Disentangling the role of prokaryotes in regulating export flux via suspended and sinking organic matter in the southern ocean
- Authors: Dithugoe, Choaro David
- Date: 2022-10-14
- Subjects: Microbial ecology , Bioinformatics , Biochemistry , Oceanography , Metagenomics , Carbon cycle (Biogeochemistry) , Prokaryotes
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/365745 , vital:65782 , DOI https://doi.org/10.21504/10962/365745
- Description: The role of phytoplankton in regulating atmospheric carbon dioxide in the marine environment has been the subject of extensive research. We lack, however, comparative insights regarding the functional contributions of bacteria, archaea, fungi, and viruses (the microbiota) to organic matter export especially in understudied polar marine environments such as the Southern Ocean. This knowledge deficit is in part due to the high levels of microbial diversity which obscures efforts to study the relationship between diversity and ecosystem functions including their roles in the sequestration of carbon and nitrogen. Elucidating their precise contributions to organic matter export may be central to potential ecosystems feedbacks to global climate change. We examined several factors which may influence organic matter export to depth including net primary production, phytoplankton biomass, temperature, and prokaryotic functional capacity in the Southern Ocean. A Marine Snow Catcher was used to collect suspended and sinking material 10 metres below mixed layer depth at Southern Ocean Time Series (SOTS) in autumn (March-April) and in the Atlantic sector of the Southern Ocean in winter (July-August) and spring (October-November) 2019. The suspended and sinking material was used to determine the particulate organic carbon and nitrogen concentrations which were then used to calculate fluxes and export ratio ((e-ratio) - particulate organic carbon flux divided by net primary production). Additionally, genomic DNA was extracted from the suspended and sinking material and sequenced to obtain Shotgun metagenomic data which was employed to reconstruct metagenome assembled genome (MAGs) and their functional capacity using bioinformatic tools such as DRAM. Data from the Atlantic sector of the Southern Ocean, demonstrate that net primary production and temperature were inversely related to the e-ratio which is consistent with previous findings from the northern region of the Southern Ocean. Genomic functional capacity from SOTS suggested that r-strategist (organisms adapted to live in unstable environments) bacteria (e.g., Gammaproteobacteria) were prominent in the suspended pool. By contrast, the sinking particle-pool appeared to be dominated by K- strategists (organisms adapted to stable environment). The opposite was true for the archaea. This finding (i.e., bacteria) differs from a previous study in the northern region of the Southern Ocean, showing that microbes with K-strategists were more abundant in the suspended fraction. K-strategists typically degrade sinking organic matter into suspended organic matter or dissolved organic matter reducing the organic carbon export efficiency. Furthermore, Data from the Atlantic sector of the Southern Ocean revealed that seasonal temperature changes might dictate the rate of regional prokaryotic degradation across the zones. Resulting in rapid degradation at the northerly warmer regions and slow degradation further south. The data further provide evidence of chemolithoautotrophic mechanisms, with prokaryotes harbouring key pathways, required to transform dissolved inorganic carbon into complex organic forms, including recalcitrant dissolved organic carbon. Collectively, the SOTS and Atlantic sector of the Southern Ocean data suggest that shifts in prokaryotic community structure and functional capacity may regulate (either degradation or synthesis of organic matter) carbon export to depth. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2022
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- Date Issued: 2022-10-14
Systematics of the Afrotropical Chalcididae (Hymenoptera: Chalcidoidea)
- Authors: Faure, Sariana
- Date: 2021-10-29
- Subjects: Uncatalogued
- Language: English
- Type: Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/192808 , vital:45266
- Description: Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2021
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- Date Issued: 2021-10-29
Functional biogeography: evaluating community assemblage patterns and ecosystem functioning in intertidal systems using trait-based approaches
- Authors: Gusha, Molline Natanah C
- Date: 2022-10-14
- Subjects: Biogeography , Marine algae , Benthic ecology , Invertebrates , Functional redundancy , Ocean temperature , Biology Classification
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/365792 , vital:65790 , DOI https://doi.org/10.21504/10962/365792
