Ixodes Arboricola Research Articles

Atlas of ticks (Acari: Argasidae, Ixodidae) in Germany: 1st data update

The first data update of the atlas of ticks in Germany published in 2021 is presented here. This atlas provides maps based on georeferenced tick locations of 21 species endemic in Germany as well as three tick species that are regularly imported to Germany. The data update includes the following numbers of newly georeferenced tick locations: 17 Argas reflexus, 79 Carios vespertilionis, 2 Dermacentor marginatus, 43 Dermacentor reticulatus, 4 Haemaphysalis concinna, 3 Haemaphysalis punctata, 3 Hyalomma rufipes, 3 Ixodes apronophorus, 9 Ixodes arboricola, 1 Ixodes ariadnae, 30 Ixodes canisuga, 3 Ixodes frontalis, 80 Ixodes hexagonus, 3 Ixodes lividus, 497 Ixodes ricinus/inopinatus, 1 Ixodes rugicollis, 17 Ixodes trianguliceps, 14 Ixodes vespertilionis, and 45 Rhipicephalus sanguineus sensu lato. Old and new tick findings were mapped, such as the northernmost occurrence of D. marginatus in Germany observed in 2021, but also the historical records from the first descriptions of I. apronophorus and I. arboricola, which were georeferenced here for the first time. The digital dataset of tick locations available for Germany is supplemented by 854 new tick locations. These records increase the number of tick species mapped in the federal states Bavaria, Brandenburg and Mecklenburg Western Pomerania by five each, those in Berlin and Schleswig-Holstein by four each, those in Hamburg by three, those in Baden-Wuerttemberg, Bremen, Lower Saxony, Northrhine-Westphalia, Rhineland Palatinate and Thuringia by two each, and those in Hesse, Saxony and Saxony-Anhalt by one each. Thus, the first data update of the tick atlas in Germany and the underlying digital dataset significantly improve our knowledge of the distribution of these tick species and helps to investigate the effects of climate change and habitat changes on them.

Hard ticks (Acari: Ixodidae) associated with birds in Europe: Review of literature data.

Hard ticks (Acari: Ixodidae) are considered the most important transmitters of pathogens in the temperate zone that covers most of Europe. In the era of climate change tick-borne diseases are predicted to undergo geographical range expansion toward the north through regions that are connected to southern areas of the continent by bird migration. This alone would justify the importance of synthesized knowledge on the association of tick species with avian hosts, yet birds also represent the most taxonomically and ecologically diverse part of urban vertebrate fauna. Birds frequently occur in gardens and near animal keeping facilities, thus playing a significant role in the dispersal of ticks and tick-borne pathogens in synanthropic environments. The primary aim of this review is to provide a comprehensive reference source (baseline data) for future studies, particularly in the context of discovering new tick-host associations after comparison with already published data. The records on the ixodid tick infestations of birds were assessed from nearly 200 papers published since 1952. In this period, 37 hard tick species were reported from 16 orders of avian hosts in Europe. Here we compile a list of these tick species, followed by the English and Latin name of all reported infested bird species, as well as the tick developmental stage and country of origin whenever this information was available. These data allowed a first-hand analysis of general trends regarding how and at which developmental stage of ticks tend to infest avian hosts. Five tick species that were frequently reported from birds and show a broad geographical distribution in the Western Palearctic (Ixodes arboricola, I. frontalis, I. ricinus, Haemaphysalis concinna and Hyalomma marginatum) were also selected for statistical comparisons. Differences were demonstrated between these tick species regarding their association with bird species that typically feed from the ground and those that rarely occur at the soil level. The ecology of these five bird-infesting tick species is also illustrated here according to avian orders, taking into account the ecology (habitat type) and activity (circadian rhythm and feeding level) of most bird species that represent a certain order.

