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.     

Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.     

Stand Structural Dynamics of North American Boreal Forests

Stand structure, the arrangement and interrelationships of live and dead trees, has been linked to forest regeneration, nutrient cycling, wildlife habitat, and climate regulation. The objective of this review was to synthesize literature on stand structural dynamics of North American boreal forests, addressing both live tree and coarse woody debris (CWD) characteristics under different disturbance mechanisms (fire, clearcut, wind, and spruce budworm), while identifying regional differences based on climate and surficial deposit variability. In fire origin stands, both live tree and CWD attributes are influenced initially largely by the characteristics of the stand replacing fire and later increasingly by autogenic processes. Differences in stand structure have also been observed between various stand cover types. Blowdown and insect outbreaks are two significant non-stand replacing disturbances that can alter forest stand structure through removing canopy trees, freeing up available growing space, and creating microsites for new trees to establish. Climate and surficial deposits are highly variable in the boreal forest due to its extensive geographic range, influencing stand and landscape structure by affecting tree colonization, stand composition, successional trajectories, CWD dynamics, and disturbance regimes including regional fire cycles. Further, predicted climate change scenarios are likely to cause regional-specific alterations in stand and landscape structure, with the implications on ecosystem components including wildlife, biodiversity, and carbon balance still unclear. Some stand structural attributes are found to be similar between clearcut and fire origin stands, but others appear to be quite different. Future research shall focus on examining structural variability under both disturbance regimes and management alternatives emulating both stand replacing and non-stand replacing natural disturbances.

     

Microsite variation and climate result in stand dynamics variation in pristine Pinus sylvestris mires: Evidence from a stand structure study

Questions: How do climate conditions and the site's ecohy‐ drological properties affect the age and size structure of natural Pinus sylvestris stands on pristine boreal mires? How do the long‐term stand dynamics on mires proceed as stands age? Do the mire stands reach a balanced, old‐growth stage? Location: Boreal mire forests in southern and northern Finland. Methods: Tree age and diameter distributions were analysed in 52 stands in two climate areas and in two mire site types with different ecohydrological properties. Temporal stand dynamics were examined by (1) comparing the graphs of the stands’ mean tree ages by diameter at breast height (1.3 m) classes and (2) describing the changes in stand characteristics and stand age and size structures as a function of stand dominant age in a chronosequence. Results: In the south, the DBH distributions were mostly unimodal and bell‐shaped in both site type groups. Age distributions were multimodal and flat in fully‐stocked sites but more uneven in sparsely forested composite sites. In the north, both the age and size distributions were clearly uneven in both site type groups. Tree age and size variation increased with stand age, but levelled out in the long term. Particularly in the south, the abundance of small trees decreased as stand age increased. Conclusions: The pine stands on pristine boreal mires are more dynamic than anticipated and are generally not characterised by a balanced, self‐perpetuating structure. Their dynamics reflect differences in climate and ecohydrology: on stocked sites in favourable boreal conditions, the stands showed structures typically resultant of inter‐tree competition processes that control tree growth and regeneration, whereas in harsh boreal climates, the tree regeneration process is ongoing diversifying the stand structure.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

The direct regeneration hypothesis in northern forests

Question: Can the direct regeneration hypothesis (DRH) be used to predict post‐disturbance regeneration after fire, wind disturbance, and clearcutting in northern forests? Do life‐history traits such as regeneration strategy and shade tolerance influence post‐disturbance regeneration success of tree species? Location: Northern forests in North America. Methods: A meta‐analysis was conducted by collecting published data on pre‐ and post‐disturbance stand compositional characteristics in the northern forests. For each tree species, compositional difference (CD) was calculated as the difference between basal area proportions of the post‐ and pre‐disturbance stands, but for post‐disturbance stands <25 years of age, post‐disturbance proportions were calculated based on relative stem density. Results: Species response to disturbances was best explained by regeneration strategy, while disturbance type had no effect on CD. The proportion of broadleaf trees with either strong or weak vegetative reproduction ability increased after all disturbances. Serotinous species had CD values not significantly different from zero after fire, while CD for semi‐serotinous species was negative. The post‐disturbance proportions of non‐serotinous conifers decreased after all forms of disturbance. Conclusions: All disturbances promote broadleaf trees, regardless of regeneration strategy (suckering, sprouting, or seeding). The DRH is supported for conifers with serotinous cones after fire. Fire causes local extinction of non‐serotinous conifers, while wind and clearcutting only decrease the proportion of non‐serotinous conifers because of partial survival of seed sources and advanced regeneration. This study suggests that increasing stand‐replacing disturbances associated with global climate change will promote broadleaf trees in northern forests.     

