Synopsis of the hard ticks (Acari: Ixodidae) of Romania with update on host associations and geographical distribution

The current paper is a synoptic review of the distribution and host associations of the 25 species of hard tick fauna (family Ixodidae) in Romania. In addition to a full literature survey, original data is presented, based on eight years of occasional or targeted sample collection. The literature data on geographical distribution was transposed digitally to the decimal degree coordinate system. For each species, an updated distribution map is given together with all historical data and new host associations. Overall, our paper records 58 new tick-host associations for Romania: 20 for Ixodes ricinus, 1 for I. apronophorus, 6 for I. arboricola, 2 for I. hexagonus, 9 for I. redikorzevi, 1 for I. trianguliceps, 2 for I. vespertilionis, 2 for Haemaphysalis punctata, 1 for H. sulcata, 2 for H. concinna, 1 for D. marginatus, 4 for Rhipicephalus sanguineus sensu lato, 1 for R. bursa and 6 for Hyalomma marginatum.     

The distribution of ticks (Acari: Ixodidae) of domestic livestock in Portugal

This paper introduces the first countrywide faunistic study of the tick parasites on ruminants in Portugal. The aim of this study was to map accurately the distribution of the ticks Dermacentor marginatus, Rhipicephalus (Boophilus) annulatus, R. bursa, Hyalomma m. marginatum, H. lusitanicum and Ixodes ricinus in Portugal. Additional information about the abiotic preferences of these species has been obtained through the use of abiotic (temperature- and vegetation-derived) variables have been recorded from remotely sensed information at a nominal resolution of 1.1 km². A further aim was the development of predictive models of distribution using Classification and Regression Trees (CART) methodologies. Four species (R. annulatus, R. bursa, D. marginatus and H. m. marginatum) are mostly restricted to south-eastern parts of the country, under hot and dry climate conditions of Mediterranean type. H. lusitanicum has been collected almost only in the southern half of Portugal. I. ricinus has a very patchy distribution and is mainly associated with vegetation of Quercus spp., found in southern zones of the country, but it is present also in the more humid western part. A variable number of abiotic variables, mainly temperature derived, are able to describe the preferences of the tick species. It is remarkable that variables derived from maximum values of the Normalized Derived Vegetation Index (yearly or summer-derived) only apply to discriminate areas where I. ricinus has been collected. CART models are able to map the distribution of these ticks with accuracy ranging within 75.3 and 96.4% of actual positive sites.     

Differential distribution of immature Ixodes dammini (Acari: Ixodidae) on rodent hosts

Ectoparasites such as ixodid ticks that remain attached to hosts for several days while feeding on blood are able to overcome the inflammatory and immune responses of some hosts and not others. The immature stages of the deer tick Ixodes dammini are found more frequently on the white-footed mouse, Peromyscus leucopus, than on other rodents. We propose that P. leucopus is more tolerant to I. dammini than is a less common host, the meadow vole, Microtus pennsylvanicus. To test this hypothesis, the distribution patterns and engorgement indices were determined for larval and nymphal I. dammini collected from wild-caught P. leucopus and M. pennsylvanicus. There were more immature ticks, which were more fully engorged, on P. leucopus than on M. pennsylvanicus. There were more and better engorged ticks on male than on female hosts. Laboratory studies on the number and weights of larval I. dammini collected off naive and previously exposed P. leucopus and M. pennsylvanicus support the results of the field study. Fewer larval ticks were recovered from previously exposed M. pennsylvanicus than P. leucopus, and the ticks weighed less. Larval and nymphal ticks aggregated among hosts in the study grid, and higher densities per male P. leucopus were correlated with higher engorgement indices, suggesting that immature I. dammini feed better at higher densities. The feeding success of I. dammini on its preferred host species might be due to its adaptation to the immune and inflammatory reactions of the host.     

Seasonal and geographical distribution of adult Ixodes scapularis say (Acari: Ixodidae) in Louisiana.

