Shoot dimorphism enables Sequoia sempervirens to separate requirements for foliar water uptake and photosynthesis.

PREMISE: Trees in wet forests often have features that prevent water films from covering stomata and inhibiting gas exchange, while many trees in drier environments use foliar water uptake to reduce water stress. In forests with both wet and dry seasons, evergreen trees would benefit from producing leaves capable of balancing rainy-season photosynthesis with summertime water absorption. METHODS: Using samples collected from across the vertical gradient in tall redwood (Sequoia sempervirens) crowns, we estimated tree-level foliar water uptake and employed physics-based causative modeling to identify key functional traits that determine uptake potential by setting hydraulic resistance. RESULTS: We showed that Sequoia has two functionally distinct shoot morphotypes. While most shoots specialize in photosynthesis, the axial shoot type is capable of much greater foliar water uptake, and its within-crown distribution varies with latitude. A suite of leaf surface traits cause hydraulic resistance, leading to variation in uptake capacity among samples. CONCLUSIONS: Shoot dimorphism gives tall Sequoia trees the capacity to absorb up to 48 kg H2 O h-1 during the first hour of leaf wetting, ameliorating water stress while presumably maintaining high photosynthetic capacity year round. Geographic variation in shoot dimorphism suggests that plasticity in shoot-type distribution and leaf surface traits helps Sequoia maintain a dominate presence in both wet and dry forests.

Millennium-scale crossdating and inter-annual climate sensitivities of standing California redwoods.

Extremely decay-resistant wood and fire-resistant bark allow California's redwoods to accumulate millennia of annual growth rings that can be useful in biological research. Whereas tree rings of Sequoiadendron giganteum (SEGI) helped formalize the study of dendrochronology and the principle of crossdating, those of Sequoia sempervirens (SESE) have proven much more difficult to decipher, greatly limiting dendroclimatic and other investigations of this species. We overcame these problems by climbing standing trees and coring trunks at multiple heights in 14 old-growth forest locations across California. Overall, we sampled 1,466 series with 483,712 annual rings from 120 trees and were able to crossdate 83% of SESE compared to 99% of SEGI rings. Standard and residual tree-ring chronologies spanning up to 1,685 years for SESE and 1,538 years for SEGI were created for each location to evaluate crossdating and to examine correlations between annual growth and climate. We used monthly values of temperature, precipitation, and drought severity as well as summer cloudiness to quantify potential drivers of inter-annual growth variation over century-long time series at each location. SESE chronologies exhibited a latitudinal gradient of climate sensitivities, contrasting cooler northern rainforests and warmer, drier southern forests. Radial growth increased with decreasing summer cloudiness in northern rainforests and a central SESE location. The strongest dendroclimatic relationship occurred in our southernmost SESE location, where radial growth correlated negatively with dry summer conditions and exhibited responses to historic fires. SEGI chronologies showed negative correlations with June temperature and positive correlations with previous October precipitation. More work is needed to understand quantitative relationships between SEGI radial growth and moisture availability, particularly snowmelt. Tree-ring chronologies developed here for both redwood species have numerous scientific applications, including determination of tree ages, accurate dating of fire-return intervals, archaeology, analyses of stable isotopes, long-term climate reconstructions, and quantifying rates of carbon sequestration.

Structural development of redwood branches and its effects on wood growth.

Redwood branches provide all the carbohydrates for the most carbon-heavy forests on Earth, and recent whole-tree measurements have quantified trunk growth rates associated with complete branch inventories. Providing all of a tree's photosynthetic capacity, branches represent an increasing proportion of total aboveground wood production as trees enlarge. To examine branch development and its effects on wood volume growth, we dissected 31 branches from eight Sequoia sempervirens (D. Don) Endl. and seven Sequoiadendron giganteum Lindl. trees. The cambium-area-to-leaf-area ratio was maintained with size and age but increased with light availability, whereas the heartwood-deposition-area-to-leaf-area ratio increased with size and age but was insensitive to light availability. The proportion of foliage mass arrayed in <1-cm-diameter epicormic shoots increased with decreasing light and was higher in Sequoia (20-60%) than in Sequoiadendron (3-16%). Well-illuminated branches concentrated leaves higher and distally, while shaded branches distributed leaves lower and proximally. In similar light environments, older branches distributed leaves lower and more proximally than younger branches. Branch size, light, species, heartwood area, a heartwood-area-species interaction, and ovulate cone mass predicted 87.5% of the variability in wood volume growth of branches. After accounting for the positive effects of size and light, wood volume growth declined with heartwood area and age. The effect of age was trivial compared to the effect of heartwood area, suggesting that heartwood expansion caused the age-related decline in wood volume growth. Additionally, Sequoiadendron branches of similar size and light environment with more ovulate cones produced less wood, even though these cones were long-lived and photosynthetic, reflecting the energetic cost of seed production. These results contributed to a conceptual model of branch development in which light availability, injury, heartwood content, gravity, and time interact to produce the high degree of branch structural variation evident within redwood crowns.