The successful implementation of habitat preservation and management demands regular and spatially explicit monitoring of conservation status at a range of scales based on indicators. Woodland condition can be described in terms of compositional and structural attributes (e.g. overstorey, understorey, ground flora), evidence of natural turnover (e.g. deadwood and tree regeneration), andanthropogenic influences (e.g.disturbance, damage). Woodland condition assessments are currently conducted via fieldwork, which is hampered by cost, spatial coverage, objectiveness and repeatability. This project evaluates the ability of airborne remote sensing (RS) techniques to assess woodland condition, utilising a sensor-fusion approach to survey a foreststudy site and develop condition indicators. Here condition is based on measures of structural and compositional diversity in the woodland vertical profile, with consideration of the presence of native species, deadwood, and tree regeneration. A 22 km2 study area was established in the New Forest, Hampshire, UK, which contained a variety of forest types, including managed plantation, semi-ancient coniferous and deciduous woodland. Fieldwork was conducted in 41 field plots located across this range of forest types, each with varying properties. The field plots were 30x30m in size and recorded a total of 39 forest metrics relating to individual elements of condition as identified in the literature. Airborne hyperspectral data (visible and near-infrared) and small footprint LiDAR capturing both discrete-return (DR) and full-waveform (FW) data were acquired simultaneously, under both leaf-on and leaf-off conditions in 2010. For the combined leaf-on and leaf-off datasets a total of 154 metrics were extracted from the hyperspectral data, 187 metrics from the DR LiDAR and 252 metrics from the FW LiDAR. This comprised both area-based and individual tree crown metrics. These metrics were entered into two statistical approaches, ordinary least squares and Akaike information criterion regression, in order to estimate each of the 39 field plot-level forest variables. These estimated variables were then used as inputs to six forest condition assessment approaches identified in the literature. In total, 35 of the 39 field plot-level forest variables could be estimated with a validated NRMSE value below 0.4 using RS data (23 of these models had NRMSE values below 0.3). Over half of these models involved the use of FW LiDAR data on its own or combined with hyperspectral data, demonstrating this to be single most able dataset. Due to the synoptic coverage of the RS data, each of these field plot variables could be estimated and mapped continuously over the entire study site at the 30x30m resolution (i.e. field plot-level scale). The RS estimated field variables were then used as inputs to six forest condition assessment approaches identified in the literature. Three of the derived condition indices were successful based on correspondence with field validation data and woodlandcompartment boundaries. The three successful condition assessment methods were driven primarily by tree size and tree size variation. The best technique for assessing woodland condition was a score-based method which combined seventeen inputs which relate to tree species composition, tree size and variability, deadwood, and understory components; all of whichwere shown to be derived successfully from the appropriate combination of airborne hyperspectral and LiDAR datasets. The approach demonstrated in this project therefore shows that conventional methods of assessing forest condition can be applied with RS derived inputs for woodland assessment purposes over landscape-scale areas.

Coastal ecosystems are undergoing unprecedented rates of environmental change. Many of these changes are anthropogenically-driven and linked to long-term, climate-related phenomena. This thesis focusses on ecological change in soft sediment intertidal habitats. One of the largest harbours in Europe, Poole Harbour, is used as a case study. It contains a variety of important habitats including intertidal mudflat and non-tidal saline lagoon. The two main themes of the thesis are 1) assessing the physical and ecological factors that determine benthic invertebrate abundance, distribution and community structure, which is examined at the scale of the whole harbour, and at the scale of individual habitats: an intertidal mudflat and a saline lagoon; and 2) predicting the response of an overwintering shorebird population, the pied avocet (Recurvirostra avosetta), to future environmental changes, such as sea-level rise and habitat loss. This is achieved by development of an individual-based model (IBM) and consideration of the species’ unique foraging behaviour. This study contributes to the understanding of the factors structuring soft sediment benthic communities, including the use of data from fine-scale hydrodynamic models. It offers a unique comparison of the spatial and temporal variables driving community structure of a saline lagoon and an intertidal mudflat. It also provides insight into the foraging ecology of the pied avocet at a level of detail that has not previously been considered, including a comparison of foraging behaviour in a tidal and non-tidal habitat, the importance of social foraging, and the novel application of an IBM to this species.

