The Wolf Intelligencer


Moose (Alces alces) – Eurasia

Basque: altze, Catalan: ant, Chinese: 駝鹿, Croatian: los, Czech: los, Danish: elg, Dutch: eland, Estonian: Põder, Finnish: hirvi, Flemish: eland, French: elan. German: Elch, Greek: έλαφος, Hungarian: jávorszarvas, Icelandic: elgur, Italian: alce, Japanese: ムース, Latvian: alnis, Lithuanian: briedis, Mongolian: Хөвөн, Norwegian: elg, Polish: łoś, Portugese: alce, Romanian: elan, Russian: лось, Slovenian: losa, Spanish: alce, Swedish: älg

Journal Articles

The return of large carnivores: Using hunter observation data to understand the role of predators on ungulate populations. Tallian A, Ordiz A, Zimmermann B, Sand H, Wikenros C, Wabakken P, Bergqvist G, Kindberg J. Global Ecology and Conservation. 2021 Apr


Large carnivores play a key ecological role in nature, yet quantifying the effects of predation at large spatiotemporal scales remains challenging. Wolves and brown bears have recovered in Sweden, where they share the same staple prey, moose. This ecosystem is representative of the Eurasian boreal realm, and makes an interesting case study for exploring the use of hunter-collected observation data to understand the ecological effects of large carnivore recovery in human-dominated ecosystems. We used over 50 million hours of moose observation data collected by “citizen scientists” (i.e., moose hunters) between 2000 and 2017 to evaluate the role of recovering bear and wolf populations on an important ungulate vital rate, calf/cow ratio (the number of observed calves per female in fall). Calf/cow ratios were negatively correlated with both wolf and bear densities, suggesting that summer calf survival decreased via predation. Calf/cow ratios decreased by 7% and 17% in the northern and southern areas where bears and wolves were allopatric, respectively. Where wolves and bears were sympatric, the effect of predator densities was additive; calf/cow ratios decreased by 18%. However, both calf/cow ratios and moose densities declined over the last ~20 years across Sweden, including areas where wolves and bears were absent. While recolonizing large carnivores affected ungulate vital rates in Sweden’s human-dominated landscape, they were likely not the primary driver of long-term moose population trends. Our results reinforce that citizen-collected data are a useful wildlife monitoring tool that help understand ecological processes, including the effects of recovering carnivores on prey populations.

Non-naivety in a long-lived ungulate Graf, L., 2021.


Habitat selection and movement are considered important factors for survival of game species when they are hunted by humans. To reduce their risk of predation, animals can adapt their behavior to their most abundant predator over time through experience. To test for learning capabilities in a long-lived ungulate, I used GPS-data of 19 male and 84 female moose (Alces alces) in two study areas in southern Sweden from 2008-2018. I matched the GPS-data with reproduction and survival data and analyzed movement rates and habitat selection in a heavily managed moose population. While not being a gregarious species, moose are expected to learn from non-lethal mortality when a female loses her calf to harvest. Hunters were more likely to harvest male moose that moved faster before the moose hunting season began. Female moose where more likely to get harvested when they selected more for open habitats. The results of my study suggest that female moose in my study areas increased their shyness in the next hunting season after losing a calf to harvest. Female moose became increased nocturnalactivity and increased their avoidance of open habitats. As they aged, female moose decreased their movement rates and avoided open habitats. Using these proxies for shyness indicates that being shyer is beneficial for moose to survive the hunting season. My study suggests that learning effects accumulate as moose age. Furthermore, my results help to understand the influence human hunters impose on behavior of moose in a heavily managed population.

Sauvala M, Laaksonen S, Laukkanen-Ninios R, Jalava K, Stephan R, Fredriksson-Ahomaa M. Microbial contamination of moose (Alces alces) and white-tailed deer (Odocoileus virginianus) carcasses harvested by hunters. Food microbiology. 2019 Apr
(Finland, Hunter hygeine. “Moose carcasses had mean mesophilic aerobic bacteria (MAB), Enterobacteriaceae (EB) and Escherichia coli (EC) values of 4.2, 2.6 and 1.2 log10 cfu/cm2,” “High EB counts on the carcasses indicated a gut hit.”)

Novel type of chronic wasting disease detected in moose (Alces alces), Norway. Pirisinu L, Tran L, Chiappini B, Vanni I, Di Bari MA, Vaccari G, Vikøren T, Madslien KI, Våge J, Spraker T, Mitchell G. Emerging infectious diseases. 2018 Dec
(Trøndelag County, Norway. “We report 3 cases of CWD detected in moose in Norway, characterized by biochemical and IHC features clearly different from CWD cases previously described in North America and Norway. Our findings suggest the involvement of a different type of CWD prion.”)

