Field experiments on the effect of ticks on breeding success and chick health of cattle egrets

Abstract Two experiments compared broods that were naturally tick-infested with an equal number that were rendered tick-free by application of an acaricide to their nests and of barriers against further infestation. In the first experiment conducted in 1991–92 nestlings in tick-infested broods had up to 159 larval ticks at once and a mean infestation of 23.6 larval ticks per chick per day. The chick's mean tick load was inversely correlated with its longevity. Chicks that survived to at least 18 days post-hatching had significantly lower larval tick loads than those that died by 18 days, excluding the third-hatched chick in each brood whose survival rate was low irrespective of tick-infestation. At 7 days post-hatching, tick-infested chicks had a lower haematocrit and higher polychromasia than tick-free chicks. I infer that blood loss anaemia caused the deaths of tick-infested chicks in their first week. None died in their second week and the demise of those in their third week may have been due to the paralysis manifested prior to their death. I conclude that heavy tick infestation of their chicks reduced the breeding success of the parent egrets below that of the egrets whose chicks were kept free of ticks. In the second experiment in the 1992–93 season the mean level of tick infestation was much lower (5.1 per chick) and these chicks survived equally with tick-free chicks through fledging.     

The effect of Amblyomma variegatum on liveweight gain of cattle in Zambia

Abstract. The effect of the tick Amblyomma variegatum Fabricius on liveweight gain (LWG) on yearling cattle over 16 weeks was evaluated under quasi-natural conditions. Early in the exposure period when animals may have been in a state of suppressed metabolism, the effect of tick feeding on LWG was negligible. Overall, however, the difference in LWG between tick-free and tick-infested groups of cattle was 4.95 kg after 16 weeks. This difference between the groups was used to estimate a lower LWG of 48–63 g due to each engorging female tick.     

Seasonal patterns of abundance and host relationships of the Australian paralysis tick,Ixodes holocyclus Neumann (Acarina: Ixodidae), in southeastern Queensland

Estimates of seasonal abundance of larvae, nymphs and females of the paralysis tick, Ixodes holocyclus, were obtained by collecting ticks that engorged on small mammals and birds trapped in two localities in southeastern Queensland: Brisbane (wet sclerophyll forest) and Tamborine Mountain (cleared rain forest). The long-tailed short-nosed bandicoot, Isoodon macrourus, was the most common mammal trapped but small numbers of other marsupials, rodents and ground frequenting birds were also captured. Small numbers of five other tick species were also collected. In both habitats there was clearly one dominant generation of I. holocyclus;7er year, although the presence of all stages at most times of the year indicated overlapping of smaller cohorts. Females were most abundant in spring and early summer, larvae in summer-autumn, and nymphs in autumn-winter. I. holocyclus was abundant on I. macrourus and rare on most other mammals and birds captured. At the peak of abundance of each instar, each bandicoot dropped from 500 to 2000 engorged larvae, 100 to 200 engorged nymphs, and four to six engorged females. Life tables were compiled for the tick in both habitats and these indicate that there was relatively high survival from engorged larva to engorged nymph and thence to engorged female and that most mortality occurred between detaching of the engorged female and the detaching of the engorged larva. The tick was more abundant on bandicoots from cleared rain forest and rain forest edge, than on those from sclerophyll woodland. The survival of engorged larvae and nymphs of I. holocyclus and the larval productivity of engorged females were examined in a warm moist climate where the tick was abundant (Tamborine Mountain) and in a hotter dryer climate where the tick was rare or absent (Amberley). In both localities engorged larvae and nymphs survived to the next instar in all seasons of the year. On most occasions engorged females produced eggs which hatched. Mature bandicoots from tick infested areas showed little or no resistance to infestation with larvae or nymphs of I. holocyclus, whereas other small mammals from the same area showed an appreciable degree of resistance to the immature stages of the tick. Feeding larvae and nymphs exposed to normal light-dark cycles in the laboratory detached during the afternoon and early evening. This behaviour and host resistance are discussed in relation to the daily activity cycles of host species, their habitat preferences, and their role as hosts for I. holocyclus.     