The distribution and seasonality of adult black-legged ticks (Ixodes scapularis Say) in Louisiana was measured. The presence of adult ticks was determined by flagging at 106 sites throughout Louisiana. It was concluded that Ixodes scapularis is widely distributed throughout Louisiana. Ticks were also collected twice per month at one site over a 15-month period by flagging and use of CO2 traps to establish the relative seasonal abundance pattern of free-living adult ticks. Host-seeking, black-legged adult ticks were collected from October to May. Peak adult abundance was observed in December. More ticks were collected by the use of CO2 traps compared to flagging in October, November, and February. No black-legged tick larvae or nymphs were collected in this study using either collection method.     

The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae)

Twenty-six species of ticks are reported from the island of Sulawesi (Celebes), Indonesia. These include two species of soft ticks (Argasidae), Carios batuensis and C. vespertilionis, and the following 24 species of hard ticks (Ixodidae): Amblyomma babirussae, A. breviscutatum, A. cordiferum, A. fimbriatum, A. helvolum, A. testudinarium, A. trimaculatum, A. varanense, Dermacentor atrosignatus, D. steini, Haemaphysalis celebensis, H. hystricis, H. kadarsani, H. papuana, H. psalistos, H. renschi, H. toxopei, H. wellingtoni, Ixodes cordifer, I. granulatus, Rhipicephalus haemaphysaloides, R. (Boophilus) microplus, R. pilans and R. sanguineus. This represents an almost three-fold increase in the number of tick species recorded (9) from Sulawesi since the last available list in 1950. The tick records reported herein represent a culmination of data from specimens in the U.S. National Tick Collection, new records of ticks from endemic tarsiers and associated vertebrates, and literature reviews. Collectively, the tick fauna of Sulawesi shows most affinities with the fauna of southeast Asia but there are distinct faunal elements that show relationships with other Indonesian islands, the Philippines or Australasia, as well as a few tick species with widespread geographical distributions. Some ticks known from Sulawesi have known or potential medical-veterinary significance. These include R. (B.) microplus which is a significant pest of cattle and a vector of the agents of bovine anaplasmosis, I. granulatus which is a vector of Langat virus and Lyme disease spirochetes and has been shown to harbor pathogenic rickettsiae in other parts of its range, and R. sanguineus which is a globally widespread ectoparasite of canines and a vector of canine pathogens and parasites.     

Vertebrate hosts of Ixodes pacificus (Acari: Ixodidae) in California.

The western black-legged tick, Ixodes pacificus Cooley & Kohls, is an important parasite and vector of disease agents that affect human and animal health in the western United States. This paper presents a review of all published California host records for I. pacificus. Unpublished data from public health, academic, and vector control agencies and researchers were reviewed as well. Host species were identified for each active life stage (larvae, nymph and adult). A total of 108 vertebrate species in three classes (Mammalia, Aves, and Reptilia) were identified as hosts for at least one life stage of I. pacificus. Adult I. pacificus were recorded from 29 species of mammals, 2 species of birds, and 1 reptile species. Nymphal I. pacificus were recorded from 30 species of mammals, 38 species of birds, and 8 reptile species. Larval I. pacificus were recorded from 29 species of mammals, 43 species of birds, and 8 species of reptiles. A table depicting the taxonomic classification of host species is provided. This review adds eight new host records to the California list of recognized vertebrate host species for I. pacificus.     

The drop-off rhythm of Amblyomma triguttatum triguttatum Koch (Acari: Ixodidae) from laboratory hosts

The pattern of drop-off of Amblyomma triguttatum triguttatum Koch was studied by infesting rabbits and rats with larvae and nymphs, and calves with adult ticks maintained at a daily photoperiodic regime of 12 h light 12 h dark. Additional information was obtained by infesting rabbits with nymphs in or out of phase with the photoperiodic regime experienced by the host.The great majority of ticks detached during the photophase: only 9.7% of larvae and 11.3% of nymphs fed on rats, 12.0% of larvae and 15.8% of nymphs fed on rabbits, respectively, and 10.0% of female ticks detached during the scotophase. The rhythm was not substantially altered by feeding nymphs in or out of phase with the photoperiodic regime of the host. The drop-off rhythm seemed to be coupled with endogenous rhythms of the host.     

