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Eastern moose (Alces alces americana) are heavily parasitized by winter ticks (Dermacentor albipictus), the dominant cause of increased calf mortality in the northeastern US. Although much work has focused on the direct negative effects of winter tick on moose, it remains unknown whether diseases transmitted by ticks may also affect moose health or pose a risk to other species. We explored the role that moose and winter ticks play in transmission of the tick-borne bacterial pathogens, Anaplasma spp., which cause mild to severe illness in humans and domestic animals. Our objectives were to 1) estimate the prevalence of Anaplasma spp. in moose and winter ticks; 2) determine the phylogenetic placement of these strains with respect to those found in other hosts and vectors; and 3) explore risk factors of Anaplasma infection in moose. A total of 157 moose (142 calves, 15 adults) were captured in western (n=83) and northern (n=74) Maine in 2017 and 2018. We screened for Anaplasma spp. in moose whole blood samples using a genus-specific PCR assay targeting the 16S rRNA gene. Over half (54%) of the moose were infected with Anaplasma bacteria, with a greater proportion of moose harboring Anaplasma-infections in the western (67%) versus northern study areas (38%). Male moose exhibited a higher prevalence than did females (63% vs. 47%). In contrast, Anaplasma spp. prevalence in winter ticks was low (<1%). Sequencing and phylogenetic analysis revealed that the single Anaplasma strain in moose was highly divergent from the strain in winter ticks and most closely related to an uncharacterized North American cervid strain. We conclude that winter ticks are unlikely to play a significant role in Anaplasma transmission to moose; however, high infection prevalence warrants further investigation into the impacts of Anaplasma spp. infection on moose health. [ABSTRACT FROM AUTHOR]

Many wildlife populations are experiencing a variety of environmental pressures due to the direct and indirect consequences of a changing climate. In the northeast, USA, moose Alces alces are declining in large part because of the increasing parasitism by winter tick Dermacentor albipictus, facilitated by high host density and optimal environmental conditions. To test this hypothesis, and better understand the influence of this interaction on the stability of the regional population, we constructed a population viability model using data collected through comprehensive survival and productivity studies in 2002–2005 and 2014–2018 in northern New Hampshire. Years of heavy tick infestation (epizootics) saw a marked reduction in calf survival (< 50%), adult calving (< 60%), twinning rate (< 5%) and complete loss of yearling productivity. We conducted population viability analysis using VORTEX ver. 10.2 to model this moose population for 40 years using mean demographics from both time periods, including environmental variation measured in the field during winter tick epizootic (2002, 2014, 2015, 2016) and non-epizootic (2003, 2004, 2005, 2017) years. This exercise highlights the influence of winter tick infestation on the trajectory of the population with the potential for rapid population growth or decline depending on the frequency of epizootics. We suggest a shift in moose management strategy focused on lowering moose density, assuming continued influence of climate change on the host–parasite relationship. [ABSTRACT FROM AUTHOR]

Moose naturally acquire soil fungi on their fur that are entomopathogenic to the winter tick, Dermacentor albipictus. Presumed to provide a measure of on-host tick control, it is unknown whether these soil fungi impact subsequent off-host stages of the tick. Eggs and resultant larvae originating from engorged, adult female winter ticks collected from dead calf moose (Alces alces) were used to investigate the presence and extent of fungal infection. Approximately 40% of eggs and larvae were infected, almost exclusively by the fungus Scopulariopsis brevicaulis (teleomorph Microascus brevicaulis: Microascaceae, Ascomycota). Eggs analysed on the day of oviposition and day of hatching had high frequency (40%) of S. brevicaulis, whereas the frequency in eggs harvested in utero was minimal (7%); therefore, exposure occurs pre-oviposition in the female's genital chamber, not by transovarial transmission. At hatching, larvae emerge containing S. brevicaulis indicating transstadial transmission. Artificial infection by topical application of eggs and larvae with a large inoculum of S. brevicaulis spores caused rapid dehydration, marked mortality; pathogenicity was confirmed by Koch's postulates. The high hatching success (>90%) and multi-month survival of larvae imply that S. brevicaulis is maintained as a natural pathobiont in winter ticks. [ABSTRACT FROM AUTHOR]

Certain moose (Alces alces) populations in the northeastern United States are in decline because of increasing frequency of winter tick (Dermacentor albipictus) epizootics characterized by high calf mortality and reduced productivity in the population. Previous research identified soil fungi in wallows of bull moose that are pathogenic to larval winter ticks. We analysed skin/hair samples of moose harvested in Maine, USA, to further understand this fungus–tick–moose relationship. A total of 32 keratinophilic fungi from moose fur and skins were isolated, predominately Penicillium, Cladosporium, Aspergillus, Alternaria, Trichoderma, Beauveria, and Scopulariopsis. Each displayed pathogenic activity against unfed winter tick larvae (sole infestation stage) by topical application of spore inoculums and Koch's postulates. Bulls had more entomopathogenic fungi, pathogenic fungal isolates to ticks, and infected larvae than cows and calves. We propose a disease cycle with interactions of soil fungi, moose, and on-host winter tick larvae killed by entomopathogenic soil fungi. Bull moose presumably experience enhanced exposure and protection from abundant soil fungi due to their wallowing behaviour during the breeding season that is coincidental with questing by winter tick larvae. [ABSTRACT FROM AUTHOR]

