Biotic and abiotic controls of nitrogen and phosphorus cycling in Central European forests

The functioning and services of Central European forests are threatened by global change and a loss of biodiversity. Nutrient cycling as a key forest function is affected by biotic drivers (e.g., dominant tree species, understory plants, soil organisms) that interact with abiotic conditions (e.g., climate, soil properties). In contrast to grassland ecosystems, evidence for the relationship of nutrient cycles and biodiversity in forests is scarce because the structural complexity of forests limits experimental control of driving factors. Alternatively, observational studies along gradients in abiotic conditions and biotic properties may elucidate the role of biodiversity for forest nutrient cycles. This thesis aims to improve the understanding of the functional importance of biodiversity for nutrient cycles in forests by analyzing water-bound fluxes of nitrogen (N) and phosphorus (P) along gradients in biodiversity in three regions of Germany. The tested hypotheses included: (1) temperate forest canopies retain atmospheric N and retention increases with increasing plant diversity, (2) N release from organic layers increases with resource availability and population size of decomposers but N leaching decreases along a gradient in plant diversity, (3) P leaching from forest canopies increases with improved P supply from recalcitrant P fractions by a more diverse ectomycorrhizal fungal community. In the canopies of 27 forest stands from three regions, 16 % to 51 % of atmospheric N inputs were retained. Regional differences in N retention likely resulted from different in N availability in the soil. Canopy N retention was greater in coniferous than in beech forests, but this was not the case on loessderived soils. Nitrogen retention increased with increasing tree and shrub diversity which suggested complementary aboveground N uptake. The strength of the diversity effect on canopy N uptake differed among regions and between coniferous and deciduous forests. The N processing in the canopy directly coupled back to N leaching from organic layers in beech forests because throughfall-derived N flushed almost completely through the mull-type organic layers at the 12 studied beech sites. The N release from organic layers increased with stand basal area but was rather low (< 10 % of annual aboveground litterfall) because of a potentially high microbial N immobilization and intensive incorporation of litter into the mineral soil by bioturbation. Soil fauna biomass stimulated N mineralization through trophic interactions with primary producers and soil microorganisms. Both gross and net leaching from organic layers decreased with increasing plant diversity. Especially the diversity but not the cover of herbs increased N uptake. In contrast to N, P was leached from the canopy. Throughfall-derived P was also flushed quickly through the mull-type organic layers and leached P was predominantly immobilized in non directly plant-available P fractions in the mineral soil. Concentrations of plant-available phosphate in mineral soil solution were low and P leaching from the canopy increased with increasing concentrations of the moderately labile P fraction in soil and increasing ectomycorrhiza diversity while leaf C:P ratios decreased. This suggested that tree P supply benefited from complementary mining of diverse mycorrhizal communities for recalcitrant P. Canopy P leaching increased in years with pronounced spring drought which could lead to a deterioration of P supply by an increasing frequency of drought events. This thesis showed that N and P cycling in Central European forests is controlled by a complex interplay of abiotic site conditions with biological processes mediated by various groups of organisms, and that diverse plant communities contribute to tightening the N cycle in Central European forests and that diverse mycorrhizal communities improve the limited P availability. Maintaining forest biodiversity seems essential to ensure forest services in the light of environmental change.

Larix decidua and other larches in Europe: distribution, habitat, usage and threats

The European larch (Larix decidua Mill.) is a pioneer, very long-lived, fast-growing coniferous tree, which occurs in the central and eastern mountains of Europe, forming open forests or pasture woods at the upper tree limits. Larch is the only deciduous conifer in Europe as an adaptation to continental alpine climates. In fact, it is able to tolerate very cold temperatures during winter and, by losing its needles, avoids foliage desiccation. It is a transitional species, colonising open terrain after natural disturbances. It forms pure stands but more often it is found with other alpine tree species, which tend to replace it if no other disturbances occur. Thanks to its adaptability and the durability of its wood, the European larch represents an important silvicultural tree species in the alpine regions, planted even outside its natural ranges. Its wood is largely used for carpentry, furniture and pulp for paper. In lower altitudes or with high precipitation rates, larch is more susceptible to fungal diseases. Outbreaks of insect defoliators, principally caused by the larch bud moth (Zeiraphera diniana), can limit tree development, with economic losses in plantations, but they rarely lead to the death of the trees.

