The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodida): A World List of Valid Tick Names

The world's argasid tick fauna comprises 183 species in four genera, namely Argas, Carios, Ornithodoros and Otobius in the family Argasidae. The ixodid tick fauna consists of 241 species in the genus Ixodes and 442 species in the genera Amblyomma, Anomalohimalaya, Bothriocroton, Cosmiomma, Dermacentor, Haemaphysalis, Hyalomma, Margaropus, Nosomma, Rhipicentor and Rhipicephalus in the family Ixodidae, with the genus Boophilus becoming a subgenus of the genus Rhipicephalus. The family Nuttalliellidae is represented by the monospecific genus Nuttalliella. The species names of these ticks, based on seven previous complete or partial listings, as well as those of recently described new species, are presented in tabular format. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ticks on humans in Ankara,Turkey

In this study, a total of 5,094 ticks found on humans were examined in terms of species, development stage, gender, host features and seasonality for a year period. Of these ticks 17 were argasid and 5,077 were ixodid. Predominantly species of the ixodid genera Hyalomma, Dermacentor, Rhipicephalus and Haemaphysalis were found on humans in Ankara (Anatolia). Most abundant were Hyalomma nymphs (29.8%) and adults (28.2%). Primary factors in terms of tick bite risk were region, habitat and season.     

Ticks (Ixodidae) on humans in South America

Twenty eight species of Ixodidae have been found on man in South America (21 Amblyomma, 1 Boophilus, 2 Dermacentor, 2 Haemaphysalis, 1 Ixodes and 1 Rhipicephalus species). Most of them are rarely found on man. However, three species frequently parasitize humans in restricted areas of Argentina (A. neumanni reported from 46 localities), Uruguay (A. triste from 21 sites) and Argentina–Brazil (A. parvum from 27 localities). The most widespread ticks are A. cajennense (134 localities in Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Suriname and Venezuela), A. ovale (37 localities in Argentina, Brazil, Ecuador, French Guiana, Guyana, Paraguay, Suriname and Venezuela) and A. oblongoguttatum (28 sites in Brazil, Colombia, French Guiana, Guyana, Suriname and Venezuela). Amblyomma aureolatum (18 localities in Argentina, Brazil, French Guiana and Paraguay), A. cajennense, and A. triste are vectors of rickettsioses to man in South America. A better understanding of the respective roles of these and other tick species in transmitting pathogens to humans will require further local investigations. Amblyomma ticks should be the main subjects of these studies followed by species of Boophilus, Dermacentor, Haemaphysalis and Rhipicephalus species. In contrast with North America, Europe and Asia, ticks of the genus Ixodes do not appear to be major players in transmitting diseases to human. Indeed, there is only one record of an Ixodes collected while feeding on man for all South America.     

Ticks (Acari: Ixodoidea: Argasidae,Ixodidae) of China

This paper presents results of an investigation and listing of tick species found in China during a survey in all 28 provinces. This will be a step towards a definitive list of tick species and their distribution. To date, the tick fauna of this area consists of 117 species in the following families: Argasidae-Argas (7 species), Carios (4 species) and Ornithodoros (2 species); Ixodidae-Amblyomma (8 species), Anomalohimalaya (2 species), Dermacentor (12 species), Haemaphysalis (44 species), Hyalomma (6 species), Ixodes (24 species) and Rhipicephalus (8 species). Some well known ticks carrying and transmitting many infectious agents to man and domestic animals are also found in China. These include Ixodes persulcatus, Haemaphysalis longicornis, Rhipicephalus sanguineus, R. (Boophilus) microplus and Hyalomma asiaticum. It is worth mentioning that Ixodes rangtangensis Teng and Haemaphysalis xinjiangensis Teng should be relegated to a synonym of I. moschiferi and Hae. danieli, respectively. The distribution of ticks over the provinces of China is also discussed. The information on ticks in some areas such as Henan is not exhaustive.     

