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1.
Jack pine budworm (Choristoneura pinus pinus Free.) (Lepidoptera: Tortricidae) is a native insect that periodically defoliates areas of jack pine (Pinus banksiana Lamb.) in the subboreal forests of North America east of the Rocky Mountains. Outbreaks of jack pine budworm generally occur at 6- to 12-year intervals and collapse after 2–4 years. Periodicity of outbreaks varies and is associated with site-related factors. Survival of early-instar larvae during spring dispersal is tied to the abundance of pollen cones (microsporangiate strobili), which provide a refuge for larvae until current-year needles expand. Jack pine trees that have been heavily defoliated produce few pollen cones in the following year, often resulting in high mortality of early-stage larvae. A diverse guild of generalist parasitoids attack jack pine budworm, but only a few species account for most mortality in any area. Collapsing jack pine budworm populations are characterized by sharp declines in early instar survival, coupled with an increased rate of parasitism in the late larval and pupal stages. The reciprocal interaction between heavy defoliation and low pollen cone production, and increased parasitism of late-stage larvae or pupae, are consistent with second-order density dependence factors identified in analysis of a long-term population data set. Since the 1950s, several jack pine budworm outbreaks have been roughly synchronous over a large geographic area, suggesting that Moran effect processes, as well as moth dispersal or other factors, may be involved in jack pine budworm dynamics. Although the short duration of outbreaks enables most trees to recover, over time dead trees and top-killed trees accumulate in jack pine stands. Jack pine is well adapted to fire and when fires ignite, the accumulation of dead trees and woody debris often leads to intense wildfires followed by prolific regeneration. The three-way interaction of jack pine, jack pine budworm, and fire ultimately serves to maintain vigorous stands and ensures continued hosts for jack pine budworm. Received: October 1, 1999 / Accepted: September 22, 2000  相似文献   

2.
Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.  相似文献   

3.
4.
Microsporidia are believed to play little or no role in outbreaks of the jack pine budworm, Choristoneura pinuspinus Freeman (Lepidoptrera: Tortricidae), because the short duration (2–4 years) of those outbreaks may not permit significant build-up of the pathogen. We conducted the first survey of Nosema sp. (Microsporidia: Nosematidae) over the course of a recent jack pine budworm outbreak in Ontario. Between 2004 and 2010 the outbreak defoliated a cumulative total of 1.78 million ha. Microscopic examination of ∼15,000 overwintering larvae collected over 6 years in sites with densities of 3 larvae per branch or more revealed widespread occurrence of Nosema at generally high infection intensities. The pathogen was present in 69.5% of the 518 plots that were monitored. Prevalence of infection was generally low (below 40% in 84% of plots with infected larvae) but reached high levels (80–95%) locally and increased rapidly in most infestations within 1–2 years of onset. We hypothesize that the habit of early-instar larvae to feed on developing male flowers (pollen cones) after spring emergence is critical in allowing rapid build-up of Nosema by increasing efficiency of horizontal transmission (higher density of both infected larvae and egested spores). Nosema infection may contribute to the complexity of jack pine budworm outbreak patterns by affecting egg recruitment and early-instar survival at the stand level in concert with known effects of budworm-induced reductions in pollen cone production on those processes.  相似文献   

5.
We report data collected over the entire course of an outbreak of jack pine budworm, Choristoneura pinus pinus Freeman (Lepidoptera: Tortricidae), between 1984 and 1988 at 12 plots in Manitoba. The positive relationship between the level of defoliation, used as a proxy for population density, and the abundance of eggs suggests local reproduction by females. The density of pollen cones apparently reduced larval fitness in 1986, when flowers were least abundant, but had limited impact in other years when pollen cones were abundant; this suggests that the relative abundance of pollen cones is more likely to influence the termination of an outbreak than its onset. Considering the conditions that prevail at the onset of the outbreak (low defoliation combined with a high abundance of eggs), a predictive tool may be developed to anticipate outbreaks of jack pine budworm based on environmental conditions that are conducive to high survival and/or fecundity of females.  相似文献   