Picea abies in Europe: distribution, habitat, usage and threats

Among the coniferous species, Norway spruce (Picea abies (L.) Karst.) is one of the most important trees in Europe both for economic and ecological aspects, with a long tradition of cultivation. It can be a big tree, reaching 50-60 m in height with a straight and regular trunk, particularly used for timber constructions, pulpwood for paper and furniture. This widespread species dominates the Boreal forests in Northern Europe and the subalpine areas of the Alps and Carpathian Mountains. Thanks to its high performances in different site conditions, it can also be found outside its natural distribution on lower elevations in more temperate forests. Norway spruce has been massively planted up to its niche limits, where it is particularly susceptible to heat and drought, due to its shallow root system. For this reason it is expected to be severely affected under global warming conditions. Disturbed and weakened plants can be easily attacked by rot fungi such as Heterobasidion annosum and Armillaria, or by the bark beetles Ips typographus, one of the most destructive spruce forest pests.

Contrasting growth and water use efficiency after thinning in mixed Abies pinsapo–Pinus pinaster–Pinus sylvestris forests

Foresters frequently lack sufficient information about thinning intensity effects to optimize semi-natural forest management and their effects and interaction with climate are still poorly understood. In an Abies pinsapo–Pinus pinaster–Pinus sylvestris forest with three thinning intensities, a dendrochronologial approach was used to evaluate the short-term responses of basal area increment (BAI), carbon isotope (δ13C) and water use efficiency (iWUE) to thinning intensity and climate. Thinning generally increased BAI in all species, except for the heavy thinning in P. sylvestris. Across all the plots, thinning increased 13C-derived water-use efficiency on average by 14.49% for A. pinsapo, 9.78% for P. sylvestris and 6.68% for P. pinaster, but through different ecophysiological mechanisms. Our findings provide a robust mean of predicting water use efficiency responses from three coniferous species exposed to different thinning strategies which have been modulated by climatic conditions over time.

Comportamiento de coníferas bajo riego : Mendoza, Argentina

La provincia de Mendoza, ubicada en el centro-oeste de la Argentina, tiene una extensión de 150 830 km2. Prácticamente todas las actividades agropecuarias y forestales están concentradas en el 3 % de su territorio que es posible irrigar. Al oeste, en el límite con Chile, está la cadena montañosa que forma parte de la región fitogeográfica del Desierto Andino que se extiende por más de 500 km, con un ancho promedio de 100 km. El objetivo del trabajo fue introducir algunas especies de coníferas y evaluar su comportamiento en dicha región fitogeográfica, con la finalidad de lograr un aprovechamiento forestal, proteger el suelo contra la erosión hídrica y/o eólica, modificar el paisaje y desarrollar áreas de explotación turística. Las especies seleccionadas en esta primera etapa fueron: Cedrus deodara, Cupressus arizonica, Cupressus macrocarpa, Juniperus virginiana, Pinus griffithii, Pinus halepensis, Pinus nigra, Pinus pinaster, Pinus pinea y Pinus radiata. Las experiencias se llevaron a cabo en dossitios, separados por más de 150 km, y situados a 1 050 y 2 000 msnm, respectivamente. Se tomaron datos de supervivencia, altura y diámetro de los individuos, y se efectuaron observaciones sobre su hábito de crecimiento y condiciones fitosanitarias. Las principales conclusiones fueron las siguientes: 1. Todas las especies mostraron buen estado fitosanitario sin haberse detectado ninguna plaga o enfermedad de importancia. 2. El hábito de crecimiento fue el correspondiente a cada especie; sólo hubo fustes bifurcados en algunos individuos de Pinus pinea. 3. Las especies que evidenciaron mayor aptitud fueron: Juniperus virginiana, Cupressus arizonica, Pinus pinea, Pinus pinaster y Cupressus macrocarpa.

