Effectiveness of three sampling methods to survey saproxylic beetle assemblages in Mediterranean woodland

The choice of sampling methods to survey saproxylic beetles is a key aspect to assessing conservation strategies for one of the most endangered assemblages in Europe. We evaluated the efficiency of three sampling methods: baited tube traps (TT), window traps in front of a hollow opening (WT), and emergence traps covering tree hollows (ET) to study richness and diversity of saproxylic beetle assemblages at species and family levels in Mediterranean woodlands. We also examined trap efficiency to report ecological diversity, and changes in the relative richness and abundance of species forming trophic guilds: xylophagous, saprophagous/saproxylophagous, xylomycetophagous, predators and commensals. WT and ET were similarly effective in reporting species richness and diversity at species and family levels, and provided an accurate profile of both the flying active and hollow-linked saproxylic beetle assemblages. WT and ET were the most complementary methods, together reporting more than 90 % of richness and diversity at both species and family levels. Diversity, richness and abundance of guilds were better characterized by ET, which indicates higher efficiency in outlining the ecological community of saproxylics that inhabit tree hollows. TT were the least effective method at both taxonomic levels, sampling a biased portion of the beetle assemblage attracted to trapping principles, however they could be used as a specific method for families such as Bostrichiidae, Biphyllidae, Melyridae, Mycetophagidae or Curculionidae Scolytinae species. Finally, ET and WT combination allows a better characterization of saproxylic assemblages in Mediterranean woodland, by recording species with different biology and linked to different microhabitat types.

Explaining the saproxylic beetle diversity of a protected Mediterranean area

Saproxylic beetle diversity is high at the Cabañeros National Park (central Spain), where woodland habitats exhibit remarkable heterogeneity. Our aim was to explain the diversity of saproxylic beetles, focusing on species turnover among mature woodland types. We surveyed five woodland types that represented the heterogeneity of the park’s woodland habitats. Beetles were collected using window traps over a period of 20 months. The Jaccard Similarity Index was used as indirect value of beta diversity among woodlands and to test the relation between species turnover and geographical distance. We also identified the contribution of species turnover to landscape diversity by using a partitioning model. Moreover, the presence of mixed woodlands (more than one tree species) allowed us to attempt to valorise the effect of tree species (coupled with their historical management) on species turnover among woodlands. Finally, we looked for different saproxylic beetle preferences for habitat and tree species using an indicator value method. We found that saproxylic beetle species composition varied significantly among the studied woodlands. The variation in species turnover was independent from the distance among woodlands, which suggested that beetle dispersal abilities could not explain this high turnover. Tree species within woodlands were a key factor that increased diversity turnover in woodlands and, consequently, the diversity of the park. Moreover, we found saproxylic beetle species that had different habitat and tree species preferences. We conclude that woodland heterogeneity (highly affected by woodland composition) seems to be the driving force for saproxylic beetle diversity in this protected area.

On the edge: Southern Levantine Epipalaeolithic–Neolithic chronological succession

The broad picture of the cultural and chronological succession from the Epipalaeolithic to the Neolithic in the southern Levant is generally well understood. However, at a more detailed, local level, many questions remain unanswered. In this paper we examine the archaeological record of cultural developments in southern Jordan and the Negev. Focusing on a series of 14C dates from the early occupation of the PPNA site of WF16, we provide a critical review of dating evidence for the region. This review suggests that while the 14C chronology is ambiguous and problematic there is good evidence for a local historical development from the Harifian variant of the Natufian to the early PPNA, well to the south of any core Mediterranean woodland zone. This stresses the importance of considering developments at local scales of analysis, and that the Neolithic transition occurred within a framework of many interacting sub-regional provinces.

Synergy of extreme drought and shrub invasion reduce ecosystem functioning and resilience in water-limited climates

Extreme drought events and plant invasions are major drivers of global change that can critically affect ecosystem functioning and alter ecosystem-atmosphere exchange. Invaders are expanding worldwide and extreme drought events are projected to increase in frequency and intensity. However, very little is known on how these drivers may interact to affect the functioning and resilience of ecosystems to extreme events. Using a manipulative shrub removal experiment and the co-occurrence of an extreme drought event (2011/2012) in a Mediterranean woodland, we show that native shrub invasion and extreme drought synergistically reduced ecosystem transpiration and the resilience of key-stone oak tree species. Ecosystem transpiration was dominated by the water use of the invasive shrub Cistus ladanifer, which further increased after the extreme drought event. Meanwhile, the transpiration of key-stone tree species decreased, indicating a competitive advantage in favour of the invader. Our results suggest that in Mediterranean-type climates the invasion of water spending species and projected recurrent extreme drought events may synergistically cause critical drought tolerance thresholds of key-stone tree species to be surpassed, corroborating observed higher tree mortality in the invaded ecosystems. Ultimately, this may shift seasonally water limited ecosystems into less desirable alternative states dominated by water spending invasive shrubs.

