Helminths of the wild rabbit (Oryctolagus cuniculus) in Macaronesia

Two hundred and four rabbits from 8 Macaronesian islands (Pico, San Jorge, San Miguel, Terceira, and Flores from Azores Archipelago; Tenerife and Alegranza from Canary Islands; and Madeira from Madeira Archipelago) were examined for helminth parasites between 1995 and 2000. Three species of cestodes, Taenia pisiformis (larvae), Andrya cuniculi, and Mosgovoyia ctenoides, and 5 species of nematodes, Trichuris leporis, Graphidium strigosum, Trichostrongylus retortaeformis, Passalurus ambiguus, and Dermatoxys hispaniensis, were identified. Only 3 species (M. ctenoides, T. retortaeformis, and P. ambiguus) were regularly distributed over the 3 archipelagos. Taenia pisiformis was not collected in Madeira, nor was A. cuniculi in the Azores and G. strigosum in the Canary Islands. Trichuris leporis and D. hispaniensis were only found in Madeira. Significant differences in the general prevalence of the nematodes G. strigosum and T. retortaeformis were detected between Azores and Madeira. The prevalence of T. retortaeformis differs significantly between the Azores and the Canaries and that of P. ambiguus was higher in Madeira than in Azores and Canaries. The helminth richness found in the wild rabbit in these Macaronesian archipelagos was very low compared with the Palearctic helminth fauna of this host. The wild rabbit was introduced from the Iberian Peninsula into different Macaronesian islands. Helminths introduced with Oryctolagus cuniculus into these islands also are commonly found in Iberian wild rabbits, which are excellent colonizers, as demonstrated in this study.     

Influence of host weight, sex and reproductive status on helminth parasites of the wild rabbit, Oryctolagus cuniculus, in Navarra, Spain.

A study was carried out in Navarra (northern Spain) on the influence of the weight, sex and reproductive status (lactant, pregnant or lactant + pregnant females and testicular weight for males) of the wild rabbit (Oryctolagus cuniculus) on two cestodes species: Andrya cuniculi and Mosgovoyia ctenoides and four intestinal nematodes: Graphidium strigosum, Trichostrongylus retortaeformis, Nematodiroides zembrae and Dermatoxys hispaniensis. A significantly higher prevalence of A. cuniculi was detected in lactant + pregnant females compared with non-breeding females. Trichostrongylus retortaeformis and N. zembrae showed a significantly higher mean intensity in lactant and lactant + pregnant females than in non-reproductive females. Trichostrongylus retortaeformis presented a higher mean intensity in females than in males, and the mean intensity of the same parasite species was significantly lower in active and inactive males compared with lactant and lactant + pregnant females. There were no significant differences between sexes in the prevalence of helminth parasites. No significant correlation was detected between host weight and the intensity (of infection) of helminths studied. No significant differences in the prevalence and mean intensity of the two cestode species were observed in the three weight categories studied (kittens, juveniles and adults). The prevalence of G. strigosum and mean intensity of T. retortaeformis were significantly higher in older heavier animals than in juveniles.     

The helminth parasites of the wild rabbit Oryctolagus cuniculus and their effect on host condition in Dunas de Mira, Portugal

The present study focuses on the helminth parasite community of the wild rabbit in a sand dune area in Portugal over a 5-year period. The influence of host sex and year on the composition of the helminth community is assessed, along with the potential effect of the detected helminths on host body condition. The basic structure of the helminth community comprises Mosgovoyia ctenoides, Graphidium strigosum, Trichostrongylus retortaeformis, Nematodiroides zembrae and Passalurus ambiguus. Mean intensities of G. strigosum varied between years. General G. strigosum intensities were also found to vary according to both year and host sex, but not according to the interaction of both factors. When assessing the effect of helminths on rabbit body condition (expressed by the kidney fat index), higher burdens of M. ctenoides, a cestode that presents a relatively large body mass, were found to induce a reduction in rabbit condition.     

