Genomic Underpinnings of Population Persistence in Isle Royale Moose.

Island ecosystems provide natural laboratories to assess the impacts of isolation on population persistence. However, most studies of persistence have focused on a single species, without comparisons to other organisms they interact with in the ecosystem. The case study of moose and gray wolves on Isle Royale allows for a direct contrast of genetic variation in isolated populations that have experienced dramatically differing population trajectories over the past decade. Whereas the Isle Royale wolf population recently declined nearly to extinction due to severe inbreeding depression, the moose population has thrived and continues to persist, despite having low genetic diversity and being isolated for ∼120 years. Here, we examine the patterns of genomic variation underlying the continued persistence of the Isle Royale moose population. We document high levels of inbreeding in the population, roughly as high as the wolf population at the time of its decline. However, inbreeding in the moose population manifests in the form of intermediate-length runs of homozygosity suggestive of historical inbreeding and purging, contrasting with the long runs of homozygosity observed in the smaller wolf population. Using simulations, we confirm that substantial purging has likely occurred in the moose population. However, we also document notable increases in genetic load, which could eventually threaten population viability over the long term. Overall, our results demonstrate a complex relationship between inbreeding, genetic diversity, and population viability that highlights the use of genomic datasets and computational simulation tools for understanding the factors enabling persistence in isolated populations.

Sporadic cases of chronic wasting disease in old moose - an epidemiological study.

Transmissible spongiform encephalopathies or prion diseases comprise diseases with different levels of contagiousness under natural conditions. The hypothesis has been raised that the chronic wasting disease (CWD) cases detected in Nordic moose (Alces alces) may be less contagious, or not contagious between live animals under field conditions. This study aims to investigate the epidemiology of CWD cases detected in moose in Norway, Sweden and Finland using surveillance data from 2016 to 2022.In total, 18 CWD cases were detected in Nordic moose. All moose were positive for prion (PrPres) detection in the brain, but negative in lymph nodes, all were old (mean 16 years; range 12-20) and all except one, were female. Age appeared to be a strong risk factor, and the sex difference may be explained by few males reaching high age due to hunting targeting calves, yearlings and males.The cases were geographically scattered, distributed over 15 municipalities. However, three cases were detected in each of two areas, Selbu in Norway and Arjeplog-Arvidsjaur in Sweden. A Monte Carlo simulation approach was applied to investigate the likelihood of such clustering occurring by chance, given the assumption of a non-contagious disease. The empirical P-value for obtaining three cases in one Norwegian municipality was less than 0.05, indicating clustering. However, the moose in Selbu were affected by different CWD strains, and over a 6 year period with intensive surveillance, the apparent prevalence decreased, which would not be expected for an ongoing outbreak of CWD. Likewise, the three cases in Arjeplog-Arvidsjaur could also indicate clustering, but management practices promotes a larger proportion of old females and the detection of the first CWD case contributed to increased awareness and sampling.The results of our study show that the CWD cases detected so far in Nordic moose have a different epidemiology compared to CWD cases reported from North America and in Norwegian reindeer (Rangifer tarandus tarandus). The results support the hypothesis that these cases are less contagious or not contagious between live animals under field conditions. To enable differentiation from other types of CWD, we support the use of sporadic CWD (sCWD) among the names already in use.

Links between three chronic and age-related diseases, osteoarthritis, periodontitis, and osteoporosis, in a wild mammal (moose) population.

