Spring Phenology of Butterflies : The role of seasonal variation in life-cycle regulation

Animals and plants in temperate regions must adapt their life cycle to pronounced seasonal variation. The research effort that has gone into studying these cyclical life history events, or phenological traits, has increased greatly in recent decades. As phenological traits are often correlated to temperature, they are relevant to study in terms of understanding the effect of short term environmental variation as well as long term climate change. Because of this, changes in phenology are the most obvious and among the most commonly reported responses to climate change. Moreover, phenological traits are important for fitness as they determine the biotic and abiotic environment an individual encounters. Fine-tuning of phenology allows for synchronisation at a local scale to mates, food resources and appropriate weather conditions. On a between-population scale, variation in phenology may reflect regional variation in climate. Such differences can not only give insights to life cycle adaptation, but also to how populations may respond to environmental change through time. This applies both on an ecological scale through phenotypic plasticity as well as an evolutionary scale through genetic adaptation. In this thesis I have used statistical and experimental methods to investigate both the larger geographical patterns as well as mechanisms of fine-tuning of phenology of several butterfly species. The main focus, however, is on the orange tip butterfly, Anthocharis cardamines, in Sweden and the United Kingdom. I show a contrasting effect of spring temperature and winter condition on spring phenology for three out of the five studied butterfly species. For A. cardamines there are population differences in traits responding to these environmental factors between and within Sweden and the UK that suggest adaptation to local environmental conditions. All populations show a strong negative plastic relationship between spring temperature and spring phenology, while the opposite is true for winter cold duration. Spring phenology is shifted earlier with increasing cold duration. The environmental variables show correlations, for example, during a warm year a short winter delays phenology while a warm spring speeds phenology up. Correlations between the environmental variables also occur through space, as the locations that have long winters also have cold springs. The combined effects of these two environmental variables cause a complex geographical pattern of phenology across the UK and Sweden. When predicting phenology with future climate change or interpreting larger geographical patterns one must therefore have a good enough understanding of how the phenology is controlled and take the relevant environmental factors in to account. In terms of the effect of phenological change, it should be discussed with regards to change in life cycle timing among interacting species. For example, the phenology of the host plants is important for A. cardamines fitness, and it is also the main determining factor for oviposition. In summary, this thesis shows that the broad geographical pattern of phenology of the butterflies is formed by counteracting environmental variables, but that there also are significant population differences that enable fine-tuning of phenology according to the seasonal progression and variation at the local scale.

Finding the way home: movement of butterflies in non-familiar habitats

Paisagens naturais tem sido afetadas dramaticamente pela perda de habitat e fragmentação, os quais transformam paisagens contínuas em manchas de habitat circundadas por áreas de não-habitat (matrizes). Essas matrizes, inóspitas ou não, afetam incontáveis processos ecológicos, como a dispersão. Uma das formas de compreender o efeito da matriz no processo de dispersão é estudando o alcance da percepção do habitat dos animais, o qual é definido como o alcance (distância) em que um animal pode perceber elementos da paisagem. Essa percepção está diretamente relacionada ao sucesso de chegada à um novo habitat, enquanto o animal navega por uma matriz. Nós avaliamos a percepção do habitat em Heliconius erato, uma borboleta tropical. Entretanto nós também estávamos interessados em avaliar o efeito da idade das borboletas e do tipo de matriz na percepção do habitat. Consequentemente, nós criamos borboletas em laboratório e pareamos com borboletas coletadas na floresta durante um experimento de soltura. Para determinar o alcance da percepção, nós realizamos solturas em duas diferentes matrizes (coqueiral e campo em regeneração) à três distâncias da floresta (0,30 e 100 metros) e medimos o ângulo final alcançado pelas borboletas. Nós encontramos que (I) borboletas soltas na borda se orientaram fortemente para a floresta; (II) quanto maior a distância de soltura, menor a habilidade de perceber o habitat e (III) há interação entre matriz e idade da borboleta. Borboletas inexperientes se orientaram melhor no campo aberto (alcance da percepção: 30-100 metros) e experientes se orientaram melhor no coqueiral (alcance da percepção entre 30-100 metros).

