Life cycle and host specificity of Amblyomma triste (Acari: Ixodidae) under laboratory conditions

We report biological data of two generations of Amblyomma triste in laboratory and compared the suitability of different host species. Infestations by larval and nymphal stages were performed on guinea pigs (Cavia porcellus), chickens (Gallus gallus), rats (Rattus norvegicus), rabbits (Oryctolagus cuniculus), wild mice (Calomys callosus), dogs (Canis familiaris) and capybaras (Hydrochaeris hydrochaeris). Infestations by adult ticks were performed on dogs, capybaras and rabbits. Tick developmental periods were observed in an incubator at 27 degrees C and RH 90%. Guinea pigs were the most suitable hosts for larvae and nymphs, followed by chickens. The remaining host species were less suitable for immature ticks as fewer engorged ticks were recovered from them. Mean larval feeding periods varied from 3.8 to 4.7 d between different host species. Mean larval premolt periods ranged from 8.9 to 10.4 d. Nymphal mean feeding periods varied from 4.2 to 6.2 d for ticks fed on different host species. Premolt period of male nymphs (mean: 15.4 d) was significantly longer than that of female nymphs (14.7 d). Female nymphs were significantly heavier than male nymphs. The overall sex ratio of the adult ticks emerged from nymphs was 0.9:1 (M:F). Capybaras were the most suitable host for the tick adult stage as significantly more engorged females were recovered from them and these females were significantly heavier than those recovered from dogs or rabbits. The life cycle of A. triste in laboratory could be completed in an average period of 155 d. The potential role of guinea pigs, birds and capybaras, as hosts for A. triste in nature, is discussed.     

Life cycle and host specificity of Amblyomma triste (Acari: Ixodidae) under laboratory conditions

We report biological data of two generations of Amblyomma triste in laboratory and compared the suitability of different host species. Infestations by larval and nymphal stages were performed on guinea pigs (Cavia porcellus), chickens (Gallus gallus), rats (Rattus norvegicus), rabbits (Oryctolagus cuniculus), wild mice (Calomys callosus), dogs (Canis familiaris) and capybaras (Hydrochaeris hydrochaeris). Infestations by adult ticks were performed on dogs, capybaras and rabbits. Tick developmental periods were observed in an incubator at 27 °C and RH 90%. Guinea pigs were the most suitable hosts for larvae and nymphs, followed by chickens. The remaining host species were less suitable for immature ticks as fewer engorged ticks were recovered from them. Mean larval feeding periods varied from 3.8 to 4.7 d between different host species. Mean larval premolt periods ranged from 8.9 to 10.4 d. Nymphal mean feeding periods varied from 4.2 to 6.2 d for ticks fed on different host species. Premolt period of male nymphs (mean: 15.4 d) was significantly longer than that of female nymphs (14.7 d). Female nymphs were significantly heavier than male nymphs. The overall sex ratio of the adult ticks emerged from nymphs was 0.9:1 (M:F). Capybaras were the most suitable host for the tick adult stage as significantly more engorged females were recovered from them and these females were significantly heavier than those recovered from dogs or rabbits. The life cycle of A. triste in laboratory could be completed in an average period of 155 d. The potential role of guinea pigs, birds and capybaras, as hosts for A. triste in nature, is discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Life-cycle and host specificity of Amblyomma tigrinum (Acari: Ixodidae) under laboratory conditions

