Interacting effects of changes in climate and forest cover on mortality and growth of the southernmost European fir forests

Aims The combined effects of changes in climate and land use on tree mortality and growth patterns have rarely been addressed. Relict tree species from the Mediterranean Basin serve as appropriate models to investigate these effects, since they grow in climatically stressed areas which have undergone intense cover changes. The aim is to use climate, aerial photographs, stand structure and radial‐growth data to explain the mortality and historical patterns of growth of Abies pinsapo in the area where this relict species was first protected. Location Sierra de las Nieves, West Baetic Range, southern Spain. Methods We assessed variations of tree cover in A. pinsapo forests through image analyses of aerial photographs spanning the last 50 years. We sampled 31 stands to assess current altitudinal patterns of forest structure and mortality. We evaluated the relationships between radial growth and regional climate using linear models in three sites at different elevations. Results Regional warming and a decrease in precipitation were detected. Forest tree cover increased at all elevations from 1957 until 1991, but it afterwards decreased below 1100 m. Currently, the likelihood of tree mortality increases downwards and is associated with dense, closed stands with a low living basal area. In contrast to previous droughts, a sharp synchronized reduction in tree growth, not fully accounted for in linear climate–growth models, occurred at low elevations in 1994–95, but not upwards. It was preceded by a weakening of the negative association between low‐elevation growth and water deficit since the late 1970s. Conclusions The intense densification of A. pinsapo forests following strict protection measures in the late 1950s enhanced the vulnerability of climate‐sensitive A. pinsapo forests to recent drier conditions. Such abrupt land‐use changes help to explain recent patterns of mortality and growth decline in low‐elevation A. pinsapo forests.     

Past and future evolution of Abies alba forests in Europe – comparison of a dynamic vegetation model with palaeo data and observations

Information on how species distributions and ecosystem services are impacted by anthropogenic climate change is important for adaptation planning. Palaeo data suggest that Abies alba formed forests under significantly warmer‐than‐present conditions in Europe and might be a native substitute for widespread drought‐sensitive temperate and boreal tree species such as beech (Fagus sylvatica) and spruce (Picea abies) under future global warming conditions. Here, we combine pollen and macrofossil data, modern observations, and results from transient simulations with the LPX‐Bern dynamic global vegetation model to assess past and future distributions of A. alba in Europe. LPX‐Bern is forced with climate anomalies from a run over the past 21 000 years with the Community Earth System Model, modern climatology, and with 21st‐century multimodel ensemble results for the high‐emission RCP8.5 and the stringent mitigation RCP2.6 pathway. The simulated distribution for present climate encompasses the modern range of A. alba, with the model exceeding the present distribution in north‐western and southern Europe. Mid‐Holocene pollen data and model results agree for southern Europe, suggesting that at present, human impacts suppress the distribution in southern Europe. Pollen and model results both show range expansion starting during the Bølling–Allerød warm period, interrupted by the Younger Dryas cold, and resuming during the Holocene. The distribution of A. alba expands to the north‐east in all future scenarios, whereas the potential (currently unrealized) range would be substantially reduced in southern Europe under RCP8.5. A. alba maintains its current range in central Europe despite competition by other thermophilous tree species. Our combined palaeoecological and model evidence suggest that A. alba may ensure important ecosystem services including stand and slope stability, infrastructure protection, and carbon sequestration under significantly warmer‐than‐present conditions in central Europe.     

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.     

Growth response of subalpine fir (Abies lasiocarpa) to climate in the Olympic Mountains, Washington, USA

Growth response of subalpine fir (Abies lasiocarpa) to climate was studied across its local geographical and elevation range in the Olympic Mountains, Washington. A dendroecological analysis of subalpine fir across a range of elevations (1350-1850 m) and annual precipitation (125-350 cm y^?1), was used to compare environmental factors affecting growth. Climate-growth relationships were explored using Pearson product-moment correlation coefficients; partial correlation analysis was used to assess relationships among site chronologies and climatic variables. Radial growth is negatively correlated with winter precipitation at high elevation and wet sites, but not at low and middle elevation dry sites. Growth is positively correlated with current growing season temperature at all sites; however, growth is negatively correlated with previous year August temperature, indicating that climate affects growth in subsequent years. Positive correlations between growth and summer precipitation during the growing season at low and middle elevation dry sites suggest that soil moisture is partially limiting to growth on these sites. If the climate of the Pacific Northwest becomes warmer and drier, then subalpine fir growth may increase at high elevation and wet sites, but may decrease at lower elevation dry sites in the Olympic Mountains. However, the growth response of subalpine fir to potentially rapid climate change will not be uniform because subalpine fir grows over a wide range of topographic features, habitats, and local climates at different geographical scales. A comparison of growth response to current growing season temperature suggests that the temperature-related growth response of subalpine fir is not adequately described by the parabolic curve used in JABOWA-based models.     