Dinámicas de ocupación urbana del Anillo Verde metropolitano, desde sus orígenes en el Plan General del Área Metropolitana de Madrid de 1963 hasta el Plan General de Ordenación Urbana de 1997

El Anillo Verde metropolitano, definido por el Plan General de Ordenación Urbana del Área Metropolitana de Madrid en 1963 siguiendo los modelos planteados por la cultura urbanística internacional, como armadura de la estructura urbana del AMM, espacio protagonista dentro del sistema de espacios libres, lugar de uso público destinado al recreo y contacto con la naturaleza de la población madrileña, se convierte en realidad en una reserva de suelo que va entrando en juego motivado por las alianzas entre el poder institucional y la clase social dominante actuando al margen del planeamiento, poniendo en evidencia la escasez de recursos legales y culturales disponibles para la salvaguarda de los intereses comunes y, donde los condicionantes geográficos y naturales del territorio madrileño han influido decisivamente en la especialización funcional y espacial del Área Metropolitana de Madrid. Así pues considerando esta idea como HIPÓTESIS, el objetivo de la TESIS sería demostrarla, para lo cual se hace necesario primero, acotar espacial y temporalmente el objeto de estudio, es decir, del Anillo Verde metropolitano1, segundo, contextualizar histórica y disciplinarmente los presupuestos teóricos que conformaban la idea del Anillo Verde, tercero, reconocer, localizar y documentar las piezas que han ido materializando la ocupación urbana del Anillo Verde, clasificándolas según parámetros temporales, funcionales, urbanísticos y, formales, lo que permite analizar la geografía, uso, instrumentación y forma de su transformación a escala general metropolitana y, cuarto, profundizar a modo de comprobación a escala municipal y urbana en dos escenarios representativos del conjunto metropolitano: el municipio de Pozuelo de Alarcón y el distrito de Hortaleza-Barajas. El contenido del documento se divide en tres bloques, el bloque I, se centra en las bases teóricas, el bloque II sitúa el hilo argumental de la tesis a escala metropolitana y el bloque III comprueba el fenómeno a escala municipal y urbana. De esta forma, se comienza por la comprensión del significado del concepto del Anillo Verde, que va más allá de la dimensión instrumental asignada de límite y contención urbana frente al crecimiento de la ciudad industrial de principios del siglo XX, basada en la descentralización de la ciudad tradicional, para adquirir un significado más complejo, como gran espacio de reserva y salvaguarda de valores naturales y culturales que se expresaban en su territorio y que permitirían alcanzar el equilibrio entre la ciudad y sus habitantes, es decir, entre el hombre y el espacio que habita. Se hace un recorrido por las principales corrientes urbanísticas que se van nutriendo de distintas disciplinas (economía, sociología, geografía, biología, ecología) para plantear teorías que permitieran materializar un nuevo orden urbano según principios de equidad social, económica y ambiental, en una secuencia donde Europa y Estados Unidos realizaban un constante intercambio -el movimiento de la Ciudad Jardín o el Regionalismo, que dieron paso a propuestas como el Greater London o el Gran Berlín, donde la figura del Anillo Verde tenía un papel protagonista, y del que también participaría nuestro país y la ciudad de Madrid, con modelos regionales como el Plan Besteiro y urbanos como el Plan Bidagor, antecedentes directos del Plan General de Ordenación Urbana del Área Metropolitana de Madrid de 1963 que pone en marcha la ordenación del crecimiento metropolitano de Madrid. El hilo argumental de la tesis se organiza en una doble aproximación: un acercamiento a escala metropolitana a partir del reconocimiento del modelo de ciudad definido en los distintos planes generales que acompañaron el desarrollo metropolitano (municipio de Madrid y de los siete términos municipales que rodeaban a este y que tenían suelo destinado a Anillo Verde), haciendo referencia además a las relaciones con el planeamiento regional, concretando en una escala de aproximación municipal que avanza hasta la interpretación urbana detallada. El primer acercamiento tiene lugar en el bloque II y se organiza en tres capítulos. El capítulo 4 se dedica al punto obligado de partida de la geografía local, describiendo las características biofísicas de los terrenos que formaban parte del Anillo Verde, que han marcado históricamente la forma de aprovechamiento del territorio, desde las extensiones de bosques mediterráneos al norte y al oeste continuación del Monte del Pardo, a los distintos tipos de cultivo que se adaptaban al sustrato geológico y la forma del terreno (de las suaves ondulaciones de sedimentos arcósicos al norte a las extensas plataformas arenosas y yesíferas del sur), además de las zonas de huertos aprovechando las depresiones y los cursos de agua (arroyo del Monte Carmelo, arroyo de Valdebebas, arroyo del Quinto, arroyo del Santo, arroyo Butarque, arroyo Meaques y arroyo Pozuelo). Una vez reconocida la realidad física, el capítulo 5, avanza en la descripción de los distintos modelos de ciudad propuestos desde el planeamiento urbanístico, en sus distintas escalas, la regional y la municipal, como respuesta a la coyuntura social, económica