A comprehensive resilience assessment of Mexican tree species and their relationship with drought events over the last century

The resilience of forests to drought events has become a major natural resource sustainability concern, especially in response to climate change. Yet, little is known about the legacy effects of repeated droughts, and tree species ability to respond across environmental gradients. In this study, we used a tree-ring database (121 sites) to evaluate the overall resilience of tree species to drought events in the last century. We investigated how climate and geography affected the response at the species level. We evaluated temporal trends of resilience using a predictive mixed linear modeling approach. We found that pointer years (e.g., tree growth reduction) occurred during 11.3% of the 20th century, with an average decrease in tree growth of 66% compared to the previous period. The occurrence of pointer years was associated with negative values of the Standardized Precipitation Index (SPI, 81.6%) and Palmer Drought Severity Index (PDSI, 77.3%). Tree species differed in their resilience capacity, however, species inhabiting xeric conditions were less resistant but with higher recovery rates (e.g., Abies concolor, Pinus lambertiana, and Pinus jeffreyi). On average, tree species needed 2.7 years to recover from drought events, with extreme cases requiring more than a decade to reach pre-drought tree growth rates. The main abiotic factor related to resilience was precipitation, confirming that some tree species are better adapted to resist the effects of droughts. We found a temporal variation for all tree resilience indices (scaled to 100), with a decreasing resistance (−0.56 by decade) and resilience (−0.22 by decade), but with a higher recovery (+1.72 by decade) and relative resilience rate (+0.33 by decade). Our results emphasize the importance of time series of forest resilience, particularly by distinguishing the species-level response in the context of legacy of droughts, which are likely to become more frequent and intense under a changing climate.     

Components of tree resilience: effects of successive low‐growth episodes in old ponderosa pine forests

Recent world‐wide episodes of tree dieback have been attributed to increasing temperatures and associated drought. Because these events are likely to become more common, improved knowledge of their cumulative effects on resilience and the ability to recover pre‐disturbance conditions is important for forest management. Here we propose several indices to examine components of individual tree resilience based on tree ring growth: resistance (inverse of growth reduction during the episode), recovery (growth increase relative to the minimum growth during the episode), resilience (capacity to reach pre‐episode growth levels) and relative resilience (resilience weighted by the damage incurred during the episode). Based on tree ring analyses, we analyzed historical patterns of tree resilience to successive drought‐induced low growth periods in ponderosa pine trees growing in unmanaged, remote forests of the Rocky Mountains. Low‐growth periods registered in tree rings were related to anomalies in the Palmer drought severity index (PDSI) and were attributed to drought. Independently of the impact of a specific event, subsequent growth after a single low‐growth episode was related to the growth prior to the event. Growth performance differed with tree age: young trees were overall more resistant to low‐growth periods, but older trees recovered better from more recent events. Regardless of tree age, recently burned sites exhibited lower post‐episode growth and lower resistance and resilience than unburned ones. We found mixed evidence for the cumulative effect of past low‐growth episodes: overall, greater impacts of a prior event and greater cumulative effects of past low‐growth periods caused a decrease in resistance. However, we did not find a progressive decrease in resilience over time in old trees. Our results highlight the value of using a combination of estimators to evaluate the different components of resilience. Specifically, while tree responses to disturbance depend on past disturbance episodes, the response is context‐specific and depends on the impact the capacity to recover after disturbance. This suggests that recent increases in forest mortality under current climate trends could relate to thresholds on specific components of resilience (resistance, recovery, resilience itself) rather than to an overall loss of resilience over time. Identifying such thresholds and their underlying mechanisms is a promising area of research with important implications for forest management.     

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.     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.

     

Forest resilience to drought varies across biomes

Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species‐level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree‐ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring‐width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994–1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi‐arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi‐arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.     