Biological data of three generations of Amblyomma tigrinum in the laboratory are reported and the suitability of different host species for immature ticks are compared. Grouping the three generations, infestations by both the larval and nymphal stages were performed on chickens (Gallus gallus), wistar rats (Rattus norvegicus), rabbits (Oryctolagus cuniculus),wild mice (Calomys callosus), dogs (Canis familiaris) and opossums (Didelphis albiventris). Only dogs were used for infestations by adult ticks. Tick developmental periods were observed in an incubator at 27°C and RH 90%. The proportion of engorged larvae recovered from chickens (21.7% of the exposed larvae) was significantly larger (p<0.001) than those from the five mammal species used in the infestations (maximum of 3.1%). A significant larger (p<0.01) proportion of engorged larvae successfully molted after being fed on chickens than on mammal hosts. The proportion of engorged nymphs recovered from chickens (28.8% of the exposed nymphs) was significantly larger (p<0.001) than those from mammal hosts (range: 0–2.1%). Larvae showed similar feeding periods on exposure to different host species, except for those larvae fed on C. callosus, which showed significantly longer (p<0.001) feeding periods. Engorged larvae detachment peaked on the 5^th feeding day, followed by the 6^th day, on all hosts except for C. callosus. Larval premolt periods were similar for engorged ticks exposed to different host species, except for larvae fed on dogs, which showed significantly longer (p<0.001) premolt periods. Host detachment of engorged nymphs peaked on the 6^th feeding day on chickens. Although nymphal detachment on rats peaked on the 8^th day, only 15 nymphs were recovered from this host species. In a sample of 144 F₃ nymphs fed on chickens no significant difference (p>0.10) was found between the feeding or premolt periods of 82 males and 62 females, but female nymphs were significantly heavier (p<0.005) than male nymphs. Sixteen engorged females (61.5% of the exposed ticks) were recovered after being fed on dogs, and all these females laid viable eggs. Chickens, the only avian host, were the most suitable host when compared with the five mammal species. Dogs were demonstrated to be a suitable host for adults of A. tigrinum, which is consistent with, several reports of adult A. tigrinum ticks parasitizing dogs in different areas of South America. Our results reinforce that in these same areas avian species are the major hosts for immature stages of this tick species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Expression of acquired immunity to immature stages of the tick Rhipicephalus evertsi evertsi by rabbits and guinea-pigs

Acquired immunity in guinea-pigs and rabbits to immature stages of the two-host tick Rhipicephalus evertsi evertsi Neumann was demonstrated. Repeated infestations of both hosts with larvae resulted in a significant reduction in the weight of later engorged nymphs. A sharp decline in the numbers of nymphs which successfully fed on both hosts was also observed. This study provides evidence for a gradual decrease in the mean weight of engorged nymphs towards the end of the detachment period suggesting that, in two-host ticks, the onset of nymphal feeding acts as an immune booster in a host already primed by the larval feed and that this results in a reduced feeding performance.     

Differential distribution of immature Ixodes dammini (Acari: Ixodidae) on rodent hosts

Ectoparasites such as ixodid ticks that remain attached to hosts for several days while feeding on blood are able to overcome the inflammatory and immune responses of some hosts and not others. The immature stages of the deer tick Ixodes dammini are found more frequently on the white-footed mouse, Peromyscus leucopus, than on other rodents. We propose that P. leucopus is more tolerant to I. dammini than is a less common host, the meadow vole, Microtus pennsylvanicus. To test this hypothesis, the distribution patterns and engorgement indices were determined for larval and nymphal I. dammini collected from wild-caught P. leucopus and M. pennsylvanicus. There were more immature ticks, which were more fully engorged, on P. leucopus than on M. pennsylvanicus. There were more and better engorged ticks on male than on female hosts. Laboratory studies on the number and weights of larval I. dammini collected off naive and previously exposed P. leucopus and M. pennsylvanicus support the results of the field study. Fewer larval ticks were recovered from previously exposed M. pennsylvanicus than P. leucopus, and the ticks weighed less. Larval and nymphal ticks aggregated among hosts in the study grid, and higher densities per male P. leucopus were correlated with higher engorgement indices, suggesting that immature I. dammini feed better at higher densities. The feeding success of I. dammini on its preferred host species might be due to its adaptation to the immune and inflammatory reactions of the host.     

