Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

Dynamics of Photosynthesis in Pine Stands

The CO₂ exchange of pine (Pinus sylvestris L.) shoots was continuously monitored over several years. The spatial and temporal variability of gas exchange was thoroughly investigated for three forest types. The net uptake of CO₂ by current-season shoots in a bilberry pine stand equaled 5.4 g CO₂ per g dry wt for the growing season. The net CO₂ assimilation by one-year-old shoots over a growing season constituted 9.9 and 2.4 g CO₂/(g dry wt year) in the upper and lower crown parts, respectively. The nighttime respiration of the current-season shoots released 0.7 g CO₂/(g dry wt year), and the respiration of one-year-old shoots in the upper and lower parts of the canopy released 0.45 and 0.36 g CO₂/(g dry wt year), respectively. Recalculation of three-year data per entire crown yielded an annual average carbon assimilation of 1.54 g C/(g dry wt year). Water relations markedly affected CO₂ fixation. Nevertheless, the daily average values of photosynthesis in the summer were similar for pine stands with bilberry, heather, and dwarf shrub–polytric as understory dominants. It is shown that the realization of changes in photosynthetic function is related in time to growth processes.     

Soil-surface carbon dioxide efflux and microbial biomass in relation to tree density 13 years after a stand replacing fire in a lodgepole pine ecosystem

The effects of fire on soil‐surface carbon dioxide (CO₂) efflux, F_S, and microbial biomass carbon, C_mic, were studied in a wildland setting by examining 13‐year‐old postfire stands of lodgepole pine differing in tree density (< 500 to > 500 000 trees ha^?1) in Yellowstone National Park (YNP). In addition, young stands were compared to mature lodgepole pine stands (～110‐year‐old) in order to estimate ecosystem recovery 13 years after a stand replacing fire. Growing season F_S increased with tree density in young stands (1.0 µmol CO₂ m^?2 s^?1 in low‐density stands, 1.8 µmol CO₂ m^?2 s^?1 in moderate‐density stands and 2.1 µmol CO₂ m^?2 s^?1 in high‐density stands) and with stand age (2.7 µmol CO₂ m^?2 s^?1 in mature stands). Microbial biomass carbon in young stands did not differ with tree density and ranged from 0.2 to 0.5 mg C g^?1 dry soil over the growing season; C_mic was significantly greater in mature stands (0.5–0.8 mg C g^?1 dry soil). Soil‐surface CO₂ efflux in young stands was correlated with biotic variables (above‐ground, below‐ground and microbial biomass), but not with abiotic variables (litter and mineral soil C and N content, bulk density and soil texture). Microbial biomass carbon was correlated with below‐ground plant biomass and not with soil carbon and nitrogen, indicating that plant activity controls not only root respiration, but C_mic pools and overall F_S rates as well. These findings support recent studies that have demonstrated the prevailing importance of plants in controlling rates of F_S and suggest that decomposition of older, recalcitrant soil C pools in this ecosystem is relatively unimportant 13 years after a stand replacing fire. Our results also indicate that realistic predictions and modeling of terrestrial C cycling must account for the variability in tree density and stand age that exists across the landscape as a result of natural disturbances.     

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.     

Photosynthetic responses of Mojave Desert shrubs to free air CO2 enrichment are greatest during wet years

