Abundance of Ticks (Acari: Ixodidae) Infesting the Western Fence Lizard, Sceloporus Occidentalis, in Relation to Environmental Factors

We examined the impact of environmental characteristics, such as habitat type, topographic exposure and presence of leaf litter, on the abundance of Ixodes pacificus ticks infesting the western fence lizard (Sceloporus occidentalis) at the University of California Hopland Research and Extension Center (HREC), Mendocino County, California. A total of 383 adult lizards were slip-noosed and examined for tick infestation in April and May 1998. At least 94% of the lizards were infested by ticks and at least 20% of the females and 33% of the males carried > 15 ticks. This intensive utilization of western fence lizards (which do not serve as natural reservoirs for Lyme disease spirochetes) by subadult ticks, is probably the primary reason for the low prevalence of infection with Borrelia burgdorferi in I. pacificus nymphs and adults previously recorded at the HREC. Tick loads were higher on male than female lizards. Also, male lizards were generally more heavily infested in late April than in late May. The prevalence of tick infestation exceeded 88% in all habitat types but males collected in woodland and grass/woodland edges had higher tick loads than those collected in open grassland. Male lizards captured in open, exposed grassland tended to carry heavier tick loads in northern/eastern, as compared to southern/western, exposures, and when leaf litter was present.     

The roles of birds, lizards, and rodents as hosts for the western black-legged tick Ixodes pacificus.

We compared the infestation by ixodid ticks of lizards, rodents, and birds collected simultaneously within areas representing common habitat types in Mendocino County, CA. Lizards were infested only by Ixodes pacificus Cooley and Kohls, birds by I. pacificus and Haemaphysalis leporispalustris (Packard), and rodents by I. pacificus, I. spinipalpis Hadwen and Nuttall, I. woodi Bishopp, Dermacentor occidentalis Marx, and D. variabilis (Say). Infestation by I. pacificus larvae and nymphs of lizards (Sceloporus occidentalis Baird and Girard; Elgaria spp.) and western gray squirrels (Sciurus griseus Ord) (means of 9-35 larvae and 5-6 nymphs per animal) was several times greater than for Neotoma fuscipes Baird woodrats, Peromyscus spp. mice, and birds (means of 0.9-3.5 larvae and 0-0.3 nymphs). Overall, Borrelia-refractory lizards accounted for 84% of I. pacificus larvae and 91% of nymphs collected from animals in dense woodlands. Bird species frequently utilizing tick-questing substrates such as leaf litter (guild I birds) were more heavily infested by I. pacificus subadults (5.2 larvae and 1.0 nymphs per bird) than guild IV birds with minimal perceived contact with tick-questing substrates (0.08 larvae and 0.06 nymphs per bird). Notably, guild I birds carried similar larval loads and at least 20-fold higher nymphal loads relative to woodrats and mice. Only guild IV birds carried as few I. pacificus nymphs as did these rodents. The ratios of larvae to nymphs suggest that, relative to birds, lizards, and squirrels (infested by 1.3-6.0 larvae per nymph), nocturnally active ground-dwelling rodents such as woodrats and mice are underutilized by the nymphal stage (69 to >100 larvae per nymph). The western gray squirrel and guild I-II birds (e.g., the dark-eyed junco, Junco hyemalis [L.]) were the only potential reservoirs of Borrelia burgdorferi Johnson, Schmid, Hyde, Steigerwalt, and Brenner (the causative agent of Lyme disease in North America) that were frequently infested with both I. pacificus larvae and nymphs and commonly utilized dense woodland habitats.     

