Wolf denning behaviour in response to external disturbances and implications for pup survival

We conducted a 15-year study (1999–2014) of wolf denning behavior in Naliboki Forest, Belarus, to determine the extent to which some external factors (rain, mosquitoes, human disturbances) and litter size are negatively correlated with pup survival until autumn.For this purpose we entirely documented denning areas of 30 breeding wolf pairs. Initial denning area includes the natal den site and (ii) secondary den sites between which the parent wolves transported sucessively the litter during the rearing season.The number of den sites per denning area ranged from 1 to 4. The distances in kilometres differed significantly between den sites. The total number of dens (surface dens and burrows) was higher in the natal den site compared with the other den sites. The number of surface dens per den site was higher compared with burrows in den sites 1, 2 and 3 but not den site 4. A total of 168 pups was found over the 15 years. Litter sizes ranged from 1 to 11 (average 5.6 ± 1.9). The rate of pup survival ranged from 0 to 100% and averaged 54.5 ± 29%.We examined the bivariate association between environmental conditions (mosquitoes, rainfall and human presence) and number of surface dens, total number of burrows and number of den sites. Environmental conditions were assessed from spring to mid-summer. Results show that there was a positive relationship between environmental conditions and number of den sites, and also that more difficult environmental conditions (rainy weather, many mosquitoes) decreased pup survival both in absolute and in relative terms. A cluster analysis showed that pup survival was also influenced by human presence. Wolves probably have a more complicated denning behaviour following stressful event which may lead to less effectiveness for feeding or protecting pups. The number of pups surviving was also found to be lower when there were many (8–10) pups because den shifting is more risky in this case. Burrows improve pup survival. Our results tend to suggest that, in our study area, a high mortality rate of wolf pups could be related to external and internal factors of stress.