Open Access
Synergies between climate change, biodiversity, ecosystem function and services, indirect drivers of change and human well-being in forests
(Springer, 2021) Imbert Rodríguez, Bosco; Blanco Vaca, Juan Antonio; Candel Pérez, David; Lo, Yueh-Hsin; González de Andrés, Ester; Yeste Yeste, Antonio; Herrera Álvarez, Ximena; Rivadeneira Barba, Gabriela; Liu, Yang; Chang, Shih-Chieh; Ciencias; Zientziak
Climate change is having impacts on the biodiversity and structure of many ecosystems. In this chapter, we focus on its impacts on forests. We will focus on how the potential climate change impacts on forest biodiversity and structure will have a reflection on the ecosystem services provided by forests, and therefore on the capacity of these ecosystems to support the Sustainable Development Goals set by the United Nations. The chapter will be organized in three sections, considering boreal, temperate, and tropical forests along each section. The first section will deal with the synergies or interactions between climate change, biodiversity, and ecosystem function with emphasis not only on plants but also on fungi, animals, and prokaryotes. Synergies between climate change and ecosystem services will be described and analyzed in the second section. To better link the first two sections, we will explore the relationships between ecosystem function, species traits, and ecosystem services. Finally, case studies for boreal, Mediterranean, and tropical forests will be presented, emphasizing the synergies between the above factors, the indirect drivers of change (demographic, economic, sociopolitical, science and technology, culture and religion), and human well-being (basic materials for a good life, health, good social relations, freedom of choice and actions) in forests.
Open Access
Tree-to-tree competition in mixed European beech-Scots pine forests has different impacts on growth and water-use efficiency depending on site condition.
(Wiley, 2018) González de Andrés, Ester; Camarero, Jesús Julio; Blanco Vaca, Juan Antonio; Imbert Rodríguez, Bosco; Lo, Yueh-Hsin; Sangüesa Barreda, G.; Castillo Martínez, Federico; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
Mixed conifer-hardwood forests can be more productive than pure forests and they are increasingly considered as ecosystems that could provide adaptation strategies in the face of global change. However, the combined effects of tree-to-tree competition, rising atmospheric CO2 concentrations and climate on such mixtures remain poorly characterized and understood.2. To fill this research gap, we reconstructed 34-year series (1980-2013) of growth (basal area increment, BAI) and intrinsic water-use efficiency (iWUE) of Scots pine (Pinus sylvestris L.)-European beech (Fagus sylvatica L.) mixed stands at two climati-cally contrasting sites located in the southwestern Pyrenees. We also gathered data on tree-to-tree competition and climate variables in order to test the hypotheses that (1) radial growth will be greater when exposed to inter- than to intraspecific competition, that is, when species complementarity occurs and (2) enhanced iWUE could be linked to improved stem radial growth.3. Growth of both species was reduced when intraspecific competition increased. Species complementarity was linked to improved growth of Scots pine at the continental site, while competition overrode any complementarity advantage at the drought-prone Mediterranean site. Beech growth did not show any significant response to pine admixture likely due to shade tolerance and the highly competitive nature of this species. Increasing interspecific competition drove recent iWUE changes, which increased in Scots pine but decreased in European beech. The iWUE enhancement did not involve any growth improvement in Scots pine. However, the positive BAIiWUE relationship found for beech suggests an enhanced beech growth in drought-prone sites due to improved water use.4. Synthesis. Complementarity may enhance growth in mixed forests. However, water scarcity can constrict light-related complementarity for shade intolerant species (Scots pine) in drought-prone sites. Basal area increment-intrinsic water-use efficiency relationships were negative for Scots pine and positive for European beech. These contrasting behaviours have got implications for coping with the expected increasing drought events in Scots pine-European beech mixtures located near ecological limit of the two species. Complementarity effects between tree species should be considered to avoid overestimating the degree of future carbon uptake by mixed conifer¿broadleaf forests.
