From left to right all through the night: Characteristics of lying rest in zoo elephants.

Despite increased research during the past years, many characteristics of resting behavior in elephants are still unknown. For example, there is only limited data suggesting elephants express longer lying bouts and increased total nightly lying durations on soft substrates as compared to hard surfaces. Additionally, it has not been investigated how frequently elephants change body sides between lying bouts. Here we present these characteristics based on observations of nighttime lying behavior in 10 zoo elephants (5 African Loxodonta africana and 5 Asian Elephas maximus elephants) living in five different European facilities. We found that elephants housed on soft substrates have significantly increased total lying durations per night and longer average lying bouts. Furthermore, at 70%-85% of all bouts, a consistently higher frequency of side change between lying bouts occurred on soft substrates, leading to an overall equal laterality in resting behavior. Deviations from this pattern became evident in elephants living on nonsand flooring or/and in nondominant individuals of nonfamily groups, respectively. Based on our findings, we consider elephants to normally have several lying bouts per night with frequent side changes, given an appropriate substrate and healthy social environment. We encourage elephant-keeping facilities to monitor these characteristics in their elephants' nighttime behavior to determine opportunities for further improvements and detect alterations putatively indicating social or health problems in individual elephants at an early stage.