Increasing temperatures resulting from climate change have within recent years been shown to advance phenological events in a large number of species worldwide. Species can differ in their response to increasing temperatures, and understanding the mechanisms that determine the response is therefore of great importance in order to understand and predict how a warming climate can influence both individual species, but also their interactions with each other and the environment. Understanding the mechanisms behind responses to increasing temperatures are however largely unexplored. The selected study system consisting of host plant species of the Brassicaceae family and their herbivore Anthocharis cardamines, is assumed to be especially vulnerable to climatic variations. Through the use of this study system, the aim of this thesis is to study differences in the effect of temperature on development to start of flowering within host plant species from different latitudinal regions (study I), and among host plant species (study II). We also investigate whether different developmental phases leading up to flowering differ in sensitivity to temperature (study II), and if small-scale climatic variation in spring temperature influence flowering phenology and interactions with A. cardamines (study III). Finally, we investigate if differences in the timing of A. cardamines relative to its host plants influence host species use and the selection of host individuals differing in phenology within populations (study IV). Our results showed that thermal reaction norms differ among regions along a latitudinal gradient, with the host plant species showing a mixture of co-, counter- and mixed gradient patterns (study I). We also showed that observed differences in the host plant species order of flowering among regions and years might be caused by both differences in the distribution of warm days during development and differences in the sensitivity to temperature in different phases of development (study II). In addition, we showed that small-scale variations in temperature led to variation in flowering phenology among and within populations of C. pratensis, impacting the interactions with the butterfly herbivore A. cardamines. Another result was that the less the mean plant development stage of a given plant species in the field deviated from the stage preferred by the butterfly for oviposition, the more used was the species as a host by the butterfly (study IV). Finally, we showed that the later seasonal appearance of the butterflies relative to their host plants, the higher butterfly preference for host plant individuals with a later phenology, corresponding to a preference for host plants in earlier development stages (study IV). For our study system, this thesis suggest that climate change will lead to changes in the interactions between host plants and herbivore, but that differences in phenology among host plants combined with changes in host species use of the herbivore might buffer the herbivore against negative effects of climate change. Our work highlights the need to understand the mechanisms behind differences in the responses of developmental rates to temperature between interacting species, as well as the need to account for differences in temperature response for interacting organisms from different latitudinal origins and during different developmental phases in order to understand and predict the consequences of climate change. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 4: Manuscript.</p>

Over the last 150 years the natural nighttime environment has been drastically altered by the proliferation of artificial light. The amount of artificial light at night is on the increase, and there is a current trend to replace older lighting with more energy efficient types such as light emitting diodes (LEDs) or ceramic metal halide; in Cornwall, UK, there has been a relatively recent replacement of the street lighting, from low pressure sodium to ceramic metal halide. Alongside the increasing amount of artificial nighttime light, recent research has highlighted declines in macro moth numbers. Given the well-known ‘flight-to-light’ behaviour of moths, and the negative effects this behaviour can have, alongside other known and potential ways in which nighttime light can affect moths, the increasing amount of artificial light in the environment is a suspected contributor to the declines. It is particularly important to understand how modern lighting technologies will impact upon moths, as different spectra of light are known to vary in terms of how attractive they are. As a means to determine the potential impact of different street lighting types on moths, particularly the ceramic metal halide lighting rolled out in Cornwall, UK, we compared the attractiveness to macro moths, of a number of increasingly used, energy efficient, street lighting types. We found that shorter wavelength metal halide lighting attracted significantly more individuals and species of moth than longer wavelength high pressure sodium lighting. In a second experiment, we also found ceramic metal halide lighting to be more attractive to macro moths than LED lighting. Reduced emissions of short wavelength UV light was deemed the likely reason behind the fewer macro moths attracted to the high pressure sodium and LED lighting. Interestingly, we also found striking differences in the relative attractiveness of the different lighting types to different moth groups. The metal halide lighting attracted significantly more Noctuidae than high pressure sodium lighting, whereas both high pressure sodium and metal halide lighting were equally attractive to Geometridae. Understanding accurately the extent to which different groups of moth are attracted to different wavelengths of light could be useful in determining the impact of artificial light on moth populations. In addition to impacting moths through attraction, artificial light