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Background Ticks and tick-borne diseases play a major role in companion animal health. Additionally, the European tick fauna is changing, for instance due to the spread of Dermacentor reticulatus , displaying a higher likelihood of winter activity than Ixodes ricinus . Therefore, we investigated current tick infestations in dogs and cats in Germany and in parts of Austria and the seasonal infestation risk. Methods Overall, 219 veterinary practices were invited to collect ticks from cats and dogs on a monthly basis. Ticks were morphologically identified and female I. ricinus specimens were measured to estimate attachment duration. Results In total, 19,514 ticks, 17,789 (91.2%) from Germany and 1506 (7.7%) from Austria, were received between March 2020 and October 2021, with 10,287 specimens (52.7%) detached from dogs, 8005 from cats (41.0%) and 1222 from other species (6.3%). In Germany, the most common tick species collected from dogs were I. ricinus (78.0%) and D. reticulatus (18.8%), while cats mainly harboured I. ricinus (91.3%) and I. hexagonus (5.5%) and only few D. reticulatus (0.6%). In Austria, collected I. ricinus reached similar proportions in dogs (90.4%) and cats (95.3%), followed by D. reticulatus in both dogs (5.2%) and cats (1.5%), with I. hexagonus (0.9%) collected only marginally from cats. The average infestation intensity amounted to 1.62 ticks/dog and 1.88 ticks/cat. The single to multiple infestation ratio was 79.1% to 20.9% in dogs and 69.0% to 31.0% in cats, with cats being significantly more often multiple infested than dogs, while the proportion of mixed-species infestations was 2.0% for both dogs and cats. The average attachment duration of female I. ricinus specimens amounted to 78.76 h for dogs and 82.73 h for cats. Furthermore, year-round tick exposure was confirmed, with 108 D. reticulatus and 70 I. ricinus received on average per month during December 2020 to February 2021. Conclusions The study shows a year-round tick infestation risk, with activity of both D. reticulatus and I. ricinus during winter, and confirms the widespread occurrence of D. reticulatus in Germany. Additionally, long average attachment durations and frequent multiple infestations underline the need for adequate year-round tick control, even during the winter months. Graphical Abstract

Background Neurotoxocarosis (NT) is induced by larvae of the dog or cat roundworm ( Toxocara canis or T . cati ) migrating and persisting in the central nervous system of paratenic hosts, including humans, and may be accompanied by severe neurological symptoms. Host- or parasite-induced immunoregulatory processes contribute to the pathogenesis, but detailed data on pathogenic mechanisms and involvement of signalling molecules during cerebral Toxocara species infections are scarce. Methods To elucidate alterations in immunomodulatory mediator pattern, comprehensive multiplex bead array assays profiling comprising 23 different cytokines and chemokines were performed during the course of T . canis - and T . cati -induced NT. To this end, cerebra and cerebella of experimentally infected C57Bl/6 J mice serving as paratenic host models were analysed at six different time points (days 7, 14, 28, 42, 70 and 98) post infectionem (pi). Results Brain-body mass ratios of T . canis and T . cati -infected mice were significantly lower than those of the uninfected control group at day 14 pi, and also at day 28 pi for T . canis -infected mice. Both infection groups showed a continuous decrease of pro-inflammatory cytokine concentrations, including TNF-α, IFN-γ, GM-CSF and IL-6, in the cerebrum over the course of infection. Additionally, T . canis but not T . cati -induced neurotoxocarosis was characterised by significantly elevated levels of anti-inflammatory IL-4 and IL-5 in the cerebrum in the acute and subacute phase of the disease. The higher neuroaffinity of T . canis led to a prominent increase of eotaxin and MIP-1α in both the cerebrum and cerebellum, while in T . cati -infected mice, these chemokines were significantly elevated only in the cerebellum. Conclusions The direct comparison of T . canis - and T . cati -induced NT provides valuable insights into key regulatory mechanisms of Toxocara species in paratenic hosts. The cerebral cyto-/chemokine milieu is shifted to a predominantly anti-inflammatory immune response during NT, possibly enabling both survival of the parasite and the neuroinfected paratenic host. Alteration of eotaxin and MIP-1α concentrations are congruent with the higher neuroaffinity of T . canis and species-specific tropism of T . canis to the cerebrum and T . cati to the cerebellum.

