The term extractives defines chemical compounds of different classes that can be extracted from wood or bark by means of polar or non-polar solvents. Extractives are derived mostly from the metabolic processes of the tree, particularly the sapwood to heartwood transformation. The first objective of the research was to study the distribution of extractives within Sitka spruce trees at different heights in the trunk, as well as the distribution between bark, rootwood, knotwood, heartwood and sapwood. The second aim of the work was to learn about the influence of yield class, site elevation, North/East location and thinning on the extractives content and composition of Sitka spruce across Scotland. The samples were sawdust obtained in different ways from either discs, knots or roots sawn from Sitka spruce trees freshly cut in the forest, or collected during the coring of trees from 64 sites all around Scotland and northern England. The extraction was carried out on Soxhlet extractors using acetone as solvent. Two analytical techniques were used: gas chromatography and Fourier transform infrared spectroscopy. The results of the research showed that the extractive content and composition of Sitka spruce differed according to the type of wood studied with the largest amount detected in bark and the lowest in heartwood and sapwood. The last two types of wood were studied in more detail, showing that the difference in extractive content between heartwood and sapwood was consistent at all heights in the trunk. The chromatographic analysis of heartwood, sapwood, knotwood, rootwood and bark showed that their compositions differed slightly from published data on Norway spruce.

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This thesis contributes to the understanding of two of the most intriguing questions that forest ecologists have faced over recent decades: 1) how high diversity is maintained in species-rich ecosystems; and 2) what is the role of spatio-temporal environmental variation in structuring forest communities. The aims of the research were to ascertain how species composition varies both spatially and temporally and how changes in the vegetation can be understood in the context of species coexistence theories (niche versus neutral). A group of 38 sympatric species, including 9 species of Quercus, on which little ecological research has been undertaken, were used in this study. The data used in this project include eleven years of periodic remeasurements of permanent plots established in high-altitude oak forests in Mexico. Adult, sapling and seedling trees were studied as well as their environmental surrounding. Spatial and temporal variations in forest composition were analysed using multivariate statistical approaches. The results show that there are discrete communities in these mixed oak forests that correspond to specific environments. At a broad scale the study area can be classified into two floristic zones, a mesic zone characterised by associations that include Quercus candicans, Q. laurina and Q. castanea and; a xeric zone dominated by Q. crassipes. However of a finer scale of analysis important variation in composition was associated with different life stages of the trees, with adult trees showing much stronger environmental associations than seedlings and saplings. Successional pathways and rates vary at relatively fine scales. This may be as a result of dominance alternation between dominant canopy species. Micro-niche zonation processes caused by a high degree of environmental heterogeneity combined with individual species traits explain the coexistence of phylogenetically similar sympatric Quercus species. A hierarchy of processes, each acting at a different spatial and temporal scale, determines species diversity and coexistence. The overall findings support the idea that niche differentiation rather than chance events such as dispersal limitation, are more important in permitting species coexistence.