It has been suggested that desert vegetation will show the strongest response to rising atmospheric carbon dioxide due to strong water limitations in these systems that may be ameliorated by both photosynthetic enhancements and reductions in stomatal conductance. Here, we report the long‐term effect of 55 Pa atmospheric CO₂ on photosynthesis and stomatal conductance for three Mojave Desert shrubs of differing leaf phenology (Ambrosia dumosa—drought‐deciduous, Krameria erecta—winter‐deciduous, Larrea tridentata—evergreen). The shrubs were growing in an undisturbed ecosystem fumigated using FACE technology and were measured over a four‐year period that included both above and below‐average precipitation. Daily integrated photosynthesis (A_day) was significantly enhanced by elevated CO₂ for all three species, although Krameria erecta showed the greatest enhancements (63% vs. 32% for the other species) enhancements were constant throughout the entire measurement period. Only one species, Larrea tridentata, decreased stomatal conductance by 25–50% in response to elevated CO₂, and then only at the onset of the summer dry season and following late summer convective precipitation. Similarly, reductions in the maximum carboxylation rate of Rubisco were limited to Larrea during spring. These results suggest that the elevated CO₂ response of desert vegetation is a function of complex interactions between species functional types and prevailing environmental conditions. Elevated CO₂ did not extend the active growing season into the summer dry season because of overall negligible stomatal conductance responses that did not result in significant water conservation. Overall, we expect the greatest response of desert vegetation during years with above‐average precipitation when the active growing season is not limited to ～ 2 months and, consequently, the effects of increased photosynthesis can accumulate over a biologically significant time period.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

Increased leaf area dominates carbon flux response to elevated CO2 in stands of Populus deltoides (Bartr.)

We examined the effects of atmospheric vapor pressure deficit (VPD) and soil moisture stress (SMS) on leaf‐ and stand‐level CO₂ exchange in model 3‐year‐old coppiced cottonwood (Populus deltoides Bartr.) plantations using the large‐scale, controlled environments of the Biosphere 2 Laboratory. A short‐term experiment was imposed on top of continuing, long‐term CO₂ treatments (43 and 120 Pa), at the end of the growing season. For the experiment, the plantations were exposed for 6–14 days to low and high VPD (0.6 and 2.5 kPa) at low and high volumetric soil moisture contents (25–39%). When system gross CO₂ assimilation was corrected for leaf area, system net CO₂ exchange (SNCE), integrated daily SNCE, and system respiration increased in response to elevated CO₂. The increases were mainly as a result of the larger leaf area developed during growth at high CO₂, before the short‐term experiment; the observed decline in responses to SMS and high VPD treatments was partly because of leaf area reduction. Elevated CO₂ ameliorated the gas exchange consequences of water stress at the stand level, in all treatments. The initial slope of light response curves of stand photosynthesis (efficiency of light use by the stand) increased in response to elevated CO₂ under all treatments. Leaf‐level net CO₂ assimilation rate and apparent quantum efficiency were consistently higher, and stomatal conductance and transpiration were significantly lower, under high CO₂ in all soil moisture and VPD combinations (except for conductance and transpiration in high soil moisture, low VPD). Comparisons of leaf‐ and stand‐level gross CO₂ exchange indicated that the limitation of assimilation because of canopy light environment (in well‐irrigated stands; ratio of leaf : stand=3.2–3.5) switched to a predominantly individual leaf limitation (because of stomatal closure) in response to water stress (leaf : stand=0.8–1.3). These observations enabled a good prediction of whole stand assimilation from leaf‐level data under water‐stressed conditions; the predictive ability was less under well‐watered conditions. The data also demonstrated the need for a better understanding of the relationship between leaf water potential, leaf abscission, and stand LAI.     

Long-term impact of a stand-replacing fire on ecosystem CO2 exchange of a ponderosa pine forest

