Re-evaluation of Zospeum schaufussi von Frauenfeld, 1862 and Z. suarezi Gittenberger, 1980, including the description of two new Iberian species using Computer Tomography (CT) (Eupulmonata, Ellobioidea, Carychiidae)

The present study aims to clarify the confused taxonomy of Z. schaufussi von Frauenfeld, 1862 and Zospeum suarezi Gittenberger, 1980. Revision of Iberian Zospeum micro snails is severely hindered by uncertainties regarding the identity of the oldest Iberian Zospeum species, Z. schaufussi von Frauenfeld, 1862. In this paper, we clarify its taxonomic status by designating a lectotype from the original syntype series and by describing its internal and external shell morphology. Using SEM-EDX, we attempt to identify the area of the type locality cave more precisely than "a cave in Spain". The shell described and illustrated by Gittenberger (1980) as Z. schaufussi appears not to be conspecific with the lectotype shell, and is considered a separate species, Z. gittenbergeri Jochum, Prieto & De Winter, sp. n. Zospeum suarezi was described from various caves in NW Spain. Study of the type material reveals that these shells are not homogenous in shell morphology. The holotype shell of Z. suarezi is imaged here for the first time. The paratype shell, illustrated by Gittenberger (1980) from a distant, second cave, is described as Zospeum praetermissum Jochum, Prieto & De Winter, sp. n. The shell selected here as lectotype of Z. schaufussi, was also considered a paratype of Z. suarezi by Gittenberger (1980). Since this specimen is morphologically very similar to topotypic shells of Z. suarezi, the latter species is considered a junior synonym of Z. schaufussi (syn. n.). The internal shell morphology of all these taxa is described and illustrated using X-ray Micro Computer Tomography (Micro-CT).Special gratitude goes to Anita Eschner (NHMW) for helping AJ access the von Frauenfeld collection and for providing valuable insights and primary literature. We are grateful to Markus Heneka and Andreas Heneka (RJL Micro & Analytic GmbH, Karlsdorf-Neuthard) for their help and technical prowess with the CT and SEM-EDX scans. We thank Katharina Jaksch-Mason (NHMW) for LM imaging the Z. schaufussi syntype material. We also gratefully acknowledge Emmanuel Tardy's (MHNG) image contributions and notes of the Gittenberger (1980) material formerly housed in the MHNG collection. Appreciation also goes to Gerald Favre for sharing his excellent forty-year-old speleological field notes with us. We acknowledge Thomas Neubauer and Michael Duda for their kind help in transporting the lectotype back and forth from Vienna. We thank the editor, Thierry Backeljau, the ZooKeys editorial team and our reviewers, Benjamin Gomez, Edmund Gittenberger and Barna Pall-Gergely for their helpful suggestions towards improving the manuscript. Lastly, we are indebted to SYNTHESYS for providing generous support in the form of three grants to AJ from the SYNTHESYS Project http://www.synthesys.info/, which is financed by the European Community Research Infrastructure Action under the FP7 "Capacities" Program

Raccoons contraband – The metazoan parasite fauna of free-ranging raccoons in central Europe

The invasive raccoon (Procyon lotor) is an abundant carnivore and considered as an important potential vector of infectious diseases and parasites in Europe. Raccoons show a broad, opportunistic, omnivorous food spectrum. Food supply and habitat quality in urban areas are very attractive for the generalist raccoon. This inevitably leads to increased interaction with humans, domestic animals and livestock, making the raccoon a potentially suitable zoonosis vector. In its autochthonous range, especially in the Eastern and Midwestern United States, the raccoon has been studied very intensively since the beginning of the 20th century. Whereas, basic field biology and parasitology studies in Germany and Europe are lacking and have only been conducted sporadically, regionally and on small sample sizes. In the presented study 234 raccoons from central Germany were comprehensively examined for their metazoan parasite fauna. The present study shows for the first time an extremely diverse parasite fauna in raccoons outside their native range and proves their essential role as intermediate hosts and hosts for ecto- and endoparasites. A total of 23 different parasite species were identified, five of which are human pathogens, 14 of which are new for the parasite fauna of raccoons in Europe. The human pathogenic raccoon roundworm Baylisascaris procyonis is the most common parasite species in this study, with a prevalence of up to 95%. The digenetic trematode Plagiorchis muris, another human pathogenic parasite species, was detected for the first time in raccoons. The ongoing spread of invasive carnivores and the associated spread and transmission of their parasites and other pathogens increases the potential health risk of wild and farmed animals as well as humans. An increase in parasitic diseases in humans (e.g. raccoon roundworm) is to be expected, especially in urban areas, where raccoons are becoming more and more abundant