Helminth parasites of the wild rabbit Oryctolagus cuniculus near Malham Tarn, Yorkshire, UK

Between 1992 and 1996, 95 rabbits from the immediate locality of Malham Tarn, North Yorkshire, UK were examined for the presence of helminth parasites. All the examinations took place in late September or October. Three species of nematodes, Graphidium strigosum, Passalurus ambiguus and Trichostrongylus retortaeformis and two species of cestodes, Taenia pisiformis and Cittotaenia pectinata were identified. There were no associations between helminth species richness and year of sampling, host weight or sex. A logistic model was fitted to the prevalence data from these helminths as was an over-dispersed Poisson model to the worm burden data. Graphidium strigosum was the most frequently identified species with an average prevalence of 78%. The mean prevalence and intensity of Graphidium infection were significantly effected by sampling year. The lower than normal rainfall recorded at the Tarn during the years 1995 and 1996 may have be one reason for this pattern. The worm burden of G. strigosum was significantly positively associated with rabbit body weight. The intensity of infection with P. ambiguus was significantly higher in female rabbits. There was a significant non-linear relationship between P. ambiguus worm burden and rabbit weight (P = 0.002) with worm burdens being highest in the 1000 g to 1499 g weight cohort. Trichostrongylus retortaeformis was only identified in 1994 and male rabbits harboured significantly higher worm burdens than females (48 vs. 7, P = 0.022). Over the five years, the average Taenia pisiformis prevalence was 31% and there was a significant positive association between worm burden and rabbit weight (P = 0.001). Cittotaenia pectinata had a prevalence of 37% over the whole study period with no interactions between prevalence or intensity and body weight, year of sampling or rabbit sex. All five helminths showed an overdispersed distribution with k values less than 1.     

The occurrence and abundance of helminth parasites of the mountain hare Lepus timidus (L.) and the wild rabbit Oryctolagus cuniculus (L.) in Aberdeenshire, Scotland

Mountain hares were collected each month between April 1984 and March 1985 and their intestines examined for helminths. 26 rabbits were also collected between July and December 1984 and examined for gastro-intestinal helminths. Three nematode and two cestode species were found in the hares while only two nematode and one cestode species were found in the rabbits. Trichostrongylus retortaeformis was recorded from 88% of the 193 hares and 92% of the rabbits while comparable figures for Passalurus ambiguus were 0.5% and 54% and for Mosgovoyia pectinata 14% and 81%. Seasonal fluctuations were observed in the T. retortaeformis populations in the hare; numbers tended to be low during the winter months and high throughout the summer. Paranoplocephala wimerosa was reported for the first time from the British Isles and Trichostrongylus axei was recorded for the first time in the mountain hare.     

Patterns of parasite aggregation in the wild European rabbit (Oryctolagus cuniculus)

Understanding the factors controlling the distribution of parasites within their host population is fundamental to the wider understanding of parasite epidemiology and ecology. To explore changes in parasite aggregation, Taylor's power law was used to examine the distributions of five gut helminths of the wild rabbit. Aggregation was found to be a dynamic process that varied with year, season, host sex, age class, and myxomatosis. Yearly and seasonal changes are thought, in the main, to be the result of variations in weather conditions acting upon infectious stages (or intermediate hosts). Evidence in support of this was the comparatively low degree of fluctuation in the aggregation of the pinworm, Passalurus ambiguus, as the infectious stage of this parasite is likely to be less susceptible to environmental variation. Host age had a marked effect on the level of aggregation of all parasites, but this effect varied between parasite species. P. ambiguus, Trichostrongylus retortaeformis and Cittotaenia denticulata aggregation were lower in adult than juvenile rabbits whilst Graphidium strigosum and Mosgovoyia pectinata aggregation tended to increase with age. Host immunity is thought to be responsible for these differences. Differences in aggregation for different parasites were also seen when the rabbit population was split into males and females. Myxomatosis had a marked effect on helminth distribution with substantially less aggregation in rabbits showing clinical signs of the disease.     

Influence of habitat modification on the intestinal helminth community ecology of cottontail rabbit populations

The influence of five brush management treatments using the herbicides tebuthiuron and triclopyr, with or without prescribed burning, on the intestinal helminth community of cottontail rabbits (Sylvilagus floridanus) was studied in 1987 on the Cross Timbers Experimental Range in Payne County, Oklahoma (USA). Six helminth species were found (Dermatoxys veligera, Trichostrongylus calcaratus, Passalurus nonanulatus, Wellcomia longejector, Taenia pisiformis cystercercus, and Mosgovoyia pectinata americana) in 102 rabbits (88 adult and 14 juveniles) collected over two seasons (winter and summer). Prevalence of M. pectinata americana in cottontail rabbits was significantly greater in untreated control pastures than herbicide treated pastures in winter, while prevalence of T. pisiformis was significantly greater in burned than unburned pastures. Abundances of helminth species in the intestinal tract of cottontail rabbits were unaffected by brush treatments. Mosgovoyia pectinata americana abundance demonstrated a highly significant increase from winter to summer; conversely, abundance of all oxyurid pinworms combined (D. veligera, P. nonanulatus, W. longejector) was significantly higher in winter than summer. Helminth community dynamics were significantly influenced by season, but were unaffected by brush treatments. Habitat modification could have influenced cestode transmission by altering the ecology of invertebrate and vertebrate hosts.     

