Characteristic features of biodiversity in European ixodid ticks (Acari, Ixodidae) as vectors of diseses with natural foci

The diapason of some biodiversity characteristics of ixodid ticks of the fauna of Europe and adjacent territories is analyzed. These characteristics include taxonomic and morphological diversity, the diversity of life cycles, host-parasite relations, species ranges, and biotopic associations. The significance of different aspects of biodiversity of ixodid ticks as a prerequisite of interrelations with pathogens with natural foci is estimated.     

Ability of successful attack in two species of ixodid ticks (Acari: Ixodidae) as a manifestation of their aggressiveness

Patterns of attacking hosts by two closely related species of ixodid ticks (Ixodes persulcatus and I. ricinus) were compared in parallel field tests. The ability of active unfed adults of both sexes to adhere to a flannel flag dragged over grass vegetation and to remain on the flag during this process were estimated. The tests were conducted under two temperature conditions, 6–10°C and 17–22°C. In all test versions, adults of I. persulcatus were more successful both in adhering to the flag and in remaining on it. There were no consistent differences between males and females of the same species. The results demonstrated a differing ability of successful attack in both tick species. This ability is a complex derivative of the tick activity and aggressiveness. An attempt is made to analyse the latter phenomenon, which is considered as a general one for all bloodsucking arthropods. The aggressiveness of arthropods should be taken into account as one of the leading factors influencing the sampling results.     

The intraspecific taxonomic structure and identification of closely related species of ixodid ticks (Ixodidae)

Two variants of intraspecific morphological differentiation of 7 palearctic species of Ixodidae and respectively two variants of their intraspecific taxonomic pattern have been revealed on the basis of the study of geographic variation of each species. Three species--Ixodes pavlovskyi Pom., 1946, Haemaphysalis erinacei Pavesi, 1884, Hyalomma asiaticum Sch. et Schl., 1929, are subdivided into subspecies, and four ones--I. ricinus (L., 1758), I. persulcatus Sch., 1930, I. crenulatus Koch, 1844, Dermacentor marginatus (Sulzer, 1776),--into morphotypes by the degree of distinction of the whole complex of active stages in ontogenesis. Acknowledging the conventional character of the categories of subspecies and morphotype their criteria have been formulated for ixodid ticks. Paleogeographic and ecological grounds are given. Some data of geographic variation of immature stages favour the identification of closely related species in concrete localities.     

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.     

New species of ixodid ticks (Ixodidae) for the fauna of Armenia

Two species of ixodids , Ixodes arboricola Schulze et Schlottke and I. frontalis ( Panzer ), new for the fauna of Armenia are given. Larvae and nymphs of these ticks were found on birds in the forest zone and gardens.     

Morpho-functional changes in the midgut of ixodid ticks (Acari: Ixodidae) during the life cycle

Morpho-functional studies performed in five species of the genus Ixodes, including I. pacificus, I. pavlovskyi, I. persulcatus, I. ricinus, and I. scapularis demonstrated that the replacement of the midgut epithelium lags in each consecutive life stage and is not synchronized with basic processes of morphogenesis and organogenesis during molts. The midgut epithelium of each preceding stage is retained and functions during feeding of the subsequent life stage.     

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.     

Characteristic features of the ranks genus and subgenus, and an intercalary rank "species complex" in ixodid ticks (Acari: Ixodidae)

The work was carries our from the standpoint of the morphological conception of species. Vast collections of the Zoological Institute of the Russian Academy of Science testify to the existence of hiatuses in both genera and subgenera of Palearctic ixodids at all active phases of their ontogenesis. The fact that the subgenera of Palearctic genera have been well studies is notes, and composition of the subgenera is presented. The question of a taxonomic intercalary rank "species complex" is considered in detail in the context of the coevolution between some complexes of closely related species of ixodid ticks and some closely related species (genospecies, strains) of pathogens. The question of the taxonomic rank "species complex" in ixodid ticks as a phyletic species association is postulated on the basis of comparative ontogenetic data. Nomenclature status of the intercalary association "species complex" is specified in conformity with the fourth edition of the International Code of Zoological Nomenclature. Species composition of most studied complexes is presented. Some variants of morphological differentiation between species complexes within subgenus are considered. Significance of the taxonomic concept "species complex" for zoological. parasitological, and medical aspects of the ixodid ticks study was evaluated. Prognostic significance of the rank "species complex" for the study of the relationships between ixodid ticks and pathogens is discussed.     

