Development of parasitic <Emphasis Type="Italic">Maculinea teleius</Emphasis> (Lepidoptera,Lycaenidae) larvae in laboratory nests of four <Emphasis Type="Italic">Myrmica</Emphasis> ant host species

Maculinea butterflies are social parasites of Myrmica ants. Methods to study the strength of host ant specificity in the Maculinea–Myrmica association include research on chemical and acoustic mimicry as well as experiments on ant adoption and rearing behaviour of Maculinea larvae. Here we present results of laboratory experiments on adoption, survival, development and integration of M. teleius larvae within the nests of different Myrmica host species, with the objective of quantifying the degree of specialization of this Maculinea species. In the laboratory, a total of 94 nests of four Myrmica species: M. scabrinodis, M. rubra, M. ruginodis and M. rugulosa were used. Nests of M. rubra and M. rugulosa adopted M. teleius larvae more readily and quickly than M. ruginodis colonies. No significant differences were found in the survival rates of M. teleius larvae reared by different ant species. Early larval growth of M. teleius larvae differed slightly among nests of four Myrmica host species. Larvae reared by colonies of M. rugulosa which were the heaviest at the beginning of larval development had the lowest mean larval body mass after 18 weeks compared to those reared by other Myrmica species. None of the M. teleius larvae was carried by M. scabrinodis or M. rubra workers after ant nests were destroyed, which suggests a lack of integration with host colonies. Results indicate that Myrmica species coming from the same site differ in their ability to adopt and rear M. teleius larvae but there was no obvious adaptation of this butterfly species to one of the host ant species. This may explain why, under natural conditions, all four ants can be used as hosts of this butterfly species. Slight advantages of particular Myrmica species as hosts at certain points in butterfly larval development can be explained by the ant species biology and colony structure rather than by specialization of M. teleius.     

Larval niche selection and evening exposure enhance adoption of a predacious social parasite, Maculinea arion (large blue butterfly), by Myrmica ants

Larvae of the butterfly genus Maculinea feed briefly on a foodplant before being adopted as social parasites into Myrmica ant nests. Each Maculinea species typically survives only with a single Myrmica species, yet the eggs are laid across the overlapping territories of 3-5 Myrmica species and several other ants. The ability of Maculinea arion - a 'predatory' species of Maculinea - to influence its adoption into host Myrmica colonies was studied for the first time in the field. Some earlier reports, involving captive non-host ants, suggested that larvae of the predatory Maculinea follow ant trails into host nests or wander some distance from their foodplant before being discovered and (after a long interaction) carried away by Myrmica foragers. No dispersal from foodplants occurred in wild Maculinea arion larvae. Instead, they increased by >100-fold their probability of being discovered and adopted by Myrmica spp., rather than by ants of other genera, by exposing themselves in the micro-niche occupied by Myrmica foragers at their time of day of peak foraging. Despite a complex, hour-long interaction with Myrmica workers before being carried to the nest, Maculinea arion did not enhance its adoption by host species of Myrmica. Eggs were laid without bias in Myrmica sabuleti (host) and Myrmica scabrinodis (non-host) territories; larval survival on Thymus was the same in both ants' territories; larvae waited to be found beneath their foodplant rather than seek their host; Myrmica sabuleti and Myrmica scabrinodis foraged in the same vertical and temporal niches, and had the same probability of discovering larvae; both ants behaved identically after finding larvae and took the same time to adopt them; and the ratio of wild larvae taken into Myrmica sabuleti or Myrmica scabrinodis nests was the same as the distribution between these ants of Thymus, eggs and pre-adoption larvae.     

