Analysis of adaptive responses of Pinus pinaster to changing environmental conditions in the Mediterranean region

Recent climate evolution studies highlight the progressive temperature increase and prevalence of seasonal drought, with specially incidence in the Mediterranean region. Although conifers are very important species regarding forest conservation, sustainability and productivity, given the large forest surface they cover in Spain and their active role in preventing soil erosion and desertification, we know little about the molecular mechanisms which control adaptation in this ancient taxonomic group

Trace Elements Distribution in Red Soils under Semiarid Mediterranean Environment

This study states the potential trace elements (TE’s) content of red soils located at the centre region of Spain, characterized by low rainfall and slight acidity over prolonged weathering periods. For this purpose, three soil profiles from a catena were described, sampled and analyzed. The most notable characteristics are the low organic matter content and the predominantly acidic pH. Illite and kaolinite are the predominant clay minerals. The fertility of the soils is sufficient to provide most of the nutrients required, with very suitable potassium levels. The geochemical characters of this soil are: only few elements remain almost invariable across the profiles and over time, however the majority of them were directly linked with the clay content. These soils are characterized by relatively low levels of some trace elements such as Sr (64.35 mg?kg–1), Ba (303.67 mg?kg–1) and Sc (13.14 mg?kg–1); high levels of other trace elements such as V (103.92 mg?kg–1), Cr (79.9 mg?kg–1), Cu (15.18 mg?kg–1), Hf (10.26 mg?kg–1), Ni (38 mg?kg–1) and Zr (337 mg?kg–1); while the levels for rare earth elements (REE’s) such as La (48.36 mg?kg–1), Ce (95.07 mg?kg–1), Th (13.33 mg?kg–1) and Nd (42.65 mg?kg–1) are significantly high. The distribution of mayor and trace elements was directly re- lated to weathering processes, parent material and anthropogenic activities.

Genes implicados en la respuesta molecular al estrés hídrico en Pinus pinaster Ait

La presente tesis doctoral se centra en el estudio de la respuesta molecular de las coníferas mediterráneas al estrés hídrico. Para ello se ha escogido como especie modelo Pinus pinaster Ait., la conífera más abundante en España, y que habita un amplio rango de situaciones ecológicas, especialmente en lo relativo a la disponibilidad de agua. En primer lugar, se ha aplicado un estrés hídrico controlado en cultivo hidropónico y se ha generando una genoteca sustractiva con objeto de identificar los genes inducidos por el estrés, analizando su expresión en raíces, tallos y acículas. A continuación, se ha analizado, la expresión de los genes anteriormente obtenidos así como de otros seleccionados de las bases de datos disponibles, durante una sequía prolongada en tierra, similar a las que las plantas deben afrontar en la naturaleza. Se ha utilizado en este caso, además de P. pinaster, P. pinea, otra conífera mediterránea adaptada a las sequías recurrentes. Este trabajo ha permitido identificar genes candidato expresionales, presumiblemente comunes en la respuesta molecular de las coníferas al déficit hídrico. Se han detectado diferencias notables en la expresión de determinados genes, que podrían ser los responsables de las diferencias exhibidas por ambas especies en el comportamiento frente a la sequía. Entre los genes identificados como inducidos por el estrés hídrico se encuentran varios miembros de la familia de las deshidrinas. Trabajos previos han utilizado deshidrinas como genes candidato; no obstante, la falta de especificidad de ciertos fragmentos y marcadores utilizados, debido a la complejidad estructural de esta familia, resta fiabilidad a algunos de los resultados publicados. Por este motivo, se ha estudiado en detalle esta familia en P. pinaster, se han identificado y caracterizado 8 miembros y se ha analizado su patrón de expresión frente a sequía. Este estudio ha permitido describir por primera vez unos segmentos conservados en la secuencia de aminoácidos de las deshidrinas de pináceas, cuya presencia y número de repeticiones parece estar relacionado con su especificidad. Por último, se han escogido tres genes implicados en distintas fases de la respuesta al estrés hídrico para su análisis exhaustivo: una deshidrina, una nodulina y un factor de transcripción tipo AP2. Se ha caracterizado su estructura exón/intrón y secuenciado su región promotora. Además, se han obtenido líneas transformadas que sobreexpresan estos genes tanto de forma heteróloga, en la especie modelo Arabidopsis thaliana, como en el propio P. pinaster. Este material facilitará la realización de futuros estudios sobre la función y el mecanismo de actuación de estos genes en la respuesta al estrés hídrico. ABSTRACT This thesis focuses in the study of the molecular response to water stress in Mediterranean conifers. For this purpose, P. pinaster was selected as model species. It’s the most abundant conifer in Spain, living in a wide range of ecological conditions, especially regarding water availability. First, we have applied a controlled polyethylene glycol-induced water stress in hydroponic culture and obtained a suppression subtractive hybridization (SSH) library, with the aim of identifying genes induced by water stress, analysing their expression in roots, stems and needles. We have then analysed the expression patterns of the identified genes, together with other genes selected from public databases. This study was conducted throughout a prolonged drought stress in soil, similar to the ones plants have to face in nature. In this case not only P. pinaster was analysed but also P. pinea, another Mediterranean conifer well adapted to recurrent droughts. This work has enabled us to identify of reliable candidate genes, presumably shared with other conifers in the response to water stress. We observed remarkable differences in the expression of some genes, which could be involved in the differential behaviour that these species show in the water stress response. Within the genes induced by water stress, several members of the dehydrin gene family were identified. Due to the structural complexity of the family, certain ambiguities and inconsistencies have been detected in previous works that have used dehydrins as candidate genes. For this reason, we have analysed thoroughly this gene family in P. pinaster, and have identified and characterized eight different members, whose expression patterns during drought have also been assessed. This study has allowed us to identify for the first time novel conserved segments in the amino acids sequences of Pinaceae. The presence and number of repetitions of these segments could be associated with the functional specificity of these proteins. Finally, three genes involved in different steps of the water stress response were selected for an exhaustive analysis: a dehydrin, a nodulin and an AP2 transcription factor. For all of them, the exon/intron structure was established and their promoter region was sequenced. Also, transformed lines were obtained both in Arabidopsis thaliana and in P. pinaster for the constitutive overexpression of these genes. This material will facilitate the development of further studies to investigate the function of these genes during the water stress response

