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Intra-sexual segregation is a form of social segregation widespread among vertebrates. In the bat Myotis daubentonii, males are disproportionately abundant at higher elevations, while females are restricted to lower altitude. Intra-male segregation is also known to occur yet its ecological and behavioural determinants are unclear. We studied male segregation along a river in Central Italy where we tested the following predictions: 1. Upstream ( > 1000 m a.s.l.) males will rely on scarcer prey; 2. To deal with this limitation and exploit a cooler roosting environment, they will employ more prolonged and deeper torpor than downstream (< 900 m a.s.l.) males; 3. Body condition will be better in downstream males as they forage in more productive areas; 4. To cope with less predictable foraging opportunities, upstream males will use more habitat types. Consistent with our predictions, we found that prey were less common at higher altitudes, where bats exhibited prolonged and deeper torpor. Body condition was better in downstream males than in upstream males but not in all summer months. This result reflected a decrease in downstream males’ body condition over the season, perhaps due to the energy costs of reduced opportunities to use torpor and/or intraspecific competition. Downstream males mainly foraged over selected riparian vegetation whereas upstream males used a greater variety of habitats. One controversial issue is whether upstream males are excluded from lower elevations by resident bats. We tested this by translocating 10 upstream males to a downstream roost: eight returned to the high elevation site in 1-2 nights, two persisted at low altitude but did not roost with resident bats. These results are consistent with the idea of segregation due to competition. Living at high altitude allows for more effective heterothermy and may thus be not detrimental for survival, but by staying at lower altitude males increase proximity to females and potentially benefit from summer mating opportunities.

Bats are primarily forest mammals and forest structure may affect their communities through the level of vegetation clutter. Pine plantations are typically even-aged managed forests that lack structural complexity. However, an understory layer can enhance the heterogeneity of these forests, making them suitable for several animal taxa. We hypothesized that species composition, richness, and foraging activity of insectivorous bats in pine plantations vary according to forest structure, specifically with the density of the understory. We measured pine density, Diameter at Breast Height (DBH), canopy closure, and vegetation cover of 29 pine (Pinus halepensis) plantations of the Judean Lowlands, Israel, and collected acoustic data on resident bats. We found that bat species richness and total activity increased in forests with large tree DBH and dense shrubs. Cluttered-habitat species foraged preferentially in forests with large tree DBH and high pine density, while open-habitat species preferred forests with well-developed canopies and dense shrubs. Pipistrellus pipistrellus and Eptesicus serotinus foraged in mature forests with well-developed bushes and these species are endangered in Israel. We conclude that mature planted pine forests with a well-developed under-canopy are suitable foraging grounds for insectivorous bats. Management plans for planted pine forests should consider our findings to support bat populations, including rare and endangered species.