Contribution of cultural heritage values to steppe conservation on ancient burial mounds of Eurasia.

Civilizations, including ancient ones, have shaped global ecosystems in many ways through coevolution of landscapes and humans. However, the cultural legacies of ancient and lost civilizations are rarely considered in the conservation of the Eurasian steppe biome. We used a data set containing more than 1000 records on localities, land cover, protection status, and cultural values related to ancient steppic burial mounds (kurgans); we evaluated how these iconic and widespread landmarks can contribute to grassland conservation in the Eurasian steppes, which is one of the most endangered biomes on Earth. Using Bayesian logistic generalized regressions and proportional odds logistic regressions, we examined the potential of mounds to preserve grasslands in landscapes with different levels of land-use transformation. We also compared the conservation potential of mounds inside and outside protected areas and assessed whether local cultural values support the maintenance of grasslands on them. Kurgans were of great importance in preserving grasslands in transformed landscapes outside protected areas, where they sometimes acted as habitat islands that contributed to habitat conservation and improved habitat connectivity. In addition to steep slopes hindering ploughing, when mounds had cultural value for local communities, the probability of grassland occurrence on kurgans almost doubled. Because the estimated number of steppic mounds is about 600,000 and similar historical features exist on all continents, our results may be applicable at a global level. Our results also suggested that an integrative socioecological approach in conservation might support the positive synergistic effects of conservation, landscape, and cultural values.

Contribución de los valores culturales para la conservación esteparia en los antiguos montículos funerarios de Eurasia Resumen Las civilizaciones modernas y antiguas han moldeado de muchas maneras los ecosistemas globales mediante la coevolución del paisaje y la humanidad. Sin embargo, pocas veces se considera el legado cultural de las civilizaciones perdidas o antiguas para la conservación del bioma de la estepa euroasiática. Usamos un conjunto de datos que contiene más de 1,000 registros de las localidades, cobertura del suelo, estado de protección y valores culturales relacionados con los antiguos montículos funerarios de esta estepa (kurgans). Después analizamos cómo estos símbolos icónicos y distribuidos extensamente pueden contribuir a la conservación de los pastizales en la estepa euroasiática, uno de los biomas en mayor peligro de extinción. Analizamos el potencial de conservación de los montículos en paisajes con diferentes niveles de transformación en el uso de suelo mediante regresiones logísticas generalizadas bayesianas y regresiones logísticas de probabilidades proporcionales. También comparamos el potencial de conservación de los montículos dentro y fuera de las áreas protegidas y evaluamos si los valores culturales locales conservan los pastizales dentro de estas mismas áreas. Los kurgans fueron de gran importancia para la conservación de los pastizales en los paisajes transformados ubicados fuera de las áreas protegidas, en donde llegaron a fungir como hábitats aislados que contribuyeron a la conservación y conectividad del hábitat. Además de que las pendientes pronunciadas impiden el arado, cuando los montículos contaban con valor cultural para las comunidades locales, la probabilidad de que el pastizal se ubicara sobre un kurgan casi se duplicó. Ya que se estima que el número de montículos esteparios ronda los 6,000 y que rasgos históricos similares existen en todos los continentes, nuestros resultados pueden aplicarse a nivel global. Nuestros resultados también sugieren que una estrategia socio-ecológica integradora para la conservación podría respaldar los efectos sinérgicos positivos de la conservación, el paisaje y los valores culturales.

Comparing ant morphology measurements from microscope and online AntWeb.org 2D z-stacked images.

