Landscape Implications of Contemporary Abandonment of Extensive Sheep Grazing in a Globally Important Agricultural Heritage System

[EN], In Southern Europe, the abandonment of the traditional transhumant system where sheep graze lowland areas during winter and distant mountain systems during summer has led to an important cultural loss and still poorly understood ecological consequences. We investigate the landscape-scale implications of contemporary sheep grazing patterns in a Globally Important Agricultural Heritage System (GIAHS). Our analysis showed a heterogeneous use of mountain grazing areas between 1990 and 2020. The areas most used by sheep had more abundance of pasture, fewer forests, and structurally different landscapes than those that had been fully abandoned by sheep. Likewise, we have detected decreasing trends in landscape diversity in those areas not used by sheep over the study period, whereas landscape heterogeneity is maintained in those areas grazed by sheep. Our study constitutes an original analysis of landscape patterns and shifts in relation to extensive sheep grazing by using novel approaches that combine interviews, updated satellite time series, and state-of-the-art landscape analysis techniques. Likewise, our results constitute a benchmark as they inform on the importance of preserving extensive sheep grazing if we aim to maintain the cultural heritage, and traditional diverse landscape and the semi-natural grasslands in the Mountains of León.SIEuropean Union Next-Generation EU fundsCESEFORMAP

The seed bank in Pinus stand regeneration in NW Spain after wildfire

P. 22-31In 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.S

Influence of high temperatures on seed germination of a special Pinus pinaster stand adapted to frequent fires

P. 129–136This 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

Disruption of trophic interactions involving the heather beetle by atmospheric nitrogen deposition

P. 436-445Elevated 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.S

Plant and vegetation functional responses to cumulative high nitrogen deposition in rear-edge heathlands

P. 980-990Elevated atmospheric nitrogen (N) deposition is a major driver of change, altering the structure/functioning of nutrient-poor Calluna vulgaris-heathlands over Europe. These effects amply proven for north-western/central heathlands may, however, vary across the ecosystem's distribution, especially at the range limits, as heathlands are highly vulnerable to land-use changes combined with present climate change. This is an often overlooked and greatly understudied aspect of the ecology of heathlands facing global change. We investigated the effects of five N-fertilisation treatments simulating a range of N deposition rates (0, 10, 20, and 50 kg N ha−1 yr−1 for 1 year; and 56 kg N ha−1 yr−1 for 9 years) on the Calluna-plants, the plant functional groups, species composition and richness of two life-cycle stages (building/young- and mature-phase) of Calluna-heathlands at their rear-edge limit. Our findings revealed a dose-related response of the shoot length and number of flowers of young and mature Calluna-plants to the addition of N, adhering to the findings from other heathland locations. However, cumulative high-N loading reduced the annual growth and flowering of young plants, showing early signs of N saturation. The different plant functional groups showed contrasting responses to the cumulative addition of N: annual/perennial forbs and annual graminoids increased with quite low values; perennial graminoids were rather abundant in young heathlands but only slightly augmented in mature ones; while bryophytes and lichens strongly declined at the two heathland life-cycle stages. Meanwhile there were no significant N-driven changes in plant species composition and richness. Our results demonstrated that Calluna-heathlands at their low-latitude distribution limit are moderately resistant to cumulative high-N loading. As north-western/central European heathlands under high-N inputs broadly experienced the loss of plant diversity and pronounced changes in plant species dominance, rear-edge locations may be of critical importance to unravel the mechanisms of heathland resilience to future global change.S

The footprint of large wildfires on the multifunctionality of fire-prone pine ecosystems is driven by the interaction of fire regime attributes

