Year-round multi-scale habitat selection by Crested Tit (Lophophanes cristatus) in lowland mixed forests (northern Italy)

Determining how animals respond to resource availability across spatial and temporal extents is crucial to un- derstand ecological processes underpinning habitat selection. Here, we used a multi-scale approach to study the year-round habitat selection of the Crested Tit (Lophophanes cristatus) in a semi-natural lowland woodland of northern Italy, analysing different habitat features at each scale. We performed Crested Tit censuses at three different spatial scales. At the macrohabitat scale, we used geolocalized observations of individuals to compute Manly's habitat selection index, based on a detailed land-use map of the study area. At the microhabitat scale, the trees features were compared between presence and absence locations. At the foraging habitat scale, individual foraging birds and their specific position on trees were recorded using focal animal sampling. Censuses were performed during both the breeding (March to May) and wintering (December to January) seasons. At the macrohabitat scale, the Crested Tits significantly selected pure and mixed pine forests and avoided woods of alien plant species, farmlands and urban areas. At the microhabitat scale, old pine woods with dense cover were selected, with no significant difference in the features of tree selection between the two phenological phases. At the foraging habitat scale, the species was observed spending more time foraging in the canopies than in the understorey, using mostly the portion of Scots Pine (Pinus sylvestris) canopies closer to the trunk in winter, while during the breeding period, the whole canopy was visited. Overall, breeding and wintering habitats largely overlapped in the Crested Tit. Based on our findings, lowland Crested Tits can be well defined as true habitat specialists: they are strictly related to some specific coniferous woodland features. Noteworthily, compared to other tit species, which normally show generalist habits during winter, the Crested Tit behaves as a habitat specialist also out of the breeding season. Our study stressed the importance of considering multi-scale (both spatial and phenological) habitat selection in birds.