Identification of the pregnancy-associated glycoprotein family (PAGs) and some aspects of placenta development in the European moose (Alces alces L.). Lipka A, Panasiewicz G, Majewska M, Bieniek-Kobuszewska M, Saveljev AP, Pankratov AP, Szafranska B. Theriogenology. 2016 Dec
(Olsztyn-Kortowo, Poland, “The crown-rump length and PLC size measurements permitted a novel pattern estimation of various pregnancy stages in wild Aa. The PLC number varied (5–21) and was the greatest at the mid and late stages of gestation in females bearing singletons or twins. The genomic existence of the identified PAG-like family was named AaPAG-L.”)

Moose anti-predator behaviour towards baying dogs in a wolf-free area. European journal of wildlife research. 2015 Aug
(Sweden, “Northern Sweden devoid of wolves, we exposed adult female moose (Alces alces) to hunting activity using a dog (Canis lupus familiaris) to study individual moose anti-predator behaviour. Moose were more active, had larger 24-h activity ranges and left the area after the disturbance… We argue that heavily harvested Scandinavian moose may be more adapted to human and bear predation in contrast to the suggested behavioural maladaption towards wolf predation.”)

Response of moose hunters to predation following wolf return in Sweden. Wikenros C, Sand H, Bergström R, Liberg O, Chapron G. PloS one. 2015 Apr

Synchronizing migration with birth: An exploration of migratory tactics in female moose. Näsén, L., 2015

Spatial structure in European moose (Alces alces): genetic data reveal a complex population history. Niedziałkowska M, Hundertmark KJ, Jędrzejewska B, Niedziałkowski K, Sidorovich VE, Górny M, Veeroja R, Solberg EJ, Laaksonen S, Sand H, Solovyev VA. Journal of Biogeography. 2014 Nov

Testing the risk of predation hypothesis: the influence of recolonizing wolves on habitat use by moose. Nicholson KL, Milleret C, Månsson J, Sand H. Oecologia. 2014 Sep

Admixture promotes genetic variation in bottlenecked moose populations in eastern Poland. Świsłocka M, Czajkowska M, Duda N, Ratkiewicz M. Mammal Research. 2015 Apr

Performance of hunting statistics as spatiotemporal density indices of moose (Alces alces) in Norway. Ueno M, Solberg EJ, Iijima H, Rolandsen CM, Gangsei LE. Ecosphere. 2014 Feb

Complex patterns of population genetic structure of moose, Alces alces, after recent spatial expansion in Poland revealed by sex-linked markers. Świsłocka M, Czajkowska M, Duda N, Danyłow J, Owadowska-Cornil E, Ratkiewicz M. Acta theriologica. 2013 Oct

Sharing the bounty—Adjusting harvest to predator return in the Scandinavian human–wolf–bear–moose system. Jonzén N, Sand H, Wabakken P, Swenson JE, Kindberg J, Liberg O, Chapron G. Ecological modelling. 2013 Sep

Predation on adult moose Alces alces by European brown bears Ursus arctos. Dahle B, Wallin K, Cederlund G, Persson IL, Selvaag LS, Swenson JE. Wildlife Biology. 2013 Jun

To run or stay: anti-hunter behaviour of female moose (Master’s thesis). Johnsen, S., 2013.

Moose (Alces alces) calf survival rates in the presence of wolves (Canis lupus) in southeast Norway. Sivertsen TR, Mysterud A, Gundersen H. European journal of wildlife research. 2012 Oct

Spatio-temporal relationship between calf body mass and population productivity in Fennoscandian moose Alces alces. Tiilikainen R, Solberg EJ, Nygrén T, Pusenius J. Wildlife biology. 2012 Sep;

Assessing the influence of prey–predator ratio, prey age structure and packs size on wolf kill rates. Sand H, Vucetich JA, Zimmermann B, Wabakken P, Wikenros C, Pedersen HC, Peterson RO, Liberg O. Oikos. 2012 Sep

Comparing body condition of moose (Alces alces) selected by wolves (Canis lupus) and human hunters: consequences for the extent of compensatory mortality. Sand H, Wikenros C, Ahlqvist P, Strømseth TH, Wabakken P. Canadian Journal of Zoology. 2012 Mar

Calving site selection by moose (Alces alces) along a latitudinal gradient in Sweden. na; Haydn A. 2012 Jan

Effect of immobilizations on the activity and space use of female moose (Alces alces). Canadian Journal of Zoology. 2011 Oct

Prevalence of antibodies against Toxoplasma gondii and Neospora caninum in moose (Alces alces) and roe deer (Capreolus capreolus) in Sweden. Malmsten J, Jakubek EB, Björkman C. Veterinary Parasitology. 2011 May

Moose Alces alces habitat use at multiple temporal scales in a human-altered landscape. Bjørneraas K, Solberg EJ, Herfindal I, Van Moorter B, Rolandsen CM, Tremblay JP, Skarpe C, Sæther BE, Eriksen R, Astrup R. Wildlife Biology. 2011 Mar