From pattern to process: linking intrinsic water‐use efficiency to drought‐induced forest decline

The rise in atmospheric CO₂ concentrations (Ca) has been related to tree growth enhancement and increasing intrinsic water‐use efficiency (iWUE). However, the extent that rising Ca has led to increased long‐term iWUE and whether climate could explain deviations from expected Ca‐induced growth enhancement are still poorly understood. The aim of this research was to use Ca and local climatic variability to explain changes during the 20th century in growth and tree ring and needle δ¹³C in declining and nondeclining Abies alba stands from the Spanish Pyrenees, near the southern distribution limit of this species. The temporal trends of iWUE were calculated under three theoretical scenarios for the regulation of plant‐gas exchange at increasing Ca. We tested different linear mixed‐effects models by multimodel selection criteria to predict basal area increment (BAI), a proxy of tree radial growth, using these scenarios and local temperature together with precipitation data as predictors. The theoretical scenario assuming the strongest response to Ca explained 66–81% of the iWUE variance and 28–56% of the observed BAI variance, whereas local climatic variables together explained less than 11–21% of the BAI variance. Our results are consistent with a drought‐induced limitation of the tree growth response to rising CO₂ and a decreasing rate of iWUE improvement from the 1980s onward in declining A. alba stands subjected to lower water availability.     

Fungi in ectomycorrhizal associations of silver fir (Abies alba Miller) in Central Italy

 Naturally occurring ectomycorrhizas of silver fir (Abies alba Miller) were studied in two stands, one natural and one artificial, situated in Central Italy. A total of 25 mycorrhizal types was classified, for eight of which the mycobiont was identified at the species level. Analysis of macroscopic and microscopic features and matching of field-collected carpophores with associated mycorrhizas led to the tentative identification of several other types encountered during this study, at least at the genus level. No significant differences were noticed between natural and artificial stands in the relative richness of mycorrhizal types found on A. alba, indicating the maturity of the artificial stand with regard to succession of ectomycorrhizal fungi. Confocal laser scanning microscopy was used for visualization of mycorrhizal structures formed by Lactarius spp., without the need for specific staining with a fluorochrome, thanks to latex autofluorescence. This technique allowed observation of several structures in greater detail than with conventional light microscopy. Accepted: 13 February 1998     

Disparate effects of global‐change drivers on mountain conifer forests: warming‐induced growth enhancement in young trees vs. CO2 fertilization in old trees from wet sites

Theory predicts that the postindustrial rise in the concentration of CO₂ in the atmosphere (c_a) should enhance tree growth either through a direct fertilization effect or indirectly by improving water use efficiency in dry areas. However, this hypothesis has received little support in cold‐limited and subalpine forests where positive growth responses to either rising c_a or warmer temperatures are still under debate. In this study, we address this issue by analyzing an extensive dendrochronological network of high‐elevation Pinus uncinata forests in Spain (28 sites, 544 trees) encompassing the whole biogeographical extent of the species. We determine if the basal area increment (BAI) trends are linked to climate warming and increased c_a by focusing on region‐ and age‐dependent responses. The largest improvement in BAI over the past six centuries occurred during the last 150 years affecting young trees and being driven by recent warming. Indeed, most studied regions and age classes presented BAI patterns mainly controlled by temperature trends, while growing‐season precipitation was only relevant in the driest sites. Growth enhancement was linked to rising c_a in mature (151–300 year‐old trees) and old‐mature trees (301–450 year‐old trees) from the wettest sites only. This finding implies that any potential fertilization effect of elevated c_a on forest growth is contingent on tree features that vary with ontogeny and it depends on site conditions (for instance water availability). Furthermore, we found widespread growth decline in drought‐prone sites probably indicating that the rise in c_a did not compensate for the reduction in water availability. Thus, warming‐triggered drought stress may become a more important direct driver of growth than rising c_a in similar subalpine forests. We argue that broad approaches in biogeographical and temporal terms are required to adequately evaluate any effect of rising c_a on forest growth.     

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.     

Regional patterns of increasing Swiss needle cast impacts on Douglas‐fir growth with warming temperatures

The fungal pathogen, Phaeocryptopus gaeumannii, causing Swiss needle cast (SNC) occurs wherever Douglas‐fir is found but disease damage is believed to be limited in the U.S. Pacific Northwest (PNW) to the Coast Range of Oregon and Washington (Hansen et al., Plant Disease, 2000, 84, 773; Rosso & Hansen, Phytopathology, 2003, 93, 790; Shaw, et al., Journal of Forestry, 2011, 109, 109). However, knowledge remains limited on the history and spatial distribution of SNC impacts in the PNW. We reconstructed the history of SNC impacts on mature Douglas‐fir trees based on tree‐ring width chronologies from western Oregon. Our findings show that SNC impacts on growth occur wherever Douglas‐fir is found and is not limited to the coastal fog zone. The spatiotemporal patterns of growth impact from SNC disease were synchronous across the region, displayed periodicities of 12–40 years, and strongly correlated with winter and summer temperatures and summer precipitation. The primary climatic factor limiting pathogen dynamics varied spatially by location, topography, and elevation. SNC impacts were least severe in the first half of the 20th century when climatic conditions during the warm phase of the Pacific Decadal Oscillation (1924–1945) were less conducive to pathogen development. At low‐ to mid‐elevations, SNC impacts were most severe in 1984–1986 following several decades of warmer winters and cooler, wetter summers including a high summer precipitation anomaly in 1983. At high elevations on the west slope of the Cascade Range, SNC impacts peaked several years later and were the greatest in the 1990s, a period of warmer winter temperatures. Climate change is predicted to result in warmer winters and will likely continue to increase SNC severity at higher elevations, north along the coast from northern Oregon to British Columbia, and inland where low winter temperatures currently limit growth of the pathogen. Our findings indicate that SNC may become a significant forest health problem in areas of the PNW beyond the coastal fog zone.     