Pollen record from Golkow, Poland

Janczyk-Kopikowa (1966): The series of the organic deposits, developed in the vicinity of Golkow near Warsaw as oil shales and peats, was laid down in a grough valley and now rests on the deposits of the Middle Polish Glaciation (Riss). The organic deposits are overlain by the fluviale deposits of the North Polish Glaciation (Würm). The locality Golkow occurs beyond the extent of the continental glacier of this glaciation. Polen analysis completed by microfloristic examinations allows to determine the age of the organic series that is thought to be Eemian. The pollen diagram from Golkow does not call in question the stratigraphical position of the deposits investigated mainly due to its characteristic features such as minimum content of coniferous trees in the climatic optimum - about 5%, high percentage of Corylus - 77.5% and well developed phase of hornbeam. It may be well compared with other Eemian diagrams from the area of Poland and reveals much similar features. The development of vegetation at Golkow has depended upon the prevailing climate. At first, the cool climate brings about the development of plants having small thermal requirements. Here belong thin, park-like forests with pine and birch (Pinus, Betula) accompanied by the heliophilic plants such as Hippohäe and Ephedra. Improvement of climate that becomes warm and humid provides for development of deciduous forests prevailing in the climatic optimum, of the interglacial. Decrease of temperature causes a repeated change in the type of forest. This latter changes into coniferous forest with prevailing spruce (Picea) and fir (Abies) at the beginning, and then with pine (Pinus) and birch (Betula). During the Eemian Interglacial, the development of plants at Golkow terminates with a new and long-lasting predominance of pine-birch forests. However, such a longevity may be apparent only. Apparent character of this phenomenon is proved by a fact that the pollen spectra of the warm climatic periods have found their reflex in the oil shale that increased considerably slower than the layers off feebly decomposed peat evidencing the existence of cool pine-birch forests from the decline of the Interglacial. The water basin, in which the polen grains were laid down from surrounding plants is characterized by a calm sedimentation as proved by the occurrence of the oil shale. An insignificant water flow left behind some thin sand laminae. The not too deep basin becomes shallower owing to the growing water vegetation, and marshy vegetation. The growing of the plants causes a complete shallowing of the basin and formation of peat bog in situ, as proved by the peat beds occurring in the section. ---- Gadomska (1966): In the vicinity of Golków a series of organic deposits occurs amounting to 6.5-9.3 m in thickness, and consisting of oil shales, lacustrine silts and sands, as well as peats and peaty silts. The organic deposits fill up an old, small, but fairly deep lake basin, probably of finger-lake origin. It may be seen to-day as a slight lowering of the relief, filled up with soaked ground, stretching from north to south. On the basis of palaeobotanical examinations the organic deposits considered are of Eemian Interglacial age (Z. Janczyk-Kopikowa, 1063). The lower part of the organic series consists of a compact oil shale horizon, the maximum thickness of which may attain up to 8 m. The oil shales contain particularly in their upper part, numerous intercalations of arenaceous silts, dark grey or black in colour, or of sands mainly of lacustrine provenance. At the top of the oil shales are found peats, up to 2.5 m in thickness, covered by black, humus silts with numerous plant remains. The Eemian Interglacial deposits are covered by a series of fluviatile sands belonging partly to the Baltic Glaciation (bottom part of the series), partly to the Holocene (top part of the series). The thickness of the sands is 0.5-3.7 m. Higher up, there are found the Holocene and present-day deposits developed as clayey alluvion, or arenaceous slide rocks, or arenaceous-silty soil.

Pollen analysis of ODP Hole 117-720A (Appendix B)

Pollen analytical studies of the ODP Site 720 cores revealed the wide development of the coniferous forest, which mainly composed by Pinus, Picea, Abies and Cedrus deodara, along the Indus river since the early Pleistocene.

