Ecosystems are often made up of interactions between large numbers of species. They are considered complex systems because the behaviour of the system as a whole is not always obvious from the properties of the individual parts. A complex system can be represented by a network: a set of interconnected objects. In the case of ecological networks and food webs, the objects are species and the connections are interactions between species. Many complex systems are dynamic and exhibit intricate time series. Time series analysis has been developed to understand a wide range of natural phenomena. This thesis deals with the structure, dynamics, and robustness of ecological networks, the spatial dynamics of fluctuations in a social system, and the analysis of cardiac time series. Biodiversity on Earth is decreasing largely due to human-induced causes. My work looks at the effect of anthropogenic change on ecological networks. In Chapter Two, I investigate predator adaptation on food-web robustness following species extinctions. I identify a new theoretical category of species that may buffer ecosystems against environmental change. In Chapter Three, I study changes in parasitoid-host (consumer-resource) interaction frequencies between complex and simple environments. I show that the feeding preferences of parasitoid species actively change in response to habitat modification. Ecological networks are embedded in spatially-heterogeneous landscapes. In Chapter Four, I assess the role of geography on population fluctuations in an analogous social system. I demonstrate that fluctuations in the number of venture capital firms registered in cities in the United States of America are consistent with spatial and temporal contagion. Understanding how physiological signals vary through time is of interest to medical practitioners. In Chapter Five, I present a technique for quickly quantifying disorder in high frequency event series. Applying the algorithm to patient cardiac time series provides a rapid way to detect the onset of heart arrhythmia. Increasingly, answers to scientific questions lie at the intersection of traditional disciplines. This thesis applies techniques developed in physics and mathematics to problems in ecology and medicine.

Biologists have long used demographic approaches to answer questions in ecology and evolution. The utility of these approaches has meant a constant development and refinement of methods. A key milestone has been the development of phenotype structured population models that link ecology and evolution. Moreover, biostatistical research steadily improves methods to coax demographic information from scarce data. In this thesis, I build upon some of the recent advances in the field. My first three studies focus on the consequences of sex differences in life history for population dynamics. Firstly, I test whether males matter for the dynamics of African lion (Panthera leo) populations via a previously unquantified mechanism: the inheritance of phenotype from father to offspring. Secondly, I develop a method to estimate age-specific mortality rates for both sexes in species where one of the sexes disperses around the age of maturity. Thirdly, I apply this method to study variation in mortality between the sexes and between two populations of African lions. After these three chapters, which make contributions to the field of sex-structured population dynamics, I focus on the integration of phenotype structured modelling and quantitative genetics. I illustrate how heritability of a quantitative character that develops with age depends on (i) viability selection, (ii) fertility selection, (iii) the development of the phenotype with age, and (iv) phenotype inheritance from parents to offspring. Our results question the adequacy of quantitative genetics methods to obtain unbiased estimates of heritability for wild populations. This thesis advances our understanding of population development over ecological time scales. This knowledge has applications in conservation and population management, but also contributes to untangling evolutionary processes in wild animals.

Semi-aquatic mammals have evolved to forage in both aquatic and terrestrial environments, and they generally lack locomotor specialisation for either. Having relatively unspecialised adaptations, some semi-aquatic species are generalist foragers, and their activity and diving patterns provide insight into constraints on their foraging behaviour. The recent miniaturisation and improved accuracy of logging devices allow remote data collection from small (<1kg), shallow-diving species. The development of analytical methods has lagged behind technological advances (and techniques developed for fully aquatic animals do not account for the variability of behaviour typical of semi-aquatic animals and their switching between terrestrial and aquatic environments). I applied and developed novel analytical techniques to identify activities and diving patterns of a semi-aquatic mustelid, the American mink (Neovison vison), fitted with Time-Depth Recorders (TDRs). Using a hidden Markov model (HMM) algorithm allowed me to classify dives into three states to identify clustering, and describe sequential diving patterns by mink. TDRs were used to determine active/inactive periods on the basis of rapid TDR temperature changes, and this method was validated empirically. Having developed this methodology, I applied it to 18 datasets collected from 14 mink in lowland England. Terrestrial activity of mink was positively related to ambient temperature (across both sexes), however aquatic activity, especially diving, appeared to be more influenced by daylight than by temperature. Mink showed intersexual differences, with males being more nocturnal and more active on land, and females more diurnal, and more persistent in diving. There was considerable variability between sexes and individuals. This is the first study to use HMM to classify the dives of a semi-aquatic animal, and the first to use TDR temperature records to identify mammalian activity patterns. These methods will be generally applicable to animals that make rapid transitions between environments and have thus far been difficult to study.

