Climate change and invasion impacts in the sub-Antarctic

Climate change and biological invasions are major threats to biodiversity. In particular, these threats are predicted to influence terrestrial systems in the sub-Antarctic, where significant ecosystem responses to both have already been seen. In this thesis, the sub-Antarctic Prince Edward Island group is used as a model system in which to investigate key questions relating to climate change and invasive species impacts. The island group comprises two islands, Marion (MI) and Prince Edward (PEI), both of which are experiencing rapid warming, yet have different invasive assemblages and in consequence are experiencing different impacts. Variation in the patterns of invasive species richness and abundance and their underlying causes are matters of considerable ecological and conservation significance. While an increase in thermal energy availability typically results in an increase in species richness, the mechanisms underlying these patterns are poorly understood. In Chapter 2 of this thesis, these relationships are explored for springtails, an important component of the soil fauna on Marion Island. Energy explains a large amount of the spatial variation in indigenous and invasive springtail species richness. Disturbance thresholds and stressful temperatures are more important than increased population sizes in determining this variation in species richness. As both indigenous and invasive springtail species richness and abundance are strongly related to temperature, a warming climate could have far-reaching consequences for these organisms. In particular, invasive species are predicted to be at an advantage relative to indigenous species under warming conditions. One species where this seems especially likely, given its physiological responses to experimental warming and drying, is the large invasive tomocerid, Pogonognathellus flavescens. Determining whether this will be the case depends on understanding the factors underlying its range limits and abundance structure. Moreover, few studies have sought to distinguish the causal basis of abundance structure and range limits, particularly for invasive species. Thus, in Chapter 3, local microclimate variables and physiological tolerances of the invasive springtail, P. flavescens (a habitat generalist), are examined. The results suggest that the species should be widely distributed across a range of habitats on MI. However, the springtail is restricted to indigenous Poa cookii tussock grassland habitats in the southeast. The current range limits are set by dispersal limitation (i.e. contingent absences) whilst abundance structure is a function of variation in soil substrate quality. However, over time, the widening distribution of P. cookii, as a consequence of a major management intervention (the eradication of feral cats), may enable P. flavescens to colonise all suitable areas. In Chapter 4, the focus changes to what has been considered the third major response to climate change, along with range and phenological responses - changing animal body sizes. Body size is one of the most significant and obvious features of animals and is of considerable ecological and physiological importance. A prediction of the temperature-size rule (TSR) is that with warming, body size of the weevil species on both MI and PEI should decline. However, predation by mice of the weevils on MI should fundamentally affect the pattern of such change, causing it to differ from neighbouring PEI, indicating synergistic impacts between climate change and invasions. Analysis of a 24-year data set indicates a decline in the body size of all weevil species on PEI with increasing temperature. However, on MI, a negative relationship between mean annual temperature and body size is found only for Palirhoeus eatoni, a species not eaten by mice. A possible explanation for the positive relationships found for the other species could be due to higher metabolic demands imposed on mice in colder years than in warmer ones. Any increase in predation coupled with a preference for larger sizes, which the mice clearly show, would lead to a decline in the mean size of the weevil species. Due to the relationship between body size and metabolic rate and the importance of the weevils in the islands' food webs, changes to the body size of these organisms could have significant consequences for the island ecosystems' functioning. The thermal environment experienced by organisms also has a direct effect on survival, growth and reproduction. The physiological response of organisms to rapidly changing climates is therefore a primary concern. Organisms may respond to variable environmental conditions through phenotypic plasticity as well as behaviour. Chapter 5 of this thesis shows that of the weevil species and populations investigated on MI, most display phenotypic plasticity, the form of which is in keeping with the 'Hotter is Better' hypothesis. This could be due to rare extreme temperature events and the advantage for the performance curves to incorporate high temperatures experienced in the environment. Mismatches between thermal optima and preferred temperatures displayed by all species could mean that these weevils are well equipped to cope with warming conditions on MI unless the prediction of an increase of rare extreme events such as extreme temperatures is realised. Rapidly changing climates and an increase in the introduction of non-indigenous species are issues of major conservation concern. This has increased the significance of studies on the impacts of these threats. However, this thesis shows that to understand such processes, it is essential that an integration of disciplines be undertaken. This thesis thus adopts a multidisciplinary approach and highlights key issues associated with both climate change and biological invasions. The patterns and predictions of species and community responses to these environmental changes are complex. Moreover, predicting such responses is likely to be problematic, especially as multiple factors will change concurrently and how these factors might change is unclear. This highlights the importance of long-term records for understanding organism responses to such changes. Furthermore, impacts on indigenous species are likely to be exacerbated by the predicted increase in the rate of introductions with climate change. This makes the case for preventing the dispersal of invasive species to new areas all the more important.