Impact attribution models to evaluate drought effects: economic impacts, risk analyses and management strategies in Spain and Chile

La sequía es un fenómeno natural que se origina por el descenso de las precipitaciones con respecto a una media, y que resulta en la disponibilidad insuficiente de agua para alguna actividad. La creciente presión que se ha venido ejerciendo sobre los recursos hídricos ha hecho que los impactos de la sequía se hayan visto agravados a la vez que ha desencadenado situaciones de escasez de agua en muchas partes del planeta. Los países con clima mediterráneo son especialmente vulnerables a las sequías, y, su crecimiento económico dependiente del agua da lugar a impactos importantes. Para reducir los impactos de la sequía es necesaria una reducción de la vulnerabilidad a las sequías que viene dada por una gestión más eficiente y por una mejor preparación. Para ello es muy importante disponer de información acerca de los impactos y el alcance de este fenómeno natural. Esta investigación trata de abarcar el tema de los impactos de las sequías, de manera que plantea todos los tipos de impactos que pueden darse y además compara sus efectos en dos países (España y Chile). Para ello se proponen modelos de atribución de impactos que sean capaces de medir las pérdidas económicas causadas por la falta de agua. Los modelos propuestos tienen una base econométrica en la que se incluyen variables clave a la hora de evaluar los impactos como es una variable relacionada con la disponibilidad de agua, y otras de otra naturaleza para distinguir los efectos causados por otras fuentes de variación. Estos modelos se adaptan según la fase del estudio en la que nos encontremos. En primer lugar se miden los impactos directos sobre el regadío y se introduce en el modelo un factor de aleatoriedad para evaluar el riesgo económico de sequía. Esto se hace a dos niveles geográficos (provincial y de Unidad de Demanda Agraria) y además en el último se introduce no solo el riesgo de oferta sino también el riesgo de demanda de agua. La introducción de la perspectiva de riesgo en el modelo da lugar a una herramienta de gestión del riesgo económico que puede ser utilizada para estrategias de planificación. Más adelante una extensión del modelo econométrico se desarrolla para medir los impactos en el sector agrario (impactos directos sobre el regadío y el secano e impactos indirectos sobre la Agro Industria) para ello se adapta el modelo y se calculan elasticidades concatenadas entre la falta de agua y los impactos secundarios. Por último se plantea un modelo econométrico para el caso de estudio en Chile y se evalúa el impacto de las sequías debidas al fenómeno de La Niña. iv Los resultados en general muestran el valor que brinda el conocimiento más preciso acerca de los impactos, ya que en muchas ocasiones se tiende a sobreestimar los daños realmente producidos por la falta de agua. Los impactos indirectos de la sequía confirman su alcance a la vez que son amortiguados a medida que nos acercamos al ámbito macroeconómico. En el caso de Chile, su diferente gestión muestra el papel que juegan el fenómeno de El Niño y La Niña sobre los precios de los principales cultivos del país y sobre el crecimiento del sector. Para reducir las pérdidas y su alcance se deben plantear más medidas de mitigación que centren su esfuerzo en una gestión eficiente del recurso. Además la prevención debe jugar un papel muy importante para reducir los riesgos que pueden sufrirse ante situaciones de escasez. ABSTRACT Drought is a natural phenomenon that originates by the decrease in rainfall in comparison to the average, and that results in water shortages for some activities. The increasing pressure on water resources has augmented the impact of droughts just as water scarcity has become an additional problem in many parts of the planet. Countries with Mediterranean climate are especially vulnerable to drought, and its waterdependent economic growth leads to significant impacts. To reduce the negative impacts it is necessary to deal with drought vulnerability, and to achieve this objective a more efficient management is needed. The availability of information about the impacts and the scope of droughts become highly important. This research attempts to encompass the issue of drought impacts, and therefore it characterizes all impact types that may occur and also compares its effects in two different countries (Spain and Chile). Impact attribution models are proposed in order to measure the economic losses caused by the lack of water. The proposed models are based on econometric approaches and they include key variables for measuring the impacts. Variables related to water availability, crop prices or time trends are included to be able to distinguish the effects caused by any of the possible sources. These models are adapted for each of the parts of the study. First, the direct impacts on irrigation are measured and a source of variability is introduced into the model to assess the economic risk of drought. This is performed at two geographic levels provincial and Agricultural Demand Unit. In the latter, not only the supply risk is considered but also the water demand risk side. The introduction of the risk perspective into the model results in a risk management tool that can be used for planning strategies. Then an extension of the econometric model is developed to measure the impacts on the agricultural sector (direct impacts on irrigated and rainfed productions and indirect impacts on the Agri-food Industry). For this aim the model is adapted and concatenated elasticities between the lack of water and the impacts are estimated. Finally an econometric model is proposed for the Chilean case study to evaluate the impact of droughts, especially caused by El Niño Southern Oscillation. The overall results show the value of knowing better about the precise impacts that often tend to be overestimated. The models allow for measuring accurate impacts due to the lack of water. Indirect impacts of drought confirm their scope while they confirm also its dilution as we approach the macroeconomic variables. In the case of Chile, different management strategies of the country show the role of ENSO phenomena on main crop prices and on economic trends. More mitigation measures focused on efficient resource management are necessary to reduce drought losses. Besides prevention must play an important role to reduce the risks that may be suffered due to shortages.