6.
The influence of landscape patterns on ecological processes is generally acknowledged, but often difficult to quantify. The objective of our study was to quantify the relation of jack pine budworm ( Choristoneura pinus pinus ) population levels to both the abundance of jack pine ( Pinus banksiana ) and of jack pine stand edges in the landscape. The 450 000 ha Pine Barrens region, located in northwestern Wisconsin, USA, experienced a severe jack pine budworm outbreak from 1990 to 1995. We calculated landscape indices on a landcover map derived from Landsat TM satellite imagery. Landscape indices were calculated on circular buffers (0.5, 0.75, 1, 1.5, and 2.5 km radius) centered on 143 budworm population sampling points for which annual budworm counts were available. Edge density was normalized for the proportion of jack pine in the landscape using random maps as a standard. Correlations between landscape patterns and budworm populations varied over time: proportion of jack pine showed strongest positive correlation with budworm population levels up to the peak of the outbreak (1993). Edge density exhibited positive correlation up to the peak of the outbreak, but negative correlation in the subsequent years as the outbreak declined. This may suggest that pollen-bearing male cones, which are more abundant along edges, support higher budworm populations in the initial phase of the outbreak, but stronger predation on budworm along edges subsequently reduces populations. We provide insight into previously inconclusive results on the relation of jack pine budworm population density to jack pine stand edges. The effects of landscape patterns, such as edge density, may vary not only in magnitude, but also in direction, being positive and negative during different phases of an insect outbreak. Therefore, caution should be taken in relating landscape patterns to process at either a single scale or point in time.  相似文献   

7.
The formerly endangered Kirtland's warbler (Setophaga kirtlandii) is among a growing number of conservation-reliant species that depend on active management to avoid reverting to endangered status. Because the Kirtland's warbler is a habitat specialist of young, even-aged jack pine (Pinus banksiana), managers of the recovery effort stressed creating new jack pine stands and monitoring numbers of singing males through an annual census using single visits to individual stands. Kirtland's warbler will occupy and breed in red pine (P. resinosa), but red pine has not been surveyed for Kirtland's warblers in the annual population census. Furthermore, the current monitoring approach cannot determine their species detection probability or individual detection probability, which is essential to evaluate both red pine use and the accuracy of the census. From 2016–2018 we estimated density and detection probabilities in jack pine and red pine stands through repeated visits to a limited number of stands rather than single visits to many stands. Estimates of species detection probability indicated that ≥1 male Kirtland's warbler would be detected on most sites when any were present, but individual detection probabilities were less and varied by stand type, indicating that single visits to sites would underestimate numbers and that accurate estimation of detection probability was important for estimation of density in different stand types. We offer quantitative estimates of detection probabilities for determination of Kirtland's warbler population size in jack pine versus red pine stands in the same areas and breeding seasons. Managers of Kirtland's warblers should incorporate detection probabilities into population surveys to achieve more accurate estimates of population size.  相似文献   

8.
We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m2 plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m2 in the plantations, while it varied between 50 and 180 saplings/100 m2 in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.  相似文献   

9.
Goubitz  S.  Nathan  R.  Roitemberg  R.  Shmida  A.  Ne’eman  G. 《Plant Ecology》2004,173(2):191-201
To assess the canopy seed bank structure of Pinus halepensis, we measured the level of serotiny and the seed bank size and density of trees in unburned stands and post-fire regenerated stands in Israel. We analysed the effects of tree size, tree density and fire history on the level of serotiny. The level of serotiny decreased with an increase in tree height. The high level of serotiny in short trees could be explained by selection to increase regeneration chances after burning at pre-mature age. Also, limitation of long-distance seed dispersal opportunities in short trees may favour high serotiny levels. The level of serotiny was higher in post-fire stands than in unburned stands, suggesting a fast selection for serotiny by fire. Unburned stands had a higher total stand seed density than post-fire regenerated stands, but the proportion of seeds in serotinous cones of the total stand seed density was higher in post-fire regenerated stands. The fact that P. halepensis bears simultaneously serotinous and non-serotinous cones reflects its dual strategy as both a post-fire obligate seeder, mainly from serotinous cones and an early coloniser during fire-free periods, mainly from non-serotinous cones. The relative investment in these strategies is dependent on fire history and varies with tree height. Furthermore, mature brown cones can contribute to post-fire regeneration in case of spring fires, and serotinous cones are known to open partially also in dry spell events. Thus, post-fire regeneration and invasion are strategies, which seem to complement each other.  相似文献   