Habitat-related variation in infestation of lizards and rodents with Ixodes ticks in dense woodlands in Mendocino County, California

During the spring and early summer of 2002, we examined the relative importance of Borrelia-refractory lizards (Sceloporus occidentalis, Elgaria spp.) versus potential Borrelia burgdorferi sensu lato (s.l.)-reservoirs (rodents) as hosts for Ixodes pacificus immatures in 14 woodland areas (six oak, five mixed oak/Douglas fir, and three redwood/tanoak areas) distributed throughout Mendocino County, California. Lizards were estimated to serve as hosts for 93-98% of all larvae and > or =99.6% of all nymphs infesting lizards or rodents in oak woodlands and oak/Douglas fir sites in the southern part of the county. In redwood/tanoak woodlands and oak/Douglas fir sites in northern Mendocino County, the contribution of rodents to larval feedings reached 36-69% but lizards still accounted for 94-100% of nymphal bloodmeals. From late April to mid-June, I. pacificus larvae were recovered from 95 to 96% of lizards and dusky-footed woodrats (Neotoma fuscipes) and from 59% of Peromyscus spp. mice. In contrast, 99% of lizards but few woodrats (15%) and none of the mice were infested by nymphs. Comparisons of tick loads for 19 lizard-Peromyscus spp. mouse pairings, where the lizard and mouse were captured within 10m of each other, revealed that the lizards harbored 36 times more larvae and >190 times more nymphs than the mice. In oak woodlands, loads of I. pacificus larvae decreased from late April/early May to late June for S. occidentalis lizards but increased for Peromyscus spp. mice. We conclude that the relative utilization of Borrelia-refractory lizards, as compared to rodents, by I. pacificus larvae was far higher in dry oak woodlands than in moister habitats such as redwood/tanoak and oak/Douglas fir woodlands in northern Mendocino County. Non-lizard-infesting potential enzootic vectors of B. burgdorferi s.l. (I. angustus and I. spinipalpis) were recorded from rodents in three of six oak woodland areas, two of five oak/Douglas fir woodland areas, and two of three redwood/tanoak woodland areas.     

Impact of the experimental removal of lizards on Lyme disease risk

The distribution of vector meals in the host community is an important element of understanding and predicting vector-borne disease risk. Lizards (such as the western fence lizard; Sceloporus occidentalis) play a unique role in Lyme disease ecology in the far-western United States. Lizards rather than mammals serve as the blood meal hosts for a large fraction of larval and nymphal western black-legged ticks (Ixodes pacificus--the vector for Lyme disease in that region) but are not competent reservoirs for the pathogen, Borrelia burgdorferi. Prior studies have suggested that the net effect of lizards is to reduce risk of human exposure to Lyme disease, a hypothesis that we tested experimentally. Following experimental removal of lizards, we documented incomplete host switching by larval ticks (5.19%) from lizards to other hosts. Larval tick burdens increased on woodrats, a competent reservoir, but not on deer mice, a less competent pathogen reservoir. However, most larvae failed to find an alternate host. This resulted in significantly lower densities of nymphal ticks the following year. Unexpectedly, the removal of reservoir-incompetent lizards did not cause an increase in nymphal tick infection prevalence. The net result of lizard removal was a decrease in the density of infected nymphal ticks, and therefore a decreased risk to humans of Lyme disease. Our results indicate that an incompetent reservoir for a pathogen may, in fact, increase disease risk through the maintenance of higher vector density and therefore, higher density of infected vectors.     

Differential detachment from resting hosts of replete larval and nymphal Ixodes ticks

To determine whether replete subadult Ixodes ticks detach more frequently from resting than from active hosts, diverse rodents and lizards were caged in an apparatus designed to record the ticks' sites of detachment relative to the resting site of the host. Replete larval Ixodes ricinus and Ixodes dammini accumulated mainly beneath the resting places of the mice (Apodemus agrarius and Peromyscus leucopus) most frequently parasitized in nature. Although nymphal I. ricinus similarly detached where these mice rested, nymphal I. dammini detached more randomly. When lizards were used as hosts, both subadult stages of I. ricinus tended to detach away from their main resting sites; these ticks detached from squirrels more randomly. Detachment ratios for other rodent hosts, that are abundantly infested by the larvae of these ticks in nature (Apodemus flavicollis and Clethrionomys glareolus), could not be derived because nymphs generally failed to attach. Our observations are consistent with reports that both subadult stages of I. dammini, but not the adult, feed on the same kind of nest-dwelling hosts and that the host range of I. ricinus is less focused. Detachment of mouse-feeding larvae from resting mice promotes subsequent nymphal attachment to conspecific hosts, and the absence of such behavior among nymphs facilitates access of the resulting adults to deer.     

