“…These studies argued that ecosystem development was a function of time since deglaciation, which was theorized via a space-for-time substitution (Matthews, 1992) as a chronosequence. Matthews (1992) proposed a more developed conceptual model where ecological succession was not simply interpreted as a function of time but also (a) abiotic processes such as soil grain size, microrelief, microclimate, hydrology, slope, and geomorphic stability (Burga et al, 2010;Garibotti, Pissolito, & Villalba, 2011;Rydgren, Halvorsen, Töpper, & Njos, 2014;Wietrzyk, Wegrzyn, & Lisowska, 2016) and (b) biotic processes including species interaction and competition (Schumann, Gewolf, & Tackenberg, 2016). Initially, the spatial heterogeneity in abiotic processes is thought to be dominant, but as the terrain becomes more stable and ecosystem succession advances, biotic factors are thought to become more influential (Matthews, 1992).…”