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Study area and samplingSmall mammals (murid rodents and shrews) were captured using mouse-type snap traps in Tatarstan, Russian Federation (Fig. 1, Table S1). Area type (urban or rural), vegetation (forest or field) and distance from trapping points to the nearest human settlement were recorded. The distinction between forest and field was made based on the UN Food and Agriculture Organization’s criteria23,24. Each administrative division in the Tatarstan was defined to be urban or rural by the Federal Service of State Statistics of Russian Federation25. Based on these criteria, Kazan city and Naberezhnye Chelny city were classified as urban districts and Vysokogorsky district, Yelabuzhsky district, Laishevsky district, Mamadyshsky district, Nizhnekamsky district, Pestrechinsky district and Tukayevsky district were classified as rural districts. Small mammals were captured during the spring and fall periods of 2016 and 2017. Fifty traps were placed in a line every 5 m in one place. Traps were baited and left for one night. Animal suffering was minimized as snap traps cause rapid death in murid rodents and shrews. Each captured small mammal’s species, age, and sex were morphologically identified using a reference guide26, and the animals were then stored at − 20 °C until their brains were isolated.EthicsAll experiments were performed in compliance with relevant Russian and Japanese and institutional laws and guidelines and were approved by the Ministry of Health of the Russian Federation and the Animal Research Committee of Gifu University (Permit Nos. MU 3.1.1029-01, and 17060, respectively). Study was carried out in compliance with the ARRIVE guidelines (https://arriveguidelines.org).DNA extraction and PCRBrain tissue samples were prepared as described previously12. Brain samples stored at − 20 °C were transferred to a − 86 °C deep freezer. Each deep-frozen whole brain sample was homogenized in 1 ml of a 0.9% saline solution. Total DNA was extracted from the brain tissues of each small mammal using a Genomic DNA Purification Kit (Promega, Madison, WI, USA), following the manufacturer’s instructions. Nested PCR was performed with the Takara PCR Amplification Kit (Takara Bio Inc., Foster City, California, USA) according to the manufacturer’s instructions. The primer sets and PCR conditions used to detect the B1 gene from T. gondii were those described previously12.MappingSpatial referencing of the sampling sites was conducted using global positioning system navigation with a Garmin eTrex 10 device. Visualization of cartographic data and measurements of the distances from the trapping points to the nearest human settlements were performed using QGIS 3.12 software27. Geodetic coordinates were projected into planar rectangular coordinates in the Universal Transverse Mercator projection on the WGS-84 ellipsoid (Universal Transverse Mercator, zone 39N). The overview map of the European part of Russia was made in the Lambert Conformal Conic Projection. Map coordinates are represented as geodetic coordinates (WGS-84, degrees and minutes north latitude and east longitude). To visualize thematic objects (administrative boundaries, forests, agricultural lands, and water bodies), a set of vector data layers, NextGIS (Russia), was purchased from OpenStreetMap and contributors, 2021 (https://data.nextgis.com). Data license: ODbL.Dataset and statistical analysesMultivariate logistic regression was performed using the R statistical software package (version 3.6.3)28 to assess the trapping point area (urban or rural), vegetation (forest or field), small mammal species type (alien or non-alien species), age (0–2 months-old juveniles, 3–6 months-old adults or ≧ 6 months old), sex (male or female) and distance from trapping points to the nearest human settlements as risk factors for PCR positivity. According to previous reports2,13,16,17,18, four species, Mi. arvalis, A. flavicollis, A. agrarius, A. uralensis, and three species, My. glareolus, S. araneus and D. nitedula are considered alien and non-alien species, respectively. Quantitative data were replaced with 0 or 1 dummy variables, and age data were replaced by 0, 1 and 2 for juveniles, adults and elders, respectively. Multicollinearity of the explanatory variables was evaluated using Spearman’s coefficient29 calculated using dplyr, FSA and psych packages30,31,32. None of the Spearman’s coefficients were  > 0.6. To find the best fit model, a forward selection procedure was used. Predictive performance and model fitting were assessed using the area under the receiver operating characteristic (ROC) curve, area under the curve (AUC) and corrected Akaike’s information criterion (AICc) with Akaike weight (Wi). AICc and Wi were calculated using the MuMIN package33, and the AUC was calculated using the R pROC package34. P-values of  More

