The combined effects of a long‐term experimental drought and an extreme drought on the use of plant‐water sources in a Mediterranean forest

Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.     

Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

Dampening effects of long‐term experimental drought on growth and mortality rates of a Holm oak forest

Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long‐term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (?66.5%) and Q. ilex (?17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005–2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra‐ and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long‐term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long‐term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species‐specific responses to drought, what may eventually lead to a partial community shift favoring the more drought‐resistant species. However, we also report a dampening of the treatment effect on the two drought‐sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.     

Rapid climate change-related growth decline at the southern range edge of Fagus sylvatica

Studies on Fagus sylvatica show that growth in populations toward the southern limit of this species' distribution is limited strongly by drought. Warming temperatures in the Mediterranean region are expected to exacerbate drought where they are not accompanied by increases in precipitation. We studied levels of annual growth in mature F. sylvatica trees over the last half‐century in the Montseny Mountains in Catalonia (northeast Spain). Our results show significantly lower growth of mature trees at the lower limit of this species' distribution when compared with trees at higher altitudes. Growth at the lower Fagus limit is characterized by a rapid recent decline starting in approximately 1975. By 2003, growth of mature trees had fallen by 49% when compared with predecline levels. This is not an age‐related phenomenon, nor is it seen in comparable populations at higher altitudes. Analysis of climate‐growth relationships suggests that the observed decline in growth is a result of warming temperatures and that, as precipitation in the region has not increased, precipitation is now insufficient to ameliorate the negative effects of increased temperatures on tree growth. As the climate‐response of the studied forest is comparable with that of F. sylvatica forests in other southern European regions, it is possible that this growth decline is a more widespread phenomenon. Warming temperatures may lead to a rapid decline in the growth of range‐edge populations and a consequent retreat of the species distribution in southern Europe. Assessment of long‐term growth trends across the southern range edge of F. sylvatica therefore merits further attention.     

Recent CO2 rise has modified the sensitivity of tropical tree growth to rainfall and temperature

Atmospheric CO₂ (c_a) rise changes the physiology and possibly growth of tropical trees, but these effects are likely modified by climate. Such c_a × climate interactions importantly drive CO₂ fertilization effects of tropical forests predicted by global vegetation models, but have not been tested empirically. Here we use tree‐ring analyses to quantify how c_a rise has shifted the sensitivity of tree stem growth to annual fluctuations in rainfall and temperature. We hypothesized that c_a rise reduces drought sensitivity and increases temperature sensitivity of growth, by reducing transpiration and increasing leaf temperature. These responses were expected for cooler sites. At warmer sites, c_a rise may cause leaf temperatures to frequently exceed the optimum for photosynthesis, and thus induce increased drought sensitivity and stronger negative effects of temperature. We tested these hypotheses using measurements of 5,318 annual rings from 129 trees of the widely distributed (sub‐)tropical tree species, Toona ciliata. We studied growth responses during 1950–2014, a period during which c_a rose by 28%. Tree‐ring data were obtained from two cooler (mean annual temperature: 20.5–20.7°C) and two warmer (23.5–24.8°C) sites. We tested c_a × climate interactions, using mixed‐effect models of ring‐width measurements. Our statistical models revealed several significant and robust c_a × climate interactions. At cooler sites (and seasons), c_a × climate interactions showed good agreement with hypothesized growth responses of reduced drought sensitivity and increased temperature sensitivity. At warmer sites, drought sensitivity increased with increasing c_a, as predicted, and hot years caused stronger growth reduction at high c_a. Overall, c_a rise has significantly modified sensitivity of Toona stem growth to climatic variation, but these changes depended on mean climate. Our study suggests that effects of c_a rise on tropical tree growth may be more complex and less stimulatory than commonly assumed and require a better representation in global vegetation models.     

种子大小和干旱胁迫对辽东栎幼苗生长和生理特性的影响

在温室内遮阴条件下,设置80%、60%、40%和20%田间持水量(对照、轻度、中度和重度干旱)4个处理,研究种子大小和干旱胁迫对盆栽辽东栎幼苗生长和生理特性的影响。结果表明:大种子(3.05±0.38 g)幼苗的单株叶面积、总干质量和根冠比在所有处理均显著大于小种子(1.46±0.27 g)幼苗,前者的株高、基径、叶片数、比叶面积、相对生长率和净同化率等生长参数在轻度、中度和重度干旱处理均不同程度大于后者。大种子幼苗叶片过氧化物酶(POD)、过氧化氢酶(CAT)和超氧化物歧化酶(SOD)活性均高于小种子幼苗,前者叶片丙二醛(MDA)、可溶性蛋白、游离脯氨酸含量和叶绿素总量在部分干旱处理显著大于后者。除根冠比外,其他生长参数均随干旱胁迫增强逐渐减小,重度干旱处理大、小种子幼苗总干质量分别比对照降低19.4%和20.0%。POD、CAT和SOD活性均随干旱胁迫增强先升后降,在中度干旱处理,大、小种子幼苗POD活性分别显著高于对照126.7%和142.1%,CAT活性分别显著高于对照170.0%和151.9%。在重度干旱处理,大、小种子幼苗MDA含量分别显著高于对照86.5%和68.9%。可...     

