The natural hosts of larvae and nymphs of Amblyomma tigrinum Koch, 1844 (Acari: Ixodidae)

The hosts of larvae and nymphs of Amblyomma tigrinum, a tick whose adults feed on wild and domestic Canidae in South America, are uncertain. A 17 months survey was carried out trapping wild vertebrates in north-western Córdoba, Argentina, to evaluate their parasitism with A. tigrinum subadults. Larvae and nymphs of this tick species were identified conventionally and by comparison of 16S rDNA sequences with GenBank deposited sequences. A total of 207 small and medium-sized rodents and 182 birds were captured and examined for ticks. Most ticks on birds were from ground forest feeding birds (BB) with a minimal contribution of birds feeding in open pastures. All ticks from rodents were obtained from representatives of the families Cricetidae (SR) and Caviidae (MR). Percent of larvae infestation was higher (P<0.01, Chi-square distribution) in BB (55.2%) and SR (46.4%) than in MR (17.4%) and the same trend was found for number of larvae on these hosts (test of Kruskal-Wallis). Caviidae (only representative Galea musteloides) rodents were extremely prone to be infested with nymphs of A. tigrinum (94.2%) followed by BB (50.6%) and SR (3.6%) (P<0.01) and the same tendency was found for number of nymphs (P<0.01). The index of aggregation for nymphs on MR was the lowest (0.409) followed by nymphs on BB (0.706) which may be a consequence of higher and homogenous exposure of G. musteloides to host-seeking nymphs. Several BB are food source for both larvae and nymphs of A. tigrinum while for rodents larvae were common only on SR (mainly on the Sigmodontinae Akodon dolores and Graomys sp.) and nymphs feed almost exclusively on MR. Therefore, both birds and rodents are of importance for the survival strategy of A. tigrinum subadults. The plasticity of A. tigrinum to colonize areas with different climates plus the capacity of their subadults to feed on hosts widely distributed indicates that this tick has the potential to become a widespread parasite but this does not seem to be the actual situation. Several proposals are presented to further understand its ecology.     

Cutaneous hypersensitivity induced in rabbits by extracts of the tick Amblyomma cajennense (Acari: Ixodidae)

The cutaneous hypersensitivity test was used to correlate host resistance to ticks and type of reactions elicited by Aomblyomma cajennense (Fabricius, 1787) tick extract in rabbits. Rabbits were divided into 3 groups of 2 animals each: naive, pre-infested and control. Cutaneous hypersensitivity was induced by intradermal inoculation of 25 microg extract in 0.03 ml of phosphate buffered saline (PBS) in rabbit ears. Control rabbits were inoculated with PBS only. The ear thickness was measured with a Mitutoyo device before and 10 min, 1, 2, 4, 18, 24, 48, 72 and 96 h post-inoculation (PI). Pre-infested rabbits showed an immediate type reaction within the 1st 10 min PI (60% increase in ear thickness) and a delayed reaction (18 h) (85% increase), whereas the naive rabbits showed only the immediate reaction within the 1st 4 h (60% increase). PBS induced only mild reactions. These results point out the crucial role of the cellular immune response of rabbits in the expression of resistance to A. cajennense.     

Morphological characterization of the ovary and vitellogenesis dynamics in the tick Amblyomma cajennense (Acari: Ixodidae)

In this work we describe the internal morphology of the female reproductive system of the cayenne tick Amblyomma cajennense. This system is represented by a panoistic ovary, which lacks nurse cells in the germarium. This ovary consists of a single tube, in which a large number of oocytes develop asynchronously, thus accompanying the processes of yolk deposition in the oocytes. The oocytes were classified into stages that varied from I to V, according to: cytoplasm appearance, presence of the germ vesicle, presence of yolk granules, and presence of chorion. The study of vitellogenesis dynamics suggest that the yolk elements are deposited in the oocyte following a preferencial sequence, in which the lipids are the first to appear, followed by proteins an finally by the carbohydrates. In this way the yolk of A. cajennense ticks have these three elements that may be free in the cytoplasm or chemically bounded forming glycoprotein or lipoprotein complexes.     

