The Effect of Male Ticks on the Feeding Performance of Immature Stages of Rhipicephalus Sanguineus and Amblyomma Americanum (Acari: Ixodidae)

A unique group of immunoglobulin-binding proteins (IGBPs), produced by ixodid male ticks during the latter half of their prolonged feeding period, improves the feeding performance of co-feeding females. As a follow-up to this observation, we investigated whether male tick feeding also affects the feeding of other developmental stages. Immature stages of Rhipicephalus sanguineus (Latreeille) and Amblyomma americanum (L.) were fed on rabbits in the presence or absence of conspecific males. The mean weight of larvae and nymphs of both species that fed around males and detached from the host on the first day of dropping was significantly higher than when the immature ticks fed on rabbits in the absence of males. However, larvae of both species and nymphs of R. sanguineus that fed slower and detached on the second day of dropping did not show significant differences in weight. A similar pattern was observed for A. americanum nymphs although, unlike R. sanguineus, the presence of males also influenced the feeding performance of the nymphs that fed slowly and detached on the second day of drop-off. The improved feeding performance demonstrated by immature ticks in the presence of males may be due to immunomodulatory saliva proteins, such as immunoglobulin-binding proteins (IGBPs) that are introduced into the co-feeding site. The results are considered in relation to the distribution of ixodid tick species on their natural hosts.     

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.     

Tick infestation on birds in an urban Atlantic Forest fragment in north-eastern Brazil

Birds are important hosts for various tick species, playing a significant role in their biological life cycle and dispersion. In this study, we investigated tick infestations on birds trapped in an urban remnant of Atlantic Forest in Pernambuco state, Brazil. From February 2015 to March 2017, 541 birds belonging to 52 species were trapped with mist nets and examined for ectoparasites. Birds trapped in the late successional forest were significantly more infested than birds trapped in the early successional forest. In the same way, ectoparasite infestation varied significantly according to bird weight and collection plot. Overall, 198 birds (36.6%) belonging to 27 species were parasitized by ectoparasites (i.e., ticks, lice and/or mites). Ectoparasites were effectively collected from 111 birds, of which 99 belonging to 20 species were infested by ticks (n?=?261), namely, Amblyomma longirostre (13 nymphs), Amblyomma nodosum (21 nymphs), Amblyomma varium (one nymph), and Amblyomma spp. (five nymphs and 221 larvae). Most of the ticks (>?90%) were collected from Passeriformes. This study provides the second record of A. varium in Pernambuco state and confirms that birds, especially Passeriformes, are important hosts for larvae and nymphs of Amblyomma spp. in the Atlantic Forest biome of Pernambuco.

     

Adaptive timing of detachment in a tick parasitizing hole-nesting birds

In non-permanent parasites with low intrinsic mobility such as ticks, dispersal is highly dependent on host movements as well as the timing of separation from the hosts. Optimal detachment behaviour is all the more crucial in nidicolous ticks as the risk of detaching in non-suitable habitat is high. In this study, we experimentally investigated the detachment behaviour of Ixodes arboricola, a nidicolous tick that primarily infests birds roosting in tree-holes. We infested great tits with I. arboricola larvae or nymphs, and submitted the birds to 2 experimental treatments, a control treatment in which birds had normal access to nest boxes and an experimental treatment, in which the birds were prevented access to their nest boxes for varying lengths of time. In the control group, most ticks detached within 5 days, whereas in the experimental group, ticks remained on the bird for as long as the bird was prevented access (up to 14 days). This prolonged attachment caused a decrease in survival and engorgement weight in nymphs, but not in larvae. The capacity of I. arboricola larvae to extend the duration of attachment in non-suitable environments with no apparent costs, may be an adaptation to unpredictable use of cavities by roosting hosts during winter, and at the same time may facilitate dispersal of the larval instars.     

Body‐mass or sex‐biased tick parasitism in roe deer (Capreolus capreolus)? A GAMLSS approach

Macroparasites feeding on wildlife hosts follow skewed distributions for which basic statistical approaches are of limited use. To predict Ixodes spp. tick burden on roe deer, we applied Generalized Additive Models for Location, Scale and Shape (GAMLSS) which allow incorporating a variable dispersion. We analysed tick burden of 78 roe deer, sampled in a forest region of Germany over a period of 20 months. Assuming a negative binomial error distribution and controlling for ambient temperature, we analysed whether host sex and body mass affected individual tick burdens. Models for larval and nymphal tick burden included host sex, with male hosts being more heavily infested than female ones. However, the influence of host sex on immature tick burden was associated with wide standard errors (nymphs) or the factor was marginally significant (larvae). Adult tick burden was positively correlated with host body mass. Thus, controlled for host body mass and ambient temperature, there is weak support for sex-biased parasitism in this system. Compared with models which assume linear relationships, GAMLSS provided a better fit. Adding a variable dispersion term improved only one of the four models. Yet, the potential of modelling dispersion as a function of variables appears promising for larger datasets.     

