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Detection of Borrelia burgdorferi in laboratory-reared Ixodes dammini (Acari: Ixodidae) fed on experimentally inoculated white-tailed deer.

Larvae and nymphs of Ixodes dammini Spielman, Piesman, Clifford & Corwin from a laboratory colony were fed on two white-tailed deer, Odocoileus virginianus (Zimmerman) inoculated with either the SH2-82 or JD-1 strains of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt & Brenner. Ticks were exposed to one deer 43 and 69 d after inoculation of the spirochete and to a second deer 35 and 61 d after inoculation. Polymerase chain reaction assays amplified the 158 bp OspA DNA target sequence in 11.1% (n = 9) of fed larvae and 3.3% (n = 30) of nymphs from the deer inoculated with the SH2-82 strain, and 22.7% (n = 22) of larvae and 0% (n = 21) of nymphs from a second deer inoculated with the JD-1 strain of B. burgdorferi. One of three females derived from nymphs fed on one of the inoculated deer showed presence of B. burgdorferi DNA, but none of four males was positive. Experimentally inoculated deer can serve as a source of at least two geographic strains of B. burgdorferi to I. dammini larvae and nymphs for at least several weeks.

Animals↗

Survival of immature Ixodes scapularis (Acari: Ixodidae) at different relative humidities.

Survival of unfed larvae and nymphs of Ixodes scapularis Say (formerly named I. dammini Spielman, Clifford, Piesman & Corwin) was tested at 27 degrees C for relative humidities ranging from 65 to 100%, which were maintained by various saturated salts or water. The times until half the larvae died (LT50) at 100, 93, 85, 75, and 65% RH were 67.1, 26.6, 8.3, 1.3, and 1.1 d, respectively. Maximal larval survival in the laboratory was 129, 59, 24, 3, and 2 d at the same relative humidities. There was a distinct difference between the mortality curves for larvae at 93% RH and 100% RH, which suggests that larvae were taking in water from the air at the higher RH whereas those at an RH < or = 93% were not. There was little difference between the 100 and 93% RH survival curves for the nymphs. All of the nymphs at 100% RH survived at least 149 d. At 93 and 100% RH, one nymph survived at each RH for 210 and 205 d, respectively. The LT50s for the nymphs of I. scapularis were 169.9 (100% RH), 168.5 (93%), 118.8 (85%), 10.7 (75%), and 3.6 (65%) d. Maximal nymphal survival in the laboratory was 162, 65, and 8 d at 85, 75, and 65% RH, respectively. The inability of the larvae to retain water effectively is probably more important than the ability to imbibe water from subsaturated air in the survival of this stage. Subadult I. scapularis require a relatively humid habitat for survival, although some individuals have the potential to seek a host for many weeks within a mesic environment.

Acclimatization↗

Effects of Babesia microti infection on feeding pattern, engorged body weight, and molting rate of immature Ixodes scapularis (Acari: Ixodidae).

Effects of Babesia microti Franca on the pattern of feeding time, the body weight of engorged ticks, and the molting rate of the blacklegged tick, Ixodes scapularis Say, were determined. Using the Syrian golden hamster, Mesocricetus auratus (Waterhouse), as an animal model, we found no significant differences in patterns of feeding time determined for both larvae and nymphs that fed on B. microti-infected and uninfected hamsters. However, an infection of B. microti in hamsters delayed engorgement of the ticks. The presence of B. microti had no adverse effects on the body weight of engorged ticks. The mean body weight of groups of 5 engorged larvae fed on infected hamsters was not significantly different from that on uninfected hamsters. The mean body weight of individual nymphs that fed on infected hosts was significantly higher than that on uninfected hosts. Larvae fed on infected hamsters molted in greater numbers than those fed on uninfected hamsters. However, there was no significant difference in molting rates of nymphs derived from infected and uninfected hamsters. In addition, greater body weights and higher molting rates were observed in both larvae and nymphs that fed on uninfected white-footed mice, Peromyscus leucopus Rafinesque, compared with those on uninfected hamsters. Nymphs fed on mice spent a significantly longer time achieving repletion than those on uninfected hamsters. We suggest a mutualistic relationship in the interactions between B. microti and I. scapularis. This mutualism may potentially enhance the long-term coexistence and survival of both species. Furthermore, we suggest that the pattern of feeding time, the body weight of engorged ticks, and the molting rate of immature I. scapularis are host-dependent.

