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.     

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.     

Host propagation permits extreme local adaptation in a social parasite of ants

The Red Data Book hoverfly species Microdon mutabilis is an extreme specialist that parasitises ant societies. The flies are locally adapted to a single host, Formica lemani , more intimately than was thought possible in host–parasite systems. Microdon egg survival plummeted in F. lemani colonies > 3 km away from the natal nest, from c. 96% to 0% to < 50%, depending on the hoverfly population. This is reflected in the life-time dispersal of females, measured at < 2 m, resulting in oviposition back into the same ant nests for generation after generation. To counter destabilizing effects on the host, Microdon manipulates the social dynamics of F. lemani by feeding selectively on ant eggs and small larvae, which causes surviving larvae to switch development into queens. Infested colonies rear double the number of new queens, thus propagating the vulnerable local genotype and compensating for damage to the host colonies. The consequences of such extreme host specificity for insect conservation are discussed.     

Social parasite distancing: RADseq reveals high inbreeding in the social parasite Microdon myrmicae but low philopatry for host ant nest

1. The hoverfly Microdon myrmicae is a rare and extremely localised social parasite of Myrmica ants, only occurring around wet grassland. Dispersal, location of the host, and oviposition are crucial steps in the life of ant parasites but are poorly known due to the challenge of studying such rare species.
2. Using genome‐wide loci obtained by RADseq, we investigated the genetic structure and relatedness of Mi. myrmicae larvae and its ant host, collected from three localities in South West England, and inferred biological and behavioural traits.
3. We found that: (a) Mi. myrmicae larvae show high inbreeding levels and severe heterozygosity deficiency, as an expression of the small population size that favours the mating between siblings or half‐siblings; (b) Mi. myrmicae adults can disperse for many kilometres, spreading much more than it is reported for its sibling species, Microdon mutabilis; (c) a single female lays small egg bunches in different ant colonies, sometimes spanning substantial distances (bet‐hedging strategy); in parallel, a single ant nest can harbour eggs from different Microdon females; (d) preliminary evidence suggest that contrary to Mi. mutabilis, host colony choice seems not to depend on the number of queens residing in a single colony.
4. These results overall strongly deviate from what previously found for Mi. mutabilis, whose females oviposit in the natal nest generation after generation. We argue that such different ecological traits and parasitic strategies between closely related species are mainly ascribable to the different selective pressure on the two ant hosts.

     

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.     

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.     

Getting carbohydrates in ants: damage of the young cereal sprouts by ants of the genus Myrmica Latreille, 1804 (Hymenoptera: Formicidae), to obtain plant sap

Summary

Carbohydrate food is of high importance for survival of ant colonies. Ants are known to use sugary excretions (honeydew) of various insects, nectar of floral and extrafloral nectaries, and even sap of some trees. However, the ability of ants to use sap of herbaceous plants has not been mentioned. This is the first evidence that ants of the genus Myrmica can intentionally ‘cut off’ young cereal sprouts to obtain plant sap. The investigation was carried out in a laboratory in 2018 and 2019 and involved three ant species of the genus Myrmica [12 colonies of M. rubra (Linnaeus, 1758); eight colonies of M. ruginodis Nylander, 1846; and five colonies of M. scabrinodis Nylander, 1846]. First observations were made occasionally in 10 ant colonies during the study of ant–aphid interactions. After three days of carbohydrate starvation, ants were supplied with the plants of wheat infested with aphids of Schizaphis graminum (Rondani, 1852). Within the first day in addition to ordinary trophobiotic relations with aphids, the workers of all the studied colonies demonstrated unexpected behaviour: they ‘cut off’ some sprouts and collected sap of these plants. The experimental investigation in 15 ant colonies of various sizes (about 150, 300 and 500 workers) supplied with the plants infested or non-infested with aphids has shown that getting sap of herbs depends greatly on ant colony needs and available resources. The number of damaged plants was much higher both in the larger colonies of ants and in the absence of aphids. This way of getting carbohydrates allows ants to quickly obtain some extra food needed to maintain colony viability and seems to be one of the mechanisms promoting survival of ants in conditions of acute carbohydrate deficiency. At the same time, ants avoid using plant sap when there are more available alternative carbohydrate resources.     

