus, B. spectabilis andB. arenarius. Therefore the target organisms of the secretion of theseBledius species (i.e. their predators) had to be identified. At the collection sites examined these were the earwigLabidura riparia, the antCataglyphis bicolor, the flyLispe candicans, different carabids of the generaPogonus, Dichirotrichus, Dyschirius, Bembidion andCalathus and the wading birdsHaematopus ostralegus andCalidris alba. The secretion of the abdominal glands contains the toxin ptoluquinone dissolved in either-dodecalactone and 1-undecene (B. furcatus andB. spectabilis) or in octanoic acid and octyloctanoate (B. arenarius). The ratio of these solvents is species-specific. Application experiments using some of the natural insect predators (L. riparia, C. bicolor, Pogonus, Di. gustavii, Dyschirius) revealed that these solvent ratios provided a more effective deterrent than other possible ratios. Thus by combining the solvents in certain ratios, the capability of cuticular penetration and therefore the effectiveness of the defensive secretions are adapted to their natural targets.     

Faecal marking behaviour in ringtails (Bassariscus astutus) during the non-breeding period: spatial characteristics of latrines and single faeces

Summary. Scent signals are the main source of information transmission in carnivores, being particularly important for those with nocturnal habits like the ringtail (Bassariscus astutus), a Procyonid widely distributed in Mexico. However, faecal marking behaviour of free-ranging ringtails has not been described previously. The aims of this study were to describe the use of latrines in ringtails and to test if single faeces and latrines have a marking function, based on the spatial characterisation of the defecation places and revisits to these sites. The study was conducted at an urban reserve within Mexico City in 2003, where 80 defecation points were analysed. The results showed that ringtails deposit faeces repeatedly in the same sites (latrines), which contain a variable number of faeces (between 2 and 19). The spatial distribution of latrines was not random, but a selection of substrates and zones that enhanced the effectiveness as faecal marks was observed. Latrines were preferentially placed on objects above ground level and at road margins. Latrines and single faeces that were in inconspicuous zones were generally deposited on conspicuous substrates and latrines and single faeces that were in conspicuous zones were deposited more frequently on inconspicuous substrates.     

Survival of first instar larvae ofDanaus plexippus (Lepidoptera: Danainae) in relation to cardiac glycoside and latex content ofAsclepias humistrata (Asclepiadaceae)

Summary Our paper addresses field survivorship of first instar monarch butterfly larvae (Danaus plexippus L., Lep.: Danainae) in relation to the dual cardenolide and latex chemical defenses of the sand hill milkweed plant,Asclepias humistrata (Asclepiadaceae) growing naturally in north central Florida. Survival of first instar larvae in the field was 11.5% in the first experiment (15–20 April 1990), and dropped to 3.4% in the second experiment (20–30 April). About 30% of the larvae were found glued to the leaf surface by the milkweed latex. Predator exclusion of non-flying inverte-brates by applying tanglefoot to the plant stems suggested that the balance of the mortality was due to volant inverte-brates, or to falling and/or moving off the plants. Regression analyses to isolate some of the other variables affecting survivorship indicated that first instar mortality was correlated with (1) increasing cardiac glycoside concentration of the leaves, (2) increasing age of the plants, and (3) the temporal increase in concentration of cardiac glycosides in the leaves. The study also provided confirmatory data of previous studies that wild monarch females tend to oviposit onA. humistrata plants containing intermediate concentrations of cardiac glycosides. Cardiac glycoside concentration in the leaves was not correlated with that in the latex. The concentration of cardenolide in the latex is extremely high, constituting an average of 1.2 and 9.5% of the mass of the wet and dry latex, respectively. The data suggest that an increase in water content of the latex is compensated for by an influx of cardenolide with the result that the cardenolide concentration remains constant in the latex systems of plants that are growing naturally. We also observed first instar larvae taking their first bite of milkweed leaves in the field. In addition to confirming other workers findings that monarch larvae possess elaborate sabotaging behaviour of the milkweed's latex system, we discovered that several larvae on their first bite involuntarily imbided a small globule of latex and instantly became cataleptic. This catalepsis, lasting up to 10 min, may have been in response to the high concentration of cardenolide present in the latex ofA. humistrata, more than 10 times that in the leaves. The results of the present study suggest that more attention should be directed to plant chemical defenses upon initial attack by first instar insect larvae, rather than attempting correlations of plant chemistry with older larvae that have already passed the early instar gauntlet. The first bite of neonate insects may be the most critical moment for coping with the chemical defenses of many plants and may play a much more important role in the evolution of insect herbivory than has previously been recognized.     

