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Received 11 December 2015, Revised 1 March 2016, Accepted 3 March 2016, Available online 12 March 2016 ; Demographic and genetic connectivity of fragmented plant populations will depend on effective propagule flow across the landscape. We analyze functional connectivity in a holm oak (Quercus ilex) fragmented landscape by considering three important stages driving recruitment: effective pollination, acorn production and acorn dispersal. We used a network approach to (1) determine if pollen-mediated gene exchange across the landscape was spatially structured; (2) estimate the effects of limited acorn dispersal on functional connectivity; (3) identify which landscape traits could drive source–sink dynamics of gene flow. Although long distance dispersal was relatively frequent, most effective pollen flow occurred over short distances (10 ha) are the main pollen sources, while small ones (<1 ha) are important pollen sinks. Thus, big fragments are critical to maintain functional connectivity, while small forest fragments may provide acorn crops better representing regional genetic diversity. In addition to area effects, less isolated and more central fragments showed higher migration rates and exchanged effective pollen with more fragments. Hence, we expected that landscapes with uniform or clumped distribution of big forest fragments would show optimal connectivity traits. However, despite that simulated gene flow was more evenly distributed across the landscape, connectance and migration rates decreased. Our results call for caution before translating patch-level management guidelines to the landscape scale. They also show that the level of functional connectivity may change throughout the recruitment process, suggesting that large-scale conservation strategies may fail if local effective seed establishment is disregarded. ; TM-L was beneficiary of a FPI grant funded by the Spanish Government (BES-2011-048346). This paper is a contribution to the Spanish-funded projects VULGLO (CGL2010e22180-C03e03), VERONICA (CGL2013-42271-P) and REMEDINAL3eCM (S2013/MAE-2719). ; Peer reviewed

Received 27 December 2015, Revised 14 March 2016, Accepted 19 March 2016, Available online 31 March 2016 ; The effects of fragmentation on acorn production should be mediated by their impacts on the physiological status of oaks during seed development particularly in water-limited systems, such as Mediterranean forests. The creation of forests edges reduces tree-to-tree competition, which may in turn temper water shortage during summer and, as a result, enhance acorn production. To test these two hypotheses we monitored acorn production and predawn water potential during the 2012–2014 period in two holm oak (Quercus ilex) forest archipelagos of the Iberian Peninsula. Acorn production and fragmentation effects did not differ between localities despite of their contrasting climatic conditions (accumulated water deficit from April to August was a 60% higher in the South). In general, forest interiors showed a high proportion of non-producing trees (∼50%) while trees at small forest fragments showed high acorn crops (acorn score ⩾3, ∼40% of studied trees). Our results confirmed the expectation that intraspecific competition in small forest fragments was reduced, which alleviated summer water shortage of the trees studied. This reduced water stress entailed an increased acorn production. Overall, our results show that local processes such as fragmentation may counteract climatic differences among localities and could even override the impacts of increased aridity on acorn crops. ; We are also grateful to Laura Barrios for her help in the statistical analysis. Teresa Morán-López was beneficiary of a FPI grant (funded by the Spanish Government (BES-2011-048346). This paper is a contribution to the Spanish-funded projects VULGLO (CGL2010–22180-C03–03), VERONICA (CGL2013-42271-P) and REMEDINAL 2 & 3 (CM S2009 AMB 1783) (S2013/MAE-2719). ; Peer reviewed