10.
The jack pine tip beetle, Conophthorus banksianae McPherson (Coleoptera: Scolytidae) and the red pine cone beetle, C. resinosae Hopkins, are doubtful sibling species. However, it is possible that these two taxa are valid species that maintain reproductive isolation because they accept different hosts. In a series of laboratory and field cage experiments, the host acceptance behaviours of these two species under choice and no choice conditions were compared. The field experiments demonstrated that the two species have a similar propensity to accept jack pine cones and shoots for feeding, but differ consistently in their acceptance of red pine cones, and variably in their acceptance of red pine shoots. However, the laboratory experiments did not indicate a difference between the two species in their propensity to accept red pine cones for feeding. In field cages, C. resinosae accepted significantly more red pine cones for oviposition than C. banksianae; the situation was reversed for jack pine shoots. In comparison to C. banksianae, C. resinosae is a more generalist feeder. The results from this study suggest that host acceptance behaviour is a permeable barrier unable to ensure reproductive isolation between the two species. Although there are differences in the host acceptance behaviours between C. banksianae and C. resinosae, we conclude that these differences do not necessarily support their designation as distinct species.  相似文献   

11.
12.
  • 1 Previous applications of the mating disruption technique to Neodiprion sertifer resulted in decreased numbers of males caught in the treated area but no effect on sex ratio or overall population density.
  • 2 The present study assessed the efficacy of mating disruption against N. sertifer outbreaks in pine stands surrounded by agricultural areas or pasture, and therefore isolated from other infested areas.
  • 3 Pine stands were treated by placing dispensers with an erythro‐mixture of the acetate ester of 3,7‐dimethyl‐2‐pentadecanol every 10 m in a grid.
  • 4 The efficacy of this technique was evaluated by comparing the number of males caught in sticky traps baited with synthetic pheromone, and by comparing the sex ratio and the population density of the sawfly in the subsequent generation between treated and control pine stands.
  • 5 The number of males caught within treated areas was significantly lower than in the control area.
  • 6 In the treated pine stands 46% of the egg clusters resulted in male cocoons only, compared to 3% of the clusters in the control stand.
  • 7 A significant reduction of the sawfly population was observed in the treated pine stands. The results contrast to the earlier mating disruption attempts with N. sertifer and can probably be ascribed to the isolation of the experimental stands.
  相似文献   

13.
Feeding behavior of third-instar bollworm, Helicoverpa zea (Boddie), and tobacco budworm, Heliothis virescens (F.), was observed in pure and mixed stands of nontransgenic and transgenic cotton (BTK), Gossypium hirsutum L., expressing an insecticidal protein CryIA(c) from a bacterium, Bacillus thuringiensis Berliner subsp. kurstaki. Five plant stands composed of BTK and non-BTK plants were evaluated; two pure stands and three mixed stands. Percentage ratios of BTK to non-BTK plants in the stands were 100:0, 75:25, 50:50, 25:75 and 0:100, respectively. In all stands with BTK plants, fewer bollworm and tobacco budworm larvae were found on BTK plants than non-BTK plants 24 h after infestation with third instars. At 48 h, significantly fewer tobacco budworm larvae, but not fewer bollworm larvae, were found on BTK plants. However, the number of larvae of either insect did not increase on non-BTK plants compared with the initial infestation density of three larvae per plant. The number of obacco budworm injured flower buds, and capsules was lower in all plant stands containing BTK plants compared with the pure stand of non-BTK at 48 h after infestation. Higher numbers of larvae on non-BTK plants were possibly the result of larval intoxication, reduced feeding, and increased plant abandonment and death on BTK plants rather than a classical feeding preference. Unexpectedly, the number of flower buds and capsules injured by bollworm and tobacco budworm when averaged per plant for all plants in a stand, differed little among the 75:25, 50:50 and 25:75 plant mixtures. These data suggest that larvae of both species frequently moved among plants, feeding indiscriminately on BTK and non-BTK plants.  相似文献   