Horse resistance to natural infestations of Anocentor nitens and Amblyomma cajennense (Acari: Ixodidae).

The objective of this study was to investigate some aspects of horse resistance to natural infestations of Anocentor nitens and Amblyomma cajennense over a 2-year period. Free-living stages of A. nitens were used to evaluate the influence of season of the year on horse resistance. Every 2 weeks, 10 selected engorged females, weighing 150-329 mg, were removed from naturally infested horses and were individually placed into glass tubes closed with a cotton stopper, and maintained in an incubator. The biological parameters evaluated varied according to the season of the year, i.e., reproductive efficiency, maximum larval survival (MLS) and egg-hatching rates were significantly lower in spring/summer than in fall/winter. At 14-day intervals, females of A. nitens and A. cajennense were counted on each side of the body of five mares. A comparison between the animal with the lowest infestation and the animal with the highest infestation showed greater differences in animals infested with A. nitens than those infested with A. cajennense. Ten animals were found to carry 64.9% of the total parasite burden of A. cajennense and 88.6% of the highest infestations of A. nitens females. A seasonal variation on intensity of horse resistance to A. nitens was observed during the study. However, this was not observed for A. cajennense. Cross-resistance was observed in animals with the highest total number of A. cajennense and the highest number of A. cajennense adults (r=0.817), larvae (r=0.756), and A. nitens females (r=0.799). These results led to the conclusion that changes in horse resistance to A. nitens occur seasonally. As far as the total population of A. cajennense and the female stage of A. nitens are concerned, horses develop cross-resistance.     

The sex ratio of Amblyomma cajennense (Acari: Ixodidae) with notes on the male feeding period in the laboratory

To evaluate the sex ratio of field collected nymphal Amblyomma cajennense ticks, we collected 5326 engorged nymphs from naturally infested horses in Pirassununga county and allowed them to molt to adults in the laboratory. They yielded a sex ratio of 1:1.83 (M:F). Three and two engorged females were collected from horses pastured at Pirassununga county and from tapirs pastured in Sorocaba county, respectively. These females were allowed to oviposit and their progeny were reared until the adult stage in the laboratory. Engorged females collected from Pirassununga yielded a sex ratio of 1:1.57 (M:F) and a sex ratio of 1.14:1 (M:F) were obtained for those ticks collected from tapirs. In addition, unfed tick larvae were collected from Pedreira county and reared in the laboratory until the adult stage. This collection yielded a sex ratio of 1.11:1 (M:F). These results showed significantly different (P<0.05) sex ratio constitutions among different tick populations. Laboratory rabbits were infested once with A. cajennense male ticks, which showed feeding periods varying from 7 to 86 days. During this period, the rabbits were re-infested regularly with A. cajennense female ticks. A total of 179 engorged females were collected from the rabbits and their engorged weight, feeding, preovioposition and egg incubation periods, weight of deposited eggs, percent of hatched eggs and egg production efficiency were compared to the male feeding period and to the number of live males present on the host. None of the female variables were affected by the male feeding period. Male ticks remained fertile for the whole feeding period. Percent of hatched eggs was the only female variable that significantly decreased as the number of live males decreased on the host. The results showed that although some A. cajennense populations are composed of more females than males after molting, this female predominance is compensated by a long male feeding period and maintenance of its reproductive performance.     

Repellent activity of plant-derived compounds against Amblyomma cajennense (Acari: Ixodidae) nymphs

Repellence responses of Amblyomma cajennense nymphs to callicarpenal, intermedeol, Hyptis suaveolens essential oil, extract of Melia azedarach, Cymbopogon nardus, Spiranthera odoratissima, Chenopodium ambrosioides, Ageratum conyzoides, Mentha pulegium, Ruta graveolens, and Memora nodosa were studied. Among these the extract of C. nardus stood out because of the long-lasting repellence, maintaining, in the highest concentration, 35 h of protection against 90% of the nymphs. The essential oil of H. suaveolens and the extracts of C. ambrosioides and A. conyzoides showed good repellence index (66%) when applied in high concentrations. However, greater protection could be obtained at higher concentrations but with a shorter repellence time. Callicarpenal, intermedeol, extract of M. Pulegium, and M. nodosa leaves showed moderate repellence in high concentrations. Extracts from M. azedarach, R. graveolens, S. odoratissima, and M. nodosa roots showed little or no repellent effect. These results show that some plant extracts may represent a promising alternative in the control of infestations by A. cajennense.     

