Tree‐ring stable isotopes record the impact of a foliar fungal pathogen on CO2 assimilation and growth in Douglas‐fir

Swiss needle cast (SNC) is a fungal disease of Douglas‐fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree‐rings of Douglas‐fir and a non‐susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ¹³C) of treated Douglas‐fir tree‐rings was greater than that of untreated Douglas‐fir tree‐rings during the fungicide treatment period. Both annual growth and tree‐ring Δ¹³C increased with treatment such that treated Douglas‐fir had values similar to co‐occurring western hemlock during the treatment period. There was no difference in the tree‐ring oxygen stable isotope ratio between treated and untreated Douglas‐fir. Tree‐ring Δ¹³C of diseased Douglas‐fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.     

Climatic stress increases forest fire severity across the western United States

Pervasive warming can lead to chronic stress on forest trees, which may contribute to mortality resulting from fire‐caused injuries. Longitudinal analyses of forest plots from across the western US show that high pre‐fire climatic water deficit was related to increased post‐fire tree mortality probabilities. This relationship between climate and fire was present after accounting for fire defences and injuries, and appeared to influence the effects of crown and stem injuries. Climate and fire interactions did not vary substantially across geographical regions, major genera and tree sizes. Our findings support recent physiological evidence showing that both drought and heating from fire can impair xylem conductivity. Warming trends have been linked to increasing probabilities of severe fire weather and fire spread; our results suggest that warming may also increase forest fire severity (the number of trees killed) independent of fire intensity (the amount of heat released during a fire).     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Silver fir and Douglas fir are more tolerant to extreme droughts than Norway spruce in south‐western Germany

Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.     

Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains

Extreme climatic events are key factors in initiating gradual or sudden changes in forest ecosystems through the promotion of severe, tree-killing disturbances such as fire, blowdown, and widespread insect outbreaks. In contrast to these climatically-incited disturbances, little is known about the more direct effect of drought on tree mortality, especially in high-elevation forests. Therefore projections of drought-induced mortality under future climatic conditions remain uncertain. For a subalpine forest landscape in the Rocky Mountains of northern Colorado (USA), we quantified lag effects of drought on mortality of Engelmann spruce Picea engelmannii , subalpine fir Abies lasiocarpa , and lodgepole pine Pinus contorta . For the period 1910–2004, we related death dates of 164 crossdated dead trees to early-season and late-season droughts. Following early-season droughts, spruce mortality increased over five years and fir mortality increased sharply over 11 years. Following late-season droughts, spruce showed a small increase in mortality within one year, whereas fir showed a consistent period of increased mortality over two years. Pine mortality was not affected by drought. Low pre-drought radial growth rates predisposed spruce and fir to drought-related mortality. Spruce and fir trees that died during a recent drought (2000–2004) had significantly lower pre-drought growth rates than live neighbour trees. Overall, we found large interspecific differences in drought-related mortality with fir showing the strongest effect followed by spruce and pine. This direct influence of climatic variability on differential tree mortality has the potential for driving large-scale changes in subalpine forests of the Rocky Mountains.     

Tree carbon allocation explains forest drought‐kill and recovery patterns

The mechanisms governing tree drought mortality and recovery remain a subject of inquiry and active debate given their role in the terrestrial carbon cycle and their concomitant impact on climate change. Counter‐intuitively, many trees do not die during the drought itself. Indeed, observations globally have documented that trees often grow for several years after drought before mortality. A combination of meta‐analysis and tree physiological models demonstrate that optimal carbon allocation after drought explains observed patterns of delayed tree mortality and provides a predictive recovery framework. Specifically, post‐drought, trees attempt to repair water transport tissue and achieve positive carbon balance through regrowing drought‐damaged xylem. Furthermore, the number of years of xylem regrowth required to recover function increases with tree size, explaining why drought mortality increases with size. These results indicate that tree resilience to drought‐kill may increase in the future, provided that CO₂ fertilisation facilitates more rapid xylem regrowth.     