Re-evaluation of Zospeum schaufussi von Frauenfeld, 1862 and Z. suarezi Gittenberger, 1980, including the description of two new Iberian species using Computer Tomography (CT) (Eupulmonata, Ellobioidea, Carychiidae)

The present study aims to clarify the confused taxonomy of Z. schaufussi von Frauenfeld, 1862 and Zospeum suarezi Gittenberger, 1980. Revision of Iberian Zospeum micro snails is severely hindered by uncertainties regarding the identity of the oldest Iberian Zospeum species, Z. schaufussi von Frauenfeld, 1862. In this paper, we clarify its taxonomic status by designating a lectotype from the original syntype series and by describing its internal and external shell morphology. Using SEM-EDX, we attempt to identify the area of the type locality cave more precisely than “a cave in Spain”. The shell described and illustrated by Gittenberger (1980) as Z. schaufussi appears not to be conspecific with the lectotype shell, and is considered a separate species, Z. gittenbergeri Jochum, Prieto & De Winter, sp. n. Zospeum suarezi was described from various caves in NW Spain. Study of the type material reveals that these shells are not homogenous in shell morphology. The holotype shell of Z. suarezi is imaged here for the first time. The paratype shell, illustrated by Gittenberger (1980) from a distant, second cave, is described as Zospeum praetermissum Jochum, Prieto & De Winter, sp. n. The shell selected here as lectotype of Z. schaufussi, was also considered a paratype of Z. suarezi by Gittenberger (1980). Since this specimen is morphologically very similar to topotypic shells of Z. suarezi, the latter species is considered a junior synonym of Z. schaufussi (syn. n.). The internal shell morphology of all these taxa is described and illustrated using X-ray Micro Computer Tomography (Micro-CT)

Hämatophage Dipteren : Makro- und mikroökologische Einflüsse auf ihre Verbreitung