Preferences of different tick species for human hosts in Turkey

The ticks removed from the patients who applied to the hospitals in Istanbul and neighboring cities, Turkey, with the complaint of tick bite were examined in this study, on account of their species, biological stages, attachment sites on the body, and the age of the affected patients. A total of 16,969 ticks were identified. Encountered species were as follows: 33.6 % Ixodes spp. immature, 25.3 % Hyalomma spp. immature, 24.3 % I. ricinus, 9.5 % Rhipicephalus sanguineus gr., 3.2 % R. bursa, 2.2 % Hyalomma marginatum, 1.96 % Haemaphysalis adults, 1.66 % Hyalomma aegyptium, 0.52 % Dermacentor marginatus, 0.39 % Rhipicephalus spp. nymphs, 0.12 % Dermacentor spp. nymphs, 0.11 % Haemaphysalis spp. nymphs, 0.09 % Hyalomma scupense, and 0.03 % Hyalomma excavatum. The distribution of attachment sites of the species and instars showed significant differences. Furthermore, age data of the patients also revealed that certain tick species were more common within certain age groups.     

Birds as hosts of ixodid ticks (Acarina,Ixodidae)

rophic relations between ixodid ticks of the world fauna parasitizing as adults on birds are analyzed. These ticks include 60 out of 650 species of the family Ixodidae and belong to the genera Ixodes, Haemaphysalis, and Amblyomma. Relations between birds and ticks of the genus Ixodes are most ancient. Colonial marine birds seem to be the first hosts of ixodid ticks in the class of birds. Trophic specialization of ticks is not associated with bird taxa, because in most cases no tick specificity is caused by either phylogenetic features of hosts, or specific environmental conditions.     

Dermal gland secretion improves the heat tolerance of the brown dog tick, Rhipicephalus sanguineus, allowing for their prolonged exposure to host body temperature

Dermal glands (sensilla sagittiformia) secrete when brown dog ticks, Rhipicephalus sanguineus, are mechanically disturbed, presumably as a defensive mechanism. Recently, we observed that these glands secrete due to the pressure stimulation of engorgement. In this study, we examine how dermal gland secretion alters the physiology of R. sanguineus, particularly if this secretion is an important mechanism during blood feeding. The ability of ticks to retain water was not modified by dermal gland secretion, but heat tolerance was enhanced. Short-term heat shock was improved slightly (1 h at 50 °C to 1 h at 56 °C) and featured reduced injury responses and greater recovery after heat shock. When exposed to their host body temperature (37 °C) for prolonged periods, individuals that had secreted survived over 1 week longer than individuals that did not secrete. Dorsal application of squalene, the main component of dermal gland secretion, did not increase temperature tolerance, suggesting that the act of secreting rather than the physical properties of the secretion itself is responsible for the increase in heat tolerance. Based on our results, dermal gland secretion may be an essential mechanism in certain tick species (Amblyomma, Dermacentor, Hyalomma, Rhipicephalus, but not Ixodes) for tolerating body temperature and not succumbing to heat stress during the extended time periods of feeding on a mammalian host, serving as a mechanism to prevent heat damage from the host during feeding.     

Preliminary Observations on Specific Adaptations of Exophilic Ixodid Ticks to Forests or Open Country Habitats

Some features of three-host exophilic ticks important for their survival differ at the generic level in relation to the types of their habitat. Ixodes and Haemaphysalis are typical forest genera, Hyalomma is an open country genus, whereas Dermacentor and Rhipicephalus, having some features of forest ticks, are closer to the ticks of open country. Forest ticks encounter rather stable and favorable temperature, humidity and illumination conditions as compared with open country ticks. A few differences determining every-day survival, host-seeking, and reproduction of ticks are considered. (1) Tolerance to desiccation is very low in forest ticks and much greater in ticks of open country. (2) Lack of eyes in forest ticks (having, however, epithelial photoreceptor cells) and formation of eyes as special morphological structures in ticks of open country. (3) Capability of mating and insemination of unfed specimens both on and off hosts in forest ticks (Ixodes) and insemination of partially engorged females only on hosts by fed males in open country ticks. (4) Reciprocal sexual dimorphism where in forest ticks females have a larger body than males and in open country ticks this correlation is opposite. (5) Linear dependence between female weight and number of eggs laid is established in forest ticks after the female weight becomes higher than 50% of mean engorgement weight whereas in open country ticks linear dependence begins to reveal itself much earlier. (6) In forest ticks the compensatory growth occurs only during adult feeding whereas in open country ticks such a growth occurs both during nymphal and adult feeding. The adaptive nature of the above differences and their evolutionary trends have been hypothesized. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ticks (Acari: Ixodida) infesting humans in the provinces of Kelkit Valley, a Crimean-congo hemorrhagic fever endemic region in Turkey