Ticks in the metropolitan area of Berlin, Germany

A high-resolution city map showing the geographic distribution of 12 tick species (Acari: Argasidae, Ixodidae) that have been recorded from the metropolitan area of Berlin, Germany is presented. A total of 237 tick locations was mapped. These include ten ixodid tick species: Dermacentor reticulatus, Haemaphysalis concinna, Hyalomma rufipes, Ixodes ricinus, Ixodes canisuga, Ixodes hexagonus, Ixodes arboricola, Ixodes frontalis, Ixodes trianguliceps and Rhipicephalus sanguineus sensu lato. The two tick species Hy. rufipes and R. sanguineus s.l. are not endemic to Berlin. Hyalomma rufipes ticks are introduced in Europe with migratory birds from Africa every spring. Rhipicephalus sanguineus s.l. are introduced to Central Europe with dogs that had travelled to or were imported from countries where this tick is endemic. In Germany, they are able to develop and reproduce inside heated buildings. Occurrences of two soft tick species, the pigeon tick Argas reflexus and the short-legged bat tick Carios vespertilionis were also mapped. Other tick species that are likely to be endemic to Berlin and its environs, but for which documented findings or geographical coordinates are lacking, are mentioned. These include the long-legged bat tick I. vespertilionis and the marten tick I. rugicollis documented in Brandenburg, the federal state surrounding Berlin. It can be assumed that if appropriate field studies are carried out, these tick species will also be found in the metropolitan area of Berlin. The high-resolution mapping of all tick species found in a city (like Berlin) forms the basis for further investigations into the impact of climate change and changing land use on ticks and tick-borne diseases, precisely in those habitats where most people will live in the future.

Plasticity in the timing of detachment of an Eurasian-African songbird tick, Ixodes frontalis

In non-permanent parasites, host detachment should take place in an environment that ensures the continuation of their life cycle. Timing of detachment - in combination with the host's space use - affects dispersal and transmission success of the parasites and of the pathogens they vector. Before reaching the adult reproductive stage, ticks need to go through multiple immature developmental stages (larva and nymph), each feeding on host blood. In between the feeding bouts, they often remain in the off-host environment for considerable periods of time. With this study, we aimed to obtain more insight in Ixodes frontalis’ off-host habitat use by comparing its detachment pattern in different life stages with that of two habitat-specialized ticks also found on birds: the endophilic tree-hole tick (Ixodes arboricola) and the exophilic sheep tick (Ixodes ricinus), the latter living in humid understory vegetation of forests. For this, we artificially infested hole-roosting (great tits, Parus major) and open-roosting (blackbirds, Turdus merula) birds with ticks under laboratory conditions, and recorded whether detachment occurred during the day or the night. We hypothesize that nocturnal detachment improves off-host mating opportunities and host localization, whereas diurnal detachment optimizes tick dispersal. Ixodes frontalis nymphs detached during the night, especially when feeding on blackbirds. This behaviour was very similar to that of I. arboricola (larva and nymph) feeding on great tits. In contrast, I. frontalis larvae detached during the day, especially when feeding on great tits, which resembles that of I. ricinus’ feeding behaviour (larva and nymph). Ixodes frontalis left the host within seven days, immediately after completion of the blood meal. This is similar to both developmental stages of I. ricinus but contrasts with the very long (up to 20 days) feeding duration in I. arboricola. Thus I. frontalis shows strong plasticity, switching from dispersal-centered (larvae) to host-centered (nymphs) detachment behaviour. Findings are discussed with regard to the ticks’ habitat use, dispersal, life history and host specificity.

Predictors of individual performance and evolutionary potential of life-history traits in a hematophagous ectoparasite.

Little is known about the intraspecific variation of parasite life‐history traits and how this variation may affect parasite fitness and evolution. We investigated how life‐history traits predict success of individual tree‐hole ticks Ixodes arboricola and estimated their evolutionary potential, as well as genetic correlations within stages and phenotypic correlations within and across stages. Ticks were followed individually over two generations while allowed to feed on great tits Parus major. After accounting for host and tick maternal effects, we found that short feeding times and high engorgement weights strongly increased molting success. Molting time was also positively correlated with feeding success in adults. In larvae and nymphs, we found negative phenotypic correlations between engorgement weight and both feeding and molting time, the latter supported by a negative genetic correlation. We found sex‐related differences in feeding time (longer in male nymphs) and molting time (longer in male larvae but shorter in male nymphs). Also, time since the last feeding event (set experimentally) reduced larval and nymphal fitness, whereas it increased adult female fitness. Furthermore, we found significant heritability and evolvability, that is, the potential to respond to selection, for engorgement weight and molting time across all stages but no significant heritability for feeding time. Our findings suggest that variation in tick fitness is shaped by consistent individual differences in tick quality, for which engorgement weight is a good proxy, rather than by life‐history trade‐offs.