Viele Gruppen der Lebewesen, insbesondere Insekten breiten sich durch steigende Temperaturen zunehmend in Gebieten aus, in denen sie ursprünglich nicht vorkommen(Novikov und Vaulin 2014; Bebber 2015). Hierbei ist die steigende Temperatur in verschiedenen Gebieten der Hauptfaktor für Expansionen dieser Arten in Richtung des nördlichen Polarkreises. Einige dieser Arten sind sehr tolerant für verschiedene Variablen und können damit ihr Verbreitungsgebiet deutlich nach Norden hin ausdehnen. Aufgrund steigender Temperaturen werden jedoch andere Arten in ihrem Verbreitungsgebiet eingeschränkt oder ihre Verbreitung verschiebt sich in nördliche Richtung (Ogden und Lindsay 2016; Lawler et al. 2009). Auch für die Verbreitung von Krankheiten spielen Temperaturen, Ausbreitungen oder Verbreitungsverschiebungen eine wichtige Rolle (Mordecai et al. 2019). So können, durch die Etablierung der passenden Vektoren, bisher nur in wärmeren Gebieten auftretende Krankheiten zukünftig auch in unseren Breitengraden eingeschleppt und verbreitet werden. Bremsen, invasive Stechmücken aber auch einheimische Mücken tragen alle ein Potential,verschiedenste Krankheitserreger zu verbreiten, auch wenn die Eignung als Vektor für jede Art unterschiedlich groß ausfällt und manche Arten daher kaum beobachtet und untersucht werden. Mit dem Augenmerk auf sich ändernde Verbreitungsgebiete hinsichtlich zukünftigen klimatischen Veränderungen und sich wandelnden anthropogenen Einflüssen sollten jedoch auch Arten mit bisher geringem Vektorpotential mit in Beobachtungsprogramme aufgenommen werden. Wir untersuchten in Projekt I auf kontinentaler Skala die Verbreitung von sechs verschiedenen Bremsenarten und konnten sowohl Rückschlüsse auf eine mangelhafte Beobachtung der Arten ziehen als auch Artpräferenzen hinsichtlich der Landschaftsnutzung, Auswirkungen des Klimas auf die Verbreitung der Art und bisher unbekannte Toleranzen hinsichtlich tiefen Temperaturen und äußerst verkürzten Wärmeperioden aufdecken. Eine Größenordnung niedriger wurde in Projekt II, basierend auf aktuellen und Vergangenen Klimadaten, die zukünftige und aktuelle Verbreitung einer invasiven, sich zukünftig ausbreitenden Stechmückenart innerhalb Deutschlands modelliert. Durch bisherig im Untersuchungsgebiet nur begrenztes Auftreten konnten noch keine Rückschlüsse auf die unterschiedlichen Präferenzen für das Habitat gezogen werden, es können jedoch für zukünftige Berechnungen Habitatpräferenzen aus anderen Gebieten hinzugezogen werden um die Art und ihre fortschreitende Ausbreitung genauer zu beobachten. Auf der kleinsten untersuchten Ebene konnten in Projekt III innerhalb eines Mikrohabitates verschiedenste Rückschlüsse auf limitierende oder förderliche abiotische Faktoren, die teilweise bisherig nicht oder nur geringfügig beobachtet wurden, gezogen werden. Ebenfalls konnten Auswirkungen der umgebenden Landschaft auf die Abundanzen der Tiere beobachtet werden. Mithilfe von verschiedenen Modellen und in Abhängigkeit von Klimakarten, Landbedeckungsdaten und Landnutzung sowie Eigenschaften und Toleranzen der untersuchten Arten lassen sich in verschiedenen Größenordnungen geeignete Habitate von einheimischen sowie invasiven Arten identifizieren und zukünftige Verbreitungen effizient vorhersagen. Insgesamt können, basierend auf all diesen Daten, dadurch für alle untersuchten Faktoren Modelle auf andere Gebiete übertragen werden um somit potentielle Verbreitungen dort vorherzusagen. Auf unseren Daten basierend können so zum Beispiel Modellierungen für die potentielle Ausbreitung der untersuchten Tabaniden innerhalb anderer Kontinente berechnet werden und Monitoringprogramme können die Ergebnisse unserer Studie als Startpunkt aufgreifen, um durch Beprobung an modellierten Standorten die Korrektheit unserer Modelle zu überprüfen und sowohl Landschaftstypen als auch Artzusammensetzung aufzunehmen um das Modell zu bestätigen oder zu verbessern. Die Modellierung der invasiven Art Aedes albopictus bietet die Möglichkeit, diese Art in Zukunft innerhalb der möglichen Ausbreitungskorridore genauer zu beobachten um ihre fortschreitende Verbreitung zu verifizieren oder eventuelle Änderungen des klimatischen Verlaufes mit einzubinden und das Modell anzupassen. Die Untersuchung des Mikrohabitats von Culex pipiens pipiens und Culex torrentium bietet, auch hinsichtlich anderer Arten in diesem Habitat, eine potente Methode, Vorhersagen für Artvorkommen innerhalb anderer Unterirdischen Objekte zu berechnen. Hier können, bei ausreichend großer Datenlage, eine Vielzahl von Faktoren in die Auswertung mit einfließen. Die durchgeführten Studien bestätigen die Notwendigkeit für verbesserte Monitoringkonzepte für alle vektorkompetenten Tiergruppen hinsichtlich der sich ändernden klimatischen Bedingungen, des globalen Handels und die sich wandelnde Nutzung der Landschaften durch den Menschen und darin begründete Veränderungen der Artenzusammensetzung eines Habitates, zeigen Möglichkeiten, diese Konzepte mit bisher ungenutzten Daten aufzubauen und zu verbessern und können gleichzeitig zu deren Verbesserung herangezogen werden

