Antonio Gazol

To die or not to die: early warnings of tree dieback in response to a severe drought

This study was supported by projects CGL2011-26654 (Spanish Ministry of Economy and Competitiveness), 1032S/2013 and 387/2011 (Organismo Autonomo Parques Nacionales, Spanish Ministry of Agriculture and Environment, Spain). We thank ARAID for supporting J.J.C. and the AEET for providing climatic data. We also thank M. Maestro and E. Lahoz for performing the chemical analyses and R. Hernandez, A.Q. Alla and E. Gonzalez de Andres for their help in the field. The authors declare no conflict of interest.

Wood anatomy and carbon-isotope discrimination support long-term hydraulic deterioration as a major cause of drought-induced dieback.

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought-induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long-term by quantifying wood-anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water-use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long-lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2-5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long-term capacity of trees to withstand drought-induced dieback.

Microfragmentation concept explains non-positive environmental heterogeneity–diversity relationships

Although recent studies have revealed that the relationship between diversity and environmental heterogeneity is not always positive, as classical niche theory predicts, scientists have had difficulty interpreting these results from an ecological perspective. We propose a new concept—microfragmentation—to explain how small-scale heterogeneity can have neutral or even negative effect on species diversity. We define microfragmentation as a community level process of splitting habitat into a more heterogeneous environment that can have non-positive effects on the diversity through habitat loss and subsequent isolation. We provide support for the microfragmentation concept with results from spatially explicit heterogeneity–diversity model simulations, in which varying sets of species (with different ratios of specialist and generalist species) were modeled at different levels of configurational heterogeneity (meaning that only the habitat structure was changed, not its composition). Our results indicate that environmental heterogeneity can affect community diversity in the same way as fragmentation at the landscape level. Although generalist species might not be seriously affected by microfragmentation, the persistence of specialist species can be seriously disturbed by small-scale patchiness. The microfragmentation concept provides new insight into community level diversity dynamics and can influence conservation and management strategies.

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 ca 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 ca 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 ca 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 ca 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.

Soil Nutrient Content Influences the Abundance of Soil Microbes but Not Plant Biomass at the Small-Scale

Small-scale heterogeneity of abiotic and biotic factors is expected to play a crucial role in species coexistence. It is known that plants are able to concentrate their root biomass into areas with high nutrient content and also acquire nutrients via symbiotic microorganisms such as arbuscular mycorrhizal (AM) fungi. At the same time, little is known about the small-scale distribution of soil nutrients, microbes and plant biomass occurring in the same area. We examined small-scale temporal and spatial variation as well as covariation of soil nutrients, microbial biomass (using soil fatty acid biomarker content) and above- and belowground biomass of herbaceous plants in a natural herb-rich boreonemoral spruce forest. The abundance of AM fungi and bacteria decreased during the plant growing season while soil nutrient content rather increased. The abundance of all microbes studied also varied in space and was affected by soil nutrient content. In particular, the abundance of AM fungi was negatively related to soil phosphorus and positively influenced by soil nitrogen content. Neither shoot nor root biomass of herbaceous plants showed any significant relationship with variation in soil nutrient content or the abundance of soil microbes. Our study suggests that plants can compensate for low soil phosphorus concentration via interactions with soil microbes, most probably due to a more efficient symbiosis with AM fungi. This compensation results in relatively constant plant biomass despite variation in soil phosphorous content and in the abundance of AM fungi. Hence, it is crucial to consider both soil nutrient content and the abundance of soil microbes when exploring the mechanisms driving vegetation patterns.

Functional diversity enhances silver fir growth resilience to an extreme drought

Summary It is expected that extreme climate events such as droughts will increase both in severity and intensity as a consequence of forecasted climate change. Complementarity among tree species in resource acquisition strategy may reduce interspecific competition and increase the occurrence of facilitative interactions, resulting in an improved tree growth and resilience to extreme climatic events. However, the response of individual trees growing in more functionally diverse stands to extreme events is still under debate. We investigate the growth response of silver fir (Abies alba) to an extreme drought event which occurred in 1985-1986 in the central Spanish Pyrenees, and how the growth and resilience of trees relate to their neighbourhood functional diversity, stand heterogeneity, and to intraspecific and interspecific competition. Dendroecological methods were used to reconstruct radial growth. We recorded all tree species living around each sampled tree and calculated indices of intra- and interspecific competition among coexisting trees. Functional diversity around each focal tree was calculated by using the Rao quadratic entropy. Silver fir growth decreased significantly in response to the drought in the vast majority of trees, but most of them showed a growth recovery three years later. Despite most trees showed negative growth trends following the drought event (70%), those trees growing in more functionally diverse stands were more resilient, recovered more quickly and displayed greater growth and growth trends. Synthesis The presented findings support the growing body of evidence that shows a positive impact of diversity on forest function. In addition, we provide a direct linkage between tree growth response to climate at the individual tree scale and a direct quantification of neighbourhood functional diversity and competition. An enhanced functional diversity facilitating a more efficient use of resources at the interspecific level and potentially increasing facilitative interactions contributes to lessening the negative impacts of extreme events as droughts on forest growth. This article is protected by copyright. All rights reserved.