Testosterone depresses innate and acquired resistance to ticks in natural rodent hosts: a force for aggregated distributions of parasites

The effects of testosterone on acquired resistance to ticks, Ixodes ricinus, in their natural rodent hosts (voles, Clethrionomys glareolus, and wood-mice, Apodemus sylvaticus) were investigated by manipulating testosterone levels and exposing the hosts to repeated tick infestations. Testosterone reduced both innate and acquired resistance to tick feeding. During primary infestations, attachment rates were higher on rodents with high testosterone levels than on oil-implanted controls. Successive infestations on voles were accompanied by a decrease in tick feeding success and survival, but this decrease was significantly greater in ticks fed on control voles than in those fed on voles implanted with testosterone. When reduced feeding success had been induced, either by vaccination with tick salivary gland extract or by 4 successive infestations, implantation with testosterone partially reversed the acquired resistance. These effects of testosterone will generate heterogeneities within the rodent population with respect to tick distribution and microparasite transmission. The lowest innate and acquired resistance to tick feeding occurs in that fraction of the host population, i.e., sexually active males, most actively involved in the transmission of both Babesia microti and Borrelia burgdorferi s.l.     

Studies on the economics of ticks in Zambia

Two herds of experimental Sanga cattle were maintained under traditional savanna grazing management for three years in the Central Province of Zambia. One herd was kept free of ticks by regular acaricide treatment, while the other was given no tick control. Milk production, growth rate, fertility and mortality were monitored throughout the trial. The tick-free herd performed significantly better than the tick-infested herd, but the value of the additional production was much less than the cost of the acaricide used. it is concluded that there is no economic justification for intensive tick control under these conditions. However, strategic tick control would be justified if the quantity of acaricide used could be reduced by 50% without any major reduction in benefits. The results indicate that treating calves below 45 days of age reduces their performance, as does treating cows during periods of very low tick challenge. Therefore, it seems likely that economically beneficial strategic tick control policies could be developed.     

Ixodes scapularis: effects of repeated infestations with pathogen-free nymphs on macrophage and T lymphocyte cytokine responses of BALB/c and C3H/HeN mice.

Schoeler, G. B., Manweiler, S. A., and Wikel, S. K. 1999. Ixodes scapularis: Effects of repeated infestations with pathogen-free nymphs on macrophage and T lymphocyte cytokine responses of BALB/c and C3H/HeN mice. Experimental Parasitology 92, 239-248. Ixodes scapularis is the principal vector in the United States of Borrelia burgdorferi, the causative agent of Lyme borreliosis, the human granulocytic ehrichiosis agent, and Babesia microti. Infestation with I. scapularis nymphs has previously been shown to modulate host T lymphocyte cytokine production. Tick-induced host immunomodulation is increasingly recognized as a contributing factor in successful transmission and/or establishment of tick-borne pathogens. This study was conducted to determine the effects of repeated infestations with pathogen-free I. scapularis nymphs on the production of the macrophage cytokines interleukin (IL)-1beta and tumor necrosis factor-alpha and the T lymphocyte cytokines IL-2, IL-4, IL-10, and interferon-gamma in both BALB/c and C3H/HeN mice. The pattern of T lymphocyte cytokine production was evaluated to determine if repeated tick infestation polarizes the immune response toward a Th-1 or Th-2 cytokine profile. Female BALB/c and C3H/HeN mice were infested one to four times with pathogen-free I. scapularis nymphs, with a 14-day tick-free period between each exposure. After each infestation, tick biology parameters were measured and macrophage and T lymphocyte cytokine production was assessed. Elaboration of T lymphocyte and macrophage cytokines was quantitated by antigen capture enzyme-linked immunosorbent assay. Acquired resistance to I. scapularis feeding was not developed by either mouse strain. Significant differences in cytokine production were observed between infested and noninfested mice, as well as between the two mouse strains, following tick infestation. Infestation of both strains with pathogen-free I. scapularis results in a polarization of the host immune response toward a Th-2, anti-inflammatory pattern, with a corresponding suppression of Th-1 responses.     