Incompletely observed : niche estimation for six frequent European horsefly species (Diptera, Tabanoidea, Tabanidae)

Background: More than 170 species of tabanids are known in Europe, with many occurring only in limited areas or having become very rare in the last decades. They continue to spread various diseases in animals and are responsible for livestock losses in developing countries. The current monitoring and recording of horseflies is mainly conducted throughout central Europe, with varying degrees of frequency depending on the country. To the detriment of tabanid research, little cooperation exists between western European and Eurasian countries. Methods: For these reasons, we have compiled available sources in order to generate as complete a dataset as possible of six horsefly species common in Europe. We chose Haematopota pluvialis, Chrysops relictus, C. caecutiens, Tabanus bromius, T. bovinus and T. sudeticus as ubiquitous and abundant species within Europe. The aim of this study is to estimate the distribution, land cover usage and niches of these species. We used a surface-range envelope (SRE) model in accordance with our hypothesis of an underestimated distribution based on Eurocentric monitoring regimes. Results: Our results show that all six species have a wide range in Eurasia, have a broad climatic niche and can therefore be considered as widespread generalists. Areas with modelled habitat suitability cover the observed distribution and go far beyond these. This supports our assumption that the current state of tabanid monitoring and the recorded distribution significantly underestimates the actual distribution. Our results show that the species can withstand extreme weather and climatic conditions and can be found in areas with only a few frost-free months per year. Additionally, our results reveal that species prefer certain land-cover environments and avoid other land-cover types. Conclusions: The SRE model is an effective tool to calculate the distribution of species that are well monitored in some areas but poorly in others. Our results support the hypothesis that the available distribution data underestimate the actual distribution of the surveyed species

Traversing worlds - dispersal potential and ecological classification of Speolepta leptogaster (Winnertz, 1863) (Diptera, Mycetophilidae)

Speolepta leptogaster (Winnertz, 1863) is frequently occurring in European subterranean environments. As for most cave animals, studies addressing non-anatomical aspects are sparse. Here we present the first molecular study on S. leptogaster. We investigated the demographic structure (i.e. COI locus) of 69 specimens from 36 underground populations in Hesse (Central German Uplands) to get first insights into the species’ dispersal ability. In total, 14 haplotypes were revealed. Haplotype diversity was relatively high, whereas nucleotide diversity was low. Furthermore, a significant but low pattern of isolation-by-distance and (a) past population expansion event(s) were detected. Our genetic results suggest a (good) active dispersal ability for Speolepta leptogaster. The occurrence of several surface records of adult specimens corroborates this hypothesis. We discuss the developmental stages of S. leptogaster in the context of the ecological classification system and regard the species as a eutroglophile. Evidence has been found to distinguish two larval types. A reconstructed life-cycle of the species is provided

An investigation of hibernating members from the Culex pipiens complex (Diptera, Culicidae) in subterranean habitats of central Germany

The Culex pipiens complex encompasses five species and subspecies of the genus Culex. Over time, a multitude of morphologically indistinguishable species has been assigned to this complex with several species being classified as important vectors for different diseases. Some species of this complex hibernate in subterranean habitats, and it has been proven that viruses can survive this phase of hibernation. However, studies focusing on the environmental requirements, ecology and spatial and temporal distribution patterns of mosquitos in underground habitats are sparse. Here, we investigate the main environmental factors and dependencies of Culex, considering the number of individuals and survival probabilities in underground habitats during the winter months. Methods. Since the State of Hesse, Germany harbors about 3500 to 4000 subterranean shelters ample availability of subterranean habitats there provides a good opportunity to conduct detailed investigations of the Culex pipiens complex. In this study, we identified a sample of 727 specimens of overwintering females within the Culex pipiens complex from 52 different underground sites collected over a period of 23 years using qPCR. A complete data set of samplings of hibernating mosquitos from 698 subterranean habitats in Central Germany over the same period was available to study the spatial and temporal patterns and the effect of temperature and precipitation conditions on these hibernating populations using a generalized linear model (GLM). Results. Our qPCR-results show, similar to aboveground studies of mosquitos, that Culex pipiens pipiens and Culex torrentium occur sympatrically. On the other hand, Culex pipiens molestus occurred very rarely. The GLM revealed no shifts in species composition over time, but different preferences for subterranean hibernacula, chemical effects on overwintering populations as well as effects of annual and seasonal mean temperature and precipitation during the active phase from March to November. Cx. p. pipiens and Cx. torrentium are the most common species within Hessian caves and other underground habitats during winter. They co-occur with different frequency without any patterns in species composition. Weather conditions influence the number of overwintering mosquitos during the activity phase. Depending on cave parameters, the number of mosquitos decreases during the winter months