1. The functional response, i.e. the feeding rate as a function of prey density, links resource availability to population level demographic rates. The functional response is often central to behavioural-based population models that predict the effect of environmental changes on populations based on the assumption that each individual tries to maximise its fitness at all times. 2. The functional response is a well studied phenomenon for some groups of foragers and many mechanistic models of the functional response have been derived to account for variations in foraging behaviour. However, only a few functional response models have been applied to granivorous farmland birds and these models often have simplifying assumptions that are likely to be inappropriate for these foragers. Consequently, the functional response models that often lie at the core of behavioural-based population models are untested and quite possibly provide a poor prediction of the functional response. 3. The objectives of this thesis are to: (i) test the ability of functional response models to predict the functional response of granivorous birds; (ii) identify incorrect assumptions in these models; and (iii) derive new functional response models that better describe the observed behaviour of these foragers. 4. Here I have found that many of the underlying assumptions of simple functional response models are inappropriate for granivorous birds and several new mechanistic models have been derived that attempt to describe the behaviour more accurately. Vigilance was shown to frequently interact with incompatible behavioural processes (i.e. searching) and affect the feeding rate at low prey densities. Handling time was divided into two components (i.e. attack and process) and a simple mechanistic model was derived to predict the attack component of prey handling. The searching mode of granivorous birds was shown to be akin to a pause-travel forager at low prey densities and, lastly, habitat structure was shown to reduce the feeding rate by restricting the searching area. 5. These finding will help in the development of more biologically 'realistic' functional response models and provide a more accurate prediction of the functional response when used to predict the effect of environmental change on the intake rate of foragers. z

Among British birds, a number of woodland specialists have undergone a serious population decline in recent decades, for reasons that are poorly understood. The Marsh Tit is one such species, experiencing a 71% decline in abundance between 1967 and 2009, and a 17% range contraction between 1968 and 1991. The factors driving this decline are uncertain, but hypotheses include a reduction in breeding success and annual survival, increased inter-specific competition, and deteriorating habitat quality. Despite recent work investigating some of these elements, knowledge of the Marsh Tit’s behaviour, landscape ecology and habitat selection remains incomplete, limiting the understanding of the species’ decline. This thesis provides additional key information on the ecology of the Marsh Tit with which to test and review leading hypotheses for the species’ decline. Using novel analytical methods, comprehensive high-resolution models of woodland habitat derived from airborne remote sensing were combined with extensive datasets of Marsh Tit territory and nest-site locations to describe habitat selection in unprecedented detail. Further fieldwork established the causes and frequency of breeding failure at the local population scale, and dispersal distances and success were quantified. Information from these studies was used to inform national-scale spatial analyses of habitat distribution in relation to the pattern of range contraction for the Marsh Tit and two other woodland bird species. The combined results indicate that Marsh Tits require extensive areas of mature woodland in order to accommodate large territories and short dispersal distances, with greatest selection for a woodland structure encompassing a tall, near-closed tree canopy and extensive understorey. The evidence suggests that nest-site competition, nest predation or deteriorating habitat quality have not driven the population decline. However, reduced connectivity between woodlands in the landscape, possibly due to hedgerow loss, may have interacted with increased mortality to precipitate population declines and local extinctions where habitat fragmentation was relatively high. The potential causes of increased mortality are discussed, along with priority areas for future research and the wider possible applications of remote sensing techniques in the field of woodland bird research.