has the potential to alter the day length as perceived by organisms, which at mid- to high latitudes is utilised by certain species as an abiotic cue to ensure the coincidence of development with favourable environmental conditions. Due to a paucity of knowledge on how raised ambient nighttime light levels affect moths and the trophic levels with which they interact, we carried out analyses into the impact of nighttime light on the winter moth and its host plant oak; a well-studied model system, where synchrony between moth egg hatch and oak budburst is important for the moth’s survival. Firstly we carried out an analysis looking at the relationship between the amount of nighttime light and the date of oak budburst. Spatially referenced budburst dates were matched with satellite imagery of nighttime lighting and average spring temperature data, and the relationship between the variables was analysed. Model predictions suggested that oak budburst occurs earlier in brighter areas. In addition, the predicted advance of budburst in brighter areas was still apparent when analysing only the data points that fell outside of large urban areas, where the urban heat island effect is likely reduced. The findings suggested that artificial nighttime light may be causing an advance in oak budburst. To follow up the spatial analysis we carried out a field experiment. We used light cages that simulated various nighttime lighting scenarios to test whether oak budburst and winter moth egg hatch were affected by low intensity light at night. In contrast to the spatial analysis, there was no significant relationship found between light treatment and the phenology of either oak budburst or winter moth egg hatch. However, there was a suggestion in the data that the higher buds of the oak saplings emerged earlier in the yellow light treatment, highlighting the need for further research into the potential impact of artificial nighttime light on phenology and species interactions. In conclusion, the findings of this research project provide information useful to those seeking ecologically sensitive lighting solutions, and also highlight a potential tool to assist in determining whether light at night is a causative factor behind apparent moth declines. In addition, they suggest that artificial light at night may be affecting the phenology of an ecological system at a national scale. Finally, this research project has highlighted the complexity of the ecological impacts of artificial light at night, and also a need for further research.

<p>The minimum ecologically viable population size (MEVP) of a species describes the minimum size at which the species itself or another species in the same ecosystem goes extinct as a result of the loss of inter-specific interactions. The MEVP shows a good potential for use as a tool for exploring the mechanisms behind species extinctions, but presently only a small amount of research has been done that takes advantage of this. In this study the MEVP is used to investigate what properties of species can be used as good indicators of ecological importance. 100 large computer generated food webs were created with an assembly model and the reduction in density that was necessary to induce an extinction event in the web was subsequently determined for each species within the webs. This change in density was then correlated with 28 different properties, measured for each species. The results show that properties that measure how well connected a species is, as well as measures of the species role as a prey item in the web are the ones with the greatest potential to find species with high MEVP. Further, the results put emphasis on the importance of regarding the web as a whole when working with species extinctions, while also highlighting the usefulness of the MEVP concept.</p>

Throughout history species has gone extinct due to anthropogenic activities. During the last century efforts have been done to reintroduce species back into the wild. Zoos that originally were created as amusement parks for people have today a new purpose; to keep and breed species in captivity for later reintroductions in the wild. However a relaxed environment such as a zoo leads to a general fitness decline of up to 40% per generation in captivity. The probability of a successful reintroduction of a species that has been bred in a zoo will be lower the longer time it has been kept in captivity. The reintroduction of a captive bred species can also cause secondary extinctions and other negative effects on the food-web. Both changes in the community caused by the loss of a species and changes in the species itself caused by captivity can be expected to affect the outcome of a reintroduction attempt. Using a modeling approach I here investigate how the reintroduction of a captive bred species (at three different trophic levels; basal, intermediate and top predator species) affects a food-web and what risks there are in reintroducing it. A Lokta-Volterra model with type II functional response is used. I investigate three scenarios: the reintroduction of a species with 0% change in its attributes, 40% change in its attributes and 75% change in its attributes. It was found that the most important factor for reintroduction success when reintroducing a species is whether it is a producer species (basal species) or a consumer species (intermediate and top predator species). The producer species were most sensitive to the changes in their attributes, whilst consumer species were more sensitive to change in the food-web (Euclidian distance). The producer species were found to cause most secondary extinctions in all scenarios, hence indicating that it is a bottom-up controlled food-web. The present study suggests that the success of a reintroduction attempt is affected both by the degree of changes in the food web caused by the initial loss of the species and the degree of change in the species itself caused by captivity.

