Aponomma Hydrosauri Research Articles

Reproductive interference

Reproductive interference

Indirect evidence of density-dependent population regulation in Aponomma hydrosauri (Acari: Ixodidae), an ectoparasite of reptiles

Abstract The extent to which density-dependent processes regulate natural populations is the subject of an ongoing debate. We contribute evidence to this debate showing that density-dependent processes influence the population dynamics of the ectoparasite Aponomma hydrosauri (Acari: Ixodidae), a tick species that infests reptiles in Australia. The first piece of evidence comes from an unusually long-term dataset on the distribution of ticks among individual hosts. If density-dependent processes are influencing either host mortality or vital rates of the parasite population, and those distributions can be approximated with negative binomial distributions, then general host–parasite models predict that the aggregation coefficient of the parasite distribution will increase with the average intensity of infections. We fit negative binomial distributions to the frequency distributions of ticks on hosts, and find that the estimated aggregation coefficient k increases with increasing average tick density. This pattern indirectly implies that one or more vital rates of the tick population must be changing with increasing tick density, because mortality rates of the tick's main host, the sleepy lizard, Tiliqua rugosa, are unaffected by changes in tick burdens. Our second piece of evidence is a re-analysis of experimental data on the attachment success of individual ticks to lizard hosts using generalized linear modelling. The probability of successful engorgement decreases with increasing numbers of ticks attached to a host. This is direct evidence of a density-dependent process that could lead to an increase in the aggregation coefficient of tick distributions described earlier. The population-scale increase in the aggregation coefficient is indirect evidence of a density-dependent process or processes sufficiently strong to produce a population-wide pattern, and thus also likely to influence population regulation. The direct observation of a density-dependent process is evidence of at least part of the responsible mechanism.

Variation in the attachment sites of ticks to Australian lizards

Abstract A number of species of reptile ticks in Australia, particularly those using the sleepy lizard (Trachydosaurus rugosus) as their dominant host, form parapatric boundaries wherever they come into contact. A variety of hypotheses have been put forward to explain these boundaries, none of which has yet been demonstrated to be correct. Two of these, one involving reproductive interference and the other competition for attachment sites to hosts, were based on field observations of differences in attachment sites between species and regional variation in attachment sites within species. These hypotheses assume that species-specific attachment site preferences are static. Here we show, using data from single-species infestations of T. rugosus, that this is not the case for either Aponomma hydrosauri males or Amblyomma albolimbatum males or females. We suggest that dynamics in attachment site choice are driven, at least in part, by male movement in search of females.

A Febrile Illness with Generalized Papular Rash Involving the Palms and Soles

Diagnosis: Rickettsia honei infection. A provisional diagnosis of rickettsial illness was made at transfer of the patient to our medical center, and oral doxycycline administered at an initial dose of 200 mg followed by 100 mg twice per day for 1 week was prescribed. Rapid plasma reagin tests and blood cultures had negative results. Results of serial rickettsial serological testing demonstrated a striking increase in antibody titer to R. honei (table 1). Rickettsial PCR and culture performed at the Australian Rickettsial Reference Laboratory (Geelong, Australia) on a blood specimen collected at day 14 of illness (before specific treatment was initiated) had negative results. A number of new spotted fever group rickettsiae have been identified in the past 3 decades. Infection due to R. honei in Australia was first described among residents of Flinders Island in Bass Strait (separating mainland Australia from Tasmania) 15 years ago [1], but in the last 3 years, this infection has also been recognized in southern mainland Australia [2, 3]. The evidence suggests that the principal vector of R. honei infection is the parasitic tick Aponomma hydrosauri, which has a variety of reptile hosts, including members of the skink (Scincidae) family (of which blue-tongue lizards are a member) [4]. The clinical illness is generally characteristic, with fever, myalgias, headaches, and rash. The rash may vary in nature, and the palms and soles are variably involved (figure 1). Eschars and regional lymphadenopathy are inconsistently found. Definitive

Not only ‘Flinders Island’ spotted fever

To demonstrate that Flinders Island spotted fever (FISF), a spotted fever group rickettsial infection caused by Rickettsia honei, is found not only on Flinders Island (Bass Strait), Tasmania, but elsewhere in south-east Australia. Cases of FISF were identified by rickettsial serology, culture and the detection of rickettsial DNA via PCR. Isolates and PCR products were sequenced to identify the aetiological agent as R. honei. Three new cases of FISF were detected outside of Flinders Island. One on Schouten Island, south of the Freycinet Peninsula, Tasmania, and two in south-eastern South Australia (McLaren Vale and Goolwa). These cases show that FISF extends beyond Flinders Island and most likely has the same distribution across south-east Australia as its vector, the reptile tick Aponomma hydrosauri. FISF should be considered as a differential diagnosis in patients from south-eastern Australia presenting with fever, headache and rash following a tick bite.