Background Changing geographical and seasonal activity patterns of ticks may increase the risk of tick infestation and tick-borne pathogen (TBP) transmission for both humans and animals. Methods To estimate TBP exposure of dogs and cats, 3000 female I. ricinus from these hosts were investigated for Anaplasma phagocytophilum and Borrelia species. Results qPCR inhibition, which was observed for ticks of all engorgement stages but not questing ticks, was eliminated at a template volume of 2 µl. In ticks from dogs, A. phagocytophilum and Borrelia spp. prevalence amounted to 19.0% (285/1500) and 28.5% (427/1500), respectively, while ticks from cats showed significantly higher values of 30.9% (464/1500) and 55.1% (827/1500). Accordingly, the coinfection rate with both A. phagocytophilum and Borrelia spp. was significantly higher in ticks from cats (17.5%, 262/1500) than dogs (6.9%, 104/1500). Borrelia prevalence significantly decreased with increasing engorgement duration in ticks from both host species, whereas A. phagocytophilum prevalence decreased only in ticks from dogs. While A. phagocytophilum copy numbers in positive ticks did not change significantly over the time of engorgement, those of Borrelia decreased initially in dog ticks. In ticks from cats, copy numbers of neither A. phagocytophilum nor Borrelia spp. were affected by engorgement. Borrelia species differentiation was successful in 29.1% (365/1254) of qPCR-positive ticks. The most frequently detected species in ticks from dogs were B. afzelii (39.3% of successfully differentiated infections; 70/178), B. miyamotoi (16.3%; 29/178), and B. valaisiana (15.7%; 28/178), while B. afzelii (40.1%; 91/227), B. spielmanii (21.6%; 49/227), and B. miyamotoi (14.1%; 32/227) occurred most frequently in ticks from cats. Conclusions The differences in pathogen prevalence and Borrelia species distribution between ticks collected from dogs and cats may result from differences in habitat overlap with TBP reservoir hosts. The declining prevalence of A. phagocytophilum with increasing engorgement duration, without a decrease in copy numbers, could indicate transmission to dogs over the time of attachment. The fact that this was not observed in ticks from cats may indicate less efficient transmission. In conclusion, the high prevalence of A. phagocytophilum and Borrelia spp. in ticks collected from dogs and cats underlines the need for effective acaricide tick control to protect both animals and humans from associated health risks. Graphical Abstract

Background Ixodes ricinus constitutes the main European vector tick for the Lyme borreliosis pathogen Borrelia burgdorferi ( sensu lato ), the relapsing fever borrelia Borrelia miyamotoi , as well as Anaplasma phagocytophilum and several Rickettsia species. Under laboratory conditions, a transovarial transmission to the next tick generation is described for Rickettsia spp. and Borrelia spp., especially regarding B. miyamotoi , whereas the efficiency of transovarial transfer under field conditions is largely unstudied. Methods In order to better estimate the potential infection risk by tick larvae for humans and animals, 1500 I. ricinus larvae from 50 collected “nests” (larvae adhering to the flag in a clumped manner) were individually examined for Borrelia , Rickettsia and A. phagocytophilum DNA using quantitative real-time PCR (qPCR). Results Thirty-nine of 50 nests each (78.0%, 95% CI: 64.0–88.5%) were positive for Borrelia spp. and Rickettsia spp. DNA, and in three nests (6.0%, 95% CI: 1.3–16.5%) A.   phagocytophilum DNA was detected. Overall, DNA from at least one pathogen could be detected in 90.0% (45/50, 95% CI: 78.2–96.7%) of the nests. Of the 1500 larvae, 137 were positive for Borrelia spp. DNA (9.1%, 95% CI: 7.7–10.7%), 341 for Rickettsia spp. DNA (22.7%, 95% CI: 20.6–24.9%) and three for A.   phagocytophilum DNA (0.2%, 95% CI: 0–0.6%). Quantity of Borrelia spp. and Anaplasma spp. DNA in positive larvae was low, with 2.7 × 10 0 Borrelia 5S-23S gene copies and 2.4 × 10 1 A. phagocytophilum msp2/p44 gene copies detected on average, while Rickettsia -positive samples contained on average 5.4 × 10 2 gltA gene copies. Coinfections were found in 66.0% (33/50, 95% CI: 51.2–78.8%) of the nests and 8.6% (38/443, 95% CI: 6.1–11.6%) of positive larvae. In fact, larvae had a significantly higher probability of being infected with Borrelia spp. or Rickettsia spp. when both pathogens were present in the nest. Conclusions This study provides evidence for transovarial transmission of Rickettsia spp. and Borrelia spp. in I. ricinus under field conditions, possibly facilitating pathogen persistence in the ecosystem and reducing the dependence on the presence of suitable reservoir hosts. Further studies are needed to prove transovarial transmission and to explain the surprisingly high proportion of nests containing Rickettsia and/or Borrelia DNA-positive larvae compared to infection rates in adult ticks commonly reported in other studies.

