Hagnosa longicapillata, gen. nov., sp. nov., a New Sordariaceous Ascomycete in the Indoor Environment, and the Proposal of Hagnosaceae fam. nov.

Hagnosa longicapillata, gen. nov., sp. nov, is described and illustrated from wooden building materials collected in Hungary and from pure culture. This species has been collected exclusively from indoor environments, where it was quite common. The ascocarps develop in a thick layer of brown, woolly mats of mycelia. The ostiolar region of the perithecia is ornamented with a five-lobed, flower-shaped crown. Asci are four-spored; ascospores are dark brown, smooth, muriform, not constricted at the septa, and liberated mostly through crackings of the thin ascomatal wall. Apparently, ascospores are dispersed by the mechanical disturbance of the mycelial web. In the phylogenetic tree, Hagnosa samples were placed as a basal lineage, independently from the other family of Sordariomycetidae, with high support. To place Hagnosa in Sordariales, the new family, Hagnosaceae, is proposed.

Ectoparasitic fungi of Myrmica ants alter the success of parasitic butterflies.

Exploitation of organisms by multiple parasite species is common in nature, but interactions among parasites have rarely been studied. Myrmica ants are rich in parasites. Among others, the ectoparasitic Rickia wasmannii fungus and the parasitic caterpillars of myrmecophilous Phengaris butterflies often infect the same Myrmica colonies. In this study, we examined the effects of R. wasmannii on the adoption, long-term development, and survival of P. alcon. In laboratory conditions, caterpillars introduced into nests of Myrmica scabrinodis uninfected with R. wasmannii survived significantly longer compared to caterpillars introduced into infected nests. In the field, joint infection was less common than expected if both parasites exploited M. scabrinodis colonies independently. Pre-pupal caterpillars of P. alcon were somewhat larger in nests infected with R. wasmannii than those found in uninfected nests. Based on these results it seems that R. wasmannii infection of M. scabrinodis affects the survival and development of P. alcon caterpillars, suggesting competition between these two ant parasites.

Ectoparasitic fungi Rickia wasmannii infection is associated with smaller body size in Myrmica ants.

Parasitism-generated negative effects on ant societies are multifaceted, implying individual and colony-level responses. Though laboratory based evidence shows that the sublethal fungus Rickia wasmannii is responsible for physiological and behavioral responses that may negatively affect individual workers' resilience and life expectancy in Myrmica ant workers, colony-level stress response to this parasite is largely unknown. Here, we focus on understanding of a long-term, colony-level effect of Rickia infection on Myrmica scabrinodis ant populations by tracking trait size-based changes. We collected worker specimens from infected and uninfected colonies from the same population in order to: (1) compare body size in response to parasitism, (2) assess the extent to which possible changes in size are associated with the severity of infection, and (3) investigate shifts in body size in response to infection over time by testing correlation of workers' ages and sizes. We found that workers from infected colonies were significantly smaller than their healthy congeners, but neither infection level nor the age of the workers showed significant correlation with the size in infected colonies. Decreasing body sizes in infected colonies can be ascribed to workers' mediated effect toward developing larvae, which are unable to attain the average body size before they pupate.

The Myrmecofauna (Hymenoptera: Formicidae) of Hungary: Survey of Ant Species with an Annotated Synonymic Inventory.

Ants (Hymenoptera: Forimicidae) are exceedingly common in nature. They constitute a conspicuous part of the terrestrial animal biomass and are also considered common ecosystem engineers. Due to their key role in natural habitats, they are at the basis of any nature conservation policy. Thus, the first step in developing adequate conservation and management policies is to build a precise faunistic inventory. More than 16,000 valid ant species are registered worldwide, of which 126 are known to occur in Hungary. Thanks to the last decade's efforts in the Hungarian myrmecological research, and because of the constantly changing taxonomy of several problematic ant genera, a new checklist of the Hungarian ants is presented here. The state of the Hungarian myrmecofauna is also discussed in the context of other European countries' ant fauna. Six species (Formica lemani, Lasius nitidigaster, Tetramorium immigrans, T. staerckei, T. indocile and Temnothorax turcicus) have been reported for the first time in the Hungarian literature, nine taxon names were changed after systematic replacements, nomenclatorial act, or as a result of splitting formerly considered continuous populations into more taxa. Two species formerly believed to occur in Hungary are now excluded from the updated list. All names are nomenclaturally assessed, and complete synonymies applied in the Hungarian literature for a certain taxon are provided. Wherever it is not self-evident, comments are added, especially to explain replacements of taxon names. Finally, we present a brief descriptive comparison of the Hungarian myrmecofauna with the ant fauna of the surrounding countries. The current dataset is a result of ongoing work on inventorying the Hungarian ant fauna, therefore it is expected to change over time and will be updated once the ongoing taxonomic projects are completed.

