Biological invasions are an important issue of global change and an increased understanding of invasion processes is of crucial importance for both conservation managers and international trade. In this thesis, I have studied the invasion of the brown seaweed Fucus evanescens, to investigate the fate and effect of a perennial, habitat-forming seaweed introduced to a coastal ecosystem. A long-term study of the spread of F. evanescens in Öresund (southern Sweden) showed that the species was able to expand its range quickly during the first 20 years after the introduction, but that the expansion has been slow during the subsequent 30 years. Both in Öresund and in Skagerrak, the species is largely restricted to sites where native fucoids are scarce. Laboratory experiments showed that the restricted spread of F. evanescens cannot be explained by the investigated abiotic factors (wave exposure and salinity), although salinity restricts the species from spreading into the Baltic Sea. Neither did I find evidence for that herbivores or epibiota provide biotic resistance to the invader. On the contrary, F. evanescens was less consumed by native herbivores, both compared to the native fucoids and to F. evanescens populations in its native range, and little overgrown by epiphytes. Instead, the restricted spread may be due to competition from native seaweeds, probably by pre-occupation of space, and the establishment has probably been facilitated by disturbance. The studies provided little support for a general enemy release in introduced seaweeds. The low herbivore consumption of F. evanescens in Sweden could not be explained by release from specialist herbivores. Instead, high levels of chemical anti-herbivore defence metabolites (phlorotannins) could explain the pattern of herbivore preference for different fucoids. Likewise, the low epibiotic colonisation of F. evanescens plants could be explained by high resistance to epibiotic survival. This shows that colonisation of invading seaweeds by native herbivores and epibionts depends on properties of the invading species. The large differences between fucoid species in their quality as food and habitat for epibionts and herbivores imply that invasions of such habitat-forming species may have a considerable effect on a number of other species in shallow coastal areas. However, since F. evanescens did not exclude other fucoids in its new range, its effect on the recipient biota is probably small.

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Invasive exotic species threaten native biodiversity, alter ecosystem structure and function, and annually cost over $100 billion in the US alone. Determining the ecological traits and interactions that affect invasion success are thus critical for predicting, preventing, and mitigating the negative effects of biological invasions. Native herbivores are widely assumed to facilitate exotic plant invasions by preferentially consuming native plants and avoiding exotic plants. Here, I use freshwater plant communities scattered broadly across the Southeastern U.S. to show that herbivory is an important force driving the ecology and evolution of freshwater systems. However, native consumers often preferentially consume rather than avoid exotic over native plants. Analyses of 3 terrestrial datasets showed similar patterns, with native herbivores generally preferring exotic plants. Thus, exotic plants appear defensively nave against these evolutionarily novel consumers, and exotic plants may escape their coevolved, specialist herbivores only to be preferentially consumed by the native generalist herbivores in their new ranges. In further support of this hypothesis, a meta-analysis of 71 manipulative field studies including over 100 exotic plant species and 400 native plant species from terrestrial, aquatic, and marine systems revealed that native herbivores strongly suppressed exotic plants, while exotic herbivores enhanced the abundance and species richness of exotic plants by suppressing native plants. Both outcomes are consistent with the hypothesis that prey are susceptible to evolutionarily novel consumers. Thus, native herbivores provide biotic resistance to plant invasions, but the widespread replacement of native with exotic herbivores eliminates this ecosystem service, facilitates plant invasions, and triggers an invasional meltdown. Consequently, rather than thriving because they escape their co-evolved specialist herbivores, exotic plants may thrive because their co-evolved generalist herbivores have stronger negative effects on evolutionarily nave, native plants.

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Harmonia axyridis is a ladybird native to Asia, but introduced widely as a biocontrol agent. It is invasive and detrimental to native species in North America, which meant its arrival in Britain was met with concern. Establishment was seen as an opportunity to track the spread of an invasive alien species (IAS) whilst also monitoring impacts on native species. The aims of this thesis were to examine the responses of native British ladybirds to the arrival of H. axyridis, to establish the effect of the IAS on native ladybirds when compared to other drivers, and to investigate the possible facilitation of the H. axyridis invasion by natural enemy release. Modelling ladybird distributions with life-history and resource-use traits found that species predatory on a wide range of prey families had larger range sizes than those which ate fewer prey types. This suggests that the wide diet breadth of the IAS is likely to have played a critical role in the species’ rapid spread. Dietary niche overlap between H. axyridis and native ladybirds showed positive correlation with declines of native ladybirds. This indicates that the IAS is playing an important role, but the significance of urbanisation suggests habitat destruction is also significant. Abundance of H. axyridis was influenced by habitat type and aphid abundance, but not by the native ladybird community, suggesting the spread of the IAS will not be slowed by biotic resistance. Harmonia axyridis is attacked by native parasitoids, but at a much lower rate than is the native Coccinella septempunctata, in line with natural-enemy release theory. There was no evidence of attack rate increasing with time since arrival in an area. Overall, H. axyridis is an extremely successful IAS, with detrimental effects on native ladybirds which are likely to continue.

Cirsium arvense (L.) Scop. (Californian, Canada, or creeping thistle) is an exotic perennial herb indigenous to Eurasia that successfully established in New Zealand (NZ) approximately 130 years ago. Presently, C. arvense is considered one of the worst invasive weeds in NZ arable and pastoral productions systems. The mechanism most commonly invoked to explain the apparent increased vigour of introduced weeds is release from natural enemies. The enemy release hypothesis (ERH) predicts that plants in an introduced range should experience reduced herbivory, particularly from specialists, and that release from this natural enemy pressure facilitates increased plant performance in the introduced range. In 2007 broad surveys were carried out in NZ and central Europe in order to quantify and compare growth characteristics of C. arvense in its native vs. introduced range. Additionally, permanent field plots were established in NZ and Europe where natural enemies were excluded with the use of insecticide and fungicide applications, and compared with controls (ambient natural enemy pressure). The impact of the specialist leaf-feeding beetle, Cassida rubiginosa Müller, which was recently released in NZ as a biological control agent against thistles, was also assessed. From the field surveys, significantly more endophagous herbivory was present in the native range compared to the introduced range, as predicted by the ERH. Endophagous herbivory in NZ was solely from the capitulum-feeding weevil, Rhinocyllus conicus (Frölich), and was only found in the North Island surveys. No stem mining attack was found anywhere in NZ. The proportion of shoots attacked by the specialised rust pathogen, Puccinia punctiformis (Str.) Röhl., was similar in both the native and introduced ranges. Interestingly, this has casted doubt on the idea that stem-mining vectors, such as Ceratapion onopordi Kirby, are important for transmission of the rust pathogen. Contrary to the ERH, there were no significant difference in plant performance between the native and introduced ranges, or differences could be explained by simple climatic factors. Climate tended to be more favourable for growth of C. arvense in NZ. In the permanent field plots in the native range, population growth of C. arvense was significantly greater where natural enemies were excluded, suggesting that insect herbivores and pathogens might have a regulating influence on the population growth of this plant. Furthermore, the probability of shoots transitioning to the reproductive growth stage was enhanced when insect herbivores were excluded, indicating that natural enemies might influence plant development. The biological control agent C. rubiginosa reduced the growth of C. arvense, although the impact of this herbivore was minimal in comparison to interspecific plant competition. Thus, although there is reduced specialist natural enemy pressure in NZ, the growth of C. arvense is not significantly different from in its native range. Nevertheless, there is some evidence that natural enemies in the native range might have a regulating influence on the population dynamics of the plant, and that the specialist herbivore, C. rubiginosa, can impact the plant in certain conditions.