Effects of decadal experimental drought and climate extremes on vegetation growth in Mediterranean forests and shrublands

Increased drought combined with extreme episodes of heatwaves is triggering severe impacts on vegetation growth, particularly for plant communities in arid and semiarid ecosystems. Although there is an abundance of short‐term field drought experiments in natural ecosystems, remaining knowledge gaps limit the understanding and prediction of vegetation growth to ongoing and future climate scenarios. Here, we assessed the impacts of long‐term (1999–2016) experimental drought (ca. ?30% rainfall) on the vegetation growth of a Mediterranean high (H) and low (L)‐canopy forests and an early‐successional shrubland, as indicated by above‐ground biomass increment (ABI) and standing density, respectively. We found habitat context (impact of historical climate change, soil depth and successional status) of the study sites significantly affected the magnitude of climate impacts; there were synergistic effects of experimental drought and meteorological drought (Standardised Precipitation–Evapotranspiration Index, SPEI) as well as extreme dry years on vegetation growth. Long‐term experimental drought decreased the ABI for the two forest canopy types and the standing density for the shrubland. Water availabilities in winter–spring (SPEIs) were positively correlated with the ABI and standing density. Moreover, experimental drought decreased the vegetation growth in extreme dry years for the shrubland. We propose that future work not only study the vegetation dynamics with physiological, phenological and demographical changes in long‐term processes and across climate gradients, but also should explore the changes of multiple functions simultaneously (e.g. multifunctionality) under long‐term processes and extremes. This type of analysis of long‐term data is essential to understand and predict biodiversity loss, composition shifts, declines in ecosystem function and carbon budgets at temporal and spatial scales, to enable policy makers to design and implement strategies for the maintenance of sustainable ecosystem function under future climate change scenarios.     

Twentieth century increase of Scots pine radial growth in NE Spain shows strong climate interactions

Stem radial growth responds to environmental conditions, and has been widely used as a proxy to study long‐term patterns of tree growth and to assess the impact of environmental changes on growth patterns. In this study, we use a tree ring dataset from the Catalan Ecological and Forest Inventory to study the temporal variability of Scots pine (Pinus sylvestris L.) stem growth during the 20th century across a relatively large region (Catalonia, NE Spain) close to the southern limit of the distribution of the species. Basal area increment (BAI) was modelled as a function of tree size and environmental variables by means of mixed effects models. Our results showed an overall increase of 84% in Scots pine BAI during the 20th century, consistent with most previous studies for temperate forests. This trend was associated with increased atmospheric CO₂ concentrations and, possibly, with a general increase in nutrient availability, and we interpreted it as a fertilization effect. Over the same time period, there was also a marked increase in temperature across the study region (0.19 °C per decade on average). This warming had a negative impact on radial growth, particularly at the drier sites, but its magnitude was not enough to counteract the fertilization effect. In fact, the substantial warming observed during the 20th century in the study area did not result in a clear pattern of increased summer drought stress because of the large variability in precipitation, which did not show any clear time trend. But the situation may change in the future if temperatures continue to rise and/or precipitation becomes scarcer. Such a change could potentially reverse the temporal trend in growth, particularly at the driest sites, and is suggested in our data by the relative constancy of radial growth after ca. 1975, coinciding with the warmer period. If this situation is representative of other relatively dry, temperate forests, the implications for the regional carbon balance would be substantial.     

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...     

土壤干旱对辽东栎、大叶细裂槭幼苗生长及水分利用的影响

在盆栽条件下研究了不同土壤干旱条件对辽东栎和大叶细裂槭幼苗生长及水分利用率的影响。结果表明,随干旱胁迫程度加剧,两树种叶水势均下降,随胁迫时间延长水势又有一回升趋势,显示出树种自身的调节能力,大叶细裂槭调节水势的能力强于辽东栎。叶水势与叶片含水量关系密切,水势的高低决定叶片水分状况,水分含量又决定了幼苗的生长状况。大叶细裂槭新梢在5、6月增长增粗迅速,而辽东栎在4月、5月生长迅速及叶面积扩展快,适宜水分下增粗可持续至9月份。不同土壤水分处理下两树种水分利用效率差异明显。在中度干旱下整体WUE有所上升,尤其是辽东栎提高约10％;大叶细裂槭的根冠比在重度干旱下可达2以上,说明它对干旱的适应性很强,在严重干旱胁迫下也能长期维持生命,成活率高于辽东栎,接近100％,可见其适应性强于辽东栎。     

Water temperature and fish growth: otoliths predict growth patterns of a marine fish in a changing climate

