Weather underground: Subsurface hydrologic processes mediate tree vulnerability to extreme climatic drought

Drought extent and severity have increased and are predicted to continue to increase in many parts of the world. Understanding tree vulnerability to drought at both individual and species levels is key to ongoing forest management and preparation for future transitions in community composition. The influence of subsurface hydrologic processes is particularly important in water‐limited ecosystems, and is an under‐studied aspect of tree drought vulnerability. With California's 2013–2016 extraordinary drought as a natural experiment, we studied four co‐occurring woodland tree species, blue oak (Quercus douglasii), valley oak (Quercus lobata), gray pine (Pinus sabiniana), and California juniper (Juniperus californica), examining drought vulnerability as a function of climate, lithology and hydrology using regional aerial dieback surveys and site‐scale field surveys. We found that in addition to climatic drought severity (i.e., rainfall), subsurface processes explained variation in drought vulnerability within and across species at both scales. Regionally for blue oak, severity of dieback was related to the bedrock lithology, with higher mortality on igneous and metamorphic substrates, and to regional reductions in groundwater. At the site scale, access to deep subsurface water, evidenced by stem water stable isotope composition, was related to canopy condition across all species. Along hillslope gradients, channel locations supported similar environments in terms of water stress across a wide climatic gradient, indicating that subsurface hydrology mediates species’ experience of drought, and that areas associated with persistent access to subsurface hydrologic resources may provide important refugia at species’ xeric range edges. Despite this persistent overall influence of the subsurface environment, individual species showed markedly different response patterns. We argue that hydrologic niche segregation can be a useful lens through which to interpret these differences in vulnerability to climatic drought and climate change.     

Forest response to increasing typhoon activity on the Korean peninsula: evidence from oak tree‐rings

The globally observed trend of changing intensity of tropical cyclones over the past few decades emphasizes the need for a better understanding of the effects of such disturbance events in natural and inhabited areas. On the Korean Peninsula, typhoon intensity has increased over the past 100 years as evidenced by instrumental data recorded from 1904 until present. We examined how the increase in three weather characteristics (maximum hourly and daily precipitation, and maximum wind speed) during the typhoon activity affected old‐growth oak forests. Quercus mongolica is a dominant species in the Korean mountains and the growth releases from 220 individuals from three sites along a latitudinal gradient (33–38°N) of decreasing typhoon activity were studied. Growth releases indicate tree‐stand disturbance and improved light conditions for surviving trees. The trends in release events corresponded to spatiotemporal gradients in maximum wind speed and precipitation. A high positive correlation was found between the maximum values of typhoon characteristics and the proportion of trees showing release. A higher proportion of disturbed trees was found in the middle and southern parts of the Korean peninsula where typhoons are most intense. This shows that the releases are associated with typhoons and also indicates the differential impact of typhoons on the forests. Finally, we present a record of the changing proportion of trees showing release based on tree‐rings for the period 1770–1979. The reconstruction revealed no trend during the period 1770–1879, while the rate of forest disturbances increased rapidly from 1880 to 1979. Our results suggest that if typhoon intensity rises, as is projected by some climatic models, the number of forest disturbance events will increase thus altering the disturbance regime and ecosystem processes.     

The adaptive potential of plant populations in response to extreme climate events

The frequency and magnitude of extreme climate events are increasing with global change, yet we lack predictions and empirical evidence for the ability of wild populations to persist and adapt in response to these events. Here, we used Fisher's Fundamental Theorem of Natural Selection to evaluate the adaptive potential of Lasthenia fremontii, a herbaceous winter annual that is endemic to seasonally flooded wetlands in California, to alternative flooding regimes that occur during El Niño Southern Oscillation (ENSO) events. The results indicate that populations may exhibit greater adaptive potential in response to dry years than wet years, and that the relative performance of populations will change across climate scenarios. More generally, our findings show that extreme climate events can substantially change the potential for populations to adapt to climate change by modulating the expression of standing genetic variation and mean fitness.     

Predicting species responses to climate change: demography and climate microrefugia in California valley oak (Quercus lobata)

Anticipating species movement under climate change is a major focus in conservation. Bioclimate models are one of the few predictive tools for adaptation planning, but are limited in accounting for (i) climatic tolerances in preadult life stages that are potentially more vulnerable to warming; and (ii) local‐scale movement and use of climatic refugia as an alternative or complement to large‐scale changes in distribution. To assess whether these shortfalls can be addressed with field demographic data, we used California valley oak (Quercus lobata Nee), a long‐lived species with juvenile life stages known to be sensitive to climate. We hypothesized that the valley oak bioclimate model, based on adults, would overpredict the species' ability to remain in the projected persisting area, due to higher climate vulnerability of young life stages; and underpredict the potential for the species to remain in the projected contracting area in local‐scale refugia. We assessed the bioclimate model projections against actual demographic patterns in natural populations. We found that saplings were more constricted around surface water than adults in the projected contracting area. We also found that the climate envelope for saplings is narrower than that for adults. Saplings disappeared at a summer maximum temperature 3 °C below that associated with adults. Our findings indicate that rather than a complete shift northward and upward, as predicted by the species bioclimate model, valley oaks are more likely to experience constriction around water bodies, and eventual disappearance from areas exceeding a threshold of maximum temperature. Ours is the first study we know of to examine the importance of discrete life stage climate sensitivities in determining bioclimate modeling inputs, and to identify current climate change‐related constriction of a species around microrefugia. Our findings illustrate that targeted biological fieldwork can be central to understanding climate change‐related movement for long‐lived, sessile species.     

