Palaeobiogeographical perspectives on pinus pinea, a controversial and enigmatic mediterranean pine

The origins of some species of economic importance occurring over the Mediterranean Basin have been a traditional matter of debate that has important implications for land management. The case of Pinus pinea L. (Stone pine) is probably one of the most controversial, due to its documented long-term interaction with humans and its presence as a symbolic tree in certain areas of the Mediterranean (e.g., southwestern Iberia and Tuscany). Among the rest of the Mediterranean pines, several features make this pine unique (it has a characteristic crown shape, an edible kernel, cones that require three years to mature, and a very depauperate genetic diversity across its range). In addition, its palaeoecological information is rather limited, as the taxonomic precision attained by pollen analysts is insufficient for this tree and macroremains (such as kernels or anatomically well preserved wood) are needed to unequivocally detect the species in the fossil record. Recent findings of macrofossils of Pinus pinea in inland Iberia (Duero Basin) extend the late- Holocene range of the species, but the palaeobiogeographical information and the exhaustive genetic data available still suggest a very limited natural area (but still not sufficiently well defined) and a long and intense history of linkage to humans.

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.

Age determination of sediments of Site LO09-14 (Table 1)

A composite record (LO09-14) of three sediment cores from the subpolar North Atlantic (Reykjanes Ridge) was investigated in order to assess surface ocean variability during the last 11 kyr. The core site is today partly under the influence of the Irminger Current (IC), a branch of the North Atlantic Drift continuing northwestward around Iceland. However, it is also proximal to the Sub-Arctic Front (SAF) that may cause extra dynamic hydrographic conditions. We used statistical methods applied to the fossil assemblages of diatoms to reconstruct quantitative sea surface temperatures (SSTs). Our investigations give evidence for different regional signatures of Holocene surface oceanographic changes in the North Atlantic. Core LO09-14 reveal relatively low and highly variable SSTs during the early Holocene, indicating a weak IC and increased advection of subpolar water over the site. A mid-Holocene thermal optimum with a strong IC occurs from 7.5 to 5 kyr and is followed by cooler and more stable late Holocene surface conditions. Several intervals throughout the Holocene are dominated by the diatom species Rhizosolenia borealis, which we suggest indicates proximity to a strongly defined convergence front, most likely the SAF. Several coolings, reflecting southeastward advection of cold and ice-bearing waters, occur at 10.4, 9.8, 8.3, 7.9, 6.4, 4.7, 4.3 and 2.8 kyr. The cooling events recorded in the LO09-14 SSTs correlate well with both other surface records from the area and the NADW reductions observed at ODP Site 980 indicating a surface-deepwater linkage through the Holocene.

Geochemistry and grain-size of sediment core PC29A, Quitralco Fjord, Chilean Patagonia

The climate of Chilean Patagonia is strongly influenced by the southern westerlies, which control the amount and latitudinal distribution of precipitation in the southern Andes. In austral summer, the Southern Westerly Wind Belt (SWWB) is restricted to the high latitudes. It expands northward in winter, which results in a strong precipitation seasonality between 35 and 45°S. Here, we present a new precipitation seasonality proxy record from Quitralco fjord (46°S), where relatively small latitudinal shifts in the SWWB result in large changes in precipitation seasonality. Our 1400 yr record is based on sedimentological and geochemical data obtained on a sediment core collected in front of a small river that drains the Patagonian Andes, which makes this site particularly sensitive to changes in river discharge. Our results show Fe/Al and Ti/Al values that are low between 600 and 1200 CE, increasing at 1200-1500 CE, and high between 1500 and 1950 CE. The increasing Fe/Al and Ti/Al values reflect a decrease in mean sediment grain-size from 30 to 20 µm, which is interpreted as a decrease in seasonal floods resulting from an equatorward shift of the SWWB. Our results suggest that, compared to present-day conditions, the SWWB was located in a more poleward position before 1200 CE. It gradually shifted towards the equator in 1200-1500 CE, where it remained in a sustained position until 1950 CE. The comparison of our record with published regional sea surface temperature (SST) reconstructions for the late Holocene shows that equatorward shifts in the SWWB are systematically coeval with decreasing SSTs and vice versa, which resembles fluctuations over glacial-interglacial timescales. We argue that the synchronicity between SST and SWWB changes during the last 1400 years represents the response of the SWWB to temperature changes in the Southern Hemisphere.