Palynology on sediment profile Tso Kar in Ladakh

Palynological investigation of a 410 cm long core section from Tso Kar (33°10'N, 78°E, 4527 m a.s.l.), an alpine lake situated in the arid Ladakh area of NW India at the limit of the present-day Indian summer monsoon, was performed in order to reconstruct post-glacial regional vegetation and climate dynamics. The area was covered with alpine desert vegetation from ca. 15.2 to 14 kyr BP (1 kyr=1000 cal. years), reflecting dry and cold conditions. High influx values of long-distance transported Pinus sylvestris type pollen suggest prevailing air flow from the west and northwest. The spread of alpine meadow communities and local aquatic vegetation is a weak sign of climate amelioration after ca. 14 kyr BP. Pollen data (e.g. influx values of Pinus roxburghii type and Quercus) suggest that this was due to a strengthening of the summer monsoon and the reduced activity of westerly winds. The further spread of Artemisia and species-rich meadows occurred in response to improved moisture conditions between ca. 12.9 and 12.5 kyr BP. The subsequent change towards drier desert-steppe vegetation likely indicates more frequent westerly disturbances and associated snowfalls, which favoured the persistence of alpine meadows on edaphically moist sites. The spread of Chenopodiaceae-dominated vegetation associated with an extremely weak monsoon occurred at ca. 12.2-11.8 kyr BP during the Younger Dryas interstadial. A major increase in humidity is inferred from the development of Artemisia-dominated steppe and wet alpine meadows with Gentianaceae after the late glacial/early Holocene transition in response to the strengthening of the summer monsoon. Monsoonal influence reached maximum activity in the Tso Kar region between ca. 10.9 and 9.2 kyr BP. The subsequent development of the alpine meadow, steppe and desert-steppe vegetation points to a moderate reduction in the moisture supply, which can be linked to the weaker summer monsoon and the accompanying enhancement of the winter westerly flow from ca. 9.2 to 4.8 kyr BP. The highest water levels of Tso Kar around 8 kyr BP probably reflect combined effect of both monsoonal and westerly influence in the region. An abrupt shift towards aridity in the Tso Kar region occurred after ca. 4.8 kyr BP, as evidenced by an expansion of Chenopodiaceae-dominated desert-steppe. Low pollen influx values registered ca. 2.8-1.3 kyr BP suggest scarce vegetation cover and unfavourable growing conditions likely associated with a further weakening of the Indian Monsoon.

Abundance and sizes of European oak species (Quercus petraea (Matt.) Liebl., Quercus robur L.) in dependence of neighborhood and environmental factors

Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.

Pedology, geochronology, palynology and macrofossils of soil profiles in the Altdarss area, german Baltic Sea coast

A new site with Lateglacial palaeosols covered by 0.8 - 2.4 m thick aeolian sands is presented. The buried soils were subjected to multidisciplinary analyses (pedology, micromorphology, geochronology, dendrology, palynology, macrofossils). The buried soil cover comprises a catena from relatively dry ('Nano'-Podzol, Arenosol) via moist (Histic Gleysol, Gleysol) to wet conditions (Histosol). Dry soils are similar to the so-called Usselo soil, as described from sites in NW Europe and central Poland. The buried soil surface covers ca. 3.4 km**2. Pollen analyses date this surface into the late Aller0d. Due to a possible contamination by younger carbon, radiocarbon dates are too young. OSL dates indicate that the covering by aeolian sands most probably occurred during the Younger Dryas. Botanical analyses enables the reconstruction of a vegetation pattern typical for the late Allerod. Large wooden remains of pine and birch were recorded.

Palynomorph concentrations in ODP Hole 105-645B samples (Table 1)

Significant changes in terrestrial and marine environments of Baffin Bay occurred throughout the late Pliocene to Holocene. Upper Pliocene and lowermost Pleistocene sediments contain abundant pollen and spores, which indicates the existence of open, coniferous, boreal forest to forest tundra in areas surrounding Baffin Bay. The late Pliocene-earliest Pleistocene also is characterized by relatively rich, dinoflagellate cyst and acritarch assemblages of boreal character, in which the dominance of Cymatiosphaera and Peridiniaceae may indicate neritic influx of sediments. In contrast, the late-early Pleistocene to Holocene interval is marked by a sparse terrestrial and marine palynoflora, with rare productive intervals. The general paucity of the Pleistocene palynoflora suggests low primary productivity, both on land and offshore, probably caused by cold, dry, high arctic conditions.

Determination of ages and fossil Coleoptera fauna from eight sites in the London region

These data sets report the fossil beetle assemblages identified from the Mesolithic to Late Bronze Age at eight sites in the London region. All but one of the study sites are within 2 km of the modern course of the Thames. The sites produced 128 faunal assemblages that yielded 218 identified species in 41 families of Coleoptera (beetles).  Beetle faunas of Mesolithic age indicate extensive wetlands near the Thames, bordered by rich deciduous woodlands. The proportion of woodland species declined in the Neolithic, apparently because of the expansion of wetlands, rather than because of human activities. The Early Bronze Age faunas contained a greater proportion of coniferous woodland and aquatic (standing water) species. An increase in the dung beetle fauna indicates the presence of sheep, cattle and horses, and various beetles associated with crop lands demonstrate the local rise of agriculture, albeit several centuries after the beginnings of farming in other regions of Britain. Late Bronze Age faunas show the continued development of agriculture and animal husbandry along the lower Thames. About 33% of the total identified beetle fauna from the London area sites have limited modern distributions or are extinct in the U.K. Some of these species are associated with the dead wood found in primeval forests; others are wetland species whose habitat has been severely reduced in recent centuries. The third group is stream-dwelling beetles that require clean, clear waters and river bottoms.