Acaricidal activity of an oleoresinous extract from Copaifera reticulata (Leguminosae: Caesalpinioideae) against larvae of the southern cattle tick, Rhipicephalus (Boophilus) microplus (Acari: Ixodidae)

The acaricidal activity of oleoresinous extract (oleoresin) from the copaiba tree, Copaifera reticulata was investigated. Oleoresin was dissolved with dimethylsulfoxide (DMSO) and mixed with distilled water at a rate of 0.4 ml of DMSO to 24.6 ml water. Rhipicephalus (Boophilus) microplus larvae were exposed to filter paper envelopes impregnated with different oleoresin concentrations. Four envelopes were impregnated with each tested solution. Thirty larvae between 14 and 21 days old were fasted and placed in each envelope. Bioassays were performed at 27 degrees+/-1 degrees C, RH>or=80% and a photoperiod of 12:12 h. Larval mortality was observed 24 h after treatment. Four replicates for each concentration were evaluated to determine lethal concentrations (LC). Oleoresin LC50 and LC99 values were 1579 and 3491 ppm, respectively. The potential use of oleoresin for control of R. (B.) microplus is discussed. The results of this study reinforce the importance of the preservation of C. reticulata in its natural Biome, as a potentially auto-sustainable medicinal resource among the flora of Amazonian forest.     

Acaricidal activity of plant essential oils against Dermanyssus gallinae (Acari: Dermanyssidae)

The acaricial activity of 56 plant essential oils against poultry house-collected adult Dermanyssus gallinae De Geer was examined using direct contact and fumigation methods. In a filter paper contact bioassay, 100% mortality at 0.07 mg cm⁻² was observed in bay, cade, cinnamon, clove bud, coriander, horseradish, lime dis 5F, mustard, pennyroyal, pimento berry, spearmint, thyme red and thyme white oils, whereas the mortality of these oils was significantly decreased at 0.02 mg cm⁻². In fumigation tests with adult D. gallinae at 0.28 mg cm⁻², cade, clove bud, coriander, horseradish and mustard oils were more effective in closed containers than in open ones, indicating that the effect of these essential oils was largely due to action in the vapour phase. Plant essential oils described herein merit further study as potential D. gallinae control agents.     

Vitellogenesis in the tick Amblyomma triste (Koch, 1844) (Acari: Ixodidae) Role for pedicel cells

This study presents new information on the vitellogenesis of the tick Amblyomma triste. In this species, the ovary consists of a layer of epithelial cells, which form the ovarian wall, oogonia and developing oocytes; and the pedicel, a cellular structure that synthesizes and provides yolk precursors for developing oocytes. The pedicel also attaches oocytes to the external surface of the epithelial wall. In this study, evidence is provided in support of pedicel cells in providing compounds for oocytes during vitellogenesis.     

Amblyomma triste Koch, 1844 (Acari: Ixodidae): hosts and seasonality of the vector of Rickettsia parkeri in Uruguay

Amblyomma triste Koch, 1844, the tick species most frequently involved in human bites in Uruguay, has been implicated as a vector of human rickettsiosis. Seasonal abundance of adult A. triste was examined by standard flagging at three sites where human tick bites and cases of the disease have been reported. Adult tick activity occurred from August to February (end of winter to mid summer in the southern hemisphere) with a peak in spring. Activity declined in step with decreasing temperatures and photoperiod during winter. This period of activity coincides with seasonal outbreaks of human rickettsiosis in the region. In a small mammal survey, the Sigmodontinae rodents Scapteromys tumidus (Waterhouse, 1837) and Oxymycterus nasutus (Waterhouse, 1837) and the small marsupial Monodelphis dimidiata (Wagner, 1847) were the main hosts for immature A. triste. Immature ticks were observed on hosts in November, well within the period of peak adult abundance. In stored collections, immature ticks were most often collected from January to March. These data suggest that one generation might be completed in 1 year. The main animal host for adult A. triste at our study sites was the domestic dog. Humans were afflicted by the tick in rural and suburban settlements where other host animals are scarce or extinct and where dogs are common.     

