The interface between invasive species science and legal regulation, using Hymenopteran species and their pathogens as a model system

<p>Biological invasions are one of the major causes of biodiversity decline on the planet. The key driver of the global movement of invasive species is international trade. As a response to trade driven invasive species risk, international and domestic regulations have been promulgated with the goal of managing the spread and impact of non-native species. My aims in this thesis were twofold. First, my goal was to review a subset of international and domestic regulations with a view to commenting on their fitness for purpose and suggesting potential improvements. Second, I used the example of non-native and invasive Hymenoptera, as well as their pathogens, to illustrate the risks posed by invasive species and gaps in their management. In order to assess international and domestic regulations, I reviewed the World Trade Organization’s (WTO) Agreement on Sanitary and Phytosanitary Measures, as well as associated disputes. I argue that the WTO’s regulatory system does, for the most part, allow domestic regulators to manage invasive species risk as they see fit. Subsequently, the focus of the thesis narrows to investigate New Zealand’s pre- and post-border regime managing invasive species. I argue that New Zealand’s pre-border approach represents international best practice, but the post-border management of species is fragmented. The power to manage invasive species has been delegated to sub-national and regional bodies, which typically approach invasive species management in different ways. This variation has led to regulatory inconsistencies in pests managed and funding allocated. There appears to be a substantial lack of planning in some spaces, such as the risk of aquatic invasions. I make recommendations to ameliorate these inconsistencies. My second aim involved the study of non-native and invasive Hymenoptera in New Zealand, as well as the pathogens they carry, in order to illustrate the risks posed by invasive species and gaps in their management. I show that the globally widespread invasive Argentine ant (Linepithema humile) may play a role in the pathogen dynamics and mortality of honey bee hives where the species occur sympatrically. Hives in the presence of Argentine ants suffered significantly higher mortality rates relative to hives without ants and always had higher levels of a honey bee pathogen Deformed wing virus. I demonstrate that honey bee pathogens are found in a range of invasive Hymenoptera in New Zealand. I amplify entire genomes of the honey bee virus Kashmir bee virus (KBV) from three species of non-native or invasive Hymenoptera (Argentine ants, common wasps and honey bees). I show that there is KBV strain variability within and between regions, but more between regions. Further, I demonstrate the result that as sampled KBV sequence length increases, so too does sampled diversity. These results highlight how ‘an’ invasive species is typically not alone: they carry a range of diseases that are almost always not considered in international and regional management plans. Patterns of non-native Hymenoptera carrying honey bee diseases were not restricted to New Zealand. I used mitochondrial DNA to find the likely origin of invasive populations of the globally distributed invasive German wasp. I demonstrate that German wasps show reduced genetic diversity in the invaded range compared to the native range. Populations in the introduced range are likely to have arrived from different source populations. In some regions there were likely multiple introductions. Other regions are genetically homogenous and represent potential areas for use of gene drive technologies. All four different honey bee pathogens assayed for were found in German wasp populations worldwide. These results highlight how the introduction of one exotic species likely brings a range of pathogens. This example of pathogens in Hymenoptera is likely to be true for nearly all non-native introductions. Many of the impacts of biological invasions, such as predation and competition, are relatively obvious and are frequently studied. However some, such as the impact of pathogens, are unseen and poorly understood. Legal regulation is often a post-hoc response implemented once a problem has already arisen. At a global level regulatory regimes operate relatively effectively. As the focus becomes more granular, such as the case of pathogens of Hymenoptera, fewer controls exists. This thesis helps to reduce uncertainty in this area as well as makes recommendations as to how these risks may be managed.</p>

The interface between invasive species science and legal regulation, using Hymenopteran species and their pathogens as a model system