The incidence of helminth parasites from the wild rabbit Oryctolagus cuniculus (L.) in eastern Scotland

Samples of wild rabbits were collected each month over the six-year period 1977-82 and their intestines and abdomens examined for helminths. Three species of nematode, four of cestode and one trematode were found. Graphidium strigosum was recorded in 33% of the 786 rabbits, Trichostrongylus retortaeformis in 79%, Passalurus ambiguus in 14.2%, Cittotaenia pectinata in 37%, C. denticulata in 11%, Coenurus pisiformis in 3%, Cysticercus serialis in 0.1% and Fasciola hepatica in 0.1%. G. strigosum and P. ambiguus infections were greatest in the older heavier animals whereas juvenile rabbits had the heaviest T. retortaeformis nematode burdens. Seasonal fluctuations were observed in G. strigosum, T. retortaeformis, P. ambiguus and Cittotaenia denticulata. The sex and reproductive state of the female rabbit did not have any significant effect on either the nematode or cestode worm burdens. Myxomatosis was shown to significantly increase the worm burden of T. retortaeformis and raise that of P. ambiguus. No nematode or cestode from domesticated animals were recovered and it was concluded that since F. hepatica was found only once rabbits did not constitute a serious hazard to the farm animals grazing the area.     

Evaluation of the association of parasitism with mortality of wild European rabbits Oryctolagus cuniculus (L.) in southwestern Australia

Abundances of the parasitic nematodes Trichostrongylus retortaeformis and Passalurus ambiguus, and 8 Eimeria species were estimated by fecal egg and oocyst output in 12 discrete free-ranging populations of wild rabbits (Oryctolagus cuniculus) in southwestern Australia. Comparisons of parasite egg and oocyst counts were made between those rabbits known to have survived at least 2 mo after fecal samples were collected and those rabbits that did not survive. There were significant negative relationships between parasite egg and oocyst counts and survival when all age groups and collection periods were pooled for several species of coccidia and for T. retortaeformis. However, when the same comparisons were made within rabbit age groups and within collection periods, there were very few significant differences even where sample sizes were quite large. The differences indicated by the pooled analysis for coccidia were most likely due to an uneven host age distribution with respect to survival, combined with an uneven distribution of the oocyst counts with rabbit age. The result for T. retortaeformis was similarly affected but by a seasonal pattern. Parasitism by nematodes and coccidia did not appear to be an important mortality factor in these rabbit populations, at least at the range of host densities we examined. This suggests that other factors must have been responsible for the observed pattern of density-dependent regulation in these rabbits.     

Observations on the seasonal incidence of myxomatosis and its interactions with helminth parasites in the European rabbit (Oryctolagus cuniculus)

European rabbits (Oryctolagus cuniculus) were collected monthly over a 10-yr period and the incidence of myxomatosis and the size of helminth populations were recorded. Myxomatosis occurred annually, always as an epidemic in the latter half of the year and was associated with both an increase in the percentage of animals infected and the size of the infections of the nematodes Trichostrongylus retortaeformis and Passalurus ambiguus and the cestode Mosgovoyia pectinata. It is suggested that myxomatosis had the effect of reducing the rabbits' overall immune response to infection and that due to the regular annual occurrence of myxomatosis, the resulting increase in the size of helminth infections must be considered an integral part of the population dynamics of these parasites.     

