Role reversal in the stand dynamics of an angiosperm-conifer forest: Colonising angiosperms precede a shade-tolerant conifer in Afrotemperate forest

In mixed angiosperm-conifer forests worldwide, infrequent landscape-level catastrophic disturbances create a mosaic of persistent and different aged forest stands in the landscape with varying levels of dominance by the conifer component. In the 'temporal stand replacement model' (TSRM), disturbance creates conditions favouring a colonising cohort that is replaced by a suite of relatively shade-tolerant canopy species, which establish following the synchronous senescence of the pioneer canopy. In most southern hemisphere mixed angiosperm-conifer forests, with the exception of those in southern Africa, the establishing cohort is usually a large and very long-lived (550-650 years) conifer that is gradually replaced by angiosperms. As an explanation of the apparent dominance of the conifer Podocarpus latifolius, we examine the efficacy of the TSRM in mixed Afrotemperate forests where the establishing cohort is not a conifer. Forest succession in Afrotemperate forests was deterministic with the successive replacement of species determined first by their establishment success in shaded environments, and second, by their relative longevity. Several angiosperm species that were common canopy dominants established a pioneer cohort but were gradually replaced by P. latifolius, a shade-tolerant species. Continuous regeneration beneath the angiosperm canopy by A latifolius eliminates synchronous canopy senescence, a key feature of the TSRM, as a mechanism driving the temporal replacement of canopy species. Senescing angiosperms created canopy gaps that were colonised by grasses and ferns, which suppressed canopy tree regeneration. In contrast, with continuous regeneration beneath the shaded canopy, P. latifolius gains a critical advantage over angiosperms at gap formation. Thus, in the absence of fairly large-scale natural disturbances, conifers come to dominate Afrotemperate forests. Commensurate with the latter, conifers in Podocarpus-forest were dated to approximately 320 years, more than 100 years older than the oldest P. latifolius in angiosperm-dominated forest. Tree life-history differences (shade tolerance, longevity) and the time since disturbance drive successional change from an angiosperm-dominated system to a stage dominated by P. latifolius. in general, the TSRM is a plausible explanation for the observed canopy tree structure and dynamics in mixed Afrotemperate forests. South African Afrotemperate forest is unusual among other southern hemisphere mixed angiosperm-conifer forests in that a suite of angiosperm canopy species, rather than a single conifer species, forms the colonising cohort. (C) 2009 Elsevier B.V. All rights reserved.