Acidification of surface waters has been recognised as the major water quality problem in the UK uplands. The adverse effects of conifer afforestation on streamwater chemistry and ecology are well documented in acid-sensitive catchments and have mainly been attributed to the enhanced deposition of atmospheric pollutants onto conifer canopies (the “scavenging effect”). Currently, international and national policies promote the expansion of native broadleaf woodland in the UK. Pollutant deposition onto broadleaf canopies is considered less than onto the more aerodynamically rough conifers, but there is concern that largescale broadleaf planting could delay the recovery of acidified waters or lead to further acidification in most sensitive areas. However, there has been limited investigation of the influence of broadleaf woodland cover on streamwater chemistry in the UK. To investigate the effect of woodland cover 14 catchments with different (0-78%) percentages of broadleaf woodland cover were identified in representative acidsensitive areas in north-western and central Scotland (Glen Arnisdale and Loch Katrine area) and northern and south-western England (Ullswater area and Devon) using spatial datasets in a GIS. Streamwater was sampled at high flow from the catchment outlets in winter and spring 2005 and 2006 and was analysed for major cations, anions and trace metals using standard methods. The number of samples ranged from two in the Glen Arnisdale catchments to 10 in the Loch Katrine area catchments which were sampled more intensively. Significant positive correlations were found between percentage broadleaf woodland cover and streamwater NO3 (rs = 0.51) and soluble Al (rs = 0.64) concentrations. The greater NO3 leaching to streamwater in the three most forested catchments (> 50%) was probably due to enhanced N deposition onto woodland canopies and nitrification by alder in the Ullswater area forested catchments. Streamwater NO3 concentrations equalled or exceeded non-marine SO4 in the above catchments indicating that NO3 was the principal excess acidifying ion in catchments with greater woodland cover. The woodland effect on streamwater chemistry in the study catchments was masked to some extent by variability in acid deposition climate and soil type composition. Seasalt inputs were found to be a more important control than woodland cover for streamwater chemistry in the maritime Glen Arnisdale catchments. A risk assessment of acid-sensitivity in the study catchments was conducted by calculating streamwater critical load exceedances using the Steady-State Water Chemistry (SSWC) and First-order Acidity Balance (FAB) models and modelled pollutant deposition for 1995-97 and 2002. Critical loads were exceeded by 0.01 to 1.74 keq H ha-1 yr-1 in two catchments which had woodland covers > 50% and in the Devon control catchment. The remaining 11 study catchments were assessed to be not at risk of acidification, probably due to significantly reduced non-marine S deposition from 1986 to 2001, but seasalt inputs to the Glen Arnisdale catchments might cause acidic streamwater episodes. Acid-sensitivity was also assessed using macroinvertebrates sampled in 11 of the study catchments and the results generally agreed with the critical load assessments. More detailed estimates of the enhancement of dry S and N deposition onto birchwoods in the Loch Katrine area catchments using calculated roughness length within FRAME showed that it posed no risk for streamwater acidification in these catchments because of the high rainfall environment. However, in acid-sensitive areas of the UK with lower rainfall and closer to major pollution sources, enhanced pollutant scavenging by broadleaf woodland canopies could pose a greater risk of acidification to freshwaters. The finding that almost all study catchments with woodland covers less than 30% are well protected from acidification suggests that this is a sensible threshold value for use in risk assessments of the effects of broadleaf woodland planting conducted within the Forests and Water Guidelines. The results of a sensitivity analysis of the Guidelines’ methodology, conducted using parameters such as numbers and timing of streamwater sampling, different runoff estimates and critical acid neutralising capacity values, showed that the Guidelines should be able to protect sensitive freshwaters from acidification in areas where broadleaf woodland is expanding.

Continuous cover forestry (CCF), an alternative forest management approach to clearfelling, is increasingly being adopted in the UK. It aims at enhancing stand structural diversity and favouring natural regeneration and subsequent seedling growth below the existing canopy of plantation forests. One area of limited knowledge is the critical level of below-canopy light for the growth of naturally regenerating seedlings. In addition, plant growth beneath canopies is influenced by other factors (e.g. herbivory). Picea sitchensis (Bong.) Carr. (Sitka spruce) seedlings under canopies have been observed to be severely damaged by Elatobium abietinum (Walker) (green spruce aphid) attacks. The combined effects of light availability and insect attack on seedling growth are not well understood, however, this understanding is crucial in order to ensure successful management of regeneration within CCF systems. A controlled experiment, which mimicked different natural understorey light levels along with repeated artificial aphid infestation, was conducted over two years to look at seedling performance through structural and physiological (chlorophyll fluorescence) measurements. Aphid population assessments showed significant increased population density under shaded conditions. Nevertheless, aphid impacts were mainly localised in extent while the impact of light was the major component that described seedling growth. Light was the primary factor affecting the whole-plant biomass, whereas aphids had only localised effects on the total dry weight of older needles and roots, and on leader extension growth. A significant interaction between light levels and presence/absence of aphid infestation was found for main leader extension growth of the seedlings during the second year of the experiment, with lower values at low light levels under infestation. Plant biomass allocation was affected strongly by light, while aphid presence did not result in significant changes. At the leaf physiologylevel, the light environment was found to be the main driving factor affecting photosynthetic response, whilst aphid presence had only a short-term localised effect on photosynthesis. The impact of light levels and aphid presence on seedling growth were also determined at an experimental field site where plots were located across two light regimes typical of CCF conditions in upland UK coniferous forests. The comparison of the photosynthetic response of the seedlings in both the controlled and field experiment highlighted the importance of considering the temporal heterogeneity of the light environment experienced by understorey seedlings in CCF stands, while aphid and aphid x light interaction effects could not be determined due to very low aphid levels during the year. Finally, a pre-existing light model was parameterised to predict the understorey light environment required to promote successful seedling growth in CCF stands. Sensitivity and validation analyses were also performed.

