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As with many other species of pine, the only possibility of regeneration for Pinus pinaster is from seed. The advantage of germination is that it increases the genetic variability and stability of populations. This species has an important natural recovery rate in the field after a wildfire. However, no germination or survival of seedlings under adult trees with a capacity for producing viable seeds has been seen to exist in the field. The hypothesis of this paper is that germination under the cover of the adult trees could be controlled by the following ecological factors: availability of light, inhibition by substances produced by the aboveground mass of the tree or inhibition produced by substances emitted by decomposing fallen litter. The effect of nutrient availability on germination was analysed. The germination percentages were high in all of the treatments carried out, with a positive effect appearing in the germination of leaf exudates and a lower percentage in the seeds left in total darkness. However, no significant differences are observed among the total germination percentages. The main difference between the effects of the different treatments appears in the mean time required to germinate: it was shortest in the seeds treated with leaf exudates and longest in the seeds left in total darkness

Conferencia impartida en el Seminario “Avances en el estudio de la severidad de los incendios forestales utilizando técnicas de teledetección”

This study examines the effect of fire on the germination of Pinus pinaster seeds from a population with special adaptations to this type of disturbance, due to the high frequency of fires to which they have been subjected. The action of fire was simulated in the laboratory using thermal shocks. Temperatures of 60, 90, 120, 200 and 300 °C were used for exposure times of 1 and 5 min. The viability of seeds of this species from the soil seed bank of a population subjected to a wildfire was also evaluated. The results show that germination is not increase by a thermal treatment. The effect is negative at temperatures above 90 °C and exposure times of 1 min with a significant decrease in, or even no, germination. Therefore the seeds cannot stand high temperatures and reproductive effort is placed on serotinous cone development and the production of high numbers of seeds. The high viability of the seeds from the area burned by wildfire, immediately after the fire and 1 year later, show the high potential of the soil seed bank to contribute to the regeneration of the burned area.

In the Cantabrian area (northwest Spain) Pinus stands occupy many of the original shrub communities that have been considered unproductive. These Pinus stands represent the vegetation which is most affected by fire. Regeneration after fire may occur in different ways: by resprouting or by germination or both. Germination was the only regeneration mechanism in Pinus species that appeared in these areas. The aim of this study is to determine the role of the soil seed bank in regeneration in this type of ecosystem. In order to carry out the study, three communities dominated by Pinus sylvestris which had suffered wildfires were chosen. In each of the three experimental sites of Pinus sylvestris stands the seed bank composition and above-ground vegetation were studied. The results allowed three species groups in the seed bank to be differentiated: those favoured by fire, amongst which some hardseeds, mainly belonging to Cistaceae and Leguminosae, were found; another group formed by outsider or opportunist species from outside the community and which used anemochory as their main dispersion mechanism; and the third group formed by those negatively affected, amongst which were species using vegetative resprout as the main regeneration mechanism. The species of greatest quantitative importance in the seed bank was Erica australis. In general, anemochorous species were predominant in the soil seed bank. During the first stages of succession chamaephytes were dominant and in the two years after fire therophytes were. No great similarity was observed between the bank composition and field vegetation from a qualitative viewpoint, due to differences in the presence of seeds of outsider plants in the bank and to the significance of the resprouting species in the field.

In the Mediterranean Basin, wildland fires are major drivers of forest ecosystem dynamics. In the current context of global change, these fires are becoming more severe and recurrent because of climatic conditions, land use changes and invasive species. In areas affected by mega-fires (burned area > 10,000 ha), the patterns of regeneration may be heterogeneous due to local variations in fire regime, community composition and environmental features. The goal of this study was to analyze the post-fire structure of both Pinus pinaster Aiton. seedlings population and understory community in a Mediterranean fire-prone ecosystem at short-term by means of high spatial resolution satellite imagery within the perimeter of a full stand replacing mega-fire that burned around 12,000 ha of a Pinus pinaster forest in NW Spain. We established 234 field plots of 2 × 2 meters to cover four recurrence-severity scenarios. In each plot, we sampled 15 vegetation structural variables at both pine seedlings population and understory community levels. From the WorldView-2 satellite imagery, we obtained three sets of spectral variables (reflectance, spectral indices and image textures) that were used as predictors of vegetation recovery in generalized linear models. At population level, the number and cover of pine seedlings were successfully modeled with spectral indices and textural information (normalized root mean square error of 16% and 17%, respectively). At understory community level, woody species cover was correlated with first order textures (normalized root mean square error of 9%). Other understory structure variables (height and richness of woody species, percentage of bare soil, necromass and leaves) were predicted with an error lower than 20%. The predictive capacity of the models was similar for all recurrence-severity scenarios. Our results highlight the usefulness of spectral indices and textural data at high spatial resolution in the analysis of post-fire recovery in larg

Fire frequency and burn severity may increase in pine forests in the Mediterranean Basin under the warmer and drier climate projected for this region. Our study aims to evaluate the role of fire frequency and burn severity in the post-fire recruitment and development of Mediterranean serotinous pines under different environmental conditions. Two pine forests representing contrasting climatic conditions and soil types that support serotinous pines in the Iberian Peninsula and affected by large wildfires in summer 2012, were selected. In these two study sites, we determined the number of wildfires between 1978 and 2012 and the burn severity of the last fire (2012 at both sites) through the dNBR spectral index. Three and four years after the wildfires, we sampled the density, cover and height of pine seedlings and the cover of woody understory species in 1296 1 m2 plots. The results indicated that the density and cover of pine seedlings was low after two fires combined with high severities, as well as after three fires, regardless of burn severity. Seedling recruitment after three fires was particularly threatened in the most arid study site (0.01 seedlings m−2), resulting in low seedling cover (0.01%). Seedling height decreased with fire frequency in both study sites, and with burn severity owing to fire-induced shifts in soil fertility and microclimatic conditions. There was a significant negative effect of the cover of woody understory species on the recruitment and cover of pine seedlings. Our results suggest that the effects of increasing fire frequency and severity on pine regeneration may be aggravated under arid conditions. Additionally, this study encourages forest managers to avoid the occurrence of frequent crown fires in order to prevent the loss of serotinous pine forest, and provides useful information to predict the scenarios in which post-fire restoration actions would be helpful.

