The enemy release hypothesis and beyond: Lagarosiphon major invasion dynamics and management options for New Zealand using native natural enemies from South Africa
- Authors: Baso, Nompumelelo Catherine
- Date: 2024-04-05
- Subjects: Uncatalogued
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/435627 , vital:73174 , DOI 10.21504/10962/435627
- Description: Numerous scientific investigations have demonstrated the destructive impact that exotic species can have on ecosystem services beyond a specific threshold. There are many explanations for why introduced plants are likely to be more successful outside their native range. One such explanation is offered by the Enemy Release Hypothesis (ERH), which states that plants automatically become superior competitors outside of their natural range due to release from top-down stressors (herbivory, parasites, and diseases) that is evident in the absence of their natural enemies. The underlying assumption of the ERH is that natural enemies are important regulators of plant species populations, and that the pressures from these natural enemies are felt more readily by native species compared to alien plants. Consequently, in the absence of such pressures, the ERH assumes that exotic plants can allocate more resources towards growth and reproduction, while effectively maintaining accumulated biomass. Classical biological control has previously been cited as evidence for the enemy release hypothesis. Therefore, the overarching aim and theme of this thesis was to investigate the role of ERH on the invasiveness of Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae) in New Zealand. Firstly, a literature search and a meta-analysis was used to synthesize existing studies in order to test for general applicability of this hypothesis to aquatic plant invasions. Furthermore, an empirical investigation was conducted in order to directly quantify enemy release in L. major populations invaded areas of New Zealand. To achieve this, various plant parameters of this plant, overall macrophyte and invertebrate diversity were measured and compared between sites in the native range in South Africa and the invaded areas in New Zealand. Although the meta-analysis showed variable evidence for this hypothesis depending on various modulating factors such as study type, plant growth form and measured parameters, for L. major, there was strong evidence of enemy release. The biogeographical comparisons showed that L. major exhibited increased fitness in most of the invaded sites, marked by elevated biomass accumulation, significantly higher shoot production, and the displacement of native plant species. The observed fitness advantages were directly correlated to a decrease in herbivory diversity and pressure upon the plant's introduction to New Zealand. Unlike the native populations, which contend with the presence of at least four co-occurring herbivores, including specialist herbivores, the invaded range had a substantially lower herbivore diversity, with only Hygraula nitens Butler (Lepidoptera: Crambidae) syn. Nymphula nitens, significantly damaging L. major. These findings emphasize the importance of understanding invasion ecology and theories such as ERH in order to advance aquatic plant management and also present valuable insights for developing effective strategies to mitigate the impact of invasive alien species on aquatic ecosystems. Specifically, results from the empirical investigation provide evidence in support of the ERH and highlight the suitability of implementing biological control strategies to manage the L. major invasion in New Zealand. Previous studies have shown the suitability of two specialist herbivores, Hydrellia lagarosiphon Deeming (Diptera: Ephydridae), and Polypedilum tuburcinatum Andersen (Diptera: Chironomidae), as potential biological control agents. This control strategy presents a sustainable and ecologically responsible approach, promoting coexistence between exotic plants and native species rather than displacement through competitive exclusion. With the apparent dominance of L. major at various New Zealand localities, the subsequent objective of this thesis was to investigate the competitive interactions between L. major and another invasive Hydrocharitaceae, Egeria densa Planchon, as driven by herbivory. Combinations of two host specific Ephydrid flies, H. lagarosiphon and H. egeriae, were used at eight different factorial combination of planting densities. The analysis of plant parameters and the application of inverse linear models revealed that L. major often exhibits relatively higher fitness, especially in low monoculture treatments when the two insects were isolated. However, multiple inverse linear models revealed that actual competitive outcomes are dependent on factors such as initial plant density and herbivory regime, with competitive interactions generally being mild. Nevertheless, the presence of H. lagarosiphon resulted in facilitation of E. densa growth. Thus, even at lower densities, these insects still had an impact on the observed interactions, further emphasizing suitability as damaging biological control agents. Lastly, focusing on the abiotic component of L. major invasion, Species Distribution Models (SDMs) were employed to map potential suitable habitat for this species, as well as predict the consequences of climate change on this. Correlative