Pollen profile from sediment core Höllerer See

(expanded by Eberhard Grüger, Göttingen) The site "Höllerer See" is a lake in the northern foreland of the Alps, about 30 km north of the city of Salzburg/Austria, situated in the south-western part of Oberösterreich/Austria. A 2 m long piston core from this locality, consisting entirely of calcareous gyttja, was studied by pollen analysis. The three lowermost samples (1.98, 1.95 and 1.92 m) were deposited during the Preboreal when Pinus and Betula were still the dominating forest trees. High pollen values of thermophilous woody species (mainly Corylus and Quercus, but also Ulmus, Tilia, Fraxinus) prove the Boreal age of the next younger sample (1.91 m). The following two pollen spectra attest that Alnus (1.89 m) and - later (1.88 m) - Fagus had become important members of the local (Alnus) and the regional (Fagus) vegetation. From this level up to the top of the profile these two tree taxa contribute - together with Betula - always 50 to 80 % to the arboreal pollen sum. The upper 1.89 m of sediment of the Höllerer See core evidently date from the Subboreal and the Subatlantic. As Preboreal sediment was stated at the base of the profile it must be concluded that most of the Boreal and the Atlantic is - for whatever reason - not represented by sediment in this core. As no radiocarbon dates are available age estimates of the distinguished pollen zones can be achieved only by correlating major changes of the former vegetation with historical events which probably influenced the then contemporary vegetation. The pollen grains of the Triticum and Hordeum type found in samples of zone 2.1 might indicate the growing of cereals in the region during the Late Bronze Age. The first pollen grains of Secale date from the boundary Hallstatt/Latène Age (zone 2.2). The cereal curves become continuous in Bavarian times (Bajuwarenzeit, Middle Ages, zone 3.3). The Plantago laceolata curve, continuous since 1.7 m depth (zone 2.1), points to animal breeding since the Early Subatlantic (Hallstattzeit). This curve reaches its absolute maximum in Roman time (zone 3.1). Roman time forest clearance caused a drastic decrease of tree pollen curves (start of zone 3.1). Values of anthropogenic indicators as high as in zone 3.1 are found again - after a distinct decrease in zone 3.2 - not till the Bavarians settled in the region (6th century). Maximal Fagus values and the simultaneous total lack of anthropogenic indicators mark the Migration Period (zone 3.2). The Younger Subatlantic (zone 4) is characterized by a decrease of deciduous forests due to medieval forest clearance. At the same time the conifers Pinus and Picea gained in importance. The lake was probably used for retting hemp in Medieval times. The distinction of the pollen grains of Cannabis and Humulus might not be certain in all cases. It is known that hemp as well as hop was cultivated in the study area. Markers were added to the samples at the beginning of pollen preparation (13500 Lycopodium spores, sample volume 0.5 cm**3) and counted together with the pollen grains. Therefore pollen concentrations can be calculated: Concentration = C * F / V (with C = number of grains of a particular pollen type, V = volume of the untreated pollen sample, F = marker added/marker counted). F ranges from 39 to 1688. Factors that large are not suited to produce reliably interpretable pollen concentrations. Consequently no use was made of the pollen concentrations in this thesis, although a concentration diagram is added.

Results of n-alkane biomarker analyses and paleoclimate reconstruction

The current study presents quantitative reconstructions of tree cover, annual precipitation and mean July temperature derived from the pollen record from Lake Billyakh (65°17'N, 126°47'E, 340 m above sea level) spanning the last ca. 50 kyr. The reconstruction of tree cover suggests presence of woody plants through the entire analyzed time interval, although trees played only a minor role in the vegetation around Lake Billyakh prior to 14 kyr BP (<5%). This result corroborates low percentages of tree pollen and low scores of the cold deciduous forest biome in the PG1755 record from Lake Billyakh. The reconstructed values of the mean temperature of the warmest month ~8-10 °C do not support larch forest or woodland around Lake Billyakh during the coldest phase of the last glacial between ~32 and ~15 kyr BP. However, modern cases from northern Siberia, ca. 750 km north of Lake Billyakh, demonstrate that individual larch plants can grow within shrub and grass tundra landscape in very low mean July temperatures of about 8 °C. This makes plausible our hypothesis that the western and southern foreland of the Verkhoyansk Mountains could provide enough moist and warm microhabitats and allow individual larch specimens to survive climatic extremes of the last glacial. Reconstructed mean values of precipitation are about 270 mm/yr during the last glacial interval. This value is almost 100 mm higher than modern averages reported for the extreme-continental north-eastern Siberia east of Lake Billyakh, where larch-dominated cold deciduous forest grows at present. This suggests that last glacial environments around Lake Billyakh were never too dry for larch to grow and that the summer warmth was the main factor, which limited tree growth during the last glacial interval. The n-alkane analysis of the Siberian plants presented in this study demonstrates rather complex alkane distribution patterns, which challenge the interpretation of the fossil records. In particular, extremely low n-alkane concentrations in the leaves of local coniferous trees and shrubs suggest that their contribution to the litter and therefore to the fossil lake sediments might be not high enough for tracing the Quaternary history of the needleleaved taxa using the n-alkane biomarker method.