Ecological modeling shows that even small, gradual changes in body size in a fish population can have large effects on natural mortality, biomass, and catch. However, efforts to model the impact of climate change on fish growth have been hampered by a lack of long‐term (multidecadal) data needed to understand the effects of temperature on growth rates in natural environments. We used a combination of dendrochronology techniques and additive mixed‐effects modeling to examine the sensitivity of growth in a long‐lived (up to 70 years), endemic marine fish, the western blue groper (Achoerodus gouldii), to changes in water temperature. A multi‐decadal biochronology (1952–2003) of growth was constructed from the otoliths of 56 fish collected off the southwestern coast of Western Australia, and we tested for correlations between the mean index chronology and a range of potential environmental drivers. The chronology was significantly correlated with sea surface temperature in the region, but common variance among individuals was low. This suggests that this species has been relatively insensitive to past variations in climate. Growth increment and age data were also used in an additive mixed model to predict otolith growth and body size later this century. Although growth was relatively insensitive to changes in temperature, the model results suggested that a fish aged 20 in 2099 would have an otolith about 10% larger and a body size about 5% larger than a fish aged 20 in 1977. Our study shows that species or populations regarded as relatively insensitive to climate change could still undergo significant changes in growth rate and body size that are likely to have important effects on the productivity and yield of fisheries.     

Photosynthesis and growth reduction with warming are driven by nonstomatal limitations in a Mediterranean semi‐arid shrub

Whereas warming enhances plant nutrient status and photosynthesis in most terrestrial ecosystems, dryland vegetation is vulnerable to the likely increases in evapotranspiration and reductions in soil moisture caused by elevated temperatures. Any warming‐induced declines in plant primary production and cover in drylands would increase erosion, land degradation, and desertification. We conducted a four‐year manipulative experiment in a semi‐arid Mediterranean ecosystem to evaluate the impacts of a ~2°C warming on the photosynthesis, transpiration, leaf nutrient status, chlorophyll content, isotopic composition, biomass growth, and postsummer survival of the native shrub Helianthemum squamatum. We predicted that warmed plants would show reduced photosynthetic activity and growth, primarily due to the greater stomatal limitation imposed by faster and more severe soil drying under warming. On average, warming reduced net photosynthetic rates by 36% across the study period. Despite this strong response, warming did not affect stomatal conductance and transpiration. The reduction of peak photosynthetic rates with warming was more pronounced in a drought year than in years with near‐average rainfall (75% and 25–40% reductions relative to controls, respectively), with no indications of photosynthetic acclimation to warming through time. Warmed plants had lower leaf N and P contents, δ¹³C, and sparser and smaller leaves than control plants. Warming reduced shoot dry mass production by 31%. However, warmed plants were able to cope with large reductions in net photosynthesis, leaf area, and shoot biomass production without changes in postsummer survival rates. Our findings highlight the key role of nonstomatal factors (biochemical and/or nutritional) in reducing net carbon assimilation rates and growth under warming, which has important implications for projections of plant carbon balance under the warmer and drier climatic scenario predicted for drylands worldwide. Projected climate warming over the coming decades could reduce net primary production by about one‐third in semi‐arid gypsum shrublands dominated by H. squamatum.     

Trophic mismatch and its effects on the growth of young in an Arctic herbivore

In highly seasonal environments, timing of breeding of organisms is typically set to coincide with the period of highest resource availability. However, breeding phenology may not change at a rate sufficient to keep up with rapid changes in the environment in the wake of climate change. The lack of synchrony between the phenology of consumers and that of their resources can lead to a phenomenon called trophic mismatch, which may have important consequences on the reproductive success of herbivores. We analyzed long‐term data (1991–2010) on climate, plant phenology and the reproduction of a long‐distance Arctic migrant, the greater snow goose (Chen caerulescens atlantica), in order to examine the effects of mismatched reproduction on the growth of young. We found that geese are only partially able to adjust their breeding phenology to compensate for annual changes in the timing of high‐quality food plants, leading to mismatches of up to 20 days between the two. The peak of nitrogen concentration in plants, an index of their nutritive quality for goslings, occurred earlier in warm springs with an early snow melt. Likewise, mismatch between hatch dates of young and date of peak nitrogen was more important in years with early snow melt. Gosling body mass and structural size at fledging was reduced when trophic mismatch was high, particularly when the difference between date of peak nitrogen concentration and hatching was >9 days. Our results support the hypothesis that trophic mismatch can negatively affect the fitness of Arctic herbivores and that this is likely to be exacerbated by rising global temperatures.     

Drought tolerance and growth in populations of a wide‐ranging tree species indicate climate change risks for the boreal north

Choosing drought‐tolerant planting stock in reforestation programs may help adapt forests to climate change. To inform such reforestation strategies, we test lodgepole pine (Pinus contorta Doug. ex Loud. var latifolia Englm.) population response to drought and infer potential benefits of a northward transfer of seeds from drier, southern environments. The objective is addressed by combining dendroecological growth analysis with long‐term genetic field trials. Over 500 trees originating from 23 populations across western North America were destructively sampled in three experimental sites in southern British Columbia, representing a climate warming scenario. Growth after 32 years from provenances transferred southward or northward over long distances was significantly lower than growth of local populations. All populations were affected by a severe natural drought event in 2002. The provenances from the most southern locations showed the highest drought tolerance but low productivity. Local provenances were productive and drought tolerant. Provenances from the boreal north showed lower productivity and less drought tolerance on southern test sites than all other sources, implying that maladaptation to drought may prevent boreal populations from taking full advantage of more favorable growing conditions under projected climate change.