Animals↗

Comparison of methods for introducing and producing artificial infection of ixodid ticks (Acari: Ixodidae) with Ehrlichia chaffeensis.

Only 29.5 +/- 8.91% of engorged Amblyomma americanum (L.) nymphs that we inoculated with Ehrlichia chaffeensis molted successfully to adults compared with 75.8 +/- 7.46% of engorged nymphs that were not inoculated. However, 65.4 +/- 6.02% of unfed nymphs of this species were exposed for 2 h to E. chaffeensis suspension introduced to them through glass capillaries gained weight. These nymphs were placed on rabbits, and approximately 50% of them completed their feeding and molted successfully to adults. Weight gained was higher (71.8 +/- 17.33% and 69.8 +/- 23.26%) for unfed A. americanum females that fed from capillaries for 2 and 24, h respectively, than for nymphs. Similar values were recorded for Dermacentor variabilis (Say) (61.0 +/- 16.23%) and Rhipicephalus sanguineus (Latreille) (59.0 +/- 18.62%) females after 24 h of capillary feeding. The amount of E. chaffeensis suspension taken in by females of A. americanum, D. variabilis, and R. sanguineus during 24 h of feeding was 11.2 +/- 3.56, 10.9 +/- 4.29 and 6.3 +/- 2.35 microliters, respectively. This volume is equivalent to approximately 12,969, 12,622, and 7,295 infected cells ingested by the species mentioned above. Positive correlation between the volume taken in by the ticks and the weight gained by the females was found, but the initial weight of the unfed females did not effect the weight they gained. The pathogen was found in the females of all 3 species by polymerase chain reaction procedures for at least 7 d, indicating that the capillary feeding method can be successfully used for infecting unfed ticks. The potential use of this method is discussed.

Animals↗

The role of Amblyomma variegatum in the transmission of heartwater with special reference to Guadeloupe.

Heartwater has been diagnosed in Guadeloupe, Marie-Galante, and Antigua; it induces important losses among goats and European or cross-bred cattle when local zebu creole are highly resistant to infection. Amblyomma variegatum is the vector of the disease in the Caribbean. The tick strain of Martinique, occurring in that island that has apparently been disease-free since 1948, has not lost its ability to experimentally transmit the disease. In Guadeloupe 97% of nymphs and nearly 100% of adults feed on cattle and goats. Some immature ticks (4.5%) feed on wildlife, including birds that may be involved in the spread of infected ticks. Only 1 to 4% of adult ticks are infected and only a proportion of infected ticks are really infective: 53%, 9%, and 50% of nymphs, males, and females, respectively. Nymphs play the major role in the transmission: they are more numerous than adults and engorge faster. Infected ticks have the same maximum longevity as noninfected ticks, that is, 18 months for nymphs and nearly 23 months for adults. Cowdria is not transmitted immediately after tick fixation but after a delay of 2-3 days for nymphs and 4 days for adults. The disease is more often fatal when transmitted by tick biting rather than by needle transmission. The daily infection rate that summarizes all the parameters is very low (0.14% and 0.20% for goats and cattle, respectively), resulting in an unstable epidemiological situation. The transmission of Cowdria by A. variegatum in Guadeloupe shows significant differences compared with the transmission by A. hebraeum in Africa.

Animals↗

Simultaneous detection and genotyping of three genomic groups of Borrelia burgdorferi sensu lato in Dutch Ixodes ricinus ticks by characterization of the amplified intergenic spacer region between 5S and 23S rRNA genes.

We developed a rapid and reliable method for the identification Borrelia burgdorferi sensu lato species in ticks. We used the DNA sequence polymorphism of the spacer region between 5S and 23S rRNA genes, which has been shown to be able to discriminate between eight genomic groups of B. burgdorferi sensu lato (D. Postic, M. Assous, P. A. D. Grimont, and G. Baranton, Int. J. Syst. Bacteriol. 44:743-752, 1994). Spacer DNA was amplified by PCR and was then hybridized to five membrane-bound oligonucleotides. The oligonucleotides were specific for B. burgdorferi sensu stricto, Borrelia garinii, Borrelia afzelii, and group VS116. A probe which reacted with all genomic groups of B. burgdorferi sensu lato was also used. Ninety-six ticks collected in the field were destructed by bead beating, and the supernatant was used directly in a PCR. B. burgdorferi sensu lato DNA was detected in 6 of 57 adult ticks (11%) and 9 of 39 nymphs (23%). B. garinii was found in three nymphs and four adults, three nymphs carried B. afzelii, and one adult and one nymph carried group VS116. Double infections with B. afzelii and group VS116 were found in two nymphs and one adult. Thus, our method can simultaneously identify three genomic groups of B. burgdorferi sensu lato in ticks collected in the field. This technique provides new ways to study the association of genomic groups present in ticks and the risk of Lyme borreliosis.