The historical formation of holm oak (Quercus ilex L.) dehesa woodlands has often resulted in either rapid deforestation episodes by means of clear cuts or slower depletion of oak woodlands due to tree natural mortality and the absence of tree natural regeneration. Consequently, holm oak woodlands have been progressively converted into rough pasturelands, cereal croplands and scrublands. Whilst this tendency of holm oak exhaustion was not socially questioned, scientists and conservationists defending oak woodland conservation providing long run economic and environmental arguments were regarded as romantics ignoring the necessity for social progress and development. Today, these romantics are perceived as nature conservation pioneers and many of their arguments and concepts have been included in science and politics. This paper shows that cereal cropping and permanent grassland uses are more profitable to Extremadura's landowners than holm oak artificial plantations and monitored holm oak natural regeneration. Increasing social demand for recreational and conservation services have been taken into account by Spanish and European public authorities. Along last decade, the European Union and the Spanish government have financed oak reforestation over extensive-use croplands of West and Southwest Spain, resulting in an unprecedented net increase in holm oak woodlands. The presented extended cost-benefit analysis shows that even after incorporating estimated income derived from public and private consumption of environmental services, uses associated to treeless cropland, pastureland and scrubland are still preferred to traditional extensive multiple use associated to dehesa holm oak woodlands. ; Peer reviewed

Forest decline is nowadays a major challenge to management and sustainability of natural ecosystems worldwide. This syndrome is a multifactorial disease influenced by several biotic and abiotic agents such as alien invasive pathogens, changes in land use and management policies, population dynamics driven by economic and politic changes, and climatic perturbations. All these factors changing due to anthropogenic influence, together with others, conformed the so-called global change. Since the 1990's decade, the oak decline has been identified as one of the most important ecological problems in Europe, affecting deciduous and evergreen Quercus species from the continental forests on Central and North Europe, to temperate forests of the Mediterranean basin. In the case of Iberian Peninsula, holm oak (Quercus ilex L.) and cork oak (Quercus suber L.) decline has been detected since the 1980's decade. These two species covered most of the forest surface of the south and central part of the Iberian Peninsula, mainly through "dehesas" and "montados" formations. This area is considered to be one of the worst affected regions in the world by climate change, worsening the effects and the consequences of oak decline in "dehesas". "Dehesas" are Mediterranean savanah-like ecosystems, which provide several economic yields and ecological services. In turn, the holm oak is the most representative tree in the Iberian Peninsula, and the main species conforming the tree layer on the Spanish "dehesas". The loss of this tree layer is a major ecological and economical constraint. The holm oak decline is mainly associated to the action of soil-borne pathogens, especially Phytophtora cinnamomi. Despite the influence of other factors in the decline, there is a strong association between root rot caused by oomycetes and the death of trees. Phytophtora cinnamomi is an aggressive alien plant pathogen widely widespread worldwide, which has been associated with the disease, die-off and death in a large list of different plant hosts. It is able to change trophic relationships with their hosts, becaming biotroph in asymptomatic hosts and hemibiotroph or necrotroph in susceptible hosts. It is considered that holm oak is the most susceptible Quercus sp. to the action of this pathogen. Additionally, other Phytophthora spp. and Pythium spp. have been recorded associated with the holm oak decline in Spain, Portugal, Italy and France. Many scientific efforts have focused to study this important hostpathogen system, obtaining great results and increasing the knowledge of the causes and effects of the interaction, improving management techniques to limit the spread and the symptoms. However, most of the reviewed works are based on empirical approaches, being the underlying mechanisms regulating the interaction between both species mostly unknown. This PhD Thesis covers part of this lack of basic knowledge, focusing on the host-pathogen interaction at histological and physiological level, and exploring the influence of the soil biota in the severity of the disease symptoms. For this purpose, the work was structured in seven chapters. Chapter 1 provides the framework in which the present PhD Thesis has been developed, and the general and specific objectives. Chapter 2 presents the methodology developed to evaluate the colonization and infection of Phytophthora cinnamomi in Quercus ilex seedling through the semi-automated quantification of pathogen structures present in histological