and mechanistic modelling was also used to simulate suitable habitat for potential biological control agents, thus addressing the potential for mismatches between host plant distribution and insect suitable range. The Maximum Entropy Species Distribution Modelling (MaxEnt) algorithm revealed that more than 90% of all freshwater ecosystems in New Zealand are susceptible to L. major invasion, with suitability projected to expand further under future climate scenarios. Moreover, correlative modelling using this method suggests limited suitable habitat for both herbivores. However, degree-day modelling, which also takes into account the physiological requirements, showed that H. lagarosiphon has the potential to produce viable populations in several parts of New Zealand. Overall, this thesis explored the intricate web of biotic and abiotic factors influencing the success of L. major outside its native range. The results emphasize the potential impacts of climate change on the invasion potential and management strategies for L. major. The findings also advocate for the implementation of sustainable and ecologically sound management solutions, such as biological control, to manage this species. , Thesis (PhD) -- Faculty of Science, Botany, 2024
- Full Text:
- Date Issued: 2024-04-05
- Authors: Baso, Nompumelelo Catherine
- Date: 2024-04-05
- Subjects: Uncatalogued
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/435627 , vital:73174 , DOI 10.21504/10962/435627
- Description: Numerous scientific investigations have demonstrated the destructive impact that exotic species can have on ecosystem services beyond a specific threshold. There are many explanations for why introduced plants are likely to be more successful outside their native range. One such explanation is offered by the Enemy Release Hypothesis (ERH), which states that plants automatically become superior competitors outside of their natural range due to release from top-down stressors (herbivory, parasites, and diseases) that is evident in the absence of their natural enemies. The underlying assumption of the ERH is that natural enemies are important regulators of plant species populations, and that the pressures from these natural enemies are felt more readily by native species compared to alien plants. Consequently, in the absence of such pressures, the ERH assumes that exotic plants can allocate more resources towards growth and reproduction, while effectively maintaining accumulated biomass. Classical biological control has previously been cited as evidence for the enemy release hypothesis. Therefore, the overarching aim and theme of this thesis was to investigate the role of ERH on the invasiveness of Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae) in New Zealand. Firstly, a literature search and a meta-analysis was used to synthesize existing studies in order to test for general applicability of this hypothesis to aquatic plant invasions. Furthermore, an empirical investigation was conducted in order to directly quantify enemy release in L. major populations invaded areas of New Zealand. To achieve this, various plant parameters of this plant, overall macrophyte and invertebrate diversity were measured and compared between sites in the native range in South Africa and the invaded areas in New Zealand. Although the meta-analysis showed variable evidence for this hypothesis depending on various modulating factors such as study type, plant growth form and measured parameters, for L. major, there was strong evidence of enemy release. The biogeographical comparisons showed that L. major exhibited increased fitness in most of the invaded sites, marked by elevated biomass accumulation, significantly higher shoot production, and the displacement of native plant species. The observed fitness advantages were directly correlated to a decrease in herbivory diversity and pressure upon the plant's introduction to New Zealand. Unlike the native populations, which contend with the presence of at least four co-occurring herbivores, including specialist herbivores, the invaded range had a substantially lower herbivore diversity, with only Hygraula nitens Butler (Lepidoptera: Crambidae) syn. Nymphula nitens, significantly damaging L. major. These findings emphasize the importance of understanding invasion ecology and theories such as ERH in order to advance aquatic plant management and also present valuable insights for developing effective strategies to mitigate the impact of invasive alien species on aquatic ecosystems. Specifically, results from the empirical investigation provide evidence in support of the ERH and highlight the suitability of implementing biological control strategies to manage the L. major invasion in New Zealand. Previous studies have shown the suitability of two specialist herbivores, Hydrellia lagarosiphon Deeming (Diptera: Ephydridae), and Polypedilum tuburcinatum Andersen (Diptera: Chironomidae), as potential biological control agents. This control strategy presents a sustainable and ecologically responsible approach, promoting coexistence between exotic plants and native species rather than displacement through competitive exclusion. With the apparent dominance of L. major at various New Zealand localities, the subsequent objective of this thesis was to investigate the competitive interactions between L. major and another invasive