[EN], Background Mediterranean ecosystems dominated by Pinus pinaster Ait. (maritime pine) are subject to a shift from fuel-limited to drought-driven fire regimes, characterized by an increasing wildfire extent, recurrence, and severity. Previous studies have not addressed the interacting effects of fire recurrence and severity on the ecosystem multifunctionality (EMF) of maritime pine forests, although complex relationships between such fire regime attributes are expected. Here, we evaluated the medium-term effects of fire recurrence and severity on the EMF response of unmanaged, native pine ecosystems dominated by Pinus pinaster in the western Mediterranean Basin. We considered four key ecosystem functions computed from functional indicators (carbon regulation, decomposition, soil fertility, and plant production), which were pooled into an EMF construct. The fire regime effects on the trade-offs and synergies between the considered ecosystem functions were also analyzed. Results Multiple ecosystem functions responded differentially to fire recurrence and severity. Fire recurrence had a strong effect on soil fertility, decomposition, and plant production functions. No significant effects of fire severity on any of the individual functions were detected. However, both fire regime attributes interacted to determine soil fertility and decomposition functions, suggesting that their performance is only impaired by fire severity when fire recurrence is low. The differing responses to the fire regime attributes among ecosystem functions fostered a significant EMF response to fire severity and its interaction with fire recurrence, indicating that the effect of fire severity on EMF was stronger under low fire recurrence scenarios, even when relationships between individual functions and fire severity were weak. Fire recurrence caused significant trade-offs between functions to emerge. However, these trade-offs were not strong enough to differ significantly from the intrinsic trade-offs (i.e., regardless of the fire regime) of maritime pine ecosystems. Conclusions Our results indicated the need to use an integrative approach to assess the response of ecosystem functioning to the fire regime in maritime pine ecosystems. Adaptive management responses are necessary towards the minimization of repeated burnings and the reduction of the fuel load in unmanaged maritime pine stands of the western Mediterranean Basin with similar characteristics to those analyzed in this study.[ES], Antecedentes Los ecosistemas mediterráneos dominados por pino marítimo (Pinus pinaster Ait.) están sujetos a cambios en regímenes de fuego limitados por el combustible hacia regímenes conducidos por la sequía, y caracterizados por un incremento en la extensión, recurrencia y severidad de los incendios. Estudios previos no han abordado los efectos interactivos de la recurrencia y severidad del fuego en la multifuncionalidad de los ecosistemas (EMF) en bosques de pino marítimo, aunque cabe esperar relaciones complejas entre estos atributos del regimen de fuego. En este trabajo, evaluamos los efectos a medio plazo de la recurrencia y severidad en la respuesta de la multifuncionalidad de los ecosistemas (EMF) de bosques nativos dominados por pino marítimo no gestionados en la cuenca Mediterránea occidental. Consideramos cuatro funciones clave calculadas a partir de indicadores funcionales (regulación del carbono, descomposición, fertilidad del suelo, y producción egetal) los cuales fueron agrupados en un constructo EMF. Los efectos del régimen de fuego sobre las sinergias y contrapartidas entre las funciones ecosistémicas también fueron analizados. Resultados Múltiples funciones ecosistémicas respondieron diferencialmente a la recurrencia y severidad. La recurrencia del fuego tuvo un efecto muy fuerte en la fertilidad del suelo, en la descomposición y en las funciones de producción. Ningún efecto significativo de la severidad del fuego fue detectado en ninguna de las funciones individuales. Sin embargo, los atributos de ambos regímenes de fuego interactuaron para determinar las funciones de fertilidad y descomposición, sugiriendo que su rendimiento es afectado por la severidad solo cuando la recurrencia del fuego es baja. Las diferentes respuestas a los atributos de los regímenes de fuego entre las funciones ecosistémicas promueven una respuesta significativa de la EMF a la severidad del fuego y su interacción con la recurrencia, indicando que el efecto de la severidad sobre la EMF fue más fuerte bajo escenarios de baja recurrencia, aun cuando las relaciones entre funciones individuales y la severidad fueran débiles. La recurrencia del fuego causó la aparición de ontrapartidas significativas entre funciones. Obviamente, estas contrapartidas no fueron lo suficientemente fuertes para diferir significativamente de aquellas intrínsecas (i.e., independientemente del régimen de fuego) en los ecosistemas de pino marítimo. Conclusiones Nuestros resultados indican la necesidad de usar una aproximación integrada para determinar la respuesta del funcionamiento al régimen de fuego en ecosistemas de pino marítimo. Respuestas de manejo adaptativo son necesarias para la minimización de quemas repetidas y la reducción de la carga de combustible en rodales de pino marítimo no gestionados en la cuenca Mediterránea, con características similares a aquellos analizados en este estudio.SIAEIBritish Ecological SocietyPortuguese Foundation for Science and Technolog