Ticks are mandatory blood feeding ectoparasites leading transmission of various tick-borne pathogens to human and animals. Since 2002, thousands of human tick bites and numerous Crimean-Congo Hemorrhagic Fever cases have been reported in several provinces in the Kelkit Valley region in Turkey. Despite increased cases of tick bites and tick-borne diseases, no taxonomic information is available about the tick species infesting humans in the region. In the present study, a tick survey on humans was performed to determine the species composition of ticks infesting humans in several provinces of Kelkit Valley. In the survey, 1,460 ticks (721 males, 516 females and 223 nymphs) were collected from tick-infested humans. A total of 19 tick species have been found on humans in the region, including 7 Hyalomma, 2 Argas, 2 Haemaphysalis, 2 Ixodes, Dermacentor and 3 Rhipicephalus species. Infestation of Dermacentor reticulatus on humans was documented for the first time in Turkey.     

Record of five new endemic small mammal hosts for four ectoparasite species from Sri Lanka

Both ticks and fleas are medically important arthropods, which act as vectors for a number of zoonotic diseases. In Sri Lanka, 35 species of ticks and 20 species of fleas have been reported from mammals, birds and reptiles. However, records of these parasites from small mammals are scanty. Here, we report ticks and fleas infesting muroid rodents and shrews including endemic species collected from around the country. Of the 149 small mammals collected, 48 individuals were infested with ticks, fleas or both. Immature stages of three tick species, Ixodes ceylonensis, Haemaphysalis hystricis and Hyalomma brevipunctata and unidentified species of Rhipicephalus were recorded from ten small mammal species. Four species of fleas, Stivalius phoberus, Stivalius aporus, Lentistivalius ferinus and Nosopsyllus tamilanus were identified from five species of small mammals. We report new host species for I. ceylonensis (Crocidura miya, Suncus montanus and Solisorex pearsoni), S. aporus (S. montanus and Mus fernandoni), N. tamilanus (S. pearsoni) and Rhipicephalus spp. (Crocidura hikmiya, S. montanus, S. pearsoni, M. fernandoni). All new host species reported here are endangered or critically endangered endemics to Sri Lanka.     

Hard ticks (Acari,Ixodidae) of Croatia

The present paper is based on original and literature data. In Croatia the first studies on the occurrence of ixodid species were made about 80 years ago. The number of tick species recorded in Croatia considerably increased during the 1950s, 60s, 70s and 80s of the past century. A total of 21 species of hard tick belonging to 5 genera have been recorded in Croatia. Ixodes is the best represented genus, with seven species recorded. Haemaphysalis is represented by six species, followed by Rhipicephalus with four species. Dermacentor and Hyalomma are represented by two species each. The ticks were collected on 47 different host species. Eleven tick species were collected on Bos taurus and Ovis aries, followed by Capra hircus and Equus caballus with 8 species and Canis lupus familiaris with 6 species. On the remaining 42 host species one, two or three tick species were collected. The most widespread tick is Ixodes ricinus which was found on 25 different host species.     