Multiple host-ant use by the predatory social parasite <Emphasis Type="Italic">Phengaris</Emphasis> (=<Emphasis Type="Italic">Maculinea</Emphasis>) <Emphasis Type="Italic">arion</Emphasis> (Lepidoptera,Lycaenidae)

Phengaris (=Maculinea) arion is an endangered social parasite of Myrmica ants, and for a very long time was considered as specific to Myrmica sabuleti. Previous studies carried out in Poland suggested some discrepancies within this assumption, and therefore a much more intensive survey was undertaken. The host ant use of P. arion was studied at five sites in different types of biotopes in Poland, i.e. xerothermal grasslands where Thymus pulegioides was used as a larval food plant by the butterfly, and more or less sandy biotopes with Thymus serpyllum. Altogether nine Myrmica species were recorded, and considerable variation in species composition and density of nests was recorded. At four localities M. sabuleti proved to be the most common ant. A total of 529 Myrmica nests were examined, and only 20 of them contained larvae and pupae of P. arion. Host ants belonged to five different species, i.e. M. sabuleti, Myrmica scabrinodis, Myrmica schencki, Myrmica lobicornis and Myrmica hellenica. Only at one site (NE Poland) was a significant heterogeneity in parasitation rates among Myrmica species detected. M. lobicornis was the most often infested ant there, which may suggest local specialisation of the butterfly. Overall low parasitism rates may explain the vulnerability of P. arion in Central Europe but further studies are also necessary.     

Patterns of host ant use by sympatric populations of <Emphasis Type="Italic">Maculinea alcon</Emphasis> and <Emphasis Type="Italic">M. ‘rebeli’</Emphasis> in the Carpathian Basin

Maculinea butterflies show social parasitism via obligatory myrmecophily as their larvae are adopted and raised to pupation by Myrmica ants. Suitable hosts differ for different Maculinea species, and host ant specificity can further differ at the population-level. Although early studies suggested single ant species as main hosts for each Maculinea species, it has recently become clear that their host ant specificity is more complex. Maculinea alcon and Maculinea ‘rebeli’ have variously been separated according to adult and larval morphology, phenology, and their use of different ecosystems, including host plant and host ant species. However, recent genetic evidence has questioned their separation as good species. Here we compare the use of host ants by M. alcon and M. ‘rebeli’ at the regional scale in NE-Hungary and Transylvania (Romania), where molecular studies have found no species-level separation between the two forms. We opened 778 nests of Myrmica ants and searched for Maculinea specimens (larvae, pupae and exuviae) shortly before imago emergence from the nest in seven M. alcon sites, six M. ‘rebeli’- sites and one site where both M. alcon and M. ‘rebeli’ are syntopic. In all, Maculinea caterpillars were found in the nests of seven different ant species (M. alcon was recorded mainly with Myrmica scabrinodis and occasionally with M. salina and M. vandeli; M. ‘rebeli’ used mainly M. scabrinodis, M. sabuleti and M. schencki and occasionally M. lonae and M. specioides). Myrmica scabrinodis was found to be a general host of both M. alcon and M. ‘rebeli’, which is the first record for a common host ant of these two closely related butterflies within the same region. However there were also differences in host ant use patterns between the sites occupied by the two Maculinea taxa, which reflect differences in Myrmica communities between the two types of habitat. Possible explanations for the similar but not identical host use patterns of M. alcon and M. ‘rebeli’, and their relevance for the question of whether they are separate species are discussed. Received 27 November 2007; revised 28 May 2008; accepted 11 June 2008.     

Local host ant specificity of Phengaris (Maculinea) teleius butterfly,an obligatory social parasite of Myrmica ants