The minimum ecologically viable population size (MEVP) of a species describes the minimum size at which the species itself or another species in the same ecosystem goes extinct as a result of the loss of inter-specific interactions. The MEVP shows a good potential for use as a tool for exploring the mechanisms behind species extinctions, but presently only a small amount of research has been done that takes advantage of this. In this study the MEVP is used to investigate what properties of species can be used as good indicators of ecological importance. 100 large computer generated food webs were created with an assembly model and the reduction in density that was necessary to induce an extinction event in the web was subsequently determined for each species within the webs. This change in density was then correlated with 28 different properties, measured for each species. The results show that properties that measure how well connected a species is, as well as measures of the species role as a prey item in the web are the ones with the greatest potential to find species with high MEVP. Further, the results put emphasis on the importance of regarding the web as a whole when working with species extinctions, while also highlighting the usefulness of the MEVP concept.

To understand the processes affecting the abundance of wild populations is a fundamental goal of ecology and a prerequisite for the management of living resources. Variable abundance, however, makes the investigation of ecological processes challenging. Recruitment, the process whereby new individuals enter a given stage of a ?sh population, is a highly variable entity. I have confronted this issue by developing methodologies speci?cally designed to account for, and ecologically interpret, patterns of variability in recruitment. To provide the necessary context, Chapter 2 begins with a review of the history of recruitment science. I focus on the major achievements as well as present limitations, particularly regarding environmental drivers. Approaches that include explicit environmental information are contrasted with time-varying parameter techniques. In Chapter 3, I ask what patterns of variability in pre-recruit survival can tell us about the strength of density-dependent mortality. I provide methods to investigate the presence of density-dependent mortality where this has previously been hindered by highly variable data. Stochastic density-independent variability is found to be attenuated via density dependence. Sources of recruitment variability are further partitioned in Chapter 4. Using time-varying parameter techniques, signi?cant temporal variation in the annual reproductive rate is found to have occurred in many Atlantic cod populations. Multivariate state space models suggest that populations in close proximity typically have a shared response to environmental change whereas marked differences occur across latitude. Hypotheses that could result in consistent changes in productivity of cod populations are tested in Chapter 5. I focus on a meta-analytical investigation of potential interactions between Atlantic cod and small pelagic species, testing aspects of the cultivation-depensation hypothesis. The ?ndings suggest that predation or competition by herring and mackerel on egg and larval cod could delay recovery of depleted cod populations. Chapter 6 concludes with a critical re?ection on: the suitability of the theories employed, the underlying assumptions of the empirical approaches, and the quality of the data used in my thesis. Application of ecological insights to ?sheries management is critically evaluated. I then propose future work on recruitment processes based on methods presented herein.

No description available.