Fire, fuels and restoration of ponderosa pine–Douglas fir forests in the Rocky Mountains, USA

Aim Forest restoration in ponderosa pine and mixed ponderosa pine–Douglas fir forests in the US Rocky Mountains has been highly influenced by a historical model of frequent, low‐severity surface fires developed for the ponderosa pine forests of the Southwestern USA. A restoration model, based on this low‐severity fire model, focuses on thinning and prescribed burning to restore historical forest structure. However, in the US Rocky Mountains, research on fire history and forest structure, and early historical reports, suggest the low‐severity model may only apply in limited geographical areas. The aim of this article is to elaborate a new variable‐severity fire model and evaluate the applicability of this model, along with the low‐severity model, for the ponderosa pine–Douglas fir forests of the Rocky Mountains. Location Rocky Mountains, USA. Methods The geographical applicability of the two fire models is evaluated using historical records, fire histories and forest age‐structure analyses. Results Historical sources and tree‐ring reconstructions document that, near or before ad 1900, the low‐severity model may apply in dry, low‐elevation settings, but that fires naturally varied in severity in most of these forests. Low‐severity fires were common, but high‐severity fires also burned thousands of hectares. Tree regeneration increased after these high‐severity fires, and often attained densities much greater than those reconstructed for Southwestern ponderosa pine forests. Main conclusions Exclusion of fire has not clearly and uniformly increased fuels or shifted the fire type from low‐ to high‐severity fires. However, logging and livestock grazing have increased tree densities and risk of high‐severity fires in some areas. Restoration is likely to be most effective which seeks to (1) restore variability of fire, (2) reverse changes brought about by livestock grazing and logging, and (3) modify these land uses so that degradation is not repeated.     

Use of exotic conifers as nesting sites by Black-faced Ibis (Theristicus melanopis melanopis) in an urban area of southern Chile

We studied the influence of seven habitat variables, including tree species, for nesting by the Black-faced Ibis (Theristicus melanopis melanopis) in an urban area of southern Chile. Variables were compared between 30 trees with nests and 30 randomly selected trees without nests. Nests were found in big trees with large diameters and heights. However, the only variable found to have a significant effect on site selection was tree species, which explained 57.9% of data variability (deviance) and suggested a selection of exotic conifers, mainly Douglas fir (Pseudotsuga menziesii). Tree species and tree diameter also had significant effects upon the number of nests per tree, jointly explaining 68.9% of data deviance. Our results suggest that in urban environments the Black-faced Ibis uses larger trees that provide greater nest stability and protection.     

Modeling the occurrence of 15 coniferous tree species throughout the Pacific Northwest of North America using a hybrid approach of a generic process‐based growth model and decision tree analysis

Question: Can we interpret how climatic variation limits photosynthesis and growth for one widely distributed species, and then relate these responses to model the geographic distributions of other species? Location: The forested region of the Pacific Northwest, United States and Canada. Methods: We first mapped monthly climatic data, averaged for the period 1950 to 1975 at 1 km resolution across the region. The recorded presence and absence of 15 native tree species were next mapped at 1 km resolution from data acquired on 22 771 field survey plots. To establish seasonal limits on photosynthesis and water use, a process‐based growth model (3‐PG, Physiological Processes to Predict Growth) was parameterized for Douglas‐fir (Pseudotsuga menziesii), one of the most widely distributed species in the region. Automated decision tree analyses were used to predict the distribution of different species by creating a suite of rules associated with the relative constraints that soil drought, atmospheric humidity deficits, suboptimal and subfreezing temperatures would impose on the growth of Douglas‐fir. Results: The 3‐PG process‐based modeling approach, combined with automated decision tree analyses, predicted presence and absence of 15 conifers on field survey plots with an average accuracy of 82±12%. Predictive models of current distribution for each species differed in the number of, order in, and physiological thresholds selected. A deficit in the soil water balance, followed by departures from optimum temperatures in the summer were the two most important variables selected in predicting species distributions. Conclusions: Although empirical models using different sampling techniques and statistical analyses may be more accurate in predicting current distribution of species, the hybrid approach presented in this paper provides a greater mechanistic understanding of the limits to growth and tree distributions. These attributes of process‐based models make them particularly useful in designing mitigating strategies to projected changes in climate.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Historic Anthropogenically Maintained Bear Grass Savannas of the Southeastern Olympic Peninsula