sections of fine roots. A workflow was tuned-up testing different fixing solutions, embedding substances and staining methods, and the results allowed the clear differentiation of pathogen structures from host tissues. Furthermore, different indices based on structure location, host tissue classification and specificity of pathogen structures were evaluated to find the easiest and statistically robust indices that are able to explain the progress of the oomycete into the root. In chapter 3, inoculation experiments in growth chamber were carried out to describe the pathogenesis of the Quercus ilex-Phytophthora cinnamomi interaction. Longitudinal sections were analysed for epidermal, cortex, parenchymatous tissue of central cylinder and vascular tissue after 1, 3, 7 and 14 days after inoculation. Total oomycete structures area, intracellular structures area, extracellular structures area, and specific structures area of the pathogen were quantified. The analysis of these data results in the description of the colonization/infection cycle of the pathogen, classified in three different stages related with their trophic behaviour. Moreover, histological changes of the root tissues as a result of the presence of defence responses and the action of the pathogen were described. In Chapter 4, the differential responses of holm oak seedlings to the inoculation with P. cinnamomi, the acute drought and combination of both stressors were assessed. Six-months old seedlings were inoculated and mock-inoculated, and half of each inoculation treatment plans were subjected to acute drought meanwhile the others were well irrigated. Photosynthesis, stomatal conductance and fluorescence were measured weekly, and total biomass and biomass allocation parameters were quantified at the end of the experiment. The resulting data showed differences in the response of seedlings to drought and inoculation, and the influence of the additive effect of both stressors in the seedlings die-off. In Chapter 5, soil samples of "dehesas" were collected and total DNA was extracted and analysed through metabarcoding techniques, to evaluate the specific composition and diversity of the fungal and oomycete communities, and to study their relationship with the disease symptoms. The fungal community included a wide range of pathogens and abundance of ectomycorrhizal key taxa. Phytophthora spp. dominated the oomycete community, but the species related to root rot did not appear as the most abundant, nor were they related directly to defoliation levels. A particular Operational Taxonomic Unit (OTU) belonging to the genus Trichoderma was strongly correlated with the scarcity of pathogenic Phytophthora spp. The differences in defoliation were related to changes in the functionality of soil microbiota and diversity levels of pathogenic species. Chapter 6 presents the general discussion of the Thesis, including some aspects limiting the results of the works carried out, ad new work lines deriving from this Thesis, and finally Chapter 7 contains the conclusions of the work. Changes in roots as a consequence of P. cinnamomi inoculation, including defence responses, and the differential response identified with pathogen colonization/inoculation, leads to new insights about the causes of tree death. Holm oak responds to the attack of the pathogen, and their physiological changes differ from the ones caused by water stress, allowing the recovery of plants if no additional stress is present. Moreover, the study of soil microbiome in declined "dehesas" showed the influence of the microbial diversity in the health status of trees, and also presented new species of oomycetes and fungi that must be considered in the management of holm oak decline in Andalusian "dehesas". ; El decaimiento forestal es hoy en día uno de los mayores desafíos para el manejo y la sostenibilidad de los ecosistemas naturales en todo el mundo. Dicho síndrome es una enfermedad multifactorial en la que intervienen diversos agentes bióticos y abióticos como los patógenos invasores, los cambios en los usos del territorio y las políticas de recursos, o las perturbaciones climáticas. El cambio producido en todos estos factores debido a la influencia del hombre entre otros motivos es lo que se ha dado en llamar cambio global. Desde la década de los 90 del siglo XX, el decaimiento de los robles se ha identificado como uno de los problemas ecológicos más relevantes en Europa, afectando a masas de Quercus caducifolios y perennifolios desde los bosques continentales de Centro Europa y Norte Europa, hasta los bosques templados de la cuenca mediterránea. En