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Recent decline in New Hampshire's moose (Alces alces) population is attributed to sustained parasitism by winter ticks (Dermacentor albipictus) causing high calf mortality and reduced productivity. Location of larval winter ticks that infest moose is dictated by where adult female ticks drop from moose in April when moose preferentially forage in early regenerating forest in the northeastern United States. The primary objectives of this study were to: 1) measure and compare larval abundance in 2 types of regenerating forest (clear-cuts and partial harvest cuts), 2) measure and compare larval abundance on 2 transect types (random and high-use) within clear-cuts and partial harvests, and 3) identify the date and environmental characteristics associated with termination of larval questing. Larvae were collected on 50.5% of 589 transects; 57.5% of transects in clear-cuts and 44.3% in partial cuts. The average abundance ranged from 0.11-0.36 ticks/m2 with abundance highest (P < 0.05) in partial cuts and on high-use transects in both cut types over a 9-week period; abundance was ~2 × higher during the principal 6-week questing period prior to the first snowfall. Abundance (collection rate) was stable until the onset of < 0°C and initial snow cover (~15 cm) in late October, after which collection rose temporarily on high-use transects in partial harvests during a brief warm-up. The higher abundance of winter ticks on high-use transects indicates that random sampling underestimates tick abundance and relative risk of infestation of moose. Calculating an annual index of infestation of winter ticks on moose is theoretically possible by integrating 3 factors: the infestation of harvested moose in October, the length of the questing period, and assuming a stable collection rate during the questing period. [ABSTRACT FROM AUTHOR]

Snow urine samples collected in northern New Hampshire, USA were used to measure urea nitrogen (UN) and creatinine (C) content to develop ratios for tracking the nutritional restriction of individual moose (Alces alces) through winter (2014-2017), inclusive of the adult winter tick (Dermacentor albipictus) engorgement period. Samples (n = 215) were collected from 55 moose (38 calves, 17 cows) on a twice monthly schedule from late January through snowmelt or calf mortality (March - early April). Early winter UN:C ratios from cows, surviving calves, and calves that ultimately died from infestation of winter ticks were similar and reflected a normal winter diet low in protein. A heightened UN:C ratio (> 3.5 mg/dL) was measured in March which aligned with peak feeding by adult winter ticks, and presumably reflected accelerated protein deficit associated with blood loss. This increase was not observed population-wide despite shared habitat, occurring only in calves with mortal weight loss and anemia associated with heavy winter tick infestation. Measurement of UN:C ratios from snow urine samples proved an effective method to measure the temporal impact of winter tick infestation, and March samples can support other metrics used to estimate calf mortality. [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]

An isolate of the soil fungus Scopulariopsis brevicaulis was identified from the surface of female winter ticks (Dermacentor albipictus) collected from recently dead moose (Alces alces) calves in New Hampshire in the northeastern United States. It was the sole isolate, and it matched with 98% nucITS similarity (molecular systematics Blast match) to S. brevicaulis species from soil and other tick species. Inoculation of tick larvae and eggs with 108 spores/mL + 0.05% Tween (aqueous inoculum) resulted in mortality, reduced survival time, and recovery of S. brevicaulis from within tick tissues. Rapid water loss and death from dehydration were the pathogenic consequences of the fungal infection. Three entomopathogenic fungal isolates from laboratory culture (Beauveria bassiana, B. caledonica, and Metarhizium anisopliae) inoculated concurrently at the same dose, were slightly less pathogenic to eggs than larvae of winter ticks. We conclude that S. brevicaulis imposes a limitation on the freeliving stages of the winter tick population in specific environmental conditions, but commercial fungal treatments as used in local situations to control ticks, are impractical as a means of controlling winter tick density across moose habitats. [ABSTRACT FROM AUTHOR]