1. Phengaris butterflies are obligatory social parasites of Myrmica ants. Early research suggested that there is a different Myrmica host species for each of the five European Phengaris social parasites, but more recent studies have shown that this was an oversimplification. 2. The pattern of host ant specificity within a Phengaris teleius metapopulation from southern Poland is reported. A combination of studying the frequency distribution of Phengaris occurrence and morphometrics on adult butterflies were used to test whether use of different host species is reflected in larval development. 3. Phengaris teleius larvae were found to survive in colonies of four Myrmica species: M. scabrinodis, M. rubra, M. ruginodis, and M. rugulosa. Myrmica scabrinodis was the most abundant species under the host plant but the percentage of infested nests was similar to other host ant species at two sites and lower in comparison to nests of M. rubra and M. ruginodis at the other two sites. Morphometric measurements of adult butterflies reared by wild colonies of M. scabrinodis and M. ruginodis showed that wing size and number of wing spots were slightly greater for adults eclosing from nests of M. ruginodis. 4. Our results suggest that P. teleius in the populations studied is less specialised than previously suggested. The results are consistent with the hypothesis that P. teleius is expected to be the least specific of the European Phengaris species, as it has the largest and best defended fourth‐instar caterpillars and, as a predatory species, it spends less time in the central larval chambers of the host colonies. The fact that individuals reared by M. ruginodis had wider hind wings may suggest that P. teleius had better access to resources in M. ruginodis than in M. scabrinodis colonies.     

Interspecific relationships in co‐occurring populations of social parasites and their host ants

Myrmica ant colonies host numerous insect species, including the larvae of Maculinea butterflies and Microdon myrmicae hoverflies. Little is known about the interspecific relationships among these social parasites and their host ants occurring in sympatric populations. We investigated communities of social parasites to assess the strategies allowing them to share the same pool of resources (i.e. Myrmica colonies). The present study was carried out at five sites inhabited by different social parasite communities, each comprising varying proportions of Maculinea teleius, Maculinea nausithous, Maculinea alcon, and Microdon myrmicae. We investigated their spatial distributions, host segregation, the degree of chemical similarity between social parasites and hosts, and temporal overlaps in colony resource exploitation. Spatial segregation among social parasites was found in two populations and it arises from microhabitat preferences and biological interactions. Local conditions can drive selection on one social parasite to use a Myrmica host species that is not exploited by other social parasites. Myrmica scabrinodis and Myrmica rubra nests infested by larvae of two social parasite species were found and the most common co‐occurrence was between Ma. teleius and Mi. myrmicae. The successful coexistence of these two species derives from their exploitation of the host colony resources at different times of the year. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 699–709.     

A mowing experiment to evaluate the influence of management on the activity of host ants of <Emphasis Type="Italic">Maculinea</Emphasis> butterflies

Maculinea butterflies obligatory parasitize certain species of Myrmica ants. Thus, the presence of the host ant species is a limiting factor for the survival of a Maculinea population. Here, we analyse the influence of vegetation structure and ground temperature on ant diversity and abundance on Maculinea habitats, with the final aim of identifying the environmental variables determining patterns of variation in species composition in order to recommend a mowing regime that will promote our three target species: Maculinea teleius, M. nausithous and M. alcon. Experimental plots with different mowing regimes were established at eight sites in South-Eastern Germany, a region which still contains a number of relatively large, stable populations of these threatened butterfly species. Among the seven different ant species recorded, four belong to the genus Myrmica (M. scabrinodis, M. rubra, M. ruginodis and M. vandeli). Among these, M. scabrinodis results most abundant at all sites. In a CCA analysis of environmental variables recorded at the studied plots, ant species diversity appears largely determined by litter cover, mean temperature, and mean grass cover. Mowing once a year, in the second half of September, after the larvae have left their host plants, enhances the abundance of Myrmica ants in the meadows, and would be the best management compromise for all three species.     

Consequences of the arms race between <Emphasis Type="Italic">Maculinea teleius</Emphasis> social parasite and <Emphasis Type="Italic">Myrmica</Emphasis> host ants for myrmecophilous butterfly conservation