Dispersal by bagworm larvae, Metisa plana: effects of population density, larval sex, and host plant attributes

Abstract 1. The work reported here investigated the incidence of dispersal by bagworm larvae Metisa plana (Walker) (Lepidoptera: Psychidae), a species with apterous females and winged males.
2. A mark–recapture study conducted in a plantation of oil palm revealed that a significant proportion of larvae undertakes multiple episodes of ballooning, and suggested that ballooning represents a strategy for foraging both within and between hosts.
3. Experiments conducted in a controlled cage environment indicated that increasing population density and defoliation of oil palm both promote dispersal by larvae.
4. Neonatal larvae typically remained on the host where they emerged, indicating that pupation site of late-instar females influences the distribution of their progeny, and suggesting a high level of genetic relatedness in localised bagworm populations.
5. Density-dependent dispersal may generate relatively uniform distributions of M. plana in plantations of oil palm, by simultaneously stabilising populations on heavily infested palms and redistributing larvae on lightly infested palms.
6. The rate of ballooning was greater for female than for male larvae, possibly resulting from the sex-specific impact of population density on reproductive success or from late-instar females seeking suitable oviposition sites for their future progeny.     

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.     

The effect of feeding immature Karoo paralysis ticksIxodes rubicundus (Acari: Ixodidae) on the metabolic rate of the rock elephant shrew

The effect of feeding immature Karoo paralysis ticks (Ixodes rubicundus) on the resting metabolic rate (RMR) of their principal natural host, the rock elephant shrew (Elephantulus myurus), was investigated under laboratory conditions. The elephant shrews were artificially infested with numbers of ticks simulating natural burdens. The RMR of the elephant shrews was quantified by measuring the oxygen consumption in an open through-flow system. The RMR of hosts infested only with larvae did not differ from that of the control group. Those infested with nymphs, or nymphs and larvae, had significantly lower RMR's compared to the control animals. There were no signs of paralysis in any of the infested hosts.     

Climate,satellite imagery and the seasonal abundance of the tick Rhipicephalus appendiculatus in southern Africa: a new perspective

Abstract. Recent predictive models for the distribution of the African tick Rhipicephalus appendiculatus Neumann, based on the computer packages CLIMEX and BIOCLIM and data derived from meteorological satellites, and for the seasonal dynamics of the same tick using the computer simulation models ECFXPERT and T3HOST, all have their limitations. Statistical analysis of the relationships between the seasonal abundance of all three life stages of this tick and climatic and satellite-derived data from five sites in southern Africa, taken from the literature, supports a new perspective that it is the timing of the questing activity of the desiccation-vulnerable larvae that determines the pattern of the tick's seasonal dynamics. The timing of the activity of nymphs and adults is determined by temperature-dependent development rates plus the delaying phenomenon of photoperiod-sensitive diapause, the timing and duration of which have evolved to achieve maximum generation survival by ensuring the occurrence of eggs and larvae during periods of optimal climatic conditions. The most important environmental factor appears to be night-time minimum temperature, determining condensation and saturation deficit and thus the tick's ability to replenish moisture lost during the daytime and so to survive while questing for hosts. It is the larvae whose numbers are correlated most closely with these factors, consistent with earlier experimental results showing larvae to be most susceptible to desiccating conditions. There is a statistical linkage between larval tick numbers and satellite imagery, arising from the correlation between larval numbers and minimum temperature and saturation deficit conditions, and in turn the relationship between these climatic conditions and the subsequent vegetational changes monitored by the satellites. Moisture availability to larvae is likely to be the critical factor throughout the geographical range of R.appendiculatus, but the precise combination of climatic conditions that optimize moisture availability and questing tick survival can be expected to vary geographically. The relationships between ticks, temperatures and satellite data in parts of equatorial Africa have yet to be established. These correlative patterns highlight both the critical life stage and environmental factors when trying to understand temporal, and ultimately spatial, variations in tick abundance.     

Resistance in cattle to Haemaphysalis (Kaiseriana) longicornis

Samples of each instar of the 3-host tick, Haemaphysalis (Kaiseriana) longicornis were collected from grazing Bos taurus cattle to investigate their survival on hosts with previous exposure to the tick and on previously unexposed ‘naive’ hosts. The ticks were allocated to categories based on their size to observe at what stage of development mortality occurred. Analysis of the age structure of the ticks showed that small but variable proportions of larvae, nymphs and females matured on previously exposed hosts and larger proportions on the naive hosts. This suggests that cattle acquire and express different levels of resistance against each instar of the tick. The technique used has potential for estimating levels of resistance of hosts to 3-host ticks under field conditions.     

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.     