Host Ant Change of a Socially Parasitic Butterfly (Phengaris alcon) through Host Nest Take-Over.

The socially parasitic Alcon blue butterfly (Phengaris alcon) starts its larval stage by feeding on the seeds of gentians, after which it completes development in the nests of suitable Myrmica ant species. The host plant and host ant species can differ at the population level within a region, and local adaptation is common, but some host switches are observed. It has been suggested that one mechanism of change is through the re-adoption of caterpillars by different ant species, either through occupation of abandoned nests or take-over of established nests by competitively superior colonies. To test this question in the lab we introduced relatively strong colonies (50 workers) of alien Myrmica species to the arenas of weaker colonies (two caterpillars with six workers), and to orphaned caterpillars (two caterpillars without ants). We used caterpillars from a xerophylic population of P. alcon, and both local hosts, M. sabuleti and M. scabrinodis, testing the possibility of host switch between these two host ant species during larval development. Most of the caterpillars were successfully readopted by alien ants, and survived well. Our results suggest higher ecological plasticity in host ant usage of this butterfly than generally thought.

Studies of Laboulbeniales on Myrmica ants (IV): host-related diversity and thallus distribution patterns of Rickia wasmannii.

Fungal species identities are often based on morphological features, but current molecular phylogenetic and other approaches almost always lead to the discovery of multiple species in single morpho-species. According to the morphological species concept, the ant-parasitic fungus Rickia wasmannii (Ascomycota, Laboulbeniales) is a single species with pan-European distribution and a wide host range. Since its description, it has been reported from ten species of Myrmica (Hymenoptera, Formicidae), of which two belong to the rubra-group and the other eight to the phylogenetically distinct scabrinodis-group. We found evidence for R. wasmannii being a single phylogenetic species using sequence data from two loci. Apparently, the original morphological description (dating back to 1899) represents a single phylogenetic species. Furthermore, the biology and host-parasite interactions of R. wasmannii are not likely to be affected by genetic divergence among different populations of the fungus, implying comparability among studies conducted on members of different ant populations. We found no differences in total thallus number on workers between Myrmica species, but we did observe differences in the pattern of thallus distribution over the body. The locus of infection is the frontal side of the head in Myrmica rubra and M. sabuleti whereas in M. scabrinodis the locus of infection differs between worker ants from Hungary (gaster tergites) and the Netherlands (frontal head). Possible explanations for these observations are differences among host species and among populations of the same species in (i) how ant workers come into contact with the fungus, (ii) grooming efficacy, and (iii) cuticle surface characteristics.

Patterns of host use by brood parasitic Maculinea butterflies across Europe.

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'.

Herpomyces ectoparasitic fungi (Ascomycota, Laboulbeniales) are globally distributed by their invasive cockroach hosts and through the pet trade industry.

The introduction of non-native animals occasionally results in the co-introduction of their microbial symbionts or parasites. The trade of exotic pets and zoo animals has inadvertently introduced several parasitic species to countries where they are non-native. Both the presence of suitable native hosts and opportunity for dispersal determine whether these non-native species become naturalized. During our studies dealing with species of Herpomyces (Ascomycota, Laboulbeniomycetes), fungi that are exclusively ectoparasitic on cockroaches (Hexapoda, Blattodea), we make use of artificial colonies. Most of our specimens originate from pet stores and laboratory populations. Although they were originally intended for transmission studies, we discovered that some cockroaches from artificial colonies carried fruiting bodies of Herpomyces. We screened a total 292 cockroaches from 11 populations that we maintained after purchase. Sources were different pet stores, a toxicological laboratory, and a biological supply company. In eight populations, we found at least some Herpomyces-infected cockroaches. Parasite prevalence varied between 8.77% and 86.33%. Host associations were Blatta orientalis with Herpomyces stylopygae, Blattella germanica with H. ectobiae, Periplaneta americana with H. periplanetae, Phoetalia pallida with H. leurolestis, and Shelfordella lateralis with an undescribed species of Herpomyces. Apart from the new reports, host associations, and consequences for taxonomy (a new species based on morphological and molecular characters), we started to think about the geographic distributions of these fungi and how we, humans, shape them through spreading hosts and through international pet trade. We reviewed the currently known records of Herpomyces-associated cockroaches and host-parasite relationships. Based on the available data, on a global scale, at least half of the currently known species of Herpomyces are spread by globally invasive host species and through international pet trade. This indicates that the distribution and host range of these obscure and often unnoticed fungi are affected by human activities.