Miocene divergence,phenotypically cryptic lineages,and contrasting distribution patterns in common lichen‐forming fungi (Ascomycota: Parmeliaceae)

Despite the recent advancements in recognizing diversity in lichen‐forming fungi, assessing the timing of diversification remains largely unexplored in these important fungal symbionts. To better understand the evolutionary processes driving diversification in common lichen‐forming fungi, we investigated the phylogeny and biogeography of the broadly distributed Melanelixia fuliginosa/M. glabratula group, using molecular data from six nuclear markers. Phylogenetic analyses of individual gene alignments and combined data provide strong evidence for five species‐level lineages within this species complex. Three of these lineages correspond to the previously described species M. fuliginosa, M. glabratula, and M. subaurifera. The remaining two lineages, ‘M. sp. 1’ and ‘M. sp. 2’, merit species recognition based on genealogical concordance. Both M. glabratula and M. subaurifera had broad intercontinental distributions, sharing identical haplotypes among intercontinental populations. Based on the current sampling, M. fuliginosa s.s. was represented exclusively by European material and was not collected in North America. ‘M. sp. 1’ was represented by collections from Scotland and Spain; and ‘M. sp. 2’ was represented by collections in California, USA. Environmental factors driving the contrasting distribution patterns in this group remain unknown. Divergence times estimated using a coalescence‐based multilocus species‐tree approach suggest that diversification within the M. fuliginosa/M. glabratula group occurred exclusively during the Miocene. The results of the present study indicate that phenotypically cryptic lichen‐forming fungal species‐level lineages may be relatively ancient and do not necessarily reflect recent divergence events. Furthermore, diagnosable phenotypic differences may be absent even millions of years after the initial divergence. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●●, ●●–●●.     

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.     

Unexpected benefit of a social parasite for a key fitness component of its ant host

Numerous invertebrates inhabit social insect colonies, including the hoverfly genus Microdon, whose larvae typically live as brood predators. Formica lemani ant colonies apparently endure Microdon mutabilis infections over several years, despite losing a considerable fraction of young, and may even produce more gynes. We present a model for resource allocation within polygynous ant colonies, which assumes that whether an ant larva switches development into a worker or a gyne depends on the quantity of food received randomly from workers. Accordingly, Microdon predation promotes gyne development by increasing resource availability for surviving broods. Several model predictions are supported by empirical data. (i) Uninfected colonies seldom produce gynes. (ii) Infected colonies experience a short-lived peak in gyne production leading to a bimodal distribution in gyne production. (iii) Low brood : worker ratio is the critical mechanism controlling gyne production. (iv) Brood : worker ratio reduction must be substantial for increased gyne production to become noticeable.     

Extreme host specificity by Microdon mutabilis (Diptera: Syrphiae), a social parasite of ants

Larvae of the syrphid fly Microdon mutabilis are social parasites which live up to two years, feeding on ant brood in nests of Formica lemani ants. We questioned why M. mutabilis is extremely localized when its host is widespread and abundant. Like endoparasitic diseases, social parasites must penetrate formidable defences before ''infecting'' their hosts. This occurs during the egg stage of M. mutabilis: females are sedentary and oviposit at F. lemani nest entrances, which workers treat as part of their nest, leaving the thin-shelled eggs vulnerable to attack for 10 to 14 days before hatching. We describe experiments which show a strong maternal effect in M. mutabilis. New-laid eggs had > 95% survival when introduced to the individual ant colony that reared each mother fly or to its close neighbours, but survival declined as a sigmoidal logistic function of distance from the mother nest, with F. lemani colonies from 2 and 30 km away killing 80 and > 99% of eggs, respectively, within 24 h. Attacks on eggs also increased in proportion to the delay before introduction to laboratory nests. We suggest that they may be coated with a mimetic chemical disguise that lasts for three to four days after oviposition. The results indicate extreme local adaptation by an M. mutabilis population not simply to one species of host, but to an individual host population and possibly to local strains or family groups within an F. lemani population. This conclusion is discussed in terms of virulence, transmission and coevolution in parasitic diseases.     