Thesis (PhD (Conservation Ecology and Entomology))--Stellenbosch University, 2008. / ENGLISH ABSTRACT: Shifts in species ranges are an important consequence of climate change, and can affect the composition, structure and functioning of ecosystems. Generally, in response to rising temperatures, species have increased their altitudinal and latitudinal distributions along their cooler boundaries, although there has been large variation between species. However, in addition to climatic factors, species range limits are also sensitive to biotic interactions. Therefore, interspecific interactions may be able to influence the impact of climate change on species distributions. In this thesis climate change, range shifts and spatial variation in plant-plant interactions are documented to examine the potential for biotic interactions to mediate climate-driven altitudinal range shifts on sub-Antarctic Marion Island. The climate on Marion Island changed considerably between 1949 and 2003, with significant trends in biologically-relevant parameters encompassing average climatic conditions, variability in climate parameters and extreme weather events. Combining these trends showed that while environmental conditions have ameliorated for some species, they have become more abiotically stressful for others (e.g. hygrophilous species). Concurrent with changes in the island’s climate, there have been rapid changes in the elevational distribution of Marion Island’s native vascular flora. On average species ranges expanded upslope (as predicted in response to the warming experienced on the island), although individual range expansion rates varied greatly. As a result of the idiosyncratic expansion rates, altitudinal patterns of species richness and community composition changed considerably. Therefore, both speciesand community-level changes occurred in the flora of Marion Island, demonstrating the community’s sensitivity to climate change. To determine the potential for interspecific interactions to have mediated these changes in species elevational ranges, spatial variation in the balance of positive and negative plant-plant interactions were examined. Increasing spatial association between four dominant species along an altitudinal severity gradient suggested that the intensity of facilitation, relative to the intensity of competition, is greater under more severe conditions. While, interaction strength varied between species pairs, the nature of relationship was consistent across the species. At a broader spatial scale, the performance of the grass Agrostis magellanica was compared in the presence and in the absence of the cushion plant Azorella selago along the entire altitudinal range of both species. The influence of the cushion plant on A. magellanica switched from negative to positive with increasing environmental severity, reaching a positive asymptote under extreme environmental conditions. Therefore, on Marion Island the spatial variation in the nature of biotic interactions is related to environmental severity, and facilitative interactions can be strong under extreme environmental conditions. These results show that positive biotic interactions are important at higher elevations on Marion Island, and could thus accelerate upslope range expansions in response to climate change. Nonetheless, competitive inhibition of upslope species movement, especially in areas of dense vegetation (i.e. mid or low altitudes), could have the opposite result, acting antagonistically with the effect of rising temperatures. This research highlights the importance of considering interspecific interactions when examining the biotic implications of climate change, both for affecting the rate at which species ranges change and the extent of species distributions. / AFRIKAANSE OPSOMMING: Die verskuiwing van spesies se verspreidingspatrone is ‘n belangrike gevolg van klimaatsverandering en kan die samestelling, struktuur en funksionering van ekosisteme affekteer. Oor die algemeen, in reaksie op verwarming, kom spesies op hoër hoogtes bo seespieël en hoër breedtegrade voor, alhoewel daar groot verskille tussen spesies in hierdie aspek is. Behalwe vir klimatologiese faktore, is spesies se verspreidingslimiete ook sensitief vir biotiese interaksies met ander spesies. Daarom kan interaksies tussen spesies die effek van klimaatsverandering op spesiesverspreiding verder beïnvloed. In hierdie verhandeling word klimaatsverandering, verandering in verspreiding van spesies en geografiese variasie in plant-plant interaksies ondersoek. Dit word gedoen ten einde die moontlikheid te toets dat biotiese interaksies die verandering in verspreidingspatrone van klimaat gedrewe spesies op sub-Antarktiese Marion Eiland beïnvloed. Marion Eiland se klimaat het aansienlik tussen 1949 