Moose (Alces alces) populations in northern New Hampshire and western Maine experienced 3 successive years of high winter tick infestations (epizootics) in 2014-2016 that resulted in late-winter calf mortality rates >70%. To assess productivity in these populations, we measured fecundity rates of yearling and adult cow moose, and neonatal and summer calf survival. Parturition, fecundity, and survival were measured via direct observation by stalking VHF and GPS radio-collared cows (n = 177) in May-August, 2014-2016. Calving rates for yearlings and adults averaged 0 and 57%, respectively; there was no twinning documented. Summer calf survival to August was high overall (83%), with 85% of the mortality occurring in the first week of life. Calving and twinning rates declined since last measured in New Hampshire in 2002-2005 and were below the North American average; conversely, summer survival of calves was considered normal. Given that optimal habitat has increased in the past 15 years in the study area that is dominated by commercial forestry, lower productivity is presumably related to the additive impacts of successive winter tick epizootics on year-round condition of cows. [ABSTRACT FROM AUTHOR]

Winter habitat use by moose (Alces alces) is typically comprised of regenerating forest and softwood cover in the northeastern United States, and globally, high winter densities are of concern relative to forest damage. Habitat variables associated with winter locations of moose collected during aerial surveys in Maine in 2011 and 2012 were compared to available habitat at multiple landscape scales. Mixed forest was the most used land cover type at both the location and 5 ha scales (35.1% and 31.3%, respectively). Although regenerating forest habitat was used only in proportion to availability, the proximity to recent clearcuts, light partial cuts, and heavy partial cuts was an important predictor of moose location. The used proportion of coarse habitat variables (i.e., mature and regenerating forest) were similar to those available in each aerial survey block, indicating that heterogeneous and productive moose habitat is widely available across the commercial forest landscape of northern Maine. Moose locations derived from aerial surveys can provide insight about spatial distribution and habitat use across the landscape, identify local density in areas where forest regeneration is of concern, and monitor population responses to commercial forest management practices. [ABSTRACT FROM AUTHOR]

Age, field-dressed body weight, and antler spread data collected from 11,566 harvested moose (Alces alces) were analyzed to assess whether temporal change has occurred in the physical characteristics of bull moose from 1980-2009 in Maine. The annual proportion and antler spread of trophy bulls (spread ≥ 137 cm; n = 851) were also analyzed. There was no evidence of a measurable decline in the body weight or antler spread of adult bull moose (≥1.5 years old), similar to findings in Vermont and New Hampshire in a recent >20 year temporal analysis. There was a slight increase in physical characteristics of yearlings that contrasted with the trend in New Hampshire and Vermont where it is speculated that parasitism by winter ticks (Dermacentor albipictus) reduces growth rate and recruitment by yearlings. The proportion of trophy bulls in the harvest declined proportionally ~26% (9.3 to 6.9%) as harvest increased >2x from 1980-1987 to 2005-2009; however, the mean spread of trophy bulls declined by only 2% (P = 0.002). Additionally, there were no differences (P > 0.05) in the proportion of harvested bulls within each age class between 1980-1987 and 2005- 2009, and the relatively stable proportion of mature bulls (>5 years old) in the harvest across time periods (30-44%) does not suggest selective harvest of older, trophy bulls. In the face of the declining regional population, continued monitoring of harvested moose is warranted to best manage the largest and longest harvested population in the northeastern United States. [ABSTRACT FROM AUTHOR]

Moose (Alces alces) play an important role in the ecological and economic resources of northern New England, a landscape dominated by commercial forests. This study measured the impact of moose browsing on forest regeneration in Wildlife Management Unit El in northeastern Vermont where moose density was considered high in the 1990-2000s. We surveyed 37 clearcuts categorized into 4 age classes (3-5, 6-10, 11-15, and 16-20 years old). The stocking rate (stems/plot) of commercial species ranged from 74-76% in the 3-5, 6-10, and 11-15 year age classes, increasing to 86% in the 16-20 year age class. The proportion of plots containing a commercial tree without severe damage was above the accepted threshold stocking level of 40-60% in all age classes. The proportion of plots containing a commercial hardwood stem declined with increasing age class; the opposite occurred with softwood stems indicating a possible shift from hardwood- to softwood-dominated stands from selective browsing pressure. Height of 11-20 year old stems was less than in New Hampshire, indicating that growth was possibly suppressed in Vermont due to higher moose density. Overall, browsing was not considered a major problem based upon stocking rates. Further study is warranted to evaluate whether compensatory growth occurs in response to reduced browsing as forests age and/or moose population density declines. [ABSTRACT FROM AUTHOR]