Studies of Laboulbeniales on Myrmica ants (III): myrmecophilous arthropods as alternative hosts of Rickia wasmannii.

Myrmecophilous arthropods and their manifold relations to host ants are interesting from an evolutionary perspective. Rickia wasmannii is an ectoparasitic fungus belonging to the Laboulbeniales order. Here, we show that inquiline mites can become infected by R. wasmannii, which was thought to be restricted to the genus Myrmica (Hymenoptera: Formicidae). This is the first report of R. wasmannii from an alternative host in another subphylum (Chelicerata). We also found immature fruiting bodies on a larva of Microdon myrmicae (Diptera: Syrphidae), which represents the first report of any Rickia species on flies. This fungus is capable of infecting alternative, unrelated host species as they co-occur in the ant nest "microhabitat". These observations provide direct evidence for ecological specificity in Laboulbeniales. The presence of R. wasmannii on inquilines in Myrmica ant nests suggests that the parasite may have adapted to the ant nest environment and is less dependent on acquiring specific nutrients from the hosts. However, the alternative cannot be excluded; these infections might also represent chance events if the fungus is incapable of fulfilling its life cycle.

Host plant use drives genetic differentiation in syntopic populations of Maculinea alcon.

The rare socially parasitic butterfly Maculinea alcon occurs in two forms, which are characteristic of hygric or xeric habitats and which exploit different host plants and host ants. The status of these two forms has been the subject of considerable controversy. Populations of the two forms are usually spatially distinct, but at Rascruci in Romania both forms occur on the same site (syntopically). We examined the genetic differentiation between the two forms using eight microsatellite markers, and compared with a nearby hygric site, Sardu. Our results showed that while the two forms are strongly differentiated at Rascruci, it is the xeric form there that is most similar to the hygric form at Sardu, and Bayesian clustering algorithms suggest that these two populations have exchanged genes relatively recently. We found strong evidence for population substructuring, caused by high within host ant nest relatedness, indicating very limited dispersal of most ovipositing females, but not association with particular host ant species. Our results are consistent with the results of larger scale phylogeographic studies that suggest that the two forms represent local ecotypes specialising on different host plants, each with a distinct flowering phenology, providing a temporal rather than spatial barrier to gene flow.

Rickia wasmannii increases the need for water in Myrmica scabrinodis (Ascomycota: Laboulbeniales; Hymenoptera: Formicidae).

The order Laboulbeniales (Fungi, Ascomycota) is a little-studied group of microscopic ectoparasites of invertebrates, mostly insects. The effects of Laboulbeniales species on their hosts are mostly unknown. Rickia wasmannii Cavara, 1899 is a common Laboulbeniales fungus occurring in Europe and is currently known to be a parasite of at least eight Myrmica ant species. R.wasmannii serves as a good model organism for Laboulbeniales-host interactions, as this species covers the host in a very high density, and infected host individuals can be easily collected in high numbers. The effect of R. wasmannii on the survival rate of its most common host species, Myrmica scabrinodis Nylander 1846, was therefore investigated in a laboratory experiment on an individual level. To enhance the results, environmental stresses were simulated by depriving infected and uninfected workers of water and food. The survival of individuals was recorded hourly until the death of the last individual. Infected specimens were significantly more sensitive to the withdrawal of food and water than uninfected specimens. When we tested for water consumption, we found that infected ants spent more time consuming water than uninfected ants. Therefore, it is possible that infected ants must replace the loss of water. Based on these results, R. wasmannii substantially decreases the chances and time of survival of infected individuals, at least in resource-limited environments, which suggest that R. wasmannii has a negative effect on its host.