Mammalian scavenging, disarticulating, scattering and removal of human remains can alter and obscure both soft tissue and skeletal remains which are essential to making interpretations and identifications during forensic investigations. The effects of scavenging vary between regions, environments, scavenger species, and crime scene scenarios due to a variety of factors. Nonetheless, there is a gap in the knowledge of scavenger species found within Northwestern Europe. The red fox (Vulpes vulpes) and Eurasian badger (Meles meles) are the largest wild mammalian scavenger species inhabiting peri-urban and rural environments within Northwestern Europe. These mammalian scavengers have dentitions and bite forces capable of heavily modifying and widely transporting human remains yet there are currently no species-typical and region-specific studies of these scavengers and their impacts on forensic investigations and physical searches for human remains. Forensic scientists, investigators and police search officers have been forced to rely on anecdotal evidence and scavenging studies focused on scavengers not found in this region. Scavenging studies have previously concentrated on scavenger species found in North America and Africa, such as coyote (Canis latrans), wolf (Canis lupus), hyena (Crocuta crocuta), lion (Panthera leo) and leopard (Panthera pardus), which have differing species-typical scavenging behaviour and patterns in comparison to foxes and badgers. Likewise, knowledge of the characteristics of the effects on bone surfaces of fox and badger scavenging is lacking, more so for the latter scavenger. The overall aim of this thesis is to aid forensic investigations by filling the gaps in the knowledge and identification of red fox and Eurasian badger species-typical scavenging behaviour and patterns. Avian scavenging can also modify soft tissue and skeletal remains. The buzzard (Buteo buteo) and carrion crow (Corvus corone) are the most common avian scavengers within this region. The scavenging behaviours of these avians modified soft tissue and affected mammalian scavengers’ scavenging behavours. A survey of police search officers within the U. K. indicated that the scavenging of surface deposited human remains within this region is common and that scavenging affects the recovery rates of remains. Despite the impact of scavenging on the recovery of scavenged remains, there is a lack of knowledge and literature available to forensic scientists, investigators, and police search officers to aid in the identification of scavenger species and scavenger species-typical scavenging behaviour and patterns. Thus these forensic professionals have been relying primarily on anecdotal evidence to identify scavengers or have not made efforts to identify scavengers. Experiments, conducted in southern England, using deposited deer (Cervus nippon; Capreolus capreolus) and the observation of captive scavengers found that within a woodland environment common scavengers include wood mouse (Apodemus sylvaticus), grey squirrel (Sciurus carolinensis), carrion crow, buzzard, Eurasian badger, and red fox. Scavenging activities by all scavenger species observed at remains were affected in various ways by seasonality, trophic resources, territoriality, insect activity, carcass size and condition, and decomposition. Of those scavengers, the red fox was the most frequent scavenger of surface deposited remains. The species-typical scavenging behaviour and pattern, as well as bite mark dimensions, of the red fox proved to differ to that of badgers and other canids, such as domestic dogs (Canis familiaris), coyotes and wolves. The benefits of the knowledge of scavenger species-typical scavenging behaviour and pattern to forensic investigations and physical searches were assessed by applying the results gained from the experiments within this research to current forensic investigations and search exercises performed with police search officers. The application of information on species-typical scavenging behaviour and patterns was found to improve police search officers’ search and recovery efforts of scavenged remains.

Secondary tropical forests are increasingly recognized for their role conserving biodiversity in agricultural landscapes and this role is especially important for seasonally dry tropical forests (SDTF), one of the most threatened tropical forested ecosystems. The conservation value of secondary forest is increased by its capacity to maintain ecosystem properties and provide services to humans; which has been hypothesized to have positive links to the species and functional diversity of ecosystems. However very little information is available on the occurrence of this relationship in secondary forests. This dissertation makes an important contribution to the ecological knowledge of secondary SDTF and describes changes in plant species and functional diversity by using a stratified design considering different successional stages along an environmental gradient in a region of the Caribbean coast of Colombia and a multi-trait approach to study functional diversity at three scales: species, communities and landscape. The analysis of the variation in functional traits of SDTF trees at the species level allowed me to support the hypothesis of coordination between leaves and stem traits. Three dimensions of correlated variation were identified: the first related to leaf and stem economy, the second to leaf thickness and organization and the third to the trade-offs between leaf size, stem density and bark thickness. Secondary forests showed high species turnover during succession and increasing diversity from early to late forests. Species composition similarity was higher among late successional forest than early and intermediate stage forests, showing that environmental characteristics are influencing successional trajectories. Frequency distributions of species in the three successional stages showed evidence of functional trait similarity among species and underlined the importance of changes in species abundances determining functional composition during succession. A shift in abundance from individuals with traits that favour survival after disturbance to individuals with denser stems and a more conservative resource use profile was observed from early to late stages of succession. Functional composition was also strongly influenced by environmental variables, especially leaf traits, and a shift of traits from acquisitive to conservative type species was observed with increasing nutrient content and flooding, proxies of water availability. Assessment of ecosystem services was conducted using two independent approaches: semistructured interviews and field data. A high richness of useful woody plants was recorded and the provision of services was related to a high variability in functional traits among species. The provision of the ecosystem services determined by the presence and abundance of useful species showed significant differences between stages. The relationships observed between ecosystem sevices and functional and species diversity indices were not consistent. In contrast species richess showed significant negative relationships at the plot level with most of the ecosystem services assessed, showing a trade-off among the conservation of species richness and the maintenance of service provision. Overall, this research provides novel insights into the dynamic relationships between biodiversity, ecosystem function and ecosystem services in this globally important, but under-researched forest type.