Insects are often infected with facultative endosymbiotic bacteria, which can have a range of important ecological effects. The grain aphid, Sitobion avenae, harbours diverse facultative symbionts, which suggests their importance in grain aphid biology. This thesis attempts to explain the ecological roles of the facultative endosymbionts in S. avenae. It also examines the question of whether the horizontal transmission of symbionts between aphid clones and species can be important for shaping the ecology and evolution of multi-species aphid communities. Novel techniques developed for research with the grain aphid study system are presented. Grain aphid clones vary in their tolerance to low temperatures, but this trait is not affected by their facultative endosymbionts. Strains of a symbiont Hamiltonella defensa do not protect grain aphids from hymenopterous parasitoids, regardless of the host genotype. However, experienced parasitoid females preferentially oviposit in aphids which do not harbour symbionts. Comparison of the fitness consequences of infection with the same Hamiltonella strains in their original and in novel grain aphid host clones reveal no consistent differences. Symbiont strains establish easily following artificial transfer between clones of the grain aphid, but the symbionts transferred from other aphid species form less stable infections. Hamiltonella strains do not affect the fecundity of their grain aphid host clones regardless of their host species of origin, but also do not generally confer protection against parasitoids. There are no clear patterns in the distribution of parasitoid-resistant phenotypes across phylogenetic trees of Hamiltonella and its bacteriophage APSE. Strains of four unrelated species of endosymbionts, Rickettsia, Spiroplasma, Rickettsiella and Regiella, confer the same pathogen-resistant phenotype to a single pea aphid clone. The same symbiont strains can confer resistance to clones of two different aphid species. Some strains in multiple infections may compensate for the costs of infections with other symbionts. The importance of these results for understanding the ecological and evolutionary role of facultative endosymbionts in aphids and other insects are discussed, and directions for further research are proposed.

Many bacteria, viruses and other parasites cause severe morbidity or mortality in their host populations, creating strong selection for physiological or behavioural mechanisms to avoid disease. Likewise, changes in host susceptibility and contact patterns can dramatically influence the spread of infectious diseases, and hence selection for traits such as virulence and infectivity range in parasites. Understanding how ecological and evolutionary changes in one population affect selection in another represents a key challenge for theoreticians and empiricists alike, and is essential for gaining further insights into host-parasite relationships. This thesis contains theoretical models that explore how genetic and environmental factors shape the evolutionary and coevolutionary dynamics of hosts and parasites. In particular, the roles of genetic specificity (i.e. genotype-by-genotype interactions) and population mixing patterns are investigated, using both mathematical models and computer simulations. A broad range of scenarios are covered, including the coevolution of broad resistance and infectivity ranges (generalism), the persistence of coevolutionary cycling and the maintenance of sex, the effects of mating behaviour on disease prevalence and evolution, and the evolution of sexual and social behaviour. The models presented herein develop our understanding of host-parasite relationships and highlight the importance of genetic interactions and ecological feedbacks.

