Origin of the lichen–spruce woodland in the closed-crown forest zone of eastern Canada

Aim We investigate the timing and factors responsible for the transformation of closed‐crown forests into lichen–spruce woodlands. Location The study area extends between 70° and 72° W in the closed‐crown forest zone from its southern limit near 47°30′ N to its northern limit at the contact with the lichen–spruce woodland zone around 52°10′ N. A total of 24 lichen–spruce woodlands were selected. Methods Radiocarbon dating of charcoals at mineral soil contact and within the organic horizons allowed the principal factors causing the degradation of the closed‐crown forest to be identified, i.e. light fires, successive fires and the occurrence of a spruce budworm epidemic followed by a fire. Results Charcoals dated in the organic horizon were less than 200 years old, suggesting a recent transformation of the closed‐crown forest following surface fires. Before their transformation into lichen–spruce woodlands, stands were occupied by old, dense forests that originated from fires dating back to 1000 yr bp . The radiocarbon dating of charcoals in the organic horizon indicated that several stands burned twice in less than 50 years, while others burned shortly after a spruce budworm epidemic. Light fires are frequent within the lichen–spruce woodlands according to multiple charcoal layers found within the organic matter horizon. Main conclusions While closed‐crown forests are predicted to expand under climate warming, compound disturbances diminish the natural regeneration of the closed‐crown forests in the south and favour the expansion of lichen–spruce woodlands. As black spruce germinates on mineral soils, surface fires accentuate the expansion of the lichen–spruce woodlands southward. Under global warming, warmer springs will lead to earlier low‐intensity fires that do not remove as much organic matter, and hence prevent conditions suitable for black spruce regeneration. Also, spruce budworm reduces seed production for a certain time. The occurrence of fire during this period is critical for regeneration of black spruce.     

Rapid expansion of lichen woodlands within the closed-crown boreal forest zone over the last 50 years caused by stand disturbances in eastern Canada

Aim Our two main goals are first to evaluate the resilience of the boreal forest according to latitude across the closed‐crown forest zone using the post‐disturbance distribution and cover of lichen woodlands and closed‐crown forests as a metric, and second to identify the disturbance factors responsible for the regeneration and degradation of the closed‐crown forest according to latitude since the 1950s. Location The study area extends between 70°00′ and 72°00′ W and throughout the closed‐crown forest zone, from its southern limit near 47°30′ N to its northern limit at the contact with the lichen woodland zone at around 52°40′ N. Methods Recent (1972–2002) and old (1954–1956) aerial photos were used to map the distribution of lichen woodlands across the closed‐crown forest zone. Forest disturbances such as fire, spruce budworm (Choristoneura fumiferana (Clemens)) outbreak, and logging were recorded on each set of aerial photos. Each lichen woodland and stand disturbance was validated by air‐borne surveys and digitized using GIS software. Results Over the last 50 years, the area occupied by lichen woodlands has increased according to latitude; that is, 9% of the area that was occupied by closed‐crown forests has shifted to lichen woodlands. Although logging activities have been concentrated in the same areas during the last 50 years, the area covered by logging has increased significantly. Outbreaks by the spruce budworm occurred predominantly in the southern (47°30′ N to 48°30′ N) and central (48°53′ N to 50°42′ N) parts of the study area, where balsam fir stands are extensive. In the northern part of the study area (51°–52°40′ N), extensive fires affected the distribution and cover of closed‐crown forests and lichen woodlands. Main conclusions Over the last 50 years, the area occupied by closed‐crown forests has decreased dramatically, and the ecological conditions that allow closed‐crown forests to establish and develop are currently less prevalent. Fire is by far the main disturbance, reducing the ability of natural closed‐crown forests to self‐regenerate whatever the latitude. Given the current biogeographical shift from dense to open forests, the northern part of the closed‐crown forest zone is in a process of dramatic change towards the dominance of northern woodlands.     