el caso de la Península Ibérica, el decaimiento de la encina (Quercus ilex L.) y el alcornoque (Quercus suber L.) se identificó en los años 80 del siglo XX. Estas dos especies cubren la mayor parte de la superficie forestal del sur y centro de la Península Ibérica, principalmente formando sistemas de dehesa y "montados". Dicha área geográfica está considerada como una de las regiones a nivel mundial que se verán peor afectadas por el cambio climático, lo que agravaría las consecuencias y los efectos del decaimiento de la encina en las dehesas. Las dehesas son ecosistemas mediterráneos semejantes a la sabana, que proveen de diversos beneficios económicos y servicios ambientales. Por su parte, la encina es el árbol más representativo de la Península Ibérica, y la especie principal del estrato arbóreo de las dehesas. La pérdida de este estrato es un problema ecológico y económico de gran relevancia. El decaimiento de la encina está asociado principalmente a la acción de patógenos de suelo, especialmente Phytophthora cinnamomi. Sin olvidar la influencia de otros factores en el síndrome, existe una fuerte asociación entre la podredumbre radicular causada por los oomicetos y la muerte del arbolado. Phytophthora cinnamomi es un patógeno invasor muy agresivo, ampliamente distribuido por todo el mundo, que ha sido asociado con la enfermedad, la decadencia y la mortalidad de una larga lista de diferentes especies vegetales. Es una especie capaz de cambiar su relación trófica con el huésped, comportándose como un organismo biótrofo en huéspedes asintomáticos, y como hemibiótrofo ó necrótrofo en huéspedes susceptibles. Se considera que la encina es la especie del género Quercus más susceptible a la acción del patógeno. De forma adicional, otras especies de los géneros Phytophthora y Pythium se han encontrado asociadas con el decaimiento de la encina en España, Portugal, Italia y Francia. Para estudiar este sistema patógeno-huésped se han llevado a cabo grandes esfuerzos científicos, que han obtenido resultados muy meritorios y han incrementado el conocimiento de las causas y los efectos de la interacción, mejorando las técnicas de gestión para limitar la dispersión y los síntomas del decaimiento. Sin embargo, la mayoría de los trabajos revisados se basaron en aproximaciones empíricas, desconociéndose gran parte de los mecanismos subyacentes que controlan la interacción entre las dos especies. Esta Tesis cubre parte de esta falta de conocimiento básico, centrándose en la interacción entre el huésped y el patógeno a nivel histológico y fisiológico, así como explorando la influencia de la biota del suelo en la severidad de los síntomas de la enfermedad. Para dicho propósito, el trabajo se estructuró en 7 capítulos. El Capítulo 1 proporciona el marco teórico en el que se desarrolla esta Tesis Doctoral, así como los objetivos generales y específicos. El Capítulo 2 muestra la metodología desarrollada para evaluar la colonización e infección de Phytophthora cinnamomi en plántulas de Quercus ilex a través de la cuantificación semiautomática de las estructuras del patógeno presentes en secciones histológicas de raíces finas. El flujo de trabajo fue puesto a punto probando diferentes soluciones de fijación, sustancias de inclusión y métodos de tinción, y los resultados permitieron la diferenciación clara de las estructuras del patógeno y de los tejidos del huésped. Además, distintos índices basados en la localización y especificidad de las estructuras del patógeno y en el tejido del huésped, se evaluaron con el fin de buscar la manera más sencilla y estadísticamente robusta de explicar el progreso del oomiceto en la raíz a través de índices. En el capítulo 3, se llevaron a cabo experimentos en cámara de crecimiento para describir la patogénesis de la interacción entre P. cinnamomi y Q. ilex. Se analizaron secciones longitudinales de epidermis, córtex, tejido parenquimático del cilindro central y tejido vascular, a los 1, 3, 7 y 14 días después de la inoculación. Se cuantificó el área total de estructuras, estructuras intracelulares, estructuras extracelulares y estructuras específicas del patógeno. El análisis de estos datos proporcionó una descripción del ciclo de colonización/infección del patógeno, clasificada en tres etapas diferentes relacionadas con su comportamiento trófico. Asimismo se describieron los cambios histológicos resultantes de la presencia del patógeno o de las respuestas desencadenadas por la planta. En el capítulo 4 se analizó la respuesta diferencial de plántulas de encina ante la inoculación con P. cinnamomi, ante la sequía severa y ante ambos estreses