Long-term dynamics of population density and species composition of pasture ixodid ticks (Parasitiformes,Ixodidae) in anthropogenic and natural areas

Long-term monitoring of population density and species composition of ixodid ticks inhabiting the territory of Tomsk and its outskirts has demonstrated that at least two species (Ixodes persulcatus and I. pavlovskyi) simultaneously inhabit all the biotopes examined. It was found that the fraction of I. pavlovskyi increased in biotopes adjacent to city buildings. I. persulcatus dominated in natural control biotopes, whereas I. pavlovskyi were found in these biotopes as solitary specimens. It was demonstrated that during 15 years of observation (1996–2011), the population density of ticks in the outskirts significantly increased. The combination of ecological and chemical methods of tick control in limited territories can result in a noticeable decrease of tick population density. Two tick species non-typical of the region (Dermacentor reticulatus and Haemaphysalis concinna) were revealed in one biotope.     

Sites of attachment and density assessment of ixodid ticks (Acari: Ixodidae) on impala (Aepyceros melampus)

During 1994 and 1995, 45 impala ewes, Aepyceros melampus were examined at Letaba Ranch in the Northern Province. Tick counts were made from 15 animals on three occasions, in July, October and February. The objective was to determine whether the total body tick counts can be estimated from counts done on specific, selected sampling sites on the skin. Twelve sites were shaved within ten predilection sites and the parasite numbers counted from these samples. These counts were compared to the total body parasite count as determined by the scrub and digestion techniques. More than 80% of all the ticks were present on the muzzle, the head, the pinna and the legs. Larger numbers of ticks were collected in October than in July or February. Two mathematical models were tested for the tick counts. The first model was made up of tick counts from a single shaved sampling site, the pinna. The correlation between the tick counts on the pinna and the total tick counts was highly significant (p values ranging between 0.0208 and 0.0001). The second model was developed based on tick counts from four regions, namely the head, the pinna and the front and hind feet. A less significant correlation was obtained between the number of ticks counted on the four sites and the total tick count.     

Evaluation of the apron as a taxonomic trait in ixodid ticks of the genus Dermacentor Koch (Ixodoidea, Ixodidae)

Study of the apron in 9 species of the genus Dermacentor from the fauna of the USSR has revealed differencies in its structure. The subgenus Dermacentor (s. str.) differs from two other subgenera both in the shape of the apron itself and in the shape of the postgenital sclerite and setae of perigenital area. Close species within each of two other subgenera differ in apron proportion, shape and size of denticles along its hind edge, and sometimes in their number. Inspite of the statistically reliable interspecific differences in apron structure a wide range of individual variability of some details and geographical specificity of samples from various places of the area were observed in species with a vast area.     

Morphological inversions during the ontogenesis of the ixodid ticks (Acari: Ixodidae) from the viewpoint of the hypermorphosis conception and their significance for systematics

Morphological inversions during the ontogenesis of the ixodid ticks consist in the deranges of species-specific tendencies in the development of some external structures by the female line. The inversions, which included a wide range of organs and their fine structures, have been described for the first time in 14 species of 4 from 7 Palearctic ixodid genera. Emphasis is placed on the study of the structures, which serve as differential characters for closely related species. It is shown, that the using of the structures subjected to inversions is unavoidable in systematics because of the great similarity of closely related species, especially in immature stages. It is established by the data on the variability of all active stages in the species distribution ranges, that the degree of the inversions manifestation may somewhat fluctuate in different subspecies or geographic morphotypes of the same species, but the peculiarity of the inversions always remains a species-specific character. The cause of inversions origin is the evolution of the Ixodidae by the way of morpho-ecological specialization, namely hypertrophic, giantism, the growth of teguments in the period of nutrition act by pasture ambushing, which led to the derange of the development coordination of the organs during ontogenesis. Biological implication of the inversions is a guarantee of the adaptive potential of species during the process of microevolution. Even though the inversions make identification of closely related species difficult, their substantial importance is that they form integral species-specific character for the whole active part of the species ontogenesis.     

Morphofunctional changes of salivary glands of female ixodid ticks of subfamilies Ixodinae and Amblyomminae (Acari: Ixodidae) during feeding and their significance

Using light and electron microscopy, a dynamics of functioning of salivary glands of female ticks of the genus Ixodes was studied. Comparative analysis of morphofunctional changes and role of the salivary gland secretions of females of subfamilies Ixodinae and Amblyomminae during feeding was performed on the basis of literature data and our own studies.     