Historia holocena de los pinares montanos de los sistemas Central y Cantábrico: Un objetivo con dos enfoques, biogeográfico y dendroecológico

Los estudios paleoecológicos holocenos basados en macro- y megafósiles encierran un gran valor debido a que su información tiene generalmente carácter local, su origen es conocido, pueden ser datados directamente mediante el método radiocarbónico, y pueden identificarse a un nivel taxonómicamente preciso. Sin embargo son pocas las áreas del Sur de Europa en las que sea conocida una alta densidad de yacimientos con restos leñosos de gran tamaño. En esta tesis, se presentan datos de 53 yacimientos de la sierra de Gredos y de la cordillera Cantábrica (Península Ibérica). Los restos fueron hallados en ambientes variados, como zonas higroturbosas, turberas erosionadas o lagos, y fueron identificadas mediante el estudio de la anatomía de la madera o mediante rasgos morfológicos. En la sierra de Gredos, la evidencia paleobotánica indica la existencia de un panorama relativamente estable a lo largo del Holoceno medio y principio del Holoceno final y sugiere la persistencia, a lo largo de milenios, de un piso de pinares ampliamente distribuído en cotas altas de la sierra. La información obtenida de piñas y frutos mejoran la información taxonómica disponible y revelan la existencia tanto de Pinus sylvestris como de Pinus nigra en estas sierras durante el Holoceno. La datación radiocarbónica, medición de anillos de crecimiento y sincronización preliminar de 26 secciones de troncos de subfósiles demuestran el potencial de este material de las montañas de Iberia central en la obtención de cronologías holocenas de pino. En la cordillera Cantábrica, los datos aportan información espacialmente precisa de distribuciones de ciertas especies arbóreas durante el Holoceno. En las zonas centrales de la cordillera, han sido hallados fundamentalmente restos de pino, mientras que en las zonas más occidentales los pinos estás ausentes y los restos encontrados corresponden a otras especies de caducifolios (Betula, Salix, Quercus) y arbustos (Erica, Fabaceae) Esta información paleobiogeográfica constrasta con la distribución natural actual de Pinus sylvestris y Pinus nigra en el área de estudio. En la sierra de Gredos, la naturalidad de las escasos rodales de pinos que aún persisten ha sido discutida, mientras que en la cordillera Cantábrica, la única especie del grupo que persiste es P. sylvestris y está localizada en unos pocos relictos. El clima pudo haber jugado un papel importante en una primera fase de declive de los pinares durante el Holoceno inicial, mostrado en numerosos registros polínicos de manera casi sincrónica y asociada a una expansión de frondosas. Sin embargo la información histórica disponible y la comparación entre las áreas de distribución de los pinares en el presente, modelizada y en momentos anteriores a la la generalización de la presión antrópica sugiere que durante los últimos dos milenios, la actividad humana ha sido responsable de la desaparición de estas especies como árboles naturales en áreas extensas. ABSTRACTMacro- and megafossil studies provide information of great value in palaeoecology because such evidence is spatially precise, directly radiocarbon dated and usually taxon-specific. However, few areas of southern Europe have a high density of sites with Holocene woody remains. Here, local data from 53 sites in the Gredos Mountains and the Cantabrian Range (Iberian Peninsula) is presented. Woody remains were recovered from mires, eroded peat bogs and lakes and were identified by their wood anatomy or morphological traits. In the Gredos Mountains, palaeobotanical evidence portrays a relatively stable picture of tree distribution over the mid- and beggining of the late-Holocene, and suggests the persistence of a widespread belt of pinewoods. Cones and fruits enlarge the taxonomic information available and reveal that both Pinus sylvestris and Pinus nigra were present locally during the Holocene. Radiocarbon dating, tree ring measurement and preliminary cross-dating of 26 pine sub-fossil logs demonstrate the potential of obtaining a long pine chronology from subfossil wood from the mountains of Central Iberia. In the Cantabrian Range the data provide spatially precise evidence of tree distribution in the region during the Holocene. Pines were mostly identified in the central areas, whereas at the western edge no pine evidence was detected and deciduous trees (Betula, Salix, Quercus) and shrubs (Erica, Fabaceae) were identified. This palaeoecological information contrasts with the current natural distribution ranges of P. sylvestris and P. nigra in the study area. In the Gredos Mountains, the naturality of the few pine stands currently growing has been heavily debated. In the Cantabrian Range P. sylvestris is the only pine species that is today present, and its natural presence is now limited to a few enclaves. Climate may have played a key role in the early-Holocene, as pollen archives document a pine demise that is synchronous with the spread of broadleaved taxa. However, available historical data and the comparison of the reconstructed distribution of pinewoods before extensive human forest disturbance with both present and modelled distributions suggests that during the last two millennia, anthropogenic activity may have removed these species as native trees from a large territory.

Dinámica vegetal desde el tardiglaciar en la cuenca del Duero: interacciones con clima, fuego e impacto humano