Rickettsia spp. in ticks (Acari: Ixodidae) from wild birds: First detection of Candidatus Rickettsia vini in Hesse, Germany

Tick-borne diseases are recognized as a growing public health concern and cause significant issues in humans and animals by serving as dispersal agents of ticks and their associated pathogens. Birds contribute to the spread of ticks and tick-borne diseases, with migratory birds playing a role in regional dispersal. Only a few studies have been carried out in Germany so far to investigate tick fauna on birds and the pathogens they harbour. In 2017 and 2018, we collected ticks from a total of 358 birds in the northern region of the Federal State of Hesse, Germany. Most ticks were Ixodes ricinus larvae and nymphs (95.4%). However, two ornithophilic species were also collected: Ixodes frontalis (larvae, nymphs and females) and Ixodes arboricola (larvae and nymphs). Ticks were tested individually or in pools, depending on the tick life stage and species for each bird host, in order to detect tick-borne encephalitis (TBE) virus and Rickettsia spp. All samples tested negative for TBE virus. Of 684 investigated tick pools, 162 tested positive for Rickettsia spp. by screening PCR. Of these, 117 carried Rickettsia helvetica detected by specific real-time PCR. The samples which were negative in the R. helvetica PCR were further investigated using five targets (gltA, 16S, ompA IV, ompB, 23S-5S intergenic spacer region) using multi locus sequence typing and phylogenetic analysis. The obtained sequences showed strong phylogenetic relationship to Candidatus Rickettsia vini, which is most closely related to R. japonica and R. heilongjiangensis, and occur in I. arboricola. Three more sequences were determined as R. helvetica in I. ricinus and I. frontalis. This is the first detection of Ca. R. vini in Germany and the first 23S-5S data published for Ca. R. vini, a useful target for Rickettsia species identification.

Ixodes arboricola

This datasheet on Ixodes arboricola covers Identity, Distribution, Hosts/Species Affected.

Heritable variation in host quality as measured through an ectoparasite's performance

Obligate parasites need one or more hosts to complete their life cycle. However, hosts might show intraspecific variation in quality with respect to the parasites themselves, thus affecting on‐host and off‐host parasite performance. High heritability in host quality for the parasite may therefore exert long‐lasting selective pressures on the parasite and influence host–parasite coevolution. However, the amount of variation and heritability in host quality are unknown for most parasite species, especially in wild populations of hosts. Both measures were estimated in a wild‐caught birdParus majorthat was experimentally infested by two developmental stages (larva and nymph) of a ectoparasite (the tickIxodes arboricola). We examined variation in host quality through variation in tick performance, namely the on‐host performance (attachment success, feeding time, engorgement weight and feeding success) and the off‐host performance (moulting time, moulting success and overall survival). Herein we also investigated the influence on tick performance of host traits linked with the bird's life history and physiology such as body condition, sex, age and haematocrit. By correlating tick performance variables between larvae and nymphs feeding on the same bird at different times, we found a significant correlation in attachment success, suggesting consistent among‐host variation for this performance measure, but no significant larva‐nymph correlations for the other tick variables. Animal models relating tick performance variables to the host pedigree showed a strong heritable signal for host quality as measured through tick feeding time, and lower but substantial estimates in other performance variables. With regard to the host traits, feeding success and survival of tick larvae were lower on female birds, and nymphal survival was higher on older birds. Larval feeding time was negatively correlated with host haematocrit. This is one of the first studies showing consistent intraspecific variation and heritability of host quality for a multi‐stage ectoparasite.

