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As the only species exceeding 90 m in height and 2000 years of age, Sequoia sempervirens and Sequoiadendron giganteum provide the optimal platform upon which to examine interactions among tree structure, age, and growth. We climbed 140 trees in oldgrowth redwood forests across California, USA, spanning a broad range of sizes and including the tallest, largest, and oldest known living individuals (i.e., 115.86 vs. 96.29 m tall, 424 vs. 582 Mg aboveground dry mass, and 2510 vs. 3240 years old for Sequoia and Sequoiadendron, respectively). We used a combination of direct measurements, hierarchical sampling, and dendrochronology to quantify tree structure and annual growth increments through old age. We also developed equations to predict aboveground attributes of standing redwoods via ground-based measurements. Compared to Sequoia, Sequoiadendron develops thicker bark on lower trunks, provisions leaves with more sapwood, and delays heartwood production throughout the crown. Main trunk wood volume growth (up to 1.6 vs. 0.9 m3/yr), aboveground biomass growth (up to 0.77 vs. 0.45 Mg/yr), and aboveground growth efficiency (0.55 ± 0.04 vs. 0.22 ± 0.01 kg annual growth per kg leaves, mean ± SE) are all higher in Sequoia. Two independent dimensions of structure-size and aboveground vigor-are the strongest predictors of tree-level productivity in both species. A third dimension, relative trunk size, is a significant predictor of growth in Sequoia such that trees with relatively large main trunks compared to their crowns produce more wood annually. Similar-size trees grow at similar rates regardless of latitude or elevation in tall forests of each species. Recent annual growth increments are higher than in the past for the majority of trees, and old trees are just as responsive to environmental changes as young trees. Negative growth-age relationships in previous centuries and positive growth-age relationships in recent decades reflect sampling bias and shifting disturbance regimes. Overall, we find little (if any) evidence for negative effects of old age on tree-level productivity in either species. Except for recovery periods following temporary reductions in crown size, annual increments of wood volume and biomass growth increase as redwoods enlarge with age until extrinsic forces cause tree death. [ABSTRACT FROM AUTHOR]

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens) and giant sequoia ( Sequoiadendron giganteum) trees of different sizes. Leaf-specific hydraulic conductivity ( kL) increased with height in S. sempervirens but not in S. giganteum, while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ13C) increased, and maximum mass-based stomatal conductance ( gmass) and photosynthesis ( Amass) decreased with height in both species. As a result, both Amass and gmass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum. In addition, Amass and gmass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO2 conductance ( gi). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat. [ABSTRACT FROM AUTHOR]

Sequoia sempervirens (redwood) is a long-lived, shade-tolerant tree capable of regeneration without disturbances and thus often present in all sizes within a single forest. In order to evaluate functional linkages among structures, plant distribution, and biodiversity in the canopy, we quantified all vascular plants from ground level to the treetops in an old-growth redwood forest (Prairie Creek Redwoods State Park, California, USA). This involved mapping terrestrial and epiphytic trees, shrubs, and ferns as well as climbing 27 trees up to 101 m tall within a 1-ha plot. We monitored canopy microclimates using sensor arrays that collected hourly data for up to 30 months. The plot held 4283 Mg/ha of aboveground dry mass in living plants, 95.4% of which was contributed by redwood. A high degree of structural complexity and individuality was evident in the crowns of the 14 largest trees in the form of reiterated trunks arising from main trunks, other trunks, and limbs. Thirteen species of vascular plants occurred as epiphytes in the plot, and all but one of these were restricted to the 14 largest trees. The evergreen shrub Vaccinium ovatum had the highest biomass of any epiphyte (212 kg/ha) followed by the evergreen fern Polypodium scouleri (196 kg/ha). The spatial aggregation of reiterated trunks was highly coincident with epiphyte masses, explaining 92% of the variation in fern and 75% of the variation in woody plant distribution along the vertical gradient. In addition to epiphyte biomass, there were 2366 kg/ha of soil in the canopy. Soils in crotches held 2-4 times as much water as soils on branches or limbs, and deeper layers held more water than shallower layers. During two years of monitoring, the volume of water stored in dead wood and soils in the canopy fluctuated between 24.7 m³ and 53.4 m³/ha. The vast majority of the plot's structure and arboreal habitats was held in the crowns of the 14 largest trees. Reiterated trunks and limbs on a small number of trees have important ecological functions in old-growth redwood forests, and it may be feasible to accelerate the development of structural complexity and biodiversity in maturing redwood stands. [ABSTRACT FROM AUTHOR]

Trees grow tall where resources are abundant, stresses are minor, and competition for light places a premium on height growth. The height to which trees can grow and the biophysical determinants of maximum height are poorly understood. Some models predict heights of up to 120?m in the absence of mechanical damage, but there are historical accounts of taller trees. Current hypotheses of height limitation focus on increasing water transport constraints in taller trees and the resulting reductions in leaf photosynthesis. We studied redwoods (Sequoia sempervirens), including the tallest known tree on Earth (112.7?m), in wet temperate forests of northern California. Our regression analyses of height gradients in leaf functional characteristics estimate a maximum tree height of 122-130?m barring mechanical damage, similar to the tallest recorded trees of the past. As trees grow taller, increasing leaf water stress due to gravity and path length resistance may ultimately limit leaf expansion and photosynthesis for further height growth, even with ample soil moisture. [ABSTRACT FROM AUTHOR]