La cuenca del Duero constituye un vasto territorio ibérico cuyo paisaje se encuentra actualmente muy alterado por la mano del hombre y es prácticamente imposible localizar alguna manifestación de su cubierta vegetal natural. Aunque la historia de la vegetación en los sectores central y oriental es relativamente bien conocida, en su mitad occidental los registros paleoecológicos estudiados hasta la fecha son prácticamente inexistentes. Esto hace que se desconozca la respuesta de la vegetación a las diferentes oscilaciones climáticas que se han producido desde el Último Máximo Glaciar, cuál fue el impacto de las diferentes culturas sobre el medio, cuándo se produjo una alteración profunda de la vegetación natural y cuál ha sido la historia de los incendios. Este último aspecto, el papel e importancia de los incendios, reviste un especial interés en la península Ibérica dada su situación geográfica y climática dentro de la cuenca Mediterránea, donde el fuego es un factor ecológico de primer nivel. Las distintas técnicas paleoecológicas son las más adecuadas para abordar todas estas preguntas. De este modo, los avatares de la vegetación a través del tiempo se han reconstruido mediante el análisis polínico y de macrofósiles, el impacto humano se ha trazado utilizando indicadores polínicos ligados a actividades antrópicas y esporas de hongos coprófilos, estudiándose los incendios a partir del registro de partículas microscópicas de carbón. La alta resolución temporal y taxonómica alcanzada en estos análisis, así como la amplia superficie abarcada con los yacimientos estudiados, hacen que la información obtenida sea altamente detallada y complete el conocimiento que se tiene sobre la cuenca del Duero. El Tardiglaciar se registra por primera vez en la Meseta Norte Ibérica en la secuencia de Ayoó de Vidriales, donde una vegetación esteparia prácticamente desarbolada domina durante los periodos fríos. Durante el interestadial Bølling/Allerød la expansión forestal (pinos, abedules) fue muy leve y tardía y fue interrumpida bruscamente por el Dryas Reciente. Al final del Dryas Reciente o al inicio del Holoceno se produjo una acusada y rápida expansión de los bosques. Esta dinámica sugiere que no hubo refugios glaciares importantes en esta zona de la Meseta durante el “Mystery Interval”, aparentemente el periodo más frío y seco. Los incendios fueron muy escasos, aumentando de forma muy brusca al inicio del Holoceno por el aumento de biomasa y las condiciones relativamente cálidas y secas. A partir de los registros de Ayoó y El Maíllo se consolida la importancia del gradiente oceanicidad-continentalidad en el Sistema Central y la Meseta Norte, que se manifiesta principalmente en la dominancia más prolongada de los pinares hacia el interior. Además, otra de las principales contribuciones de la presente tesis es proporcionar evidencia sobre la sucesión de distintos tipos de bosques en el noroeste de la Meseta, precisando el marco temporal en el que suceden. Así, se ha constatado que hubo un máximo desarrollo del bosque caducifolio durante el Holoceno medio en Ayoó de Vidriales y una baja incidencia del fuego que sugieren que el clima fue más húmedo que en el Holoceno inicial. El estudio de macrofósiles leñosos ha permitido detectar procesos que con el análisis polínico habrían pasado desapercibidos, como la persistencia hasta el Holoceno final de Pinus sylvestris en la sierra del Teleno y la sustitución de P. sylvestris por P. pinaster en la sierra de Francia durante el Holoceno inicial. También el estudio de los carbones procedentes de los arenales de Tierra de Pinares han proporcionado la prueba definitiva de la naturalidad de los pinares de P. pinaster. El impacto humano se detecta temprano en las secuencias del oeste de la cuenca del Duero, durante el Neolítico, aunque ha sido mucho más acusado desde la Edad del Hierro (ca 2700-2500 años cal BP). Para la detección del impacto humano temprano ha sido clave el análisis de esporas de hongos coprófilos, cuyo análisis fue incorporado en la secuencia de Ayoó de Vidriales. Una de sus principales consecuencias fue el establecimiento de comunidades de matorral (brezales, piornales) sobre amplias extensiones del occidente de la cuenca del Duero, vinculado al recrudecimiento de los regímenes de incendios. A pesar de que los incendios han sido ecológicamente importantes desde el inicio del Holoceno, los cambios introducidos por el hombre en sus regímenes sobrepasaron la resiliencia de los bosques originales, lo cual condujo a su sustitución sostenida por matorrales. ABSTRACT The Duero Basin constitutes a vast Iberian territory which is currently strongly disturbed due to human activities, so it is very difficult to find any remnant of the natural vegetation. Vegetation history for the eastern and western sectors of the Basin is relatively well-known but, in contrast, there is an almost complete lack of palaeoecological record in the western area. Consequently, there exists a profound ignorance about vegetation responses to the climatic oscillations occurred since the Last Glacial Maximum, the environmental impact of the different cultures, when a severe disturbance of the natural vegetation took place and fire history. The last question, the role and importance of fire, has a special interest in the Iberian Peninsula due to its geographic and climatic framework, the Mediterranean Basin, where fire is a major ecological factor. The diverse palaeoecological techmiques are the most suitable tools to address all these questions. Thus, vegetation shifts through time have been reconstructed using pollen and macrofossil analyses, human impact has been tracked by means of anthropogenic pollen indicators and dung fungal spores, while fire history has been studied from the quantification of microscopic charcoal particles. The high taxonomic and time resolution attained along with the extensive surface covered by the studied sites provide detailed information very useful to complete the knowledge on landscape dynamics in the Duero Basin. The Lateglacial is recorded for the first time in the Northern Iberian Plateau in the sequence from Ayoó de Vidriales, showing that almost treeless steppic vegetation dominated during the cold periods. Tree expansion (pines, birches) was late and slight during the Bølling/Allerød interstadial and was sharply interrupted by the Younger Dryas (YD) climatic reversal. By the end of the YD or the onset of the Holocene, a rapid forest expansion occurred. This forest dynamics suggests an absence of important glacial refugia for trees in this area of the Plateau during the Mystery Interval, apparently the coldest and driest period. Fires were fairly rare, increasing abruptly at the beginning of the Holocene due to the relatively warm and dry climate and the accumulation of biomass. The records from Ayoó and El Maíllo reinforce the role of the oceanicity-continentality gradient in the vegetation history of the Iberian Central Range and the Iberian Northern Plateau, reflected mainly in the longer dominance of pine forests towards inland areas. Further, another important contribution of this PhD Thesis is providing evidence on the succession of different forest types in the northestern fringe of the Plateau, specifying the chronological framework. A maximum of deciduous forest development and low fire activity have been detected in Ayoó de Vidriales during the mid-Holocene, suggesting that climate was wetter than in the early Holocene. The study of woody macrofossils has allowed detecting processes which would have remained unnoticed using pollen analysis alone, such as the persistence of Pinus sylvestris until the late Holocene in the Teleno Mountains and the early Holocene replacement of P. sylvestris with P. pinaster in the sierra de Francia range. The study of macroscopic charcoal fragments from palaeosols of the Tierra de Pinares has also provided the definitive proof of naturalness for the P. pinaster stands gorwing over this area at present. Early human impact, during the Neolithic, has been detected in the sequences from the western sector of the Duero Basin, although human disturbance has been more severe from the Iron Age onwards (ca 2700-2500 cal yr BP). The analysis of coprophilous fungi incorporated in the sequence of Ayoó de Vidriales has played a key role in recognizing that early human impact. One of the main consequences of human disturbance was the establishment of shrubland communities (heaths, brooms) over huge areas of the western Duero Basin, linked to severe and/or frequent fires. Despite fires has been ecologically important since the onset of the Holocene, human-induced changes in fire regimes have exceeded the resilience of original forests leading to a sustained replacement with shrublands.