Due to the high sensitivity of mountain landscapes to environmental changes, the study of land cover dynamics has become an essential tool for guiding management policies. Since the second half of the twentieth century, the Cantabrian Mountains (NW Spain) have been substantially altered by the loss of traditional management practices and, more recently, by the new environmental schemes developed by the Regional Government. This area is a biodiversity hotspot, representing the south-western-most distribution limit for a large number of species in Europe. Therefore, small changes in landscape patterns can result in biodiversity losses. In this study, we analyzed land cover changes in the Cantabrian Mountains from 1991 to 2004 by means of remote sensing techniques, identifying the main driving forces and classifying the territory according to its risk of land cover change. Forest expansion and loss of shrublands were the two major trajectories of change apparent during this period. When modeling the occurrence of these land cover changes, we found that performance of models was related to the nature of the change. The most accurate models were associated with processes of secondary succession, i.e. forest expansion (78.6%), while the least accurate models related to changes linked with management decisions, i.e. loss of shrubs (61.8%). The main drivers of change were variations in the number of goats (for the forest expansion model) and changes in the number of head of sheep and cattle (for the loss of shrubs model). Topographic conditions (altitude and slope) were relevant in both models. Our approach proposes an explicit decision support tool for landscape managers, allowing better identification of the areas where they should focus their attention.

Forest managers require reliable tools to evaluate post-fire recovery across different geographic/climatic contexts and define management actions at the landscape scale, which might be highly resource-consuming in terms of data collection. In this sense, remote sensing techniques allow for gathering environmental data over large areas with low collection effort. We aim to assess the applicability of remote sensing tools in post-fire management within and across three mega-fires that occurred in pine fire-prone ecosystems located along an Atlantic-Transition-Mediterranean climatic gradient. Four years after the wildfires, we established 120 2x2m plots in each mega-fire site, where we evaluated: (1) density of pine seedlings, (2) percentage of woody species cover and (3) percentage of dead plant material cover. These variables were modeled following a Bayesian Model Averaging approach on the basis of spectral indices and texture features derived from WorldView-2 satellite imagery at 2 m spatial resolution. We assessed model interpolation and transferability within each mega-fire, as well as model extrapolation between mega-fires along the climatic gradient. Texture features were the predictors that contributed most in all cases. The woody species cover model had the best performance regarding spatial interpolation and transferability within the three study sites, with predictive errors lower than 25% for the two approaches. Model extrapolation between the Transition and Mediterranean sites had low levels of error (from 6% to 19%) for the three field variables, because the landscape in these areas is similar in structure and function and, therefore, in spectral characteristics. However, model extrapolation from the Atlantic site achieved the weakest results (error higher than 30%), due to the large ecological differences between this particular site and the others. This study demonstrates the potential of fine-grained satellite imagery for land managers to conduct post

Land abandonment and the loss of traditional farming practices are thought to control land cover dynamics, and hence the ecosystem service supply in traditionally managed mountain landscapes. We evaluate the impact of land cover changes in Cantabrian Mountains (NW Spain), over 1990–2012, on the potential supply capacity of ecosystem services (regulating, provisioning, and cultural) at both regional and local scales. We also analyze trends in the use of ecosystem services at the local scale. Land cover changes were estimated from CORINE Land Cover database. Patterns of potential ecosystem service supply were assessed by applying an ecosystem service supply capacity matrix and trends in their actual use by using field data. Main trajectories of land cover change encompassed woody vegetation spread in semi-natural open systems and agricultural expansion in the most suitable areas. The capacity of landscape to provide ecosystem services improved during 1990–2012 at both scales. We detected trade-offs between the potential supply of ecosystem services associated to natural systems and those linked to traditional land uses, at both regional and local scales. Changes in the potential supply of ecosystem services matched trends in ecosystem service use. This study could help develop future scenarios to address upcoming challenges in ecosystem service supply

In the Cantabrian area (northwest Spain) Pinus stands occupy many of the original shrub communities that have been considered unproductive. These Pinus stands represent the vegetation which is most affected by fire. Regeneration after fire may occur in different ways: by resprouting or by germination or both. Germination was the only regeneration mechanism in Pinus species that appeared in these areas. The aim of this study is to determine the role of the soil seed bank in regeneration in this type of ecosystem. In order to carry out the study, three communities dominated by Pinus sylvestris which had suffered wildfires were chosen. In each of the three experimental sites of Pinus sylvestris stands the seed bank composition and above-ground vegetation were studied. The results allowed three species groups in the seed bank to be differentiated: those favoured by fire, amongst which some hardseeds, mainly belonging to Cistaceae and Leguminosae, were found; another group formed by outsider or opportunist species from outside the community and which used anemochory as their main dispersion mechanism; and the third group formed by those negatively affected, amongst which were species using vegetative resprout as the main regeneration mechanism. The species of greatest quantitative importance in the seed bank was Erica australis. In general, anemochorous species were predominant in the soil seed bank. During the first stages of succession chamaephytes were dominant and in the two years after fire therophytes were. No great similarity was observed between the bank composition and field vegetation from a qualitative viewpoint, due to differences in the presence of seeds of outsider plants in the bank and to the significance of the resprouting species in the field.

As with many other species of pine, the only possibility of regeneration for Pinus pinaster is from seed. The advantage of germination is that it increases the genetic variability and stability of populations. This species has an important natural recovery rate in the field after a wildfire. However, no germination or survival of seedlings under adult trees with a capacity for producing viable seeds has been seen to exist in the field. The hypothesis of this paper is that germination under the cover of the adult trees could be controlled by the following ecological factors: availability of light, inhibition by substances produced by the aboveground mass of the tree or inhibition produced by substances emitted by decomposing fallen litter. The effect of nutrient availability on germination was analysed. The germination percentages were high in all of the treatments carried out, with a positive effect appearing in the germination of leaf exudates and a lower percentage in the seeds left in total darkness. However, no significant differences are observed among the total germination percentages. The main difference between the effects of the different treatments appears in the mean time required to germinate: it was shortest in the seeds treated with leaf exudates and longest in the seeds left in total darkness