Accelerated phenology of blacklegged ticks under climate warming

The phenology of tick emergence has important implications for the transmission of tick-borne pathogens. A long lag between the emergence of tick nymphs in spring and larvae in summer should increase transmission of persistent pathogens by allowing infected nymphs to inoculate the population of naive hosts that can subsequently transmit the pathogen to larvae to complete the transmission cycle. In contrast, greater synchrony between nymphs and larvae should facilitate transmission of pathogens that do not produce long-lasting infections in hosts. Here, we use 19 years of data on blacklegged ticks attached to small-mammal hosts to quantify the relationship between climate warming and tick phenology. Warmer years through May and August were associated with a nearly three-week advance in the phenology of nymphal and larval ticks relative to colder years, with little evidence of increased synchrony. Warmer Octobers were associated with fewer larvae feeding concurrently with nymphs during the following spring. Projected warming by the 2050s is expected to advance the timing of average nymph and larva activity by 8–11 and 10–14 days, respectively. If these trends continue, climate warming should maintain or increase transmission of persistent pathogens, while it might inhibit pathogens that do not produce long-lasting infections.     

Borrelia Burgdorferi Sensu Lato in Ixodes Ricinus Ticks and Rodents in a Recreational Park in South-Western Ireland

Ixodes ricinus ticks infected with Borrelia burgdorferi sensu lato were numerous on the edges of paths and roads in a recreational park in south-western Ireland. The abundance of ticks at different sites was related to the presence of deer, but a negative relationship was shown between tick abundance and tick infection rates. This is thought to be due to the deposition of large numbers of uninfected ticks by deer, which are apparently not good reservoir hosts of B. burgdorferi s.l. Blood meal analysis only detected deer DNA in uninfected nymphs. Reservoir competent rodents, Apodemus sylvaticus and Clethrionomys glareolus, were abundant at all sites and a high proportion of captured specimens were infested with larval ticks. However, very few rodents were infected with B. burgdorferi s.l. and none of the unfed infected nymphs analysed for the identity of their larval blood meal had fed on rodents. The spirochaetes detected in I. ricinus in the study area may be poorly adapted to rodents or are not transmitted readily because of the absence of nymphal infestation. The majority of spirochaetes in these ticks were apparently acquired from non-rodent hosts, such as birds.     

Attachment site selection of ticks on roe deer, <Emphasis Type="Italic">Capreolus capreolus</Emphasis>

The spatio-temporal attachment site patterns of ticks feeding on their hosts can be of significance if co-feeding transmission (i.e. from tick to tick without a systemic infection of the host) of pathogens affects the persistence of a given disease. Using tick infestation data on roe deer, we analysed preferred attachment sites and niche width of Ixodes ticks (larvae, nymphs, males, females) and investigated the degree of inter- and intrastadial aggregation. The different development stages showed rather consistent attachment site patterns and relative narrow feeding site niches. Larvae were mostly found on the head and on the front legs of roe deer, nymphs reached highest densities on the head and highest adult densities were found on the neck of roe deer. The tick stages feeding (larvae, nymphs, females) on roe deer showed high degrees of intrastadial spatial aggregation, whereas males did not. Male ticks showed large feeding site overlap with female ticks. Feeding site overlap between larval-female and larval-nymphal ticks did occur especially during the months May–August on the head and front legs of roe deer and might allow pathogen transmission via co-feeding. Tick density, niche width and niche overlap on roe deer are mainly affected by seasonality, reflecting seasonal activity and abundance patterns of ticks. Since different tick development stages occur spatially and temporally clustered on roe deer, transmission experiments of tick-borne pathogens are urgently needed.     

The lizards Lacerta agilis and L. vivipara as hosts to larvae and nymphs of the tick Ixodes ricinus

In this paper we provide quantitative information on the occurrence of larvae and nymphs of the tick Ixodes ricinus in populations of the lizards Lacerta agilis and L. vivipara . Levels of infestation were rather low, at least when compared with those of small mammals and sheep. Hence we suppose that lizards feed only a minor fraction of the total tick population.
Differences in tick loads among lizard subpopulations are probably attributable to difference in body size and mobility among the host groups.
Ticks exhibit a markedly clumped distribution on the lizards. This distribution pattern fits with the negative binomial distribution. The overdispersed distribution of tick larvae in the field and aspects of the lizards' behaviour are considered as factors which contribute to the observed infestation patterns.
Tick larvae were active throughout summer, with peak levels occurring during June-July. Nymphs were most numerous during May-June but almost absent during the summer months. Almost always ticks were attached near the lizards' forelimbs. Possible mortality resulting from tick infestation does not contribute significantly to the overall lizard mortality. Hence, these ectoparasites seem to have but a minor impact on the lizard populations.     

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.     