Spatial and temporal patterns of human Puumala virus (PUUV) infections in Germany

Background Worldwide, the number of recorded human hantavirus infections as well as the number of affected countries is on the rise. In Europe, most human hantavirus infections are caused by the Puumala virus (PUUV), with bank voles (Myodes glareolus) as reservoir hosts. Generally, infection outbreaks have been related to environmental conditions, particularly climatic conditions, food supply for the reservoir species and land use. However, although attempts have been made, the insufficient availability of environmental data is often hampering accurate temporal and spatially explicit models of human hantavirus infections. Methods In the present study, dynamics of human PUUV infections between 2001 and 2015 were explored using ArcGIS in order to identify spatio-temporal patterns. Results Percentage cover of forest area was identified as an important factor for the spatial pattern, whereas beech mast was found explaining temporal patterns of human PUUV infections in Germany. High numbers of infections were recorded in 2007, 2010 and 2012 and areas with highest records were located in Baden-Wuerttemberg (southwest Germany) and North Rhine-Westphalia (western Germany). Conclusion More reliable data on reservoir host distribution, pathogen verification as well as an increased awareness of physicians are some of the factors that should improve future human infection risk assessments in Germany

Spatial and temporal patterns of human Puumala virus (PUUV) infections in Germany

Background: Worldwide, the number of recorded human hantavirus infections as well as the number of affected countries is on the rise. In Europe, most human hantavirus infections are caused by the Puumala virus (PUUV), with bank voles (Myodes glareolus) as reservoir hosts. Generally, infection outbreaks have been related to environmental conditions, particularly climatic conditions, food supply for the reservoir species and land use. However, although attempts have been made, the insufficient availability of environmental data is often hampering accurate temporal and spatially explicit models of human hantavirus infections. Methods: In the present study, dynamics of human PUUV infections between 2001 and 2015 were explored using ArcGIS in order to identify spatio-temporal patterns. Results: Percentage cover of forest area was identified as an important factor for the spatial pattern, whereas beech mast was found explaining temporal patterns of human PUUV infections in Germany. High numbers of infections were recorded in 2007, 2010 and 2012 and areas with highest records were located in Baden-Wuerttemberg (southwest Germany) and North Rhine-Westphalia (western Germany). Conclusion: More reliable data on reservoir host distribution, pathogen verification as well as an increased awareness of physicians are some of the factors that should improve future human infection risk assessments in Germany

Three new species of Carychium O.F. Müller, 1773 from the Southeastern USA, Belize and Panama are described using computer tomography (CT) (Eupulmonata, Ellobioidea, Carychiidae)

Three new species of the genus Carychium O.F. Müller, 1773, Carychium hardiei Jochum & Weigand, sp. n., Carychium belizeense Jochum & Weigand, sp. n. and Carychium zarzaae Jochum & Weigand, sp. n. are described from the Southeastern United States, Belize and Panama, respectively. In two consecutive molecular phylogenetic studies of worldwide members of Carychiidae, the North and Central American morphospecies Carychium mexicanum Pilsbry, 1891 and Carychium costaricanum E. von Martens, 1898 were found to consist of several evolutionary lineages. Although the related lineages were found to be molecularly distinct from the two nominal species, the consequential morphological and taxonomic assessment of these lineages is still lacking. In the present paper, the shells of these uncovered Carychium lineages are assessed by comparing them with those of related species, using computer tomography for the first time for this genus. The interior diagnostic characters are emphasized, such as columellar configuration in conjunction with the columellar lamella and their relationship in context of the entire shell. These taxa are morphologically described and formally assigned their own names