Ponderosa pine (Pinus ponderosa) forests of the southwestern United States are a mosaic of stands where undisturbed forests are carbon sinks, and stands recovering from wildfires may be sources of carbon to the atmosphere for decades after the fire. However, the relative magnitude of these sinks and sources has never been directly measured in this region, limiting our understanding of the role of fire in regional and US carbon budgets. We used the eddy covariance technique to measure the CO₂ exchange of two forest sites, one burned by fire in 1996, and an unburned forest. The fire was a high‐intensity stand‐replacing burn that killed all trees. Ten years after the fire, the burned site was still a source of CO₂ to the atmosphere [109±6 (SEM) g C m^?2 yr^?1], whereas the unburned site was a sink (?164±23 g C m^?2 yr^?1). The fire reduced total carbon storage and shifted ecosystem carbon allocation from the forest floor and living biomass to necromass. Annual ecosystem respiration was lower at the burned site (480±5 g C m^?2 yr^?1) than at the unburned site (710±54 g C m^?2 yr^?1), but the difference in gross primary production was even larger (372±13 g C m^?2 yr^?1 at the burned site and 858±37 g C m^?2 yr^?1at the unburned site). Water availability controlled carbon flux in the warm season at both sites, and the burned site was a source of carbon in all months, even during the summer, when wet and warm conditions favored respiration more than photosynthesis. Our study shows that carbon losses following stand‐replacing fires in ponderosa pine forests can persist for decades due to slow recovery of the gross primary production. Because fire exclusion is becoming increasingly difficult in dry western forests, a large US forest carbon sink could shift to a decadal‐scale carbon source.     

Earthworms, stand age, and species composition interact to influence particulate organic matter chemistry during forest succession

The landscapes colonized by invasive earthworms in the eastern U.S. are often patchworks of forest stands in various stages of successional development. We established six field sites in tulip poplar dominated forests in the Smithsonian Environmental Research Center in Edgewater, MD, that span mid (50–70 years-three plots) and late (120–150 years-three plots) successional stages where younger sites had greater earthworm density and biomass than older sites and were dominated by non-native lumbricid species. In particular Lumbricus rubellus, a litter-feeding species, was abundant in mid successional forests. Here, we separated particulate organic matter (POM) from the bulk soil by a combination of size and density fractionation and found that patterns in soil POM chemistry were similar to those found previously during litter decay: in younger forests with high abundance of earthworms, organic carbon normalized cutin- and suberin-derived substituted fatty acid (SFA) concentration was lower and lignin-derived phenols greater than in older forests where earthworms were less abundant. The chemistry of the dominant litter from mid versus late successional tree species did not fully explain the differences in POM chemistry between age classes. Instead, the differences in leaf body versus petiole and leaf versus root chemistry were the dominant drivers of POM chemistry in mid versus late successional stands, although aspects of stand age and tree species also impacted POM chemistry. Our results indicate that preferential ingestion of leaf body tissue by earthworms and the subsequent shifts in sources of plant biopolymers in soil influenced POM chemistry in mid successional forests. These results indicate that invasive earthworm activity in North American forests contributes to a shift in the aromatic and aliphatic composition of POM and thus potentially influences carbon stabilization in soil.     

Elevated CO2 alters carbon fluxes in early successional Mediterranean ecosystems

Annual carbon budgets of ecosystems are central to our understanding of the biotic control of atmospheric composition, but they are not available under elevated CO₂ for most vegetation types. Using gas exchange techniques, we assessed carbon fluxes of four early successional Mediterranean model communities, consisting of grasses, legumes and composites. The assemblages were grown on the same monoliths for three consecutive years in greenhouses tracking field conditions except for CO₂ maintained at ambient (370 μmol mol^?1) or elevated (700 μmol mol^?1) concentration. During the third year of study, CO₂ enrichment consistently shifted the annual carbon balance towards lower efflux, with displacements between 4.3 and 26.2 mol m^?2 y^?1 (one assemblage became a net CO₂ sink, another just reached equilibrium, and the remaining two remained as a CO₂ source). At least 50% of the shift under elevated CO₂ originated from a decrease in belowground respiration. This indicates that, during this year, CO₂ enrichment did not predominantly enhance C‐cycling, but on the contrary inhibited root respiration or microbial C‐utilization. Although elevated‐CO₂‐grown systems acted as a net CO₂ sink during a longer period of the year (4–7 months) compared with ambient‐CO₂‐grown systems (3–3.5 months), gross canopy photosynthesis was modified only to a limited extent (between ?5.9 and + 14.8%). Interaction between the carbon and the water cycle was apparently responsible for this weak stimulation. In particular, reduced evapotranspiration under elevated CO₂ coincided with inhibited canopy photosynthesis in early spring, most likely resulting from water saturation of the soil. In addition, only the earliest‐planted assemblages had an increased gross canopy photosynthesis during late autumn and early winter. This suggests that a longer summer drought, by delaying the establishment of such an annual type of vegetation, would reduce the positive impact of elevated CO₂ on productivity. Water regime appears to strongly govern the influence of CO₂ on the carbon fluxes in Mediterranean ecosystems with annual herbaceous vegetation.     