In the Mediterranean Basin, wildland fires are major drivers of forest ecosystem dynamics. In the current context of global change, these fires are becoming more severe and recurrent because of climatic conditions, land use changes and invasive species. In areas affected by mega-fires (burned area > 10,000 ha), the patterns of regeneration may be heterogeneous due to local variations in fire regime, community composition and environmental features. The goal of this study was to analyze the post-fire structure of both Pinus pinaster Aiton. seedlings population and understory community in a Mediterranean fire-prone ecosystem at short-term by means of high spatial resolution satellite imagery within the perimeter of a full stand replacing mega-fire that burned around 12,000 ha of a Pinus pinaster forest in NW Spain. We established 234 field plots of 2 × 2 meters to cover four recurrence-severity scenarios. In each plot, we sampled 15 vegetation structural variables at both pine seedlings population and understory community levels. From the WorldView-2 satellite imagery, we obtained three sets of spectral variables (reflectance, spectral indices and image textures) that were used as predictors of vegetation recovery in generalized linear models. At population level, the number and cover of pine seedlings were successfully modeled with spectral indices and textural information (normalized root mean square error of 16% and 17%, respectively). At understory community level, woody species cover was correlated with first order textures (normalized root mean square error of 9%). Other understory structure variables (height and richness of woody species, percentage of bare soil, necromass and leaves) were predicted with an error lower than 20%. The predictive capacity of the models was similar for all recurrence-severity scenarios. Our results highlight the usefulness of spectral indices and textural data at high spatial resolution in the analysis of post-fire recovery in larg

Due to the high sensitivity of mountain landscapes to environmental changes, the study of land cover dynamics has become an essential tool for guiding management policies. Since the second half of the twentieth century, the Cantabrian Mountains (NW Spain) have been substantially altered by the loss of traditional management practices and, more recently, by the new environmental schemes developed by the Regional Government. This area is a biodiversity hotspot, representing the south-western-most distribution limit for a large number of species in Europe. Therefore, small changes in landscape patterns can result in biodiversity losses. In this study, we analyzed land cover changes in the Cantabrian Mountains from 1991 to 2004 by means of remote sensing techniques, identifying the main driving forces and classifying the territory according to its risk of land cover change. Forest expansion and loss of shrublands were the two major trajectories of change apparent during this period. When modeling the occurrence of these land cover changes, we found that performance of models was related to the nature of the change. The most accurate models were associated with processes of secondary succession, i.e. forest expansion (78.6%), while the least accurate models related to changes linked with management decisions, i.e. loss of shrubs (61.8%). The main drivers of change were variations in the number of goats (for the forest expansion model) and changes in the number of head of sheep and cattle (for the loss of shrubs model). Topographic conditions (altitude and slope) were relevant in both models. Our approach proposes an explicit decision support tool for landscape managers, allowing better identification of the areas where they should focus their attention.

Fire frequency and burn severity may increase in pine forests in the Mediterranean Basin under the warmer and drier climate projected for this region. Our study aims to evaluate the role of fire frequency and burn severity in the post-fire recruitment and development of Mediterranean serotinous pines under different environmental conditions. Two pine forests representing contrasting climatic conditions and soil types that support serotinous pines in the Iberian Peninsula and affected by large wildfires in summer 2012, were selected. In these two study sites, we determined the number of wildfires between 1978 and 2012 and the burn severity of the last fire (2012 at both sites) through the dNBR spectral index. Three and four years after the wildfires, we sampled the density, cover and height of pine seedlings and the cover of woody understory species in 1296 1 m2 plots. The results indicated that the density and cover of pine seedlings was low after two fires combined with high severities, as well as after three fires, regardless of burn severity. Seedling recruitment after three fires was particularly threatened in the most arid study site (0.01 seedlings m−2), resulting in low seedling cover (0.01%). Seedling height decreased with fire frequency in both study sites, and with burn severity owing to fire-induced shifts in soil fertility and microclimatic conditions. There was a significant negative effect of the cover of woody understory species on the recruitment and cover of pine seedlings. Our results suggest that the effects of increasing fire frequency and severity on pine regeneration may be aggravated under arid conditions. Additionally, this study encourages forest managers to avoid the occurrence of frequent crown fires in order to prevent the loss of serotinous pine forest, and provides useful information to predict the scenarios in which post-fire restoration actions would be helpful.

This study examines the effect of fire on the germination of Pinus pinaster seeds from a population with special adaptations to this type of disturbance, due to the high frequency of fires to which they have been subjected. The action of fire was simulated in the laboratory using thermal shocks. Temperatures of 60, 90, 120, 200 and 300 °C were used for exposure times of 1 and 5 min. The viability of seeds of this species from the soil seed bank of a population subjected to a wildfire was also evaluated. The results show that germination is not increase by a thermal treatment. The effect is negative at temperatures above 90 °C and exposure times of 1 min with a significant decrease in, or even no, germination. Therefore the seeds cannot stand high temperatures and reproductive effort is placed on serotinous cone development and the production of high numbers of seeds. The high viability of the seeds from the area burned by wildfire, immediately after the fire and 1 year later, show the high potential of the soil seed bank to contribute to the regeneration of the burned area.

Forest managers require reliable tools to evaluate post-fire recovery across different geographic/climatic contexts and define management actions at the landscape scale, which might be highly resource-consuming in terms of data collection. In this sense, remote sensing techniques allow for gathering environmental data over large areas with low collection effort. We aim to assess the applicability of remote sensing tools in post-fire management within and across three mega-fires that occurred in pine fire-prone ecosystems located along an Atlantic-Transition-Mediterranean climatic gradient. Four years after the wildfires, we established 120 2x2m plots in each mega-fire site, where we evaluated: (1) density of pine seedlings, (2) percentage of woody species cover and (3) percentage of dead plant material cover. These variables were modeled following a Bayesian Model Averaging approach on the basis of spectral indices and texture features derived from WorldView-2 satellite imagery at 2 m spatial resolution. We assessed model interpolation and transferability within each mega-fire, as well as model extrapolation between mega-fires along the climatic gradient. Texture features were the predictors that contributed most in all cases. The woody species cover model had the best performance regarding spatial interpolation and transferability within the three study sites, with predictive errors lower than 25% for the two approaches. Model extrapolation between the Transition and Mediterranean sites had low levels of error (from 6% to 19%) for the three field variables, because the landscape in these areas is similar in structure and function and, therefore, in spectral characteristics. However, model extrapolation from the Atlantic site achieved the weakest results (error higher than 30%), due to the large ecological differences between this particular site and the others. This study demonstrates the potential of fine-grained satellite imagery for land managers to conduct post