In New Hampshire, winter ticks (Dermacentor albipictus) probably have more influence on the moose (Alces alces) population than other mortality factors, and predicting the frequency of tick epizootics is an important management consideration. Weather, moose density, and habitat use influence abundance and distribution of winter ticks. We evaluated the usefulness of 3 techniques to index winter tick abundance in 3 regions with variable moose density: 1) flagging for tick larvae, 2) line-transect counts of ticks on harvested moose, and 3) roadside surveys of tick-induced hairloss on moose. Although counts of tick larvae from fall flagging were not significantly different between years or regions, absolute tick abundance was measurably different (>50%) relative to moose density and years. Tick abundance on harvested moose reflected annual and regional differences; in general, abundance was correlated positively with moose density and annual trends within regions were similar. Tick abundance was highest for calves and lowest for cows. Hair-loss surveys indicated that hair loss was generally related to moose density, and similar annual differences were reflected in all regions. We suggest measuring tick abundance on harvested moose and conducting annual roadside hair-loss surveys to create indices and threshold values useful in predicting an epizootic of winter ticks. [ABSTRACT FROM AUTHOR]

Biological data collected from harvested moose (Alces alces) were analyzed to assess whether temporal change has occurred in the physical and reproductive condition of moose from 1988-2009 in New Hampshire. Measurements included age and field-dressed body weight of both sexes, number of corpora lutea (CL) and ovulation rate of females, and antler beam diameter (ABD) and antler spread of males. Similar data were obtained from Maine and Vermont for comparative analysis. The only significant changes (P <0.05) occurred in the yearling age class: mean body weight of both sexes, number of CL, and ABD all declined in New Hampshire. The current ovulation rate (~20%) and mean body weight (<200 kg) of yearling females in New Hampshire and Vermont were considered low. The declines measured in yearlings, yet relative stability in adults, are consistent with the presumption that winter ticks (Dermacentor albipictus) impact the productivity of moose populations through reduced calf survival and growth and fecundity of yearlings. Density-dependent factors related to habitat change are also discussed given the recent, rapid expansion of moose in the 3 states. Continued monitoring of physical parameters and productivity of harvested moose, particularly the yearling cohort, is warranted to better assess the relationships among winter ticks, habitat quality, and moose populations. [ABSTRACT FROM AUTHOR]

The impact of moose (Alces alces) browsing on the regeneration of commercial hard- wood and softwood tree species was evaluated in 3 regions with different moose population densities (0.26-0.83 moose/km2) in northern New Hampshire. Regeneration surveys were conducted in 4 age classes of clear-cuts (0-5, 6-10, 11-15, and 16-20 years) in June-August 2009. Stocking rate, tree height, and damage of dominant commercial stems were measured to assess regeneration and browse damage among age classes and regions. We assumed that a stocking rate of 40-60% (stems/plot) dominant commercial tree species without severe damage was an acceptable threshold of browse damage to achieve a fully stocked stand at 80 years. Mean stocking rate of all age classes was above the threshold in all regions (47-85%); the lowest stocking rates (47-52%) occurred in the 0-5 year age class but increased thereafter in all regions. The CT Lakes region (highest moose density) had more damage than both the North and White Mountain regions in the 11-15 year age class (F= 3.05; df; 6; P = 0.0002 and 0.0058, respectively). Tree heights were lower in the CT Lakes region (F = 2.30; df 6; P = 0.04). Most damage was restricted to a few isolated clear-cuts at higher elevation near moose wintering areas that were possibly shifting to conifer dominance. Regeneration of commercial tree species was not considered a regional problem at any moose density in northern New Hampshire. [ABSTRACT FROM AUTHOR]

