Abstract Habitat selection of animals is influenced by spatial heterogeneity as well as temporal environmental dynamics. In addition, human activities potentially have severe influences on the habitat selection of animals, often resulting in more nocturnal behavior. We investigated seasonal and circadian habitat selection patterns of red deer (Cervus elaphus) on a military training area in Bavaria (Germany). Individual deer ranged on two neighboring, but non-overlapping sites differing in landscape composition and human activity. Using GPS telemetry data, we visually investigated selection patterns and then fitted step-selection functions to assess multiple approaches to account for temporal (i.e., diel and seasonal) effects and for the impact of habitat conditions on selection. We first showed that the way in which time of day is considered in step-selection functions is essential for obtaining ecologically meaningful results, and that the usual ways of including time (i.e., either based on clock-time or based on night vs. day categories) can lead to incomplete or misleading conclusions. Furthermore, we found that individuals followed either circadian or seasonal habitat selection patterns, depending on the site they inhabited. This can be explained by differences in disturbance between the two sites. Except in winter, individuals selected for open habitats during the night irrespective of human activity level, but only individuals from the more disturbed site selected for covered habitats during the day. Our results highlight the importance of carefully considering the appropriate temporal scale for habitat selection analyses. Our findings also indicate that red deer are not a crepuscular or nocturnal species per se, as is often observed in human-dominated landscapes. Instead, our results imply that nocturnality in red deer is an effect of adaption to human activities.

中文翻译：

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如此接近又如此不同：考虑时间动态以了解栖息地选择的重要性

摘要 动物的栖息地选择受空间异质性和时间环境动态的影响。此外，人类活动可能对动物的栖息地选择产生严重影响，通常会导致更多的夜间行为。我们在巴伐利亚（德国）的军事训练区调查了马鹿（Cervus elaphus）的季节性和昼夜节律栖息地选择模式。个别鹿分布在两个相邻但不重叠的地点，景观构成和人类活动各不相同。使用 GPS 遥测数据，我们直观地研究了选择模式，然后拟合了步进选择函数，以评估多种方法来解释时间（即，昼夜和季节性）效应以及栖息地条件对选择的影响。我们首先表明，在步选择函数中考虑一天中的时间的方式对于获得具有生态意义的结果至关重要，并且包括时间的通常方式（即，基于时钟时间或基于夜晚与白天类别）可能会导致不完整或误导性的结论。此外，我们发现个体遵循昼夜节律或季节性栖息地选择模式，这取决于他们居住的地点。这可以通过两个站点之间干扰的差异来解释。除冬季外，无论人类活动水平如何，夜间选择开放栖息地的个体，但白天只选择来自更扰乱地点的个体作为覆盖栖息地。我们的结果强调了仔细考虑适当的时间尺度进行栖息地选择分析的重要性。我们的研究结果还表明，马鹿本身并不是一种黄昏或夜间活动的物种，正如在人类主导的景观中经常观察到的那样。相反，我们的结果意味着马鹿的夜间活动是适应人类活动的结果。