y política que ha caracterizado el proceso de ocupación del Anillo Verde al compás de la construcción del AMM. Se han reunido las propuestas de planeamiento municipal de los distintos municipios que disponían de terreno calificado como Anillo Verde: Madrid, Coslada, Getafe, Leganés, Alcorcón, Boadilla del Monte y Pozuelo de Alarcón. Además se han incorporado las distintas propuestas de ordenación territorial que han servido de referencia al planeamiento municipal, en todas sus versiones, desde las sectoriales, de mayor éxito y apoyo institucional, a los distintos intentos de ordenación integral, de mayor complejidad pero de menor calado, precisamente por la dificultad de consenso entre la ordenación física y el desarrollo económico, entre los intereses privados y el beneficio público. El primer horizonte, comienza con la formulación del Plan General de Ordenación Urbana del Área Metropolitana de Madrid de 1963, su desarrollo y la puesta en marcha de los primeros planes municipales en la década de los años setenta, donde se comprueba la necesidad de un marco regional que “ordene” el territorio de forma integral y sirva de referencia a las actuaciones sectoriales que habían marcado el primer desarrollo metropolitano. El segundo, se sitúa dos décadas más tarde con la aprobación del Plan General de Ordenación Urbana de Madrid de 1985 y el conjunto de planes municipales de los términos limítrofes, que siguen su filosofía de austeridad en cuanto a crecimiento territorial. El tercero se inicia en 1997 con la siguiente generación de planes de corte neoliberal que imponen un modelo territorial basado en las grandes operaciones metropolitanas de centralidad, infraestructuras y equipamiento, que consumen de forma indiscriminada la totalidad del territorio madrileño. Será en el último capítulo del segundo bloque (capítulo 6) donde se represente gráficamente a escala metropolitana y se analicen las 229 piezas que han ido colmatando el espacio destinado a Anillo Verde, según los parámetros de estudio, en base a las cuales se plantean las primeras conclusiones generales de la tesis, poniendo de manifiesto que las alianzas entre los agentes soberanos en la construcción de la ciudad y su entorno han trasgredido sucesivamente las determinaciones del Planeamiento en su definición de modelo de ciudad y territorio, acusando la carencia de recursos instrumentales y jurídicos que alentaron el proceso de su desmantelamiento, y revelando la influencia de los condicionantes geográficos y naturales en la especialización funcional y segregación social en el conjunto del Área Metropolitana de Madrid. Se remata el discurso metropolitano con una batería de conclusiones que interpretan el fenómeno de ocupación del anillo de verdor metropolitano confirmando las hipótesis iniciales, reconociendo los valores medioambientales y culturales trasgredidos, sus diversos actores, las numerosas operaciones urbanísticas desarrolladas con distintos usos y envergadura, así como los instrumentos de planeamiento utilizados, en base a las cuales se materializa la construcción del AMM según un modelo extendido (spread), dibujando una mancha de aceite (o grase-spots según Geddes) que precisamente había querido evitarse desde el planeamiento urbanístico con la definición de un Anillo Verde, espacio inmune a la edificación, que se aleja de su papel estructurante (equilibrador entre la ciudad y sus habitantes) para convertirse en armadura de la estructura comunicativa, que una vez consolidada se convierte en la mejor aliada de la máquina inmobiliaria. El último paso, se desarrolla en el bloque III que se divide en los capítulo 7,8 y 9 y supone la comprobación de lo descrito en el conjunto de escala metropolitana, en dos aspectos fundamentales, la falta de consideración por los valores culturales y medioambientales que han modelado el territorio, imprimiéndole un carácter singular y específico y, la estructura del dominio del suelo, donde se reconoce de forma precisa el grupo social y los agentes encargados en cada momento de comercializar los suelos del anillo, que bajo el paraguas de la urgencia social y el engañoso beneficio popular, obtienen importantes beneficios económicos. Con esa intención, se da un salto hacia la escala municipal y urbana, seleccionando dos escenarios de estudio, el municipio de Pozuelo de Alarcón, que representa la materialización del crecimiento suburbano de la élite madrileña ocupando las zonas de mayor valor ecológico del anillo, y el distrito de Hortaleza-Barajas que ofrece su territorio a las grandes operaciones metropolitanas, apoyándose en el eje de actividad marcado por la conexión Madrid-Barcelona y el sistema aeroportuario de escala global, ambos situados al norte de la línea de borde entre la Sierra y la Mancha, ocupando por tanto los lugares más valiosos de la geografía madrileña (estructura funcional anticipada por Bidagor en 1946 en su modelo de ciudad adaptada al territorio madrileño) Una vez descrito este proceso trasgresor de límites, de normas, de conductas, y desde una perspectiva del fenómeno suficientemente documentada, en el capítulo 10, se realiza una reflexión sobre la incidencia real de la propuesta urbanística del Anillo Verde en la construcción del AMM, de la misma forma que se sugieren nuevos roles al planeamiento en un formato intencionado de largo