Frequency and Intensity of drought events over Ebro River basin.

Lately, several researchers have pointed out that climate change is expected to increase temperatures and lower rainfall in Mediterranean regions, simultaneously increasing the intensity of extreme rainfall events. These changes could have consequences regarding rainfall regime, erosion, sediment transport and water quality, soil management, and new designs in diversion ditches. Climate change is expected to result in increasingly unpredictable and variable rainfall, in amount and timing, changing seasonal patterns and increasing the frequency of extreme weather events. Consequently, the evolution of frequency and intensity of drought periods is of most important as in agro-ecosystems many processes will be affected by them. Realising the complex and important consequences of an increasing frequency of extreme droughts at the Ebro River basin, our aim is to study the evolution of drought events at this site statistically, with emphasis on the occurrence and intensity of them. For this purpose, fourteen meteorological stations were selected based on the length of the rainfall series and the climatic classification to obtain a representative untreated dataset from the river basin. Daily rainfall series from 1957 to 2002 were obtained from each meteorological station and no-rain period frequency as the consecutive numbers of days were extracted. Based on this data, we study changes in the probability distribution in several sub-periods. Moreover we used the Standardized Precipitation Index (SPI) for identification of drought events in a year scale and then we use this index to fit log-linear models to the contingency tables between the SPI index and the sub-periods, this adjusted is carried out with the help of ANOVA inference.

Managing drought economic effects through insurance schemes based on local water availability.

In many arid or semi-arid Mediterranean regions, agriculture is dependent on irrigation. When hydrological drought phenomena occur, farmers suffer from water shortages, incurring important economic losses. Yet, there is not agricultural insurance available for lack of irrigation water. This work attempts to evaluate hydrological drought risk and its economic impact on crop production in order to provide the basis for the design of drought insurance for irrigated arable crops. With this objective a model that relates water availability with expected yields is developed. Crop water requirements are calculated from evapotranspiration, effective rainfall and soil water balance. FAO?s methodology and AquaCrop software have been used to establish the relationship between water allocations and crop yields. The analysis is applied to the irrigation zone ?Riegos de Bardenas?, which is located in the Ebro river basin, northeast Spain, to the main arable crops in the area. Results show the fair premiums of different hydrological drought insurance products. Whole-farm insurance or irrigation district insurance should be preferable to crop specific insurance due to the drought management strategies used by farmers.