Habitat use by parturient moose (Alces alces) may have important implications for calf survival and subsequently influence population dynamics. Because neonatal habitat may be limiting or specialized and little descriptive information exists in the northeastern United States, this study was conducted to measure the physical and vegetative characteristics associated with neonatal habitat of 30 maternal moose. There was no difference (P > 0.10 for each parameter) in 22 of 23 physical and vegetative parameters measured at neonatal (n = 30) and random sites (n = 30). However, neonatal sites were about 2X farther (P = 0.032) than random sites from cut/regeneration habitat where no neonatal site occurred. Most neonatal sites (> 63%) were located in pole or saw timber stands comprised of mixed or coniferous habitat (> 75%); conifers were the dominant canopy species at 67% of neonatal sites. Characteristics related to forage availability suggest that forage resources were probably not influential in location of neonatal habitat. Mature, mixed, and coniferous habitats may provide microhabitat that helps conceal neonates from potential predators such as black bears (Ursus americana), particularly in the absence of islands and open water that are believed to mitigate predation. [ABSTRACT FROM AUTHOR]

Moose (AIces alces) are susceptible to late winter mortality from infestation of winter ticks (Dermacentor albipictus) throughout much of North America. Calves, perhaps more so than other ages of moose, likely experience chronic, and eventually acute anemia from blood removal by adult female ticks that peaks during weeks 4 - 6 of the 8-week engorgement period. We modeled the potential metabolic impact on protein and energy balance of moose calves associated with blood loss during four levels, low to severe, of winter tick infestation. Our conservative estimates indicated that total blood loss in weeks 4 - 6, as a percent of total blood volume, ranged from 27 to 48% and 64 to 112% during moderate (30,000 ticks) and severe (70,000 ticks) infestations, respectively. The percent of the daily metabolizable energy requirement needed to replace daily blood loss during weeks 4 - 6 was 4.9- 8.2% and 11.4 - 19.2% during moderate and severe infestations, respectively. The protein deficit associated with blood loss and regeneration was the most critical metabolic impact. Daily protein loss during weeks 4 - 6 was 29 - 49% and 68 - 114% of the daily protein requirement in moderate and severe infestations, respectively. Daily protein losses of ∼ 30 to > 100% occurred for 2 continuous weeks. Energy costs associated with compensating for blood loss would likely elevate the daily energy deficit normal at end of winter, accelerate nutritional decline and weight loss, and cause increased physiological stress related to concurrent anemia. Severely infested calves are obviously susceptible to late winter mortality, and the impact of moderate infestations would be exacerbated by secondary parasitic infestations, severe winters, and poor body condition. [ABSTRACT FROM AUTHOR]

We examined behavioral response of moose to wildlife viewers and traffic stimuli at a moose viewing blind located on a roadside salt lick in northern New Hampshire during summer, 1997-1999. Feeding, fleeing, alertness, looking, grooming, and moving were measured relative to a standard viewer and a variety of stimuli associated with viewers and traffic. in general, moose were reasonably tolerant of most stimuli as moose never fled the lick> 15% of the time. Educational material likely influenced viewer behavior. Stimuli that caused a reduction in feeding and increased fleeing were loud viewers, cars stopped, and trucks passing, as well as combinations of stimuli including these factors. Viewing satisfaction and impacts can be addressed by considering these findings at moose viewing sites. [ABSTRACT FROM AUTHOR]

In northern New Hampshire, we examined the use patterns of moose visiting a roadside salt lick before (1996) and after (1997–1999) a blind was built specifically to view moose at the lick. Moose visitation patterns were monitored with trail monitors equipped with cameras placed on trails leading into the study and control salt licks. There was no difference in frequency of use and time of use at the study and control sites in any year. Nocturnal use was higher than diurnal use; use was greatest at 2200–2400 and 0400–0600 hours at both sites. Reduced use of the trail closest to the blind indicated that placement of the blind probably altered access patterns of certain moose. A trend in 1998–1999 toward more visits during early morning than peak afternoon viewing time indicated that assessment of viewing opportunity warrants further study. [ABSTRACT FROM AUTHOR]

A foreword to "Alces: A Journal Devoted to the Biology and Management of Moose" is presented.

