Jaime Madrigal-González

Stand Structure and Recent Climate Change Constrain Stand Basal Area Change in European Forests: A Comparison Across Boreal, Temperate, and Mediterranean Biomes

European forests have a prominent role in the global carbon cycle and an increase in carbon storage has been consistently reported during the twentieth century. Any further increase in forest carbon storage, however, could be hampered by increases in aridity and extreme climatic events. Here, we use forest inventory data to identify the relative importance of stand structure (stand basal area and mean d.b.h.), mean climate (water availability), and recent climate change (temperature and precipitation anomalies) on forest basal area change during the late twentieth century in three major European biomes. Using linear mixed-effects models we observed that stand structure, mean climate, and recent climatic change strongly interact to modulate basal area change. Although we observed a net increment in stand basal area during the late twentieth century, we found the highest basal area increments in forests with medium stand basal areas and small to medium-sized trees. Stand basal area increases correlated positively with water availability and were enhanced in warmer areas. Recent climatic warming caused an increase in stand basal area, but this increase was offset by water availability. Based on recent trends in basal area change, we conclude that the potential rate of aboveground carbon accumulation in European forests strongly depends on both stand structure and concomitant climate warming, adding weight to suggestions that European carbon stocks may saturate in the near future.

Geography and major host evolutionary transitions shape the resource use of plant parasites

Significance Patterns of host use by parasites are commonly thought to be limited by phylogenetic constraints, yet little is known about the role of the geographic distribution of hosts and parasites in such patterns. We show that evolutionary patterns in host use by a family of plant parasites are largely determined by the geographical distribution of hosts and parasites. Such phylogenetic lability in host use results in repeated colonizations of distantly related plant lineages, even across major plant evolutionary transitions. Still, these transitions constitute significant adaptive barriers in the evolution of host use. Our results thus show that host plant use by parasitic mites hinges more on where the plant and the mite are than on phylogenetic constraints. The evolution of resource use in herbivores has been conceptualized as an analog of the theory of island biogeography, assuming that plant species are islands separated by phylogenetic distances. Despite its usefulness, this analogy has paradoxically led to neglecting real biogeographical processes in the study of macroevolutionary patterns of herbivore–plant interactions. Here we show that host use is mostly determined by the geographical cooccurrence of hosts and parasites in spider mites (Tetranychidae), a globally distributed group of plant parasites. Strikingly, geography accounts for most of the phylogenetic signal in host use by these parasites. Beyond geography, only evolutionary transitions among major plant lineages (i.e., gymnosperms, commelinids, and eudicots) shape resource use patterns in these herbivores. Still, even these barriers have been repeatedly overcome in evolutionary time, resulting in phylogenetically diverse parasite communities feeding on similar hosts. Therefore, our results imply that patterns of apparent evolutionary conservatism may largely be a byproduct of the geographic cooccurrence of hosts and parasites.

Functional diversity underlies demographic responses to environmental variation in European forests

Aim Biodiversity loss and climate-driven ecosystem modification are leading to substantial changes in forest structure and function. However, the effects of diversity on demographic responses to the environment are poorly understood. We tested the diversity hypothesis (measured through functional diversity) and the mass ratio hypothesis (measured through functional identity) in relation to tree growth, tree mortality and sapling abundance. We sought to determine whether functional diversity underlies demographic responses to environmental variation in European forests. Location Europe (Spain, Germany, Wallonia, Finland and Sweden). Methods We used data from five European national forest inventories from boreal to Mediterranean biomes (c. 700,000 trees in 54,000 plots and 143 tree species) and the main forest types across Europe (i.e. from needle-leaved evergreen forests to broad-leaved deciduous forests). For each forest type, we applied maximum likelihood techniques to quantify the relative importance of stand structure, climate and diversity (i.e. functional diversity and functional identity) as determinants of growth, mortality and sapling abundance. We also tested whether demographic responses to environmental conditions (including stand density, evapotranspiration and temperature anomalies) varied with functional diversity. Results Our results suggest that functional diversity has a positive effect on sapling abundance and growth rates in forests across Europe, while no effect was observed on tree mortality. Functional identity has a strong effect on mortality and sapling abundance, with greater mortality rates in forests dominated by needle-leaved individuals and a greater abundance of saplings in forests dominated by broad-leaved individuals. Furthermore, we observed that functional diversity modified the effects of stand density on demographic responses in Mediterranean forests and the influence of evapotranspiration and temperature anomalies in forests widely distributed across Europe. Main conclusion Our results suggest that functional diversity may play a key role in forest dynamics through complementarity mechanisms, as well as by modulating demographic responses to environmental variation.