combinados. Plántulas de seis meses de edad fueron inoculados y sometidos a falsa inoculación, y la mitad de cada uno de estos tratamientos fue sometida a sequía severa, mientras que el resto fue regada de manera óptima. Semanalmente se midieron los valores de fotosíntesis, conductancia estomática y fluorescencia, y la biomasa total así como la compartimentación de la biomasa fueron cuantificadas al final del experimento. Los datos resultantes mostraron la existencia de diferencias en la respuesta de las plántulas ante la sequía y la inoculación, así como el efecto aditivo de ambos estreses en la muerte de las plántulas. En el Capítulo 5 se colectaron muestras de suelo de dehesas y se extrajo el ADN total, que se analizó a través de técnicas de metabarcoding, con el fin de evaluar la composición específica y la diversidad de las comunidades fúngica y de oomicetos, y para estudiar sus relaciones con los síntomas de la enfermedad. La comunidad fúngica presentó una gran variedad de patógenos y abundancia de taxones clave de ectomicorrizas. Phytophthora spp. apareció como el taxón dominante dentro de la comunidad de oomicetos, pero las principales especies relacionadas con la podredumbre radicular no fueron las más abundantes, ni presentaron relación directa con los niveles de defoliación. Una unidad taxonómica operacional (OTU) particular, perteneciente al género Trichoderma, presentó correlaciones significativas con la escasez de especies patógenas de Phytophthora spp. Las diferencias en defoliación se correlacionaron con cambios en la funcionalidad de la microbiota del suelo y con los niveles de diversidad de las especies patógenas. El Capítulo 6 presenta la discusión general de la Tesis, incluyendo algunos aspectos que limitan los resultados de los trabajos realizados, y nuevas líneas de trabajo que se derivan de esta Tesis, y finalmente, el Capítulo 7 contiene las conclusiones del trabajo. Los cambios que se producen en la raíz a consecuencia de la inoculación con P. cinnamomi, incluyendo las respuestas defensivas, así como la respuesta diferencial identificada con la colonización/infección, conducen a nuevas apreciaciones sobre la causa de la muerte del arbolado. La encina responde al ataque del patógeno, presentando variaciones en la fisiología diferentes de las causadas por el estrés hídrico, las cuales permiten la recuperación de las plantas si no se superpone un estrés adicional a la inoculación. Adicionalmente, el estudio del microbioma del suelo en dehesas con decaimiento del encinar mostró la influencia de la diversidad microbiana en el estado sanitario del arbolado, así como mostró nuevas especies de oomicetos y hongos que deben tenerse en consideración en el manejo del decaimiento de las dehesas de encina en Andalucía.

Water is the key element that modulates the provision of goods and services together with global/climate stressors affecting semiarid forests. In this sense, there is a need to improve the understanding and quantification of forest and water relationships as affected by forest management. This work addresses this issue by comparing net rainfall (Pn) redistribution into different belowground hydrological processes (BHP) in two forest types after a thinning treatment: a holm oak coppice (HU) and a post-fire Aleppo pine regeneration (CAL). The relative contribution (RI) of forest structure, antecedent soil moisture (θ st ), rainfall and meteorological conditions on the BHP was assessed through boosted regression trees models. In both sites, the RI of the forest structure itself was limited (<10%). However, θ st , which clearly increased significantly with thinning, received an average RI of 29%. Surface and subsurface lateral flows showed values <1% of gross rainfall (Pg) in either site and were not significantly affected by thinning. On the other hand, soil moisture and drainage were affected by the thinning treatment, although with different extent depending on the site: in the drier site (CAL), the increased Pn in the thinning was mainly allocated into increased soil water content, with very limited improvement in drainage (<10 mm/year); in contrast, in the wetter continental site of HU, drainage to deeper soil layers was the most remarkable effect of thinning (50 mm/year higher than in control), given the higher θ st and hence the lower soil water storage available. Thinning also improved the response of BHP during drought, making these processes more elastic and less vulnerable to climatic extremes. The results presented here complement those previously reported on rainfall partitioning in these sites and all together provide a