Acquiring accurate and precise population parameters is fundamental to the ecological understanding and management and conservation of moose (Alces alces). Moose density is challenging to measure and often estimated using winter aerial surveys; however, numerous alternative approaches exist including harvest analysis, public observation, unpiloted aerial system (UAS) surveys, and camera trapping. Given recent developments in a number of field and analytical techniques, there is value in reviewing and synthesizing the strengths and limitations of monitoring methods to best evaluate their respective tradeoffs in management scenarios. We reviewed 89 studies that included 131 estimates or indices of moose density. As expected, aerial surveys were the most common method of obtaining a moose density estimate (58%) followed by use of public data (e.g., harvest records = 27%); more recent studies employed novel methods including UAS. Most estimates (64%) failed to account for imperfect detection of moose (i.e., "sightability") and this tendency has not improved over time. Density estimates ranged from <0.1 to 10.6 moose/km² (average = 0.7) and population precision, as measured by the 90% confidence interval, ranged from 6.5 to 120.0% of the density estimate (average = 37.4%). Correlations among estimates obtained for the same populations varied widely, with R² values ranging from 0.02 to 0.99 (average = 0.58). Our review indicates that: 1) methods to estimate moose density have been dominated by aerial surveys but are diversifying, 2) precision of density estimates has been highly variable and on average lower than broadly accepted target benchmarks, and 3) many methods did not account for sightability and presumably underestimated moose density. We reflect on these trends and discuss how emerging methods, including camera trapping, UAS surveys, and integrated population modeling (IPM) can complement and improve traditional approaches. We suggest that no single "best" method exists, but rather the best method is one that accounts for sightability bias and yields target precision at reasonable cost, which vary by jurisdiction and goal. [ABSTRACT FROM AUTHOR]

Soil fungi were cultured from 24 wallows and proximal control sites in Maine and New Hampshire, USA during the autumn moose (Alces alces) breeding season of 2016 to investigate the presence of soil fungi pathogenic to winter tick larvae (Dermacentor albipictus). Twenty genera of fungi were isolated, and all are considered common in a forested ecosystem. The predominant genera isolated in wallows were pathogenic to winter tick larvae and included Aspergillus spp. (in particular A. flavus), Beauveria bassiana, Mortierella spp., Mucor spp., Paecilomyces spp., Penicillium spp., and Trichoderma spp. Wallow soils had specific characteristics and differed from proximal control sites by having: 1) lower fungal diversity, 2) a higher frequency of primary colonizers including Mortierella spp., Mucor spp., Penicillium spp., and Trichoderma spp., and 3) a more variable total amount of fungi indicative of changing (disturbed) soil conditions. We conclude that wallows are sites of soil disturbance that concentrate fungi known to be pathogenic to larval winter ticks. Fungi acquired by breeding moose using wallows might subsequently act as an on-host mechanism of tick control. [ABSTRACT FROM AUTHOR]

The New Hampshire Fish and Game Department and the University of New Hampshire initiated research in northern New Hampshire to better understand population dynamics and seasonal habitat use of a moose population that has apparently stabilized, despite optimal habitat and modest harvest levels. In total, 94 moose were captured by helicopter (81 net-gunned and 13 tranquilized) in December 2001–2003 and 2 were darted at salt-licks in July of 2002. Capture mortality attributed to myopathy and injury was 4%. In comparison to measured reproduction during capture (63 and 100%), our ability to measure pregnancy by direct observations (69 and 100%) was validated in 2002-2003. Production was 0.82 and 0.85 calves per adult cow; rate of twinning was 20 and 10%. Calf mortality 2 months post-partum was similar (26 and 27%) each year. Annual mortality of adult/yearling moose was 27 and 12%. Hunting and vehicle collision mortality was 4 (all adult cows) and 6% (all calves but 1) each year. High annual winter calf mortality (38–43%) in late March and early April was associated with the combined effects of malnutrition and winter tick/lung nematodes. Winter home range size was not restricted, and composition of available habitat was similar across seasons although overlap was minimal between seasons. Consideration of habitat and population dynamics data suggests that both density dependent and independent factors could be influencing the study population. [ABSTRACT FROM AUTHOR]