(Table 1) Tree age, post fire stand age, and soil organic matter thickness in the western Quebec black spruce boreal ecosystem

The observed long-term decrease in the regional fire activity of Eastern Canada results in excessive accumulation of organic layer on the forest floor of coniferous forests, which may affect climate-growth relationships in canopy trees. To test this hypothesis, we related tree-ring chronologies of black spruce (Picea mariana (Mill.) B.S.P.) to soil organic layer (SOL) depth at the stand scale in the lowland forests of Quebec's Clay Belt. Late-winter and early-spring temperatures and temperature at the end of the previous year's growing season were the major monthly level environmental controls of spruce growth. The effect of SOL on climate-growth relationships was moderate and reversed the association between tree growth and summer aridity from a negative to a positive relationship: trees growing on thin organic layers were thus negatively affected by drought, whereas it was the opposite for sites with deep (>20-30 cm) organic layers. This indicates the development of wetter conditions on sites with thicker SOL. Deep SOL were also associated with an increased frequency of negative growth anomalies (pointer years) in tree-ring chronologies. Our results emphasize the presence of nonlinear growth responses to SOL accumulation, suggesting 20-30 cm as a provisional threshold with respect to the effects of SOL on the climate-growth relationship. Given the current climatic conditions characterized by generally low-fire activity and a trend toward accumulation of SOL, the importance of SOL effects in the black spruce ecosystem is expected to increase in the future.

Pollen and non-pollen palynomorph records of the Holocene part of the composite sediment core TMD from lake Tso Moriri (NW India) and of modern surface samples from the Tso Moriri region

The high-altitude lake Tso Moriri (32°55'46'' N, 78°19'24'' E; 4522 m a.s.l.) is situated at the margin of the ISM and westerly influences in the Trans-Himalayan region of Ladakh. Human settlements are rare and domestic and wild animals are concentrating at the alpine meadows. A set of modern surface samples and fossil pollen from deep-water TMD core was evaluated with a focus on indicator types revealing human impact, grazing activities and lake system development during the last ca. 12 cal ka BP. Furthermore, the non-pollen palynomorph (NPP) record, comprising remains of limnic algae and invertebrates as well as fungal spores and charred plant tissue fragments, were examined in order to attest palaeolimnic phases and human impact, respectively. Changes in the early and middle Holocene limnic environment are mainly influenced by regional climatic conditions and glacier-fed meltwater flow in the catchment area. The NPP record indicates low lake productivity with high influx of freshwater between ca. 11.5 and 4.5 cal ka BP which is in agreement with the regional monsoon dynamics and published climate reconstructions. Geomorphologic observations suggest that during this period of enhanced precipitation the lake had a regular outflow and contributed large amounts of water to the Sutlej River, the lower reaches of which were integral part of the Indus Civilization area. The inferred minimum fresh water input and maximum lake productivity between ca. 4.5-1.8 cal ka BP coincides with the reconstruction of greatest aridity and glaciation in the Korzong valley resulting in significantly reduced or even ceased outflow. We suggest that lowered lake levels and river discharge on a larger regional scale may have caused irrigation problems and harvest losses in the Indus valley and lowlands occupied by sedentary agricultural communities. This scenario, in turn, supports the theory that, Mature Harappan urbanism (ca. 4.5-3.9 cal ka BP) emerged in order to facilitate storage, protection, administration, and redistribution of crop yields and secondly, the eventual collapse of the Harappan Culture (ca. 3.5-3 cal ka BP) was promoted by prolonged aridity. There is no clear evidence for human impact around Tso Moriri prior to ca. 3.7 cal ka BP, with a more distinct record since ca. 2.7 cal ka BP. This suggests that the sedimentary record from Tso Moriri primarily archives the regional climate history.