Biological aspects of <Emphasis Type="Italic">Amblyomma brasiliense</Emphasis> (Acari: Ixodidae) under laboratory conditions

Although Amblyomma brasiliense Aragão 1908 has been reported as one of the most aggressive ticks to humans in Brazil, information about the biology of this tick species is virtually inexistent. This work reports data on the life cycle of A. brasiliense fed on rabbits and pigs and maintained in an incubator at 20°C, 90% RH and 12 h of light for off-host development. Tick yield of adult females fed on pigs and rabbits was 81.2% and 58.3%, respectively. Females fed on pigs had mean engorgement weight of 862.3 mg and egg mass of 208 mg, while females fed on rabbits had mean engorgement weight of 606.1 mg and egg mass of 160 mg; these values did not differ statistically between host species. Feeding period of female ticks fed on pigs (10 days) was significantly shorter than that on rabbits (17 days). Mean preoviposition period was slightly longer (35.9 days) for ticks fed on pigs than on rabbits (30 days). The minimum incubation period of eggs of ticks from both host species was similar and over 100 days. Egg production efficiency was low for females fed on both hosts (less than 30% and 20% for ticks from pigs and rabbits, respectively). More than 55% of larvae and 79% of nymphs fed on rabbits, set free inside the feeding chambers, engorged successfully. These ticks attained an engorgement weight of 1.3 and 18.2 mg, respectively, and fed for approximately 5 days. The minimum pre-molt period was 30 days for engorged larvae and over 44 days for nymphs. Molting success was low, less than 50% in the case of larvae and less than 20% for nymphs. Further studies are required to better determine the off-host requirements of this tick species.     

Prevalence and abundance of Ixodes pacificus immatures (Acari: Ixodidae) infesting western fence lizards (Sceloporus occidentalis) in northern California: temporal trends and environmental correlates.

The prevalence and abundance of immature Ixodes pacificus ticks on western fence lizards (Sceloporus occidentalis) were examined in relation to time of year, host attributes (i.e., age, gender, and presence or absence of blood parasites), and 5 environmental characteristics, including topographic exposure and ground cover substrate, over a 2-year period in northern California. Lizards were infested with subadult ticks from early March until late July or early August, with peak median numbers of larvae and nymphs recorded in late April and early May of both years. Peak larval and nymphal abundances differed between years. The overall ratio of larvae to nymphs on adult male lizards was low, ranging from 0.80 in 1999 to 2.41 in 2000. Such intensive feeding of nymphs versus larvae on these lizards, which are reservoir-incompetent for Borrelia burgdorferi spirochetes, may explain previous observations of decreasing spirochetal infection prevalence from the nymphal to adult stage in northwestern California. Adult male lizards were more likely to be infested with nymphs and harbored greater abundances of larvae and nymphs than adult females. Lizards uninfected with blood parasites had more nymphs than infected lizards. The measured environmental characteristics could explain only a small percentage of the total variation observed in larval prevalence (22%) and in larval and nymphal abundance (12 and 3%, respectively).     

Ticks (Acari: Ixodidae) on wild animals from the Porto-Primavera Hydroelectric power station area,Brazil

From June 2000 to June 2001, a total of 741 ticks were collected from 51 free-living wild animals captured at the Porto-Primavera Hydroelectric power station area, located alongside an approximately 180 km course of the Paran river, between the states of S o Paulo and Mato Grosso do Sul, comprising 9 species of 3 genera: Ambly-omma (7 species), Boophilus (1) and Anocentor (1). A total of 421 immature Amblyomma ticks were reared in laboratory until the adult stage, allowing identification of the species. A. cajennense was the most frequent tick species (mostly immature stages) collected on 9 host species: Myrmecophaga tridactyla, Tamandua tetradactyla,Cerdocyon thous, Puma concolor,Tayassu tajacu, Mazama gouazoubira,Hydrochaeris hydrochaeris,Alouatta caraya, Cebus apella. Other tick species were less common, generally restricted to certain host taxa.     

Experimental evaluation of birds as disseminators of the cosmopolitan tick Rhipicephalus sanguineus (Acari: Ixodidae)

Rhipicephalus sanguineus is believed to be the most widespread tick species of the world and its dissemination seems to rely on the diffusion of its main host, the dog. Empirical observations indicate that several bird species in urban areas regularly steal dog food. Such circumstances create a chance for R. sanguineus ticks to climb on birds and carry ticks to another site. In this work we evaluated experimentally the likelihood of birds (chicks) to either feed and/or carry R. sanguineus ticks from an infested site to another and to infest a host (rabbit) in the new location. Chicks were not suitable hosts for R. sanguineus ticks. Not a single adult tick engorged on chicks, yield as well as weight of engorged larvae and nymphs were very low and feeding period of these ticks was very long. However, a few larvae and, chiefly, nymphs were delivered to a new location either mechanically or after attachment and engorging total or partially on chicks. A few of these ticks fed successfully on rabbits. Further evidence on the capacity of birds to introduce R. sanguineus into non-infested dog settings should be provided by systematic examination of birds from urban areas, close to tick infested households.     