Modeling the Impact of Climate and Landscape on the Efficacy of White Tailed Deer Vaccination for Cattle Tick Control in Northeastern Mexico

Cattle ticks are distributed worldwide and affect animal health and livestock production. White tailed deer (WTD) sustain and spread cattle tick populations. The aim of this study was to model the efficacy of anti-tick vaccination of WTD to control tick infestations in the absence of cattle vaccination in a territory where both host species coexist and sustain cattle tick populations. Agent-based models that included land cover/landscape properties (patch size, distances to patches) and climatic conditions were built in a GIS environment to simulate WTD vaccine effectiveness under conditions where unvaccinated cattle shared the landscape. Published and validated information on tick life cycle was used to build models describing tick mortality and developmental rates. Data from simulations were applied to a large territory in northeastern Mexico where cattle ticks are endemic and WTD and cattle share substantial portions of the habitat. WTD movements were simulated together with tick population dynamics considering the actual landscape and climatic features. The size of the vegetation patches and the distance between patches were critical for the successful control of tick infestations after WTD vaccination. The presence of well-connected, large vegetation patches proved essential for tick control, since the tick could persist in areas of highly fragmented habitat. The continued application of one yearly vaccination on days 1-70 for three years reduced tick abundance/animal/patch by a factor of 40 and 60 for R. annulatus and R. microplus, respectively when compared to non-vaccinated controls. The study showed that vaccination of WTD alone during three consecutive years could result in the reduction of cattle tick populations in northeastern Mexico. Furthermore, the results of the simulations suggested the possibility of using vaccines to prevent the spread and thus the re-introduction of cattle ticks into tick-free areas.     

Immunosuppression and feeding success of Ixodes ricinus nymphs on BALB/c mice

Abstract. The effect of repeated infestations of Ixodes ricinus (L.) nymphs on BALB/c mice was studied. Four successive infectations resulted in an increase of tick feeding success. Tick yield and mean engorged weight increased and the length of the feeding period was reduced significantly (P <0.05-0.01). The increase of specific anti-tick antibodies was not significant (P > 0.05). The blastogenic response of spleen lymphocytes to T-cell mitogens (Con A and PHA-P) was unimpaired or slightly enhanced, whereas the response to B-cell activators (LPS and PWM) was suppressed, as was the total antibody generation in vitro. The numbers of mast cells in murine skin at the tick attachment sites slightly decreased during the third infestation. The suppression of B-cell competence and of antibody generation, together with decrease of skin mast cell numbers in tick attachment sites, are considered to be responsible for enhancement of tick feeding success.     

Prevalence of Borrelia burgdorferi (the Lyme disease agent) antibodies in king penguin Aptenodytes patagonicus in Crozet Archipelago

The seabird tick Ixodes uriae (White) is common in subantarctic areas and is likely to be a vector of pathogenic agents. This paper reports on the prevalence of tick-infested adult king penguins (Aptenodytes patagonicus) and on antibodies to the Lyme disease agent (Borrelia burgdorferi sensu lato) in adult and chick penguins. Fifteen percent of the adults of the colony studied were tick infested. B. burgdorferi antibodies were detected in 14% of tick-infested adults' and in 6% of chicks' sera. The percentage of tick-infested adults differed significantly according to location in the colony, the highest density occurring on the upper slopes of the colony. The potential role of king penguins as B. burgdorferi reservoirs and the effect of this agent on king penguin populations remain unknown. The consequences for human health are discussed. Accepted: 15 March 1999     

Resistance and cross-resistance in guinea-pigs and rabbits to immature stages of ixodid ticks

Infestation of guinea-pigs and rabbits with larvae of any one of five species of ticks, Rhipicephalus appendiculatus Neumann, Rhipicephalus evertsi evertsi Neumann, Amblyomma hebrauem Koch, Amblyomma variegatum Fabricius and Ixodes ricinus L., conferred resistance in the animals when exposed to subsequent infestations with the same tick species. Resistance to infestations by other tick species was not observed.     