Hydrocharitaceae, Egeria densa Planchon, as driven by herbivory. Combinations of two host specific Ephydrid flies, H. lagarosiphon and H. egeriae, were used at eight different factorial combination of planting densities. The analysis of plant parameters and the application of inverse linear models revealed that L. major often exhibits relatively higher fitness, especially in low monoculture treatments when the two insects were isolated. However, multiple inverse linear models revealed that actual competitive outcomes are dependent on factors such as initial plant density and herbivory regime, with competitive interactions generally being mild. Nevertheless, the presence of H. lagarosiphon resulted in facilitation of E. densa growth. Thus, even at lower densities, these insects still had an impact on the observed interactions, further emphasizing suitability as damaging biological control agents. Lastly, focusing on the abiotic component of L. major invasion, Species Distribution Models (SDMs) were employed to map potential suitable habitat for this species, as well as predict the consequences of climate change on this. Correlative and mechanistic modelling was also used to simulate suitable habitat for potential biological control agents, thus addressing the potential for mismatches between host plant distribution and insect suitable range. The Maximum Entropy Species Distribution Modelling (MaxEnt) algorithm revealed that more than 90% of all freshwater ecosystems in New Zealand are susceptible to L. major invasion, with suitability projected to expand further under future climate scenarios. Moreover, correlative modelling using this method suggests limited suitable habitat for both herbivores. However, degree-day modelling, which also takes into account the physiological requirements, showed that H. lagarosiphon has the potential to produce viable populations in several parts of New Zealand. Overall, this thesis explored the intricate web of biotic and abiotic factors influencing the success of L. major outside its native range. The results emphasize the potential impacts of climate change on the invasion potential and management strategies for L. major. The findings also advocate for the implementation of sustainable and ecologically sound management solutions, such as biological control, to manage this species. , Thesis (PhD) -- Faculty of Science, Botany, 2024
- Full Text:
- Date Issued: 2024-04-05
A native weevil and an exotic planthopper: investigating potential biological control agents for nymphaea mexicana zuccarini (nymphaeaceae) and its hybrids in South Africa
- Authors: Reid, Megan Kim
- Date: 2023-03-31
- Subjects: Uncatalogued
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/422554 , vital:71957 , DOI 10.21504/10962/422554
- Description: Nymphaea mexicana Zuccarini (Nymphaeaceae) is an invasive plant originating from southern USA and Mexico that has become problematic in South Africa, invading several water bodies around the country. Manual removal of this plant is very labour intensive and is not cost efficient or effective for long term control, while the use of herbicides is damaging to the environment and expensive. Consequently, this plant is a desirable candidate for biological control, which takes advantage of enemy release of the target weed and aims to re-establish population suppression induced by host specific natural enemies. Initiating biological control requires that several steps are followed to maximise the success of the programme, and the first few of these, including overseas surveys in the native range of the plant, have already been completed. This thesis aimed to continue biological control research for this species to take further steps at effectively managing the plant. Firstly, pre-release surveys in the invaded range are necessary to: determine what factors (including enemy release) contribute to the invasiveness of the target weed; establish a baseline of information to allow for comparison after biological control agents have been released; and identify any insect herbivores that may already be present in the country. The pre-release surveys conducted in this study revealed useful information about N. mexicana invasions in South Africa and provided evidence that enemy release is applicable to this case. However, these studies determined that a native weevil, Bagous longulus Gyllenhal (Coleoptera: Curculionidae), has expanded its host range to include the exotic N. mexicana at three sites, and may thus have potential for management of the species through augmentative releases. The invasion of N. mexicana in South Africa is further complicated by the presence of several Nymphaea hybrids originating from a complex history of horticultural trade. Although previous research has shown that several hybrid groups are present in South Africa, their parentage is not known. As biological control requires the use of host specific insects adapted to overcome the unique chemical and morphological defences utilised by plant species, hybrids are notoriously difficult to manage because they possess intermediate characters inherited from parent species to which natural enemies may not have adapted. Although biological control of hybrids is challenging, other case studies have demonstrated that it is possible to find suitable agents, but the chances of success are increased if putative parents of the