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Abstract: The northern treeline is generally limited by available warmth. However, in recent years, more and more studies have identified drought stress as an additional limiting factor for tree growth in northern boreal forests and at treelines. Three growth responses to warming have been identified: increase in growth, decrease in growth, and nonsignificant correlation of tree growth with climate. Here we investigate the effect of drought stress on radial growth of white spruce at northern treelines along a longitudinal gradient spanning the entire Brooks Range in Alaska. We systematically sampled 687 white spruce at seven treeline sites. Where possible, we sampled three site types at a given site: high-density forest, low-density forest, and floodplain forest. We investigated the relationship of site and site type to tree growth responses. In the western part of our study area, we found very high numbers of trees responding with increase in growth to recent warming; while in the eastern part, trees responding with decrease in growth to recent warming are predominant. Within a given site, more trees reacting positively to warming grow on site types characterized by low tree density. These patterns coincide with precipitation decreases from west to east and local water availability gradients, therefore pointing to drought stress as the controlling factor for the distribution of trees responding with increase or decrease in growth to recent warming. Compared to 20th century climate, we project a 25–50% basal area (BA) increase in the western region for the 21st century due to climate warming as projected by five general circulation models, 4–11% in the central region and decreases (+1 to −11%) in the eastern region. The overall net change in projected 21st century BA increase at each site seems to be controlled by the relative proportion of responder groups. If these are similar, differences in the magnitude of increase versus decrease in growth control BA projections for that site. This study highlights the importance of regional-scale investigations of biosphere–climate interactions, since our results indicate a substantial gain in aboveground biomass as a result of future warming only in the western regions; while in the eastern regions, climate warming will decrease overall wood production and therefore carbon uptake potential. [Copyright &y& Elsevier]

Northern and high-latitude alpine treelines are generally thought to be limited by available warmth. Most studies of tree-growth–climate interaction at treeline as well as climate reconstructions using dendrochronology report positive growth response of treeline trees to warmer temperatures. However, population-wide responses of treeline trees to climate remain largely unexamined. We systematically sampled 1558 white spruce at 13 treeline sites in the Brooks Range and Alaska Range. Our findings of both positive and negative growth responses to climate warming at treeline challenge the widespread assumption that arctic treeline trees grow better with warming climate. High mean temperatures in July decreased the growth of 40% of white spruce at treeline areas in Alaska, whereas warm springs enhance growth of additional 36% of trees and 24% show no significant correlation with climate. Even though these opposing growth responses are present in all sampled sites, their relative proportion varies between sites and there is no overall clear relationship between growth response and landscape position within a site. Growth increases and decreases appear in our sample above specific temperature index values (temperature thresholds), which occurred more frequently in the late 20th century. Contrary to previous findings, temperature explained more variability in radial growth after 1950. Without accounting for these opposite responses and temperature thresholds, climate reconstructions based on ring width will miscalibrate past climate, and biogeochemical and dynamic vegetation models will overestimate carbon uptake and treeline advance under future warming scenarios. [ABSTRACT FROM AUTHOR]

Needle hardiness of introduced yellow pine, Pinus banksiana Lamb., lodgepole pine, P. contorta Dougl. and native white spruce, Picea glauca (Moench) Voss, were assessed by the effective prefreezing temperature method. Yellow pine needles were less hardy than lodgepole pine or white spruce needles in Alaska on each date measured. Although hardiness decreased in springtime in all species, decreases in hardiness in yellow pine began before temperatures were above -20°C, apparently in response to day length, while decreases in hardiness in lodgepole pine and white spruce began only when mean temperatures were above 0°C. Hardiness was increased by decreasing the water content of yellow pine and spruce needles. However, only the latter increased its field hardiness by decreased water contents, and only to a small degree. Large decreases in phospholipid occurred during the dehardening period, indicating the presence of major membrane-associated changes. However, changes in hardiness did not closely parallel those in phospholipid; hardiness decreased before phospholipid did in spruce and after phospholipid did in lodgepole pine. In yellow pine, changes in hardiness were more closely related to changes in phospholipid content. Decreases in phospholipid appeared to be correlated with the day length in all species. [ABSTRACT FROM AUTHOR]