recorrido en oposición a lo inmediato y circunstancial, que permita hacer una nueva lectura de los presupuestos teóricos que conformaban la idea del Anillo Verde, espacio articulador (medioambiental, social y cultural) del territorio madrileño. ABSTRACT The Metropolitan Greenbelt was defined by the 1963 Master Plan for the Madrid Metropolitan Area (MMA), following established international models of urban development, as the structural framework of the MMA, the principal open space within its network of open spaces and a public area of recreation and contact with nature for the residents of Madrid. In reality, however, it ha become a reserve of land in which various alliances between the institutional authorities and the dominant social class have been operating on the margin of the original plan, exposing a scarcity of legal and cultural resources for the safeguarding of common interests, and in which the geographical and natural characteristics of the territory itself have come to play an influential role in the functional specialization and spatial segregation of the MMA. With that idea as its HYPOTHESIS, the aim of this THESIS is to demonstrate its reality. The first step in this is to delineate, temporally and spatially, the object of study; i.e. the Metropolitan Greenbelt2. The second is to contextualize historically and disciplinarily those theoretical ideas which conform to the greenbelt concept. The third is to acknowledge, locate and document the elements which have characterized the urban occupation of the Greenbelt and classify these according to the parameters of time, function, urban development and form, which in turn would enable the geography, use, instrumentation and form of its transformation to be analysed on a general metropolitan scale. The fourth step, as a method of verification, is an in-depth analysis of two representative settings within the metropolitan network: the municipality of Pozuelo de Alarcón and the Hortaleza-Barajas district. The content of the document is divided into three parts. Part I focuses on the study’s theoretical foundations, Part II establishes a line of argument at the metropolitan level and Part III examines the phenomenon from a municipal and urban perspective. The thesis, then, begins with a study of the greenbelt concept itself and its meaning, which is far more complex than the accepted instrumental dimension of limiting and containing urbanization in response to the growth of the industrial city of the early 20th century, and which is based on a decentralization of the traditional city. This wider purpose is the setting aside of a large reserved space to safeguard the natural and cultural values of the region and thereby achieve a balance between the city and its residents; that is to say, between man and the space he inhabits. The principal currents of thought in urban planning will then be examined. These have drawn upon a variety of disciplines (economics, sociology, geography, biology, ecology) to develop theories for establishing a new urban order according to the principles of social, economic and environmental equity, and have involved a constant interchange between Europe and the United States. Thus, the City Garden and Regionalist movements would clear the way for proposals such as Greater London and Great Berlin, Chicago and Washington, in which the greenbelt would play a fundamental role. The participation of our own country and the city of Madrid is also discussed, through regional models such as the Besteiro Plan and urban ones like the Bidagor Plan, direct forerunners of 1963’s General Organizational Plan for the Madrid Metropolitan Area, which would set into motion the organization of Madrid’s metropolitan growth. The line of argument followed in this thesis is two-fold: first, an examination of metropolitan development in keeping with the city model as defined in the various General Plans for the development of both the municipality of Madrid and the seven surrounding municipalities which have land designated for its Greenbelt; and second, an examination of this growth in relation to Regional Planning measures, is detailed on a smaller scale (municipal and district), where the conditioning factors affecting the land property structure and the network of biophysical units may be analysed in depth. The first of these is dealt with in Part II and organized into three chapters (4, 5 and 6). Chapter 4 is dedicated to the obligatory starting point of the geographical setting itself. The biophysical characteristics of the territories set aside for the Greenbelt, and which historically have played a role in the area’s exploitation, are described here. These range from expanses of Mediterranean woodland to the north and west of Monte del Pardo to the various types of farmland that have been adapted to the geological substratum and the contours of the terrain (gentle undulations of arkosic sediment in the north, and wide sandy and gypsiferous tableland in the south), as well as orchards planted in low valleys and along watercourses (the creeks of Monte Carmelo, Valdebebas, Quinto, Santo, Butarque, Meaques and Pozuelo). Once this physical reality ha been detailed, in Chapter 5 will examine the various city models proposed by urban planners, both regionally and municipally, in response to the