Bringing Laboulbeniales into the 21st century: enhanced techniques for extraction and PCR amplification of DNA from minute ectoparasitic fungi.

Laboulbeniales is one of the most peculiar orders of Ascomycota. These fungi are characterized by an ectoparasitic life-style on arthropods, determinate growth, lack of an asexual stage, high species richness, and intractability to culture. The order Laboulbeniales, sister to Pyxidiophorales, has only recently been assigned a separate class, the Laboulbeniomycetes, based on very few ribosomal DNA sequences. So far, DNA isolations and PCR amplifications have proven difficult. Here, we provide details of isolation techniques and the application of commercially available kits that enable efficient and reliable genetic analyses of these fungi. We provide 43 newly generated Laboulbeniales ribosomal DNA sequences, among which are the first published sequences for species in the genera Gloeandromyces, Herpomyces, Laboulbenia, Monoicomyces, and Polyandromyces. DNA extractions were possible using from 1 to 30 thalli from hosts preserved in ethanol (70-100 %). In two cases, we successfully isolated DNA from thalli on dried insect collections. Laboulbeniales molecular systematics could be substantially enhanced through these improved methods by allowing more complete sampling of both taxa and gene regions.

The re-discovered Maculinea rebeli (Hirschke, 1904): Host ant usage, parasitoid and initial food plant around the type locality with taxonomical aspects (Lepidoptera, Lycaenidae).

The taxonomy of the myrmecophilous Maculinea alcon group (Lepidoptera: Lycaenidae) is highly debated. The host-plant and host-ant usage of these butterflies have conventionally been important in their identification. Maculinea 'rebeli' has generally been considered to be the xerophilous form of Ma. alcon (Ma. alcon X hereafter) with Gentiana cruciata as initial food plant. However, the type locality and all other known sites of Ma. rebeli are found above the coniferous zone, and are well separated from the lower regions where Ma. alcon X sites are found. Furthermore, no food plant and host ant data for the nominotypic Ma. rebeli have yet been published. Our aim was therefore to identify the host ant(s) of Ma. rebeli around the type locality and compare this with the host ant usage of nearby Ma. alcon X. Nests of Myrmica spp. (Hymenoptera: Formicidae) close to the host plants were opened on one Ma. alcon X (host plant: Gentiana cruciata) and two Ma. rebeli (host plant: Gentianella rhaetica, first record, confirmed by oviposition and emerging larvae) sites just before the flying period, to find prepupal larvae and pupae. Three Myrmica species (My. lobulicornis, My. ruginodis, My. sulcinodis) were found on the two Ma. rebeli sites, which parasitized exclusively My. sulcinodis (22 individuals in 7 nests). On the Ma. alcon X site Myrmica sabuleti and My. lonae were found, with My. sabuleti the exclusive host (51 individuals in 10 nests). Ichneumon cf. eumerus parasitized both butterflies. The results highlight the differentiation of Maculinea rebeli from Ma. alcon X, from both conservation biological and ecological points of view. Thus, it should be concluded that Ma. rebeli does not simply represent an individual form of Ma. alcon but it can be considered as at least an ecological form adapted to high mountain conditions both in its initial food plant and host ant species. In addition, it should be emphasized that Ma. alcon X (= Ma. rebeli auct. nec Hirschke) cannot be synonymised with Ma. rebeli (Hirschke, 1904).

Individual behaviour in firebugs (Pyrrhocoris apterus).

The concept of animal personalities has recently become of major interest as researchers began to wonder why animals within a given population show consistent behaviour across situations and contexts, what led to the evolution of such behavioural inflexibility and what mechanisms might underlie the phenomenon. A recent model explains individual differences in a population as the result of trade-off between present and future reproduction. We tested this model on the two wing morphs, i.e. short-winged (brachypterous) and long-winged (macropterous) specimens of the firebug (Pyrrhocoris apterus). Since it has been already demonstrated that the two wing morphs differ in their life-history strategies, this species is an ideal subject to test whether the specimens with different life-history strategies have different personalities as well. The results show that individuals behave consistently over time and across contexts, meaning observed bugs do have personalities. We also have found that in females, the two wing morphs have different personalities supporting the theoretical predictions, i.e. winged ones, which are supposed to have lower future reproductive value, are braver and more exploratory. We found no difference between the morphs in males. Differences in reproductive investment might explain this discrepancy between the sexes.