Maps of ticks (Acari: Argasidae,Ixodidae) for Austria and South Tyrol,Italy

A first compilation of georeferenced tick locations in Austria and South Tyrol, Italy, is presented here. This allows the tick fauna to be examined in the various climatic regions of the European Alps. The dataset comprises 424 tick locations of Austria and 48 tick locations of South Tyrol, which were digitized from literature and visualized in the form of geographical maps. The tick fauna of Austria includes two species of Argasidae in the genera Argas and Carios and 15 species of Ixodidae in the genera Dermacentor, Haemaphysalis, and Ixodes, altogether 17 tick species. In addition, two species of Ixodidae in the genera Hyalomma (each spring imported by migratory birds) and Rhipicephalus (occasionally imported by dogs returning from abroad with their owners) are included in the tick atlas. Of these, the georeferenced locations of 18 tick species are depicted in maps. The occurrence of the one remaining tick species, Ixodes inopinatus, is given at the level of the federal states. The first Austrian distribution map of the long-legged bat tick Ixodes vespertilionis, which was reported from 21 caves, deserves special mention. The most common and widespread tick species is Ixodes ricinus, with records in all nine federal states of Austria, followed by Ixodes canisuga, Ixodes hexagonus, and I. vespertilionis in six federal states each. Haemaphysalis concinna and Dermacentor reticulatus are only endemic in the eastern plains, while Dermacentor marginatus only occurs in the west, in the Tyrolean Alpine valleys. Eight tick species were reported from South Tyrol, Italy. There, the most frequently flagged tick from the vegetation is also I. ricinus, while D. marginatus and Haemaphysalis punctata are often collected from sheep. The locations are shown together with those from North and East Tyrol on a separate Tyrol map. The tick atlas in Austria and South Tyrol as well as the underlying digital dataset in the supplement contribute to the closing of data gaps in global distribution maps of ticks and improve the data basis for new species distribution models.

     

Inferring host specificity and network formation through agent-based models: tick–mammal interactions in Borneo

Patterns of host–parasite association are poorly understood in tropical forests. While we typically observe only snapshots of the diverse assemblages and interactions under variable conditions, there is a desire to make inferences about prevalence and host-specificity patterns. We studied the interaction of ticks with non-volant small mammals in forests of Borneo. We inferred the probability of species interactions from individual-level data in a multi-level Bayesian model that incorporated environmental covariates and advanced estimates for rarely observed species through model averaging. We estimated the likelihood of observing particular interaction frequencies under field conditions and a scenario of exhaustive sampling and examined the consequences for inferring host specificity. We recorded a total of 13 different tick species belonging to the five genera Amblyomma, Dermacentor, Haemaphysalis, Ixodes, and Rhipicephalus from a total of 37 different host species (Rodentia, Scandentia, Carnivora, Soricidae) on 237 out of 1,444 host individuals. Infestation probabilities revealed most variation across host species but less variation across tick species with three common rat and two tree shrew species being most heavily infested. Host species identity explained ca. 75 % of the variation in infestation probability and another 8–10 % was explained by local host abundance. Host traits and site-specific attributes had little explanatory power. Host specificity was estimated to be similarly low for all tick species, which were all likely to infest 34–37 host species if exhaustively sampled. By taking into consideration the hierarchical organization of individual interactions that may take place under variable conditions and that shape host–parasite networks, we can discern uncertainty and sampling bias from true interaction frequencies, whereas network attributes derived from observed values may lead to highly misleading results. Multi-level approaches may help to move this field towards inferential approaches for understanding mechanisms that shape the strength and dynamics in ecological networks.     

Validation and rapid extraction of nucleic acids from alcohol-preserved ticks

A method has been developed to extract DNA from alcohol-preserved ticks (Acari: Ixodidae and Argasidae). The method combines the lysing property of the chaotropic agent guanidinium thiocyanate (GuSCN) and the nucleic acid-binding property of diatomaceous earth (fossilized cell walls of unicellular algae). Debris from the tick is removed in several sequential washing steps. To monitor the efficiency of this method, a polymerase chain reaction (PCR) was designed to amplify the 16S mt rRNA gene of five tick genera (Dermacentor Fabricius, Haemaphysalis Koch, Rhipicephalus Koch, Argas Latreille and Ixodes Latreille). Detection of amplification products from this PCR indicated that DNA had been successfully extracted and that Taq-polymerase inhibitors were absent. The extraction method, therefore, enables purification of DNA such that enzymatic analysis is possible.     