The arms race between Maculinea butterflies and Myrmica host ants leads to local host-parasite adaptations. In our study, we assessed whether sympatric and allopatric Myrmica scabrinodis populations exhibit behavioural differences towards Maculinea teleius larvae during the adoption-period when butterfly larvae need to be taken inside the Myrmica nest. The second aim was to assess the butterfly survival rate inside ant colonies from different populations. We used one sympatric host population and three allopatric populations: one infested by M. teleius and two uninfested populations. We found that ants from the sympatric population showed a higher number of positive behaviours toward M. teleius larvae during adoption than ants from the allopatric populations. There were no differences in the number of inspection or negative behaviour events. The survival of butterfly larvae was highest inside sympatric host colonies and differed from the survival of M. teleius reared by ants from the allopatric, uninfested populations. No difference was found for the survival rate of M. teleius raised by infested, allopatric host colonies compared to sympatric host populations. Our results suggest the lack of behavioural counter-adaptations of local hosts of M. teleius that more easily adopt and rear butterfly caterpillars compared to naive M. scabrinodis colonies. Our results may also have implications for Maculinea butterfly conservation, especially for reintroduction programmes. We suggest that the existence of behavioural host defences should be checked for the source host population, as well as for the Myrmica population from the reintroduction site. It may also be reasonable to introduce several Myrmica host colonies from the source butterfly host population.     

Choice behaviour of Myrmica rubra workers between ant larvae and larvae of their Phengaris (Maculinea) nausithous nest parasites

Larvae of Phengaris (Maculinea) butterflies are adopted by Myrmica workers and are obligate myrmecophiles. Brood recognition by Myrmica rubra workers was tested for concolonial larvae (M. rubra) versus allocolonial larvae (M. rubra and P. nausithous) to assay the mimetic efficiency of P. nausithous. In addition, we tested M. rubra ant colonies from different populations with and without the presence of Phengaris, to test for potential local adaptation in adoption behaviour. We show that M. rubra can distinguish between nest-mate and foreign larvae as well as between P. nausithous and their own larvae. Workers from the allopatric population inspected and rejected more P. nausithous larvae than workers from the sympatric population. This might reflect a local host adaptation in which the social parasite more efficiently mimes its sympatric host ants than allopatric ones.     

Survival and growth of parasitic <Emphasis Type="Italic">Maculinea alcon</Emphasis> caterpillars (Lepidoptera,Lycaenidae) in laboratory nests of three <Emphasis Type="Italic">Myrmica</Emphasis> ant species

The Alcon blue butterfly (Maculinea alcon) parasitizes the nests of several Myrmica ant species. In Denmark, it uses M. rubra and M. ruginodis, but never M. scabrinodis. To further examine the basis of this specificity and local co-adaptation between host and parasite, the pattern of growth and survival of newly-adopted caterpillars of M. alcon in Myrmica subcolonies was examined in the laboratory. M. alcon caterpillars were collected from three populations differing in their host use, and reared in laboratory nests of all three ant species collected from each M. alcon population. While there were differences in the pattern of growth of caterpillars from different populations during the first few months after adoption, which depended on host ant species and the site from which the ants were collected, there was no evidence of major differences in final size achieved. Survival was, however, much higher in nests of M. rubra than in nests of M. ruginodis and M. scabrinodis, even for caterpillars from a population that is never known to use M. rubra as a host in the field. The caterpillars of M. alcon thus do not show local adaptation in their pattern of growth and survival, but instead show a pattern that may reflect different nestmate recognition abilities of the host ants, related to their sociogenetic organisation. The pattern of observed host ant use in the field seems to result from a combination of differences in local host availability and locally adapted infectivity, modulated by smaller differences in survivorship in the nests of the different host ants.     

Distribution,host specificity,and the potential for cryptic speciation in hoverfly Microdon myrmicae (Diptera: Syrphidae), a social parasite of Myrmica ants