Spatial and temporal variation in historic fire regimes in subalpine forests across the Colorado Front Range in Rocky Mountain National Park, Colorado, USA

Aim The historical variability of fire regimes must be understood in the context of drivers of the occurrence of fire operating at a range of spatial scales from local site conditions to broad‐scale climatic variation. In the present study we examine fire history and variations in the fire regime at multiple spatial and temporal scales for subalpine forests of Engelmann spruce–subalpine fir (Picea engelmannii, Abies lasiocarpa) and lodgepole pine (Pinus contorta) of the southern Rocky Mountains. Location The study area is the subalpine zone of spruce–fir and lodgepole pine forests in the southern sector of Rocky Mountain National Park (ROMO), Colorado, USA, which straddles the continental divide of the northern Colorado Front Range (40°20′ N and 105°40′ W). Methods We used a combination of dendroecological and Geographic Information System methods to reconstruct fire history, including fire year, severity and extent at the forest patch level, for c. 30,000 ha of subalpine forest. We aggregated fire history information at appropriate spatial scales to test for drivers of the fire regime at local, meso, and regional scales. Results The fire histories covered c. 30,000 ha of forest and were based on a total of 676 partial cross‐sections of fire‐scarred trees and 6152 tree‐core age samples. The subalpine forest fire regime of ROMO is dominated by infrequent, extensive, stand‐replacing fire events, whereas surface fires affected only 1–3% of the forested area. Main conclusions Local‐scale influences on fire regimes are reflected by differences in the relative proportions of stands of different ages between the lodgepole pine and spruce–fir forest types. Lodgepole pine stands all originated following fires in the last 400 years; in contrast, large areas of spruce–fir forests consisted of stands not affected by fire in the past 400 years. Meso‐scale influences on fire regimes are reflected by fewer but larger fires on the west vs. east side of the continental divide. These differences appear to be explained by less frequent and severe drought on the west side, and by the spread of fires from lower‐elevation mixed‐conifer montane forests on the east side. Regional‐scale climatic variation is the primary driver of infrequent, large fire events, but its effects are modulated by local‐ and meso‐scale abiotic and biotic factors. The low incidence of fire during the period of fire‐suppression policy in the twentieth century is not unique in comparison with the previous 300 years of fire history. There is no evidence that fire suppression has resulted in either the fire regime or current forest conditions being outside their historic ranges of variability during the past 400 years. Furthermore, in the context of fuel treatments to reduce fire hazard, regardless of restoration goals, the association of extremely large and severe fires with infrequent and exceptional drought calls into question the future effectiveness of tree thinning to mitigate fire hazard in the subalpine zone.     

Structural characteristics and aboveground biomass of old‐growth spruce–fir stands in the eastern Carpathian mountains,Ukraine

Abstract

Temperate old‐growth forests are known to have ecological characteristics distinct from younger forests, but these have been poorly described for the remaining old‐growth Picea abies–Abies alba forests in the eastern Carpathian mountains. In addition, recent studies suggest that old‐growth forests may be more significant carbon sinks than previously recognized. This has stimulated interest in quantifying aboveground carbon stocks in primary forest systems. We investigated the structural attributes and aboveground biomass in two remnant old‐growth spruce–fir stands and compared these against a primary (never logged) mature reference stand. Our sites were located in the Gorgany Nature Reserve in western Ukraine. Overstory data were collected using variable radius plots; coarse woody debris was sampled along line intercept transects. Differences among sites were assessed using non‐parametric statistical analyses. Goodness‐of‐fit tests were used to evaluate the form of diameter distributions. The results strongly supported the hypothesis that old‐growth temperate spruce–fir forests have greater structural complexity compared to mature forests, including higher densities of large trees, more complex horizontal structure, and elevated aboveground biomass. The late‐successional sites we sampled exhibited rotated sigmoid diameter distributions; these may reflect natural disturbance dynamics. Old‐growth Carpathian spruce–fir forests store on average approximately 155–165 Mg ha^?1 of carbon in aboveground tree parts alone. This is approximately 50% higher than mature stands. Given the scarcity of primary spruce–fir forests in the Carpathian region, remaining stands have high conservation value, both as habitat for late‐successional species and as carbon storage reservoirs.     