comprehensive understanding of the short-term effect (3–4 years) of water-oriented silviculture in Quercus ilex and Pinus halepensis low-biomass semiarid forests. Questions such as the long-term effects of thinning remain open for these ecosystems. © 2019 Elsevier B.V. ; This study is a component of research projects: HYDROSIL (CGL2011-28776-C02-02), SILWAMED (CGL2014-58127-C3-2), CEHYRFO-MED (CGL2017-86839-C3-2-R) funded by the Spanish Ministry of Science and Innovation (Spain), FEDER funds (EU) and LIFE17 CCA/ES/000063 RESILIENTFORESTS. The authors are grateful to the Valencia Regional Government (CMAAUV), VAERSA, ACCIONA, the "Sierra Calderona" Natural Park and the communal authority of Serra, for their support in allowing the use of the experimental forest and for their assistance in carrying out the fieldwork. A.J. Molina is beneficiary of a "Juan de la Cierva" post-doctoral fellowship funded by the Spanish Ministry of Economy and Competitiveness. Appendix A ; Peer reviewed

In Spain, Quercus open woodlands are animal ranching systems of organic production seriously threatened by the exotic pathogen Phytophthora cinnamomi. The root disease it causes kills thousands of oaks annually. Effective disease management needs to integrate different techniques, and the use of a resistance inducer such as fosetyl‐Al can play a key role, because the use of potassium phosphite is prohibited in Spain. In a woodland where the pathogen recently arrived, 60 holm oaks in three different defoliation classes (asymptomatic, slight and moderate defoliation) were selected for trunk injection with pressurised capsules containing 4% of commercial fosetyl‐Al or water (controls). Holm oaks were checked periodically for defoliation and presence of the pathogen in roots and rhizosphere soil. Three years after treatments, defoliation was significantly lower in oaks treated with fosetyl‐Al, which even increased canopy cover, in comparison with control oaks, independent of the initial defoliation class considered. Chlamydospore density in rhizosphere soil, as well as the presence of the pathogen into the roots, was not significantly influenced by fosetyl‐Al treatments, although a trend to a lower presence of P. cinnamomi in roots was observed in treated oaks at every soil inoculum density detected. This study has shown that fosetyl‐Al, a phosphonate registered as a fungicide in the European Union, provides protection to holm oaks against P. cinnamomi, even exhibiting a therapeutic effect on pre‐existing infections. Consequently, this effective measure should be considered as part of the integrated approach to control this highly destructive pathogen in holm oak woodlands.

There is growing consensus regarding the implementation of a new statistical framework for environmental-economic accounting to improve ecosystem related policies. As the standard System of National Accounts (SNA) fails to measure the economic contribution of ecosystems to the total income of individuals, governments recognize the need to expand the standard SNA through the ongoing System of Environmental Economic Accounting (SEEA). Based on the authors&rsquo ; own data, this study focuses on linking 15 economic activities and 12 ecosystem services for a holm oak (Quercus ilex L.) open woodlands (HOW) ecosystem type in Andalusia, Spain. We emphasize that overcoming the challenges of multiple use is preferable to measuring single ecosystem products for improving habitat conservation policies. The objectives of this paper are to measure and compare the environmental assets, ecosystem services, and incomes at basic and social prices by applying a refined version of the standard System of National Accounts (rSNA) and the authors&rsquo ; Agroforestry Accounting System (AAS), respectively, to HOW. Considering intermediate products and consumptions of HOW farmer and government activities, we find that the rSNA ecosystem services and environmental incomes at basic prices are 123.3 &euro ; /ha and &minus ; 28.0 &euro ; /ha, respectively, while those of the AAS at social prices are 442.2 &euro ; /ha and 250.8 &euro ; /ha. Given advances in non-market valuation techniques, we show that an expanded definition of economic activities can be applied to measure the contribution to total income of managed natural areas taking into account the multiple uses of the ecosystem type. However, HOW sustainability continues to be a challenging issue that requires ecological threshold indicators to be identified, not only because of the economic implications but also because they provide vital information on which to base policy implementation.