Silk Butterflies

The first section of this unfinished novel, titled Silk Butterflies is a diptych about a woman named Sarah, and her desire to acquire ancestral truth regarding her identity to negate the pain she feels from losing her unborn child. Her story, told in a guarded, first person point-of-view is paralleled with Ling’s story, an unconventional, ninety-two year old Shanghainese woman who, against her desires, had her feet bound in China during the early 1920’s. Ling’s story is also told from a lyrical first-person perspective that focuses especially on sensory details, and delves into the sacrifices we make to attain standards of beauty, and the loss Ling has never recovered from. As this historical fiction progresses, their stories overlap in an unexpected way, as both Sarah and Ling attempt to revitalize forgotten histories, including how Sarah’s grandparents fled to Shanghai in the 1930’s to escape Nazi persecution during World War II.

Chasing butterflies: The Conception of the Soul in Plutarch’s Works

When first approaching a topic such as the concept of material or natural soul in Greek literature, the researcher might be puzzled. While in diverse contemporary cultures, numerous theriomorphic figures (bears, ravens, mice, wasps, bees, dragonflies, and dung-beetles) serve to represent the human soul in its transmigration from life to death, this is not the case in Greek culture. At least, this is what one may conclude from the monograph written by the Dutch scholar J. Bremmer, The early Greek Concept of the Soul: "importunely, there are no other indications of a possible connection between the butterfly and the soul of the living and the dead" (1987: 64). Given Plutarch’s great interest in the soul, which can be seen in a variety of texts referring to its generation, form, internal dichotomy, material substance, origin and destination, etc., the question arises as to whether Plutarch also includes such a representation of the soul when departing from the dead body. Does the corpus plutarcheum preserve and transmit such conception of the human soul? And if it does, are we dealing with survival of ancestral beliefs or motifs or is it a simple metaphor by means of which ancients intended to express the departing of the life-breath? In the following pages I will focus on three texts that allegedly include the butterfly-motif to represent the human soul, to wit, Table Talks 636C, Consolation to his Wife 611F, and the fragment 177 Sandbach.

A butterfly eye's view of birds

The striking color patterns of butterflies and birds have long interested biologists. But how these animals see color is less well understood. Opsins are the protein components of the visual pigments of the eye. Color vision has evolved in butterflies through opsin gene duplications, through positive selection at individual opsin loci, and by the use of filtering pigments. By contrast, birds have retained the same opsin complement present in early-jawed vertebrates, and their visual system has diversified primarily through tuning of the short-wavelength-sensitive photoreceptors, rather than by opsin duplication or the use of filtering elements. Butterflies and birds have evolved photoreceptors that might use some of the same amino acid sites for generating similar spectral phenotypes across approximately 540 million years of evolution, when rhabdomeric and ciliary-type opsins radiated during the early Cambrian period. Considering the similarities between the two taxa, it is surprising that the eyes of birds are not more diverse. Additional taxonomic sampling of birds may help clarify this mystery.

Finding out about children's meaning making through drawing together

When I was seven I worked on a science project about caterpillars and moths. I was completely immersed in this project, fascinated by caterpillar body markings, the rhythmical, semi-circular pattern caterpillars adopt to eat leaves, their spiral construction of the chrysalis, and their transformation into moths or butterflies. I demonstrated my fascination, my research and study through carefully executed and detailed drawings. I could read and write well, but I wasn’t as interested in writing and produced a half-page summary to support my visual work.

Learning in the nectar foraging behaviour of Helicoverpa armigera

Abstract. 1. Learning may enable insects to obtain nectar from flowers more efficiently. Learning in nectar foraging has been shown primarily in studies of bees and butterflies. Here, learning is demonstrated in the nectar foraging behaviour of a noctuid moth, Helicoverpa armigera. 2. The present studies show that: (1) previous experience with a flowering host species increases the probability of that species being selected for nectar foraging, and (2) previous experience of a particular flower type (food source at bottom or top of the corolla tube) increases the likelihood of the food source being found when that flower type is being searched. 3. The implications of these findings for understanding the pattern of oviposition observed in wild populations of this important pest species are discussed.