Characterization of a glycosylase family gene specifically expressed during winter dormancy in woody plants

Winter dormancy is the strategy used by perennial plants to survive the harsh conditions of winter in temperate and cold regions. This complex mechanism is characterized by cessation of the meristems activity, which is accompanied by the budset, the acquisition of a high tolerance to the cold temperatures and, in the case of deciduous trees, by the senescence and leaf abscission. In long-lived forest species, the length of the dormancy period limits the growing season, affecting wood production and quality. A Suppression Subtractive Hybridization (SSH) enriched in genes overexpressed during the process of winter dormancy in chesnut stems identified a DNA glycosylase gene. In order to study its role in the establishment and maintenance of the winter dormancy, a molecular characterization and seasonal expression were performed. Furthermore, we have obtained poplar transgenic plantlets overexpressing the chesnut gene.

Urban trees and atmospheric pollutants in big cities: Effects in Madrid

Una investigación sobre la mejora de la contaminación del aire (CA) por medio de arbolado urbano se realizó en Madrid, una ciudad con casi 4 M de habitantes, 2,8 M de vehículos y casi 3 M de árboles de mantenimiento público. La mayoría de los árboles estaban en dos bosques periurbanos. Los 650.000 restantes era pies de alineación y parques. Los taxones estudiados fueron Platanus orientalis (97.205 árboles), Ulmus sp. (70.557), Pinus pinea (49.038), Aesculus hippocastanum (22.266), Cedrus sp. (13.678) y Quercus ilex (1.650), de calles y parques. Muestras foliares se analizaron en diferentes épocas del año, así como datos de contaminación por PM10 de 28 estaciones de medición de la contaminación durante 30 años, y también la intensidad del tráfico (IMD) en 2.660 calles. La acumulación de metales pesados (MP) sobre hojas y dentro de estas se estimó en relación con la CA y del suelo y la IMD del tráfico. La concentración media de Ba, Cd, Cr, Cu, Mn, Ni, Pb y Zn en suelo (materia seca) alcanzó: 489,5, 0,7, 49,4, 60,9, 460,9, 12,8, 155,9 y 190,3 mg kg-1 respectivamente. Los árboles urbanos, particularmente coníferas (debido a la mayor CA en invierno) contribuyen significativamente a mejorar la CA sobre todo en calles con alta IMD. La capacidad de las seis sp. para capturar partículas de polvo en su superficies foliares está relacionada con la IMD del tráfico y se estimó en 16,8 kg/año de MP tóxicos. Pb y Zn resultaron ser buenos marcadores antrópicos en la ciudad en relación con el tráfico, que fue la principal fuente de contaminación en los árboles y suelos de Madrid. Las especies de árboles variaron en función de su capacidad para capturar partículas (dependiendo de las propiedades de sus superficies foliares) y acumular los MP absorbidos de los suelos. Las concentraciones foliares de Pb y Zn estuvieron por encima de los límites establecidos en diferentes sitios de la ciudad. La microlocalización de Zn mediante microscópico mostró la translocación al xilema y floema. Se detectaron puntos de contaminación puntual de Cu and Cr en antiguos polígonos industriales y la distribución espacial de los MP en los suelos de Madrid mostró que en incluso en zonas interiores del El Retiro había ciertos niveles elevados de [Pb] en suelo, tal vez por el emplazamiento la Real Fábrica de Porcelana en la misma zona hace 200 años. Distintas áreas del centro de la ciudad también alcanzaron niveles altos de [Pb] en suelo. Según los resultados, el empleo de una combinación de Pinus pinea con un estrato intermedio de Ulmus sp. y Cedrus sp. puede ser la mejor recomendación como filtro verde eficiente. El efecto del ozono (O3) sobre el arbolado en Madrid fue también objeto de este estudio. A pesar de la reducción de precursores aplicada en muchos países industrializados, O3 sigue siendo la principal causa de CA en el hemisferio norte, con el aumento de [O3] de fondo. Las mayores [O3] se alcanzaron en regiones mediterráneas, donde el efecto sobre la vegetación natural es compensado por el xeromorfismo y la baja conductancia estomática en respuesta los episodios de sequía estival característicos de este clima. Durante una campaña de monitoreo, se identificaron daños abióticos en hojas de encina parecidos a los de O3 que estaban plantadas en una franja de césped con riego del centro de Madrid. Dada la poca evidencia disponible de los síntomas de O3 en frondosas perennifolias, se hizo un estudio que trató de 1) confirman el diagnóstico de daño de O3, 2) investigar el grado de los síntomas en encinas y 3) analizar los factores ambientales que contribuyeron a los daños por O3, en particular en lo relacionado con el riego. Se analizaron los marcadores macro y micromorfológicos de estrés por O3, utilizando las mencionadas encinas a modo de parcela experimental. Los síntomas consistieron en punteado intercostal del haz, que aumentó con la edad. Además de un punteado subyacente, donde las células superiores del mesófilo mostraron reacciones características de daños por O3. Las células próximas a las zonas dañadas, presentaron marcadores adicionales de estrés oxidativo. Estos marcadores morfológicos y micromorfológicos de estrés por O3 fueron similares a otras frondosas caducifolias con daños por O3. Sin embargo, en nuestro caso el punteado fue evidente con AOT40 de 21 ppm•h, asociada a riego. Análisis posteriores mostraron que los árboles con riego aumentaron su conductancia estomática, con aumento de senescencia, manteniéndose sin cambios sus características xeromórficas foliares. Estos hallazgos ponen de relieve el papel primordial de la disponibilidad de agua frente a las características xeromórficas a la hora de manifestarse los síntomas en las células por daños de O3 en encina. ABSTRACT Research about air pollution mitigation by urban trees was conducted in Madrid (Spain), a southern European city with almost 4 M inhabitants, 2.8 M daily vehicles and 3 M trees under public maintenance. Most trees were located in two urban forests, while 650'000 trees along urban streets and in parks. The urban taxa included Platanus orientalis (97'205 trees), Ulmus sp. (70’557), Pinus pinea (49'038), Aesculus hippocastanum (22’266), Cedrus sp. (13'678 and Quercus ilex (1'650) along streets and parks. Leave samples were analysed sequentially in different seasons, PM10 data from 28 air monitoring stations during 30 years and traffic density estimated from 2’660 streets. Heavy metal (HM) accumulation on the leaf surface and within leaves was estimated per tree related to air and soil pollution, and traffic intensity. Mean concentration of Ba, Cd, Cr, Cu, Mn, Ni, Pb and Zn in topsoil samples (dry mass) amounted in Madrid: 489.5, 0.7, 49.4, 60.9, 460.9, 12.8, 155.9 and 190.3 mg kg-1 respectively. Urban trees, particularly conifers (due to higher pollution in winter) contributed significantly to alleviate air pollution especially near to high ADT roads. The capacity of the six urban street trees species to capture air-born dust on the foliage surface as related to traffic intensity was estimated to 16.8 kg of noxious metals from exhausts per year. Pb and Zn pointed to be tracers of anthropic activity in the city with vehicle traffic as the main source of diffuse pollution on trees and soils. Tree species differed by their capacity to capture air-borne dust (by different leaf surface properties) and to allocate HM from soils. Pb and Zn concentrations in the foliage were above limits in different urban sites and microscopic Zn revelation showed translocation in xylem and phloem tissue. Punctual contamination in soils by Cu and Cr was identified in former industrial areas and spatial trace element mapping showed for central Retiro Park certain high values of [Pb] in soils even related to a Royal pottery 200 years ago. Different areas in the city centre also reached high levels [Pb] in soils. According to the results, a combination of Pinus pinea with understorey Ulmus sp. and Cedrus sp. layers can be recommended for the best air filter efficiency. The effects of ozone (O3) on trees in different areas of Madrid were also part of this study. Despite abatement programs of precursors implemented in many industrialized countries, ozone remained the main air pollutant throughout the northern hemisphere with background [O3] increasing. Some of the highest ozone concentrations were measured in regions with a Mediterranean climate but the effect on the natural vegetation is alleviated by low stomatal uptake and frequent leaf xeromorphy in response to summer drought episodes characteristic of this climate. During a bioindication survey, abiotic O3-like injury was identified in foliage. Trees were growing on an irrigated lawn strip in the centre of Madrid. Given the little structural evidence available for O3 symptoms in broadleaved evergreen species, a study was undertaken in 2007 with the following objectives 1) confirm the diagnosis, 2) investigate the extent of symptoms in holm oaks growing in Madrid and 3) analyse the environmental factors contributing to O3 injury, particularly, the site water supply. Therefore, macro- and micromorphological markers of O3 stress were analysed, using the aforementioned lawn strip as an intensive study site. Symptoms consisted of adaxial and intercostal stippling increasing with leaf age. Underlying stippling, cells in the upper mesophyll showed HR-like reactions typical of ozone stress. The surrounding cells showed further oxidative stress markers. These morphological and micromorphological markers of ozone stress were similar to those recorded in deciduous broadleaved species. However, stippling became obvious already at an AOT40 of 21 ppm•h and was primarily found at irrigated sites. Subsequent analyses showed that irrigated trees had their stomatal conductance increased and leaf life-span reduced whereas their leaf xeromorphy remained unchanged. These findings suggest a central role of water availability versus leaf xeromorphy for ozone symptom expression by cell injury in holm oak.