1. In 2002 Microdon myrmicae, a social parasite of Myrmica ants, was taxonomically separated from Microdon mutabilis. The original study in the U.K. found Microdon myrmicae to be specific to one ant species, Myrmica scabrinodis, yet it became apparent that the range of Microdon myrmicae includes at least the western Palaearctic. 2. Current knowledge of the European distributions of both Microdon myrmicae and Microdon mutabilis in Europe is reviewed. Also, in detailed studies of two Polish populations, Microdon myrmicae was found to survive equally well with two Myrmica ant species. We examine, however, the possibility that this reflects the presence of two separate Microdon species, each connected to one species of Myrmica. 3. Forty populations of Microdon myrmicae and 37 populations of Microdon mutabilis are currently known in Europe. All the populations in central and southern Europe that were visited after the separation of the two species were identified as Microdon myrmicae, while Microdon mutabilis’ recognised range is now restricted to the British Isles and Scandinavia. Myrmica scabrinodis was found to host Microdon myrmicae in 26 out of 31 populations investigated. Four other Myrmica species were identified to the host Microdon myrmicae: Myrmica gallienii (eight populations), Myrmica rubra (four), Myrmica vandeli (one), and Myrmica sabuleti (one). Microdon myrmicae occurs in waterlogged grassland habitats, mainly of the ‘Molinietum’ type, resulting in a patchy distribution relative to its host ants. 4. In two populations Myrmica scabrinodis and Myrmica gallienii are both abundant and rear Microdon myrmicae in equal proportions. Microdon myrmicae pupae from Myrmica gallienii nests were heavier and the anterior respiratory organs were of significantly different shape. In contrast, the comparisons of Microdon myrmicae pupae among all other populations showed no significant differences, suggesting only one species throughout the European range.     

Host specificity among Maculinea butterflies in Myrmica ant nests

Summary Ecological studies have been made of all 5 European species of Maculinea. These confirm that M. nausithous and M. rebeli live underground in Myrmica ant nests for 10 months of the year, as has long been known for the other 3 species. The main discovery was that each Maculinea species depends on a single, and different, host species of Myrmica. This specificity contradicts previous papers and scientific reviews of the relationship between Maculinea and ants. Therefore, early records are re-examined and 3 reasons are given to explain why most are misleading when applied to wild populations. Dependence on a single, rather than any, species of Myrmica explains why Maculinea populations exist in only a small minority of biotopes where their foodplants and Myrmica ants abound. It also explains the puzzling disappearance of Maculinea populations from apparently suitable sites. The discovery that M. alcon and M. rebeli depend on separate species of Myrmica that are not even closely related strengthens the argument that these butterflies are good species.     

Complexity of species conservation in managed habitats: interaction betweenMaculinea butterflies and their ant hosts

Europe's five species ofMaculinea butterfly are examples of endangered species adapted to live in traditional, cultural landscapes. All are threatened with extinction in Western Europe because of recent changes in land use. This is illustrated by an historical account of the extinction of the BritishMaculinea arion populations, despite many conservation attempts. It is shown how the failures proved to be due to ignorance of the key factor forM. arion, its specialization on a single ant host,Myrmica sabuleti. A brief account is given of research that shows how each of the five species is similarly dependent upon a separate hostMyrmica ant species and how each has an interesting and rare specific parasitoid. Steps for the practical conservation of existingMaculinea populations including the obligation, under the Bern Convention, to re-establish nationally extinct species are outlined. The procedure and problems involved in re-establishment are illustrated with reference to the successful programme forM. arion in Britain. The best way of ensuring robust populations ofMaculinea butterflies is to manage habitats to optimize the density and distribution of the required species ofMyrmica host and, secondarily, the distribution of the larval food plant. The value of single species conservation in cultural habitats is discussed. It is concluded that this is possible to achieve and that other rare organisms also often benefit, but only when conservation measures are based on the results of detailed autecological research.     

Host specificity revisited: New data on Myrmica host ants of the lycaenid butterfly Maculinea rebeli

Larvae of Maculinea rebeli, one of the most endangered European butterflies, are obligatory social parasites of Myrmica ants. At present, this relationship is thought to be highly specific, with Myrmica schencki being regarded as the primary host. Here we present data on six populations from Poland and Austria, including the first record of Myrmica specioides as a host, together with published data from other central European countries, which severely questions the inference that M. schencki is the exclusive host of M. rebeli. Our results indicate that Myrmica sabuleti is the most frequently used host ant in central Europe, whereas M. scabrinodis, M. sulcinodis, M. specioides and M. schencki are used as secondary hosts. Possible explanations for this highly variable host use include (1) regional differences in semiochemicals, behaviour or social structure of the potential Myrmica host species and (2) the existence of different ecological subspecies or cryptic species of M. rebeli. Finally, we emphasize the importance of identifying local host ant species prior to further conservation strategies in order to avoid failure of management programs or even damage to populations on the edge of extinction.     

