Comparing the influence of site quality, stand age, fire and climate on aboveground tree production in Siberian Scots pine forests.

Temporal patterns of stem and needle production and total aboveground net primary production (ANPP) were studied at the tree and stand level along four chronosequences of Siberian Scots pine (Pinus sylvestris L.) forests differing in site quality (poor lichen type or the more fertile Vaccinium type) and in frequency of surface fires (unburned, moderately burned (fire return interval of approximately 40 years), or heavily burned (fire return interval of approximately 25 years)). The maximum range of variability in aboveground production was quantified for: (1) possible long-term changes in site quality; (2) stand age; (3) non-stand-replacing, recurring surface fires; and (4) interannual climate variability. For (1) and (2), total ANPP was low in the lichen-type chronosequence, reached a maximum of 170 g C m(-2) year(-1) after 100 years and decreased to 100 g C m(-2) year(-1) in older stands. Maximum ANPP in the Vaccinium-type chronosequence was 340 g C m(-2) year(-1) and occurred earlier in the 53-year-old stand than in the other stands. Along the lichen-type chronosequences, peak ANPP was paralleled by maximum carbon allocation to stem growth. (3) In mature trees, damage by recurrent surface fires decreased stem growth by 17 +/- 19% over a 10-year period relative to pre-fire values. At longer timescales, ANPP was hardly affected by fire-related differences in mortality. (4) Needle- plus stem-NPP, reconstructed for a 3-year period, varied within a range of 15 g C m(-2) year(-1) in the lichen-type stands and 35 g C m(-2) year(-1) in the Vaccinium-type stands. For the same period, the coefficient of variance was higher for needle-NPP (20 +/- 10%) than for stem-NPP (12 +/- 7%). Needle- and stem-NPP did not covary in time. Most 30-year time series of stem-NPP at the tree level exhibited strong autocorrelation. In older trees, stem-NPP was positively correlated with growing season precipitation. Thus, the factors driving variability in ANPP ranked according to their maximum influence as: stand age (controlled by the frequency of stand-replacing fires) > site quality > growth depression because of surface fire damage approximately equal age-related reduction in ANPP > interannual variability approximately equal long-term effects of fire (stand density reduction). In lichen-type forests, we found that ANPP at the landscape level declined sharply when the interval between stand-replacing fires was less than 120 years, illustrating that fire strongly influences ANPP of boreal Scots pine forests.

Above-ground biomass and structure of pristine Siberian Scots pine forests as controlled by competition and fire.

The study presents a data set of above-ground biomass (AGB), structure, spacing and fire regime, for 24 stands of pristine Siberian Scots pine (Pinus sylvestris) forests with lichens (n = 20) or Vaccinium/mosses (n = 4) as ground cover, along four chronosequences. The stands of the "lichen" site type (LT) were stratified into three chronosequences according to stand density and fire history. Allometric equations were established from 90 sample trees for stem, coarse branch, fine branch, twig and needle biomass. The LT stands exhibited a low but sustained biomass accumulation until a stand age of 383 years. AGB reached only 6-10 kgdw m-2 after 200 years depending on stand density and fire history compared to 20 kgdw m-2 in the "Vaccinium" type (VT) stands. Leaf area index (LAI) in the LT stands remained at 0.5-1.5 and crown cover was 30-60%, whereas LAI reached 2.5 and crown cover was >100% in the VT stands. Although nearest-neighbour analyses suggested the existence of density-dependent mortality, fire impact turned out to have a much stronger effect on density dynamics. Fire scar dating and calculation of mean and initial fire return intervals revealed that within the LT stands differences in structure and biomass were related to the severity of fire regimes, which in turn was related to the degree of landscape fragmentation by wetlands. Self-thinning analysis was used to define the local carrying capacity for biomass. A series of undisturbed LT stands was used to characterise the upper self-thinning boundary. Stands that had experienced a moderate fire regime were positioned well below the self-thinning boundary in a distinct fire-thinning band of reduced major axis regression slope -0.26. We discuss how this downward shift resulted from alternating phases of density reduction by fire and subsequent regrowth. We conclude that biomass in Siberian Scots pine forests is strongly influenced by fire and that climate change will affect ecosystem functions predominantly via changes in fire regimes.

Leaf conductance and CO(2) assimilation of Larix gmelinii growing in an eastern Siberian boreal forest.