Life cycle and seasonal development of post-embryonic Argas reflexus (Acari: Argasidae) at two thermally different locations in Central Europe

Seasonal development was investigated in the pigeon tick, Argas reflexus (F.) over a 5-year period. The ticks were kept in desiccators at two deposition sites with different temperature conditions: a warmer attic and a cooler outdoor aviary. The life cycle of A. reflexus consists of the egg, larva, a variable number of two to four nymphal instars and the adult stage. In the cooler aviary, the ticks passed, on average, fewer nymphal instars than in the attic. At both locations, ecdysis of the nymphs and adults occurred only during the summer months, with peak numbers of ticks finishing the moult in August. This consistent pattern was evident irrespective of the feeding date of the preceding developmental stage or the year of observation. The results strongly suggest that nymphs II, nymphs I and larvae fed later than in mid-July, August or September, respectively, entered a state of diapause and, thus, overwintered in the engorged state. Argas reflexus nymphs II from a laboratory stock that were deposited inside the attic showed a remarkably different seasonal pattern of development, even more than 1 year after their deposition. This suggests that a circannual rhythm may be involved in the ticks' seasonal timing. Mortality of the engorged ticks (from repletion to ecdysis of the following stage/instar) was below 1.5% in most cases, irrespective of the season and the location. Unfed larvae survived for a maximum of one year inside the attic, whereas the median survival period of unfed nymphs was at least 3 years at the same location. Based on the present results, the generation time from (F1) egg deposition to oviposition in the F2 generation might be 3-11 years in Central European A. reflexus, depending on the course of development (two or three nymphal instars) and the number of gonotrophic cycles (probably up to six) of the F1. The life span of a single tick might take approximately 7-11 years or even longer.     

Development of resistance in rabbits to immature stages of the Ixodid tick Rhipicephalus appendiculatus

A study of acquired resistance in rabbits to larvae and nymphs of Rhipicephalus appendiculatus Neumann showed a positive correlation between the levels of gamma globulin in the serum and the resistance developed by the host. A negative correlation between the weight of engorged ticks and gamma globulin levels was also demonstrated during this study. Repeated infestations with ticks evoked a typical antibody response to antigenic challenge. The levels of gamma globulin stabilized after three infestations and no further decrease in the weight of ticks was observed. The mechanism of the acquisition of resistance by the rabbits, the production of antibodies and the involvement of complement are discussed.     

Influence of temperature and relative humidity on the moulting success of Amblyomma limbatum and Aponomma hydrosauri (Acari: Ixodidae) larvae and nymphs

This study compared the duration of the moulting periods of engorged larvae and nymphs of the ixodid ticks, Amblyomma limbatum and Aponomma hydrosauri, at different temperature/relative humidity regimes, and examined the relationships between the engorged weight of ticks and their weights after moulting. The results showed that for each species, there was a significant relationship between the weights of unfed nymphs and engorged larvae, and the weights of unfed adults and engorged nymphs. The weight of engorged nymphs was also a good indicator of their sex, with female ticks having heavier weights as engorged nymphs. Temperature and relative humidity had a marked effect on the moulting success of engorged ticks of both species. Aponomma hydrosauri larvae and nymphs were able to moult at lower temperatures than Amb. limbatum but most ticks, except Ap. hydrosauri larvae, failed to moult at 13 degrees C. Additionally, there was a marked decrease in the pre-moult times of ticks at higher temperatures, with larvae taking less time to moult than nymphs. At temperatures greater than 21 degrees C, Amb. limbatum took less time to moult than Ap. hydrosauri but this interspecific difference was less marked for nymphs. The interspecific differences in the responses of engorged larvae and nymphs to different temperatures and relative humidities correlated with interspecific differences in off-host behaviour and with the different climates the two species experience throughout most of their distributional range.     