Factors influencing the fecal egg and oocyst counts of parasites of wild European rabbits Oryctolagus cuniculus (L.) in Southern Western Australia

Abundance of intestinal parasites was monitored by fecal egg and oocyst counts for samples of wild rabbits Oryctolagus cuniculus with different levels of imposed female sterility from 12 populations in southwestern Australia. Differences in egg counts of Trichostrongylus retortaeformis between seasons and age groups were dependent on the sex of the host. Pregnancy may have been responsible for these differences because egg counts were consistently higher in intact females than in females surgically sterilized by tubal ligation. Egg counts for Passalurus ambiguus were influenced by season and host age but there were no differences between sexes or between intact and sterilized female rabbits. No differences were detected in the oocyst counts of the 8 species of Eimeria between male and female rabbits or between intact and sterilized females. Seasonal differences were detected in oocyst counts of Eimeria flavescens and Eimeria stiedai. The overwhelming determinant of coccidian oocyst counts was host age, with 6 species being much more abundant in rabbits up to 4 mo of age. There was a suggestion that egg counts of T. retortaeformis and oocyst counts of several species of Eimeria were reduced in populations where rabbit numbers had been depressed for at least 2 yr, but there was no evidence that short-term variations in rabbit numbers had a measurable effect on parasite abundance.     

Prevalence of helminths in a cyclic snowshoe hare population

Five species of helminths were monitored in a population of snowshoe hares (Lepus americanus) near Rochester, Alberta, during 1961-1977. Prevalence of both Obeliscoides cuniculi and Protostrongylus boughtoni among young hares averaged about 50% by age 2 mo, then tended to level off. Prevalence of Taenia pisiformis (cysticerci) and Dirofilaria scapiceps rose more slowly, but continued to increase steadily beyond their mean levels of 8% and 1% at age 2 mo. There were well defined seasonal (within-year) cycles in prevalence of O. cuniculi and P. boughtoni that were generated evidently to a major degree by arrested development of larvae in fall and renewed development in late winter. It was hypothesized that renewed larval development was triggered (in February) in O. cuniculi by the seasonal rise of circulating pituitary gonadotrophins, and (in April) in P. boughtoni by the seasonal rise of gonadal androgens and estrogens. Indices to gonadal hormone levels in hares indicated that these increased most rapidly among males, and may have accounted for the higher prevalences of P. boughtoni in males during April-May. Neither T. pisiformis nor D. scapiceps exhibited conspicuous seasonal changes in prevalence. Maximum prevalence of T. pisiformis was attained at about 1 yr of age, whereas D. scapiceps increased among adult snowshoes through age 2 yr before stabilizing. Long-term (between-year) changes in prevalence of O. cuniculi, T. pisiformis, and D. scapiceps were correlated significantly with the cyclic hare population which declined from a peak in fall 1961 to a low in 1965-1966, rose to another peak by fall 1970, and declined again to a low in 1975. There was no detectable time lage between this "10-yr" cycle in hare density and the cycles of parasite prevalence among juveniles (less than 1 yr of age). Among adult hares, the cycle of O. cuniculi prevalence was likewise synchronous with that of the hare population, but the cycles of D. scapiceps and T. pisiformis lagged by approximately 1 and 2 yr, respectively. This lag in T. pisiformis prevalence was largely inexplicable to us. Our data on P. boughtoni were not suitable for analyses of between-year trends; nor were those for the fifth helminth, Taenia serialis (coenuri), because mean prevalence was less than 1% among both juveniles and adults. An apparent decline in T. serialis after the early 1950's, and its continued scarcity thereafter, paralleled a major change in numbers of one important definitive host--the red fox (Vulpes vulpes).(ABSTRACT TRUNCATED AT 400 WORDS)     

Evaluation of host preferences by helminths and ectoparasites among black-tailed jackrabbits in northern California

Fifty-four black-tailed jackrabbits (Lepus californicus) (five juvenile males, 22 adult males, five juvenile females, and 22 adult females) from Humboldt County, California (USA) were evaluated for sex and age-specific differences in parasite prevalences and intensities, 26 February through 30 October 1996. Nematodes found included Biogastranema leporis in 42 hares (78% prevalence), Rauschia triangularis in 26 hares (48%), Trichostrongylus calcaratus in 14 hares (26%), and Trichuris sylvilagi in two hares (4%). Cestodes found included Taenia sp. cysticerci in five hares (9%) and Taenia sp. coenurus found in one hare (2%). Ectoparasites found included the ticks Dermacentor variabilis on 10 hares (19%) and Ixodes spinipalpis (= Ixodes neotomae) on nine hares (17%), as well as the anoplurid louse Haemodipsus setoni on 12 hares (22%). No significant differences in the parasite prevalences or intensities were found between male and female jackrabbits; this was for all males and females collectively, juvenile males and females only, as well as adult males and females only. Combining male and female hosts, adult jackrabbits had a significantly higher prevalence of B. leporis and R. triangularis compared to juveniles. This is the first known report of Trichostrongylus calcaratus, Rauschia triangularis, Trichuris sylvilagi, and Dermacentor variabilis among black-tailed jackrabbits and the first known report of T. calcaratus and T. sylvilagi in the western USA. This is the first published report of I. spinipalpis, the vector for Lyme disease in California, on black-tailed jackrabbits.     