Gustaf III inför sina undersåtar : Konsten att härska och politikens kulturhistoria

The dissertation analyses the political culture of Sweden during the reign of King Gustav III (1771-1792). This period commonly referred to as the Gustavian era followed the so-called Age of Liberty ending half a century of strong parliamentary rule in Sweden. The question at the heart of this study engages with the practice of monarchical rule under Gustav III, its ideological origins and power-political objectives as well as its symbolic expression. The study thereby addresses the very nature of kingship. In concrete terms, why did Gustav III, his court, and his civil service vigorously pursue projects that contemporaneous political opponents and, in particular, subsequent historiography have variously pictured as irrelevant, superficial, or as products of pure vanity? The answer, the study argues, is to be found in patterns of political practice as developed and exercised by Gustav III and his administration, which formed a significant part of the political culture of Gustavian Sweden. The dissertation is divided into three parts. The first traces the use and development of royal graces chivalric orders, medals, titles, privileges, and other gifts issued by the king. The practice of royal reward is illustrated through two case studies: the 1772 coup d état that established Gustav III s rule, and the birth and baptism of the crown prince, Gustav Adolf, in 1778. The second part deals with the establishment of the Court of Appeal in Vasa in 1776. The formation of the Appeals Court was accompanied by a host of ceremonial, rhetorical, emblematic, and architectural features solidifying its importance as one of Gustav III s most symbolic administrative reform projects and hence portraying the king as an enlightened monarch par excellence. The third and final part of the thesis engages with war as a cultural phenomenon and focuses on the Russo-Swedish War of 1788-1790. In this study, the war against Russia is primarily seen as an arena for the king and other players to stage, create and re-create as well as articulate themselves through scenes and roles adhering to a particular cultural idiom. Its codes and symbolic forms, then, were communicated by means of theatre, literature, art, history, and classical mythology. The dissertation makes use of a host of sources: protocols, speeches, letters, diaries, newspapers, poetry, art, medals, architecture, inscriptions and registers. Traditional political source material and literary and art sources are studied as totalities, not as separate entities. Also it is argued that political and non-fictional sources cannot be understood properly without acknowledging the context of genre, literary conventions, and artistic modes. The study critically views the futile, but nonetheless almost habitual juxtaposition of the reality of images, ideas, and metaphors, and the reality of supposedly factual historical events. Significantly, the thesis presumes the symbolic dimension to be a constitutive element of reality, not its cooked up misrepresentation. This presumption is reflected in a discussion of the concept of role , which should not be anachronistically understood as roles in which the king cast himself at different times and in different situations. Neither Gustav III nor other European sovereigns of this period played the roles as rulers or majesties. Rather, they were monarchs both in their own eyes and in the eyes of their contemporaries as well as in all relations and contexts. Key words: Eighteenth-Century, Gustav III, Cultural History, Monarchs, Royal Graces, the Vasa Court of Appeal, the Russo-Swedish War 1788–1790.

Isänmaamme keisari : Eliitin kansallisen identiteetin murros ja suomalaisen isänmaan rakentuminen autonomian ajan alussa

