“…Also unresolved is whether the species' niches are mostly realized in space over environmental gradients, or in time over forest succession, intended as a directional process of autogenic change in the species composition of the biotic community established after a stand-replacement disturbance. Bridge and Johnson (2000) explain the vegetation patterns of the boreal mixedwoods of Saskatchewan, Canada, in terms of moisture and nutrient availability, while Hély et al (2000) explain the variation in composition of southeastern Canadian boreal mixedwoods as a successional gradient. While the successional explanation is supported by species differences in shade tolerance (Messier et al, 1999), it is also well documented that mixedwood stands can originate immediately after stand-replacing disturbance (Thorpe, 1992;Youngblood, 1995;Hope, 2001;Peters et al, 2002), suggesting that succession only explains part of mixedwood diversity (Gutsell and Johnson, 2002).…”