Disturbance legacies and climate jointly drive tree growth and mortality in an intensively studied boreal forest

Most North American forests are at some stage of post‐disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short‐term studies in a longer term context, and particularly important for the carbon‐dense, fire‐prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine‐learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999–2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr^?1, with most mortality occurring in medium‐sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current‐ and previous‐year data exerted significant effects. Models based on these variables explained 23–44% of the ring‐width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old‐growth stands in a disturbance‐prone landscape, and the sustainable management of regional forests in a changing climate.     

Effects of fire and spruce beetle outbreak legacies on the disturbance regime of a subalpine forest in Colorado

Aim There is increasing research attention being given to the role of interactions among natural disturbances in ecosystem processes. We studied the interactions between fire and spruce beetle (Dendroctonus rufipennis Kirkby) disturbances in a Colorado subalpine forest. The central questions of this research were: (1) How does fire history influence stand susceptibility to beetle outbreak? And conversely, (2) How does prior occurrence of a beetle outbreak influence stand susceptibility to subsequent fire? Methods We reconstructed the spatial disturbance history in a c. 4600 ha area by first identifying distinct patches in the landscape on aerial photographs. Then, in the field we determined the disturbance history of each patch by dating stand origin, fire scars, dates of mortality of dead trees, and releases on remnant trees. A geographical information system (GIS) was used to overlay disturbance by fire and spruce beetle. Results and main conclusions The majority of stands in the study area arose following large, infrequent, severe fires occurring in c. 1700, 1796 and 1880. The study area was also affected by a severe spruce beetle outbreak in the 1940s and a subsequent low‐severity fire. Stands that originated following stand‐replacing fire in the late nineteenth century were less affected by the beetle outbreak than older stands. Following the beetle outbreak, stands less affected by the outbreak were more affected by low‐severity fire than stands more severely affected by the outbreak. The reduced susceptibility to low‐severity fire possibly resulted from increased moisture on the forest floor following beetle outbreak. The landscape mosaic of this subalpine forest was strongly influenced by the interactions between fire and insect disturbances.     

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.     

Tree mortality episodes in the intact Picea abies‐dominated taiga in the Arkhangelsk region of northern European Russia

Question: What were the temporal patterns and rates of tree mortality in a recent episodic tree mortality event? Have similar events occurred in the past, and does climatic variability play a role in the disturbance regime? Location: Intact Picea abies‐dominated taiga in the Arkhangelsk region, northwestern Russia. Methods: We reconstructed the past tree mortality and disturbance history by applying dendroecological methods in five forest stands and related these to climatic data. The role of other potential causes of tree mortality was assessed in a field inventory. Results: The recent episode lasted from 1999 to 2004, influenced all stands studied, and killed on average 21% of trees with a diameter of over 10 cm at 1.3‐m height. The annual tree mortality rate in the decades preceding this episode was 0.49%. During the past 200 years, the stands have experienced chronic small‐scale disturbances, with several irregular disturbances of moderate severity. The recent episode was associated with abundant signs of the bark beetle Ips typographus. Furthermore, the timing of both the recent tree mortality episode and the past disturbance events was associated with dry summers. Conclusion: The results indicate a connection between climatic variability and forest dynamics, the likely driving factors being droughts and bark beetles. In the context of the past 200 years, the recent episode was potentially at the higher end of the range of disturbance variability in terms of severity and spatial extent. This has ecological implications in a changing climate, potentially influencing ecosystem structure and long‐term dynamics.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality

Questions: How does woody vegetation abundance and diversity differ after natural disturbances causing different levels of mortality? Location: Abies balsamea–Betula papyrifera boreal mixed‐wood stands of southeast Quebec, Canada. Methods: Woody vegetation abundance and diversity were quantified and compared among three disturbance‐caused mortality classes, canopy gap, moderate‐severity disturbances, and catastrophic fire, using redundancy analysis, a constrained linear ordination technique, and diversity indices. Results: Substantial changes in canopy tree species abundance and diversity only occurred after catastrophic fire. Shade‐tolerant, late‐successional conifer species remained dominant after canopy gap and moderate‐severity disturbances, whereas shade‐intolerant, early‐successional colonizers dominated canopy tree regeneration after catastrophic fire. Density and diversity of mid‐tolerant and shade‐intolerant understory tree and shrub species increased as the impact of disturbance increased. Highest species richness estimates were observed after catastrophic fire, with several species establishing exclusively under these conditions. Relative abundance of canopy tree regeneration was most similar after canopy gap and moderate‐severity disturbances. For the sub‐canopy tree and shrub community, relative species abundances were most similar after moderate‐severity disturbances and catastrophic fire. Vegetation responses to moderate‐severity disturbances thus had commonalities with both extremes of the disturbance‐caused mortality gradient, but for different regeneration layers. Conclusions: Current spatio‐temporal parameters of natural disturbances causing varying degrees of mortality promote the development of a complex, multi‐cohort forest condition throughout the landscape. The projected increase in time intervals between catastrophic fires may lead to reduced diversity within the system.     