This thesis centres around two genera of large peltospirid gastropods (Mollusca: Neomphalina: Peltospiridae) endemic to hydrothermal vent ecosystems. One is the 'scaly-foot gastropod', an emblematic species of the Indian Ocean vents with unique dermal sclerites covering the foot like roof tiles. The other was recently discovered from expeditions to the Southern and Indian oceans, lacks sclerites and possesses large opercula. As both genera and their assigned species remained undescribed, they were formally described herein which forms a basis to understanding their biology. The 'scaly-foot gastropod' from both the Central Indian Ridge (CIR) and the Southwest Indian Ridge (SWIR) were confirmed to represent a single species and is formally named as Chrysomallon squamiferum. Through molecular genetic analyses using the COI gene, genetic differentiation between SWIR and CIR populations was detected for the 'scaly-foot gastropod'. In contrast, the peltospirids with large opercula from the East Scotia Ridge (ESR) and the SWIR proved to be two distinct species within an undescribed genus. The ESR species was formally described as Gigantopelta chessoia and the SWIR species as G. aegis. The molecular genetic analyses of the COI gene, confirmed the genetic isolation of the two and consolidated their status as separate species. A 3D tomographic model of Chrysomallon squamiferum was generated to characterise the soft anatomy and morphology as well as to understand its internal anatomy and adaptation which remained little-studied. Further to the enlarged esophageal gland already known to house chemosynthetic endosymbionts, C. squamiferum was discovered to have a hypertrophied circulatory system with a gigantic, muscular heart and large ctenidium to adapt to life in a hypoxic environment and to supply the endosymbionts with necessary chemicals. Histological examinations of the sclerites and operculum showed that it was unlikely that the sclerites originated from operculum duplication. Comparisons with polyplacophoran scales revealed starkly different secretion mechanisms despite the superficial similarity, which has implications on the placement of sclerite-bearing Cambrian taxa. Overall, the results from this thesis ascertained the systematic positions of these large-sized, enigmatic peltospirids, and led to improved understanding of their ecology and evolution. The important role of larval dispersal in maintaining metapopulations across the distribution of a vent-endemic taxa is highlighted. The adaptations of vent-endemic taxa remains little-known even in well-studied species, warranting future studies on these and other species.

Metapopulation ecology has developed to explain the population dynamics that occur in spatially structured landscapes. In this study, I combined an empirical laboratory approach, using metapopulation microcosms of Callosobruchus maculatus and its endospecific parasitoid Anisopteromalus calandrae, with mathematical population models in order to investigate several fundamental metapopulation processes. Population dynamics in these systems can be studied at two scales; the local patch-wise scale and the regional metapopulation scale. Here I demonstrate that in both homogeneous and heterogeneous landscapes knowledge of local scale demographic processes is necessary in order to understand regional metapopulation dynamics. The differences in the rate and net direction of dispersal between patches as a result of the permeability of the matrix in homogeneous systems and density-dependent dispersal in heterogeneous systems were also explored. Metapopulation dynamics rely on a balance between local extinctions and recolonisations. Therefore, increasing local mortality rates is likely to be detrimental to the persistence of the system. Here, the impact of several common harvesting strategies on the persistence of a host-parasitoid metapopulation was examined. Contrary to expectation I discovered that harvesting in these systems increased both local and regional population sizes. The increased population size as a result of increased mortality was explained in terms of a hydra effect, where harvesting relaxed density-dependence acting on local host populations. The results presented in this thesis are relevant for the monitoring, management and conservation of natural metapopulations and the development of sustainable harvesting strategies in structured landscapes.