自然生态系统响应气候变化的脆弱性评价研究进展

以气候变暖为标志的全球气候变化已引起各国政府、国际组织和科学工作者的高度重视.气候变化给人类及自然生态系统带来的风险和危害日趋增大.生态系统脆弱性分析和评价是适应和减缓气候变化的关键和基础,已成为近年来气候变化领域和生态学领域的研究热点.目前国内外学者正在不同领域、不同空间尺度上开展响应气候变化的脆弱性评价,其中以自然生态系统为评价对象的脆弱性研究也有了长足的发展.本文通过对脆弱性的概念、气候变化脆弱性评价研究现状、自然生态系统响应气候变化的脆弱性定量评价方法的综述,探讨了该研究领域存在的问题和未来的发展前景.     

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.     

Genomic signals reveal past evolutionary dynamics of Quercus schottkyana and its response to future climate change

Understanding the past and future evolutionary dynamics of dominant species in a forest is important for guiding decisions for biodiversity conservation, forest management, and vegetation restoration. This study used Quercus schottkyana, a dominant tree in subtropical evergreen broad-leaved forests in southwest China, to investigate the influences of past environmental fluctuations and future changes in climate on the dynamics of tree demographics. Genomic data were obtained for 133 samples of Q. schottkyana from 22 populations using double-digest genotyping by sequencing. The single-nucleotide polymorphisms in the genome showed a uniform distribution. Based on principal component analysis and Admixture analysis, two distinct lineages and one mixed group were identified that corresponded to their geographical distribution. Approximate Bayesian computation analyses provided evidence that the divergence among Q. schottkyana populations could be driven by the collision between the Asian and Indian plates during the Miocene, and that climatic fluctuations in the late Pleistocene led to the introgression. The analysis of genotype-environment relationships showed that annual precipitation and geographic distance were associated with spatial genetic variation. Populations of Q. schottkyana in the northern area of the Jinsha River basin were predicted to be the most vulnerable to future climate change. To increase genetic diversity in the northern Jinsha River basin and to buffer threats from future climate change, managers could use a mixture of local and alien seeds during forest restoration and management. This case study can promote further investigations into assessing how past and future climate change impacts genetic divergence and local adaptation of trees in forests.     

Early response of the platypus to climate warming

Combining a climatic envelope modelling technique with more than two centuries (1800–2009) of distribution records has revealed the effects of a changing climate on the egg‐laying monotreme, the platypus, Ornithorhynchus anatinus. We show that the main factor associated with platypus occurrence switched from aquatic habitat availability (estimated by rainfall) to thermal tolerances (estimated by annual maximum temperature) in the 1960s. This correlates directly with the change in the annual maximum temperature anomaly from cooler to warmer conditions in southeastern Australia. Modelling of platypus habitat under emission scenarios (A1B, A2, B1 and B2) revealed large decreases (>30%) in thermally suitable habitat by 2070. This reduction, compounded by increasing demands for water for agriculture and potable use, suggests that there is real cause for concern over the future status of this species, and highlights the need for restoration of thermal refugia within the platypus’ modelled range.     

Local trampling disturbance effects on alpine plant populations and communities: Negative implications for climate change vulnerability

Global change is modifying species communities from local to landscape scales, with alterations in the abiotic and biotic determinants of geographic range limits causing species range shifts along both latitudinal and elevational gradients. An important but often overlooked component of global change is the effect of anthropogenic disturbance, and how it interacts with the effects of climate to affect both species and communities, as well as interspecies interactions, such as facilitation and competition. We examined the effects of frequent human trampling disturbances on alpine plant communities in Switzerland, focusing on the elevational range of the widely distributed cushion plant Silene acaulis and the interactions of this facilitator species with other plants. Examining size distributions and densities, we found that disturbance appears to favor individual Silene growth at middle elevations. However, it has negative effects at the population level, as evidenced by a reduction in population density and reproductive indices. Disturbance synergistically interacts with the effects of elevation to reduce species richness at low and high elevations, an effect not mitigated by Silene. In fact, we find predominantly competitive interactions, both by Silene on its hosted and neighboring species and by neighboring (but not hosted) species on Silene. Our results indicate that disturbance can be beneficial for Silene individual performance, potentially through changes in its neighboring species community. However, possible reduced recruitment in disturbed areas could eventually lead to population declines. While other studies have shown that light to moderate disturbances can maintain high species diversity, our results emphasize that heavier disturbance reduces species richness, diversity, as well as percent cover, and adversely affects cushion plants and that these effects are not substantially reduced by plant–plant interactions. Heavily disturbed alpine systems could therefore be at greater risk for upward encroachment of lower elevation species in a warming world.     

