Snagfall the first decade after severe bark beetle infestation of high-elevation forests in Colorado, USA.

The persistence and fall rate of snags (standing dead trees) generated during bark beetle outbreaks have consequences for the behavior, effects, and suppression of potential wildfires, hazard tree and timber salvage operations, wildlife habitat, and numerous ecosystem processes. However, post-beetle snagfall dynamics are poorly understood in most forest types. We tagged standing live and dead lodgepole pine (Pinus contorta), subalpine fir (Abies lasiocarpa), and Engelmann spruce (Picea engelmannii), including beetle-killed pine snags following the peak of a recent mountain pine bark beetle outbreak in watersheds at the Fraser Experimental Forest in northcentral Colorado and sampled snagfall 10 and 12 years later. Bark beetle attacks began in 2003, peaked by 2006, and killed 78% of overstory lodgepole pine in 133 plots distributed across a range of stand and site conditions. Of those snags, only 17% fell between 2007 and 2018. Most snags broke at ground level, due to butt rot, and were oriented downhill. In contrast, snags that tipped up or snapped off above the ground were oriented with the prevailing winds. Equal numbers of snags fell singly and in multiple-tree groups, and equal numbers remained elevated rather than in contact with the ground. Lodgepole pine snagfall was 1.6-times higher on steep slopes (>40%) where dead pine density was higher, compared to flatter sites. Based on our findings and previous research, we estimate that one-half the beetle-killed lodgepole pine in high-elevation forests such as those at Fraser may fall within 15-20 yr of beetle infestation, but that some pine snags are likely to persist for decades longer. Post-outbreak snagfall dynamics create a multiple-decade legacy of bark beetle outbreaks that will persist longer in high-elevation compared to lower-elevation forests.

Woody-tissue respiration for Simarouba amara and Minquartia guianensis, two tropical wet forest trees with different growth habits.

We measured CO2 efflux from stems of two tropical wet forest trees, both found in the canopy, but with very different growth habits. The species were Simarouba amara, a fast-growing species associated with gaps in old-growth forest and abundant in secondary forest, and Minquartia guianensis, a slow-growing species tolerant of low-light conditions in old-growth forest. Per unit of bole surface, CO2 efflux averaged 1.24 µmol m-2 s-1 for Simarouba and 0.83 µmol m-2s-1 for Minquartia. CO2 efflux was highly correlated with annual wood production (r 2=0.65), but only weakly correlated with stem diameter (r 2=0.22). We also partitioned the CO2 efflux into the functional components of construction and maintenance respiration. Construction respiration was estimated from annual stem dry matter production and maintenance respiration by subtracting construction respiration from the instantaneous CO2 flux. Estimated maintenance respiration was linearly related to sapwood volume (39.6 µmol m-3s-1 at 24.6° C, r 2=0.58), with no difference in the rate for the two species. Maintenance respiration per unit of sapwood volume for these tropical wet forest trees was roughly twice that of temperate conifers. A model combining construction and maintenance respiration estimated CO2 very well for these species (r 2=0.85). For our sample, maintenance respiration was 54% of the total CO2 efflux for Simarouba and 82% for Minquartia. For our sample, sapwood volume averaged 23% of stem volume when weighted by tree size, or 40% with no size weighting. Using these fractions, and a published estimate of aboveground dry-matter production, we estimate the annual cost of woody tissue respiration for primary forest at La Selva to be 220 or 350 g C m-2 year-1, depending on the assumed sapwood volume. These costs are estimated to be less than 13% of the gross production for the forest.

Woody tissue maintenance respiration of four conifers in contrasting climates.

We estimate maintenance respiration for boles of four temperate conifers (ponderosa pine, western hemlock, red pine, and slash pine) from CO2 efflux measurements in autumn, when construction respiration is low or negligible. Maintenance respiration of stems was linearly related to sapwood volume for all species; at 10°C, respiration per unit sapwood volume ranged from 4.8 to 8.3 µmol CO2 m-3 s-1. For all sites combined, respiration increased exponentially with temperature (Q 10 =1.7, r 2=0.78). We estimate that maintenance respiration of aboveground woody tissues of these conifers consumes 52-162 g C m-2 y-1, or 5-13% of net daytime carbon assimilation annually. The fraction of annual net daytime carbon fixation used for stem maintenance respiration increased linearly with the average annual temperature of the site.

Effects of branch height on leaf gas exchange, branch hydraulic conductance and branch sap flux in open-grown ponderosa pine.

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.

Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees.

We tested the hypotheses that hydraulic conductance is lower in old (about 250 years old and 30 m tall) compared to young (about 40 years old and 10 m tall) Pinus ponderosa Dougl. ex Laws. trees and that lower hydraulic conductance of old trees limits their photosynthesis. Hydraulic conductance at the end of summer 1995, calculated from leaf water potential and leaf gas exchange measurements on one-year-old needles, was 44% lower in old trees compared to young trees growing in a mixed age-class stand on the east slope of the Oregon Cascades. Whole-tree sapflow per unit leaf area averaged 53% lower in old trees compared to young trees and mean hydraulic conductance calculated from sapflow and water potential data was 63% lower in old trees than in young trees. For the entire summer, stomatal conductance (g(s)) and assimilation (A) declined more steeply with air saturation deficit (D) in old trees than in young trees. For both old and young trees, mean g(s) and A were approximately 32 and 21% lower, respectively, at typical midday D values (2.5-3.0 kPa). We hypothesized that if hydraulic conductance limits g(s) and A, then increasing or decreasing the leaf specific conductance of a branch will result in proportional changes in the responses of g(s) and A with D. Removal of 50% of the foliage from a set of experimental branches on old trees caused g(s) and A to decline less steeply with D in early summer, but values were not significantly different from control values in late summer. Cutting transverse notches in branches on young trees had no effect on the responses of g(s) and A with D. Leaf nitrogen content and photosynthetic capacity were similar suggesting that differences in g(s) and A between old and young trees were not caused by differences in photosynthetic capacity.