西藏东部主要建群树种径向生长对极端干旱的响应差异

不同坡向、不同海拔树木生长对极端干旱事件的响应可能不同,然而这方面的认识不足。为此,选取西藏东部珠角拉山阴、阳坡的建群树种川西云杉 (Picea likiangensis var. rubescens) 和大果圆柏 (Juniperus tibetica),在不同海拔高度建立了树轮宽度年表,分析了径向生长的气候响应,以及对极端干旱事件的抵抗力和恢复力。结果表明:阳坡大果圆柏和阴坡川西云杉的树木生长对气候的响应存在相似性,均与前一年3-6月、11-12月气温显著正相关,与当年4-5月气温显著负相关,与当年4-5月降水和帕尔默干旱指数 (PDSI, Palmer Drought Severity Index) 显著正相关 (P <0.05)。阳坡大果圆柏的抵抗力显著低于阴坡川西云杉。随着海拔升高,阴坡川西云杉树木个体的抵抗力和恢复力均显著提高,而阳坡大果圆柏树木个体抵抗力、恢复力在不同海拔无显著区别。结合混合效应模型表明树木抵抗力主要受当年4-5月平均最高气温限制,树木恢复力主要受干旱事件后四年4-5月平均最高气温限制 (P <0.01),说明生长季高温引起的极端干旱是树木径向生长下降的主要原因。     

Tree resilience to drought increases in the Tibetan Plateau

Forests in the Tibetan Plateau are thought to be vulnerable to climate extremes, yet they also tend to exhibit resilience contributing to the maintenance of ecosystem services in and beyond the plateau. So far the spatiotemporal pattern in tree resilience in the Tibetan Plateau remains largely unquantified and the influence of specific factors on the resilience is poorly understood. Here, we study ring‐width data from 849 trees at 28 sites in the Tibetan Plateau with the aim to quantify tree resilience and determine their diving forces. Three extreme drought events in years 1969, 1979, and 1995 are detected from metrological records. Regional tree resistance to the three extreme droughts shows a decreasing trend with the proportion of trees having high resistance ranging from 71.9%, 55.2%, to 39.7%. Regional tree recovery is increasing with the proportion of trees having high recovery ranging from 28.3%, 52.2%, to 64.2%. The area with high resistance is contracting and that of high recovery is expanding. The spatiotemporal resistance and recovery are associated with moisture availability and diurnal temperature range, respectively. In addition, they are both associated with forest internal factor represented by growth consistence among trees. We conclude that juniper trees in the Tibetan Plateau have increased resilience to extreme droughts in the study period. We highlight pervasive resilience in juniper trees. The results have implications for predicting tree resilience and identifying areas vulnerable to future climate extremes.     

Declining tree growth resilience mediates subsequent forest mortality in the US Mountain West

Climate change-triggered forest die-off is an increasing threat to global forests and carbon sequestration but remains extremely challenging to predict. Tree growth resilience metrics have been proposed as measurable proxies of tree susceptibility to mortality. However, it remains unclear whether tree growth resilience can improve predictions of stand-level mortality. Here, we use an extensive tree-ring dataset collected at ~3000 permanent forest inventory plots, spanning 13 dominant species across the US Mountain West, where forests have experienced strong drought and extensive die-off has been observed in the past two decades, to test the hypothesis that tree growth resilience to drought can explain and improve predictions of observed stand-level mortality. We found substantial increases in growth variability and temporal autocorrelation as well declining drought resistance and resilience for a number of species over the second half of the 20th century. Declining resilience and low tree growth were strongly associated with cross- and within-species patterns of mortality. Resilience metrics had similar explicative power compared to climate and stand structure, but the covariance structure among predictors implied that the effect of tree resilience on mortality could partially be explained by stand and climate variables. We conclude that tree growth resilience offers highly valuable insights on tree physiology by integrating the effect of stressors on forest mortality but may have only moderate potential to improve large-scale projections of forest die-off under climate change.     