confluence of social, economic and political interests that have characterized the process of occupation in the Greenbelt area during the construction of the MMA. Municipal planning proposals will be collected and examined for the various municipalities which have land designated for the Greenbelt: Madrid, Coslada, Getafe, Leganés, Alcorcón, Boadilla del Monte and Pozuelo de Alarcón. Furthermore, the various territorial organization proposals which have served as references for municipal planning will also be addressed here, in all of their versions –from the sectorial, which have met with more success and institutional approval, to the many attempts at integration, which have been more complex but less influential, precisely for the difficulty of reconciling physical organization with economic development, and private interest with public benefit. The first period in this process was the development of the General Plan of 1963, followed by the first municipal development plans of the 1970s, in which the need for a regional framework that “organized” the territory in an integral fashion was defined. This would serve as a reference for the sectorial actions that marked the metropolitan area’s initial development. The second came two decades later with the approval of the General Plan of 1985, and the network of municipal plans for the surrounding communities, which followed the same philosophy of austerity with regard to territorial growth. The third would begin to take form in 1997, as a new generation of neo-liberal development plans imposed a territorial model based on centralized large-scale metropolitan operations of infrastructure and equipment, which would indiscriminately consume the totality of Madrid’s land. At the end of the Part II, in Chapter 6, the metropolitan area will be represented graphically and the 229 pieces that have been gradually encroaching upon land designated for the Greenbelt will be analysed. This analysis will be carried out according to the parameters defined for the study, and the first general conclusions of the thesis will be based on its findings. It will show how alliances between the various governing authorities in the construction of the city and its environment have successively violated established plans with regard to the definitions of city and territory, how shortages of instrumental and judicial resources have accentuated the dismantling process, and how natural and geographical factors have influenced functional specialization and social segregation in the Madrid Metropolitan Area. The final step, detailed in Part III, will address two fundamental aspects of what has just been described: the lack of consideration for the cultural and environmental values which have shaped this territory and imprinted upon it a specific and unique character; and the structure of land domination, with a precise identification of the social group and agents responsible at each stage of the Greenbelt’s commercialization, who, under an umbrella of social urgency and deceptive public benefit, have used it to obtain substantial financial rewards. For this purpose, a closer look is taken at two specific areas: the municipality of Pozuelo de Alarcón, representative of the suburban growth of an elite population which has occupied the Greenbelt areas of the greatest ecological value; and the Hortaleza-Barajas district, which has offered its territory to large metropolitan business interests, based on activities centred on the connection between Madrid and Barcelona and the system of international air travel. Both of these settings are located to the north of the line which divides the Sierra from La Mancha, and thus occupy the most valuable land in the Madrid region (a functional structure anticipated by Bidagor in 1946, with his city model adapted to the territory of Madrid). Finally, an attempt will be made to interpret the phenomenon of metropolitan Greenbelt occupation, confirming initial hypotheses, specifying the environmental and cultural values that have been violated, and identifying the various players involved, as well as numerous urbanization operations of varying sizes and interests, and the instruments of planning they have used. It will be seen from this that the construction of the MMA has in fact followed a “spread” model, a “grease spot” (as Geddes calls it) which, from the outset of the planning process and according to the definition of a greenbelt as a construction-free zone, was precisely to be avoided. This structural role (to provide a balance between a city and its residents) has thus been abandoned and the Greenbelt converted instead into a communicative framework which, once consolidated, has become the greatest ally of the real estate machine. After this process of violating limits, norms and established behaviour has been described and solidly documented, a reflection will be made on the real influence of the Greenbelt proposal in the construction of the MMA. At the same time, new roles will be suggested for future planning, roles which are deliberate and long term, in opposition to the immediate and circumstantial. This will enable a new interpretation of the theoretical principles behind the greenbelt concept, a space designed to connect the territory of Madrid environmentally, socially and culturally.

