Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: Successful biological invasions take place when species introduced to regions outside their natural dispersal range overcome several barriers and establish, persist, proliferate and spread potentially resulting in major threats to biodiversity. The success of invasive alien plants depends on species-specific traits and characteristics of the introduced environment. In this thesis I explore which species traits are important and which environmental barriers need to be overcome for an invasion to occur using Proteaceae as a test case. Firstly, I assessed the global introduction history and invasion ecology of Proteaceae - a large plant family with many taxa that have been widely disseminated by humans, but with few known invaders. This revealed that at least 402 species (i.e. 24% of 1674 species in this family) are known to have been moved by humans out of their native ranges, 58 species (14%) have become naturalized and 8 species (2%) are invasive. The probability of naturalization was greatest for species with large native range sizes, low susceptibility to Phytophthora root-rot disease, larger seeds, mammal-dispersed seeds and those with the capacity to resprout after fire or other disturbances. The probability of naturalized species becoming invasive was greater for species with larger range sizes, species used as barrier plants, taller species, species with smaller seeds, serotinous species, and those that regenerated mainly through re-seeding. Secondly, I looked at mechanisms underlying naturalization on a regional scale, using species which are not already classified as major invaders. At least 26 non-native Proteaceae species have been introduced to, and are cultivated in, South Africa. Propagule pressure facilitated the naturalization of Hakea salicifolia populations in climatically suitable areas, but in suboptimal climates human-mediated land disturbance and land management activities are important for naturalization. Similar drivers are important for naturalization of other alien Proteaceae: a long residence time, fire regimes, poor land management, and propagule pressure were important mechanisms for naturalization. Thirdly, I determined whether reproduction, which in part drives propagule pressure, serves as a barrier for naturalization. I examined several Australian Proteaceae species introduced to South Africa and observed that all species were heavily utilized by native nectar-feeding birds and insects. The five Banksia species that were assessed are self-compatible but four species have a significantly higher reproductive output when pollinators visit inflorescences. Fruit production in H. salicifolia does not differ between naturally-pollinated and autonomously-fertilized flowers. Moreover, no significant difference in fruit production was observed between the five pollination treatments (i.e. natural, pollen-supplementation, autonomous, hand-selfed and hand-crossed treatments) and naturalized and non-naturalized populations. However, pollen limitation was detected in non-naturalized populations which received fewer pollinator visits than naturalized populations. Thus, reproduction limits but is not a fundamental barrier to invasion for H. salicifolia. I conclude that reproductive success of the studied Proteaceae, which is a key barrier determining invasiveness, is not limited by autonomous seed set or mutualisms in the introduced range. In this thesis I highlight biogeographical characteristics, a set of life-history traits and ecological traits as important determinants of invasiveness. These traits are in turn dependent on the stage of invasion. Characteristics of the recipient environment are also important drivers of invasions. This study provides a better understanding of plant invasions in general, but the patterns and processes of invasions highlighted in this thesis will be particularly useful for the current and future management of alien Proteaceae in South Africa and elsewhere, as well as, other species that are adapted to Mediterranean and nutrient poor ecosystems. For example, combining traits of invasiveness and susceptible environments will help to identify which non-native species pose a high risk of becoming invasive (e.g. species with large home ranges and barrier plants) and which conditions in the target area are likely to facilitate or exacerbate invasions (e.g. strong climate match and high propagule pressure). / AFRIKAANSE OPSOMMING: Suksesvolle biologiese indringing vind