OBJECTIVE: Osteoarthritis, periodontitis and osteoporosis are chronic, age-related diseases which adversely impact millions of people worldwide. Because these diseases pose a major global public health challenge, there is an urgent need to better understand how these diseases are interrelated. Our objective was to document the age and sex-specific prevalence of each disease and assess interrelationships among the three diseases in a wild mammal (moose, Alces alces) population. METHODS: We examined the bones of moose dying from natural causes and recorded the severity of osteoarthritis (typically observed on the hip and lowest vertebrae), osteoporosis (osteoporotic lesions observed on the skull) and periodontitis (observed on maxilla and mandibles). RESULTS: Periodontitis was associated with a greater prevalence of both severe osteoarthritis and osteoporotic lesions in moose. We found no evidence to suggest that moose with osteoporotic lesions were more or less likely to exhibit signs of osteoarthritis or severe osteoarthritis. The prevalence of osteoarthritis, periodontitis and osteoporotic lesions was greater among males than for females. CONCLUSIONS: Our results were consistent with the hypothesis that bacterial pathogens causing periodontitis are a risk factor for osteoarthritis and osteoporosis. They are also consistent with the hypothesis that the inverse association between osteoarthritis and osteoporosis sometimes observed in humans may be influenced by shared risk factors, such as obesity, smoking or alcohol consumption, which are absent in moose. Together these results provide insights about three diseases which are expected to become more prevalent in the future and that cause substantial socio-economic burdens.

Increased summer temperature is associated with reduced calf mass of a circumpolar large mammal through direct thermoregulatory and indirect, food quality, pathways.

Climate change represents a growing ecological challenge. The (sub) arctic and boreal regions of the world experience the most rapid warming, presenting an excellent model system for studying how climate change affects mammals. Moose (Alces alces) are a particularly relevant model species with their circumpolar range. Population declines across the southern edge of this range are linked to rising temperatures. Using a long-term dataset (1988-1997, 2017-2019), we examine the relative strength of direct (thermoregulatory costs) and indirect (food quality) pathways linking temperature, precipitation, and the quality of two important food items (birch and fireweed) to variation in moose calf mass in northern Sweden. The direct effects of temperature consistently showed stronger relationships to moose calf mass than did the indirect effects. The proportion of growing season days where the temperature exceeded a 20 °C threshold showed stronger direct negative relationships to moose calf mass than did mean temperature values. Finally, while annual forb (fireweed) quality was more strongly influenced by temperature and precipitation than were perennial (birch) leaves, this did not translate into a stronger relationship to moose calf weight. The only indirect path with supporting evidence suggested that mean growing season temperatures were positively associated with neutral detergent fiber, which was, in turn, negatively associated with calf mass. While indirect impacts of climate change deserve further investigation, it is important to recognize the large direct impacts of temperature on cold-adapted species.

Distribution of large lungworms (Nematoda: Dictyocaulidae) in free-roaming populations of red deer Cervus elaphus (L.) with the description of Dictyocaulus skrjabini n. sp.

Lungworms of the genus Dictyocaulus are causative agents of parasitic bronchitis in domestic and wild ungulates. This study investigates the distribution, morphology and genetic diversity of D. cervi and a new lungworm species, Dictyocaulus skrjabini n. sp. infecting red deer Cervus elaphus, fallow deer Dama dama and moose Alces alces in Poland and Sweden. The study was conducted on 167 red deer from Poland and on the DNA of lungworms derived from 7 fallow deer, 4 red deer and 2 moose collected in Sweden. The prevalence of D. cervi and D. skrjabini n. sp. in dissected red deer in Poland was 31.1% and 7.2%, respectively. Moreover, D. skrjabini n. sp. was confirmed molecularly in 7 isolates of fallow deer lungworms and 1 isolate of red deer lungworms from Sweden. Dictyocaulus skrjabini n. sp. was established based on combination of their distinct molecular and morphological features; these included the length of cephalic vesicle, buccal capsule (BC), buccal capsule wall (BCW), distance from anterior extremity to the nerve ring, the width of head, oesophagus, cephalic vesicle, BC and BCW, as well as the dimensions of reproductive organs of male and female. Additionally, molecular analyses revealed 0.9% nucleotide sequence divergence for 1,605 bp SSU rDNA, and 16.5­17.3% nucleotide sequence divergence for 642 bp mitochondrial cytB between D. skrjabini n. sp. and D. cervi, respectively, and 18.7­19% between D. skrjabini n. sp. and D. eckerti, which translates into 18.2­18.7% amino acid sequence divergence between D. skrjabini n. sp. and both lungworms.

Harvest and density-dependent predation drive long-term population decline in a northern ungulate.