Feeding performance ofAmblyomma variegatum (Acarina: Ixodidae) fed repeatedly on rabbits

Rabbits infested with different stages ofAmblyomma variegatum Fabricius, 1794 became resistant to subsequent infestations by the same tick. Resistance was manifested by a reduction of 73.6% in the mean engorged weight of female ticks by the 3rd infestation. By the 5th infestation, only 70% of the nymphs engorged and their mean engorged weight was 57.7% of that of nymphs fed on tick-naive control rabbits. In the case of larvae, there was a 22.8% reduction in their mean engorged weight by the 6th infestation. Based on increases in body length while feeding on tick-naive rabbits, the nymphs could be divided into four feeding categories, i.e. Nl (unfed), N2, N3 and N4 (increasing states of engorgement). While 92–96% of ticks which dropped from susceptible rabbits fitted into feeding category N4, only 28% of those that dropped from tick-resistant rabbits fitted into that category. The remainder of the ticks fitted into categories N3 (60%) and N2 (12%). The majority of those ticks with reduced weight developed into adults which had an atypical scutal ornamentation pattern, in that an additional pair of lateral spots was regularly observed on the males. The females of such ticks deposited small numbers of eggs from which no larvae hatched.     

Acquired immunity to larvae of Amblyomma marmoreum and Ahebraeum by tortoises, guinea-pigs and guinea-fowl

A study of acquired resistance in guinea-pigs, guinea-fowl and tortoises to larvae of the ticks Amblyomma hebraeum Koch and A.marmoreum Koch (Acari: Ixodidae) showed that repeated infestations of the laboratory host resulted in a 29.3-49.4% reduction respectively in weight of engorged larvae. No resistance was induced in guinea-fowl, the natural host for larvae of both species. A small decline in the mean weight of engorged larvae was observed for both species of ticks after three successive infestations on tortoises. The reduction in weight was more pronounced in A.hebraeum (14.5%) which does not naturally feed on tortoises than for A.marmoreum (8.4%), a common tick on this host. The occurrence of acquired resistance in natural tick host relationships as opposed to unnatural tick host relationships is discussed.     

Immunity against Boophilus annulatus induced by the Bm86 (Tick-GARD) vaccine

Friesian cattle were immunized with two inoculations of anti-tick Bm86 (Tick-GARD) vaccine and were challenged 30 or 90 d later with Boophilus annulatus larvae derived from 1.2 g of eggs. No nymphs or adult ticks were found on the immunized cattle during four weeks after challenge. Repeated infestations (2 to 4) with larvae on three other calves during a period of 160 and 390 d after the immunization did not result in development of nymphal and adult stages. In control, non-immunized cattle infested with corresponding batches of larvae 1380 to 4653 replete adult female ticks were collected. Larvae issued from Babesia bovis-infected female ticks transmitted the infection to Bm86-immunized cattle, but the progeny of B. bigemina-infected females did not. Since B. bigemina is transmitted exclusively by nymphal stages of Bo. annulatus these results support the observation that immunity induced by Bm86 affects the larval stage of this tick.     

Life cycle and seaonal development of postembryonicArgas reflexus (Acari: Argasidae) at two thermally different locations in Central Europe