Functional response of Ulmus minor Mill. to drought, flooding and dutch elm disease

Ulmus minor es una especie arbórea originaria de Europa cuyas poblaciones han sido diezmadas por el hongo patógeno causante de la enfermedad de la grafiosis. La conservación de los olmos exige plantearse su propagación a través de plantaciones y conocer mejor su ecología y biología. Ulmus minor es un árbol de ribera, pero frecuentemente se encuentra alejado del cauce de arroyos y ríos, donde la capa freática sufre fuertes oscilaciones. Por ello, nuestra hipótesis general es que esta especie es moderadamente resistente tanto a la inundación como a la sequía. El principal objetivo de esta tesis doctoral es entender desde un punto de vista funcional la respuesta de U. minor a la inundación, la sequía y la infección por O. novo-ulmi; los factores que posiblemente más influyen en la distribución actual de U. minor. Con este objetivo se persigue dar continuidad a los esfuerzos de conservación de esta especie que desde hace años se dedican en varios centros de investigación a nivel mundial, ya que, entender mejor los mecanismos que contribuyen a la resistencia de U. minor ante la inoculación con O. novo-ulmi y factores de estrés abiótico ayudará en la selección y propagación de genotipos resistentes a la grafiosis. Se han planteado tres experimentos en este sentido. Primero, se ha comparado la tolerancia de brinzales de U. minor y U. laevis – otro olmo ibérico – a una inmersión controlada con el fin de evaluar su tolerancia a la inundación y comprender los mecanismos de aclimatación. Segundo, se ha comparado la tolerancia de brinzales de U. minor y Quercus ilex – una especie típica de ambientes Mediterránea secos – a la falta de agua en el suelo con el fin de evaluar el grado de tolerancia y los mecanismos de aclimatación a la sequía. El hecho de comparar dos especies contrastadas responde al interés en entender mejor cuales son los procesos que conducen a la muerte de una planta en condiciones de sequía – asunto sobre el que hay una interesante discusión desde hace algunos años. En tercer lugar, con el fin de entender mejor la resistencia de algunos genotipos de U. minor a la grafiosis, se han estudiado las diferencias fisiológicas y químicas constitutivas e inducidas por O. novo-ulmi entre clones de U. minor seleccionados a priori por su variable grado de resistencia a esta enfermedad. En el primer experimento se observó que los brinzales de U. minor sobrevivieron 60 días inmersos en una piscina con agua no estancada hasta una altura de 2-3 cm por encima del cuello de la raíz. A los 60 días, los brinzales de U. laevis se sacaron de la piscina y, a lo largo de las siguientes semanas, fueron capaces de recuperar las funciones fisiológicas que habían sido alteradas anteriormente. La conductividad hidráulica de las raíces y la tasa de asimilación de CO2 neta disminuyeron en ambas especies. Por el contrario, la tasa de respiración de hojas, tallos y raíces aumentó en las primeras semanas de la inundación, posiblemente en relación al aumento de energía necesario para desarrollar mecanismos de aclimatación a la inundación, como la hipertrofia de las lenticelas que se observó en ambas especies. Por ello, el desequilibrio del balance de carbono de la planta podría ser un factor relevante en la mortalidad de las plantas ante inundaciones prolongadas. Las plantas de U. minor (cultivadas en envases de 16 litros a media sombra) sobrevivieron por un prolongado periodo de tiempo en verano sin riego; la mitad de las plantas murieron tras 90 días sin riego. El cierre de los estomas y la pérdida de hojas contribuyeron a ralentizar las pérdidas de agua y tolerar la sequía en U. minor. Las obvias diferencias en tolerancia a la sequía con respecto a Q. ilex se reflejaron en la distinta capacidad para ralentizar la aparición del estrés hídrico tras dejar de regar y para transportar agua en condiciones de elevada tensión en el xilema. Más relevante es que las plantas con evidentes síntomas de decaimiento previo a su muerte exhibieron pérdidas de conductividad hidráulica en las raíces del 80% en ambas especies, mientras que las reservas de carbohidratos apenas variaron y lo hicieron de forma desigual en ambas especies. Árboles de U. minor de 5 y 6 años de edad (plantados en eras con riego mantenido) exhibieron una respuesta a la inoculación con O. novo-ulmi consistente con ensayos previos de resistencia. La conductividad hidráulica del tallo, el potencial hídrico foliar y la tasa de asimilación de CO2 neta disminuyeron significativamente en relación a árboles inoculados con agua, pero solo en los clones susceptibles. Este hecho enlaza con el perfil químico “más defensivo” de los clones resistentes, es decir, con los mayores niveles de suberina, ácidos grasos y compuestos fenólicos en estos clones que en los susceptibles. Ello podría restringir la propagación del hongo en el árbol y preservar el comportamiento fisiológico de los clones resistentes al inocularlos con el patógeno. Los datos indican una respuesta fisiológica común de U. minor a la inundación, la sequía y la infección por O. novo-ulmi: pérdida de conductividad hidráulica, estrés hídrico y pérdida de ganancia neta de carbono. Pese a ello, U. minor desarrolla varios mecanismos que le confieren una capacidad moderada para vivir en suelos temporalmente anegados o secos. Por otro lado, el perfil químico es un factor relevante en