Redefining physiological responses of moose (Alces alces) to warm environmental conditions

We tested the concept that moose (Alces alces) begin to show signs of thermal stress at ambient air temperatures as low as 14 °C. We determined the response of Alaskan female moose to environmental conditions from May through September by measuring core body temperature, heart rate, respiration rate, rate of heat loss from exhaled air, skin temperature, and fecal and salivary glucocorticoids. Seasonal and daily patterns in moose body temperature did not passively follow the same patterns as environmental variables. We used large changes in body temperature (≥1.25 °C in 24hr) to indicate days of physiological tolerance to thermal stressors. Thermal tolerance correlated with high ambient air temperatures from the prior day and with seasonal peaks in solar radiation (June), ambient air temperature and vapor pressure (July). At midday (12:00hr), moose exhibited daily minima of body temperature, heart rate and skin temperature (difference between the ear artery and pinna) that coincided with daily maxima in respiration rate and the rate of heat lost through respiration. Salivary cortisol measured in moose during the morning was positively related to the change in air temperature during the hour prior to sample collection, while fecal glucocorticoid levels increased with increasing solar radiation during the prior day. Our results suggest that free-ranging moose do not have a static threshold of ambient air temperature at which they become heat stressed during the warm season. In early summer, body temperature of moose is influenced by the interaction of ambient temperature during the prior day with the seasonal peak of solar radiation. In late summer, moose body temperature is influenced by the interaction between ambient temperature and vapor pressure. Thermal tolerance of moose depends on the intensity and duration of daily weather parameters and the ability of the animal to use physiological and behavioral responses to dissipate heat loads.     

Three approaches to estimate wolf Canis lupus predation rates on moose Alces alces populations

We employed three different methods to estimate predation rates on moose in a newly colonized wolf territory in Norway. In the first two methods, we estimated predation rates based on the difference in calf/cow ratios outside and inside the wolf pack territory from (1) hunter observations and (2) aerial surveys. In the last method, (3) we estimated loss of calves of radio-collared cows inside and outside the wolf pack territory. The difference in mortality rates estimated between the area subject to predation and the area outside the wolf pack territory essentially constitutes the additive component of predation. We also tested the sensitivity of violating the assumptions of methods 1 and 2 related to equal fecundity and mortality because of other factors than predation inside and outside the wolf pack territory. Predation rates varied considerably between years and methods used, with hunter observations (method 1) giving the lowest and aerial surveys (method 2) giving the highest estimates. Method 3 (radio telemetry) was the most direct assessment of predation and probably the best approach to estimate predation rates in moose. However, all three methods show the same yearly changes and may therefore be appropriate to question trends trough time or between areas.     

Geographical variation in body weight and sexual size-dimorphism of Norwegian moose (Alces alces)

Geographical variation in male carcass weight, and sexual dimorphism in size was studied in 19 populations of Norwegian moose ( Alces alces (L.)).
Significant age-specific variation in male carcass weight was found for all the populations studied up to the age of 4 1/2 years, but in some populations maximum weight was not reached until at least 5 1/2 years. Increase in the mean weight of females after the age of 2 1/2 years was not significant. Only a weak relationship existed between mean yearling and adult bull weights in a population. However, within both the southern (< 62°N) and northern (> 62°N) parts of the country, yearling carcass weight was a good predictor of adult bull weight in a region.
Adult bull weight in a region was best predicted from the increment in mean carcass weight observed between 1 1/2 and 3 1/2 years of age. Within a region, variation in age-specific carcass weight between cohorts of bulls from different years was also well predicted from annual variation in growth increment.
Those patterns reveal a sexual difference in strategy of body growth. The adult weight of females is probably strongly determined by the weight gained by the time of onset of reproduction. The males have available a longer period for growth in body weight. They are therefore able to compensate for low weights early in life by increased gain of weight in later years, that provide good conditions for growth.
Geographical variation in the degree of sexual dimorphism in size correlated only poorly with adult male size. We suggest that the sexual size-dimorphism is a result of reproductive constraints of the female, i.e. in populations living in poor conditions and having small body size, the onset of reproduction prevents further gain in body weight.     