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.     

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.     

Quarantining behaviour in ants: are Myrmica aphid milkers able to detect and get rid of fungus-contaminated aphids?

Fungal infections are highly dangerous for social insects including ants. Close trophobiotic interaction between ants and aphids promotes infection transmission, as aphids can be a disease vector. The ability of ants to detect fungus-infected aphids and get rid of them is important to the prosperity of both symbiotic partners. However, the diversity of quarantining behaviour among ants is still poorly studied. Here, the behaviour of honeydew foragers of two ant species – Myrmica rubra L. and Myrmica scabrinodis Nylander (Hymenoptera: Formicidae, Myrmicinae, Myrmicini) – was studied in laboratory towards Schizaphis graminum (Rondani) (Hemiptera: Aphididae, Aphidini) aphids contaminated with the generalist fungal pathogen Beauveria bassiana (Balsamo-Crivelli) Vuillemin (Hypocreales). This fungus attacks a wide range of hosts including aphids and ants. The removal of conidia-contaminated aphids from the host plant was found not to be typical of the ants studied. Aphid milkers of M. rubra and M. scabrinodis usually displayed non-aggressive behaviour (tolerance, antennation, honeydew collection, grooming) towards the experimental aphids regardless of whether they were covered with conidia or not. Neither ant species, nor the number of milkers had significant effects on their behaviour towards ‘infected’ aphids. However, some individuals were found to demonstrate quarantining behaviour in full. They quickly detected and removed contaminated aphids, placing them at some distance from the plant. Moreover, in addition to the simple carrying of ‘infected’ aphids down, the more effective technique of dropping them from the plant was noted as well. Ants of the genus Myrmica appear to have a tendency to perform a certain sequence of actions to remove conidia-contaminated aphids from the plant. It is likely that in larger colonies or under conditions of increased risk of infection with entomopathogenic fungi, some Myrmica ants are able to deploy and actively use the behavioural pattern of quarantining behaviour to increase their viability.     

Interspecific differences in cuticular hydrocarbon profiles of Myrmica ants are sufficiently consistent to explain host specificity by Maculinea (large blue) butterflies

The chemical signatures on the cuticles of five common Myrmica ant species were analysed (49 colonies of M. rubra, M. ruginodis, M. sabuleti, M. scabrinodis and M. schencki), each ant being the specific host of one of the five threatened European species of Maculinea butterfly. The cuticular hydrocarbon profile (based on the relative abundance of each chemical) of each ant species was highly distinctive, even between the morphologically similar species M. sabuleti and M. scabrinodis. There was no significant difference in the chemical profiles of workers and larvae from any colony. Nor was there much pattern in the intraspecific variation: colonies from the same populations were significantly, but only slightly, more similar to each other than to colonies from distant populations. M. rubra showed remarkably little variation between populations sampled widely from northern Russia, Ukraine, Scotland and southern England. The data were compared with published profiles of M. rubra and two North American Myrmica species, and with a quantitative reanalysis of data for Maculinea rebeli caterpillars. We conclude that the hydrocarbon profiles of Myrmica species are sufficiently and consistently different for chemical mimicry to explain the pattern of host specificity recorded for the European Maculinea butterflies. The optimum strategy for chemical mimicry in each of the two life-styles of Maculinea larvae is discussed: we suggest that predatory species might benefit from mimicking the median profile of their model whereas the "cuckoo" species would benefit when variation between siblings encompasses a large range of the variation recorded within a local population of the model species.     

First screening of bacterial communities of Microdon myrmicae and its ant host: do microbes facilitate the invasion of ant colonies by social parasites?