hybrids are known so that they can be surveyed for natural enemies. Further molecular studies including possible parents of the Nymphaea hybrids in South Africa were thus carried out in this thesis to focus future surveying efforts. Two main hybrid groups were identified with genetic similarity to two tested putative Nymphaea parents, and this will allow further investigations of these species to improve the chances of successfully managing these hybrid groups. Some of the tested hybrids showed genetic contributions from multiple groups, some of which were unidentified, so it is necessary to prioritise the most problematic hybrids for biological control. With more insight into the genetic makeup of the Nymphaea hybrids in South Africa, investigations into the host specificity of potential biological control agents can be conducted. The ideal biological control agent should have a broad enough host range to impact and survive on both N. mexicana and its hybrids, but without a host range so broad that it would pose risk to native South African species. Host specificity trials are thus necessary to determine the suitability of potential agents. The identification of B. longulus feeding on N. mexicana during pre-release surveys motivated further investigations to determine the natural distribution, field host range, and host specificity of B. longulus in experimentally controlled conditions. Further surveys were therefore conducted at native Nymphaea sites around South Africa in addition to host specificity trials using the native Nymphaea nouchali Burm. f. (Nymphaeaceae), two populations of N. mexicana, and a cultivated hybrid. Results from the surveys and host specificity tests suggest that B. longulus is widely distributed across South Africa, is specific to Nymphaea with no observed preference between N. mexicana and the native N. nouchali, and does not perform well on Nymphaea hybrids. Hence, B. longulus is promising for use in new association biological control through augmentative releases but is not suitable for management of hybrids. In addition to the potential use of the South African B. longulus, it is necessary to conduct host specificity trials for natural enemies from the native range of N. mexicana that were prioritised in previous studies. Megamelus toddi Beamer (Hemiptera: Delphacidae) is one such species that was imported into quarantined laboratory conditions from Florida, USA. Host specificity trials were conducted using the same test plants as described for the studies on B. longulus, in addition to multigeneration trials to determine how long M. toddi could survive on non-target host plants. As with the B. longulus studies, no statistically significant differences in preference were observed between N. mexicana and N. nouchali, but M. toddi could not complete development on the test hybrid, indicating that this species is also unsuitable for the management of Nymphaea hybrids. Despite suboptimal plant health, M. toddi completed development for three generations on the native N. nouchali. This lack of host specificity deems M. toddi unsafe for release in South Africa but highlights the importance of following predefined steps to develop a biological control programme. The concluding chapter of this thesis discusses the aforementioned findings in a broader context by considering the driving forces of plant invasions in general and specifically for N. mexicana in South Africa. Case studies are also consulted to provide insight into how to proceed with managing Nymphaea hybrids in South Africa, while the factors governing host specificity and host range expansion are also discussed and considered in the context of B. longulus and M. toddi. Finally, after a consideration of the limitations of these studies, recommendations are made to continue the development of biological control for N. mexicana in South Africa. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2023
- Full Text:
- Date Issued: 2023-03-31
- Authors: Reid, Megan Kim
- Date: 2023-03-31
- Subjects: Uncatalogued
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/422554 , vital:71957 , DOI 10.21504/10962/422554
- Description: Nymphaea mexicana Zuccarini (Nymphaeaceae) is an invasive plant originating from southern USA and Mexico that has become problematic in South Africa, invading several water bodies around the country. Manual removal of this plant is very labour intensive and is not cost efficient or effective for long term control, while the use of herbicides is damaging to the environment and expensive. Consequently, this plant is a desirable candidate for biological control, which takes advantage of enemy release of the target weed and aims to re-establish population suppression induced by host specific natural enemies. Initiating biological control requires that several steps are followed to maximise the success of the programme, and the first few of these, including overseas surveys in the native range of the plant, have already been completed. This thesis aimed to continue biological control research for this species to take further steps at effectively managing the plant. Firstly, pre-release surveys in the invaded range are necessary to: determine what factors (including enemy release) contribute to the invasiveness of the target weed; establish a baseline of information to allow for comparison after biological control agents have been