Statistical population reconstruction of moose (Alces alces) in northeastern Minnesota using integrated population models.

Given recent and abrupt declines in the abundance of moose (Alces alces) throughout parts of Minnesota and elsewhere in North America, accurately estimating statewide population trends and demographic parameters is a high priority for their continued management and conservation. Statistical population reconstruction using integrated population models provides a flexible framework for combining information from multiple studies to produce robust estimates of population abundance, recruitment, and survival. We used this framework to combine aerial survey data and survival data from telemetry studies to recreate trends and demographics of moose in northeastern Minnesota, USA, from 2005 to 2020. Statistical population reconstruction confirmed the sharp decline in abundance from an estimated 7,841 (90% CI = 6,702-8,933) in 2009 to 3,386 (90% CI = 2,681-4,243) animals in 2013, but also indicated that abundance has remained relatively stable since then, except for a slight decline to 3,163 (90% CI = 2,403-3,718) in 2020. Subsequent stochastic projection of the population from 2021 to 2030 suggests that this modest decline will continue for the next 10 years. Both annual adult survival and per-capita recruitment (number of calves that survived to 1 year per adult female alive during the previous year) decreased substantially in years 2005 and 2019, from 0.902 (SE = 0.043) to 0.689 (SE = 0.061) and from 0.386 (SE = 0.030) to 0.303 (SE = 0.051), respectively. Sensitivity analysis revealed that moose abundance was more sensitive to fluctuations in adult survival than recruitment; thus, we conclude that the steep decline in 2013 was driven primarily by decreasing adult survival. Our analysis demonstrates the potential utility of using statistical population reconstruction to monitor moose population trends and to identify population declines more quickly. Future studies should focus on providing better estimates of per-capita recruitment, using pregnancy rates and calf survival, which can then be incorporated into reconstruction models to help improve estimates of population change through time.

Comparing survey and multiple recruitment-mortality models to assess growth rates and population projections.

Estimation of population trends and demographic parameters is important to our understanding of fundamental ecology and species management, yet these data are often difficult to obtain without the use of data from population surveys or marking animals. The northeastern Minnesota moose (Alces alces Linnaeus, 1758) population declined 58% during 2006-2017, yet aerial surveys indicated stability during 2012-2017. In response to the decline, the Minnesota Department of Natural Resources (MNDNR) initiated studies of adult and calf survival to better understand cause-specific mortality, calf recruitment, and factors influencing the population trajectory. We estimated population growth rate (λ) using adult survival and calf recruitment data from demographic studies and the recruitment-mortality (R-M) Equation and compared these estimates to those calculated using data from aerial surveys. We then projected population dynamics 50 years using each resulting λ and used a stochastic model to project population dynamics 30 years using data from the MNDNR's studies. Calculations of λ derived from 2012 to 2017 survey data, and the R-M Equation indicated growth (1.02 ± 0.16 [SE] and 1.01 ± 0.04, respectively). However, the stochastic model indicated a decline in the population over 30 years (λ = 0.91 ± 0.004; 2014-2044). The R-M Equation has utility for estimating λ, and the supporting information from demographic collaring studies also helps to better address management questions. Furthermore, estimates of λ calculated using collaring data were more certain and reflective of current conditions. Long-term monitoring using collars would better inform population performance predictions and demographic responses to environmental variability.

Potential Vertical Transmission of Winter Ticks (Dermacentor albipictus) from Moose (Alces americanus) Dams to Neonates.

North American moose (Alces americanus) frequently become infested with winter ticks (Dermacentor albipictus). During capture of neonatal moose in northeastern Minnesota, US, in May-June 2013 and 2014, we recovered adult ticks from neonates, presumably vertically transferred from dams, heretofore, not documented. Infestations on neonates may have population-level implications.

A long-term assessment of the variability in winter use of dense conifer cover by female white-tailed deer.