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.     

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.     

Hypocrealean (Hypocreales,Ascomycota) Fungal Diversity in Different Stages of Tropical Forest Succession in Costa Rica1

The relationship between forest succession and microfungal diversity has been poorly studied. Fungi provide important ecosystem services that may deteriorate in deforested or highly disturbed forests. To determine the possible effects of deforestation and forest succession on microfungi, species diversity of hypocrealean fungi (Ascomycota) was compared in forest stands in Eastern Costa Rica representing three stages of succession: 1–2, 25–27 yr old, and an old growth forest. Species diversity in a second‐growth forest fragment surrounded by timber plantations and second‐growth forest was also compared to that of a stand surrounded by old growth forest. The results show that the overall diversity of hypocrealean fungi was inversely proportional to the age of the forest stand, and each family showed different successional trends. Clavicipitaceae was more diverse in the old‐growth forest and was positively related to the age of the forest stand. Nectriaceae was highly diverse in the 1‐ to 2‐yr‐old stand and less diverse in the old‐growth stand. Saprobic and plant pathogenic fungal species were more diverse in the 1‐ to 2‐yr‐old stand and their diversity was inversely proportional to the age of the forest stand. The diversity of insect pathogens was positively related to the age of the forest stand. The 20‐ to 22‐yr‐old forest fragment had the lowest number of species overall. Based on the data gathered in this study, hypocrealean fungal species diversity is related to the successional stage and fragmentation of tropical forest.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.     

Recent climatic drying leads to age‐independent growth reductions of white spruce stands in western Canada

Since 2001, climatic conditions have been notably drier than normal across large areas of the western Canadian interior, leading to widespread impacts on the forests of this region. This poses a major concern for the future, given climate change projections for continued warming and drying. We conducted tree‐ring analysis in 75 pure stands of white spruce (Picea glauca) across Alberta and west‐central Saskatchewan to examine the effects of recent climatic drying on the growth of this important boreal tree species. Allometric equations were used to calculate annual growth in aboveground tree biomass (G_BM) from ring width measurements. Results showed an increasing trend in G_BM from the 1960s to the 1990s, followed by a sharp decline during the severe drought of 2001–2002. Of the 75 stands, only 18 recovered sufficiently to cause an increase in mean G_BM from the predrought decade of 1991–2000 to the subsequent decade of 2001–2010. The remaining 57 stands exhibited a decline in mean G_BM between these decades. Climatic drying was a major cause of the growth decline, as shown by the significant stand‐level relationship between percentage change in decadal mean G_BM and the change in decadal mean values of a climate moisture index from 1991–2000 to 2001–2010. Subsequent analyses of boreal stands sampled across Alberta during 2015 revealed that white spruce growth had declined even further as drought conditions intensified during 2014–2015. Overall, there was a 38% decrease in mean G_BM between 1997 and 2015, but surprisingly, the percentage decrease was not significantly different for young, productive stands compared with older, less productive stands. Thus, stand ageing cannot explain the observed decline in white spruce growth during the past quarter century, suggesting that these forests are at risk if the trend towards more frequent, severe drought continues in the region.     

Contrasting patterns of tree mortality in late‐successional Picea abies stands in two areas in northern Fennoscandia

Question: What are tree mortality rates and how and why do they vary in late‐successional Picea abies‐dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long‐term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross‐dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even‐aged and originated following a stand‐replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter‐annual variation, which we attributed to senescence of the relatively even‐aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand‐replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small‐scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.     