A new focus of Rickettsia honei spotted fever in South Australia

We recently diagnosed rickettsial spotted fever in four patients from the south-eastern coastal region of South Australia near Adelaide, an area not known to be endemic for this infection. All infections were acquired within the geographic range of Aponomma hydrosauri, the tick vector of Rickettsia honei. Infection by R. honei was confirmed in two patients. This extension of the known geographic range of R. honei infection may be explained, in part, by alterations in host-parasite ecology.

APONOMMA HYDROSAURI, THE REPTILE-ASSOCIATED TICK RESERVOIR OF RICKETTSIA HONEI ON FLINDERS ISLAND, AUSTRALIA

Rickettsia honei is the etiologic agent of Flinders Island (Australia) spotted fever. The tick Aponomma hydrosauri is associated with reptiles and is the arthropod reservoir for this rickettsia on Flinders Island. The rickettsia appears to be maintained in the tick via vertical transmission. Of 46 ticks examined, 29 (63%) were positive for spotted fever group rickettsiae by detection of the citrate synthase gene by a polymerase chain reaction (PCR). From the positive tick samples, seven were sequenced and found to be 100% homologous with R. honei. Of 17 reptiles examined, none had evidence of rickettsiae by PCR or culture of blood.

Ultrastructural and genetic evidence of a reptilian tick, Aponomma hydrosauri, as a host of Rickettsia honei in Australia: possible transovarial transmission.

In 1993, a novel rickettsia was isolated from the blood of inhabitants of Flinders Island, Australia, with acute febrile illnesses. This rickettsia was found to be a new species of spotted fever group (SFG) rickettsia, eventually named Rickettsia honei. The suspected ectoparasite vector of this rickettsia has yet to be identified. The purpose of this study was to evaluate the presence of this rickettsial species in a suspected tick vector, Aponomma hydrosauri, by DNA sequencing and electron microscopy (EM). Ticks collected from an Australian blue-tongued lizard on Flinders Island and a copperhead snake in Tasmania were demonstrated to be infected with R. honei by PCR, DNA sequencing, and EM. Rickettsiae were found in ultrathin sections of salivary glands, malpighian tubules, and midgut epithelial cells. In a previous study with a R. honei-infected tick from Flinders Island, rickettsiae were found in the nuclei of midgut epithelial cells, and EM also revealed the presence of rickettsiae in the cytosol of oocytes and immature eggs, suggesting transovarial transmission. These results implicate A. hydrosauri as a possible host of R. honei on Flinders Island and Tasmania and also provide evidence favoring transovarial maintenance of R. honei.

Indirect evidence of density‐dependent population regulation in Aponomma hydrosauri (Acari: Ixodidae), an ectoparasite of reptiles

Abstract The extent to which density‐dependent processes regulate natural populations is the subject of an ongoing debate. We contribute evidence to this debate showing that density‐dependent processes influence the population dynamics of the ectoparasite Aponomma hydrosauri (Acari: Ixodidae), a tick species that infests reptiles in Australia. The first piece of evidence comes from an unusually long‐term dataset on the distribution of ticks among individual hosts. If density‐dependent processes are influencing either host mortality or vital rates of the parasite population, and those distributions can be approximated with negative binomial distributions, then general host–parasite models predict that the aggregation coefficient of the parasite distribution will increase with the average intensity of infections. We fit negative binomial distributions to the frequency distributions of ticks on hosts, and find that the estimated aggregation coefficient k increases with increasing average tick density. This pattern indirectly implies that one or more vital rates of the tick population must be changing with increasing tick density, because mortality rates of the tick's main host, the sleepy lizard, Tiliqua rugosa, are unaffected by changes in tick burdens. Our second piece of evidence is a re‐analysis of experimental data on the attachment success of individual ticks to lizard hosts using generalized linear modelling. The probability of successful engorgement decreases with increasing numbers of ticks attached to a host. This is direct evidence of a density‐dependent process that could lead to an increase in the aggregation coefficient of tick distributions described earlier. The population‐scale increase in the aggregation coefficient is indirect evidence of a density‐dependent process or processes sufficiently strong to produce a population‐wide pattern, and thus also likely to influence population regulation. The direct observation of a density‐dependent process is evidence of at least part of the responsible mechanism.