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 of Amblyomma cooperi (Acari: Ixodidae) using capybaras (Hydrochaeris hydrochaeris) as hosts

The life cycle of Amblyomma cooperi was evaluated under laboratory conditions testing different host species. Larval infestations were performed on chickens (Gallus gallus) and capybaras (Hydrochaeris hydrochaeris). Nymphal infestations were performed on G. gallus, H. hydrochaeris, guinea pigs (Cavia porcellus) and wild mice (Calomys callosus). Infestations by adult ticks were performed only on capybaras. All free-living stages were observed in darkness at 27 degrees C and RH 85%. Capybaras were significantly (p < 0.05) the most suitable hosts for immature ticks, with the highest larval (63.6%) and nymphal (48%) recovery. Larval and nymphal feeding and premolt periods were significantly different (p < 0.05) between ticks fed on different host species. Male nymphs showed premolt period significantly shorter (p < 0.05) than female nymphs. The overall sex ratio of adult ticks was 0.92:1 (M:F). Infestations by adult ticks on capybaras yielded more than 76% of engorged female recovery. Only three out of 33 engorged females fed on capybaras did not lay fertile eggs. The life cycle of A. cooperi in laboratory, reported for the first time, was completed in an average period of 189.4 days. During the premolt period, all A. cooperi engorged nymphs secreted distinct blackish drops, which seem to be inherent to this species. Our results, associated data in the literature, confirm the high suitability of capybaras for the adult stage of A. cooperi and also indicate this animal species as a primary host for immature stages of A. cooperi in nature. On the other hand, the results of larval and nymphal infestation on chickens and guinea pigs suggest that birds and wild guinea pigs, which are also present in the distribution area of A. cooperi in South America, could be potentially infested by A. cooperi immature stages in nature.     

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.     

Host association and seasonal activity of Amblyomma americanum (Acari: Ixodidae) in Missouri

From June 1993 through June 1996, 2,260 adult, 4,426 nymphal, and 2,178 larval lone star ticks Amblyomma americanum (L.) were collected in Missouri from vertebrate hosts and by dragging a cloth over vegetation. Prevalence, mean intensity, and relative abundance of each stage varied among hosts. The relative abundance of adult lone star ticks was highest on white-tailed deer, but this stage was also collected from raccoons, opossum, red fox, coyotes, and wild turkey. Nymphs were collected from gray squirrels, eastern cottontail rabbits, opossums, red fox, Carolina wren, and bobwhite quail, but the highest relative abundance occurred on wild turkey, white-tailed deer, and raccoons. Eastern cottontail rabbits, white-tailed deer, raccoons, and squirrels had the highest relative abundance of larval lone star ticks, but they were also found on opossums and wild turkey. The activity of adult lone star ticks was greatest from May through July. The activity for nymphs was highest from May through August, and for larvae, July through September.     

Vectorial capacity of North American Ixodes ticks.

Ixodes dammini, the vector of Lyme disease and babesiosis, is distributed in various locations in the northeastern quadrant of the United States and nearby Canada. The life cycle of this tick, which includes larval, nymphal, and adult stages, spans at least two years. The tick over-winters between larval and nymphal feeding. Horizontal transmission of pathogens is facilitated by a feeding pattern in which both the larval and nymphal stages feed on the white-footed mouse, Peromyscus leucopus, and by a seasonal pattern of activity in which nymphs precede larvae. The species range appears to have expanded from a single island location, and has invaded new sites since the 1940s, some as recently as 1980. This increased abundance appears to be related to the increased abundance of deer, the preferred host of the adult stage. I. muris predominated in coastal Massachusetts before I. dammini became abundant, but is probably now extinct. I. scapularis, which is present in the southern U.S., is a poor vector of mouse parasites because about 90 percent of these immature ticks feed on lizards. To the extent that horizontal transmission occurs, we suggest that mice serve as the principal reservoir for the Lyme spirochete as well as Babesia microti.     