Ticks (Acari: Ixodidae) parasitizing migrating and local breeding birds in Finland

Ticks are globally renowned vectors for numerous zoonoses, and birds have been identified as important hosts for several species of hard ticks (Acari: Ixodidae) and tick-borne pathogens. Many European bird species overwinter in Africa and Western Asia, consequently migrating back to breeding grounds in Europe in the spring. During these spring migrations, birds may transport exotic tick species (and associated pathogens) to areas outside their typical distribution ranges. In Finland, very few studies have been conducted regarding ticks parasitizing migrating or local birds, and existing data are outdated, likely not reflecting the current situation. Consequently, in 2018, we asked volunteer bird ringers to collect ticks from migrating and local birds, to update current knowledge on ticks found parasitizing birds in Finland. In total 430 ticks were collected from 193 birds belonging to 32 species, caught for ringing between 2018 and 2020. Furthermore, four Ixodes uriae were collected from two roosting islets of sea birds in 2016 and 2020. Ticks collected on birds consisted of: Ixodes ricinus (n = 421), Ixodes arboricola (4), Ixodes lividus (2) and Hyalomma marginatum (3). Ixodes ricinus loads (nymphs and larvae) were highest on thrushes (Passeriformes: Turdidae) and European robins (Erithacus rubecula). The only clearly imported exotic tick species was H. marginatum. This study forms the second report of both I. uriae and I. arboricola from Finland, and possibly the northernmost observation of I. arboricola from Europe. The importation of exotic tick species by migrating birds seems a rare occurrence, as over 97% of all ticks collected from birds arriving in Finland during their spring migrations were I. ricinus, a species native to and abundant in Finland.

Ineffective humoral anti-tick IgY-response in birds: reaction against pathogen constituents?

Background: Variation in parasite burdens among hosts is typically related to differences in adaptive immunity. Comprehension of underlying mechanisms is hence necessary to gain better insights into endemic transmission cycles. Here we investigate whether wild songbirds that have never been exposed to ticks develop adaptive humoral immunity against endemic Ixodes ricinus ticks. Methods: Blue tits were exposed three times in succession to wild Ixodes ricinus ticks. For each infestation, serum samples were obtained. An enzyme-linked immunosorbent assay was developed, using tick salivary antigens, in order to quantify the bird's IgY response against ticks. In addition, at every sampling occasion the birds' body weight (corrected for body size) and haematocrit level was determined. Results: Individual IgY levels against the ticks' salivary proteins increased over three consecutive tick infestations, and large among-individual variation was observed. The responses were specifically directed against I. ricinus; cross-reactivity against the congeneric tree-hole tick Ixodes arboricola was negligibly low. IgY responses did not impinge on tick feeding success (engorgement weight and attachment success). Yet, those birds with the highest immune responses were more capable to reduce the acute harm (blood depletions) by compensating erythrocyte loss. Furthermore, at the end of the experiment, these birds had gained more body weight than birds with lower IgY levels. Conclusions: Latter observations can be considered as an effect of host quality and/or tolerance mechanisms. Birds anticipate the (future) costs of the activation of the immune system by ticks and/or ongoing tick-borne pathogen infections. Furthermore, although unsuccessful against tick feeding, the IgY responses may indirectly protect birds against tick-borne disease by acting against salivary protein secretions on which pathogens rely for transmission.

Pathogenic microorganisms in ticks removed from Slovakian residents over the years 2008–2018