la resistencia de ciertos genotipos a la grafiosis. Futuros estudios deberían examinar como este perfil químico y la resistencia a la grafiosis se ven alteradas por el estrés abiótico. ABSTRACT Ulmus minor is a native European elm species whose populations have been decimated by the Dutch elm disease (DED). An active conservation of this species requires large-scale plantations and a better understanding of its biology and ecology. U. minor generally grows close to water channels. However, of the Iberian riparian tree species, U. minor is the one that spread farther away from rivers and streams. For these reasons, we hypothesize that this species is moderately tolerant to both flooding and drought stresses. The main aim of the present PhD thesis is to better understand the functional response of U. minor to the abiotic stresses – flooding and drought – and the biotic stress – DED – that can be most influential on its distribution. The overarching goal is to aid in the conservation of this emblematic species through a better understanding of the mechanisms that contribute to resistance to abiotic and biotic stresses; an information that can help in the selection of resistant genotypes and their expansion in large-scale plantations. To this end, three experiments were set up. First, we compared the tolerance to experimental immersion between seedlings of U. minor and U. laevis – another European riparian elm species – in order to assess their degree of tolerance and understand the mechanisms of acclimation to this stress. Second, we investigated the tolerance to drought of U. minor seedlings in comparison with Quercus ilex (an oak species typical of dry Mediterranean habitats). Besides assessing and understanding U. minor tolerance to drought at the seedling stage, the aim was to shed light into the functional alterations that trigger drought-induced plant mortality – a matter of controversy in the last years. Third, we studied constitutive and induced physiological and biochemical differences among clones of variable DED resistance, before and following inoculation with Ophiostoma novo-ulmi. The goal is to shed light into the factors of DED resistance that is evident in some genotypes of U. minor, but not others. Potted seedlings of U. minor survived for 60 days immersed in a pool with running water to approximately 2-3 cm above the stem collar. By this time, U. minor seedlings died, whereas U. laevis seedlings moved out of the pool were able to recover most physiological functions that had been altered by flooding. For example, root hydraulic conductivity and leaf photosynthetic CO2 uptake decreased in both species; while respiration initially increased with flooding in leaves, stems and roots possibly to respond to energy demands associated to mechanisms of acclimation to soil oxygen deficiency; as example, a remarkable hypertrophy of lenticels was soon observed in flooded seedlings of both species. Therefore, the inability to maintain a positive carbon balance somehow compromises seedling survival under flooding, earlier in U. minor than U. laevis, partly explaining their differential habitats. Potted seedlings of U. minor survived for a remarkable long time without irrigation – half of plants dying only after 90 days of no irrigation in conditions of high vapour pressure deficit typical of summer. Some mechanisms that contributed to tolerate drought were leaf shedding and stomata closure, which reduced water loss and the risk of xylem cavitation. Obviously, U. minor was less tolerant to drought than Q. ilex, differences in drought tolerance resulting mostly from the distinct capacity to postpone water stress and conduct water under high xylem tension among species. More relevant was that plants of both species exhibited similar symptoms of root hydraulic failure (i.e. approximately 80% loss of hydraulic conductivity), but a slight and variable depletion of non-structural carbohydrate reserves preceding dieback. Five- and six-year-old trees of U. minor (planted in the field with supplementary watering) belonging to clones of contrasted susceptibility to DED exhibited a different physiological response to inoculation with O. novo-ulmi. Stem hydraulic conductivity, leaf water potential and photosynthetic CO2 uptake decreased significantly relative to control trees inoculated with water only in DED susceptible clones. This is consistent with the “more defensive” chemical profile observed in resistant clones, i.e. with higher levels of saturated hydrocarbons (suberin and fatty acids) and phenolic compounds than in susceptible clones. These compounds could restrict the spread of O. novo-ulmi and contribute to preserving the near-normal physiological function of resistant trees when exposed to the pathogen. These results evidence common physiological responses of U. minor to flooding, drought and pathogen infection leading to xylem water disruption, leaf water stress and reduced net carbon gain. Still, seedlings of U. minor develop various mechanisms of acclimation to abiotic stresses that can play a role in surviving moderate periods of flood and drought. The chemical profile appears to be an important factor for the resistance of some genotypes of U. minor to DED. How abiotic stresses such as flooding and drought affect the capacity of resistant U. minor clones to face O. novo-ulmi is a key question that must be contemplated in future research.