Jugar a extinguir ensambles: importancia de las oquedades de los árboles para la biodiversidad de insectos saproxílicos

The assessment of the relation between species diversity, species interactions and stability is a major goal for conservation strategies of saproxylic insects in Iberian Mediterranean woodlands. In these woodlands, tree hollows in standing living trees are key microhabitats for saproxylic biodiversity. We assess the variation in the tree hollow-saproxylic insect network among three woodland sites in the Cabañeros National Park (Spain), and evaluate their implications for saproxylic assemblages’ persistence, through simulations of three possible scenarios of loss of tree hollow microhabitats. The composition of tree hollows is closely related to biotic and ecological complexity of saproxylic networks, and together exhibit positive effects on network stability. By other side, different scenarios entail different reorganizations of saproxylic diversity of species and interactions, and the variation of the main interacting attributes. Woodlands providing higher proportion of large tree hollows are less sensitive to microhabitat loss. Conservation strategies for saproxylic insects in Iberian Mediterranean woodlands should consider woodland composition and configuration as key factors in the selection of priority conservation areas.

Geographical and environmental correlates of big and small game in Andalusia (southern Spain)

In Andalusia, southern Spain, each game estate applies its own rules and presents its results in annual hunting reports, which have been mandatory for Spanish game estates since 1989. We used the information about hunting yields, included in 32134 annual hunting reports produced during the period 1993/94 to 2001/02 by 6049 game estates, to determine the current distribution of hunting yields of big and small game species in Andalusia. Using generalised linear models and a geographic information system, we determined the most favourable municipalities to big and small game, respectively, and delimited potential areas to attain good hunting yields for big and small game at a 1-km2 resolution. Municipalities and areas favourable to big game are mainly located in the Sierra Morena and the westernmost fringe of the Betic Range, while those favourable to small game occupy the upper Guadalquivir River valley. There is a clear segregation between big and small game species according to the physiography and land uses of the territory. Big game species are typical of Mediterranean woodland areas, while the most emblematic small game species prefer agricultural areas. Our results provide a territorial ordination of hunting yields in southern Spain and have several potential applications in strategic planning for hunting activities and biodiversity conservation in Andalusia that can be extrapolated to other regions.

Riparian woodland encroachment following flow regulation: a comparative study of Boreal and Mediterranean streams

Water development accompanying mankind development has turned rivers into endangered ecosystems. Improving the understanding of ecological responses to river management actions is a key issue for assuring sustainable water management. However, few studies have been published where ecological metrics have been quantified in response to various degrees of flow alteration. In this work, changes in natural distribution of trees and shrubs within the riparian corridor (as indicator of the ecological status of the fluvial ecosystem) were quantified at multiple sites along a flow alteration gradient (as indicator of impact) along two regulated river reaches, one Boreal and the other Mediterranean, each downstream of a dam. Based on the obtained relationships we evaluated differences in response trends related to local physico-climatic factors of the two biomes and regarding to differing life-forms. Woody vegetation establishment patterns represented objective indicators of ecological responses to flow alteration. We found different responses between life-forms. Both trees and shrubs migrated downwards to the channel after dam closure, but shrubs were most impacted under higher degrees of flow alteration in terms of lateral movement. In addition, our results show clear longitudinal recovery trends of natural patterns of tree and shrub distribution corresponding to a decrease in intensity of hydrologic alteration in the Boreal river. However, vegetation encroachment persisted along the entire Mediterranean study reach. This may result from a relatively low gradient of decrease of hydrologic alteration with distance from the dam, coupled with other overlapping pressures and the mediating effect of physico-climatic characteristics on vegetation responses.

Fertilizing Methods and Nutrient Balance at the End of Traditional Organic Agriculture in the Mediterranean Bioregion: Catalonia (Spain) in the 1860s

By reconstructing the nutrient balance of a Catalan v illage circa 1861-65 we examine the sustainability of organic agricultural sy stems in the northwest Mediterranean bioregion prior to the green rev olution and the question of whether the nutrients extracted f rom the soil were replenished. With a population density of 59 inhabitants per square km, similar to other northern European rural areas at that time, and a lower liv estock density per cropland unit, this v illage experienced a manure shortage. The gap was f illed by other labour-intensiv e way s of transf erring nutrients f rom uncultiv ated areas into the cropland. Key elements in this agricultural sy stem were v iney ards because they hav e f ew nutrient requirements, and woodland and scrublands as sources of relev ant amounts of nutrients collected in sev eral ways.

