The Great Irish Famine in history-writing and prose fiction : "the mutual interplay of two narrative genres"

Denna tvärvetenskapliga studie kontextualiserar den skönlitterära framställningen av hungersnöden i Irland i mitten av 1800-talet i förhållande till tre olika faser av irländsk historieskrivning ‒ den nationalistiska, den revisionistiska, och den postrevisionistiska ‒ med syfte att granska i vilken mån prosafiktionen antingen återspeglar eller motsäger historikernas tolkningar. År 1845 drabbades landet av en dittills okänd potatispest som förstörde skörden helt eller delvis under de följande fem åren. Missväxten ledde till utbredd svält och epidemiska sjukdomar som dödade åtminstone en miljon människor, medan ytterligare en och en halv miljon flydde, huvudsakligen till Förenta Staterna, England, och Kanada. I sina försök att hitta en rationell förklaring till hur potatispesten kunde utvecklas till den värsta svältkatastrofen i Europa under modern tid, har historiker påvisat ett antal bidragande faktorer, till exempel överbefolkning, de fattigaste jordlösa småbrukarnas och lantarbetarnas beroende av potatisen som sitt baslivsmedel, underutveckling inom jordbruket, det rådande jordegendomssystemet, och den dåvarande brittiska regeringens misslyckande att tillhandahålla effektiv och tillräcklig nödhjälp. Historiska förklaringar är naturligtvis nödvändiga för att vi skall kunna bilda oss en uppfattning om hungersnödens orsaker och konsekvenser, men svårigheten med att skildra offrens situation i en historiografisk analys baserad på fakta är uppenbar då deras egna vittnesmål till största delen saknas i källmaterialet. Följaktligen finns det en risk att historieskrivningen förmörkar det som onekligen var centrala realiteter för de värst drabbade, nämligen svält, vräkning, sjukdom och död. Här kan skönlitteraturen bidra till att komplettera historien. Genom att fokusera på ett specifikt (fiktivt) samhälle och dess (fiktiva) individer, kan skönlitterära verk ge en inblick i hur hungersnöden inverkade på olika samhällsskikt, vad människorna gjorde för att överleva, hur nöden och fasorna påverkade deras psyke, och vad eller vem de höll ansvariga för katastrofen. Å andra sidan kan denna fokusering innebära att författaren misslyckas med att ge en helhetsbild av hungersnödens enorma omfattning och att redogöra för alla faktorer som orsakade och förlängde den. Paul Ricoeurs teori om samspelet mellan historia och fiktion (the interweaving of history and fiction) är därför ett nyckelbegrepp för att bättre förstå denna traumatiska period i Irlands historia. Avhandlingen omfattar en textanalytisk, komparativ kritik av ett antal historiska och skönlitterära verk. Genom närläsning av dessa texter granskar jag vilka aspekter av hungersnöden (politiska, ekonomiska, sociala) de olika författarna valt att behandla, och på vilket sätt, samt hur deras synvinklar har format tolkningarna i sin helhet. I detta sammanhang tar jag också upp skillnaderna mella fakta och fiktion, och speciellt de etiska problem som är förknippade med skildringen av traumatiska händelser och mänskligt lidande. Samtidigt undersöker jag, med hänvisning till Roger D. Sells kommunikationsteori, huruvida vissa författare anslår en påstridig ton i sina verk och hur detta påverkar dialogen mellan författare och läsare. Med utgångspunkt i Ricoeurs teori argumenterar jag för att historia och fiktion inte bör ses som ömsesidigt antitetiska diskurser i skildringen och tolkningen av det förflutna, och att skönlitteraturen genom fokuseringen på offren, som ofta tenderar att reduceras till statistik i historieskrivningen, kan förmedla en bättre förståelse och en djupare känsla för den mänskliga dimensionen av den tragedi som utspelades under hungeråren.