The emperor of our fatherland The changing national identity of the elite and the construction of the Finnish fatherland at the beginning of the autonomy This study addresses the question of changing national identity of the elite at the beginning of the autonomy (1808 1814) in Finland. Russia had conquered Finland from Sweden, but Finland was not incorporated into the Russian Empire. Instead, it was governed as separately administered area, and Finland retained its own (laws of the realm of Sweden) laws. The inclusion in the Russian Empire compelled the elite of Finland to deliberate their national identity; they had to determine whether they remained Swedes or became Finns or Russians. The elite chose to become Finns, which may seem obvious from the nowadays perspective, but it cannot be taken for granted that the Swedish speaking and noble elite converted their local Finnish identity into a new national identity. The basis of this study is constructive in a sense that identity is not seen as stable and constant. Theoretical background lies on Stuart Hall s writings on national identity, which offer good practical methods to study national identity. According to Hall identity is based mainly on difference , difference to others . In practice this means how elite began to define themselves in contrast to Swedes and Russians. The Finnish national identity was constructed in contrast to Swedes due to the political reasons. In order to avoid Russians suspicions Finns had to diverge from Sweden. Sweden had also gone trough coup d état, which was disliked by the elite of Finland. However, the attitudes of the elite towards Sweden remained somewhat ambiguous. Even if it was politically and rationally thinking wisest to draw away from Sweden, emotionally it was difficult. Russia, on the other hand, had been for centuries the archenemy of the Finns as well as all the Swedes. The fear of the Russians was mainly imaginary. Russians were seen as cruel barbarians who hated and resented Finns. The Finnish national identity was constructed above all in contrast to the Russians, for the difference to Russia was seen as a precondition for the existence of Finland. Respectively, the new position of Finland also required approaching towards Russia, which was in its nature very pragmatic. The elite contrived to get rid off its prejudice against Russians on intellectual level, but not on emotional level. At the beginning of the autonomy the primary loyalty of the elite was directed into the Finnish fatherland and its habitants. This was a radical ideological change, because traditionally the loyalty of the elite had focused on monarch and monarch s realm. However, the role of Alexander I was crucial. According to the elite the emperor had granted them a new fatherland. The former native country (Finland) was seen as a new fatherland instead of Sweden. The loyalty of the elite to the emperor generated from the reciprocal gratitude; Alexander I had treated their native country so mercifully. The elite felt strong personal responsibility for Finland s existence. The elite believed that the future of Finland rested on their shoulders. Alexander I had given them fatherland, but it was in the hands of the elite to construct the Finnish state and national spirit. The study of the Finnish national identity brings forth also that the national identity was constructed by emphasizing Finns civic rights. The civic rights were essential part of the construction of the Finnish national identity, for the difference between Finns and Russians was based on Finns own laws and privileges, which the emperor of the Russia had ensured.

Martha Smith

[Excerpt] Martha Smith said goodbye to her long withstanding career as Associate Director of ILR’s Office of Career Services. Smith earned a B.S. in Business Administration from California State University, Hayward in 1969. In 1976, she came to Ithaca when her husband was a Ph.D. student at Cornell, and in 1978 Smith became a temporary assistant to Professor Larry Williams. Shortly after, Smith was offered a permanent position in the Office of Career Services where she advised and befriended students for over 26 years. After retiring, Smith plans to devote more time to her community and her church. When reflecting upon her career of assisting and nurturing in the development of ILR students, Smith said, "They come in as caterpillars; they evolve, and then they leave as butterflies. They have to learn to be students, it isn’t always easy Martha Smith but they always manage. As they open up we all learn that we are more alike than we are different and together we learn to appreciate the likenesses and the differences."

A new culture method for lesser mealworm, Alphitobius diaperinus

A new culture method for lesser mealworm, Alphitobius diaperinus (Panzer), was developed to provide large numbers of adult lesser mealworms of approximately the same age for insecticide resistance testing. Culturing entailed allowing 100 adults to reproduce for 4 days in a wheat-based culture medium contained inside a plastic culture box, removing the adults from the medium, and then rearing their progeny to adulthood therein, in approximately 56 days at 32 degrees C and 55% RH. During their development, progeny were supplied water via apple slices at 0, 21 and 35 days, and a foam substrate in which to pupate, also at 35 days. During 2004-2005, adult lesser mealworms were collected from six broiler-house populations and then cultured with this method. Each population produced 4500 adults required to complete resistance testing with one insecticide within ten culture boxes, at an average of 798 adults per culture box.