The micro-CT results from Arthrorhynchus agree perfectly with the previously known light microscope and transmission electron microscope images2. This emphasizes that microtomography is a good technique to visualize the type of fungal attachment to the host and especially the penetration of the cuticle, apart from the study of thallus in amber fossils17. As Jensen et al. (2019) demonstrated the presence of a haustorium in Arthrorhynchus using scanning electron microscopy, we are confident that the lack of penetration and haustorium in Rickia found by micro-CT is real. This is also in agreement with results from the scanning electron microscopical investigation of the attachment sites of R. gigas, which exhibits no indication of penetration and are very similar to those of R. wasmannii previously shown18.Despite the absence of a haustorium, and hence without any obvious means of obtaining nutrition, Rickia gigas is quite a successful fungus, being often abundant on several species of Afrotropical millipedes of the family Spirostreptidae10. It was originally described from Archispirostreptus gigas, and Tropostreptus (= ‘Spirostreptus’) hamatus20, and was subsequently reported from several other Tropostreptus species19.A further challenge for Laboulbeniales growing on millipedes is that infected millipedes, in some species even adults, may moult, shedding the exuviae with the fungus, as has been observed by us on an undescribed Rickia species on a millipede of the genus Spirobolus (family Spirobolidae).The question of how non-haustoriate Laboulbeniales obtain nutrients has been discussed by several authors18, including staining experiments using fungi of the non-haustoriate genus Laboulbenia on various beetles21. Whereas the surface of the main thallus was almost impenetrable to the dye applied (Nile Blue), the smaller appendages could sometimes be penetrated21. The dye injection into the beetle elytra upon which the fungi were sitting, actually spread from the elytron into the fungus, thus indicating that in spite of the lack of a haustorium, the fungus is able to extract nutrients from the interior of its host21.Such experiments have not been performed on Rickia species, but the possibility that nutrients may pass from the host into the basis of the fungus cannot be excluded. For this genus, or at least R. gigas, there may, however, be an alternative way to obtain nutrients: the small opening in the circular wall by which the thallus is attached to the host may allow nutrients from the surface of the millipede or from the environment to seep into the foot of the fungus. However, further experiments are needed in order to evaluate this hypothesis. Moreover, we should not exclude a potential role of primary and secondary appendages in Laboulbeniales nutrition, as we still do not understand exactly their functional role on the fungus life cycle11.The predominant position of the Laboulbeniales on the host might be related to the absence or presence of a haustorium. Thus, the haustoriate species of the genus Arthrorhynchus are most frequently encountered in large numbers on the arthrodial membranes of the host’s abdomen, although some thalli are found on legs2,22. At the arthrodial membranes the cuticle is more flexible and therefore might be easier to penetrate by a parasite. Furthermore, most tissues providing/storing nutrition (e.g., fat body) are located within the abdomen. In contrast, non-haustoriate fungi as are often located on more stiff and sclerotized body-parts like the genus Rickia on the legs or body-rings of millipedes7,20,23 or the genus Laboulbenia on the elytra of beetles21,24. A reason for this might be that the non-haustoriate forms, which are only superficially attached to the host need a more or less smooth surface for adherence and can easily become detached from a flexible surface, which is movable in itself, like the arthrodial membrane, while the haustoriate forms are firmly anchored within the hosts’ cuticle.Whereas the vast majority of the more than 2000 described species of Laboulbeniales show no sign of host penetration, haustoria have been reported from some other genera18, including Trenomyces parasitizing bird lice25,26, Hesperomyces growing on coccinellid beetles and Herpomyces on cockroaches (formerly a Laboulbeniales and now in the order Herpomycetales10), with pernicious consequences on the hosts’ fitness18,27. Micro-CT studies on these genera could help to understand the host penetration. In order to fully understand how Laboulbeniales obtain nourishment, although other approaches are, also needed—for the time being it remains a mystery how the non-haustoriate Laboulbeniales sustain themselves. More