Patterns of NPP,GPP, respiration,and NEP during boreal forest succession

We combined year‐round eddy covariance with biometry and biomass harvests along a chronosequence of boreal forest stands that were 1, 6, 15, 23, 40, ～74, and ～154 years old to understand how ecosystem production and carbon stocks change during recovery from stand‐replacing crown fire. Live biomass (C_live) was low in the 1‐ and 6‐year‐old stands, and increased following a logistic pattern to high levels in the 74‐ and 154‐year‐old stands. Carbon stocks in the forest floor (C_{forest floor}) and coarse woody debris (C_CWD) were comparatively high in the 1‐year‐old stand, reduced in the 6‐ through 40‐year‐old stands, and highest in the 74‐ and 154‐year‐old stands. Total net primary production (TNPP) was reduced in the 1‐ and 6‐year‐old stands, highest in the 23‐ through 74‐year‐old stands and somewhat reduced in the 154‐year‐old stand. The NPP decline at the 154‐year‐old stand was related to increased autotrophic respiration rather than decreased gross primary production (GPP). Net ecosystem production (NEP), calculated by integrated eddy covariance, indicated the 1‐ and 6‐year‐old stands were losing carbon, the 15‐year‐old stand was gaining a small amount of carbon, the 23‐ and 74‐year‐old stands were gaining considerable carbon, and the 40‐ and 154‐year‐old stands were gaining modest amounts of carbon. The recovery from fire was rapid; a linear fit through the NEP observations at the 6‐ and 15‐year‐old stands indicated the transition from carbon source to sink occurred within 11–12 years. The NEP decline at the 154‐year‐old stand appears related to increased losses from C_live by tree mortality and possibly from C_{forest floor} by decomposition. Our findings support the idea that NPP, carbon production efficiency (NPP/GPP), NEP, and carbon storage efficiency (NEP/TNPP) all decrease in old boreal stands.     

Stand age-related effects on soil respiration in a first rotation Sitka spruce chronosequence in central Ireland

The effect of stand age on soil respiration and its components was studied in a first rotation Sitka spruce chronosequence composed of 10‐, 15‐, 31‐, and 47‐year‐old stands established on wet mineral gley in central Ireland. For each stand age, three forest stands with similar characteristics of soil type and site preparation were used. There were no significant differences in total soil respiration among sites of the same age, except for the case of a 15‐year‐old stand that had lower soil respiration rates due to its higher productivity. Soil respiration initially decreased with stand age, but levelled out in the older stands. The youngest stands had significantly higher respiration rates than more mature sites. Annual soil respiration rates were modelled by means of temperature‐derived functions. The average Q ₁₀ value obtained treating all the stands together was 3.8. Annual soil respiration rates were 991, 686, 556, and 564 g C m^?2 for the 10‐, 15‐, 31‐, and 47‐year‐old stands, respectively. We used the trenching approach to separate soil respiration components. Heterotrophic respiration paralleled soil organic carbon dynamics over the chronosequence, decreasing with stand age to slightly increase in the oldest stand as a result of accumulated aboveground litter and root inputs. Root respiration showed a decreasing trend with stand age, which was explained by a decrease in fine root biomass over the chronosequence, but not by nitrogen concentration of fine roots. The decrease in the relative contribution of autotrophic respiration to total soil CO₂ efflux from 59.3% in the youngest stand to 49.7% in the oldest stand was explained by the higher activity of the root system in younger stands. Our results show that stand age should be considered if simple temperature‐based models to predict annual soil respiration in afforestation sites are to be used.     