Resilience capacity of Araucaria araucana to extreme drought events

Ongoing climate change has induced modification in the frequency and intensity of extreme climatic events, with consequent impact on tree and forest growth resilience. Araucaria araucana is an endangered Patagonian conifer, which provides several ecosystem services to local human societies and plays fundamental ecological roles in natural communities. These woodlands have historically suffered different types of anthropogenic disturbance, such as fire, logging and grazing, nevertheless the species resilience to extreme drought events remains still poorly understood. To fill this gap of knowledge, we applied dendrochronological methods to several A. araucana stands distributed along a steep bioclimatic gradient in order to reconstruct resilience capacity, in term of stem growth resistance and recovery, to three successive extreme spring-early summer droughts which occurred during the 20th century. Results showed an increase in the species recovery along the considered dry spells, whereas no clear trend emerged for resistance, suggesting no cumulative effect of drought upon resilience. Both resistance and recovery presented different values depending on bioclimatic settings, being xeric stands more sensitive to extreme episodes with respect to mesic woodlands, particularly during the more recent drought event when trees growing in drier environments were not able to reach pre-drought stem growth rates. Tree-level characteristics, such as age and growth trends prior to drought, modulated the species resilience, suggesting that future dry spells would possibly induce shifts in population dynamics, and furthermore be detrimental for fast-growing trees. Our analysis highlighted the response of a key Patagonian tree species to extreme drought events, providing bioclimatic-specific useful information for conservation plans of this natural resource.     

Diverging growth and resilience of Pinus tabulaeformis and Pinus massoniana to droughts in north-south transition zone,central China

Climate transition zone is a sensitive area of climate change and ecological transition where forests are vulnerable to climate extremes. Extreme droughts are increasing in frequency and magnitude under climate change, resulting in structure and function changes of forest ecosystems. Here, to analyze climate-growth relationships and quantify tree resilience to extreme droughts, we developed six tree-ring-width chronologies from P. tabulaeformis and P. massoniana sampling sites in Mt. Jigong region, Central China. The results indicated that all chronologies from the two species had good consistency, precipitation in current April and mean temperature in current August or mean minimum temperature from current August to October were the main limiting factors of the two tree species growth, but the responses of P. massoniana ring-width to climatic factors was more complex than that of P. tabulaeformis. The results also showed that tree growth of 1999–2005 was the lowest growing period during 1979–2018, and P. massoniana grew better than P. tabulaeformis before 2005 and vice versa after 2005. Comparing low growth years of trees, we identified to study tree growth resilience. The calculations from 1988, 1999–2005 and 2011 drought years indicated that P. tabulaeformis had more increased resilience to extreme droughts than that of P. massoniana, and the two species had stronger ecological recovery and resilience under global warming and non-extreme drought conditions in the recent 40 years. These results have implications for predicting tree resilience and identifying tree species in heterogeneous forest landscapes vulnerable to future climate change in climatic transition zone.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

气候和密度对刺槐径向生长和干旱脆弱性的影响

为评估气候和竞争对刺槐径向生长、抗性和弹性的影响,使用年轮气候学方法建立河南省民权和济源不同密度刺槐的生长年表,确定不同密度刺槐径向生长与气象因子的关联,利用胸高断面积增量变化获得干旱事件前后不同密度刺槐的干旱脆弱性,旨在确定气候和密度对刺槐径向生长和干旱脆弱性影响。结果表明：在生长前期,不同密度刺槐的径向生长无显著差异,随着树木的生长,高密度刺槐的年轮宽度和胸高断面积增量(BAI)开始显著低于低密度(P<0.05)。Pearson相关分析结果显示,生长季的标准化植被蒸散指数、降水、相对湿度、温度和饱和水汽压亏缺是影响刺槐生长的重要因素。路径分析结果显示降水和温度是年尺度上影响刺槐生长关键因素。受干旱事件的影响,刺槐的年轮宽度和BAI均下降,低密度刺槐恢复力、弹性、相对弹性均显著高于高密度(P<0.05),在第1次干旱事件发生后,不同密度刺槐均恢复生长,但无法恢复到干旱前的生长水平。在多次干旱事件后,高密度刺槐相对弹性趋于或小于0,表明受多次干旱影响,其生长不能恢复到干旱前水平。随着时间的推移,济源刺槐在经历3次干旱后仍保留一定的弹性,但民权高密度刺槐在第2次干旱事件后相对...     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Tree vigor and height growth in Black Spruce