Background Over the last decades, climatic and environmental changes have led to an expanding seasonal activity pattern and increasing distribution of ticks across Europe. In particular, Dermacentor reticulatus is now commonly found on dogs in central Europe. The present study compared attachment sites between Ixodes spp. and Dermacentor reticulatus ticks collected by veterinarians from dogs and cats, and investigated risk factors associated with tick infestation intensity and engorgement duration. Results The dataset comprised 6,335 dogs harbouring 10,287 ticks (8,095 Ixodes ricinus , 1,860 D. reticulatus , 218 Ixodes hexagonus / Ixodes canisuga , 114 of other tick species) respectively 4,248 cats harbouring 8,005 ticks (7,344 I. ricinus , 56 D. reticulatus , 505 I. hexagonus / I. canisuga , 100 of other tick species). Differing sites of tick attachment were not only found between the different host and tick species, but also between the tick developmental stages. Regarding the risk of infestation with multiple ticks, dogs and cats living in rural areas harboured significantly more often multiple than single specimens. Further, a long coat in cats was associated with a higher probability of multiple infestation, while this was not observed in dogs. However, there was a tendency towards a potential influence of the density of the undercoat ( p  = 0.051). In dogs, a tall to very tall body size as well as folded ears increased the risk of multiple infestation, while in cats, increasing age and increasing body size were negatively associated with multiple infestations. Ticks with an engorgement duration of > 48 h were found significantly more often on senior dogs and cats than on younger individuals, as well as on working/utility dog breeds, while engorgement duration was negatively correlated with infestation intensity in dogs. In cats, female gender and a rural residence were significantly associated with longer attachment duration. Conclusions Individual as well as breed specific characteristics can lead to a higher tick infestation intensity or longer engorgement duration. The knowledge of tick attachment sites and specific risk factors can help to raise awareness among owners concerning the importance of tick control with licensed acaricides, as recommended e.g. by the European Scientific Counsel Companion Animal Parasites (ESCCAP), and may aid in early tick removal to decrease the risk of pathogen transmission to dogs and cats whose owners nonetheless refuse acaricidal drugs.