This study was conducted in the Afram Headwaters, Tain Tributaries Block II and Worobong South Forest Reserves. Satellite record of fire incidence for the country over 11 years (1997 to 2007) was modelled via binary logistic regression analysis, and correlations between fire incidence and the correlates of fire used to explain the observed trends. Fire incidence was found to be correlated with multiple variables which probably covary. Rainfall, vegetation type and geology showed the strongest correlations with fire incidence. Recurrent fire has impacted negatively on forest structure, ground cover biomass and species composition in two forest reserves, but more marked in the wetter Worobong South Forest Reserve than the drier Tain II Forest Reserve. Basal area has reduced from 40 m²ha^-1 in least-degraded to <1 m²ha^-1 in heavily-degraded stand in Worobong South Forest Reserve, along with tree density, whereas canopy openness has increased from 6% in least-degraded to 83% in heavily-degraded forest. In Tain II Forest Reserve, however, the heavily-degraded forest has lost close to 50% of its maximum value in terms of basal area, tree density and canopy cover, all in approximately 20 years. Early-burning, with maximum seedling height growth rates of 130 cm yr^-1 and 40 cm yr^-1 for Worobong South and Tain II Forest Reserves respectively, might help control the fires, and restore forest canopy in about 10 to 20 years if regularly maintained, but must be accompanied by green firebreaks. Complete protection from fire (during convalescence), on the other hand, would take between five and 15 years to restore forest canopy, but at huge resource cost. Implications for sustainable forest management are discussed.

Public goods such as environmental conservation, amenity and carbon sequestration are increasingly emphasised in forest policy agendas. However, many public benefits in woodlands have occurred incidentally, rather than on the basis of socio-economic logic and often at locations relatively inaccessible to major centres of population. In fact, data reveal a concentration of privately owned woodland in densely populated areas, especially in central and southern England, and that woodland is often factored into residential location decisions and lifestyle behaviours. However, the provision of public goods is likely to be contingent on the value systems of private forest and woodland owners and their flexibility of response to measures promoted under the devolved forest strategies. A combination of qualitative and quantitative research methods were employed to construct a robust typology of private woodland owners with respect to their willingness and ability to deliver public good benefits in three study areas in England: the Lake District, Cornwall and the High Weald AONB. Building on an exploratory scoping study, Q Methodology interviews were conducted with 10 woodland owners in each study area, followed by a self-completion survey, administered using Dillman's Total Design Method. Data from 600 woodland owners was subjected to a Factor and Cluster Analysis, with the emergent model validated using Discriminant Analysis. Six discrete private woodland owner types were revealed: Individualists, Multifunctional Owners, Private Consumers, Conservationists, Investors and Amenity Owners. Important distinctions between owner groups are associated with the likely provision of particular benefits and disbenefits, and the classification suggests that a move from a production versus consumption/protection framework to one that includes intersecting goals may be more appropriate. Policy implications are discussed to facilitate use of the typology in targeting particular woodland owner groups with more nuanced policy mechanisms, including incentive schemes, market mechanisms and advisory services.