Land abandonment and the loss of traditional farming practices are thought to control land cover dynamics, and hence the ecosystem service supply in traditionally managed mountain landscapes. We evaluate the impact of land cover changes in Cantabrian Mountains (NW Spain), over 1990–2012, on the potential supply capacity of ecosystem services (regulating, provisioning, and cultural) at both regional and local scales. We also analyze trends in the use of ecosystem services at the local scale. Land cover changes were estimated from CORINE Land Cover database. Patterns of potential ecosystem service supply were assessed by applying an ecosystem service supply capacity matrix and trends in their actual use by using field data. Main trajectories of land cover change encompassed woody vegetation spread in semi-natural open systems and agricultural expansion in the most suitable areas. The capacity of landscape to provide ecosystem services improved during 1990–2012 at both scales. We detected trade-offs between the potential supply of ecosystem services associated to natural systems and those linked to traditional land uses, at both regional and local scales. Changes in the potential supply of ecosystem services matched trends in ecosystem service use. This study could help develop future scenarios to address upcoming challenges in ecosystem service supply

Se pretende determinar los cambios que se producen en ecosistemas de Pinus pinaster como consecuencia de incendios forestales. E I estudio se llevó a cabo en los pinares de la Sierra del Teleno (provincia de León), debido a que esta Sierra presenta áreas que han sido alteradas por el fuego en diferentes años. Se seleccionó una zona quemada en 1991 y otra que sufrió un gran incendio en Septiembre de 1998. En esta última y dada la magnitud del incendio el muestreo se realizó en tres áreas con diferente orientación y topografía (N umbría, S solana y NW con fuerte pendiente). Por último, se seleccionó otra área no quemada, considerada como control, con el objeto de conocer las características en una zona no perturbada. Se dispone de datos de cobertura vegetal a los 7, 9, 12, 20, 23 meses en la zona recientemente quemada, así como 8 y 9 años y en la zona control. Los muestreos de vegetación se realizaron estimando la cobertura de todas las especies presentes en 60 unidades de muestreo de 1 m 2 en cada área. Los valores de cobertura global se incrementan a lo largo del periodo de estudio, fundamentalmente debido a las especies leñosas que son el biotipo dominante desde los primeros meses de regeneración post-fuego. Para determinar los cambios en el suelo se tomaron cinco muestras en cada una de las zonas, analizando materia orgánica, pH, nitrógeno total, fósforo asimilable y cationes asimilables. Seis meses después del incendio se observa una alta va riabilidad entre las parcelas recientemente quemadas debido a los efectos puntuales del fuego sobre el suelo. Se pone de manifiesto como a largo plazo y a medida que se establece la vegetación se tienden a alcanzar características similares al control, aunque ocho años después del fuego aún se mantienen algunas diferencias.

El fuego es un factor ambiental que ha modelado el paisaje de la cordillera Cantá-brica, recorriendo de forma frecuente los ecosistemas que en ella se asientan. Entre ellos se encuentran los rebollares de Quercus pyrenaica, que históricamente han sufrido una im-portante pérdida de madurez, en parte debida a los incendios. Otros ecosistemas que ocu-pan amplias extensiones son los brezales, que en muchos casos representan etapas secundarias tras abandono de pastos o cultivos y persisten principalmente debido a la repe-tición de incendios, que frenan el avance sucesional hacia etapas forestales. La vegetación presente en el momento actual en ambos tipos de ecosistemas se caracteriza por una gran tolerancia al fuego, ya que las especies menos resistentes han sido desplazadas por aquellas capaces de sobrevivir o regenerarse rápidamente tras esta perturbación. En este contexto, en el presente trabajo se pretende realizar una revisión de los principales procesos y meca-nismos de recuperación post-fuego de los rebollares y brezales situados en la cordillera Can-tábrica. La diversidad florística del sotobosque de Quercus pyrenaica o de los brezales aumenta en los primeros años tras el fuego, debido principalmente a los efectos favorece-dores del mismo en la germinación y rebrote de las especies herbáceas y a la menor com-petencia con las especies arbustivas. El rebrote es frecuente en herbáceas que presentan bulbos, rizomas o tallos subterráneos capaces de sobrevivir debido a la mala difusión del calor en el suelo, como muchas gramínea, liliáceas, etc. En el curso del tiempo, estas es-pecies tienden a disminuir, al recuperarse las leñosas propias de la comunidad. En estos ecosistemas, las especies leñosas se recuperan de forma mayoritaria por rebrote vegetativo a partir de órganos subterráneos que sobreviven al fuego. Este tipo de recuperación aparece tanto en especies arbóreas, como Quercus pyrenaica, como en especies arbustivas, como Erica australis, Chamaespartium trident

Knowledge on the relationships between species functional traits and environmental filters is key to understanding the mechanisms underlying the current patterns of biodiversity loss from a multi-taxa perspective. The aim of this study was to identify the main environmental factors driving the functional structure of a terrestrial vertebrate community (mammals, breeding birds, reptiles and amphibians) in a temperate mountain system (the Cantabrian Mountains; NW Spain). Based on the Spanish Inventory of Terrestrial Vertebrate Species, we selected three functional traits (feeding guild, habitat use type and daily activity) and defined, for each trait, a set of functional groups considering vertebrate species with common functional characteristics. The community functional structure was evaluated by means of two functional indexes indicative of functional redundancy (species richness within each functional group) and functional diversity. Ordinary least squares regression and conditional autoregressive models were applied to determine the response of community functional structure to environmental filters (climate, topography, land cover, physiological state of vegetation, landscape heterogeneity and human influence). The results revealed that both functional redundancy and diversity of terrestrial vertebrates were non-randomly distributed across space; rather, they were driven by environmental filters. Climate, topography and human influence were the best predictors of community functional structure. The influence of land cover, physiological state of vegetation and landscape heterogeneity varied among functional groups. The results of this study are useful to identify the general assembly rules of species functional traits and to illustrate the importance of environmental filters in determining functional structure of terrestrial vertebrate communities in mountain systems.

We analyse the effects of burn severity on individual soil properties and soil quotients in Mediterranean fire-prone pine forests immediately after a wildfire. Burn severity was measured in the field through the substrate stratum of the Composite Burn Index and soil samples were taken 7–9 days after a wildfire occurred in a Pinus pinaster Ait. ecosystem. In each soil sample, we analysed physical (size of soil aggregates), chemical (pH, organic C, total N and available P) and biological (microbial biomass C, b-glucosidase, urease and acid phosphatase activities) properties. Size of aggregates decreased in the areas affected by high burn severity. Additionally, moderate and high severities were associated with increases in pH and available P concentration and with decreases in organic C concentration. Microbial biomass C showed similar patterns to organic C along the burn severity gradient. The enzymatic activities of phosphatase and b-glucosidase showed the highest sensitivity to burn severity, as they strongly decreased from the low-severity scenarios. Among the studied soil quotients, the C : N ratio, microbial quotient and b-glucosidase : microbial biomass C quotient decreased with burn severity. This work provides valuable information on the impact of burn severity on the functioning of sandy siliceous soils in fire-prone pine ecosystems.