Contest competition among Maculinea rebeli butterfly larvae in ant nests

Abstract.

• 1 Caterpillars of the myrmecophilous butterfly Maculinea rebeli showed strong evidence of contest competition when introduced at high densities to laboratory nests of Myrmica ants.
• 2 This is attributed to the direct feeding of caterpillars by workers, which select a few individuals to nurture when food or ant numbers are limiting. It contrasts with published data for a congener, Maculinea arion, which has predacious larvae and experiences scramble competition in crowded ant nests.
• 3 Worker ants from two Myrmica rubra colonies (I and II) were used to found the laboratory nests hosting Maculinea rebeli. Nests from each source reared a similar biomass of Maculinea, but whereas those containing M. rubra I workers reared eight to ten lightweight caterpillars each, cultures from colony II reared half as many caterpillars, each of about double the weight.
• 4 Differences in nest capacity may be due to the different social structures of colonies I and II at the start of the experiment.

     

Comparative study of the mandibular gland secretion of four species of Myrmica ants

The mandibular glands of the two species of ant, Myrmica ruginodis and Myrmica sabuleti contain a similar mixture of compounds, but the proportions are different. M. sabuleti produces ethanol, propanone, methylpropanal, 3-hexanone, 3-hexanol, 3-heptanone, 3-heptanol, 3-octanone, 3-octanol, 6-methyl-3-octanone, 6-methyl-3-octanol and 3-decanone. With the exception of 3-decanone these compounds were also found in M. ruginodis. These compounds were also found in M. rubra and M. scabrinodis.In both species now studied, the mandibular gland contents attract the workers and cause a large increase in their linear speed. In M. sabuleti these behavioural activities are due to 3-octanone and 3-octanol: the attraction of these two compounds in a synthetic mixture is exactly like that of an isolated mandibular gland; the compounds act in synergy to cause an increase in the ants' linear speed. Workers of M. ruginodis specifically respond to a mixture of ethanol, 3-octanone and 3-octanol: the alcohol only moderates the ethological action of the ketone, which is a true attractant and causes a very large increase in the ants' velocity; ethanol also attracts workers, acting in this respect in synergy with 3-octanone.These chemical and behavioural results are combined with those previously reported (Cammaerts-Tricot, 1973; Morganet al., 1978) to explain the responses of M. rubra, M. ruginodis, M. sabuleti and M. scabrinodis workers to isolated mandibular glands from each of these four species.     

Performance of Myrmica ant colonies is correlated with the presence of social parasites

1. The performance of ant colonies depends on different factors such as nest site, colony structure or the presence of pathogens and social parasites. Myrmica ants host various types of social parasites, including the larvae of Maculinea butterflies and Microdonmyrmicae (Schönrogge) hoverfly. How these social parasites affect host colony performance is still unexplored. 2. It was examined how the presence of Maculinea teleius Bergsträsser, Maculinea alcon (Denis & Schiffermüller), and M. myrmicae larvae, representing different feeding and growth strategies inside host colonies, is associated with worker survival, the number of foragers, and colony productivity parameters such as growth and reproduction. 3. It was found that the presence of social parasites is negatively associated with total colony production and the production of ant larvae and gynes. Male production was lower only in nests infested by M. teleius, whereas the number of worker pupae was significantly higher in all types of infested colonies than in uninfested colonies. Laboratory observations indicated that nests infested by Maculinea larvae are characterised by a higher number of foragers compared to uninfested nests but we did not find differences in worker survival among nest types. 4. The observed pattern of social parasite influence on colony productivity can be explained by the feeding strategies of parasitic larvae. The most negative effect was found for M. teleius, which feeds on the largest host brood and eliminates a high number of sexual forms. The strong, adverse influence of all studied parasite species on gyne production may result in low queen production in Myrmica populations exposed to these social parasites.     