plaas wanneer ʼn spesie geïntroduseer word in ʼn area buite sy natuurlike verspreidings area, sekere versperrings oorkom, vestig, bly voortbestaan, vermenigvuldig en versprei en potensieel ʼn groot bedreiging inhou vir biodiversiteit. Die sukses van uitheemse indringer plante hang af van spesifieke kenmerke van die spesie en kenmerke van die omgewing waarin dit geïntroduseer word. In hierdie tesis maak ek gebruik van Proteaceae om te ondersoek watter kenmerke is belangrik en watter omgewing versperrings moet oorkom word vir indringing om plaas te vind. Ten eerste assesseer ek die wêreldwye introduksie geskiedenis en indringers ekologie van Proteaceae – ʼn groot plant familie wat wyd gebruik word deur mense, maar met min indringer spesies. Dit het gewys dat mense ten minste 402 spesies (dus 24% van die 1674 spesies in die familie) uit die inheemse areas verskuif het, 58 spesies (14%) genaturaliseer het en 8 spesies (2%) indringers geword het. Die moontlikheid van naturalisasie was die grootste vir spesies met ʼn groot inheemse streek, lae vatbaarheid vir Phytophthora wortelvrot, groter sade, dier verspreide sade en die met ʼn vermoë om weer uit te spruit na ʼn vuur of ander versteuring. Die moontlikheid van genaturaliseerde spesies om indringers te word, was groter vir spesies met groter streek grootte, spesies wat as versperring plante gebruik word, hoër spesies, spesies met kleiner sade, serotiniese spesies, en die wat hoofsaaklik voortbestaan as saadspruiters. Tweedens, het ek gekyk na onderliggende meganismes op ʼn regionale skaal, deur gebruik te maak van spesies wat nie alreeds as belangrike indringers geklassifiseer is nie. Ten minste 26 nie-inheemse Proteaceae spesies is alreeds geïntroduseer en word gekultiveer in Suid Afrika. Propaguul druk fasiliteer die naturalisering van Hakea salicifolia populasies in areas met geskikte klimaat, terwyl in areas met ʼn sub optimale klimaat, versteurings deur mense en grond bestuurs aktiwiteite belangrik is vir naturalisering. Die selfde drywers is belangrik vir die naturalisering van ander uitheemse Proteaceae: lang verblyftyd, vuur bestel, swak land bestuur en propaguul druk. Derdens het ek bepaal of reproduksie, wat gedeeltelik propaguul druk dryf, ʼn versperring is vir naturalisasie. Ek het gekyk na verskeie Australiese Proteaceae spesies wat geïntroduseer is in Suid Afrika, en het gevind dat al die spesies besoek word deur inheemse nektar etende voëls en insekte. Die vyf Banksia spesies wat geassesseer is, kan self bestuif, maar vier van die spesies het ʼn betekenisvolle hoër reproduksie wanneer bloeiwyses deur bestuiwers besoek word. Vrug produksie verskil nie tussen natuurlik bestuifde en self bestuifde blomme in H. salicifolia nie. Verder was daar geen verskil tussen vrug produksie van die vyf bestuiwings behandelinge (naamlik: natuurlik, stuifmeel bygevoeg, self, hand self en hand kruis) en tussen genaturaliseerde en nie genaturaliseerde populasies. Ewenwel, stuifmeel beperking is gevind in nie-genaturaliseerde populasies wat egter ook minder besoeke ontvang het dan die genaturaliseerde populasies. Dus, reproduksie kan die verspreiding beperk maar is nie ʼn fundamentele versperring vir indringing van H. salicifolia nie. My konklusies is dat die reproduktiewe sukses, wat andersins ʼn sleutel versperring is vir indringing, in die bestudeerde Proteaceae nie beperk word deur outonomiese saad produksie of mutualismes in die geïntroduseerde gebied nie. In hierdie tesis beklemtoon ek die biogeografiese karakters, lewens geskiedenis kenmerke en ekologiese kenmerke as belangrike bepalers van indringing. Hierdie kenmerke is op hulle beurt weer afhanklik van die stadium van indringing. Karakters van die ontvangende omgewing is ook belangrike dryfvere van indringing. Hierdie studie verbeter hoe ons plant indringing in die algemeen verstaan, maar die patrone en prosesse van indringing wat beklemtoon word in hierdie tesis sal besonder bruikbaar wees vir huidige en toekomstige bestuur van uitheemse Proteaceae in Suid Afrika en op ander plekke, asook vir ander spesies wat aangepas is tot Mediterreense en nutriënt arm ekosisteme. Byvoorbeeld, die kombinasie van kenmerke van indringing en vatbare omgewings sal help om te identifiseer watter uitheemse spesies ʼn hoë risiko inhou om ʼn indringer te word (byvoorbeeld spesies met ʼn groot streek grootte en versperring spesies) en watter kondisies in die teiken area die waarskynlikste indringing fasiliteer of vererger (byvoorbeeld sterk klimaat ooreenstemming en hoë propaguul druk).