Fifty‐seven years of composition change in the eastern boreal forest of Canada

Question: In the boreal forest of eastern Canada, how does forest vegetation change in the sustained absence of fire? Location: Eastern boreal forest in Quebec's North Shore region, Canada (49°30′–50°00′N; 67°30′–68°35′W). Methods: Aerial photos from three different periods (1930, 1965 and 1987) were used to characterize changes in vegetation composition in 23 scenes of 200 ha. Time since fire, presence of secondary disturbances and data on soil and topographic variables were obtained. Ordination and clustering techniques were used to define compositional trajectories of change over the 57‐yr period. These trajectories were further grouped into pathways based on compositional changes, time since fire and preferential deposit‐drainage types. Results: Among the 26 compositional trajectories, three successional pathways were distinguished. Two start post‐fire succession with a dominance of intolerant hardwood. In one of these, this is followed by an increase in Abies balsamea, while in the second the importance of Picea mariana increases with time. In the third pathway P. mariana is an important component from the outset. In this pathway, we observed modest fluctuation in the relative dominance of P. mariana and A. balsamea and variation in stand structure. Conclusion: The boreal forest vegetation of Eastern Canada is diverse and dynamic even in the absence of fire, notably under the influence of partial disturbances. Such disturbances can be associated with changes in composition or stand structure. The development of management strategies aimed at maintaining stand diversity by emulating a broader variety of partial and secondary disturbances should be encouraged.     

Post‐fire vegetation and climate dynamics in low‐elevation forests over the last three millennia in Yellowstone National Park

Conifer forests of the western US are historically well adapted to wildfires, but current warming is creating novel disturbance regimes that may fundamentally change future forest dynamics. Stand‐replacing fires can catalyze forest reorganization by providing periodic opportunities for establishment of new tree cohorts that set the stage for stand development for centuries to come. Extensive research on modern and past fires in the Northern Rockies reveals how variations in climate and fire have led to large changes in forest distribution and composition. Unclear, however, is the importance of individual fire episodes in catalyzing change. We used high‐resolution paleoecologic and paleoclimatic data from Crevice Lake (Yellowstone National Park, Wyoming, USA), to explore the role of fire in driving low‐elevation forest dynamics over the last 2820 yr. We addressed two questions: 1) did low‐elevation forests at Crevice Lake experience abrupt community‐level vegetation changes in response to past fire events? 2) Did the interaction of short‐term disturbance events (fire) and long‐term climate change catalyze past shifts in forest composition? Over the last 2820 yr, we found no evidence for abrupt community‐level vegetation transitions at Crevice Lake, and no evidence that an interaction of climate and fire produced changes in the relative abundance of dominant plant taxa. In part, this result reflects limitations of the datasets to detect past event‐specific responses and their causes. Nonetheless, the relative stability of the vegetation to fires over the last 2820 yr provides a local baseline for assessing current and future ecological change. Observations of climate–fire–vegetation dynamics in recent decades suggest that this multi‐millennial‐scale baseline may soon be exceeded.     

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.     

Red Spruce Stand Dynamics, Simulations, and Restoration Opportunities in the Central Appalachians

Red spruce (Picea rubens)–dominated forests occupied as much as 600,000 ha in West Virginia prior to exploitive logging era of the late nineteenth and early twentieth centuries. Subsequently, much of this forest type was converted to northern hardwoods. As an important habitat type for a number of rare or sensitive species, only about 12,000 ha of red spruce forests presently remain in the state. In order to assess the prospects for restoration, we examined six northern hardwood stands containing understory red spruce to (1) characterize stand dynamics and regeneration patterns and (2) simulate the effectiveness of restoration silviculture to enhance red spruce overstory recruitment. Stands originated in the late 1800s to early 1900s and are currently in the (late) stem exclusion or understory reinitiation stages. Five of the six stands had even‐aged overstories that originated after clear‐cutting. Tree‐ring chronologies show high initial growth rates consistent with stand initiation. One stand, partially harvested in 1915, was uneven aged with older, legacy residuals in the canopy. Most stands had two cohorts of understory red spruce, with more than 40% of these individuals showing prior release. Our 100‐year growth simulation suggested that a 50% basal area thinning from above could double red spruce basal area to support a mixed spruce–hardwood stand in approximately 20–40 years. These results indicate that restoration silviculture could be an effective tool for increasing the amount and quality of this reduced forest type in the central Appalachians.     