Seasonal abundance and activity of the tick Dermacentor silvarum in northern China

Studies on the seasonal abundance and activity of larvae, nymphs and adults of Dermacentor silvarum (Ixodida: Ixodidae), both on and off the host, were carried out over a 2-year period from February 2008 to January 2010 in northern China. During the study period, feeding ticks were removed weekly from domestic sheep and free-living ticks were collected weekly by flag-dragging in four different habitat types. Rodents were captured and inspected for immature ticks from June to September 2008. The results of this study suggest that the ticks mainly reside in shrubs and complete only one generation per year with sequential seasonal distribution and little overlap between the activity of the larvae and nymphs. Adults were most prevalent from late February to late May and peaked in number in mid-April. Larvae were found from early June to early September and peaked in number in mid-July. Nymphs were mainly distributed from late June to late September and peaked in mid-August. Adult and nymphal D. silvarum were found primarily on the ears of sheep, but no larvae were found on sheep. Additionally, an overwintering male adult population was detected on sheep after October, but no free-living adults were found by dragging.     

Evaluation of 4-poster acaricide applicators to manage tick populations associated with disease risk in a Tennessee retirement community

In 1993, four residents of a retirement community in middle Tennessee were hospitalized with symptoms of ehrlichiosis causing community managers to implement mitigation methods to reduce tick numbers. For the past four years, managers have utilized 4-poster acaricide applicators that aim to reduce disease risk to residents by killing ticks that feed on deer. To determine the efficacy of this technique, we assessed Amblyomma americanum abundance in the vicinity of the devices by dragging 400 m vegetation transects once per month while ticks were active. In 2009, adult tick activity peaked in May, nymphal tick activity peaked in June, and larval activity peaked in September. Close to 4-poster devices, larval, nymphal, and adult tick abundances were reduced by 91%, 68%, and 49%, respectively (larval and nymphal p<0.001, adult p=0.005), relative to nearby untreated areas. No significant reduction in nymphal or adult A. americanum ticks was evident >300 m from 4-poster devices, however a ～90% reduction in larvae was observed to our sampling limit (400 m). At the low density at which these devices are currently being used (average distance between devices = 6.6 km), we conclude that they will have little large-scale effect on the health risk posed by ticks in this community.     

Lacerta bilineata (Reptilia,Lacertidae) as a host of Ixodes ricinus (Acari,Ixodidae) in a protected area of northern Italy

A survey on the importance of Lacerta bilineata as host of Ixodes ricinus was conducted in Northern Italy over a three-year period. A total of 202 western green lizards were captured and a total of 2349 ticks were collected. All ticks were identified as I. ricinus; 53.2% and 46.7% were at the larval and nymphal stages, respectively. Tick number and prevalence were higher in males than in females, especially from April to June during the host breeding period. The level of tick infestation increased with lizard age and size. The number of ticks collected on adult lizards peaked in June and in August. Infestation levels appear to be related to lizard activity patterns and behaviour. Tick number and prevalence also varied in relation to host habitat, infestation being higher in lizards from areas with hard vegetation cover.     

Parasitism of lizards by immature stages of the blacklegged tick,Ixodes scapularis (Acari,Ixodidae)

From 1982–1985 and 1993–1999, a total of 309 individual reptiles, mostly lizards and snakes, belonging to 12 species (American alligator, six lizard species, five snake species) was captured on St. Catherine's Island, Liberty County, Georgia, USA, and examined for ticks. Three lizard species, the broad-headed skink Eumeces laticeps, southeastern 5-lined skink Eumeces inexpectatus, and eastern glass lizard Ophisaurus ventralis, were severely infested with larvae and nymphs of the blacklegged tick, Ixodes scapularis. Ticks were not found on any of the other reptile species. Overall, 80% of 65 E. inexpectatus examined were parasitized by a mean intensity of 21.5larvae and 88% were parasitized by a mean intensity of 4.8 nymphs. Corresponding figures for E. laticeps (n=56) were 93% and 51.3 for larvae and 89% and 7.4 for nymphs, and for O. ventralis (n=3) were 67% and 22.5 for larvae and 100% and 21.3 for nymphs. Larvae and nymphs attached along the lateral grooves of O. ventralis. Nymphs attached mainly behind the ears and in the foreleg axillae whereas larvae mainly attached to these sites and on the hindlegs in Eumeces spp. Seasonally, both larvae and nymphs were recorded on lizards from April through October. A unimodal larval peak was recorded in May or June. Seasonal data for nymphs did not reveal any distinct peaks but small bimodal peaks in mean intensities may have occurred (one in early summer, the other in late summer)suggesting that some ticks complete their life cycle in one year, and others in two years, on St. Catherine's Island. Potential epidemiological consequences of these findings with respect to Lyme disease in the southeastern United States are briefly addressed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Heterogeneity of resource utilization in a population of the Australian reptile tick, Aponomma hydrosauri (Denny)