Temporal and spatial dynamics of a parapatric boundary between two Australian reptile ticks

Two tick species Aponomma hydrosauri and Amblyomma limbatum that infest large reptiles have an abrupt parapatric boundary near Mt Mary in South Australia. A previous model has suggested that the boundary is maintained at population density troughs resulting from habitat heterogeneity along a gradual environmental gradient. This paper describes the dynamics of the boundary on three transects over 17 years, 1982-98. Over the last seven years of that period there has been a significant increase in rainfall. At the same time, the boundary position has moved 1-2 km on the transects, with the more mesic adapted Ap. hydrosauri advancing into the distribution of the more xeric adapted Amb. limbatum. Also over the same time the density of ticks on lizards in regions flanking the boundary zone has increased for Ap. hydrosauri and decreased for Amb. limbatum. These results suggest that the environmental gradient has been altered, perhaps by increased rainfall, to favour Ap. hydrosauri, which has been able to colonize more successfully across the density troughs and extend its distribution.

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°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°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.

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.

Laboratory studies of ant predation on parapatric reptile ticks

Abstract Two reptile tick species, Aponomma hydrosauri and Amblyomma limbatum, have a parapatric distribution in South Australia. Predation may play a role in maintaining the boundary. Laboratory colonies of Rhytidoponera and Iridomyrmex ants were collected from near Mt Mary, South Australia, close to the tick boundary. They were tested as predators of the two tick species. In the experiments, ticks in leaf litter were more protected from predation than those on bare soil. When comparing leaf litter types from the Mt Mary area, mallee litter was more protective than bluebush litter of equivalent depth. Ticks positioned at the base of the litter layer were more protected from predation than those at the litter surface, and Amb. limbatum ticks were more resistant to predation than Ap. hydrosauri ticks. These results contribute to our understanding of the mechanisms maintaining the abrupt parapatric boundary between the two tick species. Predators may contribute to preventing the more susceptible Ap. hydrosauri from spreading further north, where bluebush litter is more common, and so predation risk is higher. Predators probably have less influence in preventing Amb. limbatum from spreading further south.

Ant predation on different life stages of two Australian ticks

Laboratory colonies of Rhytidoponera ants were allowed to prey on the fed and unfed stages of the Australian ticks Aponomma hydrosauri and Amblyomma limbatum. The unfed tick stages had a higher survival than the fed stages. The ants took longer to handle the adult ticks than the nymphs and longer to handle the nymphs than the larvae. The ants also took longer to handle the unfed than fed nymphs, but longer to handle the fed than unfed females. As well as the differences between the tick stages, there was a species effect, with the ants taking longer to handle A. limbatum, and with that tick species having a higher survival than A. hydrosauri after ant predation. These stage and species differences may influence the tick population dynamics.

Can predators maintain parapatry? Ant distribution across a tick parapatric boundary in South Australia

Abstract Ants were sampled by pitfall traps at 85 sites, 0.5 or 1.0km apart, along six transects across a parapatric boundary between the reptile ticks Aponomma hydrosauri and Amblyomma limbatum near Mt Mary in South Australia. There was no tendency for overall ant density, or for the distribution of any single species of ant, to be related to the tick distribution. Thus the survey found no support for the hypothesis that predation by ants was maintaining the boundary. Along transects there were no correlations between ant and tick densities. Thus the survey found no support for the hypothesis that ant predation was a major factor controlling tick densities. These negative results provide further evidence that single, strong ecological processes cannot explain this well‐studied parapatric boundary.

Host location by larvae of the reptile tick Amblyomma vikirri (Acari: Ixodidae)

The newly described tick Amblyomma vikirri has a narrow host range, being found mainly on the Australian lizard, Egernia stokesii and rarely on another lizard, Tiliqua rugosa. Both hosts are in the family Scincidae. Larvae of A. vikirri were as successful in locating E. stokesii as T. rugosa from a range of release distances between 20 and 120 mm from the host. Over this range the proportion of ticks which successfully located hosts declined and the time taken by successful ticks to locate hosts increased with increasing release distance. From 60 mm, larvae of A. vikirri located four other lizard species from the families Scincidae and Agamidae and two non-living targets as successfully as they did E. stokesii. The only evidence that there was any host specificity in the searching behaviour of larvae of A. vikirri was that A. vikirri larvae spent less time paused and questing when they were searching for E. stokesii than when they were searching for T. rugosa. Aponomma hydrosauri, a tick which commonly infests T. rugosa but not E. stokesii, spent less time paused and questing when it was searching for T. rugosa than when it was searching for E. stokesii. However, the results overall suggest that the narrow host range of A. vikirri cannot be explained by any ability of the larvae of that species to discriminate between their natural host and other reptile species.