Compositional and Geochemical Signatures for the Sedimentary Evolution of the Middle Triassic–Lower Jurassic Continental Redbeds from Western-Central Mediterranean Alpine Chains

Compositional and chemical analyses suggest that Middle Triassic–Lower Liassic continental redbeds (in the internal domains of the Betic, Maghrebian, and Apenninic chains) can be considered a regional lithosome marking the Triassic-Jurassic rift-valley stage of Tethyan rifting, which led to the Pangaea breakup and subsequent development of a mosaic of plates and microplates. Sandstones are quartzose to quartzolithic and represent a provenance of continental block and recycled orogen, made up mainly of Paleozoic metasedimentary rocks similar to those underlying the redbeds. Mudrocks display K enrichments; intense paleoweathering under a hot, episodically humid climate with a prolonged dry season; and sediment recycling. Redbeds experienced temperatures in the range of 100°–160°C and lithostatic/tectonic loading of more than 4 km. These redbeds represent an important stratigraphic signature to reconstruct a continental block (Mesomediterranean Microplate) that separated different realms of the western Tethys from Middle-Late Jurassic to Miocene, when it was completely involved in Alpine orogenesis.

Plant regeneration functional groups modulate the response to fire of soil enzyme activities in a Mediterranean shrubland

Soil enzymes are critical to soil nutrient cycling function but knowledge on the factors that control their response to major disturbances such as wildfires remains very limited. We evaluated the effect of fire-related plant functional traits (resprouting and seeding) on the resistance and resilience to fire of two soil enzyme activities involved in phosphorus and carbon cycling (acid phosphatase and β-glucosidase) in a Mediterranean shrublands in SE Spain. Using experimental fires, we compared four types of shrubland microsites: SS (vegetation patches dominated by seeder species), RR (patches dominated by resprouter species), SR (patches co-dominated by seeder and resprouter species), and IP (shrub interpatches). We assessed pre- and post-fire activities of the target soil enzymes, available P, soil organic C, and plant cover dynamics over three years after the fire. Post-fire regeneration functional groups (resprouter, seeder) modulated both pre- and post-fire activity of acid phosphatase and β-glucosidase, with higher activity in RR and SR patches than in SS patches and IP. However, we found no major differences in enzyme resistance and resilience between microsite types, except for a trend towards less resilience in SS patches. Fire similarly reduced the activity of both enzymes. However, acid phosphatase and β-glucosidase showed contrasting post-fire dynamics. While β-glucosidase proved to be rather resilient to fire, fully recovering three years after fire, acid phosphatase showed no signs of recovery in that period. Overall, the results indicate a positive influence of resprouter species on soil enzyme activity that is very resistant to fire. Long-lasting decrease in acid phosphatase activity probably resulted from the combined effect of P availability and post-fire drought. Our results provide insights on how plant functional traits modulate soil biochemical and microbiological response to fire in Mediterranean fire-prone shrublands.

Drought and grazing combined: Contrasting shifts in plant interactions at species pair and community level

The combined effects of drought stress and grazing pressure on shaping plant–plant interactions are still poorly understood, while this combination is common in arid ecosystems. In this study we assessed the relative effect of grazing pressure and slope aspect (drought stress) on vegetation cover and soil functioning in semi-arid Mediterranean grassland–shrublands in southeastern Spain. Moreover, we linked these two stress factors to plant co-occurrence patterns at species-pair and community levels, by performing C-score analyses. Vegetation cover and soil functioning decreased with higher grazing pressure and more south-facing (drier) slopes. At the community level, plants at south-facing slopes were negatively associated at no grazing but positively associated at low grazing pressure and randomly associated at high grazing pressure. At north-facing slopes, grazing did not result in a shift in the direction of the association. In contrast, analysis of pairwise species co-occurrence patterns showed that the dominant species Stipa tenacissima and Anthyllis cytisoides shifted from excluding each other to co-occurring with increasing grazing pressure at north-facing slopes. Our findings highlight that for improved understanding of plant interactions along stress gradients, interactions between species pairs and interactions at the community level should be assessed, as these may reveal contrasting results.