<p>Biological invasions are one of the major causes of biodiversity decline on the planet. The key driver of the global movement of invasive species is international trade. As a response to trade driven invasive species risk, international and domestic regulations have been promulgated with the goal of managing the spread and impact of non-native species. My aims in this thesis were twofold. First, my goal was to review a subset of international and domestic regulations with a view to commenting on their fitness for purpose and suggesting potential improvements. Second, I used the example of non-native and invasive Hymenoptera, as well as their pathogens, to illustrate the risks posed by invasive species and gaps in their management. In order to assess international and domestic regulations, I reviewed the World Trade Organization’s (WTO) Agreement on Sanitary and Phytosanitary Measures, as well as associated disputes. I argue that the WTO’s regulatory system does, for the most part, allow domestic regulators to manage invasive species risk as they see fit. Subsequently, the focus of the thesis narrows to investigate New Zealand’s pre- and post-border regime managing invasive species. I argue that New Zealand’s pre-border approach represents international best practice, but the post-border management of species is fragmented. The power to manage invasive species has been delegated to sub-national and regional bodies, which typically approach invasive species management in different ways. This variation has led to regulatory inconsistencies in pests managed and funding allocated. There appears to be a substantial lack of planning in some spaces, such as the risk of aquatic invasions. I make recommendations to ameliorate these inconsistencies. My second aim involved the study of non-native and invasive Hymenoptera in New Zealand, as well as the pathogens they carry, in order to illustrate the risks posed by invasive species and gaps in their management. I show that the globally widespread invasive Argentine ant (Linepithema humile) may play a role in the pathogen dynamics and mortality of honey bee hives where the species occur sympatrically. Hives in the presence of Argentine ants suffered significantly higher mortality rates relative to hives without ants and always had higher levels of a honey bee pathogen Deformed wing virus. I demonstrate that honey bee pathogens are found in a range of invasive Hymenoptera in New Zealand. I amplify entire genomes of the honey bee virus Kashmir bee virus (KBV) from three species of non-native or invasive Hymenoptera (Argentine ants, common wasps and honey bees). I show that there is KBV strain variability within and between regions, but more between regions. Further, I demonstrate the result that as sampled KBV sequence length increases, so too does sampled diversity. These results highlight how ‘an’ invasive species is typically not alone: they carry a range of diseases that are almost always not considered in international and regional management plans. Patterns of non-native Hymenoptera carrying honey bee diseases were not restricted to New Zealand. I used mitochondrial DNA to find the likely origin of invasive populations of the globally distributed invasive German wasp. I demonstrate that German wasps show reduced genetic diversity in the invaded range compared to the native range. Populations in the introduced range are likely to have arrived from different source populations. In some regions there were likely multiple introductions. Other regions are genetically homogenous and represent potential areas for use of gene drive technologies. All four different honey bee pathogens assayed for were found in German wasp populations worldwide. These results highlight how the introduction of one exotic species likely brings a range of pathogens. This example of pathogens in Hymenoptera is likely to be true for nearly all non-native introductions. Many of the impacts of biological invasions, such as predation and competition, are relatively obvious and are frequently studied. However some, such as the impact of pathogens, are unseen and poorly understood. Legal regulation is often a post-hoc response implemented once a problem has already arisen. At a global level regulatory regimes operate relatively effectively. As the focus becomes more granular, such as the case of pathogens of Hymenoptera, fewer controls exists. This thesis helps to reduce uncertainty in this area as well as makes recommendations as to how these risks may be managed.</p>

Inter-species horizontal toxicity: Experimental control of Argentine ants and their citrophilus mealybug mutualists in a viticultural system

<p>The invasive Argentine ant, Linepithema humile, is known to form a trophobiotic association with honeydew excreting homopterans Pseudococcus sp. providing protection from natural enemies in exchange for the honeydew they excrete. The vine mealybug Pseudococcus calceolariae, can transmit Grapevine leafroll- associated virus 3 (GLRaV-3) between vines as it travels and feeds with the ensuing leafroll disease negatively impacting on vine health and wine quality. Therefore, if an effective chemical control method targeting incursions of Argentine ants in vineyards contributes to the dissociation of this invasive ant species with its citrophilus mealybug mutualist, then in theory the spread of GLRaV-3 in vineyards by its mealybug vector can be stemmed. Three insecticidal treatments targeting Argentine ants in the canopy of potted Pinot Noir grapevines inoculated with citrophilus mealybugs were trialled at a field site established in Nelson during the summer of 2016/2017. Bifenthrin (1200ppm) was sprayed on vine trunks and the low- toxicity baits, thiamethoxam (0.0006%) or boric acid (0.5%) carried in polyacrylamide gel with 25% sucrose and 0.15% citric acid solution, were placed at the base of vines. A significant decline in ant activity (p < 0.001) and citrophilus mealybugs was observed for the bifenthrin treatment. A follow-on bioassay was conducted at Mt. Albert Plant and Food Research, in the absence of P. calceolariae’s natural enemies to test the hypothesis that the decline in citrophilus mealybugs in response to vines treated with bifenthrin, could in fact be due to inter-species horizontal toxicity because of Argentine ants transferring the toxicant bifenthrin to citrophilus mealybugs while tending them or contaminating the substrate that they fed on. The significant decrease in average citrophilus mealybug activity rate (p < 0.001) for bifenthrin treatments compared with the controls provides evidence for inter-species horizontal toxicity. Bifenthrin sprayed on grapevine trunks may be suitable to control Argentine ants in the vine canopy and indirectly control P. calceolariae, a known vector of GLRaV-3 between grapevine hosts. The concept of inter-species horizontal toxicity could become a model for targeted pest management by exploiting different insect mutualisms in various horticultural cropping systems.</p>