The declining occurrence of moose (Alces alces) at the southernmost edge of its range raise conservation concerns

The border region between Austria, the Czech Republic, and Germany harbors the most south‐western occurrence of moose in continental Europe. The population originated in Poland, where moose survived, immigrated from former Soviet Union or were reintroduced after the Second World War expanded west‐ and southwards. In recent years, the distribution of the nonetheless small Central European population seems to have declined, necessitating an evaluation of its current status. In this study, existing datasets of moose observations from 1958 to 2019 collected in the three countries were combined to create a database totaling 771 records (observations and deaths). The database was then used to analyze the following: (a) changes in moose distribution, (b) the most important mortality factors, and (c) the availability of suitable habitat as determined using a maximum entropy approach. The results showed a progressive increase in the number of moose observations after 1958, with peaks in the 1990s and around 2010, followed by a relatively steep drop after 2013. Mortality within the moose population was mostly due to human interactions, including 13 deadly wildlife‐vehicle collisions, particularly on minor roads, and four animals that were either legally culled or poached. Our habitat model suggested that higher altitudes (ca. 700–1,000 m a.s.l.), especially those offering wetlands, broad‐leaved forests and natural grasslands, are the preferred habitats of moose whereas steep slopes and areas of human activity are avoided. The habitat model also revealed the availability of large core areas of suitable habitat beyond the current distribution, suggesting that habitat was not the limiting factor explaining the moose distribution in the study area. Our findings call for immediate transboundary conservation measures to sustain the moose population, such as those aimed at preventing wildlife‐vehicle collisions and illegal killings. Infrastructure planning and development activities must take into account the habitat requirements of moose.     

Reproductive failure in moose (Alces alces) due to embryonic mortality and unfertilized oocytes

Knowledge on reproductive success is vital for successful management of large ungulates and is often measured by means of observing surviving offspring. In harvested ungulates, postmortem investigations of reproductive organs are used to estimate reproductive potential by obtaining ovulation rates and fetus numbers. However, there are differences in numbers of offspring observed, fetal/embryo counts, and ovulation rates. We hypothesize that the discrepancy between estimated reproductive potential and reproductive outcome in large ungulates is not only due to ova loss but also due to embryonic mortality. We investigated reproductive status in early pregnancy by sampling hunter-harvested moose (Alces alces) in southern Sweden from 2007 to 2011. In all, 213 reproductive organs were examined postmortem, and in confirmed pregnant moose (n?=?53), 25 % (19 of 76) embryos were nonviable and 6 % of ova was unfertilized. The discrepancy between the ovulation rate of all pregnant moose (1.49) and the number of expected offspring per pregnant female, when embryonic mortality and unfertilized oocytes were accounted for (1.08), was 27.5 %. An association between inflammation of the inner mucous membrane (endometritis) of the moose's uterus and embryonic mortality was observed. This is the first comprehensive report of embryonic mortality and endometritis in moose. The observed discrepancy between ovulation rates and early embryonic development/survival shows that ovulation rates are indicative but not accurate estimates of moose reproductive rate. The use of ovulation rates as a sole estimator of future offspring rates may lead to an overharvest of a managed moose population.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Dermacentor albipictus on moose (Alces alces) in Ontario.

Fifty-five moose (Alces alces) collected from 1963 to 1965 in the Chapleau Crown Game Preserve of northern Ontario were examined for ectoparasites. Dermacentor albipictus was the only parasite recovered. D. albipictus was absent from 15 moose examined during June, July and August; first observed on 1 or 2 moose examined in September; and present on all 38 moose collected from October to May. Ticks were not evenly distributed on the body. Infestations varied from 32 to 13,490 ticks/moose for 14 individuals. Levels of infestation observed did not have an obvious adverse effect on the health of the moose.     

Elaeophora schneideri in moose (Alces alces) from Colorado.

Two adult moose (Alces alces) from Colorado (USA) were naturally infected with Elaeophora schneideri. Both animals had patent infections indicating that moose may serve as definitive hosts. Gross and histological lesions were characterized by fibromuscular intimal proliferation within the carotid arteries and rete mirabile cerebri. This is the first report of Elaeophora schneideri in moose from Colorado.     

