Ancient juniper trees growing on cliffs: toward a long Mediterranean tree-ring chronology

Juniperus phoenicea is a tree that can grow on vertical cliff faces in dry and warm Mediterranean climate conditions. These trees are adapted to extreme growing conditions where the main constraints are verticality, compact hard limestone, and low water supply. They respond to these constraints via various specific features and high longevity. The objective of this study is to confirm whether or not their tree-rings are annual in order to specify growth strategies and accurately date these trees. Trunk morphology, anatomical wood anomalies and radial growth were analyzed on 53 trees in the Ardèche canyon. Crossdating of the ring widths using traditional dendrochronological techniques was unsuccessful, so radiocarbon dating of tree pith was used to assess tree age, and wiggle-match dating was used to test for differences between number of rings counted and radiocarbon dates. Radiocarbon dates span the period 2520–685 BP. Minimal difference between radiocarbon dates and ring counts was apparently small—missing rings occur, but not in large numbers. Tree-ring formation is annual and radial growth is low, which creates stunted old trees. Such old living trees are uncommon in the Mediterranean basin, especially at low elevation. They can provide long tree-ring chronologies back to 792–524 cal BC. Results from the radiocarbon dating indicate that accurate annual dating of these rings may be possible by crossdating. J. phoenicea growing on cliffs offer a valuable model to better understand cliff population ecology and the functional responses of trees that can live in harsh environmental conditions.     

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.     

Demystifying the age of old olive trees

Iconic to the Mediterranean landscape, the age of monumental olive trees with large trunks and unique silhouettes is often considered several centuries to millennia. Here we combine tree-ring and radiocarbon dating to show that most monumental olive trees may have maximum ages between 300 and 500 years. Our example helps disentangling the tree size-longevity conundrum in a major Mediterranean cultivated tree species and important cultural heritage.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Historic variability in fire‐generated landscape heterogeneity of subalpine forests in the Canadian Rockies

Question: This study evaluates historical changes in landscape structure and heterogeneity in subalpine forests. We use response to severe fires in 2001 and 2003, along with historical reconstructions to examine crown‐fire effects on landscape heterogeneity and to assess, comparatively, effects of fire exclusion management in the 20th century. Location: Subalpine forests of Kootenay National Park (KNP), Canadian Rockies. Methods: Using a landscape‐level model based on a fire‐origin stand age map, we reconstructed decadal burned areas within the landscape for 1750‐2000 (forming reconstructed landscapes). Landscape pattern was analysed for each reconstructed landscape map, and we compared landscape pattern indices (total area, number of patches, mean patch area, patch area variation, largest patch index, edge density, perimeter–area ratio, landscape shape index) with those in 2005 after recent large fires. Results: After large fires in 1926, connectivity of the KNP landscape increased and its diversity was quite low. After 2001 and 2003 fires, the post‐fire landscape of 2005 was highly heterogeneous in terms of size, variation, edge density and perimeter–area ratio of the remnant forest patches. Since the decline in occurrence of large fires after 1926 reflected a period of wet weather, fuel build‐up resulting from landscape homogenization within the 20th century landscape could not be attributed solely to fire exclusion. This period without fires greatly enhanced connectivity of late‐successional forests that finally burned in 2001/2003, but connectivity was within the historical range for these forests. The gradual increase in stand connectivity before recent large fires may indicate that fire exclusion was less responsible than often believed for fuel build‐up in these fire‐susceptible older forests. Conclusions: The large fires at the beginning of the 21st century are within the natural range of disturbances for this landscape, and do not stand out as “human‐induced disasters” in their effects on landscape patterns. Such stochastic large disturbances contribute to maintenance of highly heterogeneous landscape structure, which is important for many taxa and natural ecological processes. Identifying future probability of such large disturbances and their ecological roles should be incorporated into management of these dynamic, disturbance‐prone systems.     

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.     

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.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Effects of 38 years of wildfires on tree density in the Blue Mountains,Australia

Forests are vital for biodiversity, carbon storage and ecosystem services, but can be potentially threatened by fires. Given the significance of forests and fire in a changing climate, research into the long-term effects of fire on forests plays an important role in understanding the global carbon cycle by the forests functioning as a large terrestrial carbon sink or source. In this study, we used aerial photography from 1975 and 2013 to count the change in the number of trees in 560 dry sclerophyll plots (40 × 40m) in the Blue Mountains of Australia. We analysed the relationship between the number of fires and severe fires in that period on the change in numbers of trees. We found that the average response was an increase of 1 tree per plot over 38 years. The number of fires had a small positive effect on tree numbers; plots with 2 or 3 severe fires had 1 and 2 extra trees, respectively, than those without fire. One exception was a severe fire in 2001 that did not show this positive effect, probably because it corresponded with extensive drought. Our findings suggest that number of forest canopy trees is resilient to the number of fires and number of severe fires.     

