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Life cycle parameters of 2 closely related tick species, Rhipicephalus sanguineus Latreille and R. turanicus Pomerantsev, were studied under laboratory conditions. Both Rhipicephalus, which have small adults, demonstrated the same adaptations as large tick species inhabiting deserts and semideserts: high reproductive rate, decrease in egg size, and an increase in interstage growth to compensate for the smaller size at birth. Pronounced quantitative differences between both species were discerned in relation to these adaptations. Female R. turanicus produced twice as many eggs as R. sanguineus which was facilitated by the greater amount of blood engorged by females and by the smaller egg weight in R. turanicus as compared with R. sanguineus. In all developmental stages, the weight increase from unfed to fed ticks was greater in R. turanicus than in R. sanguineus (23% higher in larvae, 118% in nymphs, and 26% in females). The increase in weight in R. turanicus from the unfed larva (0.013 mg) to the unfed female (3.31 mg) was 254-fold, and in R. sanguineus it was 127-fold (from 0.021 to 2.54 mg). In nymphal R. turanicus, the higher density and the greater height of the dorsal epicuticular folds, as well as procuticular indentations found inside the folds allow this tick to stretch its alloscutum during blood engorgement to a greater extent than R. sanguineus. The rates of blood ingestion (for nymphs and females), egg maturation, and metamorphosis were 1.1-1.7 times greater in R. turanicus than in R. sanguineus. A life cycle strategy with both a higher reproductive rate and faster development in R turanicus may be explained by its greater dependence on environmental factors than that in R. sanguineus.

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This review analyzes 30 yr of data concerning physiological age of ixodid ticks. The level and state of nutritional reserves in unfed ticks has been proposed as an index of tick physiological age. The dynamics of the use of these substances reflect tick aging. Thus, physiological age of ixodid ticks is determined by estimating the irreversible changes caused by natural vital activity in the body of unfed ticks, which inevitably raises the probability of tick death. The 4 age groups used in most studies correspond to different periods in the life of unfed ticks: the postmolting development, the commencement of active life (= questing activity), the midperiod of active life, and the final period of active life. The aging of adult ixodid ticks has been studied in natural populations and laboratory colonies of different species belonging to several genera. The dynamics of age composition through the activity season observed in adult Ixodes persulcatus, I. ricinus, and Dermacentor reticulatus from field populations correlated well with the known data on the life history of these species. Physiological age was successfully used as an indicator of the patterns of tick activization (= beginning of questing activity). Tick susceptibility to acaricides was shown to increase with tick aging. The success of maintenance, reproduction, and transmission of pathogens was found to be strongly influenced by tick physiological age. The physiological age can provide information about the present physiological status of an individual tick or a group of ticks and, on this basis, predictions can be made (for example, life expectancy, degree of response to certain factors). The physiological age does not give reliable information about the past life of ticks, in particular their calendar age. In this respect, the meaning of the physiological age of ixodid ticks corresponds to the biological (physiological, functional) age of animals, as used in gerontology, and differs from the physiological age of mosquitoes and other bloodsucking dipterans. Balashov's histological method of tick age determination successfully used by many authors, remains the most appropriate, until now. However, there is ample room for further advancement of methodologies of tick age assessment. The future methods should be based on the quantitative estimation of a number of characters (test-battery) related to different organs or systems.

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, hostseeking. 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.

In some studies the prevalence of tick infection (infection rate) and the intensity of infection are negatively correlated with unfed tick age (in the broad sense of this term). However, no special research has been carried out to consider the phenomenon thoroughly. The infection indices of the female taiga ticks, Ixodes persulcatus, infected with Borrelia burgdorferi s.l. were related to tick physiological age, an index that more precisely reflects tick physiological state than the time of tick collection in the field or the duration of tick survival under laboratory conditions. A novel quantitative technique of physiological age determination based on the evaluation of the ratios between sizes of the stable (scutum) and the changing (alloscutum) structures of the tick body was used. The age was estimated in accordance with the classical age-grade scale introduced by Balashov and a more fractional scale determined by the new technique. In total, 131 female ticks were examined for their infection and physiological age, 46 of which were infected with B. burgdorferi s.l. (mean infection rate 35.1%). The minimal intensity of infection was 0.4 bacterial cells per 100 fields of view whereas the maximal infection was 172 cells. There was no difference between the prevalence of infection in ticks of different physiological age. The intensity of infection obviously differed between ticks of different age groups in the scale introduced by Balashov but did not significantly differ between ticks of different age groups according to the fractional age-grade scale. The data concerning the relationships between Borrelia burgdorferi and unfed Ixodes ticks are considered.

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