Summary The relationship between tree height and tree vigor was examined in order to test the hypothesis that increased height results in a reduction in photosynthate availability for growth due to an increase in respiratory load of the sapwood. Various vigor indices were measured on black spruce [Picea mariana (Mill.) B. S. P.] trees on a wide range of sites and ages. The relationship between these vigor indices and tree height and site quality were then evaluated through correlation and regression analyses. Vigor indices based on meristematic activity such as specific volume increment and height growth rate were generally more strongly correlated than foliage based indices (i.e. Waring vigor estimates). Both tree height and site index were found to have a significant effect on specific volume increment and height growth rate. However the height effect decreased with a decrease in site quality, and became insignificant (P <0.05) on the poorer sites. The full regression model, allowing the height effect to vary between different site quality groups, explained 82% and 72% of the variability in specific volume increment and height growth rate, respectively. The results support the hypothesis that increases in the ratio of respiration to photosynthesis in trees as they increase in size is largely a function of tree height. Possible explanations for the decreased height effect on poor sites are also discussed.     

Early Differentiation of the Phenotypic Space and Performance of Juniperus thurifera Across Woodland-Expanding Areas

Forest expansion after rural abandonment is changing landscapes, bringing new challenges and opportunities. Juvenile performance has an effect in the establishment of new forests and the subsequent acclimation of adult trees. However, most studies have focused on the performance of adult trees. Here, we analysed differences in the phenotypic trait space of juveniles across gradients of Juniperus thurifera woodland expansion and explored which traits shape juveniles’ performance, seeking specific drivers of such differentiation and the role of the ontogeny. Thus, we surveyed functional leaf and plant-size traits in 180 juveniles growing in 18 plots at three different tree density stages (mature woodland, recently established area and a transition zone between both) and recorded local environmental conditions around each individual. We also estimated their radial growth and resilience to drought (in terms of resistance, recovery and resilience) as performance indicators. We found no relevant effects of the study environmental variables, but we did find a shift in the phenotypic space among juveniles across the gradient, with taller crowns and higher amount of leaf N and δ¹³C at the expanding edge. This trait differentiation across the gradient implied higher performance of young trees growing in old agricultural fields, both in growth rate and drought resilience terms. Our results showed a relevant land-use legacy on early trait differentiation that shapes more efficient phenotypes in the expanding edge compared to those in mature woodlands, suggesting that the future of these new forests is not compromised under current conditions.

     

Marine protected areas increase resilience among coral reef communities

With marine biodiversity declining globally at accelerating rates, maximising the effectiveness of conservation has become a key goal for local, national and international regulators. Marine protected areas (MPAs) have been widely advocated for conserving and managing marine biodiversity yet, despite extensive research, their benefits for conserving non‐target species and wider ecosystem functions remain unclear. Here, we demonstrate that MPAs can increase the resilience of coral reef communities to natural disturbances, including coral bleaching, coral diseases, Acanthaster planci outbreaks and storms. Using a 20‐year time series from Australia's Great Barrier Reef, we show that within MPAs, (1) reef community composition was 21–38% more stable; (2) the magnitude of disturbance impacts was 30% lower and (3) subsequent recovery was 20% faster that in adjacent unprotected habitats. Our results demonstrate that MPAs can increase the resilience of marine communities to natural disturbance possibly through herbivory, trophic cascades and portfolio effects.     

