7Species distributions are determined by the interaction of multiple biotic and 8 abiotic factors, which produces complex spatial and temporal patterns of occurrence. 9 As habitats and climate change due to anthropogenic activities, there is a need to 10 develop species distribution models that can quantify these complex range dynamics.
11In this paper, we develop a dynamic occupancy model that uses a spatial generalized 12 additive model to estimate non-linear spatial variation in occupancy not accounted 13 for by environmental covariates. The model is flexible and can accommodate data 14 from a range of sampling designs that provide information about both occupancy and 15 detection probability. Output from the model can be used to create distribution maps 16 and to estimate indices of temporal range dynamics. We demonstrate the utility of 17 this approach by modeling long-term range dynamics of 10 eastern North American 18 birds using data from the North American Breeding Bird Survey. We anticipate this 19 framework will be particularly useful for modeling species' distributions over large 20 spatial scales and for quantifying range dynamics over long temporal scales. 21 Introduction 24 The distribution of each species is determined by the interaction of multiple biotic and abiotic 25 factors that vary across both space and time, including weather and climate (Barbet-Massin 26 and Jetz 2014), habitat availability (Hill et al. 1999), physiological tolerances (Kearney and 27 Porter 2009), and biotic interactions (Araújo and Luoto 2007). As a result, most species' 28 distributions are characterized by complex spatial and temporal patterns of occurrence, which 29 combined with the large scales over which distributions change, present challenges for both 30the collection and analysis of data to quantify range dynamics (Elith et al. 2010). As habitats 31 and climate change due to anthropogenic activities, there is an increasingly urgent need 32 to develop species distribution models (SDMs) that can accurately and efficiently quantify 33 complex range dynamics over large spatial and long temporal scales. 34 In response to this need, researchers have developed a range of SDM approaches that vary 35 in their data requirements and analytical methods (Elith et al. 2010, Guillera-Arroita 36 2017). Although each of these methods has strengths and weaknesses, SDMs designed to 37 quantify range dynamics require several key features. First, SDMs must include sufficient 38 flexibility to quantify highly non-linear spatial patterns in occurrence probability. Although 39 spatial variation in occurrence can in some cases be modeled using environmental covariates, 40 residual spatial variation (which is likely common in most applications of SDMs at large 41 spatial scales) can bias estimates of occurrence probability (Johnson et al. 2013, Guélat 42 and Kéry 2018). Second, because occurrence probability at a given point in time is not 43 independent of occurrence probability at earlier points in time, SDMs must e...