Sperm competition and evidence of sperm fertilization patterns in the carrion ball-roller beetle Canthon cyanellus cyanellus LeConte (Scarabaeidae: Scarabaeinae)

Field observation and laboratory experiments were conducted to investigate sperm competition and patterns of sperm fertilization under different experimental conditions in the carrion ball-roller beetle Canthon cyanellus cyanellus. Males in nature can mate with females whose spermathecae contains fertile sperm from other males. Sperm precedence was investigated using a visible genetic marker. The progeny of red (homozygous recessive) virgin females mated once with a red male and afterward, once with a green beetle (homozygous dominant) and vice versa, revealed that regardless of its color, the last male to mate fertilized c.a. 50% of the eggs, suggesting strong sperm competition. Males were able to achieve higher levels of paternity (more than 80%) when mated ad libitum with previously mated females, although large amount of variance in paternity does not exclude the possibility of first male sperm precedence or female cryptic choice. These results suggest that repeated mating and sperm replacement are the mechanisms by which last males achieve sperm precedence.     

Profiles of cuticular hydrocarbons mediate male mate choice and sexual isolation between hybridising <Emphasis Type="Italic">Chrysochus</Emphasis> (Coleoptera: Chrysomelidae)

Summary. Chemical signals frequently underlie sexual isolation between insect species. Our understanding of the evolutionary forces influencing these signaling systems is known for very few systems, challenging both our efforts to understand insect speciation, and our ability to predict long-term changes in the chemical communication systems of insects. Thus, we are in need of more systems in which both the chemical signals causing sexual isolation and the evolutionary forces driving sexual isolation are understood. Sexual isolation in the hybrid zone between Chrysochus cobaltinus and C. auratus has apparently increased in response to natural selection against hybridisation (i.e. reinforcement). Previous experiments suggested that this isolation was due, at least in part, to male preferences for conspecific females. Here, we confirm this role of male choice, and document that male mate choice in this system is influenced by cuticular hydrocarbon (CHC) profiles. Specifically, male C. cobaltinus responses to control cadavers and conspecific female cadavers painted with different cuticular hexane extracts, together with analyses of the composition of those extracts, revealed that male mate choice is governed by CHC profiles. Multivariate analyses of GC profiles demonstrated that those profiles are indeed both sex- and species-specific. Although GC-MS enabled identification and quantification of the specific cuticular hydrocarbons, we have not yet determined which individual compounds govern mate choice. Having established that CHCs influence sexual isolation in this system, we can now assess the evolutionary lability of these cues, which will inform both our understanding of speciation, and of the conditions under which the chemical signaling systems that influence mate choice in insects can evolve.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

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.     

Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Summary. Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450 s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores.     

Eggshell spotting in brood parasitic shiny cowbirds (<Emphasis Type="Italic">Molothrus bonariensis</Emphasis>) is not linked to the female sex chromosome

For avian brood parasites in which individual females are host-specialists, the arms race between hosts and parasites has favored egg color polymorphism in the parasite, with female lineages laying mimetic eggs that resemble those of the host species they parasitize. Female sex-linked inheritance of egg color fosters evolutionary stability of egg polymorphism if female lineages show both consistent eggshell color and host use. This co-evolutionary relationship is unlikely to occur if individual brood parasites use different hosts or if egg color is not maternally inherited. The shiny cowbird (Molothrus bonariensis) is an extreme generalist brood parasite that shows a very high degree of egg polymorphism. We tested whether egg spotting in this species has female sex-linked inheritance. If genetic factors controlling the expression of egg spotting were present on the female-specific W chromosome, we expected co-segregation between spotting patterns and mtDNA haplotypes, as both W and mtDNA are maternally inherited. In contrast to the known maternal inheritance of spotting patterns in great tits, we found no associations between eggshell spotting and mtDNA haplotypes, which suggests that eggshell spotting is not maternally inherited in this cowbird species.     