Vegetation history of Abies alba Mill. (silver fir) in Switzerland – pollen analytical and genetic surveys related to aspects of vegetation history of Picea abies (L.) H. Karsten (Norway spruce)

This contribution deals with some new aspects of the relationship between the vegetation history of Abies alba Mill. (silver fir) and genetic studies of this tree species in Switzerland. The results of the present study confirm the pollen analytical hypothesis that A. alba re-immigrated into Switzerland mainly from glacial refugia located in northern and central Italy. In particular, some distinct immigration routes of silver fir into the Ticino Alps, Valais, the Bernese Oberland (northwestern Alps), and Graubünden (eastern Alps) could be confirmed by genetic studies. Furthermore, the occurrence of other area-specific alleles indicates an additional influence from eastern European refugia on the Swiss gene pool. Moreover, genetic studies on Picea abies (L.) H. Karsten (Norway spruce) confirmed the general immigration routes from the eastern to the western Alps and from Savoie to the Jura Mountains. The combination of tree pollen and macrofossil analyses of Quaternary sediments with genetic studies of the same tree species represents a considerable research potential and is a new approach of floristic and genetic research. Received November 17, 2000 / Accepted April 27, 2001     

The effects of climate warming on the migratory status of early summer populations of Mythimna separata (Walker) moths: A case‐study of enhanced corn damage in central‐northern China, 1980–2016

Mythimna separata (Walker) moths captured in light traps were monitored in Luohe, central‐northern China, from 1980 to 2016. Annual average temperature recorded an increase of 0.298°C/10 years in this region in the period. Our results indicate that a rising April and May average temperature and earlier occurrences of days recording the highest day temperature (30°C) caused an advanced peak and increasing proportion of high ovarian development levels of first‐generation females in earlier summers. Results using Johnson's formulation of “oogenesis‐flight syndrome” indicate that increasing sexual maturity proportion has resulted in more emigrant individuals in the local first‐generation moth becoming residents, and then increased individuals rapidly in the local second‐generation moth since 2006. Consequences of this action have a boom in corn damage since 2007 in this region. Advanced peak dates of the first and second‐generation moth revealed the same response to increasing average monthly temperatures in the monitoring period. Increasing temperatures, the average May temperature exceeds or equal to 22°C, during the early 2000's may represent a physiological threshold for M. separata development. Our results suggest that climate warming may impact M. separata migratory status and cause a problem of crop production in this region.     

Population genetics analysis in a relic population of silver fir (Abies alba Mill.) in southern Italy: A comparison with microsatellites and reference data

We investigated the genetic diversity of the last natural population of Abies alba Mill. in Campania region (southern Italy) using nuclear and chloroplast microsatellite markers. We found high genetic diversity despite small population size and geographic isolation, especially when comparing our microsatellite data with information from the literature.     

Limited evidence for CO2‐related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO₂ may enhance photosynthesis and/or decrease stomatal conductance (g_s) thereby enhancing intrinsic water‐use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO₂ and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ¹³C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in g_s. Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing g_s may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.     

Seedling response to water stress in valley oak (Quercus lobata) is shaped by different gene networks across populations

Drought is a major stress for plants, creating a strong selection pressure for traits that enable plant growth and survival in dry environments. Many drought responses are conserved species‐wide responses, while others vary among populations distributed across heterogeneous environments. We tested how six populations of the widely distributed California valley oak (Quercus lobata) sampled from contrasting climates would differ in their response to soil drying relative to well‐watered controls in a common environment by measuring ecophysiological traits in 93 individuals and gene expression (RNA‐seq) in 42 individuals. Populations did not differ in their adjustment of turgor loss point during soil drying, suggesting a generalized species‐wide response. Differential expression analysis identified 689 genes with a common response to treatment across populations and 470 genes with population‐specific responses. Weighted gene co‐expression network analysis (WGCNA) identified groups of genes with similar expression patterns that may be regulated together (gene modules). Several gene modules responded differently to water stress among populations, suggesting regional differences in gene network regulation. Populations from sites with a high mean annual temperature responded to the imposed water stress with significantly greater changes in gene module expression, indicating that these populations may be locally adapted to respond to drought. We propose that this variation among valley oak populations provides a mechanism for differential tolerance to the increasingly frequent and severe droughts in California.     

Tolerance to multiple climate stressors: a case study of Douglas‐fir drought and cold hardiness

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Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.