树木年轮揭示的东灵山主要树种间干旱耐受性差异

近年来北京地区的森林随极端干旱加剧表现出脆弱性特征,为了解气候变化下不同树种的干旱耐受性,选择北京东灵山森林内3个乔木树种(华北落叶松、油松和辽东栎),利用树木年轮生态学方法分析了径向生长与气候的关系,以及对极端干旱事件的抵抗力和弹性。结果表明: 华北落叶松和油松与5—6月气温呈显著负相关,辽东栎与5月气温呈显著负相关;华北落叶松与6月降水量、5—6月和8—9月相对湿度呈显著正相关,油松与6—8月降水量和相对湿度呈显著正相关,辽东栎与2月和5月降水量、5月相对湿度呈显著正相关;所有树种均与当年5—7月标准化降水蒸散指数(SPEI)呈显著正相关。华北落叶松是干旱耐受性最弱的树种,径向生长在所选极端干旱事件中(1994年、2001—2002年和2007年)下降幅度最大(46.6%～69.6%),抵抗力(0.534、0.304、0.530)和弹性(0.686、0.570、0.753)显著低于辽东栎和油松,辽东栎在2007年抵抗力显著高于油松,弹性无显著差异。生长季持续的高温或降水减少引起的极端干旱是树木径向生长下降的主要原因,树种间各异的生理生态策略是干旱耐受性差异的可能原因。研究结果可为未来造林树种选择和森林管护措施制定提供新依据,以在气候压力持续增加背景下维持森林生态系统功能和服务。     

The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas

The distribution and abundance of trees can be strongly affected by disturbance such as fire. In mixed tree/grass ecosystems, recurrent grass‐fuelled fires can strongly suppress tree saplings and therefore control tree dominance. We propose that changes in atmospheric [CO₂] could influence tree cover in such metastable ecosystems by altering their postburn recovery rates relative to flammable herbaceous growth forms such as grasses. Slow sapling recovery rates at low [CO₂] would favour the spread of grasses and a reduction of tree cover. To test the possible importance of [CO₂]/fire interactions, we first used a Dynamic Global Vegetation Model (DGVM) to simulate biomass in grassy ecosystems in South Africa with and without fire. The results indicate that fire has a major effect under higher rainfall conditions suggesting an important role for fire/[CO₂] interactions. We then used a demographic model of the effects of fire on mesic savanna trees to test the importance of grass/tree differences in postburn recovery rates. We adjusted grass and tree growth in the model according to the DGVM output of net primary production at different [CO₂] relative to current conditions. The simulations predicted elimination of trees at [CO₂] typical of the last glacial period (180 ppm) because tree growth rate is too slow (15 years) to grow to a fire‐proof size of ca. 3 m. Simulated grass growth would produce an adequate fuel load for a burn in only 2 years. Simulations of preindustrial [CO₂] (270 ppm) predict occurrence of trees but at low densities. The greatest increase in trees occurs from preindustrial to current [CO₂] (360 ppm). The simulations are consistent with palaeo‐records which indicate that trees disappeared from sites that are currently savannas in South Africa in the last glacial. Savanna trees reappeared in the Holocene. There has also been a large increase in trees over the last 50–100 years. We suggest that slow tree recovery after fire, rather than differential photosynthetic efficiencies in C₃ and C₄ plants, might have been the significant factor in the Late Tertiary spread of flammable grasslands under low [CO₂] because open, high light environments would have been a prerequisite for the spread of C₄ grasses. Our simulations suggest further that low [CO₂] could have been a significant factor in the reduction of trees during glacial times, because of their slower regrowth after disturbance, with fire favouring the spread of grasses.     

Decomposing multiple dimensions of stability in global change experiments

Ecological stability is the central framework to understand an ecosystem's ability to absorb or recover from environmental change. Recent modelling and conceptual work suggests that stability is a multidimensional construct comprising different response aspects. Using two freshwater mesocosm experiments as case studies, we show how the response to single perturbations can be decomposed in different stability aspects (resistance, resilience, recovery, temporal stability) for both ecosystem functions and community composition. We find that extended community recovery is tightly connected to a nearly complete recovery of the function (biomass production), whereas systems with incomplete recovery of the species composition ranged widely in their biomass compared to controls. Moreover, recovery was most complete when either resistance or resilience was high, the latter associated with low temporal stability around the recovery trend. In summary, no single aspect of stability was sufficient to reflect the overall stability of the system.