Fires are a problematic and recurrent issue in Mediterranean forest ecosystems. Accurate discrimination of burn severity level is fundamental for the rehabilitation planning of affected areas. Though fieldwork is still necessary for measuring post-fire burn severity, remote sensing based techniques are being widely used to predict it because of their computational simplicity and straightforward application. Among them, spectral indices classification (especially difference Normalized Burn Ratio–dNBR- based ones) may be considered the standard remote sensing based method to distinguish burn severity level. In this work we show how this methodology may be improved by using land surface temperature (LST) to enhance the standard spectral indices. We considered a large wildfire in August 2012 in North Western Spain. The Composite Burn Index (CBI) was measured in 111 field plots and grouped into three burn severity levels. Relationship between Landsat 7 Enhanced Thematic Mapper (ETM+) LST-enhanced spectral indices and CBI was evaluated by using the normalized distance between two burn severity levels and spectral dispersion graphs. Inclusion of LST in the spectral index equation resulted in higher discrimination between burn severity levels than standard spectral indices (0.90, 8.50, and 17.52 NIR-SWIR Temperature version 1 vs 0.60, 2.83, and 6.46 NBR). Our results demonstrate the potential of LST for improving burn severity discrimination and mapping. Future research, however, is needed to evaluate the performance of the proposed LST-enhanced spectral indices in other fire regimes, and forest ecosystems.

We aimed to analyze the relationship between fire regime attributes and the post-fire greenness recovery of fire-prone pine ecosystems over the short (2-year) and medium (5-year) term after a large wildfire, using both a single and a combined fire regime attribute approach. We characterized the spatial (fire size), temporal (number of fires, fire recurrence, and return interval), and magnitude (burn severity of the last fire) fire regime attributes throughout a 40-year period with a long-time series of Landsat imagery and ancillary data. The burn severity of the last fire was measured by the dNBR (difference of the Normalized Burn Ratio) spectral index, and classified according to the ground reference values of the CBI (Composite Burn Index). Post-fire greenness recovery was obtained through the difference of the NDVI (Normalized Difference Vegetation Index) between pre- and post-fire Landsat scenes. The relationship between fire regime attributes (single attributes: fire recurrence, fire return interval, and burn severity; combined attributes: fire recurrence-burn severity and fire return interval-burn severity) and post-fire greenness recovery was evaluated using linear models. The results indicated that all the single and combined attributes significantly affected greenness recovery. The single attribute approach showed that high recurrence, short return interval and low severity situations had the highest vegetation greenness recovery. The combined attribute approach allowed us to identify a wider variety of post-fire greenness recovery situations than the single attribute one. Over the short term, high recurrence as well as short return interval scenarios showed the best post-fire greenness recovery independently of burn severity, while over the medium term, high recurrence combined with low severity was the most recovered scenario. This novel combined attribute approach (temporal plus magnitude) could be of great value to forest managers in the development of

Nitrogen (N) deposition has been identified as one of the main traits of terrestrial ecosystems, affecting their structure and functioning. Among terrestrial ecosystems, heathlands are characterised by low nutrient status and highly affected by N deposition. However, few studies have been developed under natural field conditions to evaluate the amount of N deposition in these ecosystems. Therefore, a field experiment was carried out to investigate the inorganic N inputs in mountainous heathlands of North-Western Spain. Two study sites (La Majúa and San Isidro) were selected on the south side of the Cantabrian Mountains, as a representative monitoring N-sensitive ecosystems. Three replicated bulk collectors and one rain gauge were installed at each study site to collect monthly rainwater samples over three years. Wet bulk N deposition was different between the study areas (2.81 kg N ha-1 year-1 in La Majúa and 4.56 kg N ha-1 year-1 in San Isidro), but showed the same temporal pattern, with the highest N deposition rate observed in April and the lowest in August-September. In general, annual wet bulk NO3--N deposition was higher than NH4+-N, reflecting the prevailing NOX emission sources. Depositions of NO3--N were mostly originated by NOX emissions from northern highly industrialized and populated areas of North Spain. The lower NH4+/NO3- ratio in rainwater observed in the study area could be due to the decline of traditional land uses (livestock grazing, heath burning and croplands) associated with NHY emissions. Despite the low rates of N deposition observed in this study, N-sensitive heathlands in North-Western Spain could be threatened by N depositions.

We analysed the relationship between burn severity indicators, from remote sensing and field observations, and soil properties after a wildfire in a fire-prone Mediterranean ecosystem. Our study area was a large wildfire in a Pinus pinaster forest. Burn severity from remote sensing was identified by studying immediate post-fire Land Surface Temperature (LST). We also evaluated burn severity in the field applying the Composite Burn Index (CBI) in a total of 84 plots (30 m diameter). In each plot we evaluated litter consumption, ash colour and char depth as visual indicators. We collected soil samples and pH, soil organic carbon, dry aggregate size distribution (MWD), aggregate stability and water repellency were analysed. A controlled heating of soil was also carried out in the laboratory, with soil from the control plots, to compare with the changes produced in soils affected by different severity levels in the field. Our results shown that changes in soil properties affected by wildfire were only observed in soil aggregation in the high severity situation. The laboratory-controlled heating showed that temperatures of about 300 °C result in a significant reduction in soil organic carbon and MWD. Furthermore, soil organic carbon showed a significant decrease when LST values increased. Char depth was the best visual indicator to show changes in soil properties (mainly physical properties) in large fires that occur in Mediterranean pine forests. We conclude that CBI and post-fire LST can be considered good indicators of soil burn severity since both indicate the impact of fire on soil properties.

