Expansion of urban areas, landscape transformation and increasing human outdoor activities strongly affect wildlife behaviour. The outbreak of the COVID-19 pandemic in particular led to drastic changes in human behaviour, exposing wildlife around the world to either reduced or increased human presence, potentially altering animal behaviour. Here, we investigate behavioural responses of wild boar (Sus scrofa) to changing numbers of human visitors to a suburban forest near Prague, Czech Republic, during the first 2.5 years of the COVID-19 epidemic (April 2019-November 2021). We used bio-logging and movement data of 63 GPS-collared wild boar and human visitation data based on an automatic counter installed in the field. We hypothesised that higher levels of human leisure activity will have a disturbing effect on wild boar behaviour manifested in increased movements and ranging, energy spent, and disrupted sleep patterns. Interestingly, whilst the number of people visiting the forest varied by two orders of magnitude (from 36 to 3431 people weekly), even high levels of human presence (>2000 visitors per week) did not affect weekly distance travelled, home range size, and maximum displacement of wild boar. Instead, individuals spent 41 % more energy at high levels of human presence (>2000 visitors per week), with more erratic sleep patterns, characterised by shorter and more frequent sleeping bouts. Our results highlight multifaceted effects of increased human activities ('anthropulses'), such as those related to COVID-19 countermeasures, on animal behaviour. High human pressure may not affect animal movements or habitat use, especially in highly adaptable species such as wild boar, but may disrupt animal activity rhythms, with potentially detrimental fitness consequences. Such subtle behavioural responses can be overlooked if using only standard tracking technology.

Department of Biosciences Swansea University Singleton Park Swansea SA2 8PP Wales UK

Department of Forestry and Wood Economics Faculty of Forestry and Wood Sciences Czech University of Life Sciences Kamýcká 129 Prague 6 Suchdol 165 00 Czech Republic

Department of Game Management and Wildlife Biology Faculty of Forestry and Wood Sciences Czech University of Life Sciences Kamýcká 129 Prague 6 Suchdol 165 00 Czech Republic

Department of Game Management and Wildlife Biology Faculty of Forestry and Wood Sciences Czech University of Life Sciences Kamýcká 129 Prague 6 Suchdol 165 00 Czech Republic; Mammal Research Institute Polish Academy of Sciences Stoczek 1 17 230 Białowieża Poland

School of Biological Sciences Queen's University Belfast 19 Chlorine Gardens Belfast BT95DL Northern Ireland UK

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