Unprecedented technological advances in digitization and the steadily expanding open-access digital repositories are yielding new opportunities to quickly and efficiently measure morphological traits without transportation and advanced/expensive microscope machinery. A prime example is the AntWeb.org database, which allows researchers from all over the world to study taxonomic, ecological, or evolutionary questions on the same ant specimens with ease. However, the reproducibility and reliability of morphometric data deduced from AntWeb compared to traditional microscope measurements has not yet been tested. Here, we compared 12 morphological traits of 46 Temnothorax ant specimens measured either directly by stereomicroscope on physical specimens or via the widely used open-access software tpsDig utilizing AntWeb digital images. We employed a complex statistical framework to test several aspects of reproducibility and reliability between the methods. We estimated (i) the agreement between the measurement methods and (ii) the trait value dependence of the agreement, then (iii) compared the coefficients of variation produced by the different methods, and finally, (iv) tested for systematic bias between the methods in a mixed modeling-based statistical framework. The stereomicroscope measurements were extremely precise. Our comparisons showed that agreement between the two methods was exceptionally high, without trait value dependence. Furthermore, the coefficients of variation did not differ between the methods. However, we found systematic bias in eight traits: apart from one trait where software measurements overestimated the microscopic measurements, the former underestimated the latter. Our results shed light on the fact that relying solely on the level of agreement between methods can be highly misleading. In our case, even though the software measurements predicted microscope measurements very well, replacing traditional microscope measurements with software measurements, and especially mixing data collected by the different methods, might result in erroneous conclusions. We provide guidance on the best way to utilize virtual specimens (2D z-stacked images) as a source of morphometric data, emphasizing the method's limitations in certain fields and applications.

From Parasitized to Healthy-Looking Ants (Hymenoptera: Formicidae): Morphological Reconstruction Using Algorithmic Processing.

BACKGROUND: Parasites cause predictable alternative phenotypes of host individuals. Investigating these parasitogenic phenotypes may be essential in cases where parasitism is common or taxa is described based on a parasitized individual. Ignoring them could lead to erroneous conclusions in biodiversity-focused research, taxonomy, evolution, and ecology. However, to date, integrating alternative phenotypes into a set of wild-type individuals in morphometric analysis poses extraordinary challenges to experts. This paper presents an approach for reconstructing the putative healthy morphology of parasitized ants using algorithmic processing. Our concept enables the integration of alternative parasitogenic phenotypes in morphometric analyses. METHODS: We tested the applicability of our strategy in a large pool of Cestoda-infected and healthy individuals of three Temnothorax ant species (T. nylanderi, T. sordidulus, and T. unifasciatus). We assessed the stability and convergence of morphological changes caused by parasitism across species. We used an artificial neural network-based multiclass classifier model to predict species based on morphological trait values and the presence of parasite infection. RESULTS: Infection causes predictable morphological changes in each species, although these changes proved to be species-specific. Therefore, integrating alternative parasitogenic phenotypes in morphometric analyses can be achieved at the species level, and a prior species hypothesis is required. CONCLUSION: Despite the above limitation, the concept is appropriate. Beyond parasitogenic phenotypes, our approach can also integrate morphometric data of an array of alternative phenotypes (subcastes in social insects, alternative morphs in polyphenic species, and alternative sexes in sexually dimorphic species) whose integrability had not been resolved before.

Formerly considered rare, the ant species Cryptoponeochracea (Mayr, 1855) can be commonly detected using citizen-science tools.

Citizen science is a valuable tool for monitoring different species, especially in cases concerning truly rare and difficult-to-detect species where time-consuming field studies are limited and long-term research projects are uncertain. To better understand the distribution of the rarely collected Cryptoponeochracea (Mayr, 1855) (Hymenoptera, Formicidae) in Hungary, we obtained the occurrence data with photos uploaded by non-professionals to the page of the largest Hungarian Facebook group dealing with ants and a citizen-science website dealing with biological data collection. In this article, we expand the known distribution of C.ochracea to include 46 new records from Hungary and one from Serbia. With two historical records, this previously undersampled species has now been found 48 times in Hungary. Our results prove that social media platforms and other websites for citizen science projects offer new and useful opportunities for researchers to involve non-professionals in scientific work and, thus, obtain large amounts of valuable data, even for understudied arthropod species.

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.

Functional composition of ant assemblages in habitat islands is driven by habitat factors and landscape composition.