Forest decline and increasing tree mortality are of global concern and the identification of the causes is necessary to develop preventive measures. Global warming is an emerging factor responsible for the increasing tree mortality in drought-prone ecosystems. In the southwestern Iberian Peninsula, Mediterranean holm oak open woodlands currently undergo large-scale population-level tree die-off. In this region, temperature and aridity have increased during recent decades, but the possible role of climate change in the current oak mortality has not been investigated. To assess the role of climate change in oak die-off in managed open woodlands in southwestern Spain, we analyzed climate change-related signals in century-long tree ring chronologies of dead holm oaks. We examined the high/lowfrequency variability in growth and the relationship between growth and climate. Similar to other Mediterranean forests, growth was favored by precipitation from autumn of the year prior to ring formation to spring of the year of ring formation, whereas high temperatures during spring limited growth. Since the 1970s, the intensity of the high-frequency response to water availability increased simultaneously with temperature and aridity. The growth trends matched those of climatic changes. Growth suppressions occurred during droughts in the 1970s, 1980s and 1990s. Widespread stand-level, age-independent mortality occurred since 2005 and affected trees that can not be considered old for the species standards. The close relationship between growth and climate indicate that climate change strongly controlled the growth patterns. This suggests that harsher climatic conditions, especially increased aridity, affected the tree performance and could have played a significant role in the mortality process. Climate change may have exacerbated or predisposed trees to the impact of other factors (e.g. intense management and pathogens). These observations could suggest a similar future increase in oak mortality which may occur in more northern oak open woodlands if aridity further increases. ; This study was supported by the Department of Innovation, Science and Business of the Regional Government of Andalusia, Spain (project ref: P07RNM02688), the Spanish Ministry of Economy and Competitiveness, the European Union (FEDER funds), the Spanish National Agriculture Research Institute (project ref: RTA2013-00011-C02-02), and the International Campus of Excellence for Environment, Biodiversity and Global Change (CeiCambio). FN thanks E. Garriga (CIFOR-INIA) and D. Martin Perez for help with sample collection and processing. The authors acknowledge the Environmental Council in Huelva and the staff of the Department for Environment of the Government of Andalusia for their assistance in locating study sites. Ulf Buntgen (WSL, Birmensdorf, Switzerland) provided useful suggestions to improve a previous version of the manuscript.

Intro -- Contents -- 1 Oaks and People: A Long Journey Together -- Abstract -- Acknowledgements -- References -- 2 An Updated Infrageneric Classification of the Oaks: Review of Previous Taxonomic Schemes and Synthesis of Evolutionary Patterns -- Abstract -- 2.1 History of Classifications of Oaks -- 2.2 Change in Criteria for Classification -- 2.3 Changing from Morphology to Molecules -- 2.4 Revised Subgeneric and Sectional Classification of Oaks -- 2.4.1 Genus Quercus -- 2.4.2 Subgenus Quercus -- 2.4.2.1 Section Protobalanus (Intermediate Oaks) -- 2.4.2.2 Section Ponticae -- 2.4.2.3 Section Virentes -- 2.4.2.4 Section Quercus (White Oaks) -- 2.4.2.5 Section Lobatae (Red Oaks) -- 2.4.3 Subgenus Cerris -- 2.4.3.1 Section Cyclobalanopsis -- 2.4.3.2 Section Ilex -- 2.4.3.3 Section Cerris -- 2.5 Infrasectional Classification: The Big Challenge -- 2.6 Fossil Record -- 2.7 Conclusion and Outlook -- Acknowledgements -- Appendix 2.1 -- References -- 3 The Fossil History of Quercus -- Abstract -- 3.1 Introduction -- 3.2 Methodology -- 3.3 Results -- 3.3.1 The Cretaceous and Paleocene record -- 3.3.1.1 North America -- 3.3.1.2 Europe and Western Asia -- 3.3.1.3 Eastern Asia -- 3.3.2 North America: Eocene through Pliocene -- 3.3.2.1 The Eocene Records -- The Early to Middle Eocene -- The Late Eocene -- 3.3.2.2 The Oligocene -- 3.3.2.3 The Neogene -- Western North America -- Eastern and Central North America -- Central America -- 3.3.3 Europe and Western Asia: Eocene through Pliocene -- 3.3.3.1 The Eocene -- 3.3.3.2 The Oligocene -- 3.3.3.3 The Neogene -- 3.3.4 East Asia: Eocene through Pliocene -- 3.3.4.1 The Eocene -- 3.3.4.2 The Oligocene -- 3.3.4.3 The Neogene -- 3.4 Biogeographical and Palaeoclimatic Implications -- 3.5 Conclusions -- Acknowledgements -- References