A total of 750 ticks feeding on humans were collected during the years 2008–2018. The majority of ticks (94.8 %) came from Slovakia, with 3.5 % from the Czech Republic, 0.9 % from Austria, and 0.3 % from Hungary. Travellers from Ukraine, Croatia, France, and Cuba also brought one tick from each of these countries. The majority of the analysed ticks were identified as Ixodes ricinus (94.3 %). Dermacentor reticulatus (0.93 %), Haemaphysalis concinna (0.1 %), Haemaphysalis sp. (0.1 %), Ixodes arboricola (0.1 %), and Rhipicephalus sp. (0.1 %) were also encountered. The most frequently found stage of I. ricinus was the nymph (69.9 %) followed by adult females (20.4 %) and larvae (8.3 %). Ticks were predominantly found on children younger than 10 years (46.3 %) and adults between 30–39 years (21.4 %). In children younger than 10 years, the ticks were usually found on the head, while in other age categories, the ticks were predominantly attached to legs. Ticks were further individually analysed for the presence of Rickettsia spp., Coxiella burnetii, Borrelia burgdorferi sensu lato, Anaplasma phagocytophilum, Neoehrlichia mikurensis, Bartonella spp. and Babesia spp. The overall prevalences of tick-borne bacteria assessed in I. ricinus ticks acquired in Slovakia were: rickettsiae 25.0 % (95 % CI: 21.7–28.2), B. burgdorferi s.l. 20.5 % (95 % CI: 17.4–23.5), A. phagocytophilum 13.5 % (95 % CI: 10.9–16.0), Babesia spp. 5.2 % (95 % CI: 3.5–6.9), C. burnetii 3.0 % (95 % CI: 1.5–4.6), and N. mikurensis 4.4 % (95 % CI: 2.0–6.8). Pathogenic species Rickettsia raoultii, Rickettsia helvetica, Rickettsia monacensis, A. phagocytophilum, Borrelia garinii, Borrelia afzelii, Borrelia valaisiana, Babesia microti, and Babesia divergens were identified in D. reticulatus and I. ricinus ticks.

Isolation of Candidatus Rickettsia vini from Belgian Ixodes arboricola ticks and propagation in tick cell lines

Candidatus Rickettsia vini was originally detected in Ixodes arboricola ticks from Spain, and subsequently reported from several other Western Palearctic countries including Belgium. Recently, the bacterium was isolated in mammalian (Vero) cell culture from macerated male I. arboricola from Czech Republic, but there have been no reports of propagation in tick cells. Here we report isolation in a tick cell line of three strains of Ca. R. vini from I. arboricola collected from nests of great tits (Parus major) in Belgium. Internal organs of one male and two engorged female ticks were dissected aseptically, added to cultures of the Rhipicephalus microplus cell line BME/CTVM23 and incubated at 28 °C. Rickettsia-like bacteria were first seen in Giemsa-stained cytocentrifuge smears between 2 and 15 weeks later. Two of the isolates grew rapidly, destroying the tick cells within 2–4 weeks of onward passage in BME/CTVM23 cells, while the third isolate grew much more slowly, only requiring subculture at 4−5-month intervals. PCR amplification of bacterial 16S rRNA and Rickettsia gltA, sca4, ompB, ompA and 17-kDa genes revealed that all three isolates were Ca. R. vini, with 100 % identity to each other and to published Ca. R. vini sequences from other geographical locations. Transmission electron microscopy revealed typical single Rickettsia bacteria in the cytoplasm of BME/CTVM23 cells. The Ca. R. vini strain isolated from the male I. arboricola tick, designated Boshoek1, was tested for ability to grow in a panel of Ixodes ricinus, Ixodes scapularis and R. microplus cell lines and in Vero cells. The Boshoek1 strain grew rapidly, causing severe cytopathic effect, in the R. microplus line BME26, the I. ricinus line IRE11 and Vero cells, more slowly in the I. ricinus line IRE/CTVM19, possibly established a low-level infection in the I. ricinus line IRE/CTVM20, and failed to infect cells of any of four I. scapularis lines over a 12-week observation period. This study confirmed the applicability of the simple tick organ-cell line co-cultivation technique for isolation of tick-borne Rickettsia spp. using BME/CTVM23 cells.