Seasonal development was investigated in the pigeon tick,Argas reflexus (F.) over a 5-year period. The ticks were kept in desiccators at two deposition sites with different temperature conditions: a warmer attic and a cooler outdoor aviary. The life cycle ofA. reflexus consists of the egg, larva, a variable number of two to four nymphal instars and the adult stage. In the cooler aviary, the ticks passed, on average, fewer nymphal instars than in the attic. At both locations, ecdysis of the nymphs and adults occurred only during the summer months, with peak numbers of ticks finishing the moult in August. This consistent pattern was evident irrespective of the feeding date of the preceding developmental stage or the year of observation. The results strongly suggest that nymphs II, nymphs I and larvae fed later than in mid-July, August or September, respectively, entered a state of diapause and, thus, overwintered in the engorged state.Argas reflexus nymphs II from a laboratory stock that were deposited inside the attic showed a remarkably different seasonal pattern of development, even more than 1 year after their deposition. This suggests that a circannual rhythm may be involved in the ticks' seasonal timing. Mortality of the engorged ticks (from repletion to ecdysis of the following stage/instar) was below 1.5% in most cases, irrespective of the season and the location. Unfed larvae survived for a maximum of one year inside the attic, whereas the median survival period of unfed nymphs was at least 3 years at the same location. Based on the present results, the generation time from (F₁) egg deposition to oviposition in the F₂ generation might be 3–11 years in Central EuropeanA. reflexus, depending on the course of development (two or three nymphal instars) and the number of gonotrophic cycles (probably up to six) of the F₁. The life span of a single tick might take approximately 7–11 years or even longer.     

Biology and life cycle of <Emphasis Type="Italic">Amblyomma incisum</Emphasis> (Acari: Ixodidae)

Amblyomma incisum Neumann is a major tick species in the Atlantic Forest of Brazil. Tapir is the main host for adult ticks and a high aggressiveness of nymphs to humans has been reported. In this work data on the biology and life cycle of this tick species is presented for the first time. It was shown that horse is a suitable host for A. incisum adults and rabbit for larvae and nymphs. It was also shown that A. incisum is a big tick species (mean engorged female weight of 1.96 g) with a long life cycle which lasts 262.3 days when maintained at 27°C and 85% RH. These laboratory conditions were, however, inappropriate and egg hatching rate (1.2%) was very low. Nevertheless egg hatching of ticks in a forest patch increased considerably (72.2%) indicating that this A. incisum population is highly dependent on a forest-like environment.     

Infestations of the bont tick Amblyomma hebraeum (Acari: Ixodidae) on different breeds of cattle in Zimbabwe

Infestations of adults and nymphs of Amblyomma hebraeum were counted on Brahman (Br), Brahman x Simmental (BS), Sanga (Sa) and Hereford (He) steers exposed to infested pastures at Mbizi in southern Zimbabwe in 1986–1987. Herefords were always the most heavily infested, while the Sanga tended to carry the fewest ticks with the Brahman and Brahman x Simmental groups being in between. The ratios of the engorged females on the four breeds were 2.3 : 1.4 : 1.4 : 1.0 for He : Br : BS : Sa. The ratios of the standard nymphs were 2.2 : 1.4 : 1.7 : 1.0 for He : Br : BS : Sa. The results confirm earlier observations in Africa and support the view that there are genetic differences between breeds in the expression of resistance to this tick species.Deceased.     

The life cycle of Hyalomma rufipes (Acari: Ixodidae) under laboratory conditions

Biological characteristics of Hyalomma rufipes parasitising on rabbits and sheep were compared under laboratory conditions in Gansu, China. Mature ticks could parasitize both rabbits and sheep, while immature ticks only fed on rabbits successfully. Adults sucked blood on sheep significantly longer than on rabbits (16 and 13 days, respectively). Other adult parasite characteristics fed on the two hosts were similar, including the weight of engorged adult, female daily oviposition, and the weight and amount of the egg mass laid. Those indicated that this tick species showed little host specificity between sheep and rabbits during its adult stage. In total, the life cycle of H. rufipes was completed in an average period of 179 days. The average developmental periods were 59 days for egg incubation, 3 and 21 days for immature tick prefeeding and feeding, 2, 12 and 40 days for adult prefeeding, female preoviposition and oviposition. The longer female fed for engorgement, the shorter preoviposition period of engorged female needed, although when the feeding period was less than 15 days, this relationship was not obvious. The results confirmed the correlation between the weight of the engorged female and the number of eggs laid (r = 0.909). The reproductive efficiency index (REI) and reproductive fitness index (RFI) in females was 10.63 and 7.22, respectively. Engorged nymphs moulting to females were significantly heavier (27.6 ± 0.89 mg) than those moulting to males (22.3 ± 0.52 mg). Males outnumbered females by 1.4:1.