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.

Glaciers and climate in northern Sweden during the 19th and 20th century

Our understanding of the climate of northern Sweden during the late Holocene is largely dependent on proxy-data series. These datasets remain spatially and temporally sparse and instrumental series are rare prior to the mid 19th century. Nevertheless, the glaciology and paleo-glaciology of the region has a strong potential significance for the exploration of climate change scenarios, past and future. The aim of this thesis is to investigate the 19th and 20th century climate in the northern Swedish mountain range. This provides a good opportunity to analyse the natural variability of the climate before the onset of the industrial epoch. Developing a temporal understanding of fluctuations in glacier front positions and glacier mass balance that is linked to a better understanding of their interaction and relative significance to climate is fundamental in the assessment of past climate. I have chosen to investigate previously unexplored temperature data from northern Sweden from between 1802 and 1860 and combined it with a temperature series from a synoptic station in Haparanda, which began operation in 1859, in order to create a reliable long temperature series for the period 1802 to 2002. I have also investigated two different glaciers, Pårteglaciären and Salajekna, which are located in different climatic environments. These glaciers have, from a Swedish perspective, long observational records. Furthermore, I have investigated a recurring jökulhlaup at the glacier Sälkaglaciären in order to analyse glacier-climate relationships with respect to the jökulhlaups. A number of datasets are presented, including: glacier frontal changes, in situ and photogrammetric mass balance data, in situ and satellite radar interferometry measurements of surface velocity, radar measurements, ice volume data and a temperature series. All these datasets are analysed in order to investigate the response of the glaciers to climatic stimuli, to attribute specific behaviour to particular climates and to analyse the 19th and 20th century glacier/climate relationships in northern Sweden. The 19th century was characterized by cold conditions in northern Sweden, particularly in winter. Significant changes in the amplitude of the annual temperature cycle are evident. Through the 19th century there is a marked decreasing trend in the amplitude of the data, suggesting a change towards a prevalence of maritime (westerly) air masses, something which has characterised the 20th century. The investigations on Salajekna support the conclusion that the major part of the 19th century was cold and dry. The 19th century advance of Salajekna was probably caused by colder climate in the late 18th and early 19th centuries, coupled with a weakening of the westerly airflow. The investigations on Pårteglaciären show that the glacier has a response time of ~200 years. It also suggests that there was a relatively high frequency of easterly winds providing the glacier with winter precipitation during the 19th century. Glaciers have very different response times and are sensitive to different climatic parameters. Glaciers in rather continental areas of the Subarctic and Arctic can have very long response times because of mass balance considerations and not primarily the glacier dynamics. This is of vital importance for analyzing Arctic and Subarctic glacier behaviour in a global change perspective. It is far from evident that the behaviour of the glacier fronts today reflects the present climate.