The relative effect of top-down versus bottom-up forces in regulating and limiting wildlife populations is an important theme in ecology. Untangling these effects is critical for a basic understanding of trophic dynamics and effective management. We examined the drivers of moose (Alces alces) population growth by integrating two independent sources of observations within a hierarchical Bayesian population model. We used one of the largest existing spatiotemporal data sets on ungulate population dynamics globally. We documented a 20% population decline over the period examined. There was negative density-dependent population growth of moose. Although we could not determine the mechanisms producing density-dependent suppression of population growth, the relatively low densities at which we documented moose populations suggested it could be due to density-dependent predation. Predation primarily limited population growth, except at low density, where it was regulating. After we simulated several harvest scenarios, it appeared that harvest was largely additive and likely contributed to population declines. Our results highlight how population dynamics are context dependent and vary strongly across gradients in climate, forest type, and predator abundance. These results help clarify long-standing questions in population ecology and highlight the complex relationships between natural and human-caused mortality in driving ungulate population dynamics.

Wolf spatial behavior promotes encounters and kills of abundant prey.

Predators use different spatial tactics to track the prey on the landscape. Three hypotheses describe spatial tactics: prey abundance for prey that are aggregated in space; prey habitat for uniformly distributed prey; and prey catchability for prey that are difficult to catch and kill. The gray wolf (Canis lupus) is a generalist predator that likely employs more than one spatial hunting tactic to match their diverse prey with distinct distributions and behavior that are available. We conducted a study on 17 GPS collared wolves in 6 packs in Riding Mountain National Park, Manitoba, Canada where wolves prey on moose (Alces alces) and elk (Cervus canadensis). We evaluated wolf selection for prey density, habitat selection and catchability on the landscape through within-territory habitat selection analysis. We reveal support for both the prey habitat and prey catchability hypotheses. For moose, their primary prey, wolves employed a mixed habitat and catchability tactic. Wolves used spaces described by the intersection of moose habitat and moose catchability. Wolves selected for the catchability of elk, their secondary prey, but not elk habitat. Counter to our predictions, wolves avoided areas of moose and elk density, likely highlighting the ongoing space race between predator and prey. We illustrate that of the three hypotheses the primary driver was prey catchability, where the interplay of both prey habitat with catchability culminate in predator spatial behaviour in a multiprey system.

Body temperature patterns during pregnancy and parturition in moose.

Gestation and lactation have high energetic requirements. Up to three-fourths of the gestation period in moose (Alces alces) overlaps with the food-scarce period in winter. During this period, moose deal with the limited forage resources available through hypometabolism with decreased heart rate and body temperature (Tb). Body temperature is also an indicator of oestrus, pregnancy and parturition, which is well documented in several domestic species. In this study, we sought to determine if moose displayed a similar Tb pattern during pregnancy and parturition to domesticated ruminants, and if we could detect parturition by combining Tb and activity data. We studied the Tb pattern of 30 free-ranging adult female moose (≥1.5 years old), equipped with ruminal temperature loggers and GPS collars. We documented a 0.13-0.19°C higher Tb in pregnant compared to non-pregnant moose, depending on the study area with the Tb difference increasing along a south-north gradient, and a drop in Tb and in activity when parturition was imminent. Detection of parturition was highly successful when combining Tb and activity data with an accuracy of 91.5%. Our findings demonstrate that Tb responses to pregnancy and parturition in a wild capital-breeding ruminant are similar to those of domesticated ruminants.

Life-history theory provides a framework for detecting resource limitation: a test of the Nutritional Buffer Hypothesis.

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity-a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose (Alces alces) populations that varied in calf recruitment (33-69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

Divergence in parturition timing and vegetation onset in a large herbivore-differences along a latitudinal gradient.