Animals↗

Low temperature acclimated populations of the grain aphid Sitobion avenae retain ability to rapidly cold harden with enhanced fitness.

In contrast to previous studies of rapid cold-hardening (RCH), which have investigated the responses of insects maintained under 'summer conditions' (20 degrees to 25 degrees C), this study focuses on the ability of low-temperature acclimated insects to undergo RCH. When the grain aphid Sitobion avenae Fabricus was low-temperature acclimated by rearing for three generations at 10 degrees C, the discriminating temperatures (temperature that results in approximately 20% survival after direct transfer from the rearing temperature to a sub-zero temperature for a period of 3 h), of first instar nymphs and adult aphids were -11.5 degrees and -12 degrees C, respectively. Maximum rapid cold-hardening was induced by cooling aphids at 0 degrees C for 2 h (nymphs) or 30 min (adults), resulting in survival at the respective discriminating temperatures increasing from 26% to 96% (nymphs) and 22% to 70% (adults). Cooling from 10 degrees to 0 degrees C at 1 degree, 0.1 degrees and 0.05 degrees C min-1 significantly increased survival of nymphs at the discriminating temperature, but not of adults. There were no ;ecological costs' associated with rapid cold-hardening at 0 degrees C, or with exposure of rapidly cold-hardened aphids to the discriminating temperatures; fecundity and longevity, in both nymphs and adults were either similar to control aphids or significantly increased. The study demonstrates that rapid cold-hardening ability is retained in aphids that have already undergone cold-acclimation, as would be the case in overwintering aphids. Both rapid cold-hardening and subsequent exposure at previously lethal temperatures can enhance fitness in surviving individuals.

Acclimatization↗

[Population statistics of Triatoma sordida Stäl 1859 (Hemiptera:Reduviidae) in experimental conditions].

Four cohorts of 100 Triatoma sordida eggs were reared under standard laboratory conditions (28 degrees C, 63% R.H.) and fed weekly on hens during 40 minutes. The numbers of dead insects, moults and eggs laid were recorded weekly. A constant survival value was observed without any important peaks in the four cohorts. Life expectancy was 37.4 weeks for males and 36.8 weeks for females. The average generation time was 61.7 weeks. The average net reproduction rate suggests that this population would increase 143.2 times during each generation. The average intrinsic rate of natural increase was 0.082 per individual per week. The highest reproductive value was observed 3 weeks after entering the adult stage. The age structure, assuming stable age distribution, would be 32% eggs, 26% of first instar nymphs, 19% of second instar nymphs, 13% of third instar nymphs, 6% of fourth instar nymphs, 3% of fifth instar nymphs and 1% of adults. Our results suggest that this species behaves as a k-strategist under experimental conditions.

Animals↗

Life cycle and reproductive parameters of Clerada apicicornis signoret (Hemiptera: Lygaeidae) under laboratory conditions.

The life cycle of Clerada apicicornis was determined under laboratory conditions. Mean development times in days were: egg 27.2, nymph I 12.5, nymph II 12, nymph III 13.4, nymph IV 16.4, nymph V 26. The life expectancy of adults ranged from 117 to 317 days (mean 196 days). Based on a cohort of 29 females of C. apicicornis, a horizontal life table was constructed. The following predictive parameters were obtained: net rate of reproduction (Ro = 48.31), intrinsic rate of population increase (r(m) = 0.153), generation time (Tc = 28.20 weeks), and finite rate of population increment (lambda = 1.16). The reproductive value (Vx) for each age class of the cohort females was calculated. The following observed parameters were calculated after mortality in each stage: net rate of reproduction (R'o=13.4), intrinsic rate of population increase (r c' =0.09 ), and finite rate of population increment (lambda' =1.1). The generation time (Tc' =27.4) was estimated using the methods of Laughlin and Bengstron. A vertical life table was elaborated and mortality was described for one generation of the cohort.

Animals↗

Host selection, development, survival, and reproduction of turnip aphid (Homoptera: Aphididae) on green and red cabbage varieties.

