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Aims Both human and non-human determinants have shaped Mediterranean forest structure over the last few millennia. The effects of recent human activities on forest composition, however, remain poorly understood. We quantified changes in forest composition during the past century in the mixed forests of Quercus suber (cork oak) and Q. canariensis (Algerian oak), and explored the effects of forest management and environmental (climate, topography) factors on forest structure at various spatial and temporal scales.Location Mountains north of the Strait of Gibraltar (southern Spain).Methods First, we quantified 20th-century changes in species composition from a series of inventories in nine mixed forests (c. 40,000 ha), and examined them in terms of the management practices and environmental conditions. Second, we analysed present-day Q. suber and Q. canariensis stand structure along environmental gradients at two spatial scales: (1) that of the forest landscape (c. 284 ha), combining local inventories and topographic variables and using a digital elevation model; and (2) regional (c. 87,600 km 2 ), combining data from the Spanish Forest Inventory for the Andalusia region and estimates of climatic variables. ResultsHistorical data indicate anthropogenic changes in stand composition, revealing a sharp increase in the density of cork oaks over the last century. Forest management has favoured this species (for cork production) at the expense of Q. canariensis. The impact of management is imprinted on the present-day forest structure. The abundance of both species increases with annual mean precipitation, and they coexist above 800 mm (the minimum threshold for Q. canariensis). Quercus suber dominates in most of the stands, and species segregation in the landscape is associated with the drainage network, Q. canariensis being clearly associated with moister habitats near streams.Main conclusions Our study quantitatively exemplifies a recent humanmediated shift in forest composition. As a result of forest management, the realized niche of the cork oak has been enlarged at the expense of that of Q. canariensis, providing further evidence for humans as major drivers of oak forest composition across the Mediterranean. Recent regeneration problems detected in Q. suber stands, a reduced demand for wood products, conservation policies, and climate change augur new large-scale shifts in forest composition.

Tree species can differ in their responses to resource availability during the critical phase of establishment, which could influence forest dynamics. In Mediterranean forests, most of the attention has focused on the effects of shade and summer drought on seedling survival, but little is known about the effect of autumn to spring rains on earlier stages of recruitment. A sowing experiment was set up along natural light and water gradients with three co-occurring oak species (Quercus suber L. (cork oak), Quercus canariensis Willd. (Algerian oak), and Quercus pyrenaica Willd. (Pyrenean oak)) that show limited natural regeneration in southern Spain. Recruitment stages were monitored for 1 year. Models of seed germination, seedling emergence, and seedling survival as well as of overall recruitment patterns were developed as functions of light, soil moisture, and soil compaction. The influence of intraspecific variation in seed mass and emergence time were also tested. Excess soil water levels during the winter reduced germination and emergence and lengthened time to emergence (in waterlogged open areas), which in turn decreased seedling survival during the dry season. Seedlings from larger seeds were more likely to germinate and emerge. The results suggest that temporal and spatial variability of soil water content, mediated by emergence time and seed size, play a crucial role in the regeneration dynamics of Mediterranean oak forests. Résumé :Les espèces d'arbre peuvent avoir des réactions différentes face à la disponibilité des ressources pendant la phase critique d'établissement, ce qui peut influencer la dynamique forestière. Dans les forêts méditerranéennes, on s'est surtout préoccupé des effets du manque de lumière et de la sécheresse estivale sur la survie des semis, mais les effets de la pluie qui tombe de l'automne au printemps sur les premiers stades de recrutement sont peu connus. Une expérience d'ensemencement a été établie le long de gradients naturels de disponibilité de lumière et d'eau avec trois espèces de chêne sympatriques (Quercus suber L., Quercus canariensis Willd. et Quercus pyrenaica Willd.) dont la régénération naturelle est peu abondante dans le sud de l'Espagne. Les stades de recrutement ont été suivis pendant une année. Des modèles de germination, d'émergence et de survie de même que des patrons généraux de recrutement ont été établis en fonction de la lumière, de l'humidité du sol et de la compaction du sol. L'influence de la variation intraspécifique du poids des semences et de l'émergence des semis a aussi été testée. Un excès d'eau du sol pendant l'hiver a diminué la germination et l'émergence et allongé la période de temps nécessaire à l'émergence (dans les endroits ouverts et saturés en eau), ce qui a diminué le taux de survie des semis pendant la saison sèche. Les semis issus de grosses semences avaient plus de chances de germer et d'émerger. Ces résultats indiquent que la variabilité temporelle et spatiale de la teneur en eau du sol, par l'intermédiaire de la période d'émergence et...