A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution

Ticks are important ectoparasites, causing a variety of serious infectious diseases in humans and domestic animals. There is very limited taxonomic information about the tick species of Turkey in the literature, even though Turkey has very suitable climate and vegetation for ticks. In the current study, species diversity, hosts and geographical distribution of the ticks present in Turkey are reviewed based on taxonomic data in the literature from 1915 to 2011 and our recent observations. The names of tick species are arranged according to the most recent check lists. The taxonomic records in the literature and our studies on actual tick samples indicated that the tick fauna of Turkey consists of 46 species; 38 species from Ixodidae and 8 species from Argasidae.     

The natural hosts for larvae and nymphs of Amblyomma neumanni and Amblyomma parvum (Acari: Ixodidae)

Based on the hypothesis that birds and rodents are important hosts for subadults of the Neotropical Amblyomma neumanni and Amblyomma parvum ticks, a survey of these type of hosts was carried out from July 2004 to March 2006, in Quilino (A. parvum) and Dean Funes (A. neumanni), Córdoba province, Argentina. Additionally, monthly tick counts were performed on cattle and goats with occasional tick search in other domestic hosts. Records of questing height of subadult ticks on vegetation were also carried out monthly. Rodents (n = 123) and birds (n = 122) captured in Dean Funes showed no infestation with A. neumanni. Apart of few nymphs found on horses, all larvae and nymphs of A. neumanni were on cattle with a larval prevalence and mean number of 22.2%, and 7.7 ± 22.52, respectively, and a prevalence of nymphs of 47.8% with a mean of 7.9 ± 18.49. The average questing height of larvae and nymphs of A. neumanni was 23.5 ± 17.1 cm and 30.7 ± 26.7 cm, respectively. A total of 138 rodents and 130 birds were captured in Quilino but the Caviidae rodent Galea musteloides carried 99.3% of larvae and 99.8% of nymphs of A. parvum, and no immature stages were detected on cattle, goat or vegetation. Tick counts on G. musteloides (n = 74) showed a prevalence of 42% and a mean number of 9.9 ± 24.83 for larvae, while nymphal infestation had a prevalence of 56.5% and a mean of 8.7 ± 11.31. Cattle appear to be suitable hosts to sustain the complete cycle of A. neumanni in nature (adult ticks infest cattle too) and questing height of subadults indicates that they are expecting to feed on medium and large-sized mammals, such as cattle and other ungulates. At least in the study site, G. musteloides is the principal host for the survival strategy of A. parvum subadults; adult ticks are common on cattle and goats. These hosts are introduced in the Neotropics but A. neumanni was able to develop a surrogate cycle independent of native hosts while A. parvum still depends on probably primeval hosts to sustain their larvae and nymphs.     

Life cycle of Amblyomma cooperi (Acari: Ixodidae) using capybaras (Hydrochaeris hydrochaeris) as hosts

The life cycle of Amblyomma cooperi was evaluated under laboratory conditions testing different host species. Larval infestations were performed on chickens (Gallus gallus) and capybaras (Hydrochaeris hydrochaeris). Nymphal infestations were performed on G. gallus, H. hydrochaeris, guinea pigs (Cavia porcellus) and wild mice (Calomys callosus). Infestations by adult ticks were performed only on capybaras. All free-living stages were observed in darkness at 27 degrees C and RH 85%. Capybaras were significantly (p < 0.05) the most suitable hosts for immature ticks, with the highest larval (63.6%) and nymphal (48%) recovery. Larval and nymphal feeding and premolt periods were significantly different (p < 0.05) between ticks fed on different host species. Male nymphs showed premolt period significantly shorter (p < 0.05) than female nymphs. The overall sex ratio of adult ticks was 0.92:1 (M:F). Infestations by adult ticks on capybaras yielded more than 76% of engorged female recovery. Only three out of 33 engorged females fed on capybaras did not lay fertile eggs. The life cycle of A. cooperi in laboratory, reported for the first time, was completed in an average period of 189.4 days. During the premolt period, all A. cooperi engorged nymphs secreted distinct blackish drops, which seem to be inherent to this species. Our results, associated data in the literature, confirm the high suitability of capybaras for the adult stage of A. cooperi and also indicate this animal species as a primary host for immature stages of A. cooperi in nature. On the other hand, the results of larval and nymphal infestation on chickens and guinea pigs suggest that birds and wild guinea pigs, which are also present in the distribution area of A. cooperi in South America, could be potentially infested by A. cooperi immature stages in nature.     

