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Aim A major question with regard to the ecology of temperate rain forests in south-central Chile is how pioneer and shade-tolerant tree species coexist in old-growth forests. We explored the correspondence between tree regeneration dynamics and life-history traits to explain the coexistence of these two functional types in stands apparently representing a non-equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600-m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree-fall gaps. We used tree-ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long-lived pioneer Eucryphia cordifolia and the shade-tolerant Aextoxicon punctatum. Shade-tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi-modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree-ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade-tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand-disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree-fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long-term coexistence of pioneer and shade-tolerant tree species in this coastal, old-growth rain forest, despite the rarity of major disturbances. [ABSTRACT FROM AUTHOR]

1 Few studies have addressed the mechanisms of coexistence of shade-tolerant and intolerant tree species in the canopy of old-growth, lowland rain forests of southern South America. We explored the hypothesis that their forest dynamics result from frequent, single tree-fall gap episodes. 2 We analysed the disturbance regime and assessed the regeneration modes of shade-tolerant and intolerant canopy trees in a lowland, old growth North Patagonian rain forest in Chiloé Island (42° S) using dendroecological methods. 3 Dominant canopy trees were the shade-intolerant Nothofagus nitida (Fagaceae), Drimys winteri (Winteraceae) and the tolerant Podocarpus nubigena (Podocarpaceae). The oldest individuals, however, were represented by Saxegothaea conspicua, Podocarpaceae (shade tolerant > 498 years) and Weinmannia trichosperma, Cunoniaceae (intolerant > 382 years). Shade-tolerant species have regenerated continuously for the past 400 years, but recruitment of shade-intolerant species has increased only in the past 50 years. These regeneration patterns suggest a lack of catastrophic disturbance, at least since 1650. 4 Analysis of tree radial growth patterns revealed frequent moderate releases of both shade-tolerant and intolerant tree species, consistent with frequent small-scale disturbances, such as individual tree-fall gaps. Increased releases, peaking in 1940 and followed by enhanced recruitment, may be a consequence of individual tree-falls induced by low-intensity windstorms or limited selective logging. Such disturbances have occurred for at least 250 years, but have had little overall effect on stand structure and canopy composition. 5 The replacement and coexistence of shade-tolerant and intolerant tree species in the canopy of this North Patagonian forest can be explained by frequent small-scale disturbances (i.e. gap-phase dynamics) and by the absence of large-scale natural or anthropogenic disturbances over the past four centuries. Journal of Ecology (2004) 92, 598–608 [ABSTRACT FROM AUTHOR]