Ecological restoration across the Mediterranean Basin as viewed by practitioners

Restoration efforts in the Mediterranean Basin have been changing from a silvicultural to an ecological restoration approach. Yet, to what extent the projects are guided by ecological restoration principles remains largely unknown. To analyse this issue, we built an on-line survey addressed to restoration practitioners. We analysed 36 restoration projects, mostly from drylands (86%). The projects used mainly soil from local sources. The need to comply with legislation was more important as a restoration motive for European Union (EU) than for non-EU countries, while public opinion and health had a greater importance in the latter. Non-EU countries relied more on non-native plant species than EU countries, thus deviating from ecological restoration guidelines. Nursery-grown plants used were mostly of local or regional provenance, whilst seeds were mostly of national provenance. Unexpected restoration results (e.g. inadequate biodiversity) were reported for 50% of the projects and restoration success was never evaluated in 22%. Long term evaluation (> 6 years) was only performed in 31% of cases, and based primarily on plant diversity and cover. The use of non-native species and species of exogenous provenances may: i) entail the loss of local genetic and functional trait diversity, critical to cope with drought, particularly under the predicted climate change scenarios, and ii) lead to unexpected competition with native species and/or negatively impact local biotic interactions. Absent or inappropriate monitoring may prevent the understanding of restoration trajectories, precluding adaptive management strategies, often crucial to create functional ecosystems able to provide ecosystem services. The overview of ecological restoration projects in the Mediterranean Basin revealed high variability among practices and highlighted the need for improved scientific assistance and information exchange, greater use of native species of local provenance, and more long-term monitoring and evaluation, including functional and ecosystem services' indicators, to improve and spread the practice of ecological restoration.

Sedimentology of core GeoTu_SL110 from the eastern Mediterranean Sea

Clay mineral assemblages in a sediment core from the distal Nile discharge plume off Israel have been used to reconstruct the late Quaternary Nile sediment discharge into the Eastern Mediterranean Sea (EMS). The record spans the last ca. 140 ka. Smectite abundances indicate the influence of the Blue Nile and Atbara that have their headwaters in the volcanic rocks of the Ethiopian highlands. Kaolinite abundances indicate the influence of wadis, which contribute periodically to the suspension load of the Nile. Due to the geographical position, the climate and the sedimentary framework of the EMS is controlled by two climate systems. The long-term climate regime was governed by the African monsoon that caused major humid periods with enhanced sediment discharge at 132 to <126 ka (AHP5), 116 to 99 ka (AHP4), and 89 to 77 ka (AHP3). They lasted much longer than the formation of the related sapropel layers S5 (>2 ka), S4 (3.5 ka) and S3 (5 ka). During the last glacial period (MIS 4-2) the long-term changes of the monsoonal system were superimposed by millennial-scale changes of an intensified mid-latitude glacial system. This climate regime caused short but pronounced drought periods in the Nile catchment, which are linked to Heinrich Events and alternate with more humid interstadials. The clay mineral record further implies that feedback mechanisms between vegetation cover and sediment discharge of the Nile are detectable but of minor importance for the sedimentary record in the southeastern Mediterranean Sea during the investigated African Humid Periods.

Available water modifications by topsoil treatments under mediterranean semiarid conditions: afforestation plan

During dry periods in the Mediterranean area, the lack of water entering the soil matrix reduces organic contribu- tions to the soil. These processes lead to reduced soil fertility and soil vegetation recovery which creates a positive feedback process that can lead to desertification. Restoration of native vegetation is the most effective way to regenerate soil health, and control runoff and sediment yield. In Mediterranean areas, after a forestry proposal, it is highly common to register a significant number of losses for the saplings that have been introduced due to the lack of rainfall. When no vegetation is established, organic amendments can be used to rapidly protect the soil surface against the erosive forces of rain and runoff. In this study we investigated the hydrological effects of five soil treatments in relation to the temporal vari- ability of the available water for plants. Five amendments were applied in an experimental set of plots: straw mulching; mulch with chipped branches of Aleppo Pine (Pinus halepensis L.); TerraCotten hydroabsobent polymers; sewage sludge; sheep manure and control. Plots were afforested following the same spatial pattern, and amendments were mixed with the soil at the rate 10 Mg ha-1. In control plots, during June, July, August and September, soils were registered below the wilting point, and therefore, in the area of water unusable by plants. These months were coinciding with the summer mediter- ranean drought. This fact justifies the high mortality found on plants after the seeding plan. Similarly, soils have never exceeded the field capacity value measured for control plots. Conversely, in the straw and pinus mulch, soils were above the wilting point during a longer time than in control plots. Thus, the soil moisture only has stayed below the 4.2 pF suction in July, July and August. Regarding the amount of water available was also higher, especially in the months of December, January and February. However, the field capacity value measured has not showed any differences regarding the control. For these treatments, the survival sapling rates measured were the highest. Sludge, manure and polymers showed a moisture retention capacity slightly more limited than straw and pinus mulch. Likewise, it has been found that the area of usable water by plants was also lower, especially during the months of January and February. This situation is especially sharpened in plots amended with manure. In this treatment, the upper part of the soil profile was below the wilting point for six months a year (from April to August). For this treatment, the survival sapling rates measured were the lowest. In conclusion, from a land management standpoint, the pinus and straw mulch treatments have been shown as effective methods reducing water stress for plants. In this research, mulching has been proved as a significant method to reduce the mortality sapling rates during the mediterranean summer drought.