Heritable pheromone blend variation in a local population of the butterbur borer moth Ostrinia zaguliaevi (Lepidoptera: Crambidae)

Summary. In a local population of Ostrinia zaguliaevi Mutuura & Munroe (Lepidoptera: Crambidae), extensive variation was found in the blend ratio of three sex pheromone components, (Z)-9-tetradecenyl acetate (Z9-14:OAc, 10.2-63.8%), (Z)-11-tetradecenyl acetate (Z11-14:OAc, 32.2-86.8%), and (E)-11-tetradecenyl acetate (E11-14:OAc, 2.1-11.9%). The variation was observed over a three-year period (2002-2004). Mother-daughter regression analyses have shown that although the heritability of the minor component E11-14:OAc was not significant, the heritability of the proportions of Z9-14:OAc and Z11-14:OAc were substantial (0.5–0.6). In artificial selection experiments, the mean % Z9-14:OAc in the sex pheromone changed significantly within three generations (37% in the control line, 48 and 52% for two lines selected for increase, and 23 and 30% for two lines selected for decrease). Despite these changes, the amounts of fatty acyl pheromone precursors, (Z)-9-, (Z)-11- and (E)-11-tetradecenoate, in the pheromone gland were not significantly affected by the selection. Taken together, variation in the pheromone blend of O. zaguliaevi is likely to be attributable to a few genes involved in the reduction or acetylation of fatty acyl pheromone precursors, the last two steps in pheromone biosynthesis.     

Herbicide exposure affects the chemical recognition of a non native predator in common toad tadpoles (<Emphasis Type="Italic">Bufo bufo</Emphasis>)

Summary. In amphibians and fishes, evidence is increasing that chemical cues from injured conspecifics can play a role in the chemical labelling and learned recognition of unfamiliar predators. In this laboratory study, we tested the prediction that prior chemical exposure to a non-native predator feeding on conspecific tadpoles will subsequently allow tadpoles of the common toad (Bufo bufo) to recognize the chemical cues specifically released by this starved predator. Furthermore, we investigated the vulnerability of this chemically-mediated process to herbicide contamination. With these aims in view, groups of tadpoles were kept either unexposed or exposed for ten days to chemical cues from Turkish crayfish (Astacus leptodactylus) previously fed on tadpoles, both in uncontaminated water and in the presence of four sublethal concentrations of amitrole (0.01, 0.1, 1 and 10 mg.l⁻¹). We then assessed the effects of the six conditioning treatments on general activity and behavioural response to chemical cues from starved crayfish. Larval treatments did not affect the general activity of the tadpoles. By contrast, the treatments had significant effects on the behavioural response to the test solution prepared form starved crayfish. The only tadpoles to show an antipredator behavioural response to the chemical stimulation from starved crayfish belonged to the groups derived from chemical exposure to tadpole-fed crayfish in uncontaminated water and in contaminated water with the lowest concentration of amitrole (0.01 mg.l⁻¹). Conversely, this chemical stimulation produced no behavioural change in the control group or in the groups derived from exposure to tadpole-fed crayfish in contaminated water containing 0.1, 1 and 10 mg.l⁻¹ of amitrole. This study demonstrates that chemical cues released during the predator’s feeding activity can subsequently be used by common toad tadpoles in the recognition of an unfamiliar predator. In addition, our results show that the presence of sublethal amitrole concentrations can impair this recognition process. Such a pesticide effect might be especially detrimental for amphibian populations threatened by invasive 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.     

Less is more: social learning of predator recognition requires a low demonstrator to observer ratio in Arctic charr (Salvelinus alpinus)

Reintroduction programs for conservation purposes suffer from low post-release survival of captive-bred animals, especially so with fish reintroductions. High mortality is most often due to weakened antipredator skills, which could be enhanced through direct experience with predators and social learning. As yet, the relative efficiency of these learning methods has not been tested, and the significance of the factors determining how efficiently antipredator responses are transmitted socially from experienced (demonstrator) to predator naive (observer) individuals is largely unknown. Using hatchery-bred stock of the endangered Saimaa Arctic charr (Salvelinus alpinus), we show for the first time that: (1) contrary to theoretical predictions, acquired predator recognition was only socially transmitted from predator-experienced to predator-naive fish in groups with a smaller proportion of experienced than naive individuals, and that (2) socially acquired avoidance of the predator odors was as strong as in those fish conditioned with a combination of odors and predator exposure. We conclude that using social transmission procedures in training fish for reintroduction programs may provide considerable ecological, economical and ethical advantages.Communicated by J. Krause     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.