Sea-surface temperature reconstruction for sediment core GeoB1023-5

Sediment core GeoB 1023-5 from the eastern South Atlantic was investigated at high temporal resolution for variations of sea-surface temperature (SST) during the past 22 kyr, using the alkenone (UK'37) method. SSTs increased by 3.5°C from about 18°C during the Last Ice Age (21±2 cal kyr BP) to about 21.5°C at 14.5 cal kyr BP. This warming trend associated with the deglaciation phase was followed by a cooling event with lowest SSTs near 20°C, persisting for about 1000 years between 13 and 12 cal kyr BP. The SSTs then continued to increase to about 22.5°C at the Holocene climatic optimum at 7 cal kyr BP, and decreased again during the Late Holocene to a core-top value of 19.8°C that is comparable to modern annual mean SST values. When compared with alkenone SST records from the eastern North Atlantic, our SST record indicates continuous warming throughout the deglaciation phase in the Benguela Current, while its northern counterpart, the Canary Current, experienced prominent cooling during 'Heinrich Event 1' (H1). On the other hand, for the time period corresponding to the 'Younger Dryas' (YD) cooling event, the Benguela SST record exhibits a cold-temperature interval that corresponds to that observed in the eastern North Atlantic SST records. This observation suggests that interhemispheric climate response in Atlantic eastern boundary current systems was different with respect to the two abrupt climate events associated with Termination I. For the H1, the eastern South Atlantic SST record strongly supports the hypothesis that an 'anti-phase' thermal behavior in South Atlantic surface waters was forced by the slowdown of the North Atlantic Deep Water formation during cold spells in the North Atlantic. In contrast, the abrupt cooling in the eastern South Atlantic coincident with the YD period was probably induced by more vigorous global atmospheric circulation, enhancing the upwelling intensity in both eastern boundary current systems. This atmospheric control may have overridden any effect caused by changes in thermohaline circulation on the South Atlantic SSTs during the YD, which leads to the assumption that the thermohaline circulation was already much closer to its interglacial mode during the YD than during the H1.

(Table 1) AMS radiocarbon ages from ODP Sites 169-1033 and 169-1034

This paper explores the paleoseismic record potentially preserved in the upper 40 m of hydraulic piston cores collected in 1996 at two sites in Saanich Inlet, British Columbia, during ocean drilling program (ODP) Leg 169S. The ODP cores are missing 1-2 m of water-rich sediment directly underlying the seafloor, but this sediment is preserved in shorter piston cores collected in 1989 and 1991. The upper part of the ODP cores consists of rhythmically laminated (varved) marine mud with intercalated massive beds, interpreted to be debris flow deposits. Some of the debris flow deposits are linked to past earthquakes, including the 1946 Vancouver Island earthquake (M7.2), a great (M8-9) plate-boundary earthquake at the Cascadia subduction zone in January 1700, and a large crustal or plate-boundary earthquake about 1000 yr ago. Earthquakes may also be responsible for debris flows in about AD 1600, 1500, 1250, 1150, 850, 450, 350, 180, and BC 200, 220, 500, 900, and 1050. If so, the average recurrence interval for moderate to large earthquakes, which trigger debris flows in Saanich Inlet, is about 150 yr. This recurrence interval is broadly consistent with the frequency of moderate to large earthquakes in the region during the historical period. Debris flows, however, can also be triggered by non-seismic processes, making it difficult to assemble a complete earthquake record from the Saanich Inlet cores. We propose that extensive debris flow deposits, emplaced by single large failures or many smaller coincident failures, probably have a seismic origin.