A combination of habitat loss, degradation and fragmentation are causing declines in butterfly species across Europe, with habitat specialists more vulnerable than generalists. The influence of climate change is serving to complicate matters, particularly at the edge of geographic ranges. Improving the quality of habitats is a key tool for the conservation of threatened species at both a local and landscape scale and this requires a thorough understanding of the ecology and habitat requirements of species. From this perspective this thesis examines and compares the ecology of two closely butterfly species, Polyommatus bellargus (the Adonis Blue) and Polyommatus coridon (the Chalkhill Blue) in southern England; calcareous grassland specialists, found in a highly fragmented ecosystem at the northern most edge of their geographic range. Chapter 2 explores whether there has been an expansion in the larval niche of P. bellargus due to a warming climate in recent decades, and finds no indication of change once concurrent local habitat change has been accounted for. Chapter 3 examines the role that differences in microhabitat requirements play in differentiating the larval niche of P. bellargus and P. coridon, showing P. coridon has broader microhabitat requirements. Chapter 4 experimentally examines the specificity of mutualistic interactions between the larvae of each butterfly species with the ants Myrmica sabuleti and Lasius niger. Chapter 5 uses microsatellites to explore the genetic structure of both P. bellargus and P. coridon, showing (as predicted) that P. coridon has a less fragmented population structure than P. bellargus, but both species have high levels of inbreeding.

Species diversity varies dramatically across the surface of the Earth. A key step in the accumulation of species diversity is the ability of species to coexist in biological communities. Thus, identifying the mechanisms underlying community assembly is a major challenge for ecologists seeking to explain patterns in species diversity and composition. Recently some consensus has been reached on the set of processes that influence community assembly: speciation, demographic stochasticity, niche-based fitness trade-offs among species and dispersal. However, it is unclear how the importance of a particular process changes with spatial scale, which interactions exist among processes at large spatial scales and the extent to which niche-based resource partitioning among species explains differences in diversity among communities. Neotropical birds offer an ideal opportunity to address these uncertainties because of their high diversity and the existence of detailed information on their evolutionary history and ecology. In this thesis, I first use trait and phylogenetic metrics of community structure to show that both habitat filtering and interspecific competition shape community composition at the scale of individual bird territories (~1-2 ha). Second, I use simulations of community assembly to show that trait-based metrics of community structure outperform phylogenetic metrics for detecting niche-based community assembly, and that both sets of metrics often have low power when multiple processes influence community composition. Third, taking a trait-based, species-level approach, I show that both habitat filtering and interspecific competition influence species occurrence at regional scales (~75000 km²), and interact with dispersal ability so that their effect on species occurrence is increased for species with greater dispersal ability. Finally, using a combination of trait- and isotope-based methods to quantify resource partitioning, I show that species' niche widths do not change and niche overlap is reduced at high compared to low species richness. Taken together, these results suggest that both habitat filtering and interspecific competition (i.e. niche-based processes) influence community assembly from local to regional scales. However, at least at regional scales, the degree to which these processes are important for determining the occurrence of any given species depends on that species's dispersal ability. They also suggest, based on niche-based interspecific competition influencing community composition, that differences in species richness among communities are in part explained by differences among sites in the breadth of available niche space, not by increased ecological specialisation or niche overlap.

Inclusive fitness theory concerns the study of social traits. Often, individuals differ in their phenotype (e.g. size, weight, nutritional state) independently of their genetic make up, that is, individuals differ in their quality. Individuals can then be classified into different “classes” according to their quality, which enable us to understand social evolution in class-structured populations. This is important because individuals in natural populations often differ in quality, either because of intrinsic factors (e.g. size), or extrinsic factors (e.g. resource availability). My thesis concerns the evolution of social traits in class-structured populations. In chapter 1, I make a brief introduction to my thesis, providing the abstract of each chapter. In chapter 2, I outline a general theory of individual quality, where I show how individual quality impacts social evolution in two fundamental ways. In chapter 3, I show that resource heterogeneity greatly influences the evolution of conditional social behaviour. In chapter 4, I show that temporal group-size heterogeneity promotes the evolution of both conditional helping and harming. In chapter 5, I analyse the effect of individual quality on kin selection. I find that individual quality has an important impact in kin selection, which can lead to extreme forms of social behaviour. In chapter 6, I show that stable environments promote the evolution of negative density-dependent dispersal, while unstable environments promote the evolution of positive density-dependent dispersal. In chapter 7, I show that budding and low local quality promote the evolution of dispersal and cooperation.