The late Miocene macroflora of the La Cerdaña Basin, Eastern Pirenees, Spain. Towards a Synthesis

The fossil plant-bearing beds of the Tortonian (late Miocene) intramontane basin of La Cerdanya (Eastern Pyrenees, Catalonia, Spain) have been investigated for more than a century, and 165 species from 12 outcrops have been described in previous publications. The sediments with rich plant fossil assemblages, which correspond to lacustrine diatomitic deposits, contain large numbers of plant remains, mainly leaf compressions and impressions. These assemblages are well preserved, a consequence of the rapid accumulation of plant remains in the sediments of the basin's ancient lake, and the often close proximity of its shores to wetland and upland vegetation. This paper provides a comprehensive taxonomic and nomenclatural review of the historic and new collections of late Miocene macroflora for the La Cerdanya Basin. Examination of the newer materials allowed emendments to be made to the diagnoses ofAbies saportana, Acer pyrenakum,Alnus occidentalis, Quercus hispanka and Tilia vidali provided by REROLLE for the basin at the end of the 19th century. In addition, 24 species of vascular plants are identified for the basin for the first time, including one horsetail, three conifers, 19 arboreal or bushy dicotyledonous angiosperms, and one monocotyledonous angiosperm. Indeed, this is the first time that Cedrela helkonia (UNGER) KNOBLOCH, Decodon sp„ Hedera cf multinervis KOLAKOVSKII, Mahonia cf pseudosimplex KVACEK & WALTHER, Smilax cf. aspera L. vm.fossilis and Ulmus cf. plurinervia UNGER have been recorded anywhere in the Iberian Peninsula. The La Cerdanya Basin plant assemblages of the late Miocene mainly consisted of conifers and deciduous broadleaved taxa of Arctotertiary origin; evergreen Palaeotropical elements were less well represented. This flora is similar to those recorded at coeval sites in northern Greece, northern Italy and central and eastern France. Within the Iberian Peninsula, the late Miocene macroflora reported for the nearby Seu d'Urgell Basin is the most similar.

Vegetation and fire history since the last glacial maximum in an inland area of the western Mediterranean Basin (Northern Iberian Plateau, NW Spain)

We reconstructed vegetation responses to climate oscillations, fire and human activities since the last glacial maximum in inland NW Iberia, where previous paleoecological research is scarce. Extremely sparse and open vegetation composed of steppic grasslands and heathlands with scattered pioneer trees suggests very cold and dry conditions during the Oldest Dryas, unsuitable for tree survival in the surroundings of the study site. Slight woodland expansion during the Bolling/Allerod was interrupted by the Younger Dryas cooling. Pinewoods dominated for most of the early Holocene, when a marked increase in fire activity occurred. Deciduous trees expanded later reaching their maximum representation during the mid-Holocene. Enhanced fire activity and the presence of coprophilous fungi around 6400-6000 cal yr BP suggest an early human occupation around the site. However, extensive deforestation only started at 4500 calyrBP, when fire was used to clear the tree canopy. Final replacement of woodlands with heathlands, grasslands and cereal crops occurred from 2700 cal yr BP onwards due to land-use intensification. Our paleoecological record can help efforts aimed at restoring the natural vegetation by indicating which communities were dominant at the onset of heavy human impact, thus promoting the recovery of currently rare oak and alder stands.

Caracterización morfofisiológica de semillas de especies leñosas distribuidas en dos zonas secas presentes en el Sur del Ecuador