In northern environments, the period of access to high-quality forage is limited, exerting strong selective pressure to optimize the timing of parturition. We analysed timing and variation in moose (Alces alces) parturition dates of 555 females at 18 study sites across 12° of latitude (56-68° N, 1350 km) in Sweden. We found evidence for a spatial match of parturition timing to vegetation onset, but no evidence that moose adjust parturition to vegetation onset in a given year. We found a breakpoint at 64° N. Despite adaptation across latitudes, temporal divergences occurred. Females below 64° N calved after vegetation onset and females above 64° N calved before. Here, parturition before vegetation onset might be a strategy to optimize forage utilization time with the very short growing season. Highly seasonal environments such as at higher latitudes may make it advantageous to adapt parturition towards long-term climatic patterns by matching the most favourable period. Given the direction of temporal divergence, our study suggests that climate change may have less of an impact on moose parturition at northern latitudes than southern latitudes.

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.

Adapt or die-Response of large herbivores to environmental changes in Europe during the Holocene.

Climate warming and human landscape transformation during the Holocene resulted in environmental changes for wild animals. The last remnants of the European Pleistocene megafauna that survived into the Holocene were particularly vulnerable to changes in habitat. To track the response of habitat use and foraging of large herbivores to natural and anthropogenic changes in environmental conditions during the Holocene, we investigated carbon (δ13 C) and nitrogen (δ15 N) stable isotope composition in bone collagen of moose (Alces alces), European bison (Bison bonasus) and aurochs (Bos primigenius) in Central and Eastern Europe. We found strong variations in isotope compositions in the studied species throughout the Holocene and diverse responses to changing environmental conditions. All three species showed significant changes in their δ13 C values reflecting a shift of foraging habitats from more open in the Early and pre-Neolithic Holocene to more forest during the Neolithic and Late Holocene. This shift was strongest in European bison, suggesting higher plasticity, more limited in moose, and the least in aurochs. Significant increases of δ15 N values in European bison and moose are evidence of a diet change towards more grazing, but may also reflect increased nitrogen in soils following deglaciation and global temperature increases. Among the factors explaining the observed isotope variations were time (age of samples), longitude and elevation in European bison, and time, longitude and forest cover in aurochs. None of the analysed factors explained isotope variations in moose. Our results demonstrate the strong influence of natural (forest expansion) and anthropogenic (deforestation and human pressure) changes on the foraging ecology of large herbivores, with forests playing a major role as a refugial habitat since the Neolithic, particularly for European bison and aurochs. We propose that high flexibility in foraging strategy was the key for survival of large herbivores in the changing environmental conditions of the Holocene.

Avoiding misleading messages: Population assessment using camera trapping is not a simple task.

Population assessment is indispensable for appropriate and socially acceptable conservation and management of wildlife populations. This article critiques the paper by Campos-Candela et al. 2018 and highlights issues that could lead to inappropriate management and conservation policies.

Sarcocystis species identification in the moose (Alces alces) from the Baltic States.

Various muscle tissue samples from 60 moose (Alces alces) in the Baltic region were examined for Sarcocystis species. Sarcocysts were detected in 49 out of 60 (81.7%) moose investigated. Six species, Sarcocystis alces, Sarcocystis hjorti, Sarcocystis linearis, Sarcocystis silva, Sarcocystis ovalis, and Sarcocystis sp., were identified using light microscopy (LM), and DNA sequence analysis (cox1 and 18S rDNA). Sarcocysts of S. alces, S. ovalis, and S. hjorti were studied using transmission electron microscopy (TEM); sarcocyst walls of S. alces, S. ovalis, and S. hjorti were type 25, type 24, and type 7a, respectively. Sarcocystis linearis previously found in roe deer and red deer was also shown to use moose as an intermediate host for the first time. The unknown Sarcocystis sp. was rare and might employ another main intermediate host. Phylogenetic results demonstrated that Sarcocystis sp. was most closely related to Sarcocystis tarandivulpes, using canids as definitive hosts.

Novel Type of Chronic Wasting Disease Detected in Moose (Alces alces), Norway.