Host selection, development, survival, and reproduction of the turnip aphid, Lipaphis erysimi (Kaltenbach), on selected green and red cabbage varieties, Brassicae oleracea L. variety capitata, were studied in the laboratory. Although numbers of alate L. erysimi on different varieties varied generally, more alate adults and nymphs were observed on green than on red varieties on day 3; whereas, these differences were not so significant on day 15. There were no significant differences in alate adults on all six varieties on day 15. Numbers of nymphs and apterous adults of L. erysimi varied among the varieties. When mean numbers of those aphids on green varieties were compared with those on red varieties, the green varieties had more nymphs and apterous adults than those on the red varieties. Developmental durations of L. erysimi nymphs also varied among all varieties, and the nymphs fed on green varieties developed longer than those fed on red varieties. Adult longevities on these cabbage varieties were not significantly different; whereas, the adult fecundities were. Survival rates of L. erysimi varied greatly among all varieties, but there are no significant differences between the green and red varieties. Factors that might affect aphid host selection, growth, longevity, fecundity, and survivorship were discussed.

Animals↗

Susceptibility of Maconellicoccus hirsutus (Homoptera: Pseudococcidae) to methyl bromide.

Eggs, crawlers, early nymphs, late nymphs, and adults of the pink hibiscus mealybug, Maconellicoccus hirsutus (Green), were tested for their susceptibility to methyl bromide in 2-h laboratory fumigations at ambient conditions (25 degrees C, 95% RH). Dose-response tests indicated that the egg was the most susceptible life stage with an LC99 of 20.2 mg/liter. Based on probit analysis of dose-response data, no significant differences were observed among susceptibilities of the crawler, early stage or late stage nymphs, or adults at either the LC50 or LC99 level, but late stage nymphs were more tolerant than early stage nymphs in a separate paired comparison test. Confirmatory tests showed that a dose of 48 mg/liter methyl bromide, the USDA-Animal Plant Health Inspection Service treatment dose schedule for mealybugs at 21-26 degrees C, produced 100% mortality of all life stages. On the basis of these results, we conclude that the methyl bromide treatment schedule for mealybugs will provide quarantine security for M. hirsutus infesting commodities for export or import.

Animals↗

Ixodes scapularis (Acari: Ixodidae): abundance and rate of infection with Borrelia burgdorferi in four state parks in Wisconsin.

Four state parks located in Lyme disease endemic regions of Wisconsin were surveyed for the presence of Ixodes scapularis Say during May and June of 1998 by drag sampling along hiking trails. Nymphal abundance varied between parks, with the average number of nymphs encountered in 1 h ranging from 6.2 +/- 3.8-47.1 +/- 36.3 (mean +/- SD). Questing nymphs were tested for the presence of Borrelia burgdorferi by culture in BSK medium and 7-12% was found to be infected. The average risk of encountering an infected nymph (entomologic risk index) ranged from 0.5 to 5.2 infected nymphs per hour. The highest entomological risk index was recorded from a small island park in northwestern Wisconsin during the last week in May (8.0 infected nymphs per hour). These results indicate a lower risk for human Lyme disease exposure in Wisconsin state parks in comparison with highly endemic areas of the northeastern United States.

Animals↗

Ticks (Acari: Ixodidae) and spirochetes (spirochaetaceae: spirochaetales) recovered from birds on a Georgia Barrier Island.

From September 1997 through July 1999, 300 individuals and 46 species of birds were mist-netted and screened for ticks and spirochetes on St. Catherine's Island, Liberty County, GA. Seventy-six (25%) of the birds were parasitized by a meal intensity of 4.6 ticks. Seasonally, more birds were infested with ticks during the summer (50% in 1998, 34% in 1999) than in spring (15% in 1998, 11% in 1999) or fall (21% in 1997, 20% in 1998), mainly because of severe infestations on some birds by immature stages of the lone star tick, Amblyomma americanum (L.), during this season. Eight species ofticks were recovered from 14 species of birds during this study: A. americanum (74 nymphs, 168 larvae); the blacklegged tick, Ixodes scapularis Say (11 nymphs, 28 larvae), the Gulf Coast tick, Amblyomma maculatum Koch (two nymphs, 29 larvae); Ixodes minor Neumann (16 larvae); the rabbit tick. Haemaphysalis leporispalustris (Packard) (one nymph, 14 larvae); the bird tick Ixodes brunneus Koch (two larvae); the American dog tick, Dermacentor variabilis (Say) (one nymph); and Ixodes affinis Neumann (one larva). The Carolina wren was parasitized by more species of ticks (seven) than any other bird species, followed by the northern cardinal (five), white-throated sparrow (four) and painted bunting (three). Spirochetes were isolated in BSK II medium from one tick (a nymphal A. americanum) and from skin biopsies of 12 (4%) of the individual birds (three downy woodpeckers, three northern waterthrushes, two Carolina wrens, one American redstart, one pine warbler, one Swainson's thrush, and one white-eyed vireo) all in fall 1997. This concentrated phenology of spirochete isolations might reflect periodic amplification or recrudescence of spirochetes in reservoir avian hosts.