Sexual transmission of the tick-borne encephalitis virus in ixodid ticks (Ixodidae)

Sexual transmission of tick-borne encephalitis virus from infected ixodid males to noninfected females is shown: in Ixodes persulcatus in 50% (6 of 12) and in Hyalomma anatolicum in 6.2% (1 of 16). The sexual transmission of tick-borne encephalitis virus is shown to provide a transmissible transfer of the virus into eggs in ixodid ticks. Electron microscope studies of the sexual system of ixodid males infected with the virus have revealed numerous morphologically mature virus particles in lumens of endoplasmic reticulum, in vacuoles of Golgi complex of spermatocytes and in association with tubular elements of spermatids.     

Two new species of African Haemaphysalis ticks (Acari: Ixodidae), carnivore parasites of the H. (Rhipistoma) leachi group

Two new tick species belonging to the African Haemaphysalis (Rhipistoma) leachi subgroup, namely H. (R.) colesbergensis n. sp. and H. (R.) oliveri n. sp., are described. Haemaphysalis (R.) colesbergensis adults are easily differentiated from the other species of the H. (R.) leachi subgroup, including H. (R.) oliveri, by the spur on coxa IV, which is considerably longer than that on coxa III. The adults of the 2 new species are equal in size, but the dental formula of the hypostome of H. (R.) colesbergensis is 4/4 compared to 5/5 for H. (R.) oliveri. The dental formula of H. (R.) oliveri also distinguishes it from other ticks in the subgroup, namely H. (R.) leachi, H. (R.) elliptica, H. (R.) moreli, and H. (R.) punctaleachi (4/4 in these species), but not from H. (R.) paraleachi, which has a 5/5 dental arrangement. However, the average total length and width of H. (R.) oliveri males (2.47 x 1.20 mm) are considerably shorter and narrower than those of H. (R.) paraleachi males (3.81 x 1.79 mm). Similar differences in size apply to the females. Nymphs and larvae of H. (R.) colesbergensis and H. (R.) oliveri can be distinguished from those of other members of the H. (R.) leachi subgroup, as well as from each other, by a combination of the following characters: size and measurement ratios, length of posterodorsal and posteroventral spurs on palpal segment II, and number of denticles per file on the hypostome. Haemaphysalis (R.) colesbergensis is known only from South Africa, where it has been collected from domestic cats and dogs and medium-sized wild felids. Haemaphysalis (R.) oliveri is recorded only from Sudan, where it has been collected from small- to medium-sized wild felids and canids and an antelope. The hosts of the immature stages of H. (R.) colesbergensis are unknown, while nymphs of H. (R.) oliveri have been collected from rodents.     

Evolution of the basic types of life cycle in ixodid ticks (Ixodidae)

The burrow-nest life scheme of ixodid ticks, which was formed at the beginning of the Chalk, gave rise to the subsequent types of parasitism: pasture (end of the Chalk), pasture-burrow (end of Paleogene) and pasture-stall one (late Neogene). Changes in the climate in different geological epochs and regions together with the hosts' evolution, natural selection in the aggregate of variability and heredity of ixodids in their populations were the cause of the evolution.     

Ecdysteroids as a component of the genital sex pheromone in two species of hard ticks,Dermacentor variabilis andDermacentor andersoni (Acari: Ixodidae)

Neutering of part-fed females eliminated copulatory behavior inDermacentor variabilis (Say) andD. andersoni Stiles males. Extracts from the anterior reproductive tracts of part-fed (7 days) females partly restored the male copulatory behavior in conspecific neutered females. Similar extracts from unfed females did not restore the behavior, suggesting that the pheromone was produced during feeding. Perception of the genital sex pheromone by sensillae on the male cheliceral digits was confirmed by electrophysiological techniques.Males ofD. variabilis andD. andersoni responsed positively to authentic ecdysone and 20-hydroxyccdysone (20HE) in neutered female bioassays. Responses to sterols were significantly lower than to ecdysteroids. Electrophysiological assays suggest a sensitivity of males to high doses of ecdysteroids. The strongest responses were to 20HE in both species. Ecdysteroids, specifically ecdysone and 20HE were shown to be present in the anterior reproductive tracts in excess of amounts that could be explained by mere hemolymph contamination. Ecdysteroids were also found in washings of the vaginal lumen of these two species.Dermacentor andersoni females contained larger amounts of ecdysteroids thanD. variabilis females. 20-hydroxyecdysone and possibly ecdysone appear to be components of the genital sex pheromone ofD. variabilis andD. andersoni. Species recognition may be facilitated by these components, but the complete mechanism is not yet fully understood.Supported by grant AI10, 986 from the National Institute of Allergy and Infectious Diseases, National Institutes of Health, U.S. Public Health Service, Department of Health and Human Services, Bethesda, Maryland.