Inter-species horizontal toxicity: Experimental control of Argentine ants and their citrophilus mealybug mutualists in a viticultural system

<p>The invasive Argentine ant, Linepithema humile, is known to form a trophobiotic association with honeydew excreting homopterans Pseudococcus sp. providing protection from natural enemies in exchange for the honeydew they excrete. The vine mealybug Pseudococcus calceolariae, can transmit Grapevine leafroll- associated virus 3 (GLRaV-3) between vines as it travels and feeds with the ensuing leafroll disease negatively impacting on vine health and wine quality. Therefore, if an effective chemical control method targeting incursions of Argentine ants in vineyards contributes to the dissociation of this invasive ant species with its citrophilus mealybug mutualist, then in theory the spread of GLRaV-3 in vineyards by its mealybug vector can be stemmed. Three insecticidal treatments targeting Argentine ants in the canopy of potted Pinot Noir grapevines inoculated with citrophilus mealybugs were trialled at a field site established in Nelson during the summer of 2016/2017. Bifenthrin (1200ppm) was sprayed on vine trunks and the low- toxicity baits, thiamethoxam (0.0006%) or boric acid (0.5%) carried in polyacrylamide gel with 25% sucrose and 0.15% citric acid solution, were placed at the base of vines. A significant decline in ant activity (p < 0.001) and citrophilus mealybugs was observed for the bifenthrin treatment. A follow-on bioassay was conducted at Mt. Albert Plant and Food Research, in the absence of P. calceolariae’s natural enemies to test the hypothesis that the decline in citrophilus mealybugs in response to vines treated with bifenthrin, could in fact be due to inter-species horizontal toxicity because of Argentine ants transferring the toxicant bifenthrin to citrophilus mealybugs while tending them or contaminating the substrate that they fed on. The significant decrease in average citrophilus mealybug activity rate (p < 0.001) for bifenthrin treatments compared with the controls provides evidence for inter-species horizontal toxicity. Bifenthrin sprayed on grapevine trunks may be suitable to control Argentine ants in the vine canopy and indirectly control P. calceolariae, a known vector of GLRaV-3 between grapevine hosts. The concept of inter-species horizontal toxicity could become a model for targeted pest management by exploiting different insect mutualisms in various horticultural cropping systems.</p>

The influence of semiochemicals on the co-occurrence patterns of a global invader and native species