Activity patterns of predator and prey: a simultaneous study of GPS-collared wolves and moose. Eriksen A, Wabakken P, Zimmermann B, Andreassen HP, Arnemo JM, Gundersen H, Liberg O, Linnell J, Milner JM, Pedersen HC, Sand H. Animal Behaviour. 2011 Feb

Differential forage use makes carrying capacity equivocal on ranges of Scandinavian moose (Alces alces). Wam HK, Hjeljord O, Solberg EJ. Canadian Journal of Zoology. 2010 Nov

Wolf predation on moose and roe deer: chase distances and outcome of encounters. Wikenros C, Sand H, Wabakken P, Liberg O, Pedersen HC. Acta Theriologica. 2009 Sep

Environmental variation and moose Alces alces density as determinants of spatio‐temporal heterogeneity in browsing. Månsson J. Ecography. 2009 Aug

The non-impact of hunting on moose Alces alces movement, diurnal activity, and activity range. Neumann W, Ericsson G, Dettki H. European Journal of Wildlife Research. 2009 Jun

Encounter frequencies between GPS‐collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory. Eriksen A, Wabakken P, Zimmermann B, Andreassen HP, Arnemo JM, Gundersen H, Milner JM, Liberg O, Linnell J, Pedersen HC, Sand H. Ecological Research. 2009 May

The role of ecological and economic factors in the management of a spatially structured moose Alces alces population. Nilsen EB, Skonhoft A, Mysterud A, Milner JM, Solberg EJ, Andreassen HP, Stenseth NC. Wildlife biology. 2009 Mar

Moose Alces alces behaviour related to human activity. Neumann W. 2009

Lack of compensatory body growth in a high performance moose Alces alces population. Solberg EJ, Garel M, Heim M, Grøtan V, Sæther BE. Oecologia. 2008 Dec

Evaluation of four methods used to estimate population density of moose Alces alces. Rönnegård L, Sand H, Andrén H, Månsson J, Pehrson Å. Wildlife Biology. 2008 Sep

Mitochondrial DNA diversity in the moose, Alces alces, from northeastern Poland: evidence for admixture in a bottlenecked relic population in the Biebrza valley. Świsłocka M, Ratkiewicz M, Borkowska A, Komenda E, Raczyński J. InAnnales Zoologici Fennici 2008 Aug

Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?. Sand H, Wabakken P, Zimmermann B, Johansson Ö, Pedersen HC, Liberg O. Oecologia. 2008 May

Three approaches to estimate wolf Canis lupus predation rates on moose Alces alces populations. Gundersen H, Solberg EJ, Wabakken P, Storaas T, Zimmermann B, Andreassen HP. European Journal of Wildlife Research. 2008 May

Quantitative estimates of tree species selectivity by moose (Alces alces) in a forest landscape. Månsson J, Kalén C, Kjellander P, Andrén H, Smith H. Scandinavian Journal of Forest Research. 2007 Oct

Fidelity to calving areas in moose (Alces alces) in the absence of natural predators. Tremblay JP, Solberg EJ, Sæther BE, Heim M. Canadian Journal of Zoology. 2007 Aug;

Multistage, long‐range natal dispersal by a global positioning system‐collared Scandinavian wolf. Wabakken P, Sand H, Kojola I, Zimmermann B, Arnemo JM, Pedersen HC, Liberg O. The Journal of Wildlife Management. 2007 Jul

Wolf movement patterns: a key to estimation of kill rate?. Zimmermann B, Wabakken P, Sand H, Pedersen HC, Liberg O. The Journal of wildlife management. 2007 Jun

Cross-continental differences in patterns of predation: will naive moose in Scandinavia ever learn?. Proceedings of the Royal Society B: Biological Sciences, Sand, H., Wikenros, C., Wabakken, P. and Liberg, O., 2006

Effects of hunting group size, snow depth and age on the success of wolves hunting moose. Sand H, Wikenros C, Wabakken P, Liberg O. Animal Behaviour. 2006 Oct

Patterns of hunting mortality in Norwegian moose (Alces alces) populations. Nilsen EB, Solberg EJ. European Journal of Wildlife Research. 2006 Aug

Risk of capture-related mortality in large free-ranging mammals: experiences from Scandinavia. Arnemo JM, Ahlqvist P, Andersen R, Berntsen F, Ericsson G, Odden J, Brunberg S, Segerström P, Swenson JE. Wildlife Biology. 2006 Mar

Moose hunting, forestry, and wolves in Sweden. Bergman M, Åkerberg S. Alces. 2006 Jan

Winter prey selection of moose (Alces alces) by Scandinavian wolves (Canis lupus) Bernelind, T.,  (Doctoral dissertation, Sveriges lantbruksuniversitet. Institutionen för naturvårdsbiologi). 2006