Elevated nitrogen (N) deposition impacts the structure and functioning of heathland ecosystems across Europe. Calluna plants under high N-inputs are very sensitive to secondary stress factors, including defoliation attacks by the heather beetle. These attacks result in serious damage or death of Calluna, its rapid replacement by grasses, and the subsequent loss of heathland. We know very little about the mechanisms that control the populations and trigger outbreaks of the heather beetle, impeding proper management measures to mitigate the damage. We investigated the effects of N deposition on the relationships between the heather beetle, its host plant, and two arthropod predators at building (rejuvenated through fire) and mature heathlands. The study combines field manipulation experiments simulating a range of N deposition rates (0, 1, 2, 5 g N m−2 year−1 for 2 years, and 5.6 g N m−2 year−1 for 10 years), and food-choice laboratory experiments testing the preferences of adults and larvae of the heather beetle for N-treated Calluna plants, and the preferences of predators for larvae grown on plants with different N-content. The larvae of the heather beetle achieved the highest abundances after the long-term (10-year) addition of N at mature Calluna plots in the field. Contrary to the adults, the larvae foraged preferentially on the most N-rich Calluna shoots under laboratory conditions. Predators showed no aggregative numerical responses to the accumulation of heather beetle larvae at high N-input experimental plots. During the feeding trials, predators consumed a small number of larvae, both in total and per individual, and systematically avoided eating the larvae reared on high-N Calluna shoots. Our study showed that the most severe defoliation damage by the heather beetle is inflicted at the larval stage under prolonged availability of high-N inputs, and that arthropod predators might not act as effective regulators of the beetle's populations.

Projections of future wildfire regimes forecast an increased frequency of large high-severity fires that create very harsh environmental conditions and constitute a challenge to post-fire ecosystem regeneration. Under these new circumstances, better knowledge of the plant interaction mechanisms underlying post-fire seedling establishment success would aid restoration management to achieve the intended targets. We evaluated the combined effect of recurrent large stand-replacing fires and conventional post-fire restoration activities (salvage logging after a single large fire, and direct seeding and linear subsoiling plus seedling planting after two subsequent large fires) on tree seedling recruitment and performance (development, annual growth, and biomass) in the early stages of succession in fire-prone maritime pine (Pinus pinaster Ait.) ecosystems. We quantified plant facilitative/competitive interactions between naturally recruited pine seedlings, neighbouring seedlings and potential nurse shrubs with different post-fire regeneration strategies (obligate seeders vs resprouters), by computing the relative interaction index (RII). The results evidenced that fire recurrence altered plant species composition and conditioned initial pine seedling recruitment and establishment, prevailing over the expected negative impact of salvage logging and positive impact of seeding. Seedling recruitment was sufficient to ensure natural tree regeneration after a single fire event and undermined by repeated fires. Both delaying burned timber removal during salvage logging operations and retaining immature dead trees without commercial value onsite in subsoiled stands enhanced seedling recruitment via facilitative interactions. Higher seedling growth and height under shrubs than in open ground resulted in lower aerial and root biomass, indicating elongation in response to shade, and net competition for resources. Inter-specific competition between naturally regenerated seedlings and

This study evaluated the opportunities and challenges of using drones to obtain multispectral orthomosaics at ultra-high resolution that could be useful for monitoring large and heterogeneous burned areas. We conducted a survey using an octocopter equipped with a Parrot SEQUOIA multispectral camera in a 3000 ha framework located within the perimeter of a megafire in Spain. We assessed the quality of both the camera raw imagery and the multispectral orthomosaic obtained, as well as the required processing capability. Additionally, we compared the spatial information provided by the drone orthomosaic at ultra-high spatial resolution with another image provided by theWorldView-2 satellite at high spatial resolution. The drone raw imagery presented some anomalies, such as horizontal banding noise and non-homogeneous radiometry. Camera locations showed a lack of synchrony of the single frequency GPS receiver. The georeferencing process based on ground control points achieved an error lower than 30 cm in X-Y and lower than 55 cm in Z. The drone orthomosaic provided more information in terms of spatial variability in heterogeneous burned areas in comparison with theWorldView-2 satellite imagery. The drone orthomosaic could constitute a viable alternative for the evaluation of post-fire vegetation regeneration in large and heterogeneous burned areas

Fires are a problematic and recurrent issue in Mediterranean forest ecosystems. Accurate discrimination of burn severity level is fundamental for the rehabilitation planning of affected areas. Though fieldwork is still necessary for measuring post-fire burn severity, remote sensing based techniques are being widely used to predict it because of their computational simplicity and straightforward application. Among them, spectral indices classification (especially difference Normalized Burn Ratio–dNBR- based ones) may be considered the standard remote sensing based method to distinguish burn severity level. In this work we show how this methodology may be improved by using land surface temperature (LST) to enhance the standard spectral indices. We considered a large wildfire in August 2012 in North Western Spain. The Composite Burn Index (CBI) was measured in 111 field plots and grouped into three burn severity levels. Relationship between Landsat 7 Enhanced Thematic Mapper (ETM+) LST-enhanced spectral indices and CBI was evaluated by using the normalized distance between two burn severity levels and spectral dispersion graphs. Inclusion of LST in the spectral index equation resulted in higher discrimination between burn severity levels than standard spectral indices (0.90, 8.50, and 17.52 NIR-SWIR Temperature version 1 vs 0.60, 2.83, and 6.46 NBR). Our results demonstrate the potential of LST for improving burn severity discrimination and mapping. Future research, however, is needed to evaluate the performance of the proposed LST-enhanced spectral indices in other fire regimes, and forest ecosystems.

We analysed the relationship between burn severity indicators, from remote sensing and field observations, and soil properties after a wildfire in a fire-prone Mediterranean ecosystem. Our study area was a large wildfire in a Pinus pinaster forest. Burn severity from remote sensing was identified by studying immediate post-fire Land Surface Temperature (LST). We also evaluated burn severity in the field applying the Composite Burn Index (CBI) in a total of 84 plots (30 m diameter). In each plot we evaluated litter consumption, ash colour and char depth as visual indicators. We collected soil samples and pH, soil organic carbon, dry aggregate size distribution (MWD), aggregate stability and water repellency were analysed. A controlled heating of soil was also carried out in the laboratory, with soil from the control plots, to compare with the changes produced in soils affected by different severity levels in the field. Our results shown that changes in soil properties affected by wildfire were only observed in soil aggregation in the high severity situation. The laboratory-controlled heating showed that temperatures of about 300 °C result in a significant reduction in soil organic carbon and MWD. Furthermore, soil organic carbon showed a significant decrease when LST values increased. Char depth was the best visual indicator to show changes in soil properties (mainly physical properties) in large fires that occur in Mediterranean pine forests. We conclude that CBI and post-fire LST can be considered good indicators of soil burn severity since both indicate the impact of fire on soil properties.