Open Access
Drought limits tree growth more than greenness and reproduction: insights from five case studies in Spain
(KeAi Communications, 2025-08-01) Camarero, Jesús Julio; Rubio-Cuadrado, Álvaro; González de Andrés, Ester; Valeriano, Cristina; Pizarro, Manuel; Imbert Rodríguez, Bosco; Lo, Yueh-Hsin; Blanco Vaca, Juan Antonio; Ciencias; Zientziak; Institute for Multidisciplinary Research in Applied Biology - IMAB
Droughts impact forests by influencing various processes such as canopy greenness, tree growth, and reproduction, but most studies have only examined a few of these processes. More comprehensive assessments of forest responses to climate variability and water shortages are needed to improve forecasts of post-drought dynamics. Iberian forests are well-suited for evaluating these effects because they experience diverse climatic conditions and are dominated by various conifer and broadleaf species, many of which exhibit masting. We assessed how greenness, evaluated using the normalized difference vegetation index (NDVI), tree radial growth, and seed or cone production responded to drought in five tree species (three conifers: silver fir (Abies alba), Scots pine (Pinus sylvestris), and stone pine (Pinus pinea); two broadleaves: European beech (Fagus sylvatica) and holm oak (Quercus ilex) inhabiting sites with different aridity. We correlated these data with the standardized precipitation evapotranspiration index (SPEI) using the climate window analysis (climwin) package, which identifies the most relevant climate window. Drought constrained growth more than greenness and seed or cone production. Dry conditions led to high seed or cone production in species found in cool, moist sites (silver fir, beech, and Scots pine). We also found negative associations of cone production with summer SPEI in the drought-tolerant stone pine, which showed lagged growth−cone negative correlations. However, in the seasonally dry holm oak forests, severe droughts constrained both growth and acorn production, leading to a positive correlation between these variables. Drought impacts on greenness, growth, seed, and cone production depended on species phenology and site aridity. A negative correlation between growth and reproduction does not necessarily indicate trade-offs, as both may be influenced by similar climatic factors.
Open Access
Complex climate constraints of upper treeline formation in the Pyrenees
(Springer, 2015) González de Andrés, Ester; Camarero, Jesús Julio; Büntgen, Ulf; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
Alpine treelines are often induced by a threshold of minimum temperatures above which tree growth is generally not possible anymore. However, this worldwide pattern may not account for more drought-prone mountains, where changes in precipitation can also affect ecosystem functioning and productivity at higher elevations. Here, we aim to determine if tree-ring formation in ecotones along the Pyrenees indeed primarily depends on temperature means or also reflects hydroclimatic changes. The mean duration of the growing season at Pyrenean treelines was 177 days, lasting from mid-May to early November, which corresponds to a mean root-zone temperature of 8.2 °C. Temperatures during the early and late growing season were most critical for mountain pine growth at highest elevations, and above average spring temperatures also contributed to radial enlargement of tracheids. Sites showing more similar soil temperatures also presented more similar long-term growth trends indicating that local conditions trace regional influences on growth. Nevertheless, more wet conditions in summer enhanced growth at treelines subjected to more Mediterranean influence and therefore having a higher summer water deficit. A broader perspective reveals that soil moisture availability during spring and summer, however, also influence ring width formation at other treelines located in mid-latitude mountains separating temperate and semi-arid biomes. Our results suggest re-evaluating the idea of purely temperature-limited treelines, especially in mid-latitude mountains that experience moisture seasonality or where precipitation decreases with increasing elevation.
Open Access
CO2 fertilization plays a minor role in long-term carbon accumulation patterns in temperate pine forests in the southwestern Pyrenees
(Elsevier, 2019) Lo, Yueh-Hsin; Blanco Vaca, Juan Antonio; González de Andrés, Ester; Imbert Rodríguez, Bosco; Castillo Martínez, Federico; Institute for Multidisciplinary Research in Applied Biology - IMAB
Isolating the long-term fertilization effect of CO 2 from other climate- and site-related effects on tree growth has been proven a challenging task. To isolate long-term effects of [CO2] on water use efficiency at ecosystem level, we used the FORECAST Climate forest model, calibrated for Scots pine (Pinus sylvestris L.) forests in the southwestern Pyrenees, growing at a Mediterranean montane site and at a continental subalpine site. Future climate scenarios (RCP 4.5 and RCP 8.5) were generated using a battery of six climate models to estimate daily values of temperature and precipitation in a 90-year series. A factorial experiment was designed to disentangle the importance on C pools of three growing limiting factors (nitrogen limitation, climate (temperature + precipitation) limitation and atmospheric CO 2 concentration). The relative importance of each factor was quantified by comparing the scenario with the limitation of each individual factor turned on with the non-limitation scenario. Positive CO 2 fertilization due to improvement in water use efficiency was detected by the model, but its quantitative impact improving tree growth was minimum: its average increase in ecosystem C pools ranged from 0.3 to 0.9%. At the site with cooler climate conditions (continental), the main limitation for tree growth was climate. Such limitation will be reduced under climate change and the ecosystem will store more carbon. At the site with milder climate conditions (Mediterranean), N availability was the main limiting factor albeit modulated by water availability. Such limitation could be reduced under climate change as N cycling could accelerate (higher litterfall production and decomposition rates) but also increase if droughts become more frequent and severe. In addition, the magnitude of the uncertainty related to climate model selection was much more important than CO 2 fertilization, indicating that atmospheric processes are more important than tree physiological processes when defining how much carbon could be gained (or lost) in forests under climate change. In conclusion, due to the small changes in forest C pools caused by variation of atmospheric CO 2 concentrations compared to changes caused by other growth limiting factors (nutrients, climate), reducing uncertainty related to climate projections seems a more efficient way to reduce uncertainty in tree growth projections than increasing forest model complexity.