- Description: Analyses of taxonomic diversity patterns within coastal systems has been critical in the development of the theory of biogeography. Increasing evidence, however, shows that the variety of functions that species perform in ecosystems (rather than their taxonomic identity) is a better predictor of the influence of the environment on the species. This information has been useful in predictive ecology leading to the development of trait-based approaches (TBA). Until the late 1970s, however, limited effort (particularly in marine systems) was channeled towards patterns in functional species traits and how they may be affected by changes in environmental gradients. Here, I mapped the functional biogeography of the South African coastline based on a suite of species' reproduction and development traits. Because species composition is one of the key tools used by zoogeographers to map species distribution patterns, I expected lower variability in trait composition within main biogeographic regions than in intervening transition zones based on the habitat templet theory and following the biomass ratio and limiting similarity hypotheses. In brief, the habitat templet theory proposes that “the habitat provides a templet upon which evolution forges species characteristics”, while the biomass ratio hypothesis assumes that the most abundant species traits determine ecosystem functioning. The limiting similarity hypothesis also sometimes referred to as the niche complementarity hypothesis, however, predicts that species can coexist if their niches complement one another. In light of the habitat being an evolutionary templet, abiotic and biotic habitat patterns were measured as nearshore SST and chlorophyll-a gradients, respectively. I expected the SST gradient to act as the stronger key filter of trait diversification because temperature is often considered the most influential environmental factor affecting species survival with seasonality of SST affecting the timing of spawning and along with food availability, possibly influencing fecundity. Functional trait data were thus compiled for macroinvertebrate species collected from fifty-two rocky shore sites from three main bioregions (east, south, and west) and two transition zones (south-west and south-east). Biological trait analysis and functional diversity indices were used to evaluate how traits related to species development and reproduction respond to temperature and chlorophyll -a (used as a proxy for food availability) gradients along the coastline. GLMM and hierarchical cluster analyses showed distinct patterns/shifts in SST and chlorophyll-a gradients across bioregions, with two main breaks in SST separating the east and south-east overlap (SEO) bioregions from the south, south-west overlap (SWO) and west bioregions. In contrast, chlorophyll-a exhibited three major breaks with the east, SEO–south–SWO, and west clustering independently of each other. The RLQ analysis (a type of co-inertia analysis) which simultaneously ordinates 3-matrix datasets [i.e., (environment × site[R]), (species × site[L]) and (species × traits [Q])] showed that the higher SST gradient on the east and SEO promoted higher abundance and biomass of simultaneous hermaphrodites while higher chlorophyll-a gradients on the SWO and west coasts strongly promoted reproductive maturity at larger-sizes. The combined fourth-corner analyses showed that the modalities within the development trait domain responding to chlorophyll-a gradients primarily included filter feeders, sessile and swimming species and also species living on the infratidal zone. In addition, the reproduction trait domain showed higher sensitivity and association to differences in chlorophyll-a and SST gradients than development traits. Overall, SST and chlorophyll-a gradients influenced the distribution of the most dominant traits as indicated by shifts in community-weighted mean trait values across bioregions. This suggests the importance of habitat filtering on coastal species reproduction. A separate study evaluating the influence of large-scale biogeographic effects vs the micro-scale biogenic habitat structure offered by coralline seaweeds across 24 sites revealed some notable effects of both factors on the diversity and abundance of macroalgal epifauna. There was a notable biogeographic influence on epifauna, with the SEO recording the highest epifaunal species richness and abundance, followed by the south coast, then the SWO and lastly the west coast. In addition, the total biomass gradient of the corallines followed a similar trend. The epifauna however, showed no host-specificity, illustrating that epifauna may not be species–centric as commonly assumed, and the higher diversity of epifaunal diversity may well be simply because those corallines are the available habitat within the sampled part of the coastline. Lastly, macroinvertebrate trait distribution on the South African coastline confirms that the habitat, particularly the biotic filter (in this case chl-a) provides a templet upon which evolution forges species traits. However, since temperature is a proxy for nutrient availability (cold upwelling brings nutrients), then temperature drives chlorophyll-a. Subsequently this means the abiotic component indirectly drives trait distribution by influencing the biotic environment (chl-a). For epifauna species, also, the coralline diversity and composition can also be regarded as a biotic filter influencing the epifaunal abundances and composition across different bioregions. Moreover, since temperature is regarded as a conservative trait in seaweeds, temperature tolerance defines the biogeographical boundaries of seaweeds, therefore temperature may be indirectly affecting epifauna abundances through coralline species diversity and biomass. In summary, considering the deterministic processes governing ecosystem functioning and community assemblage, the mass ratio and limiting similarity hypotheses showed complementary effects. Different bioregions provided variable support for these two hypotheses, but overall, the mass ratio hypothesis (weighted by species biomass) received stronger support and may be more meaningful to the interpretation of ecosystem functioning and persistence within rocky shore systems. Lastly, although, the SWO showed some of the characteristics of a subtraction zone based on the relatively low abundance, diversity, and biomass measures. Nonetheless, there was evidence of high functional redundancy across all other four bioregions. This suggests that in the context of development and reproduction traits, the rocky shore ecosystem along the SA coastline may be functionally stable at this stage. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2022
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- Date Issued: 2022-10-14
Integrative systematic structuring of the widespread psammophiid snakes (Psammophiidae): a multi-evidence species delineation approach
- Authors: Keates, Chad
- Date: 2021-10-29
- Subjects: Psammophis South Africa , Herpetology , Herpetology Africa , Molecular biology , Psammophis Classification , Psammophis Genetics , Psammophis Morphology , Psammophis Phylogeny , Morphology Mathematics , Psammophylax
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/295077 , vital:57288 , DOI 10.21504/10962/295079
- Description: Species form the foundations upon which we build our understanding of the natural world. Although a focus of much scientific attention, our understanding of species is stunted by the intrinsic ‘fuzziness’ of boundaries within nature. Due to the complexity of the evolutionary process, coupled with an ever-changing abiotic landscape, species are hard to delineate, even at the best of times. Whilst various species concepts and sophisticated delimitation methods have helped scientists tease apart species, many species complexes persist. This is because taxonomy is a discrete ordering system imposed upon the continuous and intercalated structure of life. To improve our understanding of a wide-ranging family of snakes, I investigated the taxonomy and evolutionary structuring within Psammophiidae using both molecular and morphological approaches, employing phylogenetic, phylogeographic, and morphometric analyses on the group. The systematic complexity of the family (as evidenced by past research) coupled with the group’s widespread distribution and ecological importance, made the taxon an ideal candidate for a broad-sweeping multi-level systematic analysis using multiple species delimitation methods. Additionally, in this thesis I attempted to build on the ground-breaking work of Christopher Kelly by addressing several knowledge gaps identified within the family, and in so doing, produce the most thorough evolutionary and taxonomic study of Psammophiidae possible. Given the taxonomic uncertainty associated with the family, Chapter Two used a representative sampling from every available species (near complete taxon sampling approach) in the family. The chapter used both standard and time-calibrated phylogenetic modelling and distance/threshold-based species delimitation, to elucidate the finer-level structuring within the family. Geometric morphometrics was used to determine whether there were diagnosable differences in head structure between the different genera. The final phylogenetic tree incorporated 320 samples, representing the most comprehensive phylogenetic reconstruction of the family to date. By using a near-complete taxon sampling approach, I was able to resolve previously unsupported relationships within the family whilst also identifying several novel instances of an under- and over-appreciation of species diversity within the family. Geometric morphometrics also identified clear distinctions between genera based on head shape (head width and ‘beakedness’). This chapter showcased the importance of complete taxon sampling and robust methodology for species delimitation and the deleterious effect of species concepts when implemented in isolation. In Chapter Three, I narrowed the scope of the study to focus on the genus level. Psammophylax (Fitzinger 1843) is an abundant, yet poorly studied genus of grass snakes, endemic to Africa. The generalist nature of the genus and wide-spanning distributions of the constituent species has given rise to several subspecies and a poor understanding of the taxonomic structuring within the genus. The overlapping distributions (sympatry) of many of Psammophylax species, coupled with the potential for cryptic speciation via mechanisms such as convergent evolution, made the group the ideal candidate for a broad-sweeping systematic study (as evidenced in Chapter Two). By applying the suite of analyses used in Chapter Two to the generic level, we aimed to determine the effectiveness of a multi-evidence species delineation approach when tackling systematic problems at lower taxonomic levels. A genetic phylogeny of six of the seven species was estimated using multiple phylogenetic and distance/ threshold-based species delimitation methods. To support the molecular analyses, we conducted morphological analyses on the body (traditional morphology) and head (geometric morphometrics) separately. Phylogenetic analyses recovered a similar topology to past studies, but with better resolution and