<p>The success of invasive species in their introduced range is often influenced by interactions with resident species communities. Chemical communication is one the factors which contributes to a variety of aspects of a species life cycle, ranging from mating, to food localization and interactions with members of the same and other species. In my thesis, I investigate the effects of venoms and semiochemicals on interactions between the invasive Argentine ant (Linepethima humile) with other resident ant species and demonstrate how pheromones can potentially be utilized as an area wide control mechanism of this species, by disrupting their foraging success. I studied the effects of venom composition, their toxicity and utilization on the outcome of aggressive interactions between the Argentine ant and the four Monomorium species in New Zealand occurring. The toxicity of the venom of the two species co-occurring with Argentine ants was significantly higher than the toxicity of the species which do not. However, no correlation between venom toxicity and Monomorium survival was found. For M. antipodum a significant relationship between venom utilization and its mortality was found, indicating that the way venom is used might be an important aspect of these interactions. Physical Aggression between Monomorium and Argentine ants also had strong effects on Monomorium worker mortality, which provided evidence that a variety of factors and strategies contribute to the ability of interacting organisms to withstand the pressure of a dominant invader at high abundance. I conducted bioassays with food sources and synthetic trail pheromones, providing a proof of concept on disrupting the foraging ability of Argentine ants. Other resident species benefited from the reduced success of Argentine ants, but to a varying degree. Behavioural variations between the resident species provided an explanation for observed differences in foraging success and how much each of these individual competitors was able to increase their foraging. The mechanism for the observed increase in resource acquisition of resident species appeared to be a decrease in aggressive behaviour displayed by Argentine ants. I expanded the usage of the synthetic pheromone to a commercial vineyard, were Argentine ants can have negative effects on crop development by dispersing and tending to homopteran pest species. Argentine ants’ access to the crop canopy could be significantly reduced by placing pheromone dispensers at the base of the vine plant, while dispensers in the plant canopy had little effect on Argentine ant numbers. Doubling the amount of pheromone did not result in an additional reduction of ant activity. Lastly incorporating the knowledge gained in the previous chapter, I extended the application of the pheromone to a large field trial over a three month period. Argentine ant activity and foraging success was significantly supressed compared to untreated control plots, providing evidence that this form of large scale application might be a possible way to control large invasive ant populations by disrupting their trail following and foraging behaviour for a prolonged period of time. While initial calculations have suggested that the treatment is currently not feasible (13.3 US$/mg/ha), I found a significant reduction in body fat in workers collected from treated plots compared with untreated plots, suggesting adverse effects on nest fitness. My findings provide new insights into chemical communication between invasive and resident species, support existing dominance hierarchy models in ant populations, and help to establish a target specific potential management technique of wide-spread invasive ant species.</p>

Bacterial diversity and viral discovery in the invasive Argentine ant (Linepithema humile)

<p>Invasive species can lead to major economic and ecological issues. For this reason, biological controls are being developed in order to help with invasive species population management. Pathogenic bacteria and viruses offer good biological control opportunities as both micro-organisms have played a role in natural population declines. However, beneficial bacteria and viruses associated with the targeted invasive species may interfere with biological controls, by protecting their hosts from infections. Previous knowledge on both pathogenic and beneficial bacteria and viruses present in invasive species may then support the development of an active and efficient biological control. The Argentine ant, Linepithema humile, is a South American invasive ant species that has successfully spread over five continents. The ants were introduced to New Zealand after a complex invasion path, from Argentina their home range to Europe, then to Australia and finally to New Zealand. In their new environments, invasive Argentine ants affect species diversity and can cause agricultural losses. In the absence of any biological controls, the Argentine ant population is controlled by chemical sprays and poison baits. Management of these invasive ants in New Zealand is estimated to cost NZ$ 60 million a year. The Argentine ant population in New Zealand was reported to have unexpectedly declined. It was hypothesised that pathogens were the cause of this population collapse. In this study, bacteria and viruses present in the invasive ants were investigated using 454 sequencing and Illumina sequencing for future developments of possible biological controls for the Argentine ants, and a better understanding of the observed population decline in New Zealand. Bacterial diversity present in Argentine ants either declined or diminished along the invasion pathway. At the same time, the invasive ants maintained a core of nine bacteria genera, including Lactobacillus and Gluconobacter, two bacterial genera with members known for their beneficial associations with honey bees. The presence of these core bacteria may have participated in the success of Argentine ants in their new environments. In the laboratory, the use of ampicillin and gentamicin antibiotics on the ants increased bacterial diversity present in the ants. Furthermore, ampicillin, kanamycin and spectinomycin antibiotic treatments increased ant survival but did not affect the ant fitness or intra-species aggressiveness. Only spectinomycin treated ants presented a higher level of inter-species aggressiveness. Bacterial diversity may play an important role in the ant health and at length population dynamics. Finally, Argentine ants are the hosts of two viruses: the Deformed wing virus (DWV) involved in colony collapse disorder in honey bees, and Linepithema humile virus 1 (LHUV-1), a new virus related to DWV. Both viruses actively replicate within the ants, indicating a possible reservoir role of the ants. However, the effects of the viruses on the ants are not yet known. Further viral infection in the laboratory under different stress conditions and / or antibiotic treatment will give an insight in the role played by these viruses in the observed population collapse of Argentine ants in New Zealand. LHUV-1 may offer a possibility in the development of the first biological control for Argentine ants, depending on its specificity and its effects. This dissertation provides a first insight in the core bacteria as well as potential harmful viruses present in Argentine ants. These bacteria and viruses may play a role in the ant population dynamics. Invasive species may co-introduce harmful pathogens with them, and participate to the spread of local ones. The pathogens may affect both invasive ants and native species population dynamics.</p>