Population regulation in ticks: the role of acquired resistance in natural and unnatural hosts

Attachment, engorgement and subsequent development of successive infestations of Ixodes trianguliceps larvae and nymphs on natural hosts, Apodemus sylvaticus, and unnatural hosts, laboratory mice, are compared. In laboratory mice, primary infestations above a threshold level of about 10 ticks elicit an immunological response which reduces, in a density-dependent manner, the rate of successful tick engorgement during subsequent infestations. In contrast, in A. sylvaticus successive infestations of larvae result in unchanged or slightly improved survival through to nymphs. The relevance of these results to the concept of host-parasite co-evolution and to tick population regulation is discussed.     

Populations of the ticks Ixodes (Pholeoixodes) hexagonus and Ixodes (Pholeoixodes) canisuga infesting suburban foxes, Vulpes vulpes

Only two species of tick (Ixodes hexagonus and Ixodes canisuga) were found to infest suburban foxes. The populations of these two ticks were examined, their distributions within the host population described, and infestation levels of I hexagonus discussed in relation to the sex, age and behaviour of the host. The most important factor regulating the level of tick infestation is probably the degree of den usage by the host. The tick infestations were found to have minimal effect on the host, and even an abnormally high level of infestation found on one fox was not considered to be lethal.     

Lack of resistance against the tick Ixodes ricinus in two related passerine bird species

Although many wild bird species may act as reservoir hosts for tick-transmitted diseases and/or support long-distance dispersal of infected ticks, to date no research has been done on the extent to which songbirds may acquire resistance to ixodid ticks. Here we investigate whether two passerine species belonging to the family Paridae, the blue tit (Cyanistes caeruleus) and the great tit (Parus major), are able to acquire resistance after repeated infestations with Ixodes ricinus nymphs. As blue tits are less frequently exposed to I. ricinus in the wild than great tits, we expected I. ricinus to be less adapted towards the blue tit’s resistance mechanisms. Over the three infestation sessions we observed consistently high tick attachment rates and yields, high engorgement weights, and short engorgement and moulting durations, indicating that neither of the two songbird species is able to mount effective immune responses against I. ricinus nymphs after repeated infestations. As a consequence of the lack of resistance, birds were unable to prevent the direct harm (acute blood depletion) caused by tick feeding. Birds compensated the erythrocyte loss without reduction in general body condition (body mass corrected for tarsus length). The lack of resistance suggests that I. ricinus has a long co-evolutionary history with both avian hosts, which enables the tick to avoid or suppress the host’s resistance responses.     

Can acaricide‐impregnated leg bands fitted to female red grouse reduce sheep tick parasitization of chicks and increase chick survival?

In parts of northern England, North Wales and the Scottish Highlands, increasing numbers of sheep ticks Ixodes ricinus (Ixodida: Ixodidae), and the louping ill virus they can carry, are considered to be important factors that reduce red grouse Lagopus lagopus scotica productivity. The present study tested this hypothesis by fitting adult female grouse with leg bands impregnated with the acaricide cypermethrin to experimentally control ticks on their chicks on two managed grouse moors in northeast Scotland. The chicks of females fitted with acaricide leg bands showed reduced tick infestations and improved survival in one of the two study years, relative to chicks of control females. Acaricide leg bands constitute a potential management technique that may be adopted by grouse moor managers in circumstances of high tick infestations on grouse chicks.     