Background The European hedgehog ( Erinaceus europaeus ) is known for high levels of ectoparasitism that not only represents a health risk for the animals themselves, but also for pet animals and humans as hedgehogs are frequently taken into human care. In the present study, patterns of ectoparasite infestation were assessed in hedgehogs taken into care at northern German animal rehabilitation centres. Methods Ectoparasites (ticks, fleas and mites) of 498 hedgehogs were collected over a period of 3 years from July 2018 to May 2021. Species were identified based on morphological characteristics and also via amplification and sequencing of the partial cytochrome c oxidase subunit 2 (COX-2) gene for fleas of the family Ceratophyllidae. Seasonal changes in infestation patterns as well as correlations with animal age, body weight and health status were assessed using generalised linear models. Results Infestation with ticks, fleas and mites occurred throughout the year. Overall, 86.5% (431/498) of the examined hedgehogs were infested with ticks, 91.4% (455/498) with fleas and 17.7% (88/498) with mites . Ixodes ricinus and Ixodes hexagonus / Ixodes canisuga were the most common tick species detected, with the additional occurrence of one Ixodes frontalis. Significant seasonal changes were observed for I. ricinus , but not for I. hexagonus / I. canisuga . Additionally, I. ricinus nymph prevalence declined significantly as of 2020, probably as a consequence of the climate change-related drought as of 2018. In hedgehogs with flea infestations, Archaeopsylla erinacei , Ceratophyllus sciurorum , Nosopsyllus fasciatus and Ctenocephalides felis were identified. In all cases of mite infestation, Caparinia tripilis was detected, in addition to specimens of the family Macronyssidae and free-living mites of the family Acaridae. Statistical analyses showed correlations regarding the factors month, year, body weight and age, but no correlation was evident regarding the health status of the animals. Conclusions With a detected infestation rate of 98.6%, almost all of the examined hedgehogs were infested with at least one ectoparasite species. The seasonal activity patterns of the different ectoparasite species together with the complex annual cycle of hedgehogs lead to different seasonal patterns in ectoparasite prevalence and infestation intensities. Due to the risk of transmission of zoonotic pathogens as well as the possible negative impact on the host itself, hedgehogs should be treated against ectoparasites when taken into care facilities. Graphical Abstract

Background Rickettsiales ( Rickettsia spp. and Anaplasma phagocytophilum ) transmitted by ticks are considered (re-)emerging pathogens posing a risk to public health. Nevertheless, year-long monitoring studies on prevalences of these pathogens in questing ticks to contribute to public health risk assessment are rare. Methods The current study extends previous prevalence surveillances (2005 and 2010) by 2015 to a 10-year monitoring. Therefore, 2100 questing Ixodes ricinus were collected from April to October 2015 at ten different recreation sites in the city of Hanover, Germany, to determine potential changes in tick infection rates with Rickettsiales. Results Of the collected ticks, 288 were adult females, 285 adult males and 1527 nymphs. Overall, 3.8% (79/2100) of ticks were infected with A. phagocytophilum , 50.8% (1066/2100) with Rickettsia spp. and 2.2% (46/2100) with both pathogens. Statistical analyses revealed stagnating A. phagocytophilum infection rates over the 10-year monitoring period, whereas Rickettsia infections increased significantly (33.3% in 2005 and 26.2% in 2010 vs 50.8% in 2015). This increase was also characterized by prominent seasonality with higher prevalences from July to October. Conclusions As increased tick infection rates result in an increased risk for public health, the long-term data reported here provide significant implications for the understanding of progressing Rickettsiales distribution in ticks and essentially contribute to reliable public health risk assessments.

Background Nature conservation with reduced drainage of pastures has been increasingly promoted in agriculture in recent years. However, moisture on pastures is a crucial factor for the development of free-living stages of many parasite species in ruminants. Hence, for the first time, we conducted a field study between 2015 and 2017 at the German North Sea coast to investigate the long-term effect of pasture rewetting (since 2004) on endoparasite infections in sheep and cattle. Methods We examined faecal samples of 474 sheep and 646 cattle from five farms in spring, summer and autumn each year for the presence of endoparasite infections. Animals were kept on conventionally drained, undrained and rewetted pastures. The association between pasture rewetting and endoparasite infection probability was analysed in generalized linear mixed models and including further potential confounders. Results Infection frequencies for gastrointestinal strongyles, Eimeria spp. and Strongyloides papillosus were significantly higher in sheep (62.9%, 31.7% and 16.7%) than in cattle (39.0%, 19.7% and 2.6%). Fasciola hepatica was detected with a frequency of 13.3% in sheep and 9.8% in cattle, while rumen fluke frequency was significantly higher in cattle (12.7%) than in sheep (3.8%). Nematodirus spp., lungworms (protostrongylids, Dictyocaulus viviparus ), Moniezia spp., Trichuris spp. and Dicrocoelium dendriticum were identified in less than 7% of samples. Co-infection with more than three endoparasite taxa was present significantly more often in sheep than in cattle. We identified significant positive correlations above 0.2 for excretion intensities between S. papillosus with strongyles, Eimeria spp. and Nematodirus spp. in sheep and between strongyles and Nematodirus spp. in cattle. Pasture rewetting had no long-term effect on endoparasite infections, neither in sheep nor in cattle. Interestingly, F. hepatica infections decreased significantly in sheep and cattle from 2015 (10.9% and 13.9%) to 2017 (1.4% and 2.1%). Conclusions Pasture rewetting for nature conservation did not increase endoparasite infection probability in ruminants in the long term. This finding should be confirmed in ongoing studies aimed at further animal welfare parameters. The rapid decrease in F. hepatica infections over 3 years may suggest climatic impact or competition with rumen flukes in addition to potential anthelmintic treatment after feedback of the results to the farmers. Graphical Abstract

