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N.L. Stephenson

Publications and source records attributed to N.L. Stephenson.

At least 19 recordsLinked to original sources

Rate of tree carbon accumulation increases continuously with tree size

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage - increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to understand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence.

Nature

The contribution of competition to tree mortality in old-growth coniferous forests

Competition is a well-documented contributor to tree mortality in temperate forests, with numerous studies documenting a relationship between tree death and the competitive environment. Models frequently rely on competition as the only non-random mechanism affecting tree mortality. However, for mature forests, competition may cease to be the primary driver of mortality. We use a large, long-term dataset to study the importance of competition in determining tree mortality in old-growth forests on the western slope of the Sierra Nevada of California, U.S.A. We make use of the comparative spatial configuration of dead and live trees, changes in tree spatial pattern through time, and field assessments of contributors to an individual tree's death to quantify competitive effects. Competition was apparently a significant contributor to tree mortality in these forests. Trees that died tended to be in more competitive environments than trees that survived, and suppression frequently appeared as a factor contributing to mortality. On the other hand, based on spatial pattern analyses, only three of 14 plots demonstrated compelling evidence that competition was dominating mortality. Most of the rest of the plots fell within the expectation for random mortality, and three fit neither the random nor the competition model. These results suggest that while competition is often playing a significant role in tree mortality processes in these forests it only infrequently governs those processes. In addition, the field assessments indicated a substantial presence of biotic mortality agents in trees that died. While competition is almost certainly important, demographics in these forests cannot accurately be characterized without a better grasp of other mortality processes. In particular, we likely need a better understanding of biotic agents and their interactions with one another and with competition.

California

Widespread increase of tree mortality rates in the Western United States

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.

Science

Apparent climatically induced increase of tree mortality rates in a temperate forest

We provide a first detailed analysis of long-term, annual-resolution demographic trends in a temperate forest. After tracking the fates of 21 338 trees in a network of old-growth forest plots in the Sierra Nevada of California, we found that mortality rate, but not the recruitment rate, increased significantly over the 22 years of measurement (1983-2004). Mortality rates increased in both of two dominant taxonomic groups (Abies and Pinus) and in different forest types (different elevational zones). The increase in overall mortality rate resulted from an increase in tree deaths attributed to stress and biotic causes, and coincided with a temperature-driven increase in an index of drought. Our findings suggest that these forests (and by implication, other water-limited forests) may be sensitive to temperature-driven drought stress, and may be poised for die-back if future climates continue to feature rising temperatures without compensating increases in precipitation. ?? 2007 Blackwell Publishing Ltd/CNRS.

Ecology Letters

The accuracy of matrix population model projections for coniferous trees in the Sierra Nevada, California

1 We assess the use of simple, size-based matrix population models for projecting population trends for six coniferous tree species in the Sierra Nevada, California. We used demographic data from 16 673 trees in 15 permanent plots to create 17 separate time-invariant, density-independent population projection models, and determined differences between trends projected from initial surveys with a 5-year interval and observed data during two subsequent 5-year time steps. 2 We detected departures from the assumptions of the matrix modelling approach in terms of strong growth autocorrelations. We also found evidence of observation errors for measurements of tree growth and, to a more limited degree, recruitment. Loglinear analysis provided evidence of significant temporal variation in demographic rates for only two of the 17 populations. 3 Total population sizes were strongly predicted by model projections, although population dynamics were dominated by carryover from the previous 5-year time step (i.e. there were few cases of recruitment or death). Fractional changes to overall population sizes were less well predicted. Compared with a null model and a simple demographic model lacking size structure, matrix model projections were better able to predict total population sizes, although the differences were not statistically significant. Matrix model projections were also able to predict short-term rates of survival, growth and recruitment. Mortality frequencies were not well predicted. 4 Our results suggest that simple size-structured models can accurately project future short-term changes for some tree populations. However, not all populations were well predicted and these simple models would probably become more inaccurate over longer projection intervals. The predictive ability of these models would also be limited by disturbance or other events that destabilize demographic rates. ?? 2005 British Ecological Society.