The European Union and its Northern Frontier : EUropean Geopolitics and its Arctic Context

The present study explores EUropean geopolitical agency in a distinct spatio-temporal context: the Arctic region of the early 21st century. Thus, it provides an in-depth analysis of the European Union’s process to construct EUropean legitimacy and credibility in its ‘Northern Neighbourhood’ between 2008 and 2014. Embedded in a conceptual and methodological framework using critical geopolitics, this study assesses the strategic policy reasoning of the EU and the implicit geopolitical discourses that guide and determine a particular line of argumentation so as to claim a ‘legitimate’ role in the Arctic and accordingly construct a distinct ‘EUropean Arctic space’. In doing so, it establishes a clearer picture on the (narrated) regional interests of the EU and the related developed policy and concrete steps taken in order to get hold of these interests. Eventually, the analysis gets to the conceptual bottom of what exactly fashioned the EU with geopolitical agency in the circumpolar North. As a complementary explanation, this study provides a thick description of the area under scrutiny – the Arctic region – in order to explicate the systemic context that conditioned the EU’s regional demeanour and action. Elucidated along the lines of Arctic history and identity, rights, interests and responsibility, it delineates the emergence of the Arctic as a region of and for geopolitics. The findings indicate that the sui generis character of the Arctic as EUropean neighbourhood essentially determined the EU’s regional performance. It explicates that the Union’s ‘traditional’ geopolitical models of civilian or normative power got entangled in a fluid state of Arctic affairs: a distinct regional system, characterised by few strong state actors with pronounced national Arctic interests and identities, and an indefinite local context of environmental changes, economic uncertainties and social challenges. This study applies critical geopolitics in a Political Science context and essentially contributes to a broader understanding of EU foreign policy construction and behaviour. Ultimately, it offers an interdisciplinary approach on how to analyse EU external action by explicitly taking into account the internal and external social processes that ultimately condition a certain EUropean foreign policy performance.

Patterned fen formation and development from the Great Sandy Region, south-east Queensland, Australia

The Great Sandy Region (incorporating Fraser Island and the Cooloola sand-mass), south-east Queensland, contains a significant area of Ramsar-listed coastal wetlands, including the globally important patterned fen complexes. These mires form an elaborate network of pools surrounded by vegetated peat ridges and are the only known subtropical, Southern Hemisphere examples, with wetlands of this type typically located in high northern latitudes. Sedimentological, palynological and charcoal analysis from the Wathumba and Moon Point complexes on Fraser Island indicate two periods of swamp formation (that may contain patterned fens), one commencing at 12 000 years ago (Moon Point) and the other ~4300 years ago (Wathumba). Wetland formation and development is thought to be related to a combination of biological and hydrological processes with the dominant peat-forming rush, Empodisma minus, being an important component of both patterned and non-patterned mires within the region. In contrast to Northern Hemisphere paludifying systems, the patterning appears to initiate at the start of wetland development or as part of an infilling process. The wetlands dominated by E. minus are highly resilient to disturbance, particularly burning and sea level alterations, and appear to form important refuge areas for amphibians, fish and birds (both non-migratory and migratory) over thousands of years.

A retrospective evaluation of sustainable yields for Australia's Northern Prawn Fishery

The fundamental aim in fisheries management is to determine an optimal fishing effort for sustainably harvesting from a replenishable resource. The current management objective of Australia's Northern Prawn Fishery is to maximize the long-term net economic return following Australian government policy, resulting in an average recent catch of tiger prawn species of about 1,250 tons only. However, the maximum sustainable catch stated from different studies is around 3,000-4,700 tons. We also evaluated the net profit assuming that there was no buyback scheme in 2005 and the fishing fleet was kept at 89 vessels since 2005 and concluded that 40% more catch on average (2006-2009) and an additional total profit of A$ 17 million ( excluding crew cost) could have been gained in addition to the many millions of dollars of savings in the buyback scheme. These findings have great implications for future management in Australia and elsewhere because there is a grave concern of overfishing worldwide.

Population dynamics of the European rabbit (Oryctolagus cuniculus) in north eastern Australia: simulated responses to control

Wild European rabbits are a serious problem to agriculture in Australia, with an estimated annual cost of A$ 113 million. Biological control agents (myxomatosis and rabbit haemorrhagic disease virus) have caused large and sustained declines in rabbit populations throughout Australia. A simulation model incorporates these diseases as well as warren destruction as methods of controlling rabbit populations in Queensland, north eastern Australia. These diseases reduced populations by 90-99% and the combination of these and warren destruction led to 100% control in simulations at six sites across southern Queensland. Increasing monthly pasture growth by 15% had little effect on simulated populations whereas a 15% decrease reduced populations by 0-50%. An increase in temperature of 2.5 °C would lead to a 15-60% decrease in populations. These effects suggest that climate change will lead to a decrease in the population of rabbits in Queensland and a retraction in the northern limit of their distribution in Australia.