Sudamérica es una de las zonas con mayor cantidad de bosque seco tropical a nivel mundial. No obstante, estos bosques han sido poco conocidos y la mayoría de estudios han estado orientados hacia los bosques húmedos tropicales. Los bosques secos se han reducido drásticamente y siguen muy amenazados, corriendo el riesgo de desaparecer en los próximos años. Por ello, es fundamental, generar investigación aplicada para la conservación inmediata de los ecosistemas secos tropicales. En Ecuador, la situación no es diferente y las zonas secas catalogadas como biodiversas están en constante amenaza. Los pocos estudios realizados en Ecuador sobre zonas secas, han permitido mejorar nuestro conocimiento referente a la diversidad y abundancia de las especies, relaciones planta-planta y síndromes de dispersión. No existen estudios sobre caracteres morfológicos en frutos y semillas de las especies leñosas de bosque seco. Sin embargo, nuestra comprensión de la dinámica y estructura de las comunidades ecológicas de zonas secas poco estudiadas, puede mejorar rápidamente mediante el estudio y enfoque de rasgos morfofisiológicos funcionales. El objetivo general del presente estudio fue aportar al conocimiento de la ecología y biología de semillas de zonas secas tropicales mediante el análisis y evaluación de rasgos morfofisiológicos de frutos y semillas de una comunidad de especies leñosas. El estudio se realizó en una zona de bosque y matorral seco, ubicados al sur occidente del Ecuador, a una altitud comprendida entre los 250 a 1 200 m s.n.m. caracterizada por una marcada estacionalidad ambiental, con lluvias desde diciembre a abril y una estación seca de mayo a noviembre. Precipitación media anual de 500 mm con una temperatura media anual de 20° a 26 °C. La zona de estudio forma parte de la región Tumbesina compartida entre el sur del Ecuador y el norte del Perú con gran diversidad de especies vegetales endémicas. Para el estudio se colectaron frutos con semillas maduras previamente a su dispersión de entre ocho y diez individuos de 80 especies entre árboles y arbustos más representativos de los bosques secos ecuatorianos. De los frutos colectados se utilizó una muestra al azar de 50 frutos y semillas por especie para los diferentes análisis. Se midió y evaluó 18 rasgos morfológicos y fisiológicos cuantitativos y cualitativos de frutos, semillas y de la especie. Se realizaron diferentes análisis de asociación y correlación entre los rasgos evaluados, con cinco variables ambientales registradas de las 109 parcelas establecidas en el área de estudio, además analizamos el tipo de dormición y comparamos la respuesta germinativa a la deshidratación relacionada con dos comunidades secas, matorral y bosque seco. Los resultados mostraron que las especies presentan gran heterogeneidad en rasgos continuos de las semillas. La variabilidad fue más evidente en rasgos como tamaño, volumen, masa y número de semillas por fruto. Sin embargo, una alta proporción de las especies tiende a producir una semilla por fruto. Además, la mayoría de las especies de bosque seco se caracterizan por no poseer algún tipo de apéndices o areola en sus semillas, forma ovalada y sin endospermo. La reserva nutritiva de las semillas se encuentra especialmente en los cotiledones de los embriones. Se encontraron seis tipos diferentes de embriones y la mayoría de las especies presentó embriones gruesos e invertidos. La dispersión de semillas está dominada por zoocoria en un 38 %, con relación a anemocoria (22 %) y autocoria (19 %). Sin embargo, encontramos que el 70 % de las especies posee frutos secos. Los análisis de dormición en las semillas de bosque seco, mostraron que el 60 % de las especies de bosque seco presentaron semillas con algún tipo de latencia, menor a la encontrada en especies de bosque deciduo tropical y sabanas, sin embargo, la dormición de las especies de bosque seco fue mayor al porcentaje de especies con dormición de bosque semiperenne y selva lluviosa tropical. La dormición física constituyó el 35 % de las especies de bosque seco, seguido del 12 % con dormición fisiológica, mientras que solamente una especie tuvo dormición morfológica. Encontramos que la dormición de las semillas de las especies en estudio se relaciona significativamente con el tipo y función del embrión y con el endospermo. Existieron relaciones significativas entre los rasgos morfológicos de los frutos, semillas, embriones y atributos de los individuos de 46 especies, aunque en algunos casos con coeficientes de correlación bajos. Hubo pocas relaciones entre los rasgo morfológicos de las semillas con las variables ambientales registradas. Solamente el tipo de testa y la presencia de apéndices en las semillas mostraron relación con el pH y la temperatura media del suelo. No obstante usando el modelo fouth corner-RLQ, no se encontraron asociaciones claras ni significativas entre rasgos morfológicos de semillas y frutos con variables ambientales. Al medir el efecto de la deshidratación en las semillas de los dos hábitats secos tropicales: bosque y matorral seco, los resultados determinaron que tanto las semillas de las especies leñosas de ambientes más áridos (matorral seco) están en gran medida pre-adaptadas a la desecación que las especies de ambientes menos áridos (bosque seco). Los tratamientos de deshidratación ejercieron un efecto negativo en los porcentajes de germinación en todas las especies, excepto para C. platanifolia. Los resultados más sorprendentes se registraron para Senna alata que mostró germinación extremadamente baja o incluso sin germinación a contenidos de humedad de la semillas de 0,10 g H2O g de peso seco. Las curvas de germinación difirieron significativamente entre los tratamientos de deshidratación en cada especie. Aportar al conocimiento la fisiología de la deshidratación y los límites de tolerancia de las semillas de bosque y matorral seco ayudará a entender mejor el papel de este rasgo en la ecología de las semillas y dinámica de las comunidades áridas tropicales. El estudio demostró, que la adaptación ecológica de las semillas de las especies leñosas de bosque seco a factores ambientales extremos, puede verse reflejada en una red de interacciones y correlaciones complejas entre los