BACKGROUND: Long-term studies allow capture of a wide breadth of environmental variability and a broader context within which to maximize our understanding of relationships to specific aspects of wildlife behavior. The goal of our study was to improve our understanding of the biological value of dense conifer cover to deer on winter range relative to snow depth and ambient temperature. METHODOLOGY/PRINCIPAL FINDINGS: We examined variation among deer in their use of dense conifer cover during a 12-year study period as potentially influenced by winter severity and cover availability. Female deer were fitted with a mixture of very high frequency (VHF, n = 267) and Global Positioning System (GPS, n = 24) collars for monitoring use of specific cover types at the population and individual levels, respectively. We developed habitat composites for four study sites. We fit multinomial response models to VHF (daytime) data to describe population-level use patterns as a function of snow depth, ambient temperature, and cover availability. To develop alternative hypotheses regarding expected spatio-temporal patterns in the use of dense conifer cover, we considered two sets of competing sub-hypotheses. The first set addressed whether or not dense conifer cover was limiting on the four study sites. The second set considered four alternative sub-hypotheses regarding the potential influence of snow depth and ambient temperature on space use patterns. Deer use of dense conifer cover increased the most with increasing snow depth and most abruptly on the two sites where it was most available, suggestive of an energy conservation strategy. Deer use of dense cover decreased the most with decreasing temperatures on the sites where it was most available. At all four sites deer made greater daytime use (55 to >80% probability of use) of open vegetation types at the lowest daily minimum temperatures indicating the importance of thermal benefits afforded from increased exposure to solar radiation. Date-time plots of GPS data (24 hr) allowed us to explore individual diurnal and seasonal patterns of habitat use relative to changes in snow depth. There was significant among-animal variability in their propensity to be found in three density classes of conifer cover and other open types, but little difference between diurnal and nocturnal patterns of habitat use. CONCLUSIONS/SIGNIFICANCE: Consistent with our findings reported elsewhere that snow depth has a greater impact on deer survival than ambient temperature, herein our population-level results highlight the importance of dense conifer cover as snow shelter rather than thermal cover. Collectively, our findings suggest that maximizing availability of dense conifer cover in an energetically beneficial arrangement with quality feeding sites should be a prominent component of habitat management for deer.

Winter body condition of moose (Alces alces) in a declining population in northeastern Minnesota.

Assessments of the condition of moose (Alces alces) may be particularly informative to understanding the dynamics of populations and other influential factors. During February-March 2003 to 2005, we assessed the nutritional condition of 79 moose (39 females, 40 males) in northeastern Minnesota by body condition scoring (BCS(F), scale of 0-10); 67 of these by were assessed by ultrasonographic measurements of rump fat (Maxfat), which was used to estimate ingesta-free body fat (IFBF) in all but two of the females. Scores of the BCS(F) were related (r(2)=0.34, P<0.0001) to Maxfat. Body condition scores were not affected by sex × capture-year, capture-year, or age-at-capture, but the mean body condition score of males (6.5 ± 0.2 [SE], n=40) was less (P ≤ 0.009) than that of females (7.4 ± 0.2, n=39). Overall, Maxfat ranged from 0.0 to 4.6 and 0.3 to 2.8 cm in females and males, respectively, and was unaffected by age-at-capture. There was a sex×capture-year effect (P=0.021) on Maxfat; mean values were stable for males during the winters of 2003 to 2005 but in females were lowest during 2003, consistent with the lowest pregnancy rates and lowest winter and spring survival compared to 2004 and 2005. Based on estimates of percent IFBF, late winter-early spring survival in 2003 of at least 11% of the collared animals assessed by Maxfat, 21% of the adult females, specifically, may have been seriously challenged directly by poor condition. Data from this study provide reference values and assessments of body condition of moose that will be an essential component of the additional, comprehensive research needed to better understand the influence of extrinsic and intrinsic factors on the performance of this viable, but declining, population. For future research, we will concentrate on developing a more-reliable BCS which would allow IFBF estimation once rump fat is depleted.

What time is it? Choice of time origin and scale in extended proportional hazards models.