The impacts of environmental change on ecological communities are poorly understood relative to impacts on species. Impacts on inter-community (beta-) diversity are particularly neglected. As a result, our ability to forecast the impacts of environmental change on communities, and on individual species constrained by those communities, is seriously limited. However, as conservation efforts increasingly emphasise broad-scale approaches in terms of multi-species coverage and spatial scale, it is imperative that understanding ofbiodiversity change at these scales is enhanced so that conservation can be based on appropriate scientific evidence. Within this thesis I aimed to conduct multi-species analyses over multi-decada1 temporal scales at the spatial meso-sca1e to improve our understanding of such issues in both terrestrial and marine ecosystems. I discussed my findings in the theoretical context of G1easonian and C1ementsian views of species distributions (i.e. limitations to individualistic species responses) and their impact at the community scale. The temporal extent provided the opportunity to empirically test emerging concepts, including non-analogous communities, biotic homogenization, metacommunities and climate tracking responses at an appropriate meso-sca1e. Although no evidence for nonanalogous communities was found, biotic homogenization was supported and appeared to be caused by increased nitrogen and decreased light availability. An intertidal assemblage also converged but appeared to be driven by a reduced sea surface temperature gradient. Woodland plant metacommunity structure was demonstrated to be C1ementsian for woodland plants despite experiencing biodiversity loss. Hydrodynamic features were demonstrated to act as meso-sca1e dispersal barriers that limited intertidal invertebrates in tracking of climate (sea surface temperature) over the last 20 years. These barriers appear to alter when modelled under a scenario of sea level rise. Overall, results suggest that species are responding individualistically but that these responses are bounded by extrinsic constraints.

This volume and supporting papers constitute the submission for an award of a PhD research degree by publication. Eleven works completed by the author within the last 15 years (eight published; three in press) have been submitted for consideration. All the papers are concerned with animal exploitation in late prehistoric and Roman Britain and/or Medieval north-west Russia. To put these submissions into context, Chapter 2 summarizes the author’s academic career and the history of the research projects with which he has been involved. The next two chapters provide summaries and critically evaluative comments concerning the submitted works. Chapter 3 discusses the works concerned with the exploitation of animals and their products in the late Iron Age and Roman periods in Britain. Chapter 4 considers papers principally concerned with the exploitation of animals within the Medieval town and territory of Novgorod in north-west Russia. Chapter 5 presents an evaluation of the contribution the submitted works have made to furthering knowledge, not only of the specific periods and regions involved, but also more generally to the development of urban zooarchaeology (including comparisons between urban and rural faunal assemblages), the study of carcass processing, and the integration of zooarchaeology into general research questions.