propios rasgos morfológicos y fisiológicos continuos y cuantitativos, sobre todo en rasgos internos de las semillas, quienes ejercerían una mayor influencia en toda la red de interacciones. Si bien, los rasgos de las semillas no mostraron fuertes relaciones con las variables ambientales, posiblemente las asociaciones presentes entre rasgos morfológicos pudiesen predecir en cambio interacciones entre especies y comportamientos y procesos relacionados con la tolerancia a la deshidratación y dormición de las semillas. ABSTRACT South America is one of the areas with the largest number of tropical dry forest in the world. However, these forests have been poorly understood and most studies have been directed to tropical rainforests. Dry forests have been drastically reduced and are very threatened, risking desaparecerer in the next years. It is therefore essential, generate applied research for conservation of tropical dry ecosystems. In Ecuador the situation is no different and dry areas classified as biodiverse are under constant threat. The few studies made in Ecuador on drylands have improved our knowledge concerning the diversity and abundance of species, plant-plant relationships and dispersion syndromes. Morphological studies on fruits and seeds of woody dry forest species do not exist. However, our understanding of the dynamics and structure of ecological communities dryland little studied, may improve quickly through the study and functional approach morphophysiological traits. The overall objective of this study was to contribute to the knowledge of the ecology and biology of tropical dry seeds through analysis and evaluation of morphophysiological traits of fruits and seeds of a community of woody species. The study was conducted in an area of dry scrub forest, located at the southwest of Ecuador, at an altitude between 250 to 1200 m asl. Environmental characterized by a marked seasonality, with rainfall from December to April and a dry season from May to November. Annual rainfall of 500 mm with an average annual temperature of 20° to 26 °C. The study area is part of the shared Tumbesina region between southern Ecuador and northern Peru with a great diversity of endemic plant species. For the study, we collected fruit and seed madure of eight and ten individuos of 80 species of trees and shrub most representated of the Ecuador dry forest. We selected a sample of 50 fruits and seeds for different analysis. We measure and evaluate 18 morphological and physiological traits of fruits, seeds and species. We perform analysis and correlation between traits associated with five environmental variables taken from the 109 plots established in the study area also analyze and compare the germination response to dehydration related to two dry communities, scrub and dry forest. The results showed that the species have great heterogeneity in continuous seed traits. Variability was more evident in features such as size, volume, mass, and number of seeds per fruit. However, a high proportion of species tends to produce a seed per fruit. In addition, most of the species of dry forest is characterized by not having some sort of ppendices or areola in its seeds, oval form and without endosperm. The nutrient reserves of seeds are especially in the cotyledons of the embryos. Six different embryos were found and most of the species presented thick and inverted embryos. Seed dispersal zoochory is dominated by 38 %, relative to anemochory (22 %) and autochory (19 %). However, we found that 70 % of the species has dried fruits. The analysis of dormancy from tropical dry forest, showed that 60 % of species showed seed dormancy, down from species found in tropical deciduous forest and savanna, however dormancy dry forest species was higher than the percentage of forest species dormancy semi-evergreen and tropical rain forest. Physical dormancy corresponds to 35 % of species, followed by 12 % with physiological dormancy, while only one species had morphological dormancy. We found that dormancy of the seeds was significantly related to the type and function of the embryo and the endospemo. There were significant relationships between morphological traits of fruits, seeds, embryos and attributes of individuals of 46 species, although in some cases with low correlation coefficients. There was little relationship between the morphologic traits of the seeds with the registered environmental variables. Only the type of tesla and the presence of appendages on the seeds showed relation to pH and the mean soil temperature. However, using the fourth corner-RLQ model, neither clear nor significant between morphological traits of seeds and fruits associations with environmental variables were found. The effect of dehydration on seeds of two tropical dry forest habitats was evident in dry scrub. The results determined that both the seeds of woody species forest and dry scrub are pre-adapted to drier conditions. Dehydration treatments exerted a negative effect on germination percentage in all species, except for C. platanifolia. However, all species germinated in treatments of extreme dryness, but in low percentages. The most striking results were recorded for Senna alata showed no germination when its moisture content was 0.10 g H2O g dry weight. Germination curves differ significantly between the treatments of dehydration in each species. Contribute to the knowledge of physiology and dehydration tolerance limits seeds dry scrub forest and help you better understand the role of this trait in seed ecology and dynamics of tropical arid communities. The study showed that the ecological adaptation of seeds of woody species of dry forest to extreme environmental factors may be reflected in a complex web of interactions and correlations between morphological and physiological traits continuous and quantitative themselves, especially in internal seed traits, who exerted a major influence on the entire network of interactions. While the seed traits showed strong relationships with environmental variables possibly present associations between morphological traits could predict interactions between species and change behaviors related to desiccation tolerance and seed dormancy processes.