Dietary media for mass rearing of rusty grain beetle, Cryptolestes ferrugineus (Stephens) and flat grain beetle, Cryptolestes pusillus (Schonherr) (Coleoptera: Cucujidae)

We developed a suitable diet for mass rearing of Cryptolestes ferrugineus (Stephens) populations under laboratory conditions. Recently, this pest has developed strong level of resistance to phosphine in Australia, and therefore, a significant amount of research has been directed towards its management. In total, nineteen grain-based diets, containing rolled oats, various combinations of cracked grains and flours of wheat, sorghum, maize and barley were tested. Each diet contained a small proportion of wheat germ (4.5% w/w) and torula yeast (0.5% w/w). Experiments were conducted at fixed temperature and relative humidity regimes of 30 ± 2 °C and 70 ± 2%, respectively, and replicated three times. Adults (n = 40) of a laboratory strain of C. ferrugineus were introduced into each diet, removed after 14 days and total numbers of live adult progeny were recorded. The following diets resulted in highest live progeny production: barley flour (95%) (607.67 ± 11.21) = rolled oats (75%) + cracked sorghum (20%) (597.33 ± 33.79) ≥ wheat flour (47.5%) + barley flour (47.5%) (496.67 ± 52.93) > cracked sorghum (95%) (384.00 ± 60.66). The performance of these four diets was then tested with field-collected populations of C. ferrugineus and Cryptolestes pusillus (Schonherr). The diets based on rolled oats + cracked sorghum, wheat flour + barley flour, and barley flour alone consistently produced highest progeny numbers in field-collected populations of both species, with mean progeny numbers ranging from 359.9 to 478.5. The multiplication of C. pusillus was significantly higher than C. ferrugineus on all four diets. Our findings will help in mass rearing of healthy cultures of C. ferrugineus and C. pusillus that will greatly facilitate laboratory and field research and in particular, in developing management tactics for these species.

Modeling ecological and evolutionary processes in spatially structured populations

Many species inhabit fragmented landscapes, resulting either from anthropogenic or from natural processes. The ecological and evolutionary dynamics of spatially structured populations are affected by a complex interplay between endogenous and exogenous factors. The metapopulation approach, simplifying the landscape to a discrete set of patches of breeding habitat surrounded by unsuitable matrix, has become a widely applied paradigm for the study of species inhabiting highly fragmented landscapes. In this thesis, I focus on the construction of biologically realistic models and their parameterization with empirical data, with the general objective of understanding how the interactions between individuals and their spatially structured environment affect ecological and evolutionary processes in fragmented landscapes. I study two hierarchically structured model systems, which are the Glanville fritillary butterfly in the Åland Islands, and a system of two interacting aphid species in the Tvärminne archipelago, both being located in South-Western Finland. The interesting and challenging feature of both study systems is that the population dynamics occur over multiple spatial scales that are linked by various processes. My main emphasis is in the development of mathematical and statistical methodologies. For the Glanville fritillary case study, I first build a Bayesian framework for the estimation of death rates and capture probabilities from mark-recapture data, with the novelty of accounting for variation among individuals in capture probabilities and survival. I then characterize the dispersal phase of the butterflies by deriving a mathematical approximation of a diffusion-based movement model applied to a network of patches. I use the movement model as a building block to construct an individual-based evolutionary model for the Glanville fritillary butterfly metapopulation. I parameterize the evolutionary model using a pattern-oriented approach, and use it to study how the landscape structure affects the evolution of dispersal. For the aphid case study, I develop a Bayesian model of hierarchical multi-scale metapopulation dynamics, where the observed extinction and colonization rates are decomposed into intrinsic rates operating specifically at each spatial scale. In summary, I show how analytical approaches, hierarchical Bayesian methods and individual-based simulations can be used individually or in combination to tackle complex problems from many different viewpoints. In particular, hierarchical Bayesian methods provide a useful tool for decomposing ecological complexity into more tractable components.

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.