Plant-pollinator biodiversity and pollination services in a complex Mediterranean landscape

Mediterranean landscapes comprise a complex mosaic of different habitats that vary in the diversity of their floral communities, pollinator communities and pollination services. Using the Greek Island of Lesvos as a model system, we assess the biodiversity value of six common habitats and measure ecosystemic 'health' using pollen grain deposition in three core flowering plants as a measure of pollination services. Three fire-driven habitats were assessed: freshly burnt areas, fully regenerated pine forests and intermediate age scrub; in addition we examined oak woodlands, actively managed olive groves and groves that had been abandoned from agriculture. Oak woodlands, pine forests and managed olive groves had the highest diversity of bees. The habitat characteristics responsible for structuring bee communities were: floral diversity, floral abundance, nectar energy availability and the variety of nectar resources present. Pollination services in two of our plant species, which were pollinated by a limited sub-set of the pollinator community, indicated that pollination levels were highest in the burnt and mature pine habitats. The third species, which was open to all flower visitors, indicated that oak woodlands had the highest levels of pollination from generalist species. Pollination was always more effective in managed olive groves than in abandoned groves. However, the two most common species of bee, the honeybee and a bumblebee, were not the primary pollinators within these habitats. We conclude that the three habitats of greatest overall value for plant-pollinator communities and provision of the healthiest pollination services are pine forests, oak woodland and managed olive groves. We indicate how the highest value habitats may be maintained in a complex landscape to safeguard and enhance pollination function within these habitats and potentially in adjoining agricultural areas. (c) 2005 Elsevier Ltd. All rights reserved.

Landscape typology in the Mediterranean context: a tool for habitat restoration

Despite the wide use of Landscape Character Assessment (LCA) as a tool for landscape planning in NW Europe, there are few examples of its application in the Mediterranean. This paper reports on the results from the development of a typology for LCA in a study area of northern Sardinia, Italy to provide a spatial framework for the analysis of current patterns of cork oak distribution and future restoration of this habitat. Landscape units were derived from a visual interpretation of map data stored within a GIS describing the physical and cultural characteristics of the study area. The units were subsequently grouped into Landscape Types according to the similarity of shared attributes using Two Way Indicator Species Analysis (TWINSPAN). The preliminary results showed that the methodology classified distinct Landscape Types but, based on field observations, there is a need for further refinement of the classification. The distribution and properties of two main cork oak habitats types was examined within the identified Landscape Types namely woodlands and wood pastures using Patch Analyst. The results show very clearly a correspondence between the distribution of cork oak pastures and cork oak woodland and landscape types. This forms the basis of the development of strategies for the maintenance, restoration and recreation of these habitat types within the study area, ultimately for the whole island of Sardinia. Future work is required to improve the landscape characterisation , particularly with respect to cultural factors, and to determine the validity of the landscape spatial framework for the analysis of cork oak distribution as part of a programme of habitat restoration and re-creation.

The effects of cattle grazing on plant-pollinator communities in a fragmented Mediterranean landscape

The main aims of this study were to assess grazing impacts on bee communities in fragmented mediterranean shrubland (phrygana) and woodland habitats that also experience frequent wildfires, and to explain the mechanisms by which these impacts occur. Fieldwork was carried out in 1999 and 2000 on Mount Carmel, in northern Israel, a known hot-spot for bee diversity. Habitats with a range of post-burn ages and varying intensities of cattle grazing were surveyed by transect recording, grazing levels, and the diversity and abundance of both flowers and bees were measured. The species richness of both bees and flowers were highest at moderate to high grazing intensities, and path-analysis indicated that the effects of both grazing and fire on bee diversity were mediated mainly through changes in flower diversity, herb flowers being more important than shrubs. The abundance of bees increased with intensified grazing pressure even at the highest levels surveyed. Surprisingly though, changes in bee abundance at high grazing levels were not caused directly by changes in flower cover. The variation in bee abundance may have been due to higher numbers of solitary bees from the family Halictidae in grazed sites, where compacted ground (nesting resource) and composites (forage resource) were abundant. The effects of grazing on plants were clearest in the intermediate-aged sites, where cattle inhibited the growth of some of the dominant shrubs, creating or maintaining more open patches where light-demanding herbs could grow, thus allowing a diverse flora to develop. Overall, bee communities benefit from a relatively high level of grazing in phrygana. Although bee and flower diversity may decrease under very heavy grazing, the present levels of grazing on Mount Carmel appear to have only beneficial effects on the bee community.

