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The range of hosts exploited by a parasite is determined by several factors, including host availability, infectivity and exploitability. Each of these can be the target of natural selection on both host and parasite, which will determine the local outcome of interactions, and potentially lead to coevolution. However, geographical variation in host use and specificity has rarely been investigated. Maculinea (= Phengaris ) butterflies are brood parasites of Myrmica ants that are patchily distributed across the Palæarctic and have been studied extensively in Europe. Here, we review the published records of ant host use by the European Maculinea species, as well as providing new host ant records for more than 100 sites across Europe. This comprehensive survey demonstrates that while all but one of the Myrmica species found on Maculinea sites have been recorded as hosts, the most common is often disproportionately highly exploited. Host sharing and host switching are both relatively common, but there is evidence of specialization at many sites, which varies among Maculinea species. We show that most Maculinea display the features expected for coevolution to occur in a geographic mosaic, which has probably allowed these rare butterflies to persist in Europe. This article is part of the theme issue ‘The coevolutionary biology of brood parasitism: from mechanism to pattern’.

Butterflies of the genus Maculinea are highly endangered in Europe. The cuckoo species M. rebeli has been thoroughly investigated through both empirical and modelling studies, but less is known about the population ecology of predatory Maculinea. We present the findings of a 2‐year research study on sympatric populations of two endangered butterflies: Maculinea teleius and M. nausithous in the Kraków region, southern Poland. The study comprised mark–release–recapture sampling and laboratory rearing of butterflies from larvae collected in the field. For both species the sex ratio was slightly, but consistently, female‐biased and there was little year‐to‐year change in the seasonal population sizes. Daily numbers showed greater variation between the 2 years of the study due to the differences in daily survival rate. The average life span of laboratory‐raised butterflies kept in ideal conditions was more than 6 days, compared to only 2–3 days in the field. The recruitment of both males and females consistently followed a bimodal pattern. A small proportion of individuals (maximum 25%) changed sites, in spite of relatively short distances of ca. 100 m separating them. The results indicate that populations of both species are typically stable within their sites, possibly due to larval polymorphism, but there is little inter‐site mobility and thus landscape corridors seem necessary to enhance metapopulation viability. A further problem to be considered in the conservation of Maculinea butterflies is the fact that their very short life span in relation to flight‐period length reduces the effective population size.

Abstract. Butterflies of the genus Phengaris have a highly specialised life cycle involving an obligatory relationship with Myrmica ants. A knowledge of the host ant specificity is essential for understanding the relationship between a particular Phengaris species and its hosts and also important for the conservation of these butterflies. Data on host ant specificity were collected in Poland, the Czech Republic, Slovakia and Ukraine. Five different Myrmica species were used by P. teleius as hosts (M. scabrinodis, M. rubra, M. ruginodis, M. rugulosa and M. gallienii) and at most localities it was not possible to distinguish a primary host -i.e. several Myrmica species were parasitized to similar extents. Three populations of P. nausithous were found in Poland and Ukraine. In every case, M. rubra was its primary host, although in the Kraków region (Poland) two nests of M. scabrinodis and two of M. ruginodis were infested by this butterfly species. P. alcon in the four populations investigated in Poland and Ukraine invariably only used M. scabrinodis as a host despite the presence of other Myrmica species. These results obtained suggest lack of host specificity in P. teleius and high host specificity in P. nausithous, which mainly uses M. rubra as its host across Europe. Moreover, the three populations of P. alcon investigated seem to be highly specific and use M. scabrinodis as a host, which confirms the high local specialisation of these populations. 871* Present and corresponding address:

Ants are dominant members of many terrestrial ecosystems and are regarded as indicators of environmental changes. However, little is known about the effects of invasive alien plants on ant populations, particularly as regards the density, spatial distribution and size of ant colonies, as well as their foraging behaviour. We addressed these questions in a study of grassland ant communities on five grasslands invaded by alien goldenrods (Solidago sp.) and on five non-invaded grasslands without this plant. In each grassland, seven 100 m 2 plots were selected and the ant colonies counted. Ant species richness and colony density was lower in the plots on the invaded grasslands. Moreover, both of these traits were higher in the plots near the grassland edge and with a higher number of plant species in the grasslands invaded by goldenrods but not in the non-invaded ones. On average, ant colony size was lower on the invaded grasslands than the non-invaded ones. Also, ant workers travelled for longer distances to collect food items in the invaded areas than they did in the non-invaded ones, even after the experimental removal of some ant colonies in order to exclude the effect of higher colony density in the latter. Our results indicate that invasive alien goldenrods have a profound negative effect on grassland ant communities which may lead to a cascade effect on the whole grassland ecosystem through modification of the interactions among species. The invasion diminishes a major index of the fitness of ants, which is a colony's size, and probably leads to increased foraging effort of workers. This, in turn, may have important consequences for the division of labour and reproductive strategies within ant colonies.

Myrmica ants have been model species for studies in a variety of disciplines, including insect physiology, chemical communication, ant social dynamics, ant population, community ecology, and ant interactions with other organisms. Species belonging to the genus Myrmica can be found in virtually every habitat within the temperate regions of the northern hemisphere and their biology and systematics have been thoroughly studied. These ants serve as hosts to highly diverse parasitic organisms from socially parasitic butterfly caterpillars to microbes, and many Myrmica species even evolved into parasitizing species of their own genus. These parasites have various impacts both on the individuals and on the social structure of their hosts, ranging from morphological malformations to reduction in colony fitness. A comprehensive review of the parasitic organisms supported by Myrmica and the effects of these organisms on individuals and on whole ant colonies has not yet been compiled. Here, we provide a review of the interactions of these organisms with Myrmica ants by discussing host and parasite functional, behavioral or physiological adaptations. In addition, for all "symbiont groups" of Myrmica ants described in this paper, we examine the present limitations of the knowledge at present of their impact on individuals and host colony fitness. In conclusion, we argue that Myrmica ants serve as remarkable resource for the evolution of a wide variety of associated organisms.

The presence of annual and biennial individuals within the same population has been recently demonstrated in the myrmecophilous butterflies Maculinea rebeli and Maculinea alcon, which present a cuckoo strategy inside Myrmica nests, and Maculinea arion which is a predatory species. Here, we present field and laboratory data on polymorphic larval growth in two other predatory species of Maculinea: M. teleius and M. nausithous. Body mass distributions of pre-pupation larvae were bimodal in both species. These results point to the existence of larvae that develop in 1 or 2 years. We also showed that the probability of pupation depended on larval body mass. In the case of M. teleius, the critical body mass at which larvae have a 50% probability of pupation is about 80 mg. We suggest that polymorphism in Maculinea may have evolved as an adaptation to life in ant nests, a habitat which protects them from predators and provides food. However, the quality of this resource is highly variable and unpredictable. According to the bet-hedging hypothesis, if the habitat is unpredictable, females should have an advantage by producing more variable offspring. In the case of Maculinea butterflies, this may involve maintaining larvae that develop in 1 or 2 years.

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