14.
  • 1 The disruption of host‐finding cues has been proposed as a key mechanism underlying the lower damage caused by phytophagous insects in mixed forests. We tested this hypothesis by investigating the distribution of pine processionary moth Thaumetopoea pityocampa (Denis & Schiffer‐Müller) (Lepidoptera) infestation at the edges of pure stands of Pinus pinaster (AÏton) at some distance from nonhost trees (Experiment 1) or bordered in part by a broadleaved hedgerow (Experiment 2).
  • 2 An ‘edge effect' was demonstrated, with trees at the edge of the stand being more heavily infested than those at the interior of the stand.
  • 3 The presence of a nonhost broadleaved hedgerow in front of the edge of the pine stand resulted in lower T. pityocampa infestation. There were significantly fewer T. pityocampa nests behind the hedgerow than on the exposed part of the edge. The presence of the hedgerow did not dilute or repel T. pityocampa infestation further into the pine stand, although it decreased the infestation of T. pityocampa throughout the pine stand. The decrease in T. pityocampa infestation behind the hedgerow was greater when the broadleaved hedgerow was taller than the pine trees.
  • 4 These results highlight the benefits of using nonhost tree species on the edge of monospecific forest stands to reduce insect damage. This approach could be promoted as an innovative forest pest management method.
  相似文献   

15.
1. Coniferous trees deploy a combination of constitutive (pre‐existing) and induced (post‐invasion), structural and biochemical defences against invaders. Induced responses can also alter host suitability for other organisms sharing the same host, which may result in indirect, plant‐mediated interactions between different species of attacking organisms. 2. Current range and host expansion of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) from lodgepole pine‐dominated forests to the jack pine‐dominated boreal forests provides a unique opportunity to investigate whether the colonisation of jack pine (Pinus banksiana Lamb.) by MPB will be affected by induced responses of jack pine to a native herbaceous insect species: the jack pine budworm (Choristoneura pinus pinus Freeman; JPBW). 3. We simulated MPB attacks with one of its fungal associates, Grosmannia clavigera Robinson‐Jeffrey & Davidson, and tested induction of either herbivory by JPBW or inoculation with the fungus followed by a challenge treatment with the other organism on jack pine seedlings and measured and compared monoterpene responses in needles. 4. There was clear evidence of an increase in jack pine resistance to G. clavigera with previous herbivory, indicated by smaller lesions in response to fungal inoculations. In contrast, although needle monoterpenes greatly increased after G. clavigera inoculation and continued to increase during the herbivory challenge, JPBW growth was not affected, but JPBW increased the feeding rate to possibly compensate for altered host quality. 5. Jack pine responses varied greatly and depended on whether seedlings were treated with single or multiple organisms, and their order of damage.  相似文献   

