## 4 Discussion
In summary, our results demonstrate that there was a high amount of gene flow in Scandinavian red foxes, probably due to frequent movements across long distances and across country borders, though with some limited differences in local populations. The extent of gene flow entails that Norwegian and Swedish populations are near identical in genetic structure and can be viewed as one, large, panmixing population. A previous study showed a general trend of a northward dispersal patterns in Fennoscandian red foxes [7]. In our study, we revealed contrasting patterns between historic and contemporary gene flow. We observed a high degree of historic gene flow from Hedmark North to Sweden North, whereas contemporary gene flow followed an opposite pattern, i.e., from south to north. The highest gene flow was estimated to be from Hedmark South to Hedmark North, as well as from Sweden South and North to Hedmark North. Dispersal in a northern direction was expected from climate-induced range shifts (e.g., References [59; 60]). However, the weak genetic structuring and high amount of gene flow between the geographical regions also indicated that gene flow occurred from several different source populations, and not only in a northern direction, which may indicate range expansion [7]. Changes in environmental conditions, or biotic changes, such as prey availability, may induce population increases, which again can influence the dispersal and expansion from local areas [61]. With increasing temperatures due to climate change, in addition to other anthropogenic changes, has increased the range of suitable red fox habitats northwards [62; 63]. Another influential factor may be the major outbreak of the parasitic disease sarcoptic mange (_Sarcopres scabiei_) that took place in 1975 with its origin in North-Central Sweden. The outbreak resulted in a population decrease of up to 90% in the affected red fox populations [64]. This reduction in red fox density may have created more vacant territories, inducing recent recolonization due to a population recovery [65]. Our findings are in line with the findings of [PERSON] et al. [7] in their study of the geographic patterns of expansion in the Fennoscandian red fox. By comparing historical and contemporary samples of red foxes from Fennoscandia, they revealed low genetic structuring, high connectivity, and asymmetric migration rates, which indicates source-sink dynamics in the Fennoscandian red fox population. The same pattern was found in our study, and this indicates that the weak genetic structuring and high amount of gene flow is in line with the classic signs of a range expansion or recolonization. It is likely that the major outbreak of sarcoptic mange created sink areas with a low red fox population density, and that these sink areas are now being recolonized from a range of sources located in a southerly direction. It is possible that the shifting direction in migration rates from the historical southerly direction to a more northwesterly direction in the contemporary analyses in this study was due to historic large-scale biotic changes that has induced a range shift or changed the direction of the population expansion. As the outbreak of mange in 1975 started in North-Central Sweden [64], it is a possibility that the outbreak first created sink areas available for recolonization in a southern direction before the outbreak spread northwards. However, to investigate this, thoroughly historic samples of red foxes should be analyzed. [PERSON] et al. [10] in their study of GPS-marked red foxes in Sweden and Norway found a statistically non-significant northwesterly trend in long-distance dispersal movements of red foxes that coincides with the findings in this study. Similar patterns of a preference for long-range dispersal in a northerly direction has been described for several other red fox populations [3; 66; 67].