Comparative study of the dufour gland secretions of workers of four species of Myrmica ants

The Dufour gland secretions of myrmica rubra, M. ruginodis, M. sabuleti and M. scabrinodis have been studied. The most volatile portions of the secretion of workers of all four species were found to be similar, containing C₂C₄ oxygenated compounds. The less volatile portion consists of a mixture of hydrocarbons. In M. ruginodis this is chiefly a mixture of linear saturated and mono-unsaturated hydrocarbons, similar in composition to that of M. rubra, while in M. sabuleti it consists of (Z,E)-α-farnesene and its homologues, homofarnesene, bishomofarnesene and trishomofarnesene, similar in composition to that of M. scabrinodis. Workers of each species studied were attracted to the Dufour gland volatiles of all four species, these substances chiefly causing an increase in running speed, with the workers not distinguishing between conspecific and allospecific secretions, though small quantitative differences could be demonstrated between the speed and orientation reaction of workers of each species. The less volatile fraction of the Dufour gland secretion is used for territorial marking by foraging workers. This marking is specific for each species except between M. rubra and M. ruginodis.     

Variation in Butterfly Larval Acoustics as a Strategy to Infiltrate and Exploit Host Ant Colony Resources

About 10,000 arthropods live as ants'' social parasites and have evolved a number of mechanisms allowing them to penetrate and survive inside the ant nests. Many of them can intercept and manipulate their host communication systems. This is particularly important for butterflies of the genus Maculinea, which spend the majority of their lifecycle inside Myrmica ant nests. Once in the colony, caterpillars of Maculinea “predatory species” directly feed on the ant larvae, while those of “cuckoo species” are fed primarily by attendance workers, by trophallaxis. It has been shown that Maculinea cuckoo larvae are able to reach a higher social status within the colony''s hierarchy by mimicking the acoustic signals of their host queen ants. In this research we tested if, when and how myrmecophilous butterflies may change sound emissions depending on their integration level and on stages of their life cycle. We studied how a Maculinea predatory species (M. teleius) can acoustically interact with their host ants and highlighted differences with respect to a cuckoo species (M. alcon). We recorded sounds emitted by Maculinea larvae as well as by their Myrmica hosts, and performed playback experiments to assess the parasites'' capacity to interfere with the host acoustic communication system. We found that, although varying between and within butterfly species, the larval acoustic emissions are more similar to queens'' than to workers'' stridulations. Nevertheless playback experiments showed that ant workers responded most strongly to the sounds emitted by the integrated (i.e. post-adoption) larvae of the cuckoo species, as well as by those of predatory species recorded before any contact with the host ants (i.e. in pre-adoption), thereby revealing the role of acoustic signals both in parasite integration and in adoption rituals. We discuss our findings in the broader context of parasite adaptations, comparing effects of acoustical and chemical mimicry.     

Tetracosane on the cuticle of the parasitic butterfly Phengaris (Maculinea) nausithous triggers the first contact in the adoption process by Myrmica rubra foragers

Phengaris (Maculinea) butterflies are social parasites of Myrmica ant colonies. Larvae of the parasite are adopted by the ant workers into the colonies. Apparently, chemical signals are used by Phengaris nausithous Bergsträsser larvae to mimic those of the host brood to be recognized by the ants. In the present study, chemical extracts of ant brood and butterfly larvae using four different solvents are tested in behavioural choice assays in search of compounds involved in the adoption process. Tetracosane is the main shared compound in all brood extracts of Myrmica rubra L. and in all larvae of P. nausithous. The attractiveness of tetracosane for M. rubra workers is confirmed by testing synthetic tetracosane in behavioural choice assays, suggesting that the adoption ritual may be initiated by tetracosane.