Wild birds as hosts of Amblyomma cajennense (Fabricius, 1787) (Acari: Ixodidae).

We evaluated the prevalence, mean intensity and relative density of ticks in 467 wild birds of 67 species (12 families) from forest and cerrado habitats at two protected areas of Minas Gerais, between March and September 1997. Ticks collected (n=177) were identified as larvae and nymphs of Amblyomma cajennense and four other species of Amblyomma. We report for the first time 28 bird species as hosts of the immature stages of A. cajennense, demonstrating the lack of host specificity of the larvae and nymphs. A. cajennense had 15% prevalence on birds, with a mean infestation intensity of 0.37 ticks per host sampled, and 2.5 ticks per infested bird. Prevalence varied in relation to host species, diet and between birds from forests at two successional stages. There were no differences in relation to host forest dependence, participation in mixed flocks of birds, and nest type constructed. A. cajennense is a species of medical and veterinary importance, occurring on domestic animals but is known little of its occurrence on wildlife.     

Biology of Amblyomma aureolatum (Pallas, 1772) (Acari: Ixodidae) on some laboratory hosts in Brazil

The ixodid Amblyomma aureolatum is suspected to play a role in the epidemiology of wild life-cycle hemoparasites, which frequently infect dogs in rural and hunting areas in Brazil. Little is known about its bionomics. The objective of the present study was to evaluate some bionomic aspects of A. aureolatum ticks in Brazil. One engorged female, collected from a dog (Canis familiaris) in S?o Sebasti?o das Aguas Claras, State of Minas Gerais, was used to establish a colony in the laboratory. Subsequently its parasitic stage progeny were fed on domestic dogs and laboratory animals. The free-living stages were incubated at 27 degrees C +/- 2 degrees C and minimum 70% relative humidity in a BOD incubator. The egg incubation period ranged from 31 to 34 days; the parasitic period of larvae ranged from 4 to 6 days and ecdysis to nymphs occurred from day 19 up to day 22. The parasitic period of nymphs ranged from 5 to 8 days and the period of ecdysis to adults from 31 to 33 days. The parasitic period of adults ranged from 11 to 15 days, the pre-oviposition period from 6 to 12 days, and the oviposition period from 9 to 38 days. The total duration of the life cycle ranged from 116 to 168 days.     

The midgut hemolysin of Ixodes dammini (Acari:Ixodidae)

Midgut homogenates of the hard tick, Ixodes dammini, lyse erythrocytes from rabbits, rats, hamsters, and guinea pigs. The activity displays sigmoidal kinetics, has an alkaline pH optimum, and is activated by temperature. Hemolytic activity is lost when homogenates are incubated with trypsin or heated for 1 hr at 60 C. Activity is not detectable in nonfed ticks as well as ticks attached for up to 2 days to a host, but increases during the growth phase of feeding. Such activity is postulated to help the initial process of the blood meal digestion by releasing the contents of erythrocytes for further enzymatic hydrolysis.     

Cold-hardiness of Dermacentor marginatus (Acari: Ixodidae)

The cold-hardiness of Dermacentor marginatus using laboratory-reared offspring of ticks collected in Germany was characterized. Investigations of unfed stages revealed that adult ticks suffered 50% mortality at –10°C after 4–5 months, but larvae and nymphs suffered mortality within few days, whereas –15°C was lethal for all stages within a very short period. Larval hatch and moulting of engorged larvae and nymphs did not occur at 10°C. Embryonic development of eggs with larval hatch was considerably reduced by exposure of eggs to 10°C. Engorged females did not lay eggs at 10°C, the oviposition capability, however, persisted over 6 months at 10°C, 5 months at 5°C, 3 months at 0°C and 2 months at –10°C without substantial decrease of the oviposition capacity or reduction of viable eggs. These results present evidence that unfed adult ticks are the ecoepidemiologically most effective stages, which are capable to tolerate long and extremely cold winters without substantial impairment of the population density. It is also considered that engorged females interrupt their oviposition at low and subzero temperatures delaying it for months and so contribute in bypassing winter conditions. None of the stages survived supercooling indicating that D. marginatus is freeze intolerant. Mean supercooling point (SCP) ranged between –26°C in eggs and –12, 6°C in engorged females. Compared with eggs, the SCP of the other stages was significantly higher. In conclusion, the SCP is considered to have no predictive value in the context with cold-hardiness.     