Shifts in population size structure for a drying‐tolerant fish in response to extreme drought

For freshwater systems, climate change‐induced alterations to drought regimes are a considerable threat to already threatened species. This is particularly poignant for kōwaro (or Canterbury mudfish, Neochanna burrowsius), a critically endangered fish largely restricted to drying‐prone waterways on the Canterbury Plains, New Zealand. By comparing three catchment‐wide surveys (2007, 2010, 2015) within the Waianiwaniwa Valley, we assessed the scale and magnitude of population change induced by 2 years of consecutive drought (2014–15), when compared to surveys during wetter conditions (2007, 2010). The droughts triggered a catchment‐wide switch from adult‐dominated populations to populations comprised of juveniles indicated by a significant reduction in median size (~95 mm during the wet to ~60 mm after drought). In comparison, population abundances were highly variable, indicated by no significant change in catch‐per‐unit‐effort. The large variation in catch rates and connection of median size to reproductive potential mean median size will be useful to measure in monitoring to infer potential changes to population resilience, particularly during extreme events. Furthermore, because N. burrowsius could be regarded as extremophile fish, already restricted to harsh habitats, they are likely to become increasingly threatened by climate change. Thus, tools that allow for insightful comparisons between populations, such as a population resilience framework based on both abundance and body size distribution, will be increasingly important for pragmatic decision‐making for targeted conservation measures.     

Phylogeography of two closely related species of Allium endemic to East Asia: Population evolution in response to climate oscillations

This study investigated the effects of climate oscillations on the evolution of two closely related Allium species, A. neriniflorum and A. tubiflorum. We sequenced three cp DNA (cpDNA) fragments (rps16, rpl32‐trnL, and trnD‐trnT, together approximately 2,500 bp in length) of two closely related Allium species, with samples from 367 individuals in 47 populations distributed across the total range of these species. The interspecific and intraspecific divergence times of the two species were in the Quaternary glaciation. The population divergence was high for the cpDNA variation, suggesting a significant phylogeographic structure (N_ST = 0.844, G_ST = 0.798, p < 0.05). Remarkable ecological differentiation was also revealed by Niche models and statistical analyses. Our results suggest the speciation event of the two species was triggered by violent climatic changes during the Quaternary glaciation.     

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.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

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.     

Unpacking the mechanisms captured by a correlative species distribution model to improve predictions of climate refugia

Climate refugia are regions that animals can retreat to, persist in and potentially then expand from under changing environmental conditions. Most forecasts of climate change refugia for species are based on correlative species distribution models (SDMs) using long‐term climate averages, projected to future climate scenarios. Limitations of such methods include the need to extrapolate into novel environments and uncertainty regarding the extent to which proximate variables included in the model capture processes driving distribution limits (and thus can be assumed to provide reliable predictions under new conditions). These limitations are well documented; however, their impact on the quality of climate refugia predictions is difficult to quantify. Here, we develop a detailed bioenergetics model for the koala. It indicates that range limits are driven by heat‐induced water stress, with the timing of rainfall and heat waves limiting the koala in the warmer parts of its range. We compare refugia predictions from the bioenergetics model with predictions from a suite of competing correlative SDMs under a range of future climate scenarios. SDMs were fitted using combinations of long‐term climate and weather extremes variables, to test how well each set of predictions captures the knowledge embedded in the bioenergetics model. Correlative models produced broadly similar predictions to the bioenergetics model across much of the species' current range – with SDMs that included weather extremes showing highest congruence. However, predictions in some regions diverged significantly when projecting to future climates due to the breakdown in correlation between climate variables. We provide unique insight into the mechanisms driving koala distribution and illustrate the importance of subtle relationships between the timing of weather events, particularly rain relative to hot‐spells, in driving species–climate relationships and distributions. By unpacking the mechanisms captured by correlative SDMs, we can increase our certainty in forecasts of climate change impacts on species.     

Detecting nonlinear response of spring phenology to climate change by Bayesian analysis

The impact of climate change on the advancement of plant phenological events has been heavily studied in the last decade. Although the majority of spring plant phenological events have been trending earlier, this is not universally true. Recent work has suggested that species that are not advancing in their spring phenological behavior are responding more to lack of winter chill than increased spring heat. One way to test this hypothesis is by evaluating the behavior of a species known to have a moderate to high chilling requirement and examining how it is responding to increased warming. This study used a 60‐year data set for timing of leaf‐out and male flowering of walnut (Juglans regia) cultivar ‘Payne’ to examine this issue. The spring phenological behavior of ‘Payne’ walnut differed depending on bud type. The vegetative buds, which have a higher chilling requirement, trended toward earlier leaf‐out until about 1994, when they shifted to later leaf‐out. The date of male bud pollen shedding advanced over the course of the whole record. Our findings suggest that many species which have exhibited earlier bud break are responding to warmer spring temperatures, but may shift into responding more to winter temperatures (lack of adequate chilling) as warming continues.