Technical efficiency of blood component preparation in blood centres of 10 European countries

Various reasons, such as ethical issues in maintaining blood resources, growing costs, and strict requirements for safe blood, have increased the pressure for efficient use of resources in blood banking. The competence of blood establishments can be characterized by their ability to predict the volume of blood collection to be able to provide cellular blood components in a timely manner as dictated by hospital demand. The stochastically varying clinical need for platelets (PLTs) sets a specific challenge for balancing supply with requests. Labour has been proven a primary cost-driver and should be managed efficiently. International comparisons of blood banking could recognize inefficiencies and allow reallocation of resources. Seventeen blood centres from 10 countries in continental Europe, Great Britain, and Scandinavia participated in this study. The centres were national institutes (5), parts of the local Red Cross organisation (5), or integrated into university hospitals (7). This study focused on the departments of blood component preparation of the centres. The data were obtained retrospectively by computerized questionnaires completed via Internet for the years 2000-2002. The data were used in four original articles (numbered I through IV) that form the basis of this thesis. Non-parametric data envelopment analysis (DEA, II-IV) was applied to evaluate and compare the relative efficiency of blood component preparation. Several models were created using different input and output combinations. The focus of comparisons was on the technical efficiency (II-III) and the labour efficiency (I, IV). An empirical cost model was tested to evaluate the cost efficiency (IV). Purchasing power parities (PPP, IV) were used to adjust the costs of the working hours and to make the costs comparable among countries. The total annual number of whole blood (WB) collections varied from 8,880 to 290,352 in the centres (I). Significant variation was also observed in the annual volume of produced red blood cells (RBCs) and PLTs. The annual number of PLTs produced by any method varied from 2,788 to 104,622 units. In 2002, 73% of all PLTs were produced by the buffy coat (BC) method, 23% by aphaeresis and 4% by the platelet-rich plasma (PRP) method. The annual discard rate of PLTs varied from 3.9% to 31%. The mean discard rate (13%) remained in the same range throughout the study period and demonstrated similar levels and variation in 2003-2004 according to a specific follow-up question (14%, range 3.8%-24%). The annual PLT discard rates were, to some extent, associated with production volumes. The mean RBC discard rate was 4.5% (range 0.2%-7.7%). Technical efficiency showed marked variation (median 60%, range 41%-100%) among the centres (II). Compared to the efficient departments, the inefficient departments used excess labour resources (and probably) production equipment to produce RBCs and PLTs. Technical efficiency tended to be higher when the (theoretical) proportion of lost WB collections (total RBC+PLT loss) from all collections was low (III). The labour efficiency varied remarkably, from 25% to 100% (median 47%) when working hours were the only input (IV). Using the estimated total costs as the input (cost efficiency) revealed an even greater variation (13%-100%) and overall lower efficiency level compared to labour only as the input. In cost efficiency only, the savings potential (observed inefficiency) was more than 50% in 10 departments, whereas labour and cost savings potentials were both more than 50% in six departments. The association between department size and efficiency (scale efficiency) could not be verified statistically in the small sample. In conclusion, international evaluation of the technical efficiency in component preparation departments revealed remarkable variation. A suboptimal combination of manpower and production output levels was the major cause of inefficiency, and the efficiency did not directly relate to production volume. Evaluation of the reasons for discarding components may offer a novel approach to study efficiency. DEA was proven applicable in analyses including various factors as inputs and outputs. This study suggests that analytical models can be developed to serve as indicators of technical efficiency and promote improvements in the management of limited resources. The work also demonstrates the importance of integrating efficiency analysis into international comparisons of blood banking.

Genetic diversity of Atlantic salmon (Salmo salar) and European whitefish (Coregonus lavaretus) in the Baltic Sea