Modeling Pinus pinaster forest structure after a large wildfire using remote sensing data at high spatial resolution

P. 257-271In 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 large and heterogeneous burnt areas. Given the accuracy and predictive capacity of the models obtained in this study, high spatial resolution satellite imagery together with field data provide useful information in post-fire decision making in fire prone ecosystems.S

Soil-vegetation relationships in Mediterranean forests after fire

P.1-13Background Wildfires are one of the major environmental concerns in Mediterranean ecosystems. Thus, many studies have addressed wildfire impacts on soil and vegetation in Mediterranean forests, but the linkages between these ecosystem compartments after fire are not well understood. The aim of this work is to analyze soil-vegetation relationships in Mediterranean burned forests as well as the consistency of these relationships among forests with different environmental conditions, at different times after fire, and among vegetation with different functional traits. Results Our results indicate that study site conditions play an important role in mediating soil-vegetation relationships. Likewise, we found that the nature of soil-vegetation relationships may vary over time as fire effects are less dominant in both ecosystem compartments. Despite this, we detected several common soil-vegetation relationships among study sites and times after fire. For instance, our results revealed that available P content and stoichiometry (C:P and N:P) were closely linked to vegetation growth, and particularly to the growth of trees. We found that enzymatic activities and microbial biomass were inversely related to vegetation growth rates, whereas the specific activities of soil enzymes were higher in the areas with more vegetation height and cover. Likewise, our results suggest that resprouters may influence soil properties more than seeders, the growth of seeders being more dependent on soil status. Conclusions We provide pioneer insights into how vegetation is influenced by soil, and vice-versa, in Mediterranean burned areas. Our results reflect variability in soil-vegetation relationships among study sites and time after fire, but consistent patterns between soil properties and vegetation were also detected. Our research is highly relevant to advance in forest science and could be useful to achieve efficient post-fire management.S

Provenance and seed mass determine seed tolerance 5 to high temperatures associated to forest fires in Pinus pinaster

P. 381-391We show that Pinus pinaster provenance affects both seed germination and seedling recruitment after fire. This information is crucial for managers to select the best seed-provisioning populations when implementing reforestation programs to assist the natural post-fire regeneration of the speciesS

Radar and multispectral remote sensing data accurately estimate vegetation vertical structure diversity as a fire resilience indicator

The structural complexity of plant communities contributes to maintaining the ecosystem functioning in fire-prone landscapes and plays a crucial role in driving ecological resilience to fire. The objective of this study was to evaluate the resilience to fire off several plant communities with reference to the temporal evolution of their vertical structural diversity (VSD) estimated from the data fusion of C-band synthetic aperture radar (SAR) backscatter (Sentinel-1) and multispectral remote sensing reflectance (Sentinel-2) in a burned landscape of the western Mediterranean Basin. We estimated VSD in the field 1 and 2 years after fire using Shannon’s index as a measure of vertical heterogeneity in vegetation structure from the vegetation cover in several strata, both in burned and unburned control plots. Random forest (RF) was used to model VSD in the control (analogous to prefire scenario) and burned plots (1 year after fire) using as predictors (i) Sentinel-1 VV and VH backscatter coefficients and (ii) surface reflectance of Sentinel-2 bands. The transferability of the RF model from 1 to 2 years after wildfire was also evaluated. We generated VSD prediction maps across the study site for the prefire scenario and 1 to 4 years postfire. RF models accurately explained VSD in unburned control plots (R2 = 87.68; RMSE = 0.16) and burned plots 1 year after fire (R2 = 80.48; RMSE = 0.13). RF model transferability only involved a reduction in the VSD predictive capacity from 0.13 to 0.20 in terms of RMSE. The VSD of each plant community 4 years after the fire disturbance was significantly lower than in the prefire scenario. Plant communities dominated by resprouter species featured significantly higher VSD recovery values than communities dominated by facultative or obligate seeders. Our results support the applicability of SAR and multispectral data fusion for monitoring VSD as a generalizable resilience indicator in fire-prone landscapes.SIEuropean Regional Development Fund (ERDF)Spanish Ministry of Economy and CompetitivenessRegional Government of Castilla and LeónBritish Ecological Societ