This paper documents the existence and character of a little known fire‐maintained anthropogenic ecosystem in the southeastern Olympic Peninsula of Washington State, U.S.A. Due to cessation of anthropogenic burning, there is no longer an intact example of this ecosystem. We present evidence from Skokomish oral tradition, historical documents, floral composition, tree‐ring analysis, stand structure, and site potential to describe former savanna structure and function. We believe this system was a mosaic of prairies, savannas, and woodlands in a forest matrix maintained through repeated burning to provide culturally important plants and animals. The overstory was dominated by Douglas‐fir (Pseudotsuga menziesii). Bear grass (Xerophyllum tenax) likely was a dominant understory component of the savannas, woodlands, and prairie edges. These lands grew forests in the absence of anthropogenic burning. Wide spacing of older trees or stumps in former stands and rapid invasion by younger trees in the late 1800s and early 1900s suggest a sudden change in stand structure. Shade‐intolerant prairie species are still present where openings have been maintained but not in surrounding forests. Bark charcoal, fire scars, tree establishment patterns, and oral traditions point to use of fire to maintain this system. A common successional trajectory for all these lands leads to forested vegetation. These findings suggest that frequent application of prescribed burning would be necessary to restore this ecosystem.     

Radial and stand‐level thinning treatments: 15‐year growth response of legacy ponderosa and Jeffrey pine trees

Restoration efforts to improve vigor of large, old trees and decrease risk to high‐intensity wildland fire and drought‐mediated insect mortality often include reductions in stand density. We examined 15‐year growth response of old ponderosa pine (Pinus ponderosa) and Jeffrey pine (Pinus jeffreyi) trees in northeastern California, U.S.A. to two levels of thinning treatments compared to an untreated (control) area. Density reductions involved radial thinning (thinning 9.1 m around individual trees) and stand thinning. Annual tree growth in the stand thinning increased immediately following treatment and was sustained over the 15 years. In contrast, radial thinning did not increase growth, but slowed decline compared to control trees. Available soil moisture was higher in the stand thinning than the control for 5 years post‐treatment and likely extended seasonal tree growth. Our results show that large, old trees can respond to restoration thinning treatments, but that the level of thinning impacts this response. Stand thinning must be sufficiently intensive to improve old tree growth and health, in part due to increasing available soil moisture. Importantly, focusing stand density reductions around the immediate neighborhood of legacy trees was insufficient to elicit a growth response, calling into question treatments attempting to increase vigor of legacy trees while still maintaining closed canopies in dry, coniferous forest types. Although radial thinning did not affect tree growth rates, this treatment may still achieve other resource objectives not studied here, such as protecting wildlife habitat, reducing the risk of severe fire injury, and decreasing susceptibility to bark beetle attacks.     

Impact of feral water buffalo and fire on growth and survival of mature savanna trees: An experimental field study in Kakadu National Park,northern Australia