Experimental study of micro-habitat selection by ixodid ticks feeding on avian hosts

Mechanisms of on-host habitat selection of parasites are important to the understanding of host-parasite interactions and evolution. To this end, it is important to separate the factors driving parasite micro-habitat selection from those resulting from host anti-parasite behaviour. We experimentally investigated whether tick infestation patterns on songbirds are the result of an active choice by the ticks themselves, or the outcome of songbird grooming behaviour. Attachment patterns of three ixodid tick species with different ecologies and host specificities were studied on avian hosts. Ixodes arboricola, Ixodes ricinus and Ixodes frontalis were put on the head, belly and back of adult great tits (Parus major) and adult domestic canaries (Serinus canaria domestica) which were either restricted or not in their grooming capabilities. Without exception, ticks were eventually found on a bird’s head. When we gave ticks full opportunities to attach on other body parts – in the absence of host grooming – they showed lower attachment success. Moreover, ticks moved from these other body parts to the host's head when given the opportunity. This study provides evidence that the commonly observed pattern of ticks feeding on songbirds’ heads is the result of an adaptive behavioural strategy. Experimental data on a novel host species, the domestic canary, and a consistent number of published field observations, strongly support this hypothesis. We address some proximate and ultimate causes that may explain parasite preference for this body part in songbirds. The link found between parasite micro-habitat preference and host anti-parasite behaviour provides further insight into the mechanisms driving ectoparasite aggregation, which is important for the population dynamics of hosts, ectoparasites and the micro-pathogens for which they are vectors.

Ticks (Acari: Ixodida) on birds (Aves) migrating through the Polish Baltic coast

Seasonal bird (Aves) migration between breeding and wintering areas, often located on different continents, can facilitate the spreading of tick species (Acari: Ixodida) and of tick-borne pathogens. The aim of the study was to analyse the occurrence of ticks dispersed by birds migrating along the Polish Baltic coast during spring and autumn migration. Field research was conducted at the bird ringing station in Wicie, located on the middle of the Polish Baltic coast, in 2011 and 2012 during spring and autumn migration. A total of 2657 birds from 45 species was examined. The most common species inspected were European robin (Erithacus rubecula) (63.3%), song thrush (Turdus philomelos) (5.13%), and goldcrest (Regulus regulus) (4.5%). Overall, 3129 ticks belonging to six species were collected: Ixodes ricinus (1650 larvae, 1390 nymphs and 1 male), Ixodes frontalis (20 larvae, 20 nymphs), Ixodes arboricola (35 larvae), Dermacentor reticulatus (1 larva), and Haemaphysalis punctata (1 nymph). Ten larvae and one nymph could only be identified to the genus level Ixodes. Ticks were located on various parts of the head: on the corner of the beak (75.0%), near the eyes (14.6%), on the chin (4.4%), near the ears (4.4%), on the neck (1.1%), and in the beak (0.5%). The overall tick prevalence was 40.5%. The highest prevalence was for bird species feeding on the ground, covering a medium distance to wintering grounds and migrating at night. Statistically significant differences between the number of ticks and the sex of the host species were detected in blackbirds: males carried more parasites than females, both, during spring and autumn migration. The fact that I. ricinus and other ticks parasitize birds migrating through Poland extends the possibility of the spread of tick-borne diseases.

Isolated populations of Ixodes lividus ticks in the Czech Republic and Belgium host genetically homogeneous Rickettsia vini

In the last two decades, the advent of molecular methods has revealed a remarkable diversity of rickettsiae (Rickettsiales: Rickettsiaceae) in invertebrates. Several species of these obligate intracellular bacteria are known to cause human infections, hence more attention has been directed towards human-biting ectoparasites. A spotted fever group Rickettsia sp. was previously detected in Ixodes lividus ticks (Ixodidae) associated with sand martins (Hirundinidae: Riparia riparia). In order to identify whether this rickettsia varies among isolated tick populations, a total of 1758 I. lividus ticks and five Ixodes ricinus ticks (Ixodidae) were collected in the Czech Republic and 148 I. lividus ticks were collected in Belgium, from nests of sand martins, European bee-eaters (Meropidae: Merops apiaster), Eurasian tree sparrows (Passeridae: Passer montanus), and from captured sand martins. We screened 165 and 78 I. lividus ticks (from the Czech Republic and Belgium, respectively) and all five I. ricinus ticks for the presence of rickettsial DNA. Only I. lividus samples were positive for Rickettsia vini, a spotted fever group rickettsia that commonly infects the tree-hole tick Ixodes arboricola (Ixodidae). Maximum likelihood analysis of the rickettsial sequences showed that the most closely related organism to R. vini corresponds to an uncharacterized rickettsia detected in Argas lagenoplastis (Argasidae), a nidicolous soft tick of the fairy martin (Hirundinidae: Petrochelidon ariel) in Australia. The observed variability of R. vini sequences from isolated tick populations was low; all 85 sequenced samples were identical to each other in five out of six partial rickettsial genes, except for the sca4 sequence (99.9% identity, 808/809 nt) that differed in I. lividus ticks from two sampling sites in the Czech Republic.