The interactions between vegetation and climate are complex and critical to our ability to predict and mitigate climate change. Savanna ecosystems, unique in their structure and composition, are particularly dynamic and their carbon cycling has been identified as highly significant to the global carbon budget. Understanding the responses of these dynamic ecosystems to environmental conditions is therefore central to both ecosystem management and scientific knowledge. Longleaf pine ecosystems are highly biodiverse and unique savanna ecosystems located in the south-eastern USA – an important current carbon sink and key area identified for future carbon sequestration. These ecosystems depend on fire to maintain their structure and function, and the longleaf pine tree itself (Pinus palustris Mill.) has been noted for its resilience to drought, fire, pests and storms and is thus becoming increasingly attractive as both a commercial forestry species and a provider of other ecosystem services. Previous process-based models tested in the south-eastern USA have been shown to fail in conditions of drought or rapid disturbance. Consequently, in order to inform management and understand better the physiology of these ecosystems, there is a need for a process-based model capable of upscaling leaf-level processes to the stand scale to predict GPP of longleaf pine savannas. P.palustris exists across a wide range of soil moisture conditions, from dry sandy well-drained soils (xeric) to claypan areas with higher moisture content (mesic). Previous work has demonstrated that this species adjusts many aspects of its physiology in response to these differing soil conditions, even under identical climate. The research in this thesis supports these previous findings, but additionally explores, with the assistance of the Soil Plant Atmosphere model (SPA), the productivity response of P. palustris across the soil moisture gradient. Contrary to expectations, measurements, field observations and modelling suggest that P. palustris trees growing in already water-limited conditions cope better with exceptional drought than their mesic counterparts. At the leaf-level, xeric P. palustris trees were found to have higher measured net photosynthesis, but the lower stand density and leaf area at this site meant that in non-drought conditions mesic P. palustris annual gross primary productivity (GPP) was 23% greater than xeric annual GPP. Initial upscaling of leaf-level processes to the canopy scale using the SPA model found that, during the growing season when other components of longleaf pine ecosystems are active, the longleaf pine may only be responsible for around 65% of the total productivity. Other important components of longleaf pine savannas are oaks and grasses which, with pine, constitute 95% of longleaf pine ecosystem biomass. Each of these groups, however, responds differently to fire and water availability. Despite this, the other components of longleaf pine savannas have received limited research attention and have never been modelled using a process-based model such as SPA. As integral components of longleaf pine carbon budgets, it is essential that the physiology and productivity of oaks and grasses in this system are better understood. The research in this thesis studied the productivity response of these groups during drought across a soil moisture gradient, and found that oak and pines at each site appear to fill separate ecohydrological niches depending on whether or not they are growing in a xeric or mesic habitat. As expected, the highest drought tolerance was found in the C4 grass, wiregrass (Aristida stricta), at both xeric and mesic sites. In order to further explore the contributions of the different functional groups in longleaf pine savannas, the SPA model was adapted to run with concurrent functional groups and to represent the different photosynthetic pathways of the understorey grasses (C4) and the canopy trees (C3). The aim of this part of the thesis was to represent better a savanna ecosystem in a process-based model and explore and quantify the contributions of each functional group diurnally, seasonally, annually and interannually. Modelling results suggest that accurately representing the phenology not only of trees but of grasses, is critical to capturing ecosystem GPP and its variability. This phenology may not only be seasonally controlled, but also dictated by fire. Overall, this research highlights the importance of continued research into savanna and savanna-like ecosystems. Additionally, it provides an insight into the responses of multiple ecosystem components to an extreme drought, and how these responses differ at leaf, stand and landscape scales. The thesis also employs a little-used method of combining eddy-covariance data with a process-based model to separate out different ecosystem components, a method becoming more common but not yet widely tested.