Fasciola hepatica and Ostertagia ostertagi are internal parasites of cattle compromising physiology, productivity, and well-being. Parasites are complex in their effect on hosts, sometimes making it difficult to identify clear directions of associations between infection and production parameters. Therefore, unsupervised approaches not assuming a structure reduce the risk of introducing bias to the analysis. They may provide insights which cannot be obtained with conventional, supervised methodology. An unsupervised, exploratory cluster analysis approach using the k–mode algorithm and partitioning around medoids detected two distinct clusters in a cross-sectional data set of milk yield, milk fat content, milk protein content as well as F . hepatica or O . ostertagi bulk tank milk antibody status from 606 dairy farms in three structurally different dairying regions in Germany. Parasite–positive farms grouped together with their respective production parameters to form separate clusters. A random forests algorithm characterised clusters with regard to external variables. Across all study regions, co–infections with F . hepatica or O . ostertagi , respectively, farming type, and pasture access appeared to be the most important factors discriminating clusters (i.e. farms). Furthermore, farm level lameness prevalence, herd size, BCS, stage of lactation, and somatic cell count were relevant criteria distinguishing clusters. This study is among the first to apply a cluster analysis approach in this context and potentially the first to implement a k–medoids algorithm and partitioning around medoids in the veterinary field. The results demonstrated that biologically relevant patterns of parasite status and milk parameters exist between farms positive for F . hepatica or O . ostertagi , respectively, and negative farms. Moreover, the machine learning approach confirmed results of previous work and shed further light on the complex setting of associations a between parasitic diseases, milk yield and milk constituents, and management practices.

Ticks transmit a variety of pathogens affecting both human and animal health. In temperate and cold regions of Europe (Western, Central, Eastern, and Northern Europe), the most relevant zoonotic tick-borne pathogens are tick-borne encephalitis virus (TBEV), Borrelia spp. and Anaplasma phagocytophilum . More rarely, Rickettsia spp., Neoehrlichia mikurensis , and zoonotic Babesia spp. are identified as a cause of human disease. Domestic animals may also be clinically affected by these pathogens, and, furthermore, can be regarded as sentinel hosts for their occurrence in a certain area, or even play a role as reservoirs or amplifying hosts. For example, viraemic ruminants may transmit TBEV to humans via raw milk products. This review summarizes the role of domestic animals, including ruminants, horses, dogs, and cats, in the ecology of TBEV, Borrelia spp., A. phagocytophilum, Rickettsia spp., N. mikurensis , and zoonotic Babesia species. It gives an overview on the (sero-)prevalence of these infectious agents in domestic animals in temperate/cold regions of Europe, based on 148 individual prevalence studies. Meta-analyses of seroprevalence in asymptomatic animals estimated an overall seroprevalence of 2.7% for TBEV, 12.9% for Borrelia burgdorferi sensu lato (s.l.), 16.2% for A. phagocytophilum and 7.4% for Babesia divergens , with a high level of heterogeneity. Subgroup analyses with regard to animal species, diagnostic test, geographical region and decade of sampling were mostly non-significant, with the exception of significantly lower B. burgdorferi s.l. seroprevalences in dogs than in horses and cattle. More surveillance studies employing highly sensitive and specific test methods and including hitherto non-investigated regions are needed to determine if and how global changes in terms of climate, land use, agricultural practices and human behavior impact the frequency of zoonotic tick-borne pathogens in domestic animals.