Contents lists available at ScienceDirect Ecological Complexity journal homepage: www.elsevier.com/locate/ecocom Patterns of landscape seasonality influence passerine diversity: Implications for conservation management under global change Emilio Civantos a, ⁎ , Antonio T. Monteiro a , João Gonçalves a,b , Bruno Marcos a,b , Paulo Alves a , João P. Honrado a,b a CIBIO/InBIO. Research Center in Biodiversity and Genetic Resources, University of Porto, Campus Agrário de Vairão, 4485-661 Vairão, Portugal b Faculdade de Ciências, Universidade do Porto, Rua do Campo Alegre FC4, 4169-007 Porto, Portugal ARTICLE INFO Keywords: Landscape energy Landscape heterogeneity Remote sensing Species richness Land abandonment ABSTRACT The importance of environmental heterogeneity for biodiversity across scales is widely recognized in ecological theory and profusely supported by evidence. However, our understanding of the effects of spatiotemporal pat- terns of landscape functional properties on biodiversity is still rather limited. We examined the relationship between common passerine species richness and ecosystem functioning dynamics, namely seasonality, measured by satellite remote sensing. We focused on rural landscapes of a mountain National Park in Portugal undergoing rapid reshaping from agro-pastoral mosaics to early successional landscapes. We applied multi-model inference to compare the hypothesis of landscape seasonality as a driver of species richness with three competing hy- potheses representing structural habitat heterogeneity, disturbance, and availability of food resources. We found support for landscape seasonality and its spatial heterogeneity in explaining passerine richness in mountain rural landscapes. Conversely, no significant support of the remaining hypotheses was found. These results highlight the role of ecosystem functioning variability in space and time. They also stress the importance of considering species-energy relationships for conservation at the landscape level. Specifically, they provide support and guidance to the identification of meaningful functional attributes of the landscape that shape its biodiversity. Our results further demonstrate the utility of remote sensing approaches and products to measure those attri- butes and follow their trends through time. Spatially-explicit measures of energy variability, such as the func- tional amplitude between winter and summer retrieved from earth observations, can link global socio-en- vironmental change to species’ responses and support the inclusion of landscape seasonality on conservation and monitoring frameworks. 1. Introduction Understanding the relationship between biological diversity and landscape patterns and dynamics is crucial to face ongoing environ- mental change. Mediterranean landscapes of Southern Europe, shaped by human management through a combination of fire, husbandry and agriculture, sustain diverse mosaics that promote the local increase of species richness (Grove and Rackham, 2001). These landscapes typi- cally include patches of relatively natural habitat in a mosaic of human land uses. When compared to more uniform environments, these het- erogeneous landscapes allow more species to coexist locally. In fact, both species richness and diversity of bird communities were shown to be higher in heterogeneous landscapes (Benton et al., 2003; Smith et al., 2010). The importance of this heterogeneity for biodiversity across scales is widely recognized in ecological theory and supported by abundant evidence (reviewed in (Rosenzweig, 1995; Statzner and Moss, 2004). The positive relationship between biodiversity and habitat hetero- geneity at landscape scales is generally accepted, particularly in agri- cultural landscapes (Benton et al., 2003; Fahrig et al., 2011). The re- lationship between structural habitat heterogeneity and species diversity is a well-documented pattern in landscape ecology (Tews et al., 2004). Moreover, structural modifications in habitat mo- saics are known to develop alongside with important changes in func- tional attributes of ecosystems, such as energy balance, which may ultimately affect biodiversity (Hurlbert, 2004). Therefore, in addition to structural heterogeneity, energy variations in space and time may also influence species diversity in heterogeneous mosaics, especially in dy- namic landscapes submitted to driving forces such as land use change (Evans et al., 2005; Hurlbert and Haskell, 2003). https://doi.org/10.1016/j.ecocom.2018.07.001 Received 4 January 2018; Received in revised form 1 June 2018; Accepted 7 July 2018 ⁎ Corresponding author. E-mail address: ecivantos@cibio.up.pt (E. Civantos). Ecological Complexity 36 (2018) 117–125 Available online 24 August 2018 1476-945X/ © 2018 Elsevier B.V. All rights reserved. T