Questions: What is the current distribution of pine and oak species along environmental gradients in southern Spain? Do pine and oak regeneration niches differ from the environmental niches of adults? Is oak species regeneration favoured under the canopy of pine forests?Location: Forest areas of Andalusia ($87 600 km 2 , southern Spain). Methods:We compiled extensive forest inventory data to explore differences in abundance (basal area, m 2 ha À1 ) patterns of adults (dbh 4 7.4 cm) and regeneration (dbh 7.4 cm) of five pine and five oak species. Canonical correspondence analysis (CCA) and generalized linear models were applied to explore species-environment relationships along climatic, edaphic, topographic and fire-frequency gradients.Results: Both pines and oaks segregated along complex environmental gradients, with pines generally dominating in more severe (colder and drier) environments, while oaks dominated in milder, wetter winter areas. In 40-55% of mature pine stands there was a lack of regeneration in the understorey, while in two oak species (Q. suber and Q. canariensis) 70% of stands did not show regeneration. Pine recruits were found at a higher frequency and abundance under the canopy of their congeners, whereas some oaks (Q. ilex) had greater regeneration under mixed pine-oak canopies.Conclusions: Climatic limitations and soil properties partly explained the regional distribution of pines and oaks. We found evidence for an upward shift of Q. ilex recruits towards areas with colder conditions in pine forests, which could be explained by a possible facilitative effect of the pine canopy on seedling establishment.

Abstract. Wildfires are a major concern on the Iberian Peninsula, and the establishment of effective prevention and early warning systems are crucial to reduce impacts and losses. Fire weather indices are daily indicators of fire danger based upon meteorological information. However, their application in many studies is conditioned to the availability of sufficiently large climatological time series over extensive geographical areas and of sufficient quality. Furthermore, wind and relative humidity, important for the calculation of fire spread and fuel flammability parameters, are relatively scarce data. For these reasons, different reanalysis products are often used for the calculation of surrogate fire danger indices, although the agreement with those derived from observations remains as an open question to be addressed.In this study, we analyze this problem focusing on the Canadian Fire Weather Index (FWI) -and the associated Seasonal Severity Rating (SSR) -and considering three different reanalysis products of varying resolutions on the Iberian Peninsula: NCEP, ERA-40 and ERA-Interim. Besides the inter-comparison of the resulting FWI/SSR values, we also study their correspondence with observational data from 7 weather stations in Spain and their sensitivity to the input parameters (precipitation, temperature, relative humidity and wind velocity).As a general result, ERA-Interim reproduces the observed FWI magnitudes with better accuracy than NCEP, with lower/higher correlations in the coast/inland locations. For instance, ERA-Interim summer correlations are above 0.5 in inland locations -where higher FWI magnitudes are attained -whereas the corresponding values for NCEP are below this threshold. Nevertheless, departures from the observed distributions are generally found in all reanalysis, with a general tendency to underestimation, more pronounced in the case of NCEP. In spite of these limitations, ERA-Interim may still be useful for the identification of extreme fire danger events.(e.g. those above the 90th percentile value) and for the definition of danger levels/classes (with level thresholds adapted to the observed/reanalysis distributions).