Sex ratios of the tick Ixodes arboricola are strongly female-biased, but there are no indications of sex-distorting bacteria

Studies on sex ratio are of fundamental importance for understanding the biology of populations and biological control of pests and pathogens. In most Ixodes tick species, only females feed in the adult stage and, hence, contribute to pathogen transmission. The tree-hole tick Ixodes arboricola infests cavity-nesting birds and has limited dispersal possibilities. It plays an important role in the maintenance of zoonotic disease cycles. Here, we quantified the sex ratio of 718 adult I. arboricola ticks obtained from a laboratory stock at nine distinct periods (cohorts) from 2008 to 2015. In addition, we screened 93 specimens, collected from four study sites in 2011 and 2012, for the presence of six maternally inherited bacterial parasites known to manipulate arthropod sex ratios. We found significantly female-biased sex ratios in seven out of nine cohorts. There were no infections with members of the Wolbachia, Arsenophonus or Cardinium bacterial genera, whereas 96.8% of the screened ticks were infected with Rickettsia vini, 22.6% with Rickettsiella sp., and 14.0% with Spiroplasma ixodetis. Male and female I. arboricola were found equally infected. Our results suggest skewed sex ratios in I. arboricola are not caused by these bacterial infections, although there may be other, untested candidates driving sex ratios. Alternatively, female-biased sex ratios may be an adaptation in females to high local densities and low dispersal, where the production of daughters has a selective advantage because a few sons can fertilise all daughters.

Genera, subgenera, species and subspecies of hard ticks (Acari: Ixodidae) described, named, renamed or given new rank by Paul Schulze (1887–1949) and their current status

The names for 19 genera, 17 subgenera, 150 species and 150 subspecies of Ixodidae described by Paul Schulze, names changed by him after the revision of certain taxa, new ranks for names originally described by other authors, a few lapsus calamorum and some incorrect names ascribed to Schulze by other workers are discussed. Two genera (Cosmiomma and Nosomma) are valid, but most of Schulze’s subgenera are difficult to assess because of the current disarray that prevails at this classificatory level in the Ixodidae. Thirty-six of 150 species names discussed by Schulze are considered valid herein (3 species of Amblyomma, 1 Bothriocroton, 1 Cosmiomma, 6 Dermacentor, 4 Haemaphysalis, 5 Hyalomma, 13 Ixodes, 1 Nosomma and 2 Rhipicephalus), while 28 are valid species epithets that are currently assigned to different genera (e.g., Acarus undatus was formerly classified in the genus Aponomma but is currently known as Bothriocroton undatum). Twenty-nine of these valid species names were authored by Schulze or Schulze & Schlottke. The remaining species names are mostly synonyms or names that are incertae sedis, nomen nudum or lapsus calamorum. The 150 subspecific names are treated as synonyms of valid species or as nomina nuda, nomina dubia, incertae sedis or lapsus calamorum. Many names used by Schulze have been synonymized based on sound systematic research, but the justification for synonymizing other names is occasionally weak. It will be important to take these names into account if, as has already happened, future molecular and morphological studies result in the reinstatement of Schulze names currently considered synonyms. This situation has particular relevance for Amblyomma geoemydae, the Amblyomma marmoreum species complex, Amblyomma testudinarium, Oriental species of Dermacentor, Hyalomma in general, Ixodes arboricola, Ixodes canisuga, Ixodes lividus, Ixodes trianguliceps and Ixodes uriae. In conclusion, the prolific contributions of Paul Schulze have their weak points —the many species and subspecies of Hyalomma described by him are unjustified as is the alleged worldwide distribution of the former genus Aponomma—but his perception of the complexity of the genus Dermacentor (under Indocentor) in the Oriental and Australasian Zoogeographic Regions, as well as his understanding of tick biodiversity in these regions constitute seminal scientific achievements.