Helminths from introduced small mammals on Kerguelen, Crozet, and Amsterdam Islands (southern Indian Ocean).

Four monoxenous nematodes and 1 heteroxenous cestode were found in 4 species of introduced small mammals on isolated sub-Antarctic islands of the Indian Ocean. In the Kerguelen Archipelago, Syphacia obvelata, Passalurus ambiguus (Nematoda: Oxyuridae), and Rodentolepis straminea (Cestoda: Cyclophyllidae) were respectively found in the house mouse Mus musculus, the rabbit Oryctolagus cuniculus, and the black rat Rattus rattus. One accidental nematode, Trichostrongylus sp. (Nematoda: Trichostrongylidae), was also found in a black rat on Kerguelen. On Possession Island (Crozet Archipelago), R. straminea was present in the black rat. On Amsterdam Island, the brown rat R. norvegicus harbored 2 species, R. straminea and Nippostrongylus brasiliensis (Nematoda: Heligmonellidae). The small number of founder hosts and the depauperate terrestrial communities on these remote islands explain the low diversity in the helminth communities of these introduced mammals compared with continental populations.     

Parasite co-infection and interaction as drivers of host heterogeneity

We examined the hypothesis that the interaction between concomitant infecting parasites modifies host susceptibility, parasite intensity and the pattern of parasite distribution within the host population. We used a 26 year time series of three common parasites in a natural population of rabbits: two gastrointestinal nematodes (Trichostrongylus retortaeformis and Graphidium strigosum) and the immunosuppressive myxoma virus. The frequency distribution of nematodes in the host population and the relationship between host age and nematode intensity were explored in rabbits with either single or dual nematode infections and rabbits infected with the nematodes and myxoma virus. The aggregation of T. retortaeformis and G. strigosum among the rabbits varied with the nature of the co-infection both in male and female hosts. The two nematodes exhibited different age-intensity profiles: G. strigosum intensity increased exponentially with host age while T. retortaeformis intensity exhibited a convex shape. The presence of a secondary infection did not change the age-intensity profile for G. strigosum but for T. retortaeformis co-infection (either both nematodes or myxoma-nematodes) resulted in significantly greater intensities in adult hosts. Results suggest that multi-species infections contributed to aggregation of parasites in the host population and to seasonal variation in intensity, but also enhanced differences in parasitism between sexes. This effect was apparent for T. retortaeformis, which appears to elicit a strong acquired immune response but not for G. strigosum which does not produce any evident immune reaction. We concluded that concomitant infections mediated by host immunity are important in modifying host susceptibility and influencing heterogeneity amongst individual hosts.     

Parasitism by helminths in the grey-sided vole (Clethrionomys rufocanus) in northern Finland: influence of density, habitat and sex of the host

We have studied helminths from 532 grey-sided voles (Clethrionomys rufocanus) in two localities in northern Finland, representing different biogeographic zones, during 1978-1983. The helminth communities in the two study areas were similar, characterized by a small number of species (eight) compared to eastern Siberia and Japan, and by the dominance of a single anoplocephalid cestode Andrya kalelai. The prevalence of this helminth varied significantly among habitats, possibly because of differences in the distribution and abundance of the intermediate hosts, oribatid mites. Vole density did not explain the habitat differences in A. kalelai, nor did the prevalence of A. kalelai increase between 2 yr of sustained high density in the host population. A between-year increase in the prevalence of the larval cestode Taenia tenuicollis at Kilpisj?rvi was probably due to a simultaneous increase in the abundance of its definitive hosts, mustelids. The prevalence of A. kalelai was always higher in males; no sexual differences were detected in the larval T. tenuicollis.     