Reduced Weight Gain Due to Subclinical Anaplasma phagocytophilum (Formerly Ehrlichia phagocytophila) Infection

Tick-borne fever (TBF) is caused by the rickettsia Anaplasma phagocytophilum (formerly Ehrlichia phagocytophila) and is a common disease in sheep in areas of Norway infested by Ixodes ricinus ticks. TBF can cause both direct and indirect losses to sheep kept on tick-infested pastures. In the present work we studied a sheep flock of 26 ewes and 50 lambs on pasture from May until September. No cases of TBF had earlier been observed on this pasture. Blood samples from lambs with a reduced weekly weight gain were collected and analysed for A. phagocytophilum infection by blood smear examination. In addition, at the end of the study, sera from all lambs were analysed by an indirect fluorescent antibody assay (IFA) to determine the antibody titre to E. equi. No clinical signs of tick-borne infections were observed, except in one lamb. However, 30 (60%) of the lambs grazing on this pasture became infected with A. phagocytophilum, and the infected lambs had a reduced weight gain (mean) of 3.8 kg compared with the uninfected lambs. The present study indicates that A. phagocytophilum infection may be widespread and contribute to considerable productivity losses even on pastures with no apparent tick infestation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Increased mortality of black-browed albatross chicks at a colony heavily-infested with the tick Ixodes uriae

At Bird Island, South Georgia, we studied the effects of the tick Ixodes uriae on survival of chicks at two colonies of the black-browed albatross Diomedea melanophrys, one where most chicks were infested with ticks, the other where most chicks were tick-free. When the two colonies were compared, it was found that the colony heavily-infested with ticks had significantly greater chick mortality than the colony lightly-infested with ticks. However, within each of the two colonies, there was no significant difference in survival between chicks with ticks and those without ticks.     

A comparison of infestation patterns by Ixodes ticks in urban and rural populations of the Common Blackbird Turdus merula

Although spatial variation in the patterns of parasite infestations among host populations may have important ecological and epidemiological consequences, the causes underlying such variation are poorly known. In the context of a long-term study on the population biology of Common Blackbirds Turdus merula , we examined the prevalence and intensity of infestation by Ixodes ticks between birds living in rural vs. urban habitats. The overall prevalence of tick infestations was significantly higher in the rural habitat where 74% of individuals ( n  = 130) were infested. This result contrasted markedly with the situation in the urban habitat where less than 2% of individuals ( n  = 360) carried ticks. There was no significant effect of the sex of the host on the intensity or prevalence of tick infestations. There was a significant effect of the age of the host on tick infestations essentially due to the absence of ticks on nestlings. Possible mechanisms responsible for the differences between habitats could include differences in tick survival and/or host resistance towards ticks. Previous studies have shown higher population densities and suggested longer survival for Blackbirds in urban than in rural habitats. Given that ixodid ticks are known to transmit pathogens like Borrelia spp. to wild birds, and that Blackbirds can act as reservoirs for these pathogens, the infection patterns observed in our study area provide a suitable situation to study the interrelations between ticks, Blackbirds and pathogens.     

The impact of tick parasites on the behaviour of the lizard Tiliqua rugosa

Populations of the Australian sleepy lizard, Tiliqua rugosa, near Mt. Mary, South Australia carry natural infestations of two tick species Aponomma hydrosauri and Amblyomma limbatum. In field experiments at two sites, 18 km apart, lizards with experimentally increased tick loads had smaller home ranges, moved shorter distances in a day, and were found basking more but moving less often than lizards from which ticks were experimentally removed. The results were consistent for adult lizards in two years, and for sub-adults in a third year. Laboratory trials showed that juvenile lizards that had tick infestations had lower sprint speeds than uninfested siblings, and that adults with tick infestations had less endurance than those that were uninfested. The results contrast with those of a previous survey that showed that lizards with high tick loads had greater body size and remained longer at a site, but indicate that there may be a balance, for lizards, between the fitness advantages in occupying habitats with high-quality resources, and the costs from parasites that also prefer those habitats. Received: 02 March 1999 / Accepted: 07 October 1999