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.     

Succession after stand replacing disturbances by fire,wind throw,and insects in the dark Taiga of Central Siberia

The dark taiga of Siberia is a boreal vegetation dominated by Picea obovata, Abies sibirica, and Pinus sibirica during the late succession. This paper investigates the population and age structure of 18 stands representing different stages after fire, wind throw, and insect damage. To our knowledge, this is the first time that the forest dynamics of the Siberian dark taiga is described quantitatively in terms of succession, and age after disturbance, stand density, and basal area. The basis for the curve–linear age/diameter relation of trees is being analyzed. (1) After a stand-replacing fire Betula dominates (4,000 trees) for about 70 years. Although tree density of Betula decreases rapidly, basal area (BA) reached >30 m²/ha after 40 years. (2) After fire, Abies, Picea, and Pinus establish at the same time as Betula, but grow slower, continue to gain height and eventually replace Betula. Abies has the highest seedling number (about 1,000 trees/ha) and the highest mortality. Picea establishes with 100–400 trees/ha, it has less mortality, but reached the highest age (>350 years, DBH 51 cm). Picea is the most important indicator for successional age after disturbance. Pinus sibirica is an accompanying species. The widely distributed “mixed boreal forest” is a stage about 120 years after fire reaching a BA of >40 m²/ha. (3) Wind throw and insect damage occur in old conifer stands. Betula does not establish. Abies initially dominates (2,000–6,000 trees/ha), but Picea becomes dominant after 150–200 years since Abies is shorter lived. (4) Without disturbance the forest develops into a pure coniferous canopy (BA 40–50 m²/ha) with a self-regenerating density of 1,000 coniferous canopy trees/ha. There is no collapse of old-growth stands. The dark taiga may serve as an example in which a limited set to tree species may gain dominance under certain disturbance conditions without ever getting monotypic.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

Carbon distribution of a well- and poorly-drained black spruce fire chronosequence

The objective of this study was to quantify carbon (C) distribution for boreal black spruce (Picea mariana (Mill.) BSP) stands comprising a fire chronosequence in northern Manitoba, Canada. The experimental design included seven well‐drained (dry) and seven poorly‐drained (wet) stands that burned between 1998 and 1850. Vegetation C pools (above‐ground + below‐ground) steadily increased from 1.3 to 83.3 t C ha^?1 for the dry chronosequence, and from 0.6 to 37.4 t C ha^?1 for the wet chronosequence. The detritus C pools (woody debris + forest floor) varied from 10.3 to 96.0 t C ha^?1 and from 12.6 to 77.4 t C ha^?1 for the dry and wet chronosequence, respectively. Overstorey biomass, mean annual biomass increment (MAI), woody debris mass, and litterfall were significantly greater (α = 0.05) for the dry stands than for the wet stands, but the bryophyte, understorey, and forest floor C pools were significantly less for the dry than for the wet stands. The root mass ratio decreased with stand age until 37 years after fire, was fairly constant thereafter, and was not significantly affected by soil drainage. The C pools of the overstorey and bryophyte tended to increase with stand age. Foliage biomass, litterfall, and MAI (for the dry stands) peaked at 71 years after fire and declined in the oldest stands. The results from this study illustrate that the effects of disturbance and edaphic conditions must be accounted for in boreal forest C inventories and C models. The appropriateness of using chronosequences to examine effects of wildfire on ecosystem C distribution is discussed.     

The Contribution of Litterfall to Net Primary Production During Secondary Succession in the Boreal Forest

Litterfall is a fundamental process in the nutrient cycle of forest ecosystems and a major component of annual net primary production (NPP). Despite its importance for understanding ecosystem energetics and carbon accounting, the dynamics of litterfall production following disturbance and throughout succession remain poorly understood in boreal forest ecosystems. Using a replicated chronosequence spanning 209 years following fire and 33 years following logging in Ontario, Canada, we examined the dynamics of litterfall production associated with stand development, overstory composition type (broadleaf, mixedwood, and conifer), and disturbance origin. We found that total annual litterfall production increased with stand age following fire and logging, plateauing in post-fire stands approximately 98 years after fire. Neither total annual litterfall production nor any of its constituents differed between young fire- or logging-originated stands. Litterfall production was generally higher in broadleaf stands compared with mixedwood and conifer stands, but varied seasonally, with foliar litterfall highest in broadleaf stands in autumn, and epiphytic lichen litterfall highest in conifer stands in spring. Contrary to previous assumptions, we found that the contribution of litterfall production to net primary production increased with stand age, highlighting the need for modeling studies of net primary productivity to account for the effects of stand age on litterfall dynamics.     

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha^‐1) than either Populus (25 500 kg‐C.ha^‐1) or Pinus (19 400 kg‐C.ha^‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.