9 páginas. 7 figuras ; A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests. ; This research was financially supported by MEC projects CLI97-0344, REN2003-04871, and CGL 2004-01402/BOS from the Spanish Government, and by the European project ALARM (Contract 506675, EU sixth framework programme). ; Peer reviewed

4 páginas, 2 figuras. ; Ultrasonic acoustic emissions were measured in Quercus ilex trees of a Mediterranean forest in Catalonia (NE Spain) each season from summer of 2004 to autumn of 2005. Acoustic emissions were maximum during hot and dry summer periods. Acoustic emissions started below 17% soil moisture, 0.85 RWC, and 2.5 MPa leaf water potential. They were negatively correlated with soil moisture and leaf water potential. The relationship between acoustic emissions and leaf water potential was the strongest, indicating that xylem tension is the most important factor inducing both cavitation (acoustic emissions) and a decrease in leaf water potential. Future increase of xylem cavitation derived from climate change may result in growth and survival limitations for this species in the drier southern limits of its current distribution. ; This research was financially supported by MEC REN2003-04871 and CGL 2004-01402/BOS grants from the Spanish Government, a Fundacio´n BBVA 2004 grant, a Catalan Government SGR2005-00312 grant, and the European project ALARM (Contract 506675, EU sixth framework programme). ; Peer reviewed

10 páginas, 4 figuras. ; Several volatile organic compounds (VOCs) have been reported as having a communication role between plants and also between plants and animals. We aimed to test whether methanol, a short-chain oxygenated VOC, could also have a signalling role between plants. We monitored photosynthetic performance and VOC exchange rates of Quercus ilex L. saplings before and after two different treatments: (a) clipping of some leaves to simulate an attack by herbivores and (b) fumigation with gaseous methanol for 5 h to simulate the amount of methanol a plant could receive from surrounding plants if those had been already attacked by herbivores. The clipping treatment enhanced the photosynthetic rates, the chlorophyll a to b ratio and the carotenoid to chlorophyll ratio of nonclipped leaves, suggesting an activation of plant protective metabolism. Also, a small but interesting systemic (in nonclipped leaves) increase in methanol emission rates was observed, which agrees with the possibility that methanol may act as a signalling cue. The methanol fumigation treatment induced an increase in the actual photochemical efficiency of PSII and also in the carotenoid to chlorophyll ratio. Methanol fumigation also promoted a 14% increase in the monoterpene emission rate, 1 day after the treatment, a similar response to the ones induced by other signalling VOCs. The enhanced monoterpene emissions could add to the blend of VOCs emitted after stress and be part of further signalling pathways, thus forwarding the message started by methanol. This study suggests that clipping and methanol fumigation at natural concentrations elicit significant neighbour plant physiological responses and further BVOC emissions. ; This study was supported by the Spanish Government grants CGL2006-04025/BOS, CGL2010-17172, and Consolider-Ingenio Montes CSD2008-00040, the CSIC grant PIF08-006-3 and the Catalan Government grant SGR 2009-458. Roger Seco gratefully acknowledges a FPI fellowship (BES-2005-6989) from MEC (Spanish Government). ; Peer reviewed