The analysis of telemetry data offers many unique challenges due to both the observation process and the complexity of the underlying system (e.g., risk of mortality may be influenced by both age and a wide range of environmental variables). Although semi-parametric proportional hazards (SPPH) models have been proposed for analyzing ecological data, recent applications have failed to address the importance of choosing an appropriate time origin and scale for analysis. We compared models fit to a long-term deer (Odocoileus spp.) survival data set using three alternative survival timescales: age, time since start of study, and time since 6 June (with a seasonally recurrent timescale). Temporal variability in risk resulted from multiple sources (e.g., changes in hunting pressure, winter severity), and the risk of mortality varied nonlinearly with age (highest risk for young and older individuals). Age-varying hazards were represented well using regression splines, but temporal variability was more difficult to model using parametric assumptions. Annual survival estimates using the three timescales differed considerably. The model using a study-based timescale most closely tracked temporal patterns in risk. Given the difficulties in modeling temporal variability using parametric assumptions, we recommend this approach over an age-based or recurrent timescale when using SPPH models to evaluate the impact of large (naturally occurring or experimental) disturbances or to estimate annual age-specific survival rates. Lastly, we discuss the strengths and limitations of SPPH models relative to fully parametric approaches.

Birth, morphologic, and blood characteristics of free-ranging white-tailed deer neonates.

Winter severity is a primary factor influencing deer survival and reproduction in northern climates. Prolonged, harsh winters can adversely affect body condition of does, resulting in depressed morphologic development of neonates. In this study, we captured 59 white-tailed deer (Odocoileus virginianus) neonates (28 in 2001 and 31 in 2002), following two distinctly different winters, one severe and the other historically mild. Vaginal implant transmitters allowed exact age to be determined for 73% of the neonates; new hoof growth was used to estimate age (days) of the other 27%. Birthdate and morphologic measurements of neonates (i.e., birth mass, new hoof growth, hoof length) were compared by sex and capture year. For known-age neonates (n=43), there was a year-by-sex interaction effect (P=0.01) on birthdate, being later for females during spring 2001 compared with 2002, which was consistent with a significant (P=0.03) year-by-sex interaction for total hoof length (22.3 mm [SE=0.9] and 20.3 [SE=0.8] for females and males in 2001; 19.9 [SE=1.0] and 22.1 [SE=1.0] for females and males in 2002). Interestingly, there was no effect of year on birth mass or birthdate of known-age neonates. A year-by-sex interaction (P=0.04) was determined for birthdates of estimated age (5 yr old were born later (P<0.01) than fawns born to dams

Mycobacterium avium subsp. paratuberculosis from free-ranging deer and rabbits surrounding Minnesota dairy herds.

The objectives of this study were to estimate the prevalence of Mycobacterium avium subsp. paratuberculosis (MAP) among deer and rabbits surrounding infected and noninfected Minnesota dairy farms using fecal culture, and to describe the frequency that farm management practices were used that could potentially lead to transmission of infection between these species. Fecal samples from cows and the cow environment were collected from 108 Minnesota dairy herds, and fecal pellets from free-ranging white-tailed deer and eastern cottontail rabbits were collected from locations surrounding 114 farms; all samples were tested using bacterial culture. In addition, a questionnaire was administered to 114 herd owners. Sixty-two percent of the dairy herds had at least 1 positive fecal pool or environmental sample. A total of 218 rabbit samples were collected from 90% of the herds, and 309 deer samples were collected from 47% of the herds. On 2 (4%) of the farms sampled, 1 deer fecal sample was MAP positive. Both farms had samples from the cow fecal pool and cow environment that were positive by culture. On 2 (2%) other farms, 1 rabbit fecal sample was positive by culture to MAP, with one of these farms having positive cow fecal pools and cow environmental samples. Pasture was used on 79% of the study farms as a grazing area for cattle, mainly for dry cows (75%) and bred or prebred heifers (87%). Of the 114 farms, 88 (77%) provided access to drylot for their cattle, mainly for milking cows (77/88; 88%) and bred heifers (87%). Of all study farms, 90 (79%) used some solid manure broadcasting on their crop fields. Of all 114 farms, the estimated probability of daily physical contact between cattle manure and deer or rabbits was 20% and 25%, respectively. Possible contact between cattle manure and deer or rabbits was estimated to occur primarily from March through December. The frequency of pasture or drylot use and manure spreading on crop fields may be important risk factors for transmission of MAP among dairy cattle, deer, and rabbits. Although the MAP prevalence among rabbits and deer is low, their role as MAP reservoirs should be considered.