Wildfires affect tree growth and resilience in Northeastern China natural Dahurian larch forests: A dendrochronological perspective

Wildfires are natural and ubiquitous disturbances in boreal forests. Assessing their impacts on tree growth and resilience are particularly important to recognize the adaptation strategies of fire-tolerant species and forest succession in fire conditions. To date, the growth resilience of fire-tolerant species in boreal forests remains largely unquantified, and the drivers of resilience are poorly understood. Here, we measured the tree-ring widths of 99 fire-scarred trees from three sites in natural Dahurian larch (Larix gmelinii) forests. Three moderate-severity fire events in years 1987, 1990, and 2000 occurring at three sites were detected from the records of local forestry bureau. Based on tree-ring width data, we calculated resilience components (i.e., resistance, recovery, resilience and relative resilience) to quantify the responses of growth resilience in the larch trees to fires and analyzed their drivers at three sites. Results indicated that fires significantly reduced the tree growth. With the increasing tree age, these reductions were more pronounced. As for resilience components, our study showed a limited resistance but high recovery of tree growth against fires, and resistance tended to increase northwards but recovery showed the opposite, suggesting a growth-survival tradeoff was exhibited in Dahurian larch trees. With an increasing tree age, regional resistance and resilience showed a decreasing trend, whereas recovery and relative resilience showed an increasing trend. Resilience components were mainly affected by the climatic factors in spring. An increase in moisture availability enhanced resistance, a reduction in diurnal temperature range enhanced recovery, and an increase in mean temperature enhanced resilience and relative resilience. This study reveals that Dahurian larch could be even less favorable when faced with moderate or severe fire events, but a high capacity of recovery enables this species to adapt to the fire-prone condition. Moreover, this work highlights that the resilience of tree growth should be considered to understand tree behaviors and survival strategies of boreal forests following fires across fire-prone regions under future climate warming.     

Changes in fire regime break the legacy lock on successional trajectories in Alaskan boreal forest

Predicting plant community responses to changing environmental conditions is a key element of forecasting and mitigating the effects of global change. Disturbance can play an important role in these dynamics, by initiating cycles of secondary succession and generating opportunities for communities of long‐lived organisms to reorganize in alternative configurations. This study used landscape‐scale variations in environmental conditions, stand structure, and disturbance from an extreme fire year in Alaska to examine how these factors affected successional trajectories in boreal forests dominated by black spruce. Because fire intervals in interior Alaska are typically too short to allow relay succession, the initial cohorts of seedlings that recruit after fire largely determine future canopy composition. Consequently, in a dynamically stable landscape, postfire tree seedling composition should resemble that of the prefire forest stands, with little net change in tree composition after fire. Seedling recruitment data from 90 burned stands indicated that postfire establishment of black spruce was strongly linked to environmental conditions and was highest at sites that were moist and had high densities of prefire spruce. Although deciduous broadleaf trees were absent from most prefire stands, deciduous trees recruited from seed at many sites and were most abundant at sites where the fires burned severely, consuming much of the surface organic layer. Comparison of pre‐ and postfire tree composition in the burned stands indicated that the expected trajectory of black spruce self‐replacement was typical only at moist sites that burned with low fire severity. At severely burned sites, deciduous trees dominated the postfire tree seedling community, suggesting these sites will follow alternative, deciduous‐dominated trajectories of succession. Increases in the severity of boreal fires with climate warming may catalyze shifts to an increasingly deciduous‐dominated landscape, substantially altering landscape dynamics and ecosystem services in this part of the boreal forest.     

Age of A2 Horizon Charcoal and Forest Structure near Porto Trombetas,Pará, Brazil1

To study the structure and composition of old‐growth forest in the Saracá‐Taquera National Forest near Porto Trombetas, Brazil, we established 36 0.25 ha plots and described the vegetation. We collected charcoal from the A2 soil horizon of each plot for radiocarbon dating. Although fires have been very rare in this forest during historic times, the presence of charcoal in these soils indicates fire at some earlier period. The ages (conventional radiocarbon age adjusted to 1997) of the charcoal ranged from 177 to 1547 years. These ages, however, did not correlate significantly with any of several measures of biodiversity or stand characteristics. The relative uniformity of the current old‐growth forest indicates that either the prehistoric fires were of such low intensity that they had little long‐term effect on the vegetation or that the present stands have progressed to near steady state.     