The soil fauna community in pure and mixed stands of beech and spruce of different age: trophic structure and structuring forces

This study investigates the response of the soil fauna community to replacement of beech by spruce or by mixed stands of beech and spruce. Stands of different age were investigated in a factorial design with the factors tree species (beech and spruce) and stand age (30 and 120 yr). The input of leaf/needle litter did not differ significantly between the study sites. By contrast, the amount of organic matter in upper soil layers (L/F, H/Ah) of spruce forests strongly exceeded that of beech forests particularly in mature stands. The increase in organic matter in spruce stands was not associated by an increase in the amount of microbial biomass. Biomass of eight (bacterivorous, fungivorous and omnivorous nematodes, enchytraeids, earthworms, isopodes, mycetophilid and cecidomyiid Diptera) of the twelve microbi‐detritivorous soil animal groups studied was significantly increased in beech forests; only that of one group (elaterid beetles) was increased in spruce forests and three groups did not respond significantly (collembolans, oribatid mites, sciarid Diptera). This indicates that in the forests studied neither habitat space (amount of organic matter in L/F and H/Ah layers) nor the amount of microbial biomass controlled microbi‐detritivores. Rather, the quality of litter materials and the concentration of microbial biomass therein appeared to be most important. Herbivores and predators also were favoured by beech: the biomass of one (rhizophagous nematodes) of the three herbivorous groups studied were significantly increased in beech stands and none in spruce stands; the biomass of four (predatory nematodes, centipedes, carabid and cantharid beetles) of the seven carnivorous groups studied were increased in beech stands, none in spruce stands. Generally, the biomass ratio between prey and predators was at a minimum in mature beech and mixed stands indicating more intense top‐down control in these forests. Overall, the study documents that replacement of beech by spruce strongly alters the soil food web. Mixed stands were more similar to spruce stands in respect to the biomass of soil animal groups but predator–prey interactions appeared to be more similar in mature beech and mixed stands. Differences between tree species usually were more pronounced in 120 compared to 30 yr old stands indicating that the development of stand characteristics is slow.     

Spatial and temporal variations in boreal forest fire frequency in northern Alberta

Abstract. Spatial and temporal variations in fire frequency in the boreal forest of Wood Buffalo National Park (WBNP) were assessed using forest stand age, fire scar and historical data. I test the hypotheses that (1) fire frequency is higher in jack pine forests and aspen forests than in black spruce forests and white spruce forests, (2) these variations in fire frequency can be related to the mean waterbreak distance (MWD) around a site and (3) fire frequency has changed over the past 300 years. The fire cycles (the time required to burn an area equal in size to the entire study area) in jack pine forests (39 years) and in aspen forests (39 years) were significantly shorter than those in black spruce forests (78 years) and in white spruce forests (96 years). The length of the fire cycle varies inversely with the MWD around a site, and the MWD was significantly higher in jack pine and aspen forests than in black or white spruce forests. It is suggested that covariations between soil type and the MWD influence, respectively, variations in forest dominant and fire frequency. A change in fire frequency at 1860 was apparent in the fire history for all of WBNP, the black spruce dominated stands, and the near and medium MWD classes. The fire cycle estimates for these classes were all significantly shorter during the period 1750 to 1859 (fire cycles = 25–49 years) than they were in the period 1860 to 1989 (fire cycles = 59–89 years). The possible roles of changes in climate and aboriginal burning practices in causing the temporal change in fire frequency are discussed.     