High calf mortality attributed to winter tick (Dermacentor albipictus) parasitism occurs in moose (Alces alces) populations along their southern range in the northeastern United States. We analyzed habitat use of cow and calf moose during the critical drop-off and questing periods in the winter tick life cycle to determine a potential relationship between tick density and habitat. We measured habitat use using geospatial analyses of locational data from > 200 radio-marked animals at 3 sites in New Hampshire and Maine. Moose selected for optimal habitat, defined as 4-16 year-old forest openings, regardless of season or site; this was the only land cover type used more than available (1.1-2.1X availability in home range, 1.2-3.1X availability in core range). Further, the proportional availability of optimal habitat within overlapping portions of seasonal home and core ranges exceeded the absolute proportion of optimal habitat within any one range. Temporal use of optimal habitat, which is available in relatively low proportion (15-20%) across the landscape, likely exceeds the geospatial estimates of use because moose spend 30-40% of daily activity foraging. We conclude that disproportionally abundant densities of winter ticks exist in this preferred cover type because of its selective use during the drop-off and questing periods of winter ticks. [ABSTRACT FROM AUTHOR]

We report that larvae ofDermacentor albipictuscan detect CO2from a dry ice source at distances up to 3.0 m and initiate increased activation, movement and questing behaviour. The maximum distance that larvae spread was ~0.5 m after 24 h, compared to <0.1 m without CO2as an excitant. In Petri dish bioassays, 2,6-dichlorophenol, methyl salicylate ando-nitrophenol acted as attractants, albeit with mixed results at 1.0 m, 2.0 m and 3.0 m, respectively. The 2,6-dichlorophenol target was the most effective, although few larvae crawled >1.0 m and CO2was required as a carrier. In Petri dish bioassays, larvae arrested into dense aggregations on eggshells collected after hatching. This arrestment was associated with the eggshell lipids cholesteryl oleate and cholesteryl palmitate, implying that cholesteryl esters act as assembly pheromones for larvae. Faster response was evident at 65% relative humidity (RH) versus 95% RH. Larvae were not arrested by tick excreta, guanine or related purines. Although larvae can be activated from long distances, in the field they remain fairly localized around the egg mass (hatching location), have limited dispersal and are more responsive in drier conditions. Given their arrestant properties, cholesteryl oleate and cholesteryl palmitate may prove useful in pheromone-assisted control methods becauseD. albipictuslarvae are the single stage of infestation to the host. [ABSTRACT FROM AUTHOR]

Camera-traps are increasingly used to estimate wildlife abundance, yet few studies exist for small-sized carnivores or comparing efficacy against traditional methods. We developed a camera-trap to identify the unique ventral patches of American martens (Martes americana). Our method was designed to: (1) determine the optimal trap configuration to photograph ventral patches; (2) evaluate the use of temporally clustered photographs to determine independence and improve identification; and (3) determine factors that influence identification probability. We tested our method by comparing camera- and live-trap density estimates using spatial capture-recapture (SCR) models. The ventral patches of radio-collared martens were most visible when traps were placed 15-20 cm above a feeding platform. Radio-collared martens (n = 14) visited camera-traps for long periods (median = 7min) with long intervals between visits (median = 419min), and visits by different martens at the same trap <15 min apart was infrequent (n = 3) during both years. Similarly, there was complete agreement among observers that clustered photos of un-collared martens were always of the same individual. Pairwise agreement was high between observers; eight un-collared martens were identifiable by consensus on 90% (54 of 60) of recorded visits. Factors influencing identification probability were directly related to the time martens spent feeding at traps (β = 0.143, P = 0.01) and inversely proportional to the time that elapsed since traps were baited (β = -0.344, P = 0.006). Density estimates were higher and more precise for camera-trapping (0.60, 0.35-1.01 martens/km²) than live-trapping (0.45, 0.16-1.22 martens/km²), providing evidence that SCR density estimates may be biased when capture heterogeneity is present, yet cannot be accounted for due to small sample size. Our camera-trap method provides a minimally invasive and accurate tool for monitoring marten populations. [ABSTRACT FROM AUTHOR]