Being a parasitoid of parasites: host finding in the tick wasp Ixodiphagus hookeri by odours from mammals

The parasitic wasp Ixodiphagus hookeri Howard (Hymenoptera: Encyrtidae) parasitizes larvae and nymphs of a number of tick species worldwide. Ticks themselves are parasitic on vertebrate hosts. To study the specificity and reliability of vertebrate odours used by I. hookeri for host location, we conducted bioassays in a four‐chamber olfactometer. Wasps were arrested by carbon dioxide and by odours from roe deer faeces and odours from hair of roe deer and wild boar. Odours from faeces of cattle, rabbit, and field mouse as well as odours from hair of cattle and field mouse had no effect. Odours from faeces of the host tick species Ixodes ricinus L. (Acari: Ixodidae) were attractive only up to a distance of 1 cm. Thus, I. hookeri reacts to general (carbon dioxide) and specific vertebrate odours from wild boar and deer. Examination of freshly shot specimens demonstrates that deer and wild boars are infested with a sufficient number of tick nymphs to tap the full reproductive potential of an I. hookeri female, which makes cues from these mammal species reliable. These results indicate that I. hookeri locates its hosts using specific and reliable mammal odours and that ticks are parasitized on their vertebrate hosts. The implications of this host‐finding strategy and its benefits for the parasitoid are discussed.     

Biology of Ixodes (Pholeoixodes) hexagonus under laboratory conditions. Part I. Immature stages

A rearing method for Ixodes (Pholeoixodes) hexagonus, the hedgehog tick, was established which enabled the life cycle of immature stages to be studied under laboratory conditions. Larvae were fed on Swiss mice and nymphs on the cars of New Zealand rabbits. The feeding time of the larvae and nymphs on both hosts was 4–17 days. Larvae moulted to nymphs 15–21 days after detachment from mice. The premoulting period was 13–26 days for newly emerged males and 15–27 days for females. Engorged nymphs which developed into males weighed less (5.64±0.91 mg) than those that developed into females (6.019±88mg). The sex ratio (male: female) under laboratory conditions was 1:1.13.     

Feeding patterns of immature stages of Hyalomma truncatum and Hyalomma marginatum rufipes on different hosts

In this study we examine the feeding patterns of immature stages of Hyalomma truncatum and Hyalomma marginatum rufipes ticks on different hosts. Larvae of H. truncatum developed through a three-host pattern on two species of field mice, Rhabdomys pumilio and Lemniscomys rosalia. On guinea-pigs, both Hyalomma species followed a mixed two-host and three-host pattern, with the latter route being preferred, since more than 70% of the fully fed larvae dropped off from their hosts. H. truncatum was a two-host tick on rabbits. Larvae of H. marginatum rufipes did not prefer R. pumilio and L. rosalia as hosts. On guinea-pigs, H. marginatum rufipes immatures showed a mixed two-host and three-host pattern with a bias towards the three-host life cycle, since approximately 58% of the fully fed larvae dropped off. On rabbits, H. marginatum rufipes was exclusively a two-host tick. Mean engorgement weights and blood quantities ingested by H. truncatum nymphs that developed through a three-host pattern on mice were significantly higher (p<0.0001) than for those that developed through a two-host pattern on guinea-pigs and rabbits. For H.marginatum rufipes, there were no significant differences (p>0.05) between engorgement weights of nymphs that developed through two-host and three-host patterns. However, there were significant differences (p<0.0001) in blood quantities ingested by nymphs of this tick species following feeding on different hosts.     

Population ecology of the commensal chironomid Epoicocladius flavens on its mayfly host Ephemera danica

SUMMARY. 1. Epoicocladius flavens (Diptera: Chironomidae) has a 1-year life cycle in the River Tud in eastern England. Pupation and adult emergence occurred from the end of May until the beginning of September. Larval growth was rapid during the summer but thereafter almost ceased until the following April.1 Unlike other chironomids, the population density of E. flavens was very stable and remained at around 1000m ⁻² all through the year.
2. Ephemera danica (Ephemeroptera: Ephemeridae), the host of E. flavens , has a 2-year life cycle. The proportion infested was about 90% for the older host year-class and less than 30% for the younger year-class from autumn until the following summer. A rapid increase in infestation occurred during the summer as a result of hatching of the E. flavens eggs. Of those infested, the older year-class hosts harboured two or three larvae, on average, whereas the younger ones had only one.
3. When host year-classes were treated separately, the frequency distribution of the number of E. flavens per host each month did not depart significantly from a Poisson distribution. Younger instars departed from a Poisson distribution more frequently than older instars.
4. The majority of E. flavens larvae attached to the gills of their hosts. As the instars progressed, increasing proportions were found on the thorax, legs and tails. The thorax was the preferred site for fourth instar larvae.
5. Enhanced chance of survival, rather than the supply of detrital food particles which collect on the host's body, is considered as an important benefit to accrue to E. flavens from this commensalism.