16.
Population reduction and disturbances may alter dispersal, mating patterns and gene flow. Rather than taking the common approach of comparing different populations or sites, here we studied gene flow via wind‐mediated effective pollen dispersal on the same plant individuals before and after a fire‐induced population drop, in a natural stand of Pinus halepensis. The fire killed 96% of the pine trees in the stand and cleared the vegetation in the area. Thirteen trees survived in two groups separated by ~80 m, and seven of these trees had serotinous (closed) prefire cones that did not open despite the fire. We analysed pollen from closed pre and postfire cones using microsatellites. The two groups of surviving trees were highly genetically differentiated, and the pollen they produced also showed strong among‐group differentiation and very high kinship both before and after the fire, indicating limited and very local pollen dispersal. The pollen not produced by the survivors also showed significant prefire spatial genetic structure and high kinship, indicating mainly within‐population origin and limited gene flow from outside, but became spatially homogeneous with random kinship after the fire. We suggest that postfire gene flow via wind‐mediated pollen dispersal increased by two putative mechanisms: (i) a drastic reduction in local pollen production due to population thinning, effectively increasing pollen immigration through reduced dilution effect; (ii) an increase in wind speeds in the vegetation‐free postfire landscape. This research shows that dispersal can alleviate negative genetic effects of population size reduction and that disturbances might enhance gene flow, rather than reduce it.  相似文献   

17.
The pinewood nematode, Bursaphelenchus xylophilus, is the causative agent of pine wilt of Pinus thunbergii and P. densiflora in Japan. The nematode is vectored by cerambycid beetles of the genus Monochamus. It is inferred to have been introduced from North America early in the 1900s and then to have distributed in China, Korea, and Taiwan. Intensive and/or long-term studies of pine wilt systems have elucidated the pattern and mechanism of the nematode’s spread within a pine stand, dispersal of vector beetles, and spread pattern of pine wilt within a prefecture. The modeling of nematode spread over pine stands, which involves beetle reproduction within a pine stand, has been developing and should elucidate the factors influencing the rate at which the nematode range expands. In this review, we summarize the biologies of the nematode, beetle, and tree, and then characterize the spread of the nematode within a pine stand, locally over pine stands, and regionally over unit administrative districts. Local and regional spreading of the nematode is related primarily to long-distance dispersal by insect vectors and to the artificial transportation of pine logs infested with the nematode and its vector, respectively.  相似文献   

18.
  • 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
  • 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
  • 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
  • 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.
  相似文献   

19.
  • 1 Quantifying dispersal in predator–prey systems can improve our understanding of how these species interact in space and time, as well as their relative distributions across complex landscapes.
  • 2 We measured the dispersal abilities of three forest insects associated with red pine decline: the eastern five spined pine engraver Ips grandicollis (Coleoptera: Curculionidae), its main predator Thanasimus dubius (Coleoptera: Cleridae) and the basal stem and root colonizer Dendroctonus valens (Coleoptera: Curculionidae). We also examined the edge behaviours of these species and the predator Platysoma spp (Coleoptera: Histeridae) between red pine stands (habitat) and clearings (nonhabitat).
  • 3 Thanasimus dubius dispersed 12 times farther than its prey I. grandicollis, with 50% of predators dispersing farther than 1.54 km. This profound difference in dispersal behaviour between prey and predator may contribute to the clumped distribution of I. grandicollis.
  • 4 Most T. dubius and D. valens were confined in the pine forest, thus showing strong edge behaviour. This differed from I. grandicollis and Platysoma spp., which were commonly found in open areas adjacent to red pine plantations.
  • 5 The bark beetle I. grandicollis and one of its main predators, T. dubius, exhibited different patterns of movement within a fragmented landscape. Despite a greater dispersal ability of T. dubius within forests, the spatial distribution of this predator may be restricted by fragmentation of its habitat, and provide an opportunity for partial escape of its prey.
  • 6 The present study contributes to our knowledge of top‐down forces within red pine stands undergoing decline. Differences of dispersal patterns and edge behaviour could contribute to the initiation of new pockets of decline, as well as the connectedness among existing ones.
  相似文献   

20.
Carbon sequestration in boreal jack pine stands following harvesting   总被引:2,自引:0,他引:2  
A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO2 exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m?2 yr?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m?2 yr?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.  相似文献   

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