The establishment of exotic species in novel environments is a major environmental concern, however, few long-term studies have examined the effects of these species on their host environment and community, especially in forest ecosystems. The arrival and subsequent spread of the greater European pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Curculionidae, Scolytinae), into southern Ontario pine forests provided a natural experiment to assess biotic interactions between an exotic species and its new community over several years. Reproductive success of Tomicus piniperda colonies of various ages was studied between 2001 and 2004. The size and composition of competitor and natural enemy complexes present in these communities were also quantified over time. The impact of the natural enemy complexes on T. piniperda reproduction was assessed, Brood production (no. eggs and galleries/female) by T. piniperda populations rapidly approached those reported from its native range in Europe, with lower densities of parental adults. Thus, reproduction remained consistently above the replacement level for this beetle over all four years of study suggesting that these recently-introduced populations were growing rapidly and at a greater rate than in their country of origin. Tomicus piniperda successfully integrated into a large bark beetle community, and appeared to be capable of displacing native beetles to more marginal bark habitats, however, these competitors were not eliminated during the course of the study. The long-term effect of this marginalization on populations of native beetles is uncertain. Tomicus piniperda rapidly acquired natural enemies in the introduced areas, however, natural enemy-caused mortality did not show a regulating effect on its populations. It seems that intraspecific competition, rather than predation, regulates T. piniperda populations following introduction. The implications of these findings for the establishment and spread of exotic species in forest systems are examined, especially with reference to a prominent theory for success, the Enemy Release Hypothesis. In short, the Enemy Release Hypothesis is not applicable to an exotic species that is not regulated by natural enemies in its native range, and assessments of the Enemy Release Hypothesis should always include a determination of enemy regulation of the exotic in its native range.

The establishment of exotic species in novel environments is a major environmental concern, however, few long-term studies have examined the effects of these species on their host environment and community, especially in forest ecosystems. The arrival and subsequent spread of the greater European pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Curculionidae, Scolytinae), into southern Ontario pine forests provided a natural experiment to assess biotic interactions between an exotic species and its new community over several years. Reproductive success of Tomicus piniperda colonies of various ages was studied between 2001 and 2004. The size and composition of competitor and natural enemy complexes present in these communities were also quantified over time. The impact of the natural enemy complexes on T. piniperda reproduction was assessed, Brood production (no. eggs and galleries/female) by T. piniperda populations rapidly approached those reported from its native range in Europe, with lower densities of parental adults. Thus, reproduction remained consistently above the replacement level for this beetle over all four years of study suggesting that these recently-introduced populations were growing rapidly and at a greater rate than in their country of origin. Tomicus piniperda successfully integrated into a large bark beetle community, and appeared to be capable of displacing native beetles to more marginal bark habitats, however, these competitors were not eliminated during the course of the study. The long-term effect of this marginalization on populations of native beetles is uncertain. Tomicus piniperda rapidly acquired natural enemies in the introduced areas, however, natural enemy-caused mortality did not show a regulating effect on its populations. It seems that intraspecific competition, rather than predation, regulates T. piniperda populations following introduction. The implications of these findings for the establishment and spread of exotic species in forest systems are examined, especially with reference to a prominent theory for success, the Enemy Release Hypothesis. In short, the Enemy Release Hypothesis is not applicable to an exotic species that is not regulated by natural enemies in its native range, and assessments of the Enemy Release Hypothesis should always include a determination of enemy regulation of the exotic in its native range.

Invasive plants are widely cited as a major threat to native plant communities, and the correlation between plant invasions and a subsequent decline in native species is well documented at some scales. However, one outstanding question is the degree to which invasive species are a driver of native plant declines versus a correlate of other drivers, such as habitat disturbance. These two hypotheses to explain the dominance of invasive species in communities have been termed the ‘driver’ and ‘passenger’ models, respectively. In order to understand the impacts of plant invasion on native plants we need more studies that consider the role of correlated environmental predictors, which may play unseen roles in the response and recovery of native plant communities frequently attributed to invasion alone. Using a large database of plant community and environmental data from sites across Southern Ontario, I used path analyses to examine the direct and indirect relationships between disturbance, exotic and native plant richness, and relative abundance. Counter to my initial predictions, I found support for both the partial passenger and partial driver models of invasive dominance, while full passenger models were outright rejected. The causal hypotheses consistent with the data indicated significant relationships between native and exotic species richness and native and exotic relative abundance across models. An exploratory analysis, which examined species-specific models, found that the data was consistent with seven out of twelve causal hypotheses. Models that could not be rejected were split almost evenly across full passenger, partial passenger, and partial driver models. Model support varied according to the species included in the dataset suggesting that the best fit underlying model of invasive dominance likely varies by species. While the partial passenger and partial driver models were recurrently consistent with the data, no single model described the underlying patterns of invasive dominance across all systems.