Acquired resistance of horses to Amblyomma cajennense (Fabricius, 1787) ticks

Acquired immunity of horses to larvae, nymphs and adults of the Amblyomma cajennense tick was evaluated through three consecutive experimental infestations of tick-bite na?ve hosts. Data from these infestations were compared to those from field-sensitized horses and donkeys. It was observed that tick-bite na?ve horses developed a low level of resistance after two infestations as shown by a significant decrease in larval yield and a tendency for lower engorged weight of nymphs during third infestation. Ticks fed on field-sensitized horses had a similar biological performance to that observed on the third infestation of tick-bite na?ve horses but the mean engorged nymph weight was significantly lower than that of the first infestation from tick-bite na?ve horses. Donkeys presented the strongest resistance with significantly lower engorged weights of all instars and of the egg mass compared to the first infestation of tick-bite na?ve horses. Donkeys also displayed a significantly higher resistance than field-sensitized horses as demonstrated by significantly lower egg mass weights. Overall these results indicate that donkeys but not horses maintain a strong resistance to A. cajennense ticks. The importance of these findings in relation to vectoring of tick-borne diseases is discussed.     

Acaricidal effects of the essential oil of Origanum minutiflorum (Lamiaceae) against Rhipicephalus turanicus (Acari: Ixodidae)

The acaricidal effects of the volatile essential oil Origanum minutiflorum O. Schwarz & P.H. Davis (Lamiaceae) against adult Rhipicephalus turanicus was evaluated at a variety of concentrations and exposure times. Generally tick mortality increased with concentration and exposure. Ticks exposed to vapors from cotton wicks containing at least 10 μl/L resulted in complete (100%) mortality at 120 min. The major constituent of essential oil obtained from the plant material of O. minutiflorum was carvacrol.     

Population dynamics of the free-living stages of Amblyomma cajennense (Fabricius, 1787) (Acari: ixodidae) on pastures of Pedro Leopoldo, Minas Gerais State, Brazil

A study on the population behavior of Amblyomma cajennense larvae, nymphs and adult free-living stages was carried out in Pedro Leopoldo County, Minas Gerais State, Brazil, from July 1995 to July 1997. Larvae were collected from pasture from April to October using the technique of drag sampling with flannelette. The maximal larval population was observed during autumn (May) representing, respectively, 48 and 82% of the total numbers recovered during the first and the second years of the experiment. Nymphs and adults were captured throughout the year by the use of traps with carbon dioxide as chemical attractant. The highest population of nymphs occurred during the winter (July), representing 39.5% of the total collected specimens. Adults had the highest population density on pasture during spring and summer, i.e., from August to May, with peaks in January and February, representing 46 and 38.3% of the total collected specimens during the first and the second year of study, respectively.     

Acaricidal properties of two extracts from Guiera senegalensis J.F. Gmel. (Combrataceae) against Hyalomma anatolicum (Acari: Ixodidae)

Laboratory test were carried out on eggs, larvae, nymphs and adults of Hyalomma anatolicum to determine the acaricidal activities of petroleum ether (PE) and crude ethanolic extracts (EE) from the leaves of Guiera senegalensis J.F. Gmel. (Combrataceae) using immersion method.Stock solutions, of 300 mg/ml (30%) of the each two extract, were prepared. Six two-fold serial dilutions each with three replicates were used. Both extracts, at the highest concentration 150 mg/ml (15%), induced 100% failure of hatching of the treated eggs. The concentrations of PE and EE that induced 50% inhibition of the hatchability (IC₅₀) were 1.71 and 0.508%, respectively. In the larval immersion test (LIT), EE at 15% concentration caused complete mortality while the same concentration of PE resulted in 96% mortality. The mortalities increased with concentrations. There was a correlation between the mortalities and increased concentrations, the values of the linear correlation coefficient (r) for PE and EE were 0.93 and 0.79, and The LC₅₀ and LC₉₉ were 2.08 and 14.09, and 0.787 and 11.054, respectively. At the concentrations of 3.75%, 7.5% and 15%, PE inhibited the molting of the nymphs by 40, 55 and 65%, respectively, while EE induced 46.49, 64.3 and 71.4% inhibition, respectively. The effectiveness of the treatment against unfed adult females was assessed by measuring the feeding performance and egg production using adult immersion test (AIT). Although, there was no mortality in unfed adults, PE and EE inhibited feeding and egg-laying of the survived females by 35–100% and 6.16–100%, respectively. Our results indicated that G. senegalensis is a promising biocontrol candidate as an acaricidal agent against H. anatolicum.     