Long-term Wood Production in Water-Limited Forests: Evaluating Potential CO2 Fertilization Along with Historical Confounding Factors

Increased aridity may have severe effects on productivity of dry forests. However, it remains unclear to what degree the positive effects of elevated CO2 (both increased carboxylation rates and enhanced water-use efficiency) may offset the negative effects of drought and climate warming. In forest ecosystems, it is particularly challenging to evaluate CO2 effects on productivity because the impacts of climate variability, competition, and management, combine to have longlasting effects on stand-level productivity. Here we address this problem using a unique long-term database containing repeated inventories of wood biomass for every decade from 1912 to 2002 in a pine forest (Pinus pinaster Ait.) in central Spain (≈7,500 ha.). The approach is based upon a combination of statistical analyses of long-term historical management data and mechanistic modeling which allows us to evaluate the effects of potential CO2 fertilization, climate, and stand structure on woody net primary production (W-NPP). We found a significant negative effect of drought on W-NPP during the first half of the twentieth century that diminishes at the turn of the century. Simulations with a process-based ecosystem model, ORCHIDEE, suggest that wood production under conditions that included CO2 fertilization produced a more highly correlated long-term W-NPP than simulations keeping CO2 values in preindustrial levels. Interestingly, however, the CO2 effect was only apparent when accounting for confounding factors such as competition and management legacies. Identifying CO2 fertilization on forest growth is a critical issue, and requires partitioning CO2 effects from confounding factors that have jointly shaped stand dynamics and carbon balance during the twentieth century.

Climate- and successional-related changes in functional composition of European forests are strongly driven by tree mortality

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late-successional short-statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm-like strategies at low water availability in forests formerly dominated by broad-leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm-like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.

Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe

Neglecting tree size and stand structure dynamics might bias the interpretation of the diversity-productivity relationship in forests. Here we show evidence that complementarity is contingent on tree size across large-scale climatic gradients in Europe. We compiled growth data of the 14 most dominant tree species in 32,628 permanent plots covering boreal, temperate and Mediterranean forest biomes. Niche complementarity is expected to result in significant growth increments of trees surrounded by a larger proportion of functionally dissimilar neighbours. Functional dissimilarity at the tree level was assessed using four functional types: i.e. broad-leaved deciduous, broad-leaved evergreen, needle-leaved deciduous and needle-leaved evergreen. Using Linear Mixed Models we show that, complementarity effects depend on tree size along an energy availability gradient across Europe. Specifically: (i) complementarity effects at low and intermediate positions of the gradient (coldest-temperate areas) were stronger for small than for large trees; (ii) in contrast, at the upper end of the gradient (warmer regions), complementarity is more widespread in larger than smaller trees, which in turn showed negative growth responses to increased functional dissimilarity. Our findings suggest that the outcome of species mixing on stand productivity might critically depend on individual size distribution structure along gradients of environmental variation.

Functional diversity underlies demographic responses to environmental variation in European forests (Forthcoming/Available Online)

Aim Biodiversity loss and climate-driven ecosystem modification are leading to substantial changes in forest structure and function. However, the effects of diversity on demographic responses to the environment are poorly understood. We tested the diversity hypothesis (measured through functional diversity) and the mass ratio hypothesis (measured through functional identity) in relation to tree growth, tree mortality and sapling abundance. We sought to determine whether functional diversity underlies demographic responses to environmental variation in European forests. Location Europe (Spain, Germany, Wallonia, Finland and Sweden). Methods We used data from five European national forest inventories from boreal to Mediterranean biomes (c. 700,000 trees in 54,000 plots and 143 tree species) and the main forest types across Europe (i.e. from needle-leaved evergreen forests to broad-leaved deciduous forests). For each forest type, we applied maximum likelihood techniques to quantify the relative importance of stand structure, climate and diversity (i.e. functional diversity and functional identity) as determinants of growth, mortality and sapling abundance. We also tested whether demographic responses to environmental conditions (including stand density, evapotranspiration and temperature anomalies) varied with functional diversity. Results Our results suggest that functional diversity has a positive effect on sapling abundance and growth rates in forests across Europe, while no effect was observed on tree mortality. Functional identity has a strong effect on mortality and sapling abundance, with greater mortality rates in forests dominated by needle-leaved individuals and a greater abundance of saplings in forests dominated by broad-leaved individuals. Furthermore, we observed that functional diversity modified the effects of stand density on demographic responses in Mediterranean forests and the influence of evapotranspiration and temperature anomalies in forests widely distributed across Europe. Main conclusion Our results suggest that functional diversity may play a key role in forest dynamics through complementarity mechanisms, as well as by modulating demographic responses to environmental variation.