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.     

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.     

Response of broadleaved evergreen Mediterranean forest vegetation to fire disturbance during the Holocene: insights from the peri-Adriatic region

Aim To test whether fire contributed to the expansion and compositional change of evergreen forests in the Mediterranean region during the Holocene. Location The peri‐Adriatic region, encompassing the Italian peninsula, Sicily and the western and southern Balkans between latitudes 46° and 37° N. Methods New high‐resolution pollen and microscopic charcoal data from Lago dell’Accesa (Tuscany, Italy) were used to estimate the response of the evergreen oak, Quercus ilex L., to fire during its expansion phase at 8500 cal. yr bp . The data were compared with the pollen and charcoal series from other Mediterranean sites (Lago di Massaciuccoli in Tuscany, Malo Jezero in Croatia, Biviere di Gela in Sicily) and analysed using numerical techniques (redundancy analysis, detrended canonical correspondence analysis) to identify long‐term fire–vegetation linkages and the degree of compositional change. Results Microscopic charcoal and pollen of evergreen oaks were negatively correlated during the period of quasi‐natural fire regime (Mesolithic, 10,000–8000 cal. yr bp ). In addition, there was no such positive correlation during periods when the fire regime was potentially more influenced by people (Neolithic–Bronze Age, 8000–3000 cal. yr bp ). Compared with inland sites, coastal sites that are currently located at a distinct ecotone showed more compositional change. Main conclusions The analyses suggest that climatic change, without an additional effect of fire regimes, favoured the expansion and compositional change of evergreen forests across the peri‐Adriatic region. Strikingly different patterns occurred along a north–south gradient. In the north (Tuscany and Croatia, meso‐Mediterranean belt), Q. ilex replaced deciduous forests when conditions became drier; in the south (Sicily, thermo‐Mediterranean belt) the species replaced maquis or steppe vegetation when climatic conditions became moister. We conclude that the projected increase in fire activity may lead to the loss of most of the remaining relict forests of Q. ilex in southern Europe.     

Fire severity mediates climate‐driven shifts in understorey community composition of black spruce stands of interior Alaska

Question: How do pre‐fire conditions (community composition and environmental characteristics) and climate‐driven disturbance characteristics (fire severity) affect post‐fire community composition in black spruce stands? Location: Northern boreal forest, interior Alaska. Methods: We compared plant community composition and environmental stand characteristics in 14 black spruce stands before and after multiple, naturally occurring wildfires. We used a combination of vegetation table sorting, univariate (ANOVA, paired t‐tests), and multivariate (detrended correspondence analysis) statistics to determine the impact of fire severity and site moisture on community composition, dominant species and growth forms. Results: Severe wildfires caused a 50% reduction in number of plant species in our study sites. The largest species loss, and therefore the greatest change in species composition, occurred in severely burned sites. This was due mostly to loss of non‐vascular species (mosses and lichens) and evergreen shrubs. New species recruited most abundantly to severely burned sites, contributing to high species turnover on these sites. As well as the strong effect of fire severity, pre‐fire and post‐fire mineral soil pH had an effect on post‐fire vegetation patterns, suggesting a legacy effect of site acidity. In contrast, pre‐fire site moisture, which was a strong determinant of pre‐fire community composition, showed no relationship with post‐fire community composition. Site moisture was altered by fire, due to changes in permafrost, and therefore post‐fire site moisture overrode pre‐fire site moisture as a strong correlate. Conclusions: In the rapidly warming climate of interior Alaska, changes in fire severity had more effect on post‐fire community composition than did environmental factors (moisture and pH) that govern landscape patterns of unburned vegetation. This suggests that climate change effects on future community composition of black spruce forests may be mediated more strongly by fire severity than by current landscape patterns. Hence, models that represent the effects of climate change on boreal forests could improve their accuracy by including dynamic responses to fire disturbance.