Efficacy of two 65% permethrin spot-on formulations against induced infestations of Ctenocephalides felis (Insecta: Siphonaptera) and Amblyomma americanum (Acari: Ixodidae) on beagles

The efficacy of two formulations of a topically applied 65% permethrin spot-on (Defend Exspot Treatment for Dogs, Schering-Plough Animal Health) was evaluated against experimental infestations of the cat flea Ctenocephalides felis and the lone star tick Amblyomma americanum in dogs. Eighteen dogs were randomly assigned to treatment with 65% permethrin in either diethylene glycol monomethyl ether (DGME; original formulation) or propylene glycol monomethyl ether (PGME) or to be untreated as a control. Treated dogs received either 1 (body weight < 15 kg) or 2 ml (body weight > or =15 kg) of the assigned formulation on Day 0. One hundred unfed, adult C. felis were placed on each dog on Days -6, -1, 4, 11, 18, 25, and 32. Fifty unfed, adult ticks were placed on each dog on Days -1, 3, 9, 16, 23, and 30. Live fleas and ticks were counted and removed on Days 3, 7, 14, 21, and 28. Treatment of dogs with the 65% permethrin in DGME reduced flea numbers by 90.4% to 99.9% from Days 3 through 21 (P < or =.05) and by 48.2% 28 days after treatment. Treatment of dogs with 65% permethrin in PGME reduced flea numbers by 93.7% to 99.7% from Days 3 through 28 and by 78.4% 35 days after treatment (P < or =.05). Treatment with 65% permethrin in DGME reduced tick numbers by 90% or more only on Day 7, whereas treatment with 65% permethrin in PGME reduced the number of live ticks by 90%or more on Days 7 and 14 and approached 90%(87.9%) on Day 21. Efficacy against fleas and ticks for the PGME formulation was significantly better (P < or =.05) than for the DGME formulation on Day 28. Findings in this study indicate that both the DGME and PGME formulations of 65% permethrin performed well in reducing numbers of live C. felis and A. americanum on laboratory beagles; however, the PGME formulation was effective approximately 1 to 2 weeks longer than the DGME formulation.     

Larval Gulf Coast ticks (Amblyomma maculatum) [Acari: Ixodidae] as host for Hepatozoon americanum [Apicomplexa: Adeleorina].

Laboratory-reared larval Gulf Coast ticks (GCTs) (Amblyomma maculatum) were exposed experimentally and found to acquire Hepatozoon americanum infection while feeding on parasitemic dogs. These ticks supported gamogonic and sporogonic development of the apicomplexan, and oocysts from newly molted nymphs were infectious for a dog. Other nymphs from this cohort that were allowed to feed on a blood-parasite naive sheep molted normally; the resulting adult ticks contained oocysts that were infectious for another dog. Merogonic development of H. americanum in the dogs and the resulting lesions/disease appeared similar, irrespective of whether infectious oocysts were derived from nymphal or adult ticks that acquired infection as larvae. In the system previously known, nymphal ticks acquire infection and adults harbor infective oocysts, which vertebrate hosts ingest. Given that larval A. maculatum can acquire infection and nymphs can harbor viable oocysts as demonstrated by this study, the potential variety of vertebrate hosts that can alternate with GCTs in maintaining an endemic cycle is considerably expanded.     