Abstract The impact of feral Asian water buffalo (Bubalus bubalis) and season of fire on growth and survival of mature trees was monitored over 8 years in the eucalypt savannas of Kakadu National Park. Permanently marked plots were paired on either side of a 25‐km‐long buffalo‐proof fence at three locations on an elevational gradient, from ridge‐top to the edge of a floodplain; buffalo were removed from one side of the fence. All 750 trees ≥ 1.4 m height were permanently marked; survival and diameter of each tree was measured annually; 26 species were grouped into four eco‐taxonomic groups. The buffalo experiment was maintained for 7 years; trees were monitored an additional year. Fires were excluded from all sites the first 3 years, allowed to occur opportunistically for 4 years and excluded for the final year. Fires were of two main types: low‐intensity early dry season and high‐intensity late dry season. Growth rates of trees were size‐specific and positively related to diameters as exponential functions; trees grew slowest on the two ends of the gradient. Eucalypt mortality rates were 1.5 and 3 times lower than those of pantropics and of arborescent monocots, respectively, but the relative advantage was lost with fires or buffalo grazing. Without buffalo grazing, ground level biomass was 5–8 t ha^?1 compared with 2–3 t ha^?1, within 3 years. In buffalo‐absent plots, trees grew significantly slower on the dry ridge and slope, and had higher mortality across the entire gradient, compared with trees in buffalo‐present plots. At the floodplain margin, mortality of small palms was higher in buffalo‐present sites, most likely due to associated heavy infestations of weeds. Low‐intensity fires produced tree growth and mortality values similar to no‐fire, in general, but, like buffalo, provided a ‘fertilization’ effect for Eucalyptus miniata and Eucalyptus tetrodonta, increasing growth in all size classes. High‐intensity fires reduced growth and increased mortality of all functional groups, especially the smallest and largest (>35 cm d.b.h.) trees. When buffalo and fires were excluded in the final year, there were no differences in growth or mortality between paired sites across the environmental gradient. After 8 years, the total numbers of trees in buffalo‐absent plots were only 80% of the number in buffalo‐present plots, due to relatively greater recruitment of new trees in buffalo‐present plots; fire‐sensitive pantropics were particularly disadvantaged. Since the removal of buffalo is disadvantageous, at least over the first years, to savanna tree growth and survival due to a rebound effect of the ground‐level vegetation and subsequent changes in fire‐vegetation interactions, process‐orientated management aimed at reducing fuel loads and competitive pressure may be required in order to return the system to a previous state. The ‘footprint’ of 30 years of heavy grazing by buffalo has implications for the interpretation of previous studies on fire‐vegetation dynamics and for current research on vegetation change in these savannas.     

Variables associated with the occurrence of Ips beetles,red turpentine beetle and wood borers in live and dead ponderosa pines with post‐fire injury

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Ecological effects of changes in fire regimes in Pinus ponderosa ecosystems in the Colorado Front Range

Question: What is the relative importance of low‐ and high‐severity fires in shaping forest structure across the range of Pinus ponderosa in northern Colorado? Location: Colorado Front Range, USA. Methods: To assess severities of historic fires, 24 sites were sampled across an elevation range of 1800 to 2800 m for fire scars, tree establishment dates, tree mortality, and changes in tree‐ring growth. Results: Below 1950 m, the high number of fire scars, scarcity of large post‐fire cohorts, and lack of synchronous tree mortality or growth releases, indicate that historic fires were of low severity. In contrast, above 2200 m, fire severity was greater but frequency of widespread fires was substantially less. At 18 sites above 1950 m, 34 to 80% of the live trees date from establishment associated with the last moderate‐ to high‐severity fire. In these 18 sites, only 2 to 52% of the living trees pre‐date these fires suggesting that fire severities prior to any effects of fire suppression were sufficient to kill many trees. Conclusions: These findings for the P. ponderosa zone above ca. 2200 m (i.e. most of the zone) contradict the widespread perception that fire exclusion, at least at the stand scale of tens to hundreds of hectares, has resulted in unnaturally high stand densities or in an atypical abundance of shade‐tolerant species. At relatively mesic sites (e.g. higher elevation, north‐facing), the historic fire regime consisted of a variable‐severity regime, but forest structure was shaped primarily by severe fires rather than by surface fires.     

