“…Spatial Markov models (SMMs; Bolster & Dentz, 2012;Kang et al, 2014;Sund et al, 2016) are a family of models that represent memory of speed and direction through the use of a transition matrix. While these models have been applied to a broad range of systems (e.g., Bolster et al, 2014;Kang et al, 2014Kang et al, , 2015aKang et al, , 2016Le Borgne et al, 2008b, 2008aSund et al, 2015bSund et al, , 2015aSund et al, , 2017bSund et al, , 2017a, they typically rely on parameterizing a transition matrix, which can be difficult to do, although approaches applicable to real data have emerged recently (Kang et al, 2015a;Sherman et al, 2017). Another approach has involved sampling of particle trajectories (Sund et al, 2017b), which in turn can be used for mixed upscaling and downscaling models.…”