Insect herbivory, plant defense, and early Cenozoic climate change

Insect damage on fossil leaves from the Central Rocky Mountains, United States, documents the response of herbivores to changing regional climates and vegetation during the late Paleocene (humid, warm temperate to subtropical, predominantly deciduous), early Eocene (humid subtropical, mixed deciduous and evergreen), and middle Eocene (seasonally dry, subtropical, mixed deciduous and thick-leaved evergreen). During all three time periods, greater herbivory occurred on taxa considered to have short rather than long leaf life spans, consistent with studies in living forests that demonstrate the insect resistance of long-lived, thick leaves. Variance in herbivory frequency and diversity was highest during the middle Eocene, indicating the increased representation of two distinct herbivory syndromes: one for taxa with deciduous, palatable foliage, and the other for hosts with evergreen, thick-textured, small leaves characterized by elevated insect resistance. Leaf galling, which is negatively correlated with moisture today, apparently increased during the middle Eocene, whereas leaf mining decreased.

A natural experiment on plant acclimation: lifetime stomatal frequency response of an individual tree to annual atmospheric CO2 increase.

Carbon dioxide (CO2) has been increasing in atmospheric concentration since the Industrial Revolution. A decreasing number of stomata on leaves of land plants still provides the only morphological evidence that this man-made increase has already affected the biosphere. The current rate of CO2 responsiveness in individual long-lived species cannot be accurately determined from field studies or by controlled-environment experiments. However, the required long-term data sets can be obtained from continuous records of buried leaves from living trees in wetland ecosystems. Fine-resolution analysis of the lifetime leaf record of an individual birch (Betula pendula) indicates a gradual reduction of stomatal frequency as a phenotypic acclimation to CO2 increase. During the past four decades, CO2 increments of 1 part per million by volume resulted in a stomatal density decline of approximately 0.6%. It may be hypothesized that this plastic stomatal frequency response of deciduous tree species has evolved in conjunction with the overall Cenozoic reduction of atmospheric CO2 concentrations.