Indian summer monsoon variations could have affected the early Holocene woodland expansion in the Near East

Postglacial expansion of deciduous oak woodlands of the Zagros—Anti-Taurus Mountains, a major biome of the Near East, was delayed until the middle Holocene at ~6300 cal. yr BP. The current hypotheses explain this delay as a consequence of a regional aridity during the early Holocene, slow migration rates of forest trees, and/or a long history of land use and agro-pastoralism in this region. In the present paper, support is given to a hypothesis that suggests different precipitation seasonalities during the early Holocene compared with the late Holocene. The oak species of the Zagros—Anti-Taurus Mts, particularly Quercus brantii Lindl., are strongly dependent on spring precipitation for regeneration and are sensitive to a long dry season. Detailed analysis of modern atmospheric circulation patterns in SW Asia during the late spring suggests that the Indian Summer Monsoon (ISM) intensification can modify the amount of late spring and/or early summer rainfall in western/northwestern Iran and eastern Anatolia, which could in turn have controlled the development of the Zagros—Anti-Taurus deciduous oak woodlands. During the early Holocene, the northwestward shift of the Inter-Tropical Convergence Zone (ITCZ) could have displaced the subtropical anticyclonic belt or associated high pressure ridges to the northwest. The latter could, in turn, have prevented the southeastward penetration of low pressure systems originating from the North Atlantic and Black Sea regions. Such atmospheric configuration could have reduced or eliminated the spring precipitation creating a typical Mediterranean continental climate characterized by winter-dominated precipitation. This scenario highlights the complexity of biome response to climate system interactions in transitional climatic and biogeographical regions.

Molecular phylogeography of the Woodland Ringlet (Erebia medusa [Denis and Schiffermüller] 1775) in Europe

Phylogeography is a recent field of biological research that links phylogenetics to biogeography through deciphering the imprint that evolutionary history has left on the genetic structure of extant populations. During the cold phases of the successive ice ages, which drastically shaped species’ distributions since the Pliocene, populations of numerous species were isolated in refugia where many of them evolved into different genetic lineages. My dissertation deals with the phylogeography of the Woodland Ringlet (Erebia medusa [Denis and Schiffermüller] 1775) in Central and Eastern Europe. This Palaearctic butterfly species is currently distributed from central France and south eastern Belgium over large parts of Central Europe and southern Siberia to the Pacific. It is absent from those parts of Europe with mediterranean, oceanic and boreal climates. It was supposed to be a Siberian faunal element with a rather homogeneous population structure in Central Europe due to its postglacial expansion out of a single eastern refugium. An already existing evolutionary scenario for the Woodland Ringlet in Central and Eastern Europe is based on nuclear data (allozymes). To know if this is corroborated by organelle evolutionary history, I sequenced two mitochondrial markers (part of the cytochrome oxydase subunit one and the control region) for populations sampled over the same area. Phylogeography largely relies on the construction of networks of uniparentally inherited haplotypes that are compared to geographic haplotype distribution thanks to recent developed methods such as nested clade phylogeographic analysis (NCPA). Several ring-shaped ambiguities (loops) emerged from both haplotype networks in E. medusa. They can be attributed to recombination and homoplasy. Such loops usually avert the straightforward extraction of the phylogeographic signal contained in a gene tree. I developed several new approaches to extract phylogeographic information in the presence of loops, considering either homoplasy or recombination. This allowed me to deduce a consistent evolutionary history for the species from the mitochondrial data and also adds plausibility for the occurrence of recombination in E. medusa mitochondria. Despite the fact that the control region is assumed to have a lack of resolving power in other species, I found a considerable genetic variation of this marker in E. medusa which makes it a useful tool for phylogeographic studies. In combination with the allozyme data, the mitochondrial genome supports the following phylogeographic scenario for E. medusa in Europe: (i) a first vicariance, due to the onset of the Würm glaciation, led to the formation of several major lineages, and is mirrored in the NCPA by restricted gene flow, (ii) later on further vicariances led to the formation of two sub-lineages in the Western lineage and two sub-lineages in the Eastern lineage during the Last Glacial Maximum or Older Dryas; additionally the NCPA supports a restriction of gene flow with isolation by distance, (iii) finally, vicariance resulted in two secondary sub-lineages in the area of Germany and, maybe, to two other secondary sub-lineages in the Czech Republic. The last postglacial warming was accompanied by strong range expansions in most of the genetic lineages. The scenario expected for a presumably Siberian faunal element such as E. medusa is a continuous loss of genetic diversity during postglacial westward expansion. Hence, the pattern found in this thesis contradicts a typical Siberian origin of E. medusa. In contrast, it corroboratess the importance of multiple extra-Mediterranean refugia for European fauna as it was recently assumed for other continental species.