Fragmented natural habitats within human-transformed landscapes play a key role in preserving biodiversity. Ants as keystone species are essential elements of terrestrial ecosystems; thus, it is important to understand the factors influencing their presence. In a large-scale multi-site study, we surveyed ant assemblages using sweep netting and D-vac sampling on 158 ancient burial mounds preserving grassland habitats in agricultural landscapes in East-Hungary. We asked the following questions: (1) How do habitat factors and landscape composition affect species richness and functional diversity of ants? (2) Which ant traits are affected by habitat factors and landscape composition? Despite their small sizes, mounds as permanent and relatively undisturbed landscape elements could provide safe havens for diverse ant assemblages even in transformed agricultural landscapes. The complex habitat structure of wooded mounds supported high species and functional diversity of ant assemblages. Ant species on wooded mounds had small or medium-sized colonies, enabling the co-existence of more species. The effect of landscape composition on ant assemblages was mediated by habitat factors: steep slopes buffered the negative effect of the cropland matrix and enabled higher ant diversity.

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.

Linking environmental heterogeneity and plant diversity: The ecological role of small natural features in homogeneous landscapes.

Small natural features (SNFs), such as road verges, midfield islets, rocky outcrops and ancient burial mounds, provide safe havens for species of natural habitats in human-modified landscapes; therefore, their great ecological importance is in contrast to their small size. SNFs often have a high topographical heterogeneity and abiotic conditions, which differ from their surroundings; therefore, they provide a unique opportunity for establishing links between environmental heterogeneity (EH) and biodiversity. However, no study has so far investigated the EH components of topographically heterogeneous SNFs in a comprehensive framework, by linking environmental and biotic parameters. To fill this knowledge gap, we evaluated the EH components and their effect on biodiversity on ancient mounds covered by semi-natural grasslands in the Pannonian (Hungary) and Continental (Bulgaria) biogeographical regions. We designated 16 study sites, each containing a few-metre-high mounds with five microsites (top, north-, east-, south- and west-facing slopes) and a nearby plain grassland. At each microsite, we measured soil moisture, soil chemical properties, solar radiation and microclimate; and recorded the cover of vascular plants in a total of 480 plots. On the mounds, topographical heterogeneity was associated with sharp differences in microclimate and soil properties. Besides the contrast between mild north-facing and harsh south-facing slopes, east- and west-facing slopes also sustained unique microsites characterised by dynamic diurnal changes in air temperature and vapour pressure deficit. Various combinations of the EH components resulted in unique plant species compositions within the microsites, and supported the co-occurrence of species typical of contrasting habitat types, even within a couple of metres. By combining high-resolution measurements of abiotic factors with fine-scale vegetation sampling, our study provides evidence that widespread SNFs with complex topography harbour several grassland-specialist plant species and introduce a high level of EH to otherwise homogeneous plain landscapes, which cover one third of the global land area.

Consequences of rapid development owing to cohort splitting: just how costly is it to hurry?

In cohort splitting, diverging sub-cohorts may show substantial differences in their growth and developmental rates. Although in the past, causes and adaptive value of cohort splitting were studied in detail, individual-level consequences of cohort splitting are still rather overlooked. Life history theory predicts that considerably increased growth and developmental rates should be traded off against other costly life history traits. However, it is not clear whether one should expect such associations in adaptive developmental plasticity scenarios, because natural selection might have promoted genotypes that mitigate those potential costs of rapid development. To address these contrasting propositions, we assessed life history traits in the wolf spider Pardosa agrestis, both collected from natural habitat and reared in laboratory. We found that some traits are negatively associated with developmental rates in spiders collected from the wild, but these associations were relaxed to a considerable extent in laboratory-reared specimens. In general, we observed no consistent trend for the presence of developmental costs, although some results might suggest higher relative fecundity costs in rapidly developing females. Our study provides a detailed approach to the understanding of individual-level consequences of cohort splitting, and to the associations between key life history traits in adaptive developmental plasticity scenarios.

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.

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.

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.