蜱类系统学研究进展

本文介绍了近年来蜱的编目和分类及系统发育和演化等方面的研究进展。对软蜱科(Argasidae)说明了属的变动;硬蜱科(Ixodidae)介绍了璃眼蜱亚科(Hyalomminae)和凹沟蜱亚科(Bothriocrotoninae)以及相应变动的属(新建的凹沟蜱属Bothriocroton和须角蜱属Cornupalpatum,合并的牛蜱属Boophilus、暗眼蜱属Anocentor和盲花蜱属Aponomma)。根据新的分类变更对已知种类进行了分析。至2006年,世界已知蜱类有3科18属897种,中国有2科10属119种。阐明了有关蜱类系统发育研究的主要观点,并讨论了其不足和有待深入研究的问题。目前,把形态学和分子生物学数据结合在一起的全证据方法,并结合蜱类和不同宿主之间的关系、动物地理学、古生物学以及比较寄生虫学的资料,成为解决蜱类系统发育问题的有效途径。     

Identification of Tick-Borne Pathogens in Ticks Feeding on Humans in Turkey

Background

The importance of tick-borne diseases is increasing all over the world, including Turkey. The tick-borne disease outbreaks reported in recent years and the abundance of tick species and the existence of suitable habitats increase the importance of studies related to the epidemiology of ticks and tick-borne pathogens in Turkey. The aim of this study was to investigate the presence of and to determine the infection rates of some tick-borne pathogens, including Babesia spp., Borrelia burgdorferi sensu lato and spotted fever group rickettsiae in the ticks removed from humans in different parts of Ankara.

Methodology/Principal Findings

A total of 169 ticks belonging to the genus Haemaphysalis, Hyalomma, Ixodes and Rhipicephalus were collected by removing from humans in different parts of Ankara. Ticks were molecularly screened for Babesia spp., Borrelia burgdorferi sensu lato and spotted fever group rickettsiae by PCR and sequencing analysis. We detected 4 Babesia spp.; B. crassa, B. major, B. occultans and B. rossi, one Borrelia spp.; B. burgdorferi sensu stricto and 3 spotted fever group rickettsiae; R. aeschlimannii, R. slovaca and R. hoogstraalii in the tick specimens analyzed. This is the report showing the presence of B. rossi in a region that is out of Africa and in the host species Ha. parva. In addition, B. crassa, for which limited information is available on its distribution and vector species, and B. occultans, for which no conclusive information is available on its presence in Turkey, were identified in Ha. parva and H. marginatum, respectively. Two human pathogenic rickettsia species (R. aeschlimannii and R. slovaca) were detected with a high prevalence in ticks. Additionally, B. burgdorferi sensu stricto was detected in unusual tick species (H. marginatum, H. excavatum, Hyalomma spp. (nymph) and Ha. parva).

Conclusions/Significance

This study investigates both the distribution of several tick-borne pathogens affecting humans and animals, and the presence of new tick-borne pathogens in Turkey. More epidemiological studies are warranted for B. rossi, which is very pathogenic for dogs, because the presented results suggest that B. rossi might have a wide distribution in Turkey. Furthermore, we recommend that tick-borne pathogens, especially R. aeschlimannii, R. slovaca, and B. burgdorferi sensu stricto, should be taken into consideration in patients who had a tick bite in Turkey.     

New foci of <Emphasis Type="Italic">Rhipicephalus microplus</Emphasis> in West Africa

The invasive character of Rhipicephalus microplus was observed in Benin, the second West-African country from which this ticks species has been collected after the initial confirmed record in Ivory Coast in 2007. A cross-sectional study was carried out in the Department of Mono to examine the presence of the tick R. microplus. The survey covered 9 herds (villages) in an agro-ecological zone inhabited by agro-pastoralists, including the State Farm of Kpinnou that imported Girolando cattle from Brazil. Almost 800 ticks were sampled from 36 cattle, on average four cattle per village. The morphological identification revealed ticks of two different genera: Rhipicephalus and Amblyomma. Rhipicephalus microplus was the only representative of the species previously known as Boophilus or blue ticks. Its taxonomic identity was confirmed molecularly by PCR–RFLP. A comparison was made with the situation of R. microplus in Brazil.     