Open Access
Land use change effects on carbon and nitrogen stocks in the Pyrenees during the last 150 years: a modelling approach
(Elsevier, 2015) Lo, Yueh-Hsin; Blanco Vaca, Juan Antonio; Canals Tresserras, Rosa María; González de Andrés, Ester; San Emeterio Garciandía, Leticia; Imbert Rodríguez, Bosco; Castillo Martínez, Federico; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
In the southern Pyrenees, human population and therefore land uses have changed from forests to pastures, then crops, and back to pastures and secondary forests during the last two centuries. To understand what such rapid land use changes have meant for carbon (C) and nitrogen (N) stocks, we used data from two forest sites in the western Pyrenees, combined with regional data on pastures and crop production (potato, cereal), to calibrate the ecosystem-level model FORECAST. Then, we simulated 150-year of land use for each site, emulating historical changes. Our estimates show that the conversion from forests into pastures and crops created C and N deficits (378-427 Mg C ha-1, 4.0-4.6 Mg N ha-1) from which these sites are still recovering. The main ecological process behind the creation of these deficits was the loss of the ecological legacy of soil organic matter (SOM) created by the forest, particularly during conversion to farming. Pastures were able to reverse, stop or at least slow down the loss of such legacy. In conclusion, our work shows the deep impact of historical land use in ecosystem attributes, both in magnitude of removed C and N stocks and in duration of such impact. Also, the usefulness of ecological modelling in absence of historical data to estimate such changes is showcased, providing a framework for potential C and N stocks to be reached by climate change mitigation measures such as forest restoration.
Open Access
¿Están los bosques mixtos pirenaicos de pino silvestre y haya en el camino hacia la saturación por nitrógeno?
(Asociación Española de Ecología Terrestre, 2017) Blanco Vaca, Juan Antonio; San Emeterio Garciandía, Leticia; González de Andrés, Ester; Imbert Rodríguez, Bosco; Larrainzar Rodríguez, Estíbaliz; Peralta de Andrés, Francisco Javier; Lo, Yueh-Hsin; Castillo Martínez, Federico; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
Las actividades humanas causan altos niveles de deposición atmosférica crónica de N que pueden estar trastornando el ciclo del N en los bosques de los Pirineos occidentales. Para probar esta hipótesis, se han investigado los efectos de la deposición de N atmosférico en el ciclo de N en dos bosques mixtos de pino silvestre y haya en Navarra. Un bosque está situado a 1350 m de altitud y tiene un clima continental, mientras que el otro está situado a 650 m y tiene un clima mediterráneo húmedo. Pruebas preliminares indicaron una fijación biológica de N 2 atmosférico indetectable, así como la casi nula presencia de plantas con simbiontes fijadores como en la actividad de fijadores libres. Por lo tanto se asumió que la principal entrada de N en estos bosques es la deposición atmosférica. Se estimó la dependencia de la productividad de estos ecosistemas de la deposición por medio del modelo ecológico FORECAST, calibrado para estos sitios. Se simularon seis escenarios con tasas de deposición en un rango de 5 a 30 kg ha -1 año -1 . Los resultados indicaron que la productividad de estos bosques es dependiente de la deposición de N, pero indicios de saturación por N (aumento de lixiviación y carencia de aumento de productividad) indican que pueden saturarse a partir de 20-25 kg N ha -1 año -1 , unos 5-10 kg N ha -1 año -1 por encima de los niveles observados actualmente.
Open Access
Increased complementarity in water-limited environments in Scots pine and European beech mixtures under climate change
(Wiley, 2017) González de Andrés, Ester; Seely, Brad; Blanco Vaca, Juan Antonio; Imbert Rodríguez, Bosco; Lo, Yueh-Hsin; Castillo Martínez, Federico; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
Management of mixedwoods is advocated as an effective adaptation strategy to increase ecosystem resiliency in the context of climate change. While mixedwoods have been shown to have greater resource use efficiency relative to pure stands, considerable uncertainty remains with respect to the underlying ecological processes. We explored species interactions in Scots pine / European beech mixedwoods with the process-based model FORECAST Climate. The model was calibrated for two contrasting forests in the southwestern Pyrenees (northern Spain): a wet Mediterranean site at 625 m.a.s.l. and a subalpine site at 1335 m.a.s.l. Predicted mixedwood yield was higher than that for beech stands but lower than pine stands. When simulating climate change, mixedwood yield was reduced at the Mediterranean site (-33%) but increased at the subalpine site (+11%). Interaction effects were enhanced as stands developed. Complementarity dominated the Mediterranean stand but neutral or net competition dominated the subalpine stand, which had higher stand density and water availability. Reduced water demand and consumption, increased canopy interception, and improved water-use efficiency in mixtures compared to beech stands suggest a release of beech intra-specific competition. Beech also facilitated pine growth through better litter quality, non-symbiotic nitrogen fixation and above- and belowground stratification, leading to higher foliar nitrogen content and deeper canopies in pines. In conclusion, mixtures may improve water availability and use efficiency for beech and light interception for pine, the main limiting factors for each species, respectively. Encouraging pine-beech mixtures could be an effective adaptation to climate change in drought-prone sites in the Mediterranean region.