We described the patterns and extent of microsatellite DNA variation in historical and present-day Atlantic salmon (Salmo salar L.) stocks in the Baltic Sea and neighbouring areas, and in European whitefish (Coregonus lavaretus) ecotypes, populations and run-timing types in Finland. Moreover, the amount and pattern of genetic diversity in historical salmon populations before human impact were described, and the proportion of diversity maintained in the present hatchery stocks evaluated. Salmon populations in the Baltic Sea were, on average, significantly less variable than eastern Atlantic populations, and the diversity of landlocked populations (Lakes Vänern, Saimaa, Onega and Ladoga) was in turn significantly lower than that of anadromous salmon populations in the Baltic Sea populations. Within the Baltic Sea, the anadromous populations of Atlantic salmon formed three clear groups, corresponding to the northern (Gulf of Bothnia), eastern (Gulf of Finland and eastern Baltic Main Basin) and southern (western Baltic Main Basin) regions. Based on microsatellite data, three salmon population groups in the Baltic Sea were considered potentially different colonization lineages. In short- and long-term breeding programmes of Atlantic salmon, the average observed rate of loss of alleles was 4.9% and 2.0% per generation and the average rate of loss of heterozygosity was 1.4% and 1% per generation, respectively. When comparing the genetic parameters of stocks before and after hatchery breeding of several successive generations (Rivers Iijoki and Oulujoki), statistically significant changes in allele frequencies were common, while large wild stock in the Teno River has remained temporally very stable over 56 years. Despite the observed losses of genetic diversity in broodstock breeding, a large proportion of the genetic resources of the extirpated stocks are still conserved in the broodstocks. Genetic differentiation among European whitefish ecotypes was generally low, thus giving support to the hypothesis of one native European whitefish species in Fennoscandia. Among the ecotypes, the northern, large sparsely rakered, bottom-dwelling whitefish was the most unique. The known genetic differences in quantitative traits have thus either developed independently of potential phylogenetic lineages, or the lineages have mixed and the quantitative traits of the ecotypes, like gill-raker number, have later changed according to environment and selection pressures. Overall, genetic distances between the anadromous whitefish populations along the Finnish coast, especially in the Bothnian Bay area, were small. Wild whitefish populations studied had slightly higher allelic diversity than hatchery-reared populations in corresponding rivers.

Are high-impact species predictable? An analysis of naturalised grasses in northern Australia

Predicting which species are likely to cause serious impacts in the future is crucial for targeting management efforts, but the characteristics of such species remain largely unconfirmed. We use data and expert opinion on tropical and subtropical grasses naturalised in Australia since European settlement to identify naturalised and high-impact species and subsequently to test whether high-impact species are predictable. High-impact species for the three main affected sectors (environment, pastoral and agriculture) were determined by assessing evidence against pre-defined criteria. Twenty-one of the 155 naturalised species (14%) were classified as high-impact, including four that affected more than one sector. High-impact species were more likely to have faster spread rates (regions invaded per decade) and to be semi-aquatic. Spread rate was best explained by whether species had been actively spread (as pasture), and time since naturalisation, but may not be explanatory as it was tightly correlated with range size and incidence rate. Giving more weight to minimising the chance of overlooking high-impact species, a priority for biosecurity, meant a wider range of predictors was required to identify high-impact species, and the predictive power of the models was reduced. By-sector analysis of predictors of high impact species was limited by their relative rarity, but showed sector differences, including to the universal predictors (spread rate and habitat) and life history. Furthermore, species causing high impact to agriculture have changed in the past 10 years with changes in farming practice, highlighting the importance of context in determining impact. A rationale for invasion ecology is to improve the prediction and response to future threats. Although our study identifies some universal predictors, it suggests improved prediction will require a far greater emphasis on impact rather than invasiveness, and will need to account for the individual circumstances of affected sectors and the relative rarity of high-impact species.

The great errand 1939-1955

The novel was written in 2001. Description of post-war Germany from the viewpoint of a German, Fritz Meyer, who was a member of a local Nazi Youth organization in Sonneborn. He fought as a soldier and fell into the hands of the English in Northern France. He was taken to Canada as a prisoner of war. He escaped the camp and found refuge at a German family. Description of erotic encounters. Reflection on Nazi ideology. At the request of the family he returns to Germany for something subscribed as "the great errand", taking up the identity of a former American G.I. Desolation of post-war Germany. Confrontation with British emigre soldiers. Identifying with the anger of his German countrymen. Reflection on the Bible and the denial of the Jewish roots of Christianity. Creating an underground network of conspiracy with former Nazi leaders and high members of the Catholic church in order to continue the ideals of National Socialism. Donations from secret supporters abroad. Connections with the political leaders in the newly established German Republic. Revisionist history.