To explore how the one-host tick Dermacentor albipictus survives off-host, we determined water balance characteristics of eggs and larvae. In contrast to eggs, larvae lost water fast, absorbed water vapour and died from a low amount of water loss. Placing mated females under near water-saturated conditions yielded larvae that absorbed and survived only at high relative humidities. We observed larvae forming clusters. Water loss rate of larvae decreased as group size increased. In response to day length, larvae lost water slower under long day than short day conditions. Larvae could switch quickly between fast water loss rate (short day) and slow water loss rate (long day) within a day, independent of the photoperiodic experience of the egg or mother. Long day exposure had no effect on water relations, incubation and survival of eggs. We conclude that eggs are modified for water conservation. Larvae survive by having a water balance maternal effect; i.e. capacity for water vapour absorption in the larva is determined by the relative humidity experienced by the mother. Larvae regulate water loss behaviourally through larval clustering and summer quiescence. Understanding these water balance attributes and effects of desiccation on egg and larval survival during summer are important to better predict the potential population impacts of winter ticks on moose. [ABSTRACT FROM AUTHOR]

Abundance indices suggested that the moose Alces alces population in northern New Hampshire was stable despite favourable habitat and conservative harvest. Causes and rates of mortality were unknown because moose reproduction and survival was unstudied in the region. Our study was designed to investigate the dynamics of the regional population in 2002-2005.Atotal of 92 moose (33 cows and 59 calves) were captured and fitted with radio-collars (VHF=83, GPS= 9). Parturition ranged from 8 May to 13 July (median = 19 May) with 78% of births occurring during 13-27 May. Calving rate of yearlings and adults (>2 years old) averaged 30 and 85%, respectively; twinning rate was 11%. Analysis of reproductive data from harvested cows (1988-2004) indicated that the average weight of adult cows increased but their corpora lutea count declined from ; 1.4 to 1.2/cow. Both ovulation rate and average weight of yearling cows declined about 25 and 4%, respectively. There were 39 mortalities (49% calves) with winterkill/parasite (41%), vehicle collision (26%) and hunting (18%) as the leading causes. Major sources of mortality of radio-marked cows were human-related; survival was 0.87. Annual calf survival was 0.45. Unmarked calf (0-2 months of age) survival was 0.71 with 76% of mortality in the first month of life. Radio-marked calves (∼7-12 months of age) had a survival rate of 0.67; 74% of the mortality was winterkill/parasite related. Calf mortality was concentrated (88%) in late winter-early spring. The unseasonably warm and snowless fall in 2001 probably favoured high tick transmission and increased tick loads on moose that resulted in high calf mortality (0.51) and measurable cow mortality (10%) in mild winter-spring 2002. Documentation of substantial tick-related mortality of radio-marked moose calves was unique to this study. The stability of the moose population probably reflects the variation in annual recruitment and lower fecundity of yearling cows associated with heavy infestations and epizootics of winter tick. Given that fertility, calving rate and body condition of adult cows, and summer calf survival are annually high, the population should recover from tick epizootics that periodically inhibit population growth. [ABSTRACT FROM AUTHOR]

• Agent-based models, parameterized using existing studies on moose (Alces alces) and winter ticks (Dermacentor albipictus), can be used to simulate populational trends similar to those observed in the field and are therefore a promising tool for managing moose populations in northern New England against epizootic events. • Highly concentrated optimal habitat is an important driver of winter tick epizootics. • Winter tick infestations are positively correlated with moose density. • At densities ≤ 0.40 moose/km2, weather conditions including summer drought and late onset of snow become less important in predicting epizootics than at densities > 0.40 moose/km2. • The number of winter ticks aggregating in a cluster is an important variable to consider when modeling winter tick epizootics and warrants focus in field studies. North American moose (Alces alces) populations along the southern extent of their range have been experiencing high levels of calf mortality in recent years. In New England, this phenomenon has been linked to extensive blood loss resulting from extreme winter tick (Dermacentor albipictus) parasitism. Moose are symbolic of the region and generate income through tourism and the auction of hunting permits; thus, successive years of greater than 50% calf mortality (epizootics) are of concern to wildlife managers and others. It is hypothesized that high localized moose density coupled with climate change are the driving forces behind moose-winter tick epizootics; however, the impact that variable combinations of these and other factors have on the occurrence and magnitude of epizootics has not been previously simulated. We, therefore, developed and implemented a spatially explicit agent-based model with two model environments, each representative of a distinct location within an ongoing field study site in northern New Hampshire that differed in the proportional availability of optimal moose habitat. Three experiments were devised to test the sensitivity of the outcome variables, calf infestation level and calf mortality, to 1) winter tick abundance, 2) winter tick aggregation, and 3) moose density, for the length of the winter tick questing period. Each experiment was conducted in both model environments. This model generated similar mortality levels to those measured in the field study under representative moose density and weather conditions. Additionally, the modeled moose agents and the radio-collared moose in the field reflect comparable habitat usage. While the infestation levels reported by calf agents are considered conservative, we believe that future versions of this model, parameterized with more accurate estimates of winter tick abundance and moose density, may be an effective tool for managing moose against winter tick parasitism. [ABSTRACT FROM AUTHOR]