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.     

Disturbance history and stand dynamics in tall open forest and riparian rainforest in the Central Highlands of Victoria

Abstract Spatial heterogeneity in the intensity of past disturbances has directly influenced the structure and composition of present‐day forests around the world. In south‐eastern Australia infrequent, high‐intensity wildfires are a major part of the historical disturbance regime. While these fires are often assumed to produce even‐aged stands, spatial heterogeneity in fire intensity due to highly variable topography may lead to more complex forest age structures. Our study describes the influence of disturbance on the age structure and dynamics of a mosaic of tall, open eucalypt forest, cool temperate rainforest and mixed species forest surrounding Bellel Creek in the Central Highlands of Victoria using dendrochronological techniques. We were particularly interested in the impacts of the 1939 Black Friday fire and its effects on forest age structure and subsequent stand development patterns. Within our study site tall open forest displayed two distinct age cohorts: (i) trees that established immediately after the 1939 fire and accounted for the majority of individuals in the forest, and (ii) scattered groups of older trees estimated to be approximately 200–250 years old. Cool temperate rainforest and mixed forest were also dominated by the post‐1939 fire age cohort. However, a greater proportion of trees in these forest types survived the 1939 fire relative to the tall open forest. The impact of the 1939 fire on the growth of surviving trees was highly variable but generally short‐lived. In most cases growth decreased after the 1939 fire, but generally returned to prefire levels within 1–3 years. Non‐fire disturbances were limited to small‐scale branch‐ and tree‐fall events, although the extreme snowstorm of 1977 appears to have caused extensive damage to rainforest communities. Our study demonstrates the opportunities for dendroecological studies to reconstruct historical dynamics and disturbance patterns in Australian forests and provides important insights into variation in landscape‐scale fire impacts and their effect on subsequent forest development patterns.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Californian mixed-conifer forests under unmanaged fire regimes in the Sierra San Pedro Mártir, Baja California, Mexico

Aim This study appraises historical fire regimes for Californian mixed‐conifer forests of the Sierra San Pedro Mártir (SSPM). The SSPM represents the last remaining mixed‐conifer forest along the Pacific coast still subject to uncontrolled, periodic ground fire. Location The SSPM is a north–south trending fault bound range, centred on 31°N latitude, 100 km SE of Ensenada, Baja California. Methods We surveyed forests for composition, population structure, and historical dynamics both spatially and temporally over the past 65 years using repeat aerial photographs and ground sampling. Fire perimeter history was reconstructed based on time‐series aerial photographs dating from 1942 to 1991 and interpretable back to 1925. A total of 256 1‐ha sites randomly selected from aerial photographs were examined along a chronosequence for density and cover of canopy trees, density of snags and downed logs, and cover of non‐conifer trees and shrubs. Twenty‐four stands were sampled on‐the‐ground by a point‐centred quarter method which yielded data on tree density, basal area, frequency, importance value, and shrub and herb cover. Results Forests experience moderately intense understory fires that range in size to 6400 ha, as well as numerous smaller, low intensity burns with low cumulative spatial extent. SSPM forests average 25–45% cover and 65–145 trees per ha. Sapling densities were two to three times that of overstory trees. Size‐age distributions of trees ≥ 4 cm dbh indicate multi‐age stands with steady‐state dynamics. Stands are similar to Californian mixed conifer forests prior to the imposition of fire suppression policy. Livestock grazing does not appear to be suppressing conifer regeneration. Main conclusions Our spatially‐based reconstruction shows the open forest structure in SSPM to be a product of infrequent, intense surface fires with fire rotation periods of 52 years, rather than frequent, low intensity fires at intervals of 4–20 years proposed from California fire‐scar dendrochronology (FSD) studies. Ground fires in SSPM were intense enough to kill pole‐size trees and a significant number of overstory trees. We attribute long fire intervals to the gradual build‐up of subcontinuous shrub cover, conifer recruitment and litter accumulation. Differences from photo interpretation and FSD estimates are due to assumptions made with respect to site‐based (point) sampling of fire, and nonfractal fire intensities along fire size frequency distributions. Fire return intervals determined by FSD give undue importance to local burns which collectively use up little fuel, cover little area, and have little demographic impact on forests.     

