“…Thinking about the implications across scales is important but is also challenging as the relationships are often complex, nonlinear and, therefore, un-intuitive. Previously, theoretical models of multi-scale phenomena have been reviewed (Mideo, Alizon & Day, 2008; Reiner et al, 2013; Dorratoltaj et al, 2017; Murillo, Murillo & Perelson, 2013; Severins, 2012). Repeated themes of these works and others over the past two decades have included: the need for more data (Alizon & Van Baalen, 2008; Alizon, Luciani & Regoes, 2011; Handel & Rohani, 2015; Lavine, Poss & Grenfell, 2008; Pollitt et al, 2011); the challenge of integrating scales (Frost et al, 2015; Perelson et al, 1996; Handel & Rohani, 2015; Mideo et al, 2013); and the role of heterogeneity (Lavine, Poss & Grenfell, 2008; VanderWaal & Ezenwa, 2016).…”