en 2003 verander, met betekenisvolle neigings in biologies-relevante parameters, insluitend gemiddelde klimaatstoestande, variasie in klimaat en uiterste weerstoestande. As die veranderinge saam beskou word, wys dit dat, terwyl omgewingstoestande vir sommige spesies verbeter het, toestande vir ander spesies abioties meer ongunstig geword het (bv. akwatiese plantsoorte). Tesame met die veranderinge in die eiland se klimaat, het die hoogte bo seespieël verspreiding van Marion Eiland se inheemse vaatplante vinnig verander. Gemiddeld het spesies-verspreiding se hoogte bo seespieël na hoër hoogtes verskuif (soos voorspel in reaksie op verwarming), alhoewel die tempo van verskuiwing na hoër hoogtes tussen spesies verskil het. As gevolg van hierdie idiosinkratiese reaksies, het die aantal en samestelling van plant spesies op verskillende hoogtes op die eiland aansienlik verander. Die sensitiwiteit van Marion Eiland se flora ten opsigte van klimaatsverandering word deur hierdie verandering gedemonstreer. Om te bepaal of die interaksies tussen spesies verspreidingspatrone kan beïnvloed, is die geografiese variasie in die balans van positiewe en negatiewe interaksies ondersoek. Vier dominante vaatplante het meer geassosieerd geraak hoe hoër hulle bo seespieël voorgekom het. Dit is ‘n aanduiding dat die sterkte van die positiewe interaksies (fasilitering), relatief tot die sterkte van die negatiewe reaksie (kompetisie), groter onder uiterste omgewingstoestande is. Die sterkte van die interaksie het verskil tussen spesies pare, maar die verhouding tussen omgewingstoestande en die sterkte van die interaksie was dieselfde vir alle spesies. Op ‘n breër geografiese vlak is die opbrengs van die gras Agrostis magellanica in die teenwoordigheid en afwesigheid van die polsterplant Azorella selago op alle hoogtes waar die plante saam voorkom vergelyk. Die invloed van A. selago op A. magellanica was negatief op lae hoogtes bo seespieël, maar het meer positief geraak met ‘n toename in hoogte bo seespieël, tesame met omgewingshardheid. Die invloed van die polsterplant op die gras het ‘n positiewe asimptoot onder uiterste omgewingstoestande bereik. Dus, op Marion Eiland is die geografiese variasie in biotiese interaksies verwant aan omgewingstoestande, en positiewe interaksies kan selfs onder die uiterste omgewingstoestande sterk wees. Hierdie resultate wys dat positiewe biotiese interaksies tussen plante belangrik is by hoër hoogtes bo seespieël op Marion Eiland. Die interaksies kan dus die opwaartse verspreiding van spesies in reaksie op klimaatsverandering versnel. Nogtans kan negatiewe interaksies die teenoorgestelde effek hê aangesien kompetisie tussen plante, veral in plekke met digte plantegroei (d.w.s. lae of middel hoogtes bo seespieël) opwaartse verskuiwing van spesies kan verhinder. Hierdie navorsing dui aan hoe belangrik dit is om interaksies tussen spesies in ag te neem, aangesien die interaksies die tempo en omvang van veranderinge in verspreiding kan beïnvloed.

Community genetics research investigates the influence of intra-specific genetic variation on species interactions. This rapidly growing research field consists of more than one approach to explore how a significant portion of the environment of a focal species is differentially defined by the expressed genomes of other interacting species. While the basic concept of community genetics is well supported empirically, there is still a set of pertinent issues in need of further investigation. The initial research addressed herein focused on the extent to which the magnitude of a community genetic effect can be moderated when acting in concert with other forces in nature, i.e. the interaction between community genetic effects and the effects of other eco-evolutionary processes such as competition and parasitism. Subsequent research investigated the impact of genetic variation of host plants in agro-ecosystems on the performance (reproductive success) and behaviour (distribution and feeding-site choice) of plant-associated pests such as aphids, especially when pests and their hosts were subject to plant-mediated interactions. In addition, the differential effects of Indirect Ecological Effects (IEEs) and Indirect Genetic effects (IGEs) on the emergence of shared (extended) phenotypes between natural enemies (i.e. biological control agents and phytophagous insects) were examined. I provide clear evidence for significant effects of the genetic variation of host plant on aphid performance, behaviour and intra- and inter-specific competition. My findings also give credence to the concept of reciprocal moderation between plant genotype and aphid competition. I also provide observations on competition that segues into less antagonistic and possibly into a more cooperative form of interaction. In