The ecology,distribution, conservation and management of large old trees

Large old trees are some of the most iconic biota on earth and are integral parts of many terrestrial ecosystems including those in tropical, temperate and boreal forests, deserts, savannas, agro‐ecological areas, and urban environments. In this review, we provide new insights into the ecology, function, evolution and management of large old trees through broad cross‐disciplinary perspectives from literatures in plant physiology, growth and development, evolution, habitat value for fauna and flora, and conservation management. Our review reveals that the diameter, height and longevity of large old trees varies greatly on an inter‐specific basis, thereby creating serious challenges in defining large old trees and demanding an ecosystem‐ and species‐specific definition that will only rarely be readily transferable to other species or ecosystems. Such variation is also manifested by marked inter‐specific differences in the key attributes of large old trees (beyond diameter and height) such as the extent of buttressing, canopy architecture, the extent of bark micro‐environments and the prevalence of cavities. We found that large old trees play an extraordinary range of critical ecological roles including in hydrological regimes, nutrient cycles and numerous ecosystem processes. Large old trees strongly influence the spatial and temporal distribution and abundance of individuals of the same species and populations of numerous other plant and animal species. We suggest many key characteristics of large old trees such as extreme height, prolonged lifespans, and the presence of cavities – which confer competitive and evolutionary advantages in undisturbed environments – can render such trees highly susceptible to a range of human influences. Large old trees are vulnerable to threats ranging from droughts, fire, pests and pathogens, to logging, land clearing, landscape fragmentation and climate change. Tackling such diverse threats is challenging because they often interact and manifest in different ways in different ecosystems, demanding targeted species‐ or ecosystem‐specific responses. We argue that novel management actions will often be required to protect existing large old trees and ensure the recruitment of new cohorts of such trees. For example, fine‐scale tree‐level conservation such as buffering individual stems will be required in many environments such as in agricultural areas and urban environments. Landscape‐level approaches like protecting places where large old trees are most likely to occur will be needed. However, this brings challenges associated with likely changes in tree distributions associated with climate change, because long‐lived trees may presently exist in places unsuitable for the development of new cohorts of the same species. Appropriate future environmental domains for a species could exist in new locations where it has never previously occurred. The future distribution and persistence of large old trees may require controversial responses including assisted migration via seed or seedling establishment in new locales. However, the effectiveness of such approaches may be limited where key ecological features of large old trees (such as cavity presence) depend on other species such as termites, fungi and bacteria. Unless other species with similar ecological roles are present to fulfil these functions, these taxa might need to be moved concurrently with the target tree species.     

Frequent fires reduce tree growth in northern Australian savannas: implications for tree demography and carbon sequestration

Tropical savannas are typically highly productive yet fire‐prone ecosystems, and it has been suggested that reducing fire frequency in savannas could substantially increase the size of the global carbon sink. However, the long‐term demographic consequences of modifying fire regimes in savannas are difficult to predict, with the effects of fire on many parameters, such as tree growth rates, poorly understood. Over 10 years, we examined the effects of fire frequency on the growth rates (annual increment of diameter at breast height) of 3075 tagged trees, at 137 locations throughout the mesic savannas of Kakadu, Nitmiluk and Litchfield National Parks, in northern Australia. Frequent fires substantially reduced tree growth rates, with the magnitude of the effect markedly increasing with fire severity. The highest observed frequencies of mild, moderate and severe fires (1.0, 0.8 and 0.4 fires yr^?1, respectively) reduced tree growth by 24%, 40% and 66% respectively, relative to unburnt areas. These reductions in tree growth imply reductions in the net primary productivity of trees by between 0.19 t C ha^?1 yr^?1, in the case of mild fires, and 0.51 t C ha^?1 yr^?1, in the case of severe fires. Such reductions are relatively large, given that net biome productivity (carbon sequestration potential) of these savannas is estimated to be just 1–2 t C ha^?1 yr^?1. Our results suggest that current models of savanna tree demography, that do not account for a relationship between severe fire frequency and tree growth rate, are likely to underestimate the long‐term negative effects of frequent severe fires on tree populations. Additionally, the negative impact of frequent severe fires on carbon sequestration rates may have been underestimated; reducing fire frequencies in savannas may increase carbon sequestration to a greater extent than previously thought.