Rangewide climatic sensitivities and non-timber values of tall Sequoia sempervirens forests

Heavily exploited for its reddish, decay-resistant heartwood, the tallest conifer, Sequoia sempervirens, is a major component of coastal forests from extreme southwestern Oregon to California's Santa Lucia Mountains. Primary Sequoia forests are now restricted to < 5 % of their former distribution, and mature secondary forests with trees over 60 m tall are even scarcer due to repeated logging. Leveraging allometric equations recently derived from intensive work in both forest types, we climbed, measured, and core-sampled 235 trees in 45 locations distributed across the species range to examine growth trends and understand how tall Sequoia are responding to recent environmental changes. Paired samples of sapwood and heartwood collected along the height gradient were used to quantify Sequoia investment in decay resistance. During the 20th century, trees in most locations began producing more wood than expected for their size with this growth surge becoming pronounced after 1970 and ending around 2000. Radial increments-ring widths-correlate with climatic variables related to water availability, and these relationships are strengthening as temperatures rise. Sensitivity to drought increased from north to south along a 6 degrees latitudinal gradient of decreasing precipitation and summer fog frequency. Sequoia trees north of 40 degrees were least sensitive to drought, producing similar biomass annually during dry and wet years, whereas trees farther south produced less biomass during individual drought years. Hotter 21st century drought barely affected Sequoia growth efficiency (biomass increment per unit leaf mass) north of 40 degrees until the fourth consecutive year (2015), when growth efficiency dropped precipitously, recovering within two years. South of 40 degrees, Sequoia trees exhibited steadily declining growth efficiency during the multi-year drought followed by recovery, but recovery did not occur south of 37 degrees despite ample precipitation in 2017. Sequoia growth efficiency is currently highest in secondary forests north of 40 degrees, where trees produce relatively small amounts of heartwood with the lowest decay resistance (least fungicide) while receiving the most nocturnal summer fog. Increasing sink limitations, whereby rising temperatures, drier air at night, and extreme tree height collectively lower turgor pressure to inhibit cambial activity, may reduce Sequoia growth efficiency while contributing to more durable biomass production. Heartwood and fungicide increments are higher in primary than secondary forests across the species range. Crown structural complexity promotes development of vascular epiphytes and arboreal soil habitats in Sequoia forests with sufficient moisture availability. These habitats are lacking in secondary forests and rare in primary forests south of 40 degrees. After logging, restoration of tall Sequoia forests can be achieved via silviculture that maximizes height increments during early stand development and then retains some dominant trees in perpetuity, allowing them to gain full stature, produce increasingly decay-resistant heartwood, and support significant arboreal biodiversity.