Abstract. 1. The distribution of adult males, nymphs and larvae of the reptile tick Aponoma hydrosauri on their host, Trachydosaurus rugosus , in a study area near Tickera, South Australia, did not follow the Poisson distribution predicted if hosts were infested at random.
2. The non-random distribution of males can be accounted for by geographical heterogeneity in distribution. Adult females also showed geographical heterogeneity in their distribution pattern, but infested their hosts at random over all.
3. Larvae and nymphs show distribution patterns which change both with the time and the locality. However, when these biases are removed the distribution of larvae and nymphs on their hosts is still non-random.
4. Larger lizards tend to have larger numbers of ticks, but this cannot alone account for the clumped distribution.
5. A model is proposed relating the tick distribution pattern to the particular problems of a resource which is hard to find, but plentiful once found.     

Respiratory metabolism of the soft tick,Ornithodoros turicata (Dugès)

The rate of oxygen consumption was investigated in fed larval, nymphal and adult Ornithodoros turicata ticks and in starved nymphal and adult ticks. Oxygen consumption rate of fed adult ticks increased with increasing temperature. The metabolic rate of adult ticks was affected by starvation whereby starved adult ticks showed a significantly lower oxygen consumption than their fed counterparts. The oxygen consumption rate of fed female ticks was significantly higher than that of fed males but, there was no significant difference between the oxygen consumption rates of starved female versus starved male ticks. Oxygen consumption of fed larvae was significantly greater than those of fed first through third instar nymphs. Fed and starved nymphal ticks as well as fed adult ticks ventilated continuously. In contrast, starved adults ventilated discontinuously. The ability to reduce metabolic rate, plus the capability to ventilate discontinuously allow O. turicata adults to cope with prolonged starvation.     

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.     

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.     

Questing activity of Rhipicephalus appendiculatus (Acari: Ixodidae) nymphs: a random process?

Abstract.  In Zambia, an experiment under quasi-natural conditions indicated that Rhipicephalus appendiculatus nymphs react to prevailing climatic conditions. Higher temperatures and higher vapour pressure deficits lead to decreased activity. The majority of nymphs (>75%) were recorded at ground level. Simulations showed that larval phenology and temperature during the nymphal premoulting period largely explain the seasonal abundance patterns of nymphs, as observed on cattle, given the absence of a behavioural diapause. Consequently, the effect of climate, as observed in our studies, is masked. However, the results of the present study indicate that daily climatic conditions probably have a much larger effect on the transmission dynamics of Theileria parva . The vertical distribution of questing instars is a function of temperature and humidity. In years of unfavourable conditions, nymphs might feed mainly on hosts other than cattle, and this could govern the infection prevalence in the adult population. This suggestion is supported by previous epidemiological studies.     

Growth and development of winter tick, Dermacentor albipictus, on moose, Alces alces

Moose, Alces alces, were infested with 21,000 or 42,000 larval Dermacentor albipictus at the end of September. Larvae grew rapidly and molted to the nymphal stage 10-22 days after infestation. The nymphal stage lasted approximately 3 mo until mid-January and was characterized by a diapause. The diapause is likely an adaptation to survival in cold climates. Nymphs started engorging in January and adults were seen with increasing abundance from mid-January to March and April. The minimum parasitic period was 175 days. Growth of larvae and nymphs was similar on moose given different numbers of larvae and was generally similar between a moose infested in November and moose infested earlier. Dimensions and stages of development throughout the parasitic phase are given. Game enforcement officers are encouraged to use these data for determination of season of death of moose.