In July 1993, we measured leaf conductance, carbon dioxide (CO(2)) assimilation, and transpiration in a Larix gmelinii (Rupr.) Rupr. ex Kuzen forest in eastern Siberia. At the CO(2) concentration of ambient air, maximum values (mean of 10 highest measured values) for CO(2) assimilation, transpiration and leaf conductance for water vapor were 10.1 micro mol m(-2) s(-1), 3.9 mmol m(-2) s(-1) and 365 mmol m(-2) s(-1), respectively. The corresponding mean values, which were much lower than the maximum values, were 2.7 micro mol m(-2) s(-1), 1.0 mmol m(-2) s(-1) and 56 mmol m(-2) s(-1). The mean values were similar to those of Vaccinium species in the herb layer. The large differences between maximum and actual performance were the result of structural and physiological variations within the tree crowns and between trees that reduced maximum assimilation and leaf conductance by about 40 and 60%, respectively. Thus, maximum assimilation and conductance values averaged over the canopy were 6.1 micro mol m(-2) s(-1) and 146 mmol m(-2) s(-1), respectively. Dry air caused stomatal closure, which reduced assimilation by an additional 26%. Low irradiances in the morning and evening had a minor effect (-6%). Daily canopy transpiration was estimated to be 1.45 mm day(-1), which is higher than the value of 0.94 mm day(-1) measured by eddy covariance, but similar to the value of 1.45 mm day(-1) calculated from the energy balance and soil evaporation, and less than the value of 2.1 mm day(-1) measured by xylem flux. Daytime canopy carbon assimilation, expressed on a ground area basis, was 0.217 mol m(-2) day(-1), which is higher than the value measured by eddy flux (0.162 mol m(-2) day(-1) including soil respiration). We discuss the regulation of leaf gas exchange in Larix under the extreme climatic conditions of eastern Siberia (temperature > 35 degrees C and vapor pressure deficit > 5.0 kPa).

Environmental regulation of xylem sap flow and total conductance of Larix gmelinii trees in eastern Siberia.

Xylem sap flow and environmental variables were measured on seven consecutive midsummer days in a 130-year-old Larix gmelinii (Rupr.) Rupr. forest located 160 km south of Yakutsk in eastern Siberia, Russia (61 degrees N, 128 degrees E, 300 m asl). The site received 20 mm of rainfall during the 4 days before measurements, and soil samples indicated that the trees were well watered. The tree canopy was sparse with a one-sided leaf area index of 1.5 and a tree density of 1760 ha(-1). On a clear day when air temperature ranged from 9 to 29 degrees C, and maximum air saturation deficit was 3.4 kPa, daily xylem sap flux (F) among 13 trees varied by an order of magnitude from 7 l day(-1) for subcanopy trees (representing 55% of trees in the forest) to 67 l day(-1) for emergent trees (representing 18% of trees in the forest). However, when based on xylem sap flux density (F'), calculated by dividing F by projected tree crown area (a surrogate for the occupied ground area), there was only a fourfold range in variability among the 13 trees, from 1.0 to 4.4 mm day(-1). The calculation of F' also eliminated systematic and large differences in F among emergent, canopy and subcanopy trees. Stand-level F', estimated by combining half-hourly linear relationships between F and stem cross-sectional area with tree size distribution data, ranged between 1.8 +/- 0.4 (standard deviation) and 2.3 +/- 0.6 mm day(-1). These stand-level F' values are about 0.6-0.7 mm day(-1) (30%) larger than daily tree canopy transpiration rates calculated from forest energy balance and understory evaporation measurements. Maximum total tree conductance for water vapor transfer (G(tmax), including canopy and aerodynamic conductances), calculated from the ratio of F' and the above-canopy air saturation deficit (D) for the eight trees with continuous data sets, was 9.9 +/- 2.8 mm s(-1). This is equivalent to a leaf-scale maximum stomatal conductance (g(smax)) of 6.1 mm s(-1), when expressed on a one-sided leaf area basis, which is comparable to the published porometer data for Larix. Diurnal variation in total tree conductance (G(t)) was related to changes in the above-canopy visible irradiance (Q) and D. A saturating upper-boundary function for the relationship between G(t) and Q was defined as G(t) = G(tmax)(Q/[Q + Q(50)]), where Q(50) = 164 +/- 85 micro mol m(-2) s(-1) when G(t) = G(tmax)/2. Accounting for Q by excluding data for Q < Q(85) when G(t) was at least 85% of G(tmax), the upper limit for the relationship between G(t) and D was determined based on the function G(t) = (a + blnD)(2), where a and b are regression coefficients. The relationship between G(t) and D was curvilinear, indicating that there was a proportional decrease in G(t) with increasing D such that F was relatively constant throughout much of the day, even when D ranged between about 2 and 4 kPa, which may be interpreted as an adaption of the species to its continental climate. However, at given values of Q and D, G(t) was generally higher in the morning than in the afternoon. The additional environmental constraints on G(t) imposed by leaf nitrogen nutrition and afternoon water stress are discussed.