Microhabitat choice and its role in determining the distribution of the reptile tick Amblyomma vikirri

Abstract A newly described tick species, Amblyomma vikirri, infests two lizard species in the Flinders Ranges of South Australia. Although one of the hosts, Tiliqua rugosa, has a wide distribution, the tick distribution is restricted to a subset of rocky habitats where the other host, Egernia stokesii, lives. Experiments were conducted with unfed ticks, the stage when the tick is waiting for a host and is susceptible to desiccation. Amblyomma vikirri had a significantly stronger preference for rock microhabitats than two other tick species (Aponomma hydrosauri and Amblyomma limbatum) that infest T. rugosa. Unfed Amb. vikirri were also more active than the other two species, and more likely to be in the upper pan of the substrate. At warm temperatures, larvae of Amb. vikirri survived for shorter periods under desiccation stress than larvae of the other two species, although as nymphs Amb. vikirri survived as well or better. The rock crevice microhabitat experienced lower maximum temperatures than other potential, non‐rock refuge sites. The behaviour of Amb. vikirri, its susceptibility to desiccation in the larval stage and the relatively benign conditions in the rock crevices may act together to prevent Amb. vikirri establishing populations beyond rock habitat.

Host-seeking behaviour by Australian ticks (Acari: Ixodidae) with differing host specificities

Ticks generally detach from their hosts into sites where they are later exposed to host species which may or may not be suitable for infestation. The question of how four species of ixodid ticks, with differing specificities, behave towards such potential host species was examined. Observations of the generalists, Aponomma hydrosauri and Amblyomma limbatum and the specialists, Aponomma fimbriatum and Aponomma concolor, showed that the generalists were attracted to a wider range of potential host species than the specialists and that the larval stages of all tick species, regardless of specificity, contacted more potential host species than the adults. The interspecific differences may be attributed to reliance on different sets of host cues, while the intraspecific differences may reflect the developmental state of the sensory system.

Reproductive interactions between two Australian reptile tick species.

In South Australia the two tick species Amblyomma limbatum and Aponomma hydrosauri share the same common reptile host species, but have allopatric distributions which abut along a narrow parapatric boundary. Reproductive interference is a mechanism that has previously been suggested could contribute to maintaining the boundary. Populations of each species were established in pens within the range of Aponomma hydrosauri. Pens held either each species alone, or the two species together. The performance of females in those pens was monitored over 28 months. There was no indication that the proportion of attached females which mated and engorged was reduced by the presence of heterospecifics. There was no indication that the time taken to mate, engorge and detach was any longer in the presence of heterospecifics. The experiment did not support the hypothesis that reproductive interference contributes to maintaining the parapatric boundary. However, Amblyomma limbatum in the pens had a shorter season of reproductive activity, and achieved much smaller numbers of reproductive females. This may inhibit successful colonization of cooler habitats to the south of its distribution.

Influence of environmental factors on oviposition and egg development in Amblyomma limbatum and Aponomma hydrosauri (Acari: Ixodidae)

This study examined the influence of temperature and light on the length of the pre-oviposition period of engorged females of two Australian ixodid ticks, Amblyomma limbatum and Aponomma hydrosauri. The hatching success and development time of eggs of both species were also compared at different temperatures and relative humidities. Darkness was found to have no effect on the duration of the pre-oviposition time or reproductive output of females of either species. In contrast, the preoviposition period of females of both species decreased with increasing temperature. Amb. limbatum females had shorter pre-oviposition periods than Ap. hydrosauri at all temperatures examined. Temperature and relative humidity had a marked effect on the hatching success of eggs. Eggs of both species had reduced hatching success at low relative humidities. Eggs failed to hatch at temperatures below 21°C. Ap. hydrosauri eggs also failed to hatch at 34°C while Amb. limbatum eggs failed to hatch at 36°C. Within the range of temperatures suitable for egg development, the hatching times of eggs of both species decreased with increasing temperature. Amb. limbatum eggs developed faster than Ap. hydrosauri eggs at temperatures greater than 25°C, but slower at cooler temperatures. These differences in the duration of their preoviposition period, and the responses of females and their eggs to different temperatures and relative humidities correlate with the different climates the two species experience throughout most of their distributional range.