Chronic wasting disease (CWD) persists in cervid populations of North America and in 2016 was detected for the first time in Europe in a wild reindeer in Norway. We report the detection of CWD in 3 moose (Alces alces) in Norway, identified through a large scale surveillance program. The cases occurred in 13-14-year-old female moose, and we detected an abnormal form of prion protein (PrPSc) in the brain but not in lymphoid tissues. Immunohistochemistry revealed that the moose shared the same neuropathologic phenotype, characterized by mostly intraneuronal deposition of PrPSc. This pattern differed from that observed in reindeer and has not been previously reported in CWD-infected cervids. Moreover, Western blot revealed a PrPSc type distinguishable from previous CWD cases and from known ruminant prion diseases in Europe, with the possible exception of sheep CH1641. These findings suggest that these cases in moose represent a novel type of CWD.

Climate warming is associated with smaller body size and shorter lifespans in moose near their southern range limit.

Despite the importance of body size for individual fitness, population dynamics and community dynamics, the influence of climate change on growth and body size is inadequately understood, particularly for long-lived vertebrates. Although temporal trends in body size have been documented, it remains unclear whether these changes represent the adverse impact of climate change (environmental stress constraining phenotypes) or its mitigation (via phenotypic plasticity or evolution). Concerns have also been raised about whether climate change is indeed the causal agent of these phenotypic shifts, given the length of time-series analysed and that studies often do not evaluate - and thereby sufficiently rule out - other potential causes. Here, we evaluate evidence for climate-related changes in adult body size (indexed by skull size) over a 4-decade period for a population of moose (Alces alces) near the southern limit of their range whilst also considering changes in density, predation, and human activities. In particular, we document: (i) a trend of increasing winter temperatures and concurrent decline in skull size (decline of 19% for males and 13% for females) and (ii) evidence of a negative relationship between skull size and winter temperatures during the first year of life. These patterns could be plausibly interpreted as an adaptive phenotypic response to climate warming given that latitudinal/temperature clines are often accepted as evidence of adaptation to local climate. However, we also observed: (iii) that moose with smaller skulls had shorter lifespans, (iv) a reduction in lifespan over the 4-decade study period, and (v) a negative relationship between lifespan and winter temperatures during the first year of life. Those observations indicate that this phenotypic change is not an adaptive response to climate change. However, this decline in lifespan was not accompanied by an obvious change in population dynamics, suggesting that climate change may affect population dynamics and life-histories differently.

Effective thiafentanil immobilization and physiological responses of free-ranging moose (Alces alces) in northern Sweden.

OBJECTIVE: To evaluate clinical and physiological responses in moose to thiafentanil administration for immobilization. STUDY DESIGN: Cross-sectional clinical study. ANIMALS: Eleven (six males and five females) free-ranging adult moose (Alces alces). METHODS: Each moose was darted from a helicopter with 7.5 mg thiafentanil during March 2014 in northern Sweden. Physiological evaluation included vital signs and blood gases. Arterial blood was collected after induction and again after 10 minutes of intranasal oxygen administration and analyzed immediately with an i-STAT analyzer. A total of 10 mg naltrexone per milligram of thiafentanil was administered to all animals for reversal. Data were analyzed using descriptive statistics. RESULTS: All moose were sufficiently immobilized with a single dart injection. Induction occurred within 3 minutes in 10 of 11 moose. One individual became recumbent while crossing a river and naltrexone was immediately administered. Animals maintained sternal recumbency with their head raised and vital signs were stable. Nine of 10 moose were hypoxemic before oxygen administration, with seven becoming markedly hypoxemic [partial pressure of arterial oxygen (PaO2) between 40 and 59 mmHg (5.3-7.9 kPa)]. The PaO2 increased significantly between samples, but six moose remained hypoxemic despite therapy. Hypercapnia was seen in all moose, with eight having marked hypercapnia [partial pressure of arterial carbon dioxide (PaCO2) > 60 mmHg (>8.0 kPa)]. All moose were acidemic, with nine showing marked acidemia (pH < 7.20). The pH increased significantly with time and lactate decreased. Recoveries were rapid and uneventful, and all moose were living 6 months after capture. CONCLUSIONS: Thiafentanil provided rapid and sufficient immobilization of moose and its effects were rapidly reversed with naltrexone. As with other opioids, moose showed hypoxemia and varying degrees of respiratory and metabolic acidosis. Arterial oxygenation of moose improved following intranasal oxygen, but hypoxemia was not fully resolved despite therapy. CLINICAL RELEVANCE: Thiafentanil (7.5 mg per adult) is effective for immobilization of free-ranging moose. Supplemental oxygen may be of benefit when using this regimen; however, further investigation is required to confirm these results.