Dendrochronology Course In Valsaín Forest, Segovia, Spain

Abstract This report describes an international summer course, “Tree Rings, Climate, Natural Resources, and Human Interaction”, held in Valsaín, Spain, in summer of 2012. The course, with 14 participants from three countries (Spain, Algeria, and Russia), included basic training in dendrochronology skills as well as applied projects in dendroclimatology, dendroecology and dendrogeomorphology.

Climatic Stress during Stand Development Alters the Sign and Magnitude of Age-Related Growth Responses in a Subtropical Mountain Pine

The modification of typical age-related growth by environmental changes is poorly understood, In part because there is a lack of consensus at individual tree level regarding age-dependent growth responses to climate warming as stands develop. To increase our current understanding about how multiple drivers of environmental change can modify growth responses as trees age we used tree ring data of a mountain subtropical pine species along an altitudinal gradient covering more than 2,200 m of altitude. We applied mixed-linear models to determine how absolute and relative age-dependent growth varies depending on stand development; and to quantify the relative importance of tree age and climate on individual tree growth responses. Tree age was the most important factor for tree growth in models parameterised using data from all forest developmental stages. Contrastingly, the relationship found between tree age and growth became non-significant in models parameterised using data corresponding to mature stages. These results suggest that although absolute tree growth can continuously increase along tree size when trees reach maturity age had no effect on growth. Tree growth was strongly reduced under increased annual temperature, leading to more constant age-related growth responses. Furthermore, young trees were the most sensitive to reductions in relative growth rates, but absolute growth was strongly reduced under increased temperature in old trees. Our results help to reconcile previous contrasting findings of age-related growth responses at the individual tree level, suggesting that the sign and magnitude of age-related growth responses vary with stand development. The different responses found to climate for absolute and relative growth rates suggest that young trees are particularly vulnerable under warming climate, but reduced absolute growth in old trees could alter the species’ potential as a carbon sink in the future.

Indirect facilitation by a liana might explain the dominance of a small tree in a temperate forest

Aims Lianas are expected to influence composition, structure and functioning of forest systems due to unequal distribution across the potential set of host plants. However, our understanding of mechanisms associated with preferences for specific hosts is still limited, and so is our ability to discern between endogenous and exogenous forces driving forest dynamics in the long run. In this paper, we evaluated whether the dominant liana Hedera helix can indirectly contribute to the eventual dominance of the small multi-stemmed tree Corylus avellana in a remnant temperate forest in central Iberian Peninsula from comparatively reduced liana infestation on C. avellana relative to co-occurring woody species. Methods through principal component analysis and co-occurrence analysis, we studied the distribution and spatial association between woody species and the liana H. helix. We analyzed the relationship between the number of species in a plot and the number of species infested by the liana to test the hypothesis that H. helix is a generalist liana. through generalized linear mixed models, we tested the dynamic-multi-stemmed growth form of C. avellana as a plausible life strategy to withstand, in the long run, the liana infestation. In particular, we tested (i) the relationship between stem size and the probability of H. helix infestation including all the tree species within plots and (ii) the relationship between stem size and mortality as evidence of the stem turn over in the tree C. avellana. Important Findings Our results indicate that H. helix and C. avellana significantly cooccur in mature stands of this remnant temperate forest where pioneer woody species are absent. Hedera helix severely infests all the woody species whenever stem size exceed ≈25 cm perimeter and there is physical contact at the base of the stem. this implies that all the trees in the community are potential hosts for H. helix. Mixed models indicate that both, infestation by H. helix and stem mortality, are positively related to C. avellana stem perimeter. Reduced longterm infestation of the liana by means of a multi-stemmed growth form with high stem turnover in C. avellana might be an advantage with respect to unipodial tree species. thus, the liana-tree coexistence pattern may be interpreted as an indirect positive interaction that, contrary to previous findings, results here in species dominance instead of species coexistence.