We aimed to analyze the relationship between fire regime attributes and the post-fire greenness recovery of fire-prone pine ecosystems over the short (2-year) and medium (5-year) term after a large wildfire, using both a single and a combined fire regime attribute approach. We characterized the spatial (fire size), temporal (number of fires, fire recurrence, and return interval), and magnitude (burn severity of the last fire) fire regime attributes throughout a 40-year period with a long-time series of Landsat imagery and ancillary data. The burn severity of the last fire was measured by the dNBR (difference of the Normalized Burn Ratio) spectral index, and classified according to the ground reference values of the CBI (Composite Burn Index). Post-fire greenness recovery was obtained through the difference of the NDVI (Normalized Difference Vegetation Index) between pre- and post-fire Landsat scenes. The relationship between fire regime attributes (single attributes: fire recurrence, fire return interval, and burn severity; combined attributes: fire recurrence-burn severity and fire return interval-burn severity) and post-fire greenness recovery was evaluated using linear models. The results indicated that all the single and combined attributes significantly affected greenness recovery. The single attribute approach showed that high recurrence, short return interval and low severity situations had the highest vegetation greenness recovery. The combined attribute approach allowed us to identify a wider variety of post-fire greenness recovery situations than the single attribute one. Over the short term, high recurrence as well as short return interval scenarios showed the best post-fire greenness recovery independently of burn severity, while over the medium term, high recurrence combined with low severity was the most recovered scenario. This novel combined attribute approach (temporal plus magnitude) could be of great value to forest managers in the development of

Nitrogen (N) deposition has been identified as one of the main traits of terrestrial ecosystems, affecting their structure and functioning. Among terrestrial ecosystems, heathlands are characterised by low nutrient status and highly affected by N deposition. However, few studies have been developed under natural field conditions to evaluate the amount of N deposition in these ecosystems. Therefore, a field experiment was carried out to investigate the inorganic N inputs in mountainous heathlands of North-Western Spain. Two study sites (La Majúa and San Isidro) were selected on the south side of the Cantabrian Mountains, as a representative monitoring N-sensitive ecosystems. Three replicated bulk collectors and one rain gauge were installed at each study site to collect monthly rainwater samples over three years. Wet bulk N deposition was different between the study areas (2.81 kg N ha-1 year-1 in La Majúa and 4.56 kg N ha-1 year-1 in San Isidro), but showed the same temporal pattern, with the highest N deposition rate observed in April and the lowest in August-September. In general, annual wet bulk NO3--N deposition was higher than NH4+-N, reflecting the prevailing NOX emission sources. Depositions of NO3--N were mostly originated by NOX emissions from northern highly industrialized and populated areas of North Spain. The lower NH4+/NO3- ratio in rainwater observed in the study area could be due to the decline of traditional land uses (livestock grazing, heath burning and croplands) associated with NHY emissions. Despite the low rates of N deposition observed in this study, N-sensitive heathlands in North-Western Spain could be threatened by N depositions.

New fire disturbance regimes under accelerating global environmental change can have unprecedented consequences for ecosystem resilience, lessening ecosystem natural regeneration. In the Mediterranean Basin, fire-dependent obligate seeder forests that are prone to increasingly frequent stand-replacing fires and then salvaged logged repeatedly can be vulnerable to additional disturbances for decades. In this study, we investigated, for the first time, the cumulative and interactive effects of two large high-severity fires at a short (<15-year) return interval and the subsequent burned timber harvesting with biomass removal on the post-disturbance recovery of such forests. We further assessed the type and amount of the material legacies (deadwood) that persisted through the different post-disturbance successional trajectories, as well as the influence of these legacies on forest regeneration. The early recovery of the studied forests after two consecutive large fires and post-fire logging was, in the first place, driven by fire repetition, which led to reduced seedling recruitment and enhanced regrowth of resprouter shrubs. Despite no interactive effects between fire and logging were detected after a single large fire event, two repeated fires followed by salvage harvesting had a greater negative impact than two fires alone (synergistic effects) on seedling establishment; while a lower positive impact (subadditive effects) on the recovery of resprouter shrubs. There was also an interaction modification effect in which fire repetition worsened the per-unit impact of salvage logging on forest regeneration. Nonetheless, the residual legacies, i.e., fine and coarse woody debris (unburned needles, downed branches, pieces of deadwood, and burned pine cones) that remained after the manual harvesting of the burned trees, aided seedling re-establishment and hindered the regrowth of the shrubby understorey. These findings indicate that high-intensity salvage logging after two l

Multispectral imagery is a widely used source of information to address post-fire ecosystem management. The aim of this study is to evaluate the ability of remotely sensed indices derived from Landsat 8 OLI/TIRS to assess initial burn severity (overall, on vegetation and on soil) in fire-prone pine forests along the Mediterranean-Transition-Oceanic climatic gradient in the Mediterranean Basin. We selected four large wildfires which affected pine forests in a climatic gradient within the Iberian Peninsula. In each wildfire we established CBI plots to obtain field values of three burn severity metrics: site, vegetation and soil burn severity. The ability of 13 spectral indices to match these three field burn severity metrics was compared and their transferability along the climatic gradient assessed using linear regression models. Specifically, we analysed the performance of 12 indices previously used for burn severity assessments (8 reflective, 2 thermal, 2 mixed) and a new reflective index (dNBR-EVI). The results showed that Landsat spectral indices have a greater ability to determine site and vegetation burn severity than soil burn severity. We found large differences in indices performances among the three different climatic regions, since most indices performed better in the Mediterranean and Transition regions than in the Oceanic one. In general, the dNBR-EVI showed the best fit to site, vegetation and soil burn severity in the three regions, demonstrating broad transferability along the entire climatic gradient.