Climate changes and suitability for the ticks Amblyomma hebraeum and Amblyomma variegatum (Ixodidae) in Zimbabwe (1974-1999)

The spread of Amblyomma hebraeum has been reported in Zimbabwe. At the same time there was little or no spread in the distribution of Amblyomma variegatum. This paper examines the climatic cycles and their trends in the period 1974-1999 with a view to explaining the abiotic causes of this spread, and of forecasting the likely tendency in climate suitability for both tick species. An annual data-set of rainfall and air temperature was used as a source for climate, together with a habitat-modeling algorithm to estimate climate suitability for both ticks. Long-term suitable habitat for A. hebraeum exists mainly in the south and southeast of the country. Areas of adequate habitat for A. variegatum exist across the country, between approximately 17 degrees S and 18.5 degrees S, and are most suitable in regions of the northwest. The climate niches of the two species differ, and account for their almost allopatric distributions, as observed in the duration and intensity of the dry period and in total annual rainfall. Cyclic changes in both temperature and rainfall drive the periodic modifications in the distributions of the ticks. More intense periods of drought in the highveld, drive the expansion of A. hebraeum in this region. Temperature does not have any effect on the tendency in this area. Areas in south and southeast show a trend towards an increase in climatic suitability because of an increase in temperature. Zones in which habitat suitability is increasing for A. variegatum are restricted to the northwestern parts of the country, because warmer temperatures and a slight decrease in the intensity of the dry season. The progressive increase in temperatures seems to be forcing the dispersion of A. variegatum towards areas outside of zones that have a prolonged dry period. On the other hand A. hebraeum is compelled to spread northwards, following areas with adequate rainfall patterns, but halted by temperature limits and perhaps competition with A. variegatum. Without adequate control measures, invasive waves of A. hebraeum may occur over a background swell of northward expansion.     

Preliminary survey of ticks (Acari: Ixodidae) on cattle in northern Sudan

In a cross sectional survey conducted during the period June 2001 to July 2002, the geographical distribution of ticks on cattle in the Sudan was determined. Seventeen locations were surveyed from Northern, Central, Eastern, Western, Blue Nile and White Nile Provinces. Total body collections of ticks were made from 20 cattle at each location. Four tick genera and 11 species were identified. The tick species collected included Amblyomma lepidum, Amblyomma variegatum, Boophilus decoloratus, Hyalomma anatolicum anatolicum, Hyalomma dromedarii, Hyalomma impeltatum, Hyalomma marginatum rufipes, Hyalomma truncatum, Rhipicephalus evertsi evertsi, Rhipicephalus sanguineus group and Rhipicephalus simus simus. Major ecological changes have occurred due to extensive animal movement, deforestation, desertification and establishment of large mechanized agricultural schemes. These factors have certainly affected the distribution of ticks and tick-borne diseases in the Sudan. The absence of A. variegatum and A. lepidum in northern Sudan was not surprising, since these tick species are known to survive in humid areas and not in the desert and semi-desert areas of northern Sudan. The absence of B. annulatus in northern and central Sudan is in accordance with the finding that this tick species is restricted to the southern parts of the central Sudan. The presence of H. anatolicum anatolicum in Um Benin in relatively high abundance is an interesting finding. The present finding may indicate that the southern limit of this species has changed and moved southwards to latitude 13 degrees N. It is concluded that major changes in tick distribution have taken place in the Sudan.     

Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa

The suitability of present and future climates for 30 Rhipicephalus species in Africa are predicted using a simple climate envelope model as well as a Division of Atmospheric Research Limited-Area Model (DARLAM). DARLAM's predictions are compared with the mean outcome from two global circulation models. East Africa and South Africa are considered the most vulnerable regions on the continent to climate-induced changes in tick distributions and tick-borne diseases. More than 50% of the species examined show potential range expansion and more than 70% of this range expansion is found in economically important tick species. More than 20% of the species experienced range shifts of between 50 and 100%. There is also an increase in tick species richness in the south-western regions of the sub-continent. Actual range alterations due to climate change may be even greater since factors like land degradation and human population increase have not been included in this modelling process. However, these predictions are also subject to the effect that climate change may have on the hosts of the ticks, particularly those that favour a restricted range of hosts. Where possible, the anticipated biological implications of the predicted changes are explored.