A TECHNIQUE FOR DEPLOYMENT OF RUMEN BOLUS TRANSMITTERS IN FREE-RANGING MOOSE ( ALCES ALCES).

Recent uses for rumen boluses, such as mortality implant transmitters (MITs), in wildlife have made it necessary to adapt deployment techniques developed for livestock. In 29 and 30 attempts to place MITs in Minnesota free-ranging moose ( Alces alces) in 2013 and 2014, respectively, success was achieved 83% and 63% of the time. In 2014, new methods for MIT deployment were evaluated in captive moose in Alaska. Mandible measurements provided guidance for selection of an appropriate-sized bolus applicator. A Schulze mouth gag was used to aid insertion of the applicator, and canola oil was used to lubricate the bolus to facilitate swallowing. Time to first swallow and time to continuous swallow following sedative reversal was measured to gauge appropriate timing for bolus administration. Using the adapted technique with trained personnel, success rates for MIT deployment were 100% (10/10) for captive moose and 88% (21/24) for free-ranging moose in Minnesota in 2015.

MOOSE ( ALCES ALCES) MORTALITY ASSOCIATED WITH CAPRINE HERPESVIRUS 2 (CPHV-2) IN A ZOOLOGICAL COLLECTION.

Malignant catarrhal fever (MCF) can affect both domestic and wild artiodactyls. In a zoological setting, in which subclinical carriers and susceptible species are often housed in close proximity, the disease can prove fatal. This report describes a case of goat-associated MCF in a captive moose ( Alces alces). The diagnosis was confirmed by histopathology, which showed lymphocytic vasculitis in the brain and panuveitis, and by detection of caprine herpesvirus 2 DNA in tissues. Identical viral DNA sequences amplified from the clinically affected moose and from domestic, petting goats ( Capra aegagrus hircus) housed in the zoo suggest that the goats were the source for the virus transmutation. This is the first report, to our knowledge, of confirmed goat-associated MCF in any moose in North America and of the surveillance measures and procedures put in place to prevent additional spread of the disease.

Group or ungroup - moose behavioural response to recolonization of wolves.

BACKGROUND: Predation risk is a primary motivator for prey to congregate in larger groups. A large group can be beneficial to detect predators, share predation risk among individuals and cause confusion for an attacking predator. However, forming large groups also has disadvantages like higher detection and attack rates of predators or interspecific competition. With the current recolonization of wolves (Canis lupus) in Scandinavia, we studied whether moose (Alces alces) respond by changing grouping behaviour as an anti-predatory strategy and that this change should be related to the duration of wolf presence within the local moose population. In particular, as females with calves are most vulnerable to predation risk, they should be more likely to alter behaviour. METHODS: To study grouping behaviour, we used aerial observations of moose (n = 1335, where each observation included one or several moose) inside and outside wolf territories. RESULTS: Moose mostly stayed solitary or in small groups (82% of the observations consisted of less than three adult moose), and this behavior was independent of wolf presence. The results did not provide unequivocal support for our main hypothesis of an overall change in grouping behaviour in the moose population in response to wolf presence. Other variables such as moose density, snow depth and adult sex ratio of the group were overall more influential on grouping behaviour. However, the results showed a sex specific difference in social grouping in relation to wolf presence where males tended to form larger groups inside as compared to outside wolf territories. For male moose, population- and environmentally related variables were also important for the pattern of grouping. CONCLUSIONS: The results did not give support for that wolf recolonization has resulted in an overall change in moose grouping behaviour. If indeed wolf-induced effects do exist, they may be difficult to discern because the effects from moose population and environmental factors may be stronger than any change in anti-predator behaviour. Our results thereby suggest that caution should be taken as to generalize about the effects of returning predators on the grouping behaviour of their prey.