Animals↗

Investigation of relationships between temperature and developmental rates of tick Ixodes scapularis (Acari: Ixodidae) in the laboratory and field.

Relationships between temperature and preoviposition, preeclosion, and premolt developmental periods for the tick Ixodes scapularis Say were investigated by holding field-collected ticks in the laboratory at temperatures of 0 to 32 degrees C at constant daylength. The duration of these developmental periods decreased significantly with increasing temperature. Host of origin, prior storage at 4 degrees C, and season of collection of the ticks were also significantly associated with variations in the duration of the preoviposition period. For each developmental stage, the effect of temperature on development rate was best described as a power relationship. Laboratory-derived relationships were used to predict dates for molting, oviposition, and eclosion of engorged larvae and nymphs, engorged adult females and egg masses, respectively, placed in the field during 1989-1992. Predicted dates for oviposition by adult females, eclosion of eggs, and molting of engorged larvae were within 2 wk of the observed dates, and field-observed seasonal activity of questing larvae and nymphs also was predicted well by laboratory data. Molting of engorged nymphs and seasonal activity of questing adult ticks were, however, poorly predicted. Our findings suggest that duration of development in the field, of larvae from engorged adult females, and of nymphs from engorged larvae, may be explained largely by temperature effects alone, whereas emergence of adult I. scapularis from engorged nymphs may depend on temperature-independent diapause phenomena. The significance of these findings for understanding current and future distributions of I. scapularis, and of the pathogens it transmits, is discussed.

Animals↗

Life stage-related differences in density of questing ticks and infection with Borrelia burgdorferi sensu lato within a single cohort of Ixodes pacificus (Acari: Ixodidae).

The primary aims of this study were to quantify the density of Ixodes pacificus Cooley and Kohls nymphs and adults of the same generational cohort collected within a single year in six oak or madrone leaf litter habitats and to compare the prevalence of infection with Borrelia burgdorferi sensu lato (s.l.) in adults originating from nymphal cohorts with a low (<1%) versus high (>10%) infection prevalence. Because adult densities were very low both in and adjacent to several sites, direct comparisons of infection prevalence between nymphs and adults were possible only for two sites. Mean density in these sites decreased from 11.95/100 m2 for nymphs to 0/100 m2 for adults in leaf litter, and infection prevalence with B. burgdorferi s.l. was four-fold higher in nymphs (7.4%) versus adults (1.6%) of the same generational cohort collected in ecotones bordering the leaf litter areas. Assuming a density of adults in leaf litter of 0.04/100 m2 (mean for all six examined sites) and an infection prevalence similar to that found in adults collected from litter ecotones, the risk of encountering infected ticks in leaf litter decreased >1,000-fold from the nymphal to adult stage. Regardless of site-specific infection prevalence in the nymphal stage (n = 2 sites; 0.7 versus 14%), the infection prevalence for the adults of the same generational cohort was similarly low (1.5-1.6%). Peak densities of adult I. pacificus were 0-0.1/100 m2 in leaf litter, 0-6.5/100 m2 in ecotonal grasslands, and 2.0-39.0/100 m2 in ecotonal chaparral. Despite more intensive sampling efforts in leaf litter, the vast majority of the 282 adults collected came from grass or chaparral ecotones (98.9%, n = 279) rather than leaf litter (1.1%, n = 3). The study yielded eight B. burgdorferi s.l.-infected adults; four of these carried B. burgdorferi sensu stricto Johnson, Schmidt, Hyde, Steigerwalt, and Brenner, and the remaining four were infected with currently undescribed B. burgdorferi s.l. spirochetes. This is the first study comparing both density of I. pacificus nymphs and adults and prevalence of infection with B. burdorferi s.l in these ticks within the same generational cohort and sampling area.

Aging↗

Development cycle of Panstrongylus rufotuberculatus (Hemiptera: Reduviidae) under laboratory conditions.