Low-dose meningeal worm (Parelaphostrongylus tenuis) infections in moose (Alces alces)

Parelaphostrongylosis has a rapid onset and is lethal in neonatal moose (Alces alces) when large numbers of third-stage Parelaphostrongylus tenuis larvae (L3) are given experimentally. Little is known, however, about the severity and prognosis of infections acquired naturally by accidentally ingesting terrestrial gastropods which are rarely infected and have few larvae. To investigate the relationship between infecting dose, age of moose, and severity of disease, five calves were given low doses of three to 10 L3 when five (n = 2) or 9.5 mo old (n = 3). Each of two animals initially given low doses were later challenged with a dose of 15 L3. As positive controls, two calves were given doses of 15 and 30 L3, considered to be high. All five calves given low doses showed abnormal locomotory signs at 20-28 days postinoculation (DPI) that progressively became more pronounced with hind quarter weakness and front lameness. However, after 77-130 DPI, signs diminished markedly in two of these animals and disappeared in another two. Challenge infections of 15 L3 given 199 days after initial infections had no noticeable effects although an immature worm, probably resulting from the challenge, was found in the spinal cord of one animal killed 51 days later. Two positive control animals given the high doses of 15 and 30 L3 showed moderate to severe, non-resolving, locomotory signs and had to be euthanized. Results demonstrate that single, low doses of three to 10 P. tenuis L3 cause moderate disease in moose calves but over time, some worms die and animals can recover. A degree of protection may develop against future infection.     

Geographical variation of antler morphology of moose (Alces alces) in Russia

We attempt to identify geographical subdivisions within the range of moose according to antler size for a more correct trophy evaluation in the СIC system (The International Council for Game and Wildlife Conservation). Traditionally, this division coincides with subspecific division of trophy animal species. That is why we partially touch upon the problem of the geography of moose subspecies in Eurasia. Catalogs of hunters’ trophies that were available to us served as the research material (1,047 trophies in total). We were choosing the same measurements for moose antlers evaluated by different methods. They are overall spread, circumference of beam, width of palm, and number of tines. We applied multivariate statistic methods to the findings, enlarging the geographical division groups step by step. As a result, we suggest four geographical groups: European, Siberian, the north of the Far East, and the south of the Far East. In principle, this division corresponds to the subspecies: European (Alces alces alces L., 1758), east Siberian (A. a. pfizenmayeri Zukowsky, 1910), Chukotka or Kolyma (A. a. buturlini Chernyavski et. Zhelesnov, 1982), and Ussuri (А. а. cameloides Milne-Edwards, 1867). We consider it incorrect to draw the line between the European and Siberian groups along the river Yenisei. It is more correct to move it to the Urals.     

Tick (Dermacentor albipictus)-induced winter hair-loss in captive moose (Alces alces)

Five captive moose calves each infested with 42,000 larval Dermacentor albipictus, six calves each infested with 19,000-21,000, and five control moose were observed for changes in hair-loss, body condition and number, stages, and distribution of the tick. Winter hair-loss was observed only in moose infested with ticks and was correlated positively with the total number of adult ticks. Hair-loss associated with ticks was minimal from October to January, but rapidly increased from February to April when up to 44% of hair had been removed. The pattern of hair-loss was similar in all moose with the neck, shoulders, withers, and perianal areas losing the most hair. Moose with extensive premature hair-loss had less pericardial and abdominal visceral fat than moose with little or no hair-loss.     