Sedimentological, geochemical, micropaleontological and rock magnetic proxies of sediment core GeoB6211-2

Surface currents and sediment distribution of the SE South American upper continental margin are under influence of the South American Monsoon System (SAMS) and the Southern Westerly Wind Belt (SWWB). Both climatic systems determine the meridional position of the Subtropical Shelf Front (STSF) and probably also of the Brazil-Malvinas Confluence (BMC). We reconstruct the changing impact of the SAMS and the SWWB on sediment composition at the upper Rio Grande Cone off southern Brazil during the last 14 cal kyr combining sedimentological, geochemical, micropaleontological and rock magnetic proxies of marine sediment core GeoB 6211-2. Sharp reciprocal changes in ferri- and paramagnetic mineral content and prominent grain-size shifts give strong clues to systematic source changes and transport modes of these mostly terrigenous sediments. Our interpretations support the assumption that the SAMS over SE South America was weaker than today during most of the Late Glacial and entire Early Holocene, while the SWWB was contracted to more southern latitudes, resembling modern austral summer-like conditions. In consequence, the STSF and the BMC were driven to more southern positions than today's, favoring the deposition of Fe-rich but weakly magnetic La Plata River silts at the Rio Grande Cone. During the Mid Holocene, the northern boundary of the SWWB migrated northward, while the STSF reached its northernmost position of the last 14 cal kyr and the BMC most likely arrived at its modern position. This shift enabled the transport of Antarctic diatoms and more strongly magnetic Argentinean shelf sands to the Rio Grande Cone, while sediment contributions from the La Plata River became less important. During the Late Holocene, the modern El Niño Southern Oscillation set in and the SAMS and the austral tradewinds intensified, causing a southward shift of the STSF to its modern position. This reinforced a significant deposition of La Plata River silts at the Rio Grande Cone. These higher magnetic silts with intermediate Fe contents mirror the modern more humid terrestrial climatic conditions over SE South America.

Distribution of grain size and clay minerals in surface sediments of the Kara Sea (Fig 4, 5)

In this paper, we summarize data on terrigenous sediment supply in the Kara Sea and its accumulation and spatial and temporal variability during Holocene times. Sedimentological, organic-geochemical, and micropaleontological proxies determined in surface sediments allow to characterize the modern (riverine) terrigenous sediment input. AMS-14C dated sediment cores from the Ob and Yenisei estuaries and the adjacent inner Kara Sea were investigated to determine the terrigenous sediment fluxes and their relationship to paleoenvironmental changes. The variability of sediment fluxes during Holocene times is related to the post-glacial sea-level rise and changes in river discharge and coastal erosion input. Whereas during the late/middle Holocene most of the terrigenous sediments were deposited in the estuaries and the areas directly off the estuaries, huge amounts of sediments accumulated on the Kara Sea shelf farther north during the early Holocene before about 9 cal kyr BP. The maximum accumulation at that time is related to the lowered sea level, increased coastal erosion, and increased river discharge. Based on sediment thickness charts, echograph profiles and sediment core data, we estimate an average Holocene (0-11 cal kyr BP) annual accumulation of 194,106 t/yr of total sediment for the whole Kara Sea. Based on late Holocene (modern) sediment accumulation in the estuaries, probably 12,106 t/yr of riverine suspended matter (i.e. about 30% of the input) may escape the marginal filter on a geological time scale and is transported onto the open Kara Sea shelf. The high-resolution magnetic susceptibility record of a Yenisei core suggests a short-term variability in Siberian climate and river discharge on a frequency of 300-700 yr. This variability may reflect natural cyclic climate variations to be seen in context with the interannual and interdecadal environmental changes recorded in the High Northern Latitudes over the last decades, such as the NAO/AO pattern. A major decrease in MS values starting near 2.5 cal kyr BP, being more pronounced during the last about 2 cal kyr BP, correlates with a cooling trend over Greenland as indicated in the GISP-2 Ice Core, extended sea-ice cover in the North Atlantic, and advances of glaciers in western Norway. Our still preliminary interpretation of the MS variability has to be proven by further MS records from additional cores as well as other high-resolution multi-proxy Arctic climate records.