Canopy transpiration in a chronosequence of Central Siberian pine forests

Tree transpiration was measured in 28, 67, 204 and 383‐y‐old uniform stands and in a multicohort stand (140–430 y) of Pinus sylvestris ssp. sibirica Lebed. in Central Siberia during August 1995. In addition transpiration of three codominant trees was monitored for two years in a 130‐y‐old stand. All stands established after fire. Leaf area index (LAI) ranged between 0.6 (28‐y‐old stand) and 1.6 for stands older than 67‐y. Stand xylem area at 1.3 m height increased from 4 cm² m^?2 (28‐y) to 11.5 cm² m^?2 (67‐y) and decreased again to 7 cm² m^?2 in old stands. Above‐ground living biomass increased from 1.5 kg dry weight m^?2 (28‐y) to 14 kg dry weight m^?2 (383‐y). Day‐to‐day variation of tree transpiration in summer was dependent on net radiation, vapour pressure deficit, and soil water stress. Tree‐to‐tree variation of xylem flux was small and increased with heterogeneity in canopy structure. Maximum rates of xylem flux density followed the course of net radiation from mid April when a constant level of maximum rates was reached until mid September when low temperatures and light strongly reduced flux density. Maximum sap flux density (60 g m^?2 s^?1) and canopy transpiration (1.5 mm d^?1) were reached in the 67‐y stand. Average canopy transpiration of all age classes was 0.72 ± 0.3 mm d^?1. Canopy transpiration (E) was not correlated with LAI but related to stand sapwood area SA (E = ? 0.02 + 1.15SA R²) which was determined by stand density and tree sapwood area.     

Carbon dioxide,methane, and nitrous oxide exchanges in an age‐sequence of temperate pine forests

We investigated soil carbon dioxide (CO₂), methane (CH₄), and nitrous oxide (N₂O) exchanges in an age‐sequence (4, 17, 32, 67 years old) of eastern white pine (Pinus strobus L.) forests in southern Ontario, Canada, for the period of mid‐April to mid‐December in 2006 and 2007. For both CH₄ and N₂O, we observed uptake and emission ranging from ?160 to 245 μg CH₄ m^?2 h^?1 and ?52 to 21 μg N₂O m^?2 h^?1, respectively (negative values indicate uptake). Mean fluxes from mid‐April to mid‐December across the 4, 17, 32, 67 years old stands were similar for CO₂ fluxes (259, 246, 220, and 250 mg CO₂ m^?2 h^?1, respectively), without pattern for N₂O fluxes (?3.7, 1.5, ?2.2, and ?7.6 μg N₂O m^?2 h^?1, respectively), whereas the uptake rates of CH₄ increased with stand age (6.4, ?7.9, ?10.8, and ?23.3 μg CH₄ m^?2 h^?1, respectively). For the same period, the combined contribution of CH₄ and N₂O exchanges to the global warming potential (GWP) calculated from net ecosystem exchange of CO₂ and aggregated soil exchanges of CH₄ and N₂O was on average 4%, <1%, <1%, and 2% for the 4, 17, 32, 67 years old stand, respectively. Soil CO₂ fluxes correlated positively with soil temperature but had no relationship with soil moisture. We found no control of soil temperature or soil moisture on CH₄ and N₂O fluxes, but CH₄ emission was observed following summer rainfall events. LFH layer removal reduced CO₂ emissions by 43%, increased CH₄ uptake during dry and warm soil conditions by more than twofold, but did not affect N₂O flux. We suggest that significant alternating sink and source potentials for both CH₄ and N₂O may occur in N‐ and soil water‐limited forest ecosystems, which constitute a large portion of forest cover in temperate areas.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Short-term effects of extensive fertilization on community composition and carbon uptake in a Pannonian loess grassland