Genetic diversity of moose (Alces alces L.) in Eurasia

Polymorphism of nucleotide sequence of D-loop fragment of the mitochondrial DNA was studied in 20 moose from several local populations on the territory of Eurasia. Three main haplotype variants of D-loop were detected by molecular phylogenetic method, which formed three clusters named European, Asian and American. Intraspecies variation in the length of HVSI of D-loop of the mitochondrial DNA of moose was revealed. In the Far Eastern and Yakutian moose, haplotypes with a 75-bp deletion were found, which were most similar with haplotypes (also with the deletion), earlier observed in North American moose [1]. The highest diversity of the haplotypes of mitochondrial DNA is characteristic of Yakutia and the Far East (where three haplotype variants were found), which demonstrates the probable role of the region as the center of the species or as the region of ancient population mixture. The geographic region might be considered as a probable source of ancient moose migrations from Asia to America, basing on the data of distribution of mitochondrial haplotypes of D-loop and alleles of MhcAlal-DRB1. Divergence of nucleotide sequences of haplotypes with the 75-bp deletion (forming the American cluster on the phylogenetic tree) was the lowest (0.4%), which evidences respectively recent origin of the group of haplotypes. In Europe, only haplotypes of mitochondrial DNA referred to European variant were observed. Basing on analysis of variation of nucleotide sequences of D-loop, exon 4 of kappa-Casein and exon 2 of MhcALal-DRB1, we demonstrated that Eurasian moose studied belong to the unique species, which has probably passed through a bottle neck. The time of the origin of modern diversity of D-loop haplotypes of the species was estimated as 0.075-0.15 Myr ago.     

Aspects of the life cycle and pathogenesis of Elaphostrongylus alces in moose (Alces alces).

Aspects of the migratory life cycle and pathogenesis of Elaphostrongylus alces were studied in 7 randomly selected moose calves and 7 yearlings killed during August to November. One calf and 1 yearling were uninfected. The 6 infected calves had recent infections, whereas the 6 yearlings showed older infections from the summer of the previous year. The 2 calves killed in September had a total of 26 adult E. alces in the epidural space of the caudal vertebral canal and none in the skeletal muscles, whereas the remaining calves killed 1 to 2 mo later had 25 nematodes in the caudal and cranial vertebral canal and 7 in the skeletal muscles. The yearlings had a total of 101 adult E. alces in the skeletal muscles and 2 in the vertebral canal. There were no findings indicating involvement of the central nervous system in the life cycle of E. alces. Our findings suggest that E. alces migrates directly from the gut to the epidural space of the caudal vertebral canal where development to the adult stage takes place. During development, the nematode produces inflammation of the epidural tissue and spinal nerves. Development in the caudal vertebral canal is followed by some anterior dispersion of nematodes along the canal, and migration into skeletal muscles. Here the nematodes seem to live in reproductive pairs and groups. The predilection site for E. alces in moose is the muscles of the thigh.     

Telemetered heart rate as an index of energy expenditure in moose (Alces alces)

Metabolic rate, heart rate and respiratory rate of three adult moose were monitored simultaneously on selected days between January 1982 and October 1983. Heart rate was successfully monitored using two types of implanted radio transmitters. Metabolic rate was significantly (P less than 0.001) correlated with heart rate for individual animals (r = 0.94-0.98). The relationship was best described by an exponential equation. A mean weighted equation predicted metabolic rate within 9% of individual equations. In adult moose cows, heart rate was a more reliable predictor of energy expenditure than respiratory rate.     

Hematology and serum chemistry reference ranges of free-ranging moose (Alces alces) in Norway

Baseline reference ranges of serum chemistry and hematology data can be important indicators for the status of both individuals or populations of wild animals that are affected by emerging pathogens, toxicants, or other causes of disease. Frequently, reference ranges for these values are not available for wildlife species or subspecies. We present hematologic and serum chemistry reference ranges for moose (Alces alces) adults, yearlings, and calves in Norway sampled from 1992-2000. Additionally, we demonstrated that both induction time and chase time were correlated with initial rectal temperature, although they were not significantly correlated with cortisol, aspartate aminotransferase, glucose, or creatine kinase. Overall, the reference ranges given here are similar to those given for American moose, with a few differences that can be attributed to environment, testing methodology, or subspecies or species status. This is the first report, to our knowledge, of reference ranges for moose in Norway.     

Echinococcus granulosus (Cestoda: Taeniidae) infections in moose (Alces alces) from southwestern Quebec

Investigation of the distribution of larval Echinococcus granulosus in a moose population from southwestern Quebec revealed a distinct and stable pattern of infection with a prevalence of 44% (n = 580). Positive correlations between moose age and the intensity, mean cyst weight and biomass of the hydatid cysts suggested a process of continued parasite acquisition and cyst growth. The distribution of cyst sizes within individual moose provided circumstantial evidence of interaction between cysts, perhaps mediated through the host's immunological response.