The influence of semiochemicals on the co-occurrence patterns of a global invader and native species

<p>The success of invasive species in their introduced range is often influenced by interactions with resident species communities. Chemical communication is one the factors which contributes to a variety of aspects of a species life cycle, ranging from mating, to food localization and interactions with members of the same and other species. In my thesis, I investigate the effects of venoms and semiochemicals on interactions between the invasive Argentine ant (Linepethima humile) with other resident ant species and demonstrate how pheromones can potentially be utilized as an area wide control mechanism of this species, by disrupting their foraging success. I studied the effects of venom composition, their toxicity and utilization on the outcome of aggressive interactions between the Argentine ant and the four Monomorium species in New Zealand occurring. The toxicity of the venom of the two species co-occurring with Argentine ants was significantly higher than the toxicity of the species which do not. However, no correlation between venom toxicity and Monomorium survival was found. For M. antipodum a significant relationship between venom utilization and its mortality was found, indicating that the way venom is used might be an important aspect of these interactions. Physical Aggression between Monomorium and Argentine ants also had strong effects on Monomorium worker mortality, which provided evidence that a variety of factors and strategies contribute to the ability of interacting organisms to withstand the pressure of a dominant invader at high abundance. I conducted bioassays with food sources and synthetic trail pheromones, providing a proof of concept on disrupting the foraging ability of Argentine ants. Other resident species benefited from the reduced success of Argentine ants, but to a varying degree. Behavioural variations between the resident species provided an explanation for observed differences in foraging success and how much each of these individual competitors was able to increase their foraging. The mechanism for the observed increase in resource acquisition of resident species appeared to be a decrease in aggressive behaviour displayed by Argentine ants. I expanded the usage of the synthetic pheromone to a commercial vineyard, were Argentine ants can have negative effects on crop development by dispersing and tending to homopteran pest species. Argentine ants’ access to the crop canopy could be significantly reduced by placing pheromone dispensers at the base of the vine plant, while dispensers in the plant canopy had little effect on Argentine ant numbers. Doubling the amount of pheromone did not result in an additional reduction of ant activity. Lastly incorporating the knowledge gained in the previous chapter, I extended the application of the pheromone to a large field trial over a three month period. Argentine ant activity and foraging success was significantly supressed compared to untreated control plots, providing evidence that this form of large scale application might be a possible way to control large invasive ant populations by disrupting their trail following and foraging behaviour for a prolonged period of time. While initial calculations have suggested that the treatment is currently not feasible (13.3 US$/mg/ha), I found a significant reduction in body fat in workers collected from treated plots compared with untreated plots, suggesting adverse effects on nest fitness. My findings provide new insights into chemical communication between invasive and resident species, support existing dominance hierarchy models in ant populations, and help to establish a target specific potential management technique of wide-spread invasive ant species.</p>

Bacterial diversity and viral discovery in the invasive Argentine ant (Linepithema humile)