released; and identify any insect herbivores that may already be present in the country. The pre-release surveys conducted in this study revealed useful information about N. mexicana invasions in South Africa and provided evidence that enemy release is applicable to this case. However, these studies determined that a native weevil, Bagous longulus Gyllenhal (Coleoptera: Curculionidae), has expanded its host range to include the exotic N. mexicana at three sites, and may thus have potential for management of the species through augmentative releases. The invasion of N. mexicana in South Africa is further complicated by the presence of several Nymphaea hybrids originating from a complex history of horticultural trade. Although previous research has shown that several hybrid groups are present in South Africa, their parentage is not known. As biological control requires the use of host specific insects adapted to overcome the unique chemical and morphological defences utilised by plant species, hybrids are notoriously difficult to manage because they possess intermediate characters inherited from parent species to which natural enemies may not have adapted. Although biological control of hybrids is challenging, other case studies have demonstrated that it is possible to find suitable agents, but the chances of success are increased if putative parents of the hybrids are known so that they can be surveyed for natural enemies. Further molecular studies including possible parents of the Nymphaea hybrids in South Africa were thus carried out in this thesis to focus future surveying efforts. Two main hybrid groups were identified with genetic similarity to two tested putative Nymphaea parents, and this will allow further investigations of these species to improve the chances of successfully managing these hybrid groups. Some of the tested hybrids showed genetic contributions from multiple groups, some of which were unidentified, so it is necessary to prioritise the most problematic hybrids for biological control. With more insight into the genetic makeup of the Nymphaea hybrids in South Africa, investigations into the host specificity of potential biological control agents can be conducted. The ideal biological control agent should have a broad enough host range to impact and survive on both N. mexicana and its hybrids, but without a host range so broad that it would pose risk to native South African species. Host specificity trials are thus necessary to determine the suitability of potential agents. The identification of B. longulus feeding on N. mexicana during pre-release surveys motivated further investigations to determine the natural distribution, field host range, and host specificity of B. longulus in experimentally controlled conditions. Further surveys were therefore conducted at native Nymphaea sites around South Africa in addition to host specificity trials using the native Nymphaea nouchali Burm. f. (Nymphaeaceae), two populations of N. mexicana, and a cultivated hybrid. Results from the surveys and host specificity tests suggest that B. longulus is widely distributed across South Africa, is specific to Nymphaea with no observed preference between N. mexicana and the native N. nouchali, and does not perform well on Nymphaea hybrids. Hence, B. longulus is promising for use in new association biological control through augmentative releases but is not suitable for management of hybrids. In addition to the potential use of the South African B. longulus, it is necessary to conduct host specificity trials for natural enemies from the native range of N. mexicana that were prioritised in previous studies. Megamelus toddi Beamer (Hemiptera: Delphacidae) is one such species that was imported into quarantined laboratory conditions from Florida, USA. Host specificity trials were conducted using the same test plants as described for the studies on B. longulus, in addition to multigeneration trials to determine how long M. toddi could survive on non-target host plants. As with the B. longulus studies, no statistically significant differences in preference were observed between N. mexicana and N. nouchali, but M. toddi could not complete development on the test hybrid, indicating that this species is also unsuitable for the management of Nymphaea hybrids. Despite suboptimal plant health, M. toddi completed development for three generations on the native N. nouchali. This lack of host specificity deems M. toddi unsafe for release in South Africa but highlights the importance of following predefined steps to develop a biological control programme. The concluding chapter of this thesis discusses the aforementioned findings in a broader context by considering the driving forces of plant invasions in general and specifically for N. mexicana in South Africa. Case studies are also consulted to provide insight into how to proceed with managing Nymphaea hybrids in South Africa, while the factors governing host specificity and host range expansion are also discussed and considered in the context of B. longulus and M. toddi. Finally, after a consideration of the limitations of these studies, recommendations are made to continue the development of biological control for N. mexicana in South Africa. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2023
- Full Text:
- Date Issued: 2023-03-31
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