Sigmodontinae rodents as hosts for larvae and nymphs of Ixodes loricatus Neumann, 1899 (Acari: Ixodidae)

Larvae and nymphs of Ixodes loricatus Neumann, 1899 ticks (confirmed by morphological characters and by comparison of 16S mitochondrial rDNA sequences) were collected from Sigmodontinae Wagner, rodents in central and northern Argentina and Uruguay. A total of 100 larvae and 38 nymphs of I. loricatus were collected on the genera Akodon Meyen (n = 36 individuals), Calomys Waterhouse (n = 2), Oligoryzomys Bang in = 12), Oxymycterus Waterhouse (n = 9), and Scapteromys Waterhouse (n = 13). 72 larvae and 18 nymphs were collected on Akodon. Adults of I. loricatus were found in central Argentina and Uruguay on Didelphimorphia of the genera Didelphis Linnaeus and Lutreolina Thomas. Ixodes loricatus has been considered a species with strict total specificity to Didelphimorphia. Our results show that this statement may not be justified. Sigmodontinae rodents are sympatric and share habitats with the phylogenetically distant Didelphimorphia; infestation with I. loricatus seems to be its consequence. We tentatively consider I. loricatus moderately specific to Didelphimorphia.     

Crossbreeding between different geographical populations of the brown dog tick, Rhipicephalus sanguineus (Acari: Ixodidae)

Brown dog ticks are distributed world-wide, and their systematics and phylogeny are the subject of an ongoing debate. The present study evaluates the reproductive compatibility between Rhipicephalus sanguineus ticks from North America, Israel, and Africa. Female ticks of the parent generation were mated with males from the same and alternate colonies. Every pure and hybrid cohort was maintained separately into the F2 generation with F1 females being allowed to mate only with males from the same cohort. The following survival parameters were measured and recorded for every developmental stage: feeding duration and success; engorgement weight, fertility, and fecundity of females; molting and hatching success. Ticks from North American and Mediterranean populations hybridized successfully. The survival parameters of all their hybrid lines were similar to those in pure lines throughout the F1 generation, and F1 adults were fully fertile. Parent adult ticks from the African population hybridized with either North American or Mediterranean ticks and produced viable progenies whose survival parameters were also similar to those in pure lines throughout the F1 generation. However, F1 adults in the four hybrid lines that included African ancestry were infertile. No parthenogenesis was observed in any pure or hybrid lines as proportion of males in F1 generation ranged from 40 to 60 %. Phylogenetic analysis of the 12S rDNA gene sequences placed African ticks into a separate clade from those of the North American or Mediterranean origins. Our results demonstrate that Rh. sanguineus ticks from North America and Israel represent the same species, whereas the African population used in this study is significantly distant and probably represents a different taxon.     

Interspecific variations in attachment sites and density assessment in femaleIxodes rubicundus (Acari: Ixodidae) on domestic and natural hosts

The attachment sites of female Karoo paralysis ticks (Ixodes rubicundus) and equations to assess tick burdens on domestic (sheep and goats) and natural (mountain reedbuck) hosts were determined. No intraspecific differences in attachment sites for Merino sheep and Angora goats could be observed, but interspecific differences were evident.Ixodes rubicundus in general attaches on the ventral aspects of the host. More than 50% of these ticks attached to the fron quarters of the body of the domestic host, and 60% to the hindquarters of the natural host. The length of Merino sheep wool had no affect on either the attachment site or number of attached ticks. In general, more ticks were found attached to the woollen than non-woollen body areas. Equations are provided which can estimate tick burdens on hosts by counting the ticks on two or three selected body areas of the host. To facilitate sampling procedures, approximate sample sizes for the various hosts needed to estimate average tick burdens have been provided. Depending on the level of precision required (with bounds on the error of estimation equal to 1, 2 or 5 ticks), the number of animals in a flock of 200 Angora goats that must be examined varied from 26 to 85.