Seasonal infestation of birds with immature stages of Ixodes ricinus and Ixodes arboricola

This study assessed the parasitization of cavity-nesting birds and ground-nesting/foraging birds with larvae and nymphs of two Ixodes species, Ixodes ricinus and Ixodes arboricola.Totals of 679 (52.3%) I. ricinus and 619 (47.7%) I. arboricola ticks were collected from 15 species of passerine birds which were caught during the nesting and non-nesting periods of 2003–2006, in the south-eastern part of the Czech Republic, the Drahanská Vrchovina Uplands. In the non-nesting period from October to March, 6.8% (101/1492) of birds were infested with ticks, mainly with I. arboricola larvae. In the non-nesting period, the average intensity of infestation by I. arboricola and I. ricinus was 8.5 and 1.5 individuals per infested bird, respectively. In the nesting period from April to June, 21.6% (50/232) of birds were infested by both tick species but mainly with I. ricinus nymphs. The average intensity of infestation by I. ricinus and I. arboricola was 13.3 and 10.8 individuals per infested bird, respectively. Altogether, 23.2% of the infested birds were parasitized by both immature life stages of one or both tick species. From an enzootic perspective, co-feeding and co-infestation of I. ricinus and I. arboricola subadults on passerine birds might happen and may be important for the dissemination of tick-borne agents.

Ixodes kangdingensis (Acari: Ixodidae), a new species from the Siberian weasel, Mustela sibirica (Carnivora: Mustelidae) in China

SUMMARY The female and nymph of Ixodes (Pholeoixodes) kangdingensis n. sp. are described based on both morphology and analysis of 16S rRNA gene sequences. Specimens of this new tick species were collected from a Siberian weasel (Mustela sibirica) in Kangding County, Sichuan Province, China. The morphological features of the female and nymph are unique to distinguish I. kangdingensis n. sp. from other members of the subgenus Pholeoixodes, including the presence of distinctly shaped cornua, anterior and posterior processes on palpal article I and a large angular projection on each side of the hypostome. Partial sequence of 16S rRNA gene grouped this species with Ixodes arboricola and Ixodes lividus with sequence divergence of new species from I. arboricola 4·16% and from I. lividus 8·49%. Data on the phylogenetic position, hosts, geographic distribution and key to females of closely related species are also provided.

Urban Breeding Corvids as Disseminators of Ticks and Emerging Tick-Borne Pathogens.

Crows (Corvidae) are common city dwellers worldwide and are increasingly important subjects of epidemiology studies. Although their importance as hosts and transmitters of a number of zoonotic parasites and pathogens is well known, there are no studies on their importance as tick hosts. After mosquitoes, ticks are the most important vectors of zoonotic pathogens, especially for those causing emerging zoonotic diseases. Pathogenic bacteria, especially Borrelia spp., Rickettsia spp., and Anaplasma spp., vectored by ticks, are the cause for most vector-borne diseases in Europe. Here we report on ticks and tick-borne pathogens harbored by urban breeding crows. A total of 36 birds (33.33%, n = 108) hosted ticks, with 91 individual ticks belonging to 6 species (Haemaphysalis concinna, Haemaphysalis parva, Haemaphysalis punctata, Hyalomma marginatum, Ixodes arboricola, and Ixodes ricinus). Rickettsia spp. DNA was found in 6.6% of ticks and 1.9% of bird tissues, whereas Anaplasma phagocytophilum was found in 5.9% of ticks and 0.9% of birds. Two rickettsial genospecies were located, Rickettsia helvetica and Rickettsia monacensis. This is the first study to determine such a diverse tick spectrum feeding on urban corvids, while highlighting their importance as tick hosts and raising concerns about their potential risk to human health.