Among the most extended ecosystems of the temperate zone, the seminatural, dry grasslands constitute a substantial proportion in the Carpathian Basin. The aim of our present study was to investigate the short-term effect of extensive fertilization on the species composition and CO₂ exchange of loess grassland at community level. The in situ investigation of the latter parameter have not been yet carried out in Pannonian loess grasslands. Most of the parameters studied showed a considerable interannual variation both in the fertilized and in the control stands. As a result of the treatment, the average species number of the fertilized stand decreased by 22%, which was more significant in the autumn (26%) than in the spring. Diversity values, including Shannon index and species richness, increased by nearly 1.5 times in the year with adequate rainfall compared with the initial values. In general, species richness and the ratio of dicots decreased, while the ratio of therophytes, alien competitors, and C₄ plants increased with the addition of fertilizers. Significant carbon sequestration potential was only detected during wet periods in the fertilized grass. The rate of CO₂ uptake was found to be nearly five times higher in the fertilized stand and nearly three times higher in the control stand during the wet year compared with the previous, extremely dry year. The CO₂ uptake potential of the fertilized grassland exceeded that of the control stand by 12% in the year with high rainfall, while the rate of CO₂ exchange dropped by 50% in the dry year in the fertilized stand. Our study reinforced the idea that the decline in species richness was not necessarily followed by the reduction of stand level carbon uptake in a short period due to an insignificant change in ecophysiological functional groups.     

Elevated CO2 affects photosynthetic responses in canopy pine and subcanopy deciduous trees over 10 years: a synthesis from Duke FACE

Leaf responses to elevated atmospheric CO₂ concentration (C_a) are central to models of forest CO₂ exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated C_a requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up‐scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated C_a in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of Pinus taeda, but also including deciduous species, primarily Liquidambar styraciflua. We combined over one thousand controlled‐response curves of photosynthesis as a function of environmental drivers (light, air C_a and temperature) measured at canopy heights up to 20 m over 11 years (1996–2006) to generate parameterizations for leaf‐scale models for the Duke free‐air CO₂ enrichment (FACE) experiment. The enhancement of leaf net photosynthesis (A_net) in P. taeda by elevated C_a of +200 μmol mol^?1 was 67% for current‐year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P. taeda at the leaf‐scale increased by two‐fold from the driest to wettest growing seasons. Current‐year pine foliage A_net was sensitive to temporal variation, whereas previous‐year foliage A_net was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N (N_area), but statistically significant. In contrast, co‐dominant and subcanopy L. styraciflua trees showed A_net enhancement of 62% and no A_net–N_area adjustments. Various understory deciduous tree species showed an average A_net enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased C_a has the potential to enhance the mixed‐species composition of planted pine stands and, by extension, naturally regenerating pine‐dominated stands.     

Native and exotic plant cover vary inversely along a climate gradient 11 years following stand‐replacing wildfire in a dry coniferous forest,Oregon, USA

Community re‐assembly following future disturbances will often occur under warmer and more moisture‐limited conditions than when current communities assembled. Because the establishment stage is regularly the most sensitive to climate and competition, the trajectory of recovery from disturbance in a changing environment is uncertain, but has important consequences for future ecosystem functioning. To better understand how ongoing warming and rising moisture limitation may affect recovery, we studied native and exotic plant composition 11 years following complete stand‐replacing wildfire in a dry coniferous forest spanning a large gradient in climatic moisture deficit (CMD) from warm and dry low elevation sites to relatively cool and moist higher elevations sites. We then projected future precipitation, temperature and CMD at our study locations for four scenarios selected to encompass a broad range of possible future conditions for the region. Native perennials dominated relatively cool and moist sites 11 years after wildfire, but were very sparse at the warmest and driest (high CMD) sites, particularly when combined with high topographic sun exposure. In contrast, exotic species (primarily annual grasses) were dominant or co‐dominant at the warmest and driest sites, especially with high topographic sun exposure. All future scenarios projected increasing temperature and CMD in coming decades (e.g., from 4.5% to 29.5% higher CMD by the 2080's compared to the 1971–2000 average), even in scenarios where growing season (May‐September) precipitation increased. These results suggest increasing temperatures and moisture limitation could facilitate longer term (over a decade) transitions toward exotic‐dominated communities after severe wildfire when a suitable exotic seed source is present.     