Postglacial climate changes and vegetation responses in northern Europe

Postglacial climate changes and vegetation responses were studied using a combination of biological and physical indicators preserved in lake sediments. Low-frequency trends, high-frequency events and rapid shifts in temperature and moisture balance were probed using pollen-based quantitative temperature reconstructions and oxygen-isotopes from authigenic carbonate and aquatic cellulose, respectively. Pollen and plant macrofossils were employed to shed light on the presence and response rates of plant populations in response to climate changes, particularly focusing on common boreal and temperate tree species. Additional geochemical and isotopic tracers facilitated the interpretation of pollen- and oxygen-isotope data. The results show that the common boreal trees were present in the Baltic region (~55°N) during the Lateglacial, which contrasts with the traditional view of species refuge locations in the south-European peninsulas during the glacial/interglacial cycles. The findings of this work are in agreement with recent paleoecological and genetic evidence suggesting that scattered populations of tree species persisted at higher latitudes, and that these taxa were likely limited to boreal trees. Moreover, the results demonstrate that stepwise changes in plant communities took place in concert with major climate fluctuations of the glacial/interglacial transition. Postglacial climate trends in northern Europe were characterized by rise, maxima and fall in temperatures and related changes in moisture balance. Following the deglaciation of the Northern Hemisphere and the early Holocene reorganization of the ice-ocean-atmosphere system, the long-term temperature trends followed gradually decreasing summer insolation. The early Holocene (~11,700-8000 cal yr BP) was overall cool, moist and oceanic, although the earliest Holocene effective humidity may have been low particularly in the eastern part of northern Europe. The gradual warming trend was interrupted by a cold event ~8200 cal yr BP. The maximum temperatures, ~1.5-3.0°C above modern values, were attained ~8000-4000 cal yr BP. This mid-Holocene peak warmth was coupled with low lake levels, low effective humidity and summertime drought. The late Holocene (~4000 cal yr BP-present) was characterized by gradually decreasing temperatures, higher lake levels and higher effective humidity. Moreover, the gradual trends of the late Holocene were probably superimposed by higher-frequency variability. The spatial variability of the Holocene temperature and moisture balance patterns were tentatively attributed to the differing heat capacities of continents and oceans, changes in atmospheric circulation modes and position of sites and subregions with respect to large water bodies and topographic barriers. The combination of physical and biological proxy archives is a pivotal aspect of this work, because non-climatic factors, such as postglacial migration, disturbances and competitive interactions, can influence reshuffling of vegetation and hence, pollen-based climate reconstructions. The oxygen-isotope records and other physical proxies presented in this work manifest that postglacial climate changes were the main driver of the establishment and expansion of temperate and boreal tree populations, and hence, large-scale and long-term vegetation patterns were in dynamic equilibrium with climate. A notable exception to this pattern may be the postglacial invasion of Norway spruce and the related suppression of mid-Holocene temperate forest. This salient step in north-European vegetation history, the development of the modern boreal ecosystem, cannot be unambiguously explained by current evidence of postglacial climate changes. The results of this work highlight that plant populations, including long-lived trees, may be able to respond strikingly rapidly to changes in climate. Moreover, interannual and seasonal variation and extreme events can exert an important influence on vegetation reshuffling. Importantly, the studies imply that the presence of diffuse refuge populations or local stands among the prevailing vegetation may have provided the means for extraordinarily rapid vegetation responses. Hence, if scattered populations are not provided and tree populations are to migrate long distances, their capacity to keep up with predicted rates of future climate change may be lower than previously thought.

Eurasian large mammal dynamics in response to changing environments during the Late Neogene

Nisäkkäiden levinneisyyteen, niiden morfologisiin ja ekologisiin piirteisiin vaikuttavat ympäristön sekä lyhyet että pitkäkestoiset muutokset, etenkin ilmaston ja kasvillisuuden vaihtelut. Työssä tutkittiin nisäkkäiden sopeutumista ilmastonmuutoksiin Euraasiassa viimeisen 24 miljoonan vuoden aikana. Tutkimuksessa keskityttiin varsinkin viimeiseen kahteen miljoonaan vuoteen, jonka aikana ilmasto muuttui voimakkaasti ja ihmisen toiminta alkoi tulla merkittäväksi. Tämän takia on usein vaikea erottaa, kummasta em. seikasta jonkin nisäkäslajin sukupuutto tai häviäminen alueelta johtui. Aineistona käytettiin laajaa venäjänkielistä kirjallisuutta, josta löytyvät tiedot ovat kääntämättöminä jääneet aiemmin länsimaisen tutkimuksen ulkopuolelle. Työssä käytettiin myös NOW-tietokantaa, jossa on fossiilisten nisäkkäiden löytöpaikat sekä niiden iät.