Many studies have highlighted how numerous bacteria provide their hosts essential nutrients or protection against pathogens, parasites and predators. Nevertheless, the role of symbiotic microorganisms in the interactions between social insects and their parasites is still poorly known. Microdon (Diptera, Syrphidae) is a peculiar fly genus whose larvae are able to successfully infiltrate ant colonies and feed upon the ant brood. Using high throughput 16S rRNA gene amplicon sequencing, we provide the first microbiome survey of Mi. myrmicae larvae and larvae and workers of its host, Myrmica scabrinodis, collected from two sites in England. We analyzed the microbiome of the external surface of the cuticle and the internal microbiome of the body separately. The results clearly show that the Mi. myrmicae microbiome significantly differs from that of its host, while no substantial dissimilarity was detected across the microbiome of ant workers and ant larvae. Microdon myrmicae microbiome varies across the two analyzed sites suggesting that bacteria communities of Mi. myrmicae are derived from the environment rather than by horizontal transmission between hosts and parasites. Families Streptococcaceae, Carnobacteriaceae and Rizhobiaceae are dominant in My. scabrinodis, and Spiroplasma is dominant in ant workers. Microbiome of Mi. myrmicae larvae is mainly characterized by the family Anaplasmataceae, with Wolbachia as predominant genus. Interestingly, we found Serratia within both Mi. myrmicae and Myrmica larvae. Bacteria of this genus are known to produce a family of pyrazines commonly involved in ant communication, which could play a role in Microdon/ant interaction.     

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.     

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.     

Body size variation and caste ratios in geographically distinct populations of the invasive big‐headed ant,Pheidole megacephala (Hymenoptera: Formicidae)

Body size is an important life history trait that can evolve rapidly as a result of how species interact with each other and their environment. Invasive species often encounter vastly different ecological conditions throughout their introduced range that can influence relative investment in growth, reproduction and defence among populations. In this study, we quantified variation in worker size, morphology and proportion of majors among five populations of a worldwide invasive species, the big‐headed ant, Pheidole megacephala (Fabricius). The sampled populations differed in ant community composition, allowing us to examine if P. megacephala invests differently in the size and number of majors based on the local ant fauna. We also used genetic data to determine if these populations of P. megacephala represented cryptic species or if morphological differences could be attributed to change following introduction. We found significant variation in worker mass among the populations. Both major and minor workers were largest in Australia, where the ant fauna was most diverse, and minor workers were smallest in Hawaii and Mauritius, where P. megacephala interacted with few to no other ants. We also found differences in major and minor worker morphology among populations. Majors from Mauritius had significantly larger heads (width and length) relative to whole body size than those from Hawaii and Florida. Minors had longer heads and hind tibias in South Africa compared with populations from Australia, Hawaii and Florida. The proportion of majors did not differ among populations, suggesting that these populations may not be subject to trade‐offs in investment in major size versus number. Our molecular data place all samples within the same clade, supporting that these morphologically different populations represent the same species. These results suggest that the variation in shape and morphology of major and minor workers may therefore be the result of rapid adaptation or plastic responses to local conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 423–438.     

Patterns of morphological evolution in the mandible of the house mouse Mus musculus (Rodentia: Muridae)

The worldwide distributed house mouse, Mus musculus, is subdivided into at least three lineages, Mus musculus musculus, Mus musculus domesticus, and Mus musculus castaneus. The subspecies occur parapatrically in a region considered to be the cradle of the species in Southern Asia (‘central region’), as well as in the rest of the world (‘peripheral region’). The morphological evolution of this species in a phylogeographical context is studied using a landmark‐based approach on mandible morphology of different populations of the three lineages. The morphological variation increases from central to peripheral regions at the population and subspecific levels, confirming a centrifugal sub‐speciation within this species. Furthermore, the outgroup comparison with sister species suggests that M. musculus musculus and populations of all subspecies inhabiting the Iranian plateau have retained a more ancestral mandible morphology, suggesting that this region may represent one of the relevant places of the origin of the species. Mus musculus castaneus, both from central and peripheral regions, is morphologically the most variable and divergent subspecies. Finally, the results obtained in the present study suggest that the independent evolution to commensalism in the three lineages is not accompanied by a convergence detectable on jaw morphology. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 635–647.