An eddy covariance mesonet to measure the effect of forest age on land–atmosphere exchange

We deployed a mesonet of year‐round eddy covariance towers in boreal forest stands that last burned in ～1850, ～1930, 1964, 1981, 1989, 1998, and 2003 to understand how CO₂ exchange and evapotranspiration change during secondary succession. We used MODIS imagery to establish that the tower sites were representative of the patterns of secondary succession in the region, and Landsat images to show that the individual stands have changed over the last 22 years in ways that match the spatially derived trends. The eddy covariance towers were well matched, with similar equipment and programs, which maximized site‐to‐site precision and allowed us to operate the network in an efficient manner. The six oldest sites were fully operational for ～90% of the growing season and ～70% of the dormant season from 2001 or 2002 to 2004, with most of the missing data caused by low battery charge or bad signals from the sonic anemometers. The rates of midday growing‐season CO₂ uptake recovered to preburn levels within 4 years of fire. The seasonality of land–atmosphere exchange and growing‐season length changed markedly with stand age. The foliage in the younger stands (1989, 1998, and 2003 burns) was almost entirely deciduous, which resulted in comparatively short growing seasons that lasted ～65 days. In contrast, the older stands (1850, 1930, 1964, and 1981) were mostly evergreen, which resulted in comparatively long growing seasons that lasted ～130 days. The eddy covariance mesonet approach we describe could be used within the context of other ecological experimental designs such as controlled manipulations and gradient comparisons.     

Photosynthesis drives anomalies in net carbon-exchange of pine forests at different latitudes

The growth rate of atmospheric CO₂ exhibits large temporal variation that is largely determined by year‐to‐year fluctuations in land–atmosphere CO₂ fluxes. This land–atmosphere CO₂‐flux is driven by large‐scale biomass burning and variation in net ecosystem exchange (NEE). Between‐ and within years, NEE varies due to fluctuations in climate. Studies on climatic influences on inter‐ and intra‐annual variability in gross photosynthesis (GPP) and net carbon uptake in terrestrial ecosystems have shown conflicting results. These conflicts are in part related to differences in methodology and in part to the limited duration of some studies. Here, we introduce an observation‐driven methodology that provides insight into the dependence of anomalies in CO₂ fluxes on climatic conditions. The methodology was applied on fluxes from a boreal and two temperate pine forests. Annual anomalies in NEE were dominated by anomalies in GPP, which in turn were correlated with incident radiation and vapor pressure deficit (VPD). At all three sites positive anomalies in NEE (a reduced uptake or a stronger source than the daily sites specific long‐term average) were observed on summer days characterized by low incident radiation, low VPD and high precipitation. Negative anomalies in NEE occurred mainly on summer days characterized by blue skies and mild temperatures. Our study clearly highlighted the need to use weather patterns rather than single climatic variables to understand anomalous CO₂ fluxes. Temperature generally showed little direct effect on anomalies in NEE but became important when the mean daily air temperature exceeded 23 °C. On such days GPP decreased likely because VPD exceeded 2.0 kPa, inhibiting photosynthetic uptake. However, while GPP decreased, the high temperature stimulated respiration, resulting in positive anomalies in NEE. Climatic extremes in summer were more frequent and severe in the South than in the North, and had larger effects in the South because the criteria to inhibit photosynthesis are more often met.