Learning from the past to predict the future: an historical analysis of grass invasions in northern Australia

An important focus of biosecurity is anticipating future risks, but time lags between introduction, naturalisation, and (ultimately) impact mean that future risks can be strongly influenced by history. We conduct a comprehensive historical analysis of tropical grasses (n = 155) that have naturalised in Australia since European settlement (1788) to determine what factors shaped historical patterns of naturalisation and future risks, including for the 21 species that cause serious negative impacts. Most naturalised species were from the Old World (78 %), were introduced for use in pasture (64.5 %), were first recorded prior to 1940 (84.5 %) and naturalised before 1980 (90.3 %). Patterns for high-impact species were similar, with all being first recorded in Australia by 1940, and only seven naturalised since then-five intentionally introduced as pasture species. Counter to expectations, we found no evidence for increased naturalisation with increasing trade, including for species introduced unintentionally for which the link was expected to be strongest. New pathways have not emerged since the 1930s despite substantial shifts in trading patterns. Furthermore, introduction and naturalisation rates are now at or approaching historically low levels. Three reasons were identified: (1) the often long lag phase between introduction and reported naturalisation means naturalisation rates reflect historical trends in introduction rates; (2) important introduction pathways are not directly related to trade volume and globalisation; and (3) that species pools may become depleted. The last of these appears to be the case for the most important pathway for tropical grasses, i.e. the intentional introduction of useful pasture species. Assuming that new pathways don't arise that might result in increased naturalisation rates, and that current at-border biosecurity practices remain in place, we conclude that most future high-impact tropical grass species are already present in Australia. Our results highlight the need to continually test underlying assumptions regarding future naturalisation rates of high-impact invasive species, as conclusions have important implications for how best to manage future biosecurity risks.

The range expansion of the European map butterfly in Finland

Climate change will influence the living conditions of all life on Earth. For some species the change in the environmental conditions that has occurred so far has already increased the risk of extinction, and the extinction risk is predicted to increase for large numbers of species in the future. Some species may have time to adapt to the changing environmental conditions, but the rate and magnitude of the change are too great to allow many species to survive via evolutionary changes. Species responses to climate change have been documented for some decades. Some groups of species, like many insects, respond readily to changes in temperature conditions and have shifted their distributions northwards to new climatically suitable regions. Such range shifts have been well documented especially in temperate zones. In this context, butterflies have been studied more than any other group of species, partly for the reason that their past geographical ranges are well documented, which facilitates species-climate modelling and other analyses. The aim of the modelling studies is to examine to what extent shifts in species distributions can be explained by climatic and other factors. Models can also be used to predict the future distributions of species. In this thesis, I have studied the response to climate change of one species of butterfly within one geographically restricted area. The study species, the European map butterfly (Araschnia levana), has expanded rapidly northwards in Finland during the last two decades. I used statistical and dynamic modelling approaches in combination with field studies to analyse the effects of climate warming and landscape structure on the expansion. I studied possible role of molecular variation in phosphoglucose isomerase (PGI), a glycolytic enzyme affecting flight metabolism and thereby flight performance, in the observed expansion of the map butterfly at two separate expansion fronts in Finland. The expansion rate of the map butterfly was shown to be correlated with the frequency of warmer than average summers during the study period. The result is in line with the greater probability of occurrence of the second generation during warm summers and previous results on this species showing greater mobility of the second than first generation individuals. The results of a field study in this thesis indicated low mobility of the first generation butterflies. Climatic variables alone were not sufficient to explain the observed expansion in Finland. There are also problems in transferring the climate model to new regions from the ones from which data were available to construct the model. The climate model predicted a wider distribution in the south-western part of Finland than what has been observed. Dynamic modelling of the expansion in response to landscape structure suggested that habitat and landscape structure influence the rate of expansion. In southern Finland the landscape structure may have slowed down the expansion rate. The results on PGI suggested that allelic variation in this enzyme may influence flight performance and thereby the rate of expansion. Genetic differences of the populations at the two expansion fronts may explain at least partly the observed differences in the rate of expansion. Individuals with the genotype associated with high flight metabolic rate were most frequent in eastern Finland, where the rate of range expansion has been highest.