Ixodid Tick Infestation in Cattle and Wild Animals in Maswa and Iringa,Tanzania

Ticks and tick-borne diseases are important in human and livestock health worldwide. In November 2012, ixodid ticks were collected and identified morphologically from cattle and wild animals in the Maswa district and Iringa urban, Tanzania. Amblyomma gemma, A. lepidum, and A. variegatum were identified from Maswa cattle, and A. variegatum was the predominant species. A. marmoreum, Hyalomma impeltatum, and Rhipicephalus pulchellus were identified from Iringa cattle in addition to the above 3 Amblyomma species, and A. gemma was the most abundant species. Total 4 Amblyomma and 6 Rhipicephalus species were identified from wild animals of the 2 areas. A. lepidum was predominant in Maswa buffaloes, whereas A. gemma was predominant in Iringa buffaloes. Overall, A. variegatum in cattle was predominant in the Maswa district and A. gemma was predominant in Iringa, Tanzania.     

Ixodid tick salivary gland products target host wound healing growth factors

For successful blood-feeding, ticks must confront the host immune system comprising many cells and signaling molecules, mainly cytokines and growth factors. These factors bind to specific receptors on the cell membranes, thereby initiating a signaling cascade that leads to distinct cellular activities. Ticks are able to manipulate host immune responses via molecules secreted from their salivary glands. Saliva of ixodid ticks contains factors binding important cytokines and their subgroup, chemokines. Here we demonstrate that constituents of tick salivary gland extract (SGE) also appear to bind growth factors: transforming growth factor beta (TGF-β1), platelet-derived growth factor (PDGF), fibroblast growth factor (FGF-2), and hepatocyte growth factor (HGF), depending on tick species. SGE derived from Amblyommavariegatum reacted with TGF-β1, PDGF, FGF-2 and HGF; Dermacentorreticulatus and Rhipicephalusappendiculatus with TGF-β1, FGF-2 and HGF; and Ixodes ricinus and Ixodesscapularis with PDGF. SGE from the species targeting PDGF (A. variegatum and I. ricinus) also inhibited cell proliferation in vitro and induced a change in morphology of different cell lines. These effects correlated with disruption of the actin cytoskeleton. Such effects were not observed with SGE of the two species that did not target PDGF. Targeting of wound healing growth factors appears to be yet another strategy ixodid ticks adopt for suppression of inflammation and successful haematophagy.     

Ixodid ticks (Acari: Ixodidae) infesting humans in Tokat Province of Turkey: species diversity and seasonal activity

Ixodid ticks (Acari: Ixodidae) are the major vectors of pathogens threatening animal and human health. Tokat Province, Turkey, is a suitable habitat for extended tick activity with its moderate climate and vegetation. In the present study, we surveyed humans visiting health centers to determine the species diversity, geographical distribution, and seasonal activity of ixodid ticks infesting them. Out of 5,999 adult ticks collected from humans from April to September, 2008, 800 ticks were identified to species, while the remaining were identified to genus according to their distinct morphological characteristics. Hyalomma marginatum, Hyalomma detritum, Rhipicephalus bursa, Rhipicephalus (Boophilus) annulatus, Haemaphysalis punctata, Dermacentor marginatus, and Ixodes laguri were the most prevalent tick species among 24 ixodid tick species infesting humans in the region. One of these tick species, Hyalomma isacii was identified as a new record for the ixodid tick fauna of Turkey. Hyalomma species were the most abundant in summer, while Dermacentor and Ixodes species displayed the lowest frequency. Hyalomma aegyptium infestation was very common on humans in the province. Results indicated that a variety of ixodid tick species infest humans depending on the season in the target area. It is possible that a variety of ixodid tick species may contribute to the spread of tick‐borne diseases such as Crimean‐Congo haemorrhagic fever virus (CCHFV), which is endemic in the region.