Impact of alien pines on local arbuscular mycorrhizal fungal communities—evidence from two continents

The introduction of alien plants can influence biodiversity and ecosystems. However, its consequences for soil microbial communities remain poorly understood. We addressed the impact of alien ectomycorrhizal (EcM) pines on local arbuscular mycorrhizal (AM) fungal communities in two regions with contrasting biogeographic histories: in South Africa, where no native EcM plant species are present; and in Argentina, where EcM trees occur naturally. The effect of alien pines on AM fungal communities differed between these regions. In South Africa, plantations of alien EcM pines exhibited lower AM fungal richness and significantly altered community composition, compared with native fynbos. In Argentina, the richness and composition of local AM fungal communities were similar in plantations of alien EcM pines and native forest. However, the presence of alien pines resulted in slight changes to the phylogenetic structure of root AM fungal communities in both regions. In pine clearcut areas in South Africa, the richness and composition of AM fungal communities were intermediate between the native fynbos and the alien pine plantation, which is consistent with natural regeneration of former AM fungal communities following pine removal. We conclude that the response of local AM fungal communities to alien EcM pines differs between biogeographic regions with different histories of species coexistence.

Mediterranean dwarf shrubs and coexisting trees present different radial-growth synchronies and responses to climate

Due to their diversity and dominance in environmentally harsh sites, Mediterranean dwarf shrubs are a valuable tool to understand the consequences of climatic variability on radial growth in woody plants. We evaluate the dendrochronological potential of three Mediterranean dwarf shrubs versus three coexisting tree species inhabiting cold- (Hormathophylla spinosa vs. Pinus sylvestris), mesic- (Ononis fruticosa vs. Abies alba), and xeric sites (Linum suffruticosum vs. Pinus halepensis). Cross-sectional wood sections of the three shrub species and cores in the case of trees were visually cross-dated and ring-widths were measured and converted into residual growth indices. We used linear mixed-effects models to assess how growth indices respond to local factors and climatic variables. The radial growth of the three dwarf shrub species was more asynchronous, i.e., ring-width series differed among conspecific individuals, than that of coexisting tree species. Growth asynchrony was higher for H. spinosa than for O. fruticosa and L. suffruticosum. Similarly, the ring-width series of O. fruticosa and L. suffruticosum was strongly correlated with that of coexisting tree species, while growth series of H. spinosa and P. sylvestris was not related at all. The growth of the three dwarf shrub species was influenced by the regional climatic conditions, but to a lesser degree than coexisting tree species. The highest responsiveness of growth to climate was observed in Mediterranean dwarf shrubs from xeric sites. However, local conditions are also major drivers of growth in Mediterranean dwarf shrubs as indicated by the stronger asynchrony in ring formation of these species as compared with coexisting trees, particularly in cold sites.

Scale-specific determinants of a mixed beech and oak seedling–sapling bank under different environmental and biotic conditions

The persistence of seedlings in the forest understorey is of major importance for the maintenance and regeneration of canopy trees in several forested ecosystems. In the present study, we examine the small-scale spatial pattern of a mixed beech and oak seedling–sapling bank in two areas of an unmanaged temperate deciduous forest with different environmental conditions. We used environmental, biotic and spatial variables to establish the main factors that explain the spatial pattern of these seedling–sapling banks at different scales. The stand structure in both areas was similar, but while in plot A beech dominated the canopy, plot B was dominated by oaks. In both areas, established beech individuals showed a clear reverse J-shaped distribution, whereas established oaks showed a unimodal distribution with only a few young individuals. Seedlings of beech and oak were distributed in aggregates, whereas beech saplings had a random distribution. At broader scales, the abundance of seedlings and saplings is affected by the environment as well as by inter-species competition, while at finer scales the spatial pattern is mainly influenced by stochastic processes, probably related to seed predation and establishment. The structure of the seedling–sapling bank indicates an advantage of beech over oak as far as regeneration is concerned. Beech seedlings and saplings tolerate the stress induced by the canopy and the understorey and persist for many years, while oak seedlings decline in a few years. Therefore, if current conditions persist, after canopy opening beech seedlings and saplings can grow rapidly into the canopy and the stands will move towards beech dominance.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.