Using GPS technology and GIS cluster analyses to estimate kill rates in wolf‐ungulate ecosystems. Sand H, Zimmermann B, Wabakken P, Andrèn H, Pedersen HC. Wildlife Society Bulletin. 2005 Sep

Change-in-sex ratio as an estimator of population size for Norwegian moose Alces alces. Solberg EJ, Grøtan V, Rolandsen CM, Brøseth H, Brainerd S. Wildlife Biology. 2005 Jun

Moose harvesting strategies in the presence of wolves. Nilsen EB, Pettersen T, Gundersen H, Milner JM, Mysterud A, Solberg EJ, Andreassen HP, Stenseth NC. Journal of Applied Ecology. 2005 Apr

Moose recovery plan in Poland: main objectives and tasks. Bobek B, Merta D, Sułkowski P, Siuta A. Alces. 2005 Jan

Supplemental feeding of migratory moose Alces alces: forest damage at two spatial scales. Gundersen H, Andreassen HP, stein Storaas T. Wildlife Biology. 2004 Sep

Habitat selection of adult moose Alces alces at two spatial scales in central Finland. Nikula A, Heikkinen S, Helle E. Wildlife Biology. 2004 Jun

Status of moose populations and challenges to moose management in Fennoscandia. Lavsund S, Nygrén T, Solberg EJ. Alces. 2003 Jan

Wolf predation on moose––a case study using hunter observations. Wam HK, Hjeljord O. Alces. 2003 Jan

Management and regulated harvest of moose (Alces alces) in Sweden. Sylvén S. 2003

Genetic diversity of moose (Alces alces L.) in Eurasia. Udina IG, Danilkin AA, Boeskorov GG.Russian Journal of Genetics. 2002 Aug

Browsing damage on pine (Pinus sylvestris and P. contorta) by a migrating moose (Alces alces) population in winter: Relation to habitat composition and road barriers. Ball JP, Dahlgren J. Scandinavian Journal of Forest Research. 2002 Jan

Moose Alces alces hunting in Finland-an ecological risk analysis. Luoma A, Ranta E, Kaitala V. Wildlife Biology. 2001 Sep

Changes in population density of moose (Alces alces) and damage to forests in Sweden. Hörnberg S. Forest ecology and management. 2001 Aug

Age‐related reproductive effort and senescence in free‐ranging moose, Alces alces. Ericsson G, Wallin K, Ball JP, Broberg M. Ecology. 2001 Jun

Age-specific moose (Alces alces) mortality in apredator-free environment: Evidence for senescence in females. Ericsson G, Wallin K. Ecoscience. 2001 Jan

Patterns of stem damage by moose (Alces alces) in young Pinus sylvestris stands in Sweden. Bergqvist G, Bergström R, Edenius L. Scandinavian Journal of Forest Research. 2001 Jan

Age-specific harvest mortality in a Norwegian moose Alces alces population. Solberg EJ, Loison A, Sæther BE, Strand O. Wildlife Biology. 2000 Mar

Diet composition of wolves Canis lupus in east-central Finland. Gade-Jorgensen I, Stagegaard R. Acta Theriologica. 2000

Moose (Alces alces) survival in three populations in northern Norway. Stubsjøen T, Sæther BE, Solberg EJ, Heim M, Rolandsen CM. Canadian Journal of Zoology. 2000 Oct

Hunter observations of moose Alces alces as a management tool. Solberg EJ, Saether BE. Wildlife Biology. 1999 Jun

Ecological correlates of regional variation in life history of the moose Alces alces: comment. Crête M. Ecology. 1998 Jul

Wolf Canis lupus predation on moose Alces alces and roe deer Capreolus capreolus in south-central Scandinavia. Olsson O, Wirtberg J, Andersson M, Wirtberg I. Wildlife biology. 1997 Mar

Ecological correlates of regional variation in life history of the moose Alces alces. Saether BE, Andersen R, Hjeljord O, Heim M. Ecology. 1996 Jul

Life history patterns in female moose (Alces alces): the relationship between age, body size, fecundity and environmental conditions. Sand H. Oecologia. 1996 Apr

Male traits as life-history variables: annual variation in body mass and antler size in moose (Alces alces). Solberg EJ, Sæther BE. Journal of Mammalogy. 1994 Nov

Moose browsing on pine and willow in the Biebrza Valley, Poland. Borkowska A, Konopko A. Acta Theriologica. 1994 Jan

 Genetic differentiation of Scandinavian moose Alces alces populations over short geographical distances. Chesser RK, Reuterwall C, Ryman N.Oikos. 1982 Aug

The occurrence and some ecological parameters of the moose in Poland. Tomek A. Acta Theriologica. 1977 Jan

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