Current woodland restoration programs are increasingly focussing on the creation of habitat networks in order to increase woodland cover and connectivity. However, the basic assumptions underpinning such strategies are largely untested for species associated with woodland habitat. For many woodland invertebrate species, local scale processes are potentially more important than processes operating at the landscape scale in terms of species persistence, especially for those species that show high dependence on woodland habitat conditions and have limited dispersal ability. The applicability of landscape-scale approaches to woodland restoration therefore needed to be evaluated in relation to the ecological characteristics of invertebrates. This thesis examines these issues and provides a quantitative analysis of the factors influencing presence of wood cricket (Nemobius sylvestris) at multiple scales within the landscape. The investigation was conducted in woodland habitats on the Isle of Wight in the south of the United Kingdom. A landscape-scale survey indicated that wood cricket was found predominantly in large woodland fragments situated in close proximity to each other, with ancient woodland characteristics and with a high amount of edge habitat. The current pattern of distribution of wood cricket suggested that most woodland fragments in the agricultural matrix are effectively isolated from each other, indicating the importance of maintaining a high level of connectivity between habitats for this invertebrate species. An investigation within woodlands indicated that locations with permanent low cover of ground vegetation, low canopy closure and high availability of leaf litter were the preferred habitat conditions for wood cricket. Ride and track edges, woodland peripheries and open areas created and maintained by management activities were found to be the main habitat locations for wood cricket. It was further found that wood cricket was mainly present at permanent edges or in close proximity to these locations, indicating the importance of maintaining these habitat features for this species. The mean dispersal rate for dispersing wood cricket obtained from a series of field experiments was found to be similar to that of other ground-dwelling invertebrate species that were strongly associated with woodland. This level of habitat specialism was consistent with the habitat preferences found for wood cricket, and therefore wood cricket can be seen as representative of this particular group of wood land-associated invertebrates. Comparable to wood cricket, the dispersal ability for species of this group was found to be limited. Few individuals of nymph (i. e. juvenile) and adult wood cricket populations were found to disperse. Wood cricket was found able to disperse up to 55 m into non-woodland habitat and mature habitat corridors were found to be used by wood cricket, but not new immature woodland plantings. The results of this investigation indicate that the overall success of woodland conservation for woodland invertebrates lies in adopting a multi-scale and multimanagement strategic approach. The current initiatives focussing on restoration and re-instatement of traditional management activities within existing woodlands were found to be highly beneficial for wood cricket. Corridors were found to facilitate movement if suitable woodland habitat conditions were provided. Creation of woodland habitat networks might therefore be beneficial for wood cricket if given enough time to develop.

Testing the correlation of morphological and genetic marker variation enables the investigation of evolutionary processes. Knowledge of evolutionary processes can be used to identify those morphological characters that could be used to produce evolutionary meaningful taxonomies. This thesis aims to test the correlation between morphological and genetic marker variation to further understand the evolution of species within the genus Arum and identify those morphological characters that correspond with evolutionary groups. The investigation is carried out at the intraspecific level, intrageneric level and in a putative hybrid zone. At the intraspecific level, genetic (ISSR) and morphological variation was quantified in populations of the morphologically similar species A. maculatum and A. italicum. Populations of A. maculatum showed evidence of isolation by distance, presumably a result of pollinator behaviour and seed dispersal. Leaf patterning in A. maculatum did not correspond to evolutionary lineages. However, similar leaf patterning characters in A. italicum are used to classify the two subspecies neglectum and italicum and the ISSR analysis confirmed that these taxa are genetically distinct. These two subspecies were shown to be interbreeding in sympatric populations. The interbreeding has created a morphological and genetic difference between subsp. neglectum in sympatric populations compared with allopatric populations. At the intrageneric level, a phylogenetic analysis of Arum (using trnL and ITS 1 sequences) indicated that both vegetative and reproductive characters are convergent within the genus. The apparent convergent evolution of reproductive and vegetative characters indicates that both have been important during the diversification of the genus. These convergent characters are not useful for producing classifications that reflect evolutionary groups as the groups they produce are polyphyletic. In the putative hybrid zone, ISSR markers confirmed the presence of A. creticum and A. idaeum hybrids. There appears to be introgression of the A. idaeum genome into A. creticum; this could have implications for the future genetic integrity of A. creticum. Within this hybrid zone, continuous characters were found to be representative of genetic variation, however categorical characters were not. In conclusion, this thesis has shown that even within a single genus, the correlation between morphological and genetic marker variation is influenced by both the taxa being studied and the nature of the morphological trait. In particular, if morphological characters are found to be adaptively important, their correspondence to genetic groups should be tested before their use in taxonomies. The findings of this thesis also suggest there is great value in the complementary use of genetic and morphological analysis for taxonomic studies as well as evolutionary studies. For example, the importance of reproductive characters in the diversification of Arum species has produced a wide range of morphological variation, with limited taxonomic utility due to a tendency for homoplasy. Vegetative characters were also found to need careful testing before use in taxonomies as leaf patterning was found to correspond to sub-species status for one species of Arum but not another. Finally, this thesis has shown that, if closely related taxa are hybridising, variation of continuous reproductive characters may be used as an indicator of hybridisation, even if the morphological characters are potentially polygenic.