Fire frequency effects in a grassy woodland: Trees and grasses

Our study investigated the effects of fire frequency on the tree and grass components of Cumberland Plain Woodland, an endangered grassy eucalypt woodland in eastern Australia. We located three sites within each of three fire frequency classes (high, moderate, low) based on fire history and a similar time since last fire. For trees, we asked whether fire frequency affected density, eucalypt population structure, basal area and spatial patterning. For grasses, we tested for fire frequency effects on total abundance and the abundance of two common species. Density of trees was not significantly affected by fire frequency for juveniles, saplings or adult trees and neither was basal area per hectare. Features of a recruitment bottleneck model were present for eucalypts. There was a large pool of suppressed juveniles making up more than half the population at each fire frequency. Saplings were the smallest group and were susceptible to fire‐induced stem mortality, with particularly low numbers relative to juveniles and small trees where fire was frequent or intense. Despite this difference in sapling mortality, numbers of small trees did not differ with fire frequency. Saplings could transition to small trees in equal numbers if they did so more quickly at high than at lower fire frequencies, and if recruitment of saplings into the tree layer was controlled independently of fire frequency. The size hierarchy of small to medium eucalypt trees was homogeneous over eight of the nine sites, spatial patterning of adult trees was random tending to regular, and mean tree size decreased with density at all sites. These features of eucalypt population structure are indicative of possible resource competition which could regulate tree recruitment. Total grass cover index was high across all fire frequencies, with Themeda triandra dominating at high and moderate fire frequencies and Microlaena stipoides at low fire frequency.     

长白山主要针叶树种胸径和树高变异系数与竞争因子的关系

选取长白山地区8个皆伐标准地的1139株标准木,利用变异系数(CV)分析了云杉、冷杉和红松3个主要针叶树种的胸径和树高随龄阶、树高随径阶的变化.结果表明：在研究地区,3个主要针叶树种树高随径阶的变异相对较小,胸径和树高随龄阶的变异较大,胸径随龄阶的变异系数最大.在天然林中,传统的以径阶为自变量的树高曲线能较好地反映实际生长,以龄阶为自变量的胸径和树高生长模型对实际生长程度的反映较差.竞争是引起树高和胸径变动的主要因子.若将竞争因子添加到传统的生长方程和树高曲线中,可以大幅度提高模型的精度.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Post-fire regeneration strategies and tree bark resistance to heating in frequently burning tropical savanna woodlands and grasslands in Ethiopia

Regeneration mechanisms of vegetation and the role of tree bark resistance to frequent fire were studied in savanna woodlands and grasslands in Gambella, Western Ethiopia. Data were collected from four sites, each with three replicate plots. The variation between sites in species composition and biomass correlated with the differences in fire intensity. Foliar cover was recorded for individual plant species regenerating by sprouting from older parts of plants that had survived fire or by seedlings; records were made during the dry season and at the beginning of the wet season. Data on bark thickness and tree diameters of 12 dominant tree species were also recorded. Both facultative and obligate sprouters significantly contributed to post‐fire recovery, comprising 98.5 % of total vegetation cover. The contribution of seedlings to cover and abundance immediately following fire was negligible, but seedling density increased in the beginning of the rainy season, 4 to 5 months after fire. The importance of the sprouting and seeding strategies varied between the different plant growth forms. The highest contribution to cover and frequency was made by the most abundant grass species, which reproduced in both ways. Facultative sprouters made up 67.3 % of the vegetation cover, out of which 54 % consisted of grasses. Broad‐leaved herbs and trees/shrubs regenerating mainly by sprouting made up 31.3 % of the vegetation cover. Adaptations to fire in tree species seemed to include the development of a thick bark, once the tree has passed seedling stages. Tree bark thickness and tree diameter at breast height were strongly correlated with the time taken for cambium to reach an assumed lethal temperature of 60°C when exposed to fire, which indicated that mature trees with thick barks might resist stronger fire better than, e.g., small or young trees and trees with thin bark. However, for a given bark thickness the cambium resistance to heat varied three‐fold among species. Hence, site differences in fire intensity seemed to influence the distribution of trees depending on their bark characteristics and resistance to fire.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.