The success of an invasive alien species is often attributed to the ecological advantage gained from natural enemy release. Numerous factors have been suggested as contributing to the success of Harmonia axyridis as an invasive alien species, including enemy release. This thesis studied the interactions of several parasites with H. axyridis, investigating parasite transmission, growth and virulence as well as host immune responses, thereby shedding light on the potential role of enemy release in the invasion biology of this ladybird. Benefits gained by invasive alien species from enemy release diminish if parasites of native species shift hosts to exploit the novel invader. The fungal ectoparasite Hesperomyces virescens began infecting H. axyridis shortly after it invaded the UK, probably as a result of a host shift from Adalia bipunctata. This study found a rapid increase in H. virescens prevalence over three years in London H. axyridis populations. Laboratory study showed H. virescens transmission and growth to be more efficient on A. bipunctata than the novel host. In addition, reciprocal interspecific transfers of H. virescens strains isolated from A. bipunctata and H. axyridis revealed that the infection characteristics of the fungi from these two hosts differed, suggesting strains may have diverged after the initial shift from A. bipunctata to better exploit the host from which they were derived. Laboulbenialian fungi were previously thought to have negligible impacts on host fitness. A detailed examination of H. virescens infecting H. axyridis found distinct virulence, with infections resulting in a 50% reduction in host lifespan. In addition, chronic H. virescens infection in males caused acceleration in the age-associated decline in body condition while for females, infection triggered fecundity senescence and a faster age-related decline in fertility. While their role in accelerating ageing is debated, the results presented here provide evidence that infectious diseases can drive the ageing process in this insect species. In nature, multiple parasites affecting a single host are common. The effect of co-infection on the virulence caused by two fungal infections was characterised using H. axyridis and A. bipunctata hosts. The ability of two ladybird species to defend against an acute fungal parasite, while infected with the relatively avirulent H. virescens was found to be sex-specific. While for females, the presence of co-infection did not alter the virulence seen in singly infected females, a higher mortality rate existed for co-infected males compared with those infected singly. Previously, H. virescens has been considered to be avirulent, however, this study provides evidence that this chronic fungal parasite may be important when considering the mortality associated with co-infections in the field. The invasive success of H. axyridis has, in part, been attributed to a more vigorous immune ability compared with other competitor species. Previously, field studies have shown that the prevalence of the parasitoid wasp Dinocampus coccinellae in H. axyridis is considerably lower than in the UK primary host of this wasp, Coccinella septempunctata. The extent to which the prevalence asymmetry in the field is driven by differences in host encapsulation response was tested by first comparing the encapsulation ability of C. septempunctata and H. axyridis directed against an artificial implant. Following this, the encapsulation response of D. coccinellae parasitized individuals was assessed and compared between the two host species. While encapsulation ability did not differ between the host species, and D. coccinellae did not affect the immune response of H. axyridis, wasp parasitism did alter the encapsulation ability of C. septempunctata, although it was inconsistent across sexes and populations. Overall, this thesis furthers our understanding of the fungal parasite H. virescens and its association with the notorious invader H. axyridis. The research presented here also demonstrates the use of H. axyridis as a model system in areas other than invasion ecology and furthermore, contributes to understanding the role of infectious disease in the rate of ageing. Finally, sex-specific effects were found across the chapters of this thesis, demonstrating the use of H. axyridis in the study of sex-specific effects of infections.