addition, I establish novel systems of economically important crop genotypes, noxious sap-feeding aphid species and root-galling nematodes. I also devise an amalgamated approach to interpret the interwoven set of mechanisms that underpin the observations presented and conclusions drawn. I also provide further investigation on the role of Indirect Ecological Effects (IEEs) between root-knot nematodes and sap-feeding aphids, and demonstrate the influence of in-plant variation on the interaction between the spatially separated plant consumers. Furthermore, I use a quantitative genetic experimental design in order to demonstrate a differential impact of parasitoid genotype on the behaviour of its aphid host. As such, I provide some of the clearest evidence to date that the phenotype of an organism can be the product of the genes expressed in another organism via Inter-specific Indirect Genetic Effects (IIGEs). Finally, I conducted research on epiphytic bromeliads and their associated faunal communities in the tropics. Here I demonstrate that the influence of intra-specific genetic variation of the host plant on the associated ecological communities may be more universal than previously conceived, with a plausible role for such variation in the maintenance of biological diversity. My research provides evidence for the genetic basis of species interactions and, interestingly, a genetic basis for the evolutionary arms-race between foragers and their hosts. My doctoral work adds new evidence to the increasing literature on the evolutionary importance of (Genotype x Genotype) interactions and (Genotype x Genotype x Environment) interactions in shaping the dynamics of pest communities, which in turn can affect plant phenotype and can influence the properties and services of the focal ecosystem in which the inter-players live and interact.

Studier visar att dagens elever känner oro inför framtiden och de klimatförändringar vår värld utsätts för. Skolverkets nationella kartläggning utav miljöundervisning i skolan visar dock att cirka hälften av de tillfrågade gymnasielärarna inte bedriver någon undervisning i ämnet. Denna studie beskriver en alternativ modell för undervisning i klimatfrågor. Elever ges genom modellering av ekosystem möjlighet att fördjupa sin förståelse kring klimatförändringar i samband med exploatering av topp-predatorer. Detta är ett realistiskt problem då en mängd studier visar att dagens ekosystem utsätts för en mängd stressfaktorer, med ursprung i klimatförändringar, och att exploatering av topp-predatorer kan leda till konsekvenser med artutdöende som följd. Modellen som används i undersökningen är en generell Lotka-Volterra-modell vilken fungerar som analytiskt verktyg. Skolverket betonar matematikens roll som ämnesöverskridande verktyg och modellekosystem ger elever möjlighet att använda matematiska kunskaper i ett verkligt scenario. I studien undersöks även hur pass stor exploatering topp-predatorer kan utsättas för i takt med ökade klimatförändringar. De undersökta områdena inbegriper tre- och fyr-artssystem i marina och terrestra miljöer. Resultaten visar att exploateringen måste minska i takt med ökade klimatorsakade förändringar i samtliga fall, undantag terrestra tre-artsystem som utsätts för förändrade interaktionsstyrkor. / Studies show that today's students are worried about the future and the increasing climate change. A survey made by the Swedish National Agency for Education about environmental education shows that approximately half of the surveyed secondary school teachers are not engaged in teaching in the field. This study describes an alternative model for teaching environmental issues. Students are by the modeling of ecosystem given an opportunity to deepen their understanding of climate change associated with the exploitation of top predators. This is a realistic problem, numerous studies show that today's ecosystems are exposed to a variety of stressors, with the origin of climate change, and the exploitation of top predators can lead to serious consequences with extinction as a result. The model used in this study is a generalized Lotka-Volterra model which works as an analytical tool. The Swedish National Agency for Education emphasizes the role of mathematics as interdisciplinary tools and model ecosystem gives students the opportunity to use math skills in a real scenario. The study also investigated how much exploitation top predators can be exposed to in line with increased climate change. The research sites include three-and four-species system of marine and terrestrial environments. The results show that the exploitation must be reduced in line with increased climate-caused changes in all cases, except terrestrial three-species system exposed to changes in interaction strengths.