“…A metapopulation model can be used to estimate the geographical range expansion when only the occurrence or absence of the alien organism in a region needs to be considered (e.g., Colizza et al, 2007;Koike, 2006). Although metapopulation models are relatively simple, they still require quantitative pathway data, such as the traffic density between sites or the length of the paths through continuous habitats (Colizza et al, 2007;Koike & Iwasaki, 2011). However, as in the cases of the H5N1 influenza epidemic and the introduced red fire ant, these traffic pathways are sometimes complicated owing to factors such as natural waterfowl migration and poultry trading, as with H5N1 (Vandegrift, Sokolow, Daszak, & Kilpatrick, 2010), or unidentified modes of conveyance, as with the fire ant (Ascunce et al, 2011), even though world trading is one of the most important factors (Chapman, Purse, Roy, & Bullock, 2017).…”