Journal of Ecology

Forest turnover rates follow global and regional patterns of productivity

Using a global database, we found that forest turnover rates (the average of tree mortality and recruitment rates) parallel broad-scale patterns of net primary productivity. First, forest turnover was higher in tropical than in temperate forests. Second, as recently demonstrated by others, Amazonian forest turnover was higher on fertile than infertile soils. Third, within temperate latitudes, turnover was highest in angiosperm forests, intermediate in mixed forests, and lowest in gymnosperm forests. Finally, within a single forest physiognomic type, turnover declined sharply with elevation (hence with temperature). These patterns of turnover in populations of trees are broadly similar to the patterns of turnover in populations of plant organs (leaves and roots) found in other studies. Our findings suggest a link between forest mass balance and the population dynamics of trees, and have implications for understanding and predicting the effects of environmental changes on forest structure and terrestrial carbon dynamics.

Ecology Letters

Does coring contribute to tree mortality?

We assess the potential of increment coring, a common method for measuring tree ages and growth, to contribute to mortality. We used up to 21 years of annual censuses from two cored and two uncored permanent plots in the Sierra Nevada of California, to detect changes in mortality rates 12 years following coring for individuals >5 cm DBH from two coniferous species, Abies concolor (Gordon & Glend.) Lindl. (white fir) and Abies magnifica A. Murr. (red fir). Using a randomized before-after control impact (BACI) design, we found no differences in mortality rates following coring for 825 cored and 525 uncored A. concolor and 104 cored and 66 uncored A. magnifica . These results support the view that collecting tree cores can be considered nondestructive sampling, but we emphasize that our 12-year postcoring records are short compared with the maximum life-span of these trees and that other species in different environments may prove to be more sensitive to coring.

California

Actual evapotranspiration and deficit: Biologically meaningful correlates of vegetation distribution across spatial scales

Correlative approaches to understanding the climatic controls of vegetation distribution have exhibited at least two important weaknesses: they have been conceptually divorced across spatial scales, and their climatic parameters have not necessarily represented aspects of climate of broad physiological importance to plants. Using examples from the literature and from the Sierra Nevada of California, I argue that two water balance parameters—actual evapotranspiration (AET) and deficit (D)—are biologically meaningful, are well correlated with the distribution of vegetation types, and exhibit these qualities over several orders of magnitude of spatial scale (continental to local). I reach four additional conclusions. (1) Some pairs of climatic parameters presently in use are functionally similar to AET and D; however, AET and D may be easier to interpret biologically. (2) Several well-known climatic parameters are biologically less meaningful or less important than AET and D, and consequently are poorer correlates of the distribution of vegetation types. Of particular interest, AET is a much better correlate of the distributions of coniferous and deciduous forests than minimum temperature. (3) The effects of evaporative demand and water availability on a site's water balance are intrinsically different. For example, the ‘dry’ experienced by plants on sunward slopes (high evaporative demand) is not comparable to the ‘dry’ experienced by plants on soils with low water-holding capacities (low water availability), and these differences are reflected in vegetation patterns. (4) Many traditional topographic moisture scalars—those that additively combine measures related to evaporative demand and water availability—are not necessarily meaningful for describing site conditions as sensed by plants; the same holds for measured soil moisture. However, using AET and D in place of moisture scalars and measured soil moisture can solve these problems.

Journal of Biogeography

Estimated ages of some large giant sequoias: General Sherman keeps getting younger

During 1975-77, 168 bald eagles (Haliaeetus leucocephalus) found moribund or dead in 29 states were analyzed for organochlorine pesticides and polychlorinated biphenyls (PCBs); 32 specimens from 13 states were analyzed for polybrominated biphenyls (PBBs). PCBs were present in 166 bald eagle carcasses and DDE was found in 165. TDE and dieldrin were identified in 137 samples, trans-nonachlor in 118, and oxychlordane in 90. Brains of five eagles contained possible lethal levels of dieldrin, and two eagles possibly died of endrin poisoning. Nine eagle livers, analyzed because of suspected lead poisoning, contained high levels of lead. Twenty percent of the eagles died from shooting, the most common cause of death; this cause of death, however, has declined.

Madroño