Abstract Context Ungulate prey can use increased vigilance to reduce their risk of predation, but little is known of the combined and interactive risk effects from humans and wolves in determining ungulate behaviour across time and space. Understanding the interplay between these risk effects is increasingly important, considering the recolonisation of several large carnivores to more human-dominated landscapes in Europe. Aim The aim of the present study was to assess the vigilance behaviour expressed by red deer (Cervus elaphus) in response to both humans and wolves in the Polish Bialowieza Forest. Methods Using a camera-trap transect, the effect of distance to human settlements, hunting season, patterns of space use by wolves (Canis lupus), canopy openness, canopy height, time of day, as well as sex/age of individuals, on the vigilance behaviour observed in red deer was studied using a model-selection approach. Key results We did not find a clear effect of patterns of space use by wolves or distance to human settlements on red deer vigilance behaviour at the landscape scale. However, red deer showed increased vigilance during the hunting season and during the day outside of protected areas and reserves, because disturbance from human hunters is highest. Conversely, we also found that red deer were more vigilant at night within more protected areas, which is likely to be explained by the increased activity of wolves because human activity is strictly limited. Conclusions Our study showed that vigilance behaviour of red deer in Bialowieza Primeval Forest is more driven by human hunting than by the frequency of wolf presence at a landscape scale. This could be explained by the higher temporal and spatial predictability of human hunting activities than wolf risk. We found that patterns of wolf space use, as opposed to the omnipresent fear effects from humans, had only localised effects by increasing vigilance levels during night hours in non-hunting areas of the forest. The reverse was observed outside of protected reserves. Understanding how prey species respond to this new combination of risk from natural predators and humans, is increasingly important in a landscape where human risk is becoming ever more potent and carnivores recolonise.

中文翻译：

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马鹿根据人类猎人和狼的时空风险模式不同地分配警惕性

摘要 背景 有蹄类动物可以提高警惕来降低被捕食的风险，但人们对人类和狼在确定有蹄类动物跨时空行为的综合和互动风险影响知之甚少。考虑到几种大型食肉动物重新殖民到欧洲更多由人类主导的景观，了解这些风险影响之间的相互作用变得越来越重要。目的 本研究的目的是评估马鹿 (Cervus elaphus) 对波兰 Bialowieza 森林中的人类和狼的警觉行为。方法 使用相机陷阱横断面、与人类住区的距离、狩猎季节、狼（Canis lupus）空间使用模式、树冠开放度、树冠高度、一天中的时间以及个体的性别/年龄的影响，使用模型选择方法研究了在马鹿中观察到的警惕行为。主要结果 我们没有发现狼的空间使用模式或与人类住区的距离在景观尺度上对马鹿警惕行为的明显影响。然而，马鹿在狩猎季节以及保护区和保护区外的白天表现出更高的警惕性，因为人类猎人的干扰最大。相反，我们还发现，在更多保护区内，马鹿在夜间更加警惕，这可能是由于人类活动受到严格限制，狼的活动增加了。结论 我们的研究表明，比亚沃维耶扎原始森林中马鹿的警戒行为更多地是由人类狩猎而不是狼在景观尺度上出现的频率所驱动。这可以通过人类狩猎活动比狼风险更高的时间和空间可预测性来解释。我们发现，与人类无所不在的恐惧效应相反，狼空间的使用模式仅通过提高夜间森林非狩猎区的警惕水平而产生局部影响。在保护区之外观察到相反的情况。了解猎物物种如何应对来自自然捕食者和人类的这种新的风险组合，在人类风险变得越来越强大和食肉动物重新殖民的景观中变得越来越重要。仅通过在夜间在森林的非狩猎区域提高警惕水平而产生局部影响。在保护区之外观察到相反的情况。了解猎物物种如何应对来自自然捕食者和人类的这种新的风险组合，在人类风险变得越来越强大和食肉动物重新殖民的景观中变得越来越重要。仅通过提高夜间在森林非狩猎区的警戒水平而产生局部影响。在保护区之外观察到相反的情况。了解猎物物种如何应对来自自然捕食者和人类的这种新的风险组合，在人类风险变得越来越强大和食肉动物重新殖民的景观中变得越来越重要。