Open Access
Los precedentes y las respuestas de los árboles a sequías extremas revelan los procesos involucrados en el decaimiento de bosques mediterráneos de coníferas
(Asociación Española de Ecología Terrestre, 2012) Camarero, Jesús Julio; Sangüesa Barreda, G.; Alla, Arben Q.; González de Andrés, Ester; Maestro Martínez, M.; Vicente Serrano, S.M.; Ciencias del Medio Natural; Natura Ingurunearen Zientziak
¿Qué determina la vulnerabilidad de las especies de árboles frente a eventos climáticos extremos en el contexto actual de calentamiento global? Sabemos que las sequías severas contribuyen al decaimiento del bosque pero desconocemos los procesos involucrados en el declive de crecimiento y la pérdida de vigor del árbol que pueden conducir a su muerte. Para profundizar en estos procesos combinamos datos retrospectivos de crecimiento secundario, obtenidos mediante dendrocronología, y variables funcionales potencialmente indicadoras de cambios de vigor (frondosidad, producción de albura, cambios en las concentraciones de carbohidratos no estructurales en acículas y albura del tronco) en bosques mediterráneos afectados por decaimiento inducido por sequía. Evaluamos las respuestas de tres especies de árboles (Abies alba, Pinus sylvestris y Pinus halepensis) a la sequía extrema del año 2012 caracterizada mediante un índice de sequía que considera la temperatura. En primavera y verano del año 2012 la sequía fue muy intensa debido a que las elevadas temperaturas máximas condujeron a condiciones de aridez que carecen de precedentes similares desde 1950. En las tres especies los árboles menos frondosos (más defoliados) formaban menos albura y ésta contenía una menor concentración de azúcares solubles que en los árboles más frondosos. Los árboles más defoliados presentaron también mayor similitud de crecimiento secundario entre años consecutivos que los poco defoliados. Los patrones de crecimiento difirieron entre especies y sugieren que en P. sylvestris el decaimiento responde a sequías extremas recurrentes durante el siglo pasado, mientras que A. alba y P. halepensis muestran patrones correspondientes a un declive durante la pasada década relacionado con el aumento de temperaturas que ha producido el decaimiento de individuos que durante episodios climáticamente benignos pasados mostraron altas tasas de crecimiento. Los diversos patrones descritos indican la necesidad de definir conceptos como la muerte del árbol o su recuperación post-decaimiento dentro de un marco de investigación objetivo, cuantitativo y verificable.
Open Access
Monitoring rainfed alfalfa growth in semiarid agrosystems using Sentinel-2 imagery
(MDPI, 2021) Echeverría Obanos, Andrés; Urmeneta, Alejandro; González de Audícana Amenábar, María; González de Andrés, Ester; Zientziak; Ingeniaritza; Institute for Multidisciplinary Research in Applied Biology - IMAB; Institute on Innovation and Sustainable Development in Food Chain - ISFOOD; Ciencias; Ingeniería; Gobierno de Navarra / Nafarroako Gobernua
The aim of this study was to assess the utility of Sentinel-2 images in the monitoring of the fractional vegetation cover (FVC) of rainfed alfalfa in semiarid areas such as that of Bardenas Reales in Spain. FVC was sampled in situ using 1 m2 surfaces at 172 points inside 18 alfalfa fields from late spring to early summer in 2017 and 2018. Different vegetation indices derived from a series of Sentinel-2 images were calculated and were then correlated with the FVC measurements at the pixel and parcel levels using different types of equations. The results indicate that the normalized difference vegetation index (NDVI) and FVC were highly correlated at the parcel level (R 2 = 0.712), where as the correlation at the pixel level remained moderate across each of the years studied. Based on the findings, another 29 alfalfa plots (28 rainfed; 1 irrigated) were remotely monitored operationally for 3 years (2017–2019), revealing that location and weather conditions were strong determinants of alfalfa growth in Bardenas Reales. The results of this study indicate that Sentinel-2 imagery is a suitable tool for monitoring rainfed alfalfa pastures in semiarid areas, thus increasing the potential success of pasture management.