Determinant aspects of the development cycle of Panstrongylus rufotuberculatus (Champion, 1899) were evaluated under laboratory conditions. Fertility, duration of life cycle, mortality rate, feeding time, defecation time, thermal reaction time to food source, and volume of blood ingested were studied. From an initial cohort of fifth instars, nine adult pairs were formed, the females of which oviposited 2,338 eggs over 231 d. The eclosion rate was 84.4%; female and male progeny lived 223.56 (14.41) and 180.78 (14.41) d, respectively. The average total life cycle of immature stages (138.82+/-51.42 d) was distributed as egg, 24.22+/-0.42 d; nymph I, 18.94+/-5.15 d; nymph II, 15.80+/-3.29 d; nymph III, 16.55+/-4.36 d; nymph IV, 25.50+/-7.32 d; and nymph V, 36.71+/-2.75 d. Given these values, 2.6 generations per year are possible. Because of its longevity, rapid response to the presence of a host, large volume of blood ingested, and frequent defecation during feeding, this species constitutes a potentially important vector of Chagas' disease in nonendemic rural areas.

Animals↗

Age structure of a population of Ixodes ricinus (Acari: Ixodidae) in relation to its seasonal questing.

Three sites in south west Scotland, each having one deciduous and one coniferous woodland with populations of roe deer, were sampled over three years for Ixodes ricinus Linnaeus ticks using cloth drags. Nymphs and adult ticks were age graded by dissection of gut and Malpighian tubules and staining for lipid with Sudan red. Temperature and relative humidity were recorded in developmental and questing microclimates. Two annual cohorts of both nymphs and adults were found. For both instars one cohort became apparent in early spring and the other in autumn. The timing of the cohorts of nymphs corresponded to peaks of numbers questing, but neither females nor males had any clear seasonal pattern of questing. The maximum life of questing nymphs and adults was three to four months. Nymphs and adults were found questing at all months of the year and at temperatures ranging from 3.5 degrees C to 29.3 degrees C. Questing of larvae in summer peaks of numbers was positively correlated with temperature and negatively correlated with relative humidity. These results are explained by a descriptive model which invokes behavioural and morphogenetic diapause. Tests of the model are discussed and it is proposed that the main adaptive advantage of the complex life cycle is to permit moulting and oviposition to occur at favourable summer temperatures in ticks that have engorged at any time of the year.

Age Factors↗

Hosts, seasonal occurrence and life cycle of Rhipicentor nuttalli (Acari: Ixodidae).

There are only two species worldwide within the genus Rhipicentor, namely Rhipicentor bicornis and Rhipicentor nuttalli and both occur only in Africa. Rhipicentor nuttali has a widespread distribution in South Africa and the present investigation was initiated to elucidate its host preference, seasonality and life cycle. Rock elephant shrews, Elephantulus myurus were examined for ticks at four localities in the Free State Province, one in Gauteng Province and two in Limpopo Province, South Africa. Cape elephant shrews, Elephantulus edwardii were examined at two places in the Western Cape Province, and a single specimen of the bushveld elephant shrew, Elephantulus intufi was examined in central Namibia. Small mammals of other species were also examined at two of these localities. The majority of E. myurus at two sites in the Free State, at the locality in Gauteng and both sites in Limpopo Province were infested with larvae and/or nymphs of R. nuttalli, while the single E. edwardii examined at one site in the Western Cape Province and the single E. intufi examined in Namibia were infested with nymphs of this tick. Not one of the other small animals was infested. Although larvae and nymphs of R. nuttalli were present on E. myurus throughout the year, the former were generally most numerous during the period March to September, and the latter during May to October. The preferred hosts of the adults are domestic dogs, leopards, Panthera pardus and South African hedgehogs, Atelerix frontalis. Adult females engorged on Atelerix frontalis in 16-32 days and, after a preoviposition period of 2-4 days, produced approximately 170,00 eggs during the following 60-70 days. The average incubation period of the eggs was 59 days. Larvae engorged on E. myurus in 4-10 days and moulted to nymphs 12-20 days later. Nymphs required 11-15 days to engorge on E. myurus and moulted to adults 32-47 days later. Allowing 14 days for the exoskeletons and mouthparts of each of the three parasitic stages to harden before they can attach to a host, the life cycle took approximately 214 days to complete in the laboratory. The length of this period, considered in conjunction with the times of maximum seasonal occurrence of the immature stages, indicates that the life cycle probably takes a year to complete in the field.

Animals↗