Available water modifications by topsoil treatments under mediterranean semiarid conditions: afforestation plan

During dry periods in the Mediterranean area, the lack of water entering the soil matrix reduces organic contribu- tions to the soil. These processes lead to reduced soil fertility and soil vegetation recovery which creates a positive feedback process that can lead to desertification. Restoration of native vegetation is the most effective way to regenerate soil health, and control runoff and sediment yield. In Mediterranean areas, after a forestry proposal, it is highly common to register a significant number of losses for the saplings that have been introduced due to the lack of rainfall. When no vegetation is established, organic amendments can be used to rapidly protect the soil surface against the erosive forces of rain and runoff. In this study we investigated the hydrological effects of five soil treatments in relation to the temporal vari- ability of the available water for plants. Five amendments were applied in an experimental set of plots: straw mulching; mulch with chipped branches of Aleppo Pine (Pinus halepensis L.); TerraCotten hydroabsobent polymers; sewage sludge; sheep manure and control. Plots were afforested following the same spatial pattern, and amendments were mixed with the soil at the rate 10 Mg ha-1. In control plots, during June, July, August and September, soils were registered below the wilting point, and therefore, in the area of water unusable by plants. These months were coinciding with the summer mediter- ranean drought. This fact justifies the high mortality found on plants after the seeding plan. Similarly, soils have never exceeded the field capacity value measured for control plots. Conversely, in the straw and pinus mulch, soils were above the wilting point during a longer time than in control plots. Thus, the soil moisture only has stayed below the 4.2 pF suction in July, July and August. Regarding the amount of water available was also higher, especially in the months of December, January and February. However, the field capacity value measured has not showed any differences regarding the control. For these treatments, the survival sapling rates measured were the highest. Sludge, manure and polymers showed a moisture retention capacity slightly more limited than straw and pinus mulch. Likewise, it has been found that the area of usable water by plants was also lower, especially during the months of January and February. This situation is especially sharpened in plots amended with manure. In this treatment, the upper part of the soil profile was below the wilting point for six months a year (from April to August). For this treatment, the survival sapling rates measured were the lowest. In conclusion, from a land management standpoint, the pinus and straw mulch treatments have been shown as effective methods reducing water stress for plants. In this research, mulching has been proved as a significant method to reduce the mortality sapling rates during the mediterranean summer drought.

Adaptive strategies of two Portuguese grapevines varieties (Aragonês. syn. Tempranillo and Trincadeira) face drought: physiological and structural responses.

At Mediterranean regions and particularly in southern Portugal, it is imperative to identify grape varieties more adapted to warm and dry climates in order to overcome future climatic changes. Two Vitis vinifera genotypes, Aragonez (syn. Tempranillo) and Trincadeira, were selected to assess their physiological responses to soil water stress. Vines were subjected to four irrigation regimes: irrigated during all phenological cycle, non-irrigated during all phenological cycle, non irrigated until veraison, irrigated after veraison. Predawn leaf water potential was much higher in Trincadeira than Aragonez in non- irrigated plants. This result is in accordance with its higher stomatal control efficiency in this variety (Trincadeira). Photosynthetic capacity (Amax at saturating light intensity) decreased due to stomatal and biochemical limitations under water stress. However, recovery capacity of leaf water status after irrigation was faster in Trincadeira. Yield and yield x Brix increased when irrigation occurred after veraison, particularly in Trincadeira. These results show that Trincadeira presents a drought adaptation than Aragonez. Ratio of variable to maximum fluorescence Fv/Fm and total leaf chlorophyll related with leaf water potential for both species. Reflectance Normalized Difference Vegetation Index (NDVI705), Red Edge Inflexion Point Index and Photochemical Reflectance Index were related with irrigation treatment. Relative water content and specific leaf area were similar between varieties. In conclusion, we suggested that there is variation among the genotypes and the main physiological parameters for variety selection, for drought, were leaf water potential, stomatal conductance and reflectance indexes.