Seed predation and dispersal can critically influence plant community structure and dynamics. Inter-specific differences arising at these early stages play a crucial role on tree recruitment patterns, which in turn could influence forest dynamics and species segregation in heterogeneous environments such as Mediterranean forests. We investigated removal rates from acorns set onto the ground in two coexisting Mediterranean oak species Á Quercus canariensis and Q. suber Á in southern Spain. We developed maximum likelihood estimators to investigate the main factors controlling probabilities of seed removal and to describe species-specific functional responses. To account for inter-specific differences in seed-drop timing, two experiments were established: a simultaneous exposure of acorns of the two species (synchronous experiments) and a seed exposure following their natural seed-drop phenology (diachronic experiments). A total of 1536 acorns were experimentally distributed along a wide and natural gradient of plant cover, and removal was periodically monitored for three months at two consecutive years (with contrasting differences in seed production and thus seed availability on the ground). The probability of seed removal increased with plant cover (leaf area index, LAI) for the two oak species. Inter-specific differences in acorn removal were higher in open areas and disappeared in closed microhabitats, especially during a non-mast year. Despite later seed-drop, Q. suber acorns were removed faster and at a higher proportion than those of Q. canariensis. The higher probability of seed removal for this species could be attributed to its larger seed size compared to Q. canariensis, as inter-specific differences were less pronounced when similar sized acorns were exposed. Inter-specific differences in seed removal, arising from seed size variability and microsite heterogeneity, could be of paramount importance in oak species niche separation, driving stand dynamics and composition along environmental gradients.

Summary1. In heterogeneous environments, species segregate spatially in response to selective abiotic and biotic filters occurring throughout plant ontogeny. Ontogenetic conflicts in recruitment may lead to spatially discordant patterns of regeneration among microhabitats with different plant cover. In addition, species differing in seed size may be subjected to opposing ecological and evolutionary pressures throughout the life cycle of the plant. 2. We used a multi-stage demographic approach aimed at characterizing the main stage-specific probabilities of recruitment (seed survival, seed germination, seedling emergence and survival during the first 3 years of life) in two Mediterranean oak species coexisting at southern Spain. We calibrated linear and nonlinear likelihood models for each of these consecutive life history stages and calculated overall probabilities of recruitment along a wide range of plant cover and seed size variation. 3. Seed predation and seedling mortality over the dry season were the most limiting processes for the two studied oak species. However, species ranking diverged substantially through the life history stages considered in this study due to different ontogenetic trends among species. 4. At the intraspecific level, recruitment-driving processes during the seed and the seedling stages showed opposing tendencies along the explored range of plant cover and seed size. Thus, small-sized acorns and open areas were favoured for the seed stage, whereas large acorns and dense microhabitats did for the seedling stage. 5. The existence of opposing selective pressures on seed mass and their differential influence on the two studied oak species determined the occurrence of species-specific optimal seed sizes (small acorns for Quercus canariensis vs. acorns of large or intermediate size for Quercus suber). 6. The spatial patterns predicted by our overall-recruitment models provided some evidence of regeneration niche partitioning in the two coexisting oak species, supporting their current distribution patterns as saplings and adults at the study area. 7. Synthesis. We conclude that within-and among-species differences through plant ontogeny, arising from species differential response to microhabitat heterogeneity and seed size variation, could be of great importance for oak species niche segregation, driving stand dynamics and spatial pattern distribution along the landscape. The information provided by this study could be also applied to optimize management and restoration programmes since it has enabled us to identify the most favourable conditions and traits for recruitment in oak species that exhibit serious constraints for natural regeneration.

Abstract. Understanding how fire weather danger indices changed in the past and how such changes affected forest fire activity is important in a changing climate. We used the Canadian Fire Weather Index (FWI), calculated from two reanalysis data sets, ERA-40 and ERA Interim, to examine the temporal variation of forest fire danger in Europe in 1960–2012. Additionally, we used national forest fire statistics from Greece, Spain and Finland to examine the relationship between fire danger and fires. There is no obvious trend in fire danger for the time period covered by ERA-40 (1960–1999), whereas for the period 1980–2012 covered by ERA Interim, the mean FWI shows an increasing trend for southern and eastern Europe which is significant at the 99% confidence level. The cross correlations calculated at the national level in Greece, Spain and Finland between total area burned and mean FWI of the current season is of the order of 0.6, demonstrating the extent to which the current fire-season weather can explain forest fires. To summarize, fire risk is multifaceted, and while climate is a major determinant, other factors can contribute to it, either positively or negatively.