An investigation of the effects of commercial mechanised selective logging on rain forest vegetation and mammals, was undertaken in the Lope Reserve, central Gabon, between January 1989 to July 1991. Vegetation in Lope is mostly semi-evergreen lowland tropical rain forest, but there are some localised patches of savanna., which are thought to be natural in origin, but which are maintained today by regular fires started by humans. Study sites were established in areas of forest logged 20-25, 10-15, and 3-5 years previously, a fourth was logged during the study, and a fifth remained unexploited. None of the study sites had been subject to hunting in the recent past. A line-transect five kilometres in length was cut across the drainage in each site. Forest composition and structure was assessed along each transect, by identifying and measuring trees and lianes in botanical plots, counting stem density of herbaceous vegetation in the families Marantaceae and Zingiberaceae, and by measuring canopy cover at three heights. At total of 4885 trees and lianes of 327 species occurred in five 2.5 ha samples of plants greater than 10 cm dbh, whilst 1832 individuals of 137 species were found in five 25ha samples of trees greater than 70cm dbh. There were marked differences in structure and species composition both between and within sites. A model was developed to attempt to explain this variation, based upon the theory that much of Lope had been covered by savanna vegetation during a previous cool, dry climatic phase, and that forest structure and composition reflected recolonisation of the savanna by forest. Physical features such as swamps, rocky outcrops and altitude were also considered. Two types of multivariate analysis were applied to botanical data and supported the model. The effects of logging on forest vegetation were assessed by returning after logging to botanical plots established before exploitation. Damage levels were low, compared to other parts of the World, resulting in about a 10% reduction in canopy cover. Patterns of fruit production were studied by counting fallen ripe and unripe fruit on transects. Fruits encountered were classified on the basis of their morphology and dispersal mechanism. There was a period of low fruit production during the major dry season, when frugivores are likely to suffer dietary stress. A number of plant species which did produce fruit at this time were identified as species which might represent 'keystone' resources. Over 70% of fruit species were animal-dispersed, demonstrating the the important role animals play in the ecology of tropical rain forests. Forest elephant (Loxodonta africana cyclotis) diet, ecological role and group structure were analysed. The bulk of the diet consisted of the bark and leaves of trees, and some monocotyledons in the families Marantaceae and Zingiberaceae, but fruit was also important. Large-scale seasonal movements in response to fruit availability were detected. Elephants were important seed dispersers f o r many plant species, and were responsible for less than 1% of natural tree mortality. The social structure of these forest elephants differed from that of populations that have been studied i n east and southern Africa. Average group size was 2.8, and no groups of more than 10 individuals were encountered. Densi ties of primates, ungulates and squirrels were assessed using standard line-transect censuses. Resolution was poor, but statistical differences were detected between sites for some species. Chimpanzee, (Pan t. troglodytes) densities declined in logged forest, but no other species could be shown to decline after logging. Some other differences between sites were related to vegetation composition. Biomass was high, estimated a t up to 4692.6 kg km-2, but was dominated by elephants, which made up 25-82%. Conservation implications of this study are discussed, and recommendations made.

Genetic differentiation in phenotypic traits among populations from heterogeneous environments is often observed in common-garden studies on forest trees, but data on adaptive variation in Scots pine (Pinus sylvestris L.) in Scotland are limited. As a result, current seed transfer guidelines are based on earlier molecular marker studies and do not take into account environmental or adaptive genetic variation. An analysis of spatial variation in climate showed substantial differences in temperature and precipitation among the native Scots pine sites in Scotland. To investigate whether differentiation in response to environmental variation has occurred in Scotland, a glasshouse-based common-garden trial of ~3,360 seedlings from 21 populations and 84 open-pollinated families was established in 2007. At the beginning of the 2nd growing season, timing of bud flush showed evidence of genetic differentiation among populations, with those from cooler origins generally flushing earlier. Variation was also found among families within populations, suggesting that the trait is genetically controlled. Populations and families showed different levels of variability in this trait which could be partly due to variable levels of temporal climate fluctuation in different parts of Scotland. Chlorophyll fluorescence was used to examine drought response in three-year old seedlings from five populations on sites that experience contrasting levels of annual rainfall. It was found that the response was not related to rainfall, but possibly to more complex moisture variables that also take into account additional factors such as evaporation. Also, photosynthetic capacity in response to cold winter temperatures varied significantly among eight populations that were kept outdoors, and the largest reduction was seen in seedlings from the mildest, most maritime coastal site. The following spring, height growth and needle flush started earlier in seedlings from cooler locations. Earlier studies on genetic diversity of native pinewoods have shown high levels of selectively neutral variation in this predominantly outcrossing conifer, and a mating system analysis with a limited number of microsatellite markers supported this pattern. Together, these data suggest that despite significant historic population size decrease, environmental gradients have resulted in genetic differentiation among native pinewoods. In order to minimise the risk of planting poorly-adapted stock and to maximise the success of replanting programmes, it is important that the origins of planting stock are carefully considered in management guidelines for the species.