Helminths of sympatric black-tailed jack rabbits (Lepus californicus) and desert cottontails (Sylvilagus audubonii) from the high plains of eastern New Mexico

Thirty-five desert cottontails (Sylvilagus audubonii) and 35 black-tailed jack rabbits (Lepus californicus), occurring sympatrically near the Clovis-Portales area of eastern New Mexico were infected with four species of Eucestoda (adults of Raillietina salmoni and Raillietina selfi, larvae of Taenia pisiformis and Taenia serialis). Raillietina salmoni and T. pisiformis more commonly infected S. audubonii. Raillietina selfi was found in near equal prevalence in both host species. Taenia serialis was recovered only from L. californicus. Thus, three of the four helminth species were shared by both lagomorphs (Jaccard's coefficient = 75). Female hosts were most heavily infected with R. selfi and Taenia serialis.     

Taenia spp.: 18S rDNA microsatellites for molecular systematic diagnosis

The 18S rDNA gene of adult worms of Taenia parva found in Genetta genetta in the Iberian Peninsula and larval stages of T. pisiformis from the wild rabbit (Oryctolagus cuniculus) in Tenerife (Canary Islands) were amplified and sequenced. The sequences of the 18S rDNA gene of T. parva (1768 bp) and T. pisiformis (1760 bp) are reported for the first time (GenBank accession nos. AJ555167-AJ555168 and AJ555169-AJ555170, respectively). In 168 alignment positions microsatellites in the 18S rDNA of both taxa were detected for the first time (TGC in T. parva and TGCT in T. pisiformis) and differences in their sequences with different repetition numbers were observed. The use of nucleotide sequences of this gene in the resolution of systematic problems in cestodes is discussed with reference to the systematic status of Taenia spp. and mainly in human taeniids such as T. solium, T. saginata, and Asian human isolates of Taenia.     

Seasonal variation in gut morphology in wild rabbits (Oryctolagus cuniculus)

It is shown experimentally that diet composition affected the sizes of several digestive organs in juvenile rabbits collected from the wild. Stomach weight, small intestine volume and colon length were larger in rabbits reared on diets higher in fibre. Small intestine weight, however, was lower.
Analysis of natural variation in wild adult rabbits showed that all digestive organs (except for the appendix) were larger in rabbits living on chalk soils than in those living on sand or clay. Seasonal variation was more marked in males than in females, probably because of the sexes' different breeding requirements. Thus most male digestive organs were smaller between December and June than at other times of year. Female digestive organs did not show this pattern, and were largest in May/June, when the demands of pregnancy and lactation are highest. However, food quality may have been important too. Food quality is probably highest in April, and most digestive organs were then smallest (both sexes). The dry weights of stomach and small intestine did not fit these patterns. Stomach weight increased with the presence of the nematode parasite Graphidium strigosum and small intestine and caecal volumes with the presence of cestodes. Numbers of nematodes (Trichostrongylus retortaeformis and Passalurus ambiguus) in the caecum or colon, however, had no effects on organ sizes.
Data are presented on variation in intensity and prevalence of these helminth parasites with season and soil type.     

The potential role of habitat on intestinal helminths of mountain hares, Lepus timidus

Over the last century in the uplands of Scotland, the extent of heather moorland which supports high densities of mountain hares Lepus timidus has diminished and has gradually been replaced by large-scale commercial forestry plantations or expanding natural woodlands. The potential impact of such a change in land use on host-parasite interactions was investigated by comparing the intensity and prevalence of infection of hares by parasites in two separate habitats: a large hare-fenced young forestry plantation and the adjacent open moorland. Carcasses were collected in November 1990 from within both habitats and after the woodland had been enclosed for nine months. Age, sex, fatness (kidney fat index) and degree of infection of hares were noted. Two parasites were recorded: the nematode Trichostrongylus retortaeformis and the cestode Mosgovoyia pectinata. Clear differences in the intensity of infection of adults occupying the different habitats had occurred in the nine months since woodland enclosure. Adult mountain hares in the woodland had levels of infections approaching four times that observed in hares occupying the open moorland and although not significant, the prevalence of infection was greater in hosts inhabiting the woodland than the open moorland. It is suggested that the parasite-host relationship differs between the two habitats and as heather-dominated moorland landscapes become more fragmented with the increasing establishment of woodlands, the impact of parasites on the life history strategies of mountain hares needs to be reconsidered.