Elevated nitrogen (N) deposition impacts the structure and functioning of heathland ecosystems across Europe. Calluna plants under high N-inputs are very sensitive to secondary stress factors, including defoliation attacks by the heather beetle. These attacks result in serious damage or death of Calluna, its rapid replacement by grasses, and the subsequent loss of heathland. We know very little about the mechanisms that control the populations and trigger outbreaks of the heather beetle, impeding proper management measures to mitigate the damage. We investigated the effects of N deposition on the relationships between the heather beetle, its host plant, and two arthropod predators at building (rejuvenated through fire) and mature heathlands. The study combines field manipulation experiments simulating a range of N deposition rates (0, 1, 2, 5 g N m−2 year−1 for 2 years, and 5.6 g N m−2 year−1 for 10 years), and food-choice laboratory experiments testing the preferences of adults and larvae of the heather beetle for N-treated Calluna plants, and the preferences of predators for larvae grown on plants with different N-content. The larvae of the heather beetle achieved the highest abundances after the long-term (10-year) addition of N at mature Calluna plots in the field. Contrary to the adults, the larvae foraged preferentially on the most N-rich Calluna shoots under laboratory conditions. Predators showed no aggregative numerical responses to the accumulation of heather beetle larvae at high N-input experimental plots. During the feeding trials, predators consumed a small number of larvae, both in total and per individual, and systematically avoided eating the larvae reared on high-N Calluna shoots. Our study showed that the most severe defoliation damage by the heather beetle is inflicted at the larval stage under prolonged availability of high-N inputs, and that arthropod predators might not act as effective regulators of the beetle's populations.

Projections of future wildfire regimes forecast an increased frequency of large high-severity fires that create very harsh environmental conditions and constitute a challenge to post-fire ecosystem regeneration. Under these new circumstances, better knowledge of the plant interaction mechanisms underlying post-fire seedling establishment success would aid restoration management to achieve the intended targets. We evaluated the combined effect of recurrent large stand-replacing fires and conventional post-fire restoration activities (salvage logging after a single large fire, and direct seeding and linear subsoiling plus seedling planting after two subsequent large fires) on tree seedling recruitment and performance (development, annual growth, and biomass) in the early stages of succession in fire-prone maritime pine (Pinus pinaster Ait.) ecosystems. We quantified plant facilitative/competitive interactions between naturally recruited pine seedlings, neighbouring seedlings and potential nurse shrubs with different post-fire regeneration strategies (obligate seeders vs resprouters), by computing the relative interaction index (RII). The results evidenced that fire recurrence altered plant species composition and conditioned initial pine seedling recruitment and establishment, prevailing over the expected negative impact of salvage logging and positive impact of seeding. Seedling recruitment was sufficient to ensure natural tree regeneration after a single fire event and undermined by repeated fires. Both delaying burned timber removal during salvage logging operations and retaining immature dead trees without commercial value onsite in subsoiled stands enhanced seedling recruitment via facilitative interactions. Higher seedling growth and height under shrubs than in open ground resulted in lower aerial and root biomass, indicating elongation in response to shade, and net competition for resources. Inter-specific competition between naturally regenerated seedlings and

This study evaluated the opportunities and challenges of using drones to obtain multispectral orthomosaics at ultra-high resolution that could be useful for monitoring large and heterogeneous burned areas. We conducted a survey using an octocopter equipped with a Parrot SEQUOIA multispectral camera in a 3000 ha framework located within the perimeter of a megafire in Spain. We assessed the quality of both the camera raw imagery and the multispectral orthomosaic obtained, as well as the required processing capability. Additionally, we compared the spatial information provided by the drone orthomosaic at ultra-high spatial resolution with another image provided by theWorldView-2 satellite at high spatial resolution. The drone raw imagery presented some anomalies, such as horizontal banding noise and non-homogeneous radiometry. Camera locations showed a lack of synchrony of the single frequency GPS receiver. The georeferencing process based on ground control points achieved an error lower than 30 cm in X-Y and lower than 55 cm in Z. The drone orthomosaic provided more information in terms of spatial variability in heterogeneous burned areas in comparison with theWorldView-2 satellite imagery. The drone orthomosaic could constitute a viable alternative for the evaluation of post-fire vegetation regeneration in large and heterogeneous burned areas

Fires are one of the main causes of environmental alteration in Mediterranean forest ecosystems. Albedo varies and evolves seasonally based on solar illumination. It is greatly influenced by changes on vegetation: vegetation growth, cutting/planting forests or forest fires. This work analyzes albedo variations due to a large forest fire that occurred on 19- 21 September 2012 in northwestern Spain. From this area, albedo post-fire images (immediately and 1-year after fire) were generated from Landsat 7 Enhanced Thematic Mapper (ETM+) data. Specifically we considered total shortwave albedo, total-, direct-, and diffuse-visible, and near-infrared albedo. Nine to twelve weeks after fire, 111 field plots were measured (27 unburned plots, 84 burned plots). The relationship between albedo values and thematic class (burned/unburned) was evaluated by one-way analysis of variance. Our results demonstrate that albedo changes were related to burned/unburned variable with statistical significance, indicating the importance of forestry areas as regulators of land surface energy fluxes and revealing the potential of post-fire albedo for assessing burned areas. Future research, however, is needed to evaluate the persistence of albedo changes.

The increasing occurrence of large and severe fires in Mediterranean forest ecosystems produces major ecological and socio-economic damage. In this study, we aim to identify the main environmental factors driving fire severity in extreme fire events in Pinus fire prone ecosystems, providing management recommendations for reducing fire effects. The study case was a megafire (11,891 ha) that occurred in a Mediterranean ecosystem dominated by Pinus pinaster Aiton in NW Spain. Fire severity was estimated on the basis of the differenced Normalized Burn Ratio from Landsat 7 ETM +, validated by the field Composite Burn Index. Model predictors included pre-fire vegetation greenness (normalized difference vegetation index and normalized difference water index), pre-fire vegetation structure (canopy cover and vertical complexity estimated from LiDAR), weather conditions (spring cumulative rainfall and mean temperature in August), fire history (fire-free interval) and physical variables (topographic complexity, actual evapotranspiration and water deficit). We applied the Random Forest machine learning algorithm to assess the influence of these environmental factors on fire severity. Models explained 42% of the variance using a parsimonious set of five predictors: NDWI, NDVI, time since the last fire, spring cumulative rainfall, and pre-fire vegetation vertical complexity. The results indicated that fire severity was mostly influenced by pre-fire vegetation greenness. Nevertheless, the effect of pre-fire vegetation greenness was strongly dependent on interactions with the pre-fire vertical structural arrangement of vegetation, fire history and weather conditions (i.e. cumulative rainfall over spring season). Models using only physical variables exhibited a notable association with fire severity. However, results suggested that the control exerted by the physical properties may be partially overcome by the availability and structural characteristics of fuel biomass. Furthermore