<p>Invasive species can lead to major economic and ecological issues. For this reason, biological controls are being developed in order to help with invasive species population management. Pathogenic bacteria and viruses offer good biological control opportunities as both micro-organisms have played a role in natural population declines. However, beneficial bacteria and viruses associated with the targeted invasive species may interfere with biological controls, by protecting their hosts from infections. Previous knowledge on both pathogenic and beneficial bacteria and viruses present in invasive species may then support the development of an active and efficient biological control. The Argentine ant, Linepithema humile, is a South American invasive ant species that has successfully spread over five continents. The ants were introduced to New Zealand after a complex invasion path, from Argentina their home range to Europe, then to Australia and finally to New Zealand. In their new environments, invasive Argentine ants affect species diversity and can cause agricultural losses. In the absence of any biological controls, the Argentine ant population is controlled by chemical sprays and poison baits. Management of these invasive ants in New Zealand is estimated to cost NZ$ 60 million a year. The Argentine ant population in New Zealand was reported to have unexpectedly declined. It was hypothesised that pathogens were the cause of this population collapse. In this study, bacteria and viruses present in the invasive ants were investigated using 454 sequencing and Illumina sequencing for future developments of possible biological controls for the Argentine ants, and a better understanding of the observed population decline in New Zealand. Bacterial diversity present in Argentine ants either declined or diminished along the invasion pathway. At the same time, the invasive ants maintained a core of nine bacteria genera, including Lactobacillus and Gluconobacter, two bacterial genera with members known for their beneficial associations with honey bees. The presence of these core bacteria may have participated in the success of Argentine ants in their new environments. In the laboratory, the use of ampicillin and gentamicin antibiotics on the ants increased bacterial diversity present in the ants. Furthermore, ampicillin, kanamycin and spectinomycin antibiotic treatments increased ant survival but did not affect the ant fitness or intra-species aggressiveness. Only spectinomycin treated ants presented a higher level of inter-species aggressiveness. Bacterial diversity may play an important role in the ant health and at length population dynamics. Finally, Argentine ants are the hosts of two viruses: the Deformed wing virus (DWV) involved in colony collapse disorder in honey bees, and Linepithema humile virus 1 (LHUV-1), a new virus related to DWV. Both viruses actively replicate within the ants, indicating a possible reservoir role of the ants. However, the effects of the viruses on the ants are not yet known. Further viral infection in the laboratory under different stress conditions and / or antibiotic treatment will give an insight in the role played by these viruses in the observed population collapse of Argentine ants in New Zealand. LHUV-1 may offer a possibility in the development of the first biological control for Argentine ants, depending on its specificity and its effects. This dissertation provides a first insight in the core bacteria as well as potential harmful viruses present in Argentine ants. These bacteria and viruses may play a role in the ant population dynamics. Invasive species may co-introduce harmful pathogens with them, and participate to the spread of local ones. The pathogens may affect both invasive ants and native species population dynamics.</p>

Invasion Impact and Biotic Resistance by Invertebrate Communities

<p>Invasive species have been recognized as one of the greatest threats to global biodiversity and can have dire economic consequences. Yet rates of invasion are increasing due to the fast and growing network of transportation across the globe. The establishment, spread and impact of invasive species are affected by environmental conditions as well as resident species. Species respond differently to the same abiotic factors and different native species can respond either positively or negatively to invasion. The interaction between invasive and resident species, as well as the effect of temperature on invasive species, has gained much attention. The synergistic effect of suboptimal temperature and biotic resistance could have a much stronger limiting or controlling effect on invasive species than either factor alone. Linepithema humile (Argentine ants) are invasive species originally from a Mediterranean climate, but successfully spreading into extra range habitats. The establishment and spread of these ants in temperate New Zealand represents an ideal model system for studying invasion biology in terms of temperature limits and biotic resistance effects. I investigated the changing distribution of the invasive species the Argentine ants over multiple years at five sites in New Zealand. To test whether their rate of spread corresponds with microclimate I investigated their fine-scare distribution patterns and evaluated the number of generations they may develop seasonally and annually in different microhabitat types. I also evaluated their impact on other arthropod species. I conducted a laboratory experiment to evaluate the effect of temperature on their aggression towards other species, walking speed, and foraging abundance. Similarly, I tested the effect of biotic resistance from other ant species (Monomorium antarcticum and Prolasius advenus) with varying colony sizes. I investigated whether there was any interactive effect of temperature and biotic resistance on the Argentine ants. The distribution of Argentine ants had declined across many invasion fronts over the past 7-8 years. They were more likely to be found in concrete, short grass and sandy habitats, which provide warm microsites. Degree-day calculations predicted that they could develop between 2.5 to 3 generations in each of the above microhabitats per year in urban and rural sites while they were predicted to be unable to develop one generation under tree habitats. In tall grass microhabitats they were predicted to develop between 1-1.5 generations per year. The Argentine ants were hypothesised to adversely affect many other arthropod species. Richness and abundance of resident beetle species were negatively correlated with the invasion of the Argentine ants. Areas invaded by the Argentine ants were also associated with a greater number of exotic beetle species, which may imply secondary invasion. Laboratory experiments showed that lowering temperatures below 17°C negatively affected the Argentine ants‟ walking speed, foraging abundance, aggression and their resource control. A high colony size of M. antarcticum (the competing ant species) affected the foraging success of Argentine ants, and the effect was stronger when coupled with unsuitable temperature (17°C and below). Therefore, Argentine ants are weak competitors at low temperature levels. The results of my thesis underline the importance of biotic and abiotic resistances, their interactive effect as well as the effect of the Argentine ants on other species. Based on climatic considerations and the habitat preferences of resident species it may be possible to predict future spread of the Argentine ants. More importantly, knowledge of microhabitat preferences and biotic resistance may help future control measures against Argentine ants based on management of vegetation structure and microhabitat availability.</p>