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Blue Catfish were introduced in the tidal tributaries of the Chesapeake Bay in the 1970s and 1980s to establish new fisheries during a time period when many fisheries were in decline due to pollution, habitat alteration, disease, overfishing, and environmental catastrophes. Having expanded their range to most Bay tributaries, the species has drawn concern from many stakeholders and scientists for its effects on at-risk and economically important native and naturalized species. My study focused on understanding the dynamics of this species based on multiple long-term monitoring data and evaluating potential management strategies to meet stakeholder needs. I sought to understand how is growth variability was partitioned over time and space, how Blue Catfish populations changed from 1994 to 2016, and how predation on native species and fishery-based performance measures may respond to management intervention. As Blue Catfish length-at-age is exceptionally variable in Virginia tributaries of the Chesapeake Bay, I evaluated the variability in growth using candidate non-linear mixed effects models that described variability in growth over time and space. Linear trend tests supported declines in growth over time within river systems, but did not support the presence of synchronous growth responses among river systems. To better understand population dynamics of Blue Catfish in the Chesapeake Bay watershed, I developed a statistical catch-at-length model for the James River to estimate population size, instantaneous fishing mortality, and size structure over time. The statistical catch-at-length model estimated that Blue Catfish abundance increased slowly and peaked in the mid-2000s before undergoing a recent decline. The model estimated a large spike in abundance due to an estimated large recruitment event in 2011, but may be an artifact of missing data in 2012 in both relative abundance indices examined. The newly developed statistical catch-at-length model provides most detailed information on population dynamics of Blue Catfish in the James River and can be expanded and updated as new data become available. Based on results of the statistical catch-at-length model, I examined population responses to unregulated, maximum length limit (60 cm), and harvest slot limit regulations (harvest allowed 25 –60 cm) in a management strategy evaluation framework. The management strategy evaluation supported that the James River Blue Catfish population could be reduced with increased harvest, but trophy-size fish would decline. Consequently, fishery managers tasked with invasive species management must consider this tradeoff of fishery economic benefits and predation on native populations, especially those prey in which population sizes are unknown. / Doctor of Philosophy / Blue Catfish are non-native to the Chesapeake Bay watershed, but were stocked in the 1970s and 1980s to provide fishing opportunities to the region. Unknowingly, Blue Catfish expanded downstream and beyond the boundaries of the rivers to which they were originally stocked and now exist in extremely dense populations in places. This expansion in population size and distribution has generated concern for the health of the Chesapeake Bay and calls for population control. I wanted to learn more about Blue Catfish in Virginia, specifically Blue Catfish growth rates, population dynamics, and how they might respond to control efforts. I examined Blue Catfish growth rates and found growth rates differed over time and across river systems. Blue Catfish tended to grow more slowly over time as their populations matured. As growth rates declined, population size increased with maximum population sizes in the late 2000s in the James River with a subsequent decline in abundance. Many invasive species exhibit this sort of phenomenon, where population sizes increase and reach a maximum before declining. Finally, I looked at Blue Catfish responses to different fishing regulations and harvest levels, finding that increased harvest could help control Blue Catfish population sizes. However, Blue Catfish management objectives are in conflict as regulations that limit predation of native species of interest also reduce the proportion of large fish in populations. Blue Catfish management will require stakeholder-driven approaches to ensure buy-in and reduce user conflicts.

Human activities worldwide have altered nature in ways that create new combinations of species and environmental processes. To understand so-called "novel ecosystems" it is important to consider both the natural and the societal factors that shape them, and how those factors are interconnected or "coupled." We used such an approach to explore options for managing a non-native invasive frog, the coqui, which has become established on the island of Hawaii and threatens to spread to other parts of the state. The nighttime calls of the coqui create a nuisance for property owners when populations become dense enough, as often occurs in Hawaii where the frogs have no natural enemies. Humans have tried various ways to eliminate coqui on the island of Hawaii with little success. Therefore we studied how property owners cope with their presence, both through management practices and psychological coping strategies. We also examined results of those efforts. People whose properties had more frogs were more likely to take action to reduce their numbers, but also attitudes toward the coqui were less negative when people had grown used to having to share their properties with the frogs. For those who cannot cope psychologically, we found it would be possible to manage properties to reduce densities but only when leaf litter and low shrubs were completely removed from near a home. Information campaigns about managing coqui should be different when targeting people that already host frogs and those that do not.