Invasion Impact and Biotic Resistance by Invertebrate Communities

<p>Invasive species have been recognized as one of the greatest threats to global biodiversity and can have dire economic consequences. Yet rates of invasion are increasing due to the fast and growing network of transportation across the globe. The establishment, spread and impact of invasive species are affected by environmental conditions as well as resident species. Species respond differently to the same abiotic factors and different native species can respond either positively or negatively to invasion. The interaction between invasive and resident species, as well as the effect of temperature on invasive species, has gained much attention. The synergistic effect of suboptimal temperature and biotic resistance could have a much stronger limiting or controlling effect on invasive species than either factor alone. Linepithema humile (Argentine ants) are invasive species originally from a Mediterranean climate, but successfully spreading into extra range habitats. The establishment and spread of these ants in temperate New Zealand represents an ideal model system for studying invasion biology in terms of temperature limits and biotic resistance effects. I investigated the changing distribution of the invasive species the Argentine ants over multiple years at five sites in New Zealand. To test whether their rate of spread corresponds with microclimate I investigated their fine-scare distribution patterns and evaluated the number of generations they may develop seasonally and annually in different microhabitat types. I also evaluated their impact on other arthropod species. I conducted a laboratory experiment to evaluate the effect of temperature on their aggression towards other species, walking speed, and foraging abundance. Similarly, I tested the effect of biotic resistance from other ant species (Monomorium antarcticum and Prolasius advenus) with varying colony sizes. I investigated whether there was any interactive effect of temperature and biotic resistance on the Argentine ants. The distribution of Argentine ants had declined across many invasion fronts over the past 7-8 years. They were more likely to be found in concrete, short grass and sandy habitats, which provide warm microsites. Degree-day calculations predicted that they could develop between 2.5 to 3 generations in each of the above microhabitats per year in urban and rural sites while they were predicted to be unable to develop one generation under tree habitats. In tall grass microhabitats they were predicted to develop between 1-1.5 generations per year. The Argentine ants were hypothesised to adversely affect many other arthropod species. Richness and abundance of resident beetle species were negatively correlated with the invasion of the Argentine ants. Areas invaded by the Argentine ants were also associated with a greater number of exotic beetle species, which may imply secondary invasion. Laboratory experiments showed that lowering temperatures below 17°C negatively affected the Argentine ants‟ walking speed, foraging abundance, aggression and their resource control. A high colony size of M. antarcticum (the competing ant species) affected the foraging success of Argentine ants, and the effect was stronger when coupled with unsuitable temperature (17°C and below). Therefore, Argentine ants are weak competitors at low temperature levels. The results of my thesis underline the importance of biotic and abiotic resistances, their interactive effect as well as the effect of the Argentine ants on other species. Based on climatic considerations and the habitat preferences of resident species it may be possible to predict future spread of the Argentine ants. More importantly, knowledge of microhabitat preferences and biotic resistance may help future control measures against Argentine ants based on management of vegetation structure and microhabitat availability.</p>