node support. I found substantial genetic structuring within the genus, supported by significantly different head shapes between Ps. a. acutus and other Psammophylax species. Psammophylax a. acutus was recovered as sister to its congeners, and sequence divergence values and morphometrics supported its recognition as a new genus. Increased sampling in East Africa (Tanzania, Kenya, and Ethiopia) revealed that Psammophylax multisquamis is polyphyletic, necessitating the description of a new, morphologically cryptic, species from northern Tanzania. The distribution of Ps. multisquamis sensu stricto is likely restricted to Kenya and Ethiopia. Within this chapter, taxon-specific phylogenetic analyses yielded stronger intrageneric support as compared to Chapter Two, allowing for more defensible conclusions about taxonomical amendments. Geometric morphometrics proved similarly useful (as compared to Chapter Two) in teasing apart genera within the family but lacked the robustness to delineate species within Psammophylax with confidence, highlighting the apparent convergence of form within the genus. In Chapter Four, I investigated the evolutionary structuring within the Southern African endemic Psammophylax rhombeatus. The structural and environmental heterogeneity within the region has given rise to many morphological forms distributed throughout the country, with previous studies neglecting the associated molecular significance of these forms. Irrespective of their small sample sizes, both Chapter Two and Three identified substantial phylogenetic structuring within the species, making Ps. rhombeatus the ideal candidate for a multi-faceted systematic review, using a combination of phylogenetics, geometric morphometrics and, for the first time in this species, phylogeographic analyses. By investigating a single species, in detail, I was able to assess the effectiveness of the methodologies implemented in previous chapters on systematic sorting using the multi-evidence species delineation approach. Phylogenetic and haplotype analysis retrieved four well-supported clades: southeast South Africa (SESA), southwest South Africa (SWSA), north-eastern South Africa (NESA) and western South Africa (WSA). Although not variable enough to warrant taxonomic re-evaluation, the clades represented important genetic hotspots, with relatively high intraspecific genetic divergence values separating them, irrespective of the small geographic distances separating populations. This is likely a product of the taxon’s habitat-generalist lifestyle, enabling them to bypass vicariant barriers that might otherwise cause speciation in less versatile species. The clades are also geographically distinct, with little overlap, indicating previous vicariance, a finding that is supported by the split of Ps. rhombeatus from Ps. ocellatus in the mid-Pliocene, followed by the diversification of Ps. rhombeatus into four clades throughout the Pleistocene. The genetic structuring observed in Ps. rhombeatus may be a product of population expansion following ancient refugial isolation (potentially Last Glacial Maximum [LGM]). The molecular distinctiveness of the clades was not replicated in the morphological component of this chapter, with neither dorsal nor lateral geometric morphometric analyses of head shape showing any discernible distinctiveness based on geography. Whilst head shape has not been shown to be an effective delineator of evolutionary units at the species level (within this taxon), body colour, scalation, and snout-vent length has been linked to morphotypes within the species based on the work of Broadley (1966). These morphological groupings are loosely attributable to the molecular clades identified in the phylogenetic analyses, highlighting the complex interplay of genetic and morphological characteristics in the process of speciation, and their representation in systematic accounts. This thesis represents the most thorough evolutionary and systematic study of the family currently possible. In addition to identifying and describing both a new genus and species, this thesis also highlighted several instances of an over- and under-appreciation of species diversity within Psammophiidae. By applying a multi-evidence species delineation approach to this thesis, I show the intricacy of the evolutionary process (at various taxonomic levels) and showcase the ease to which species boundaries can be confounded when species concepts are implemented in isolation. These findings also highlighted the importance of sample size, sample range, species delimitation method on the outcome of taxonomic analyses, and their interpretation. Lastly, this thesis addressed the knowledge gaps left by Christopher Kelly’s PhD work and investigated the findings of recent papers that attempted to do the same. Whilst this study answers the questions of old, the taxon-intensive focus revealed several new knowledge gaps within the family, highlighting how much we know about snake systematics, and furthermore, how much we still need to learn about evolutionary structuring. , Thesis (PhD) -- Faculty of Science, Environmental Science, 2021
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- Date Issued: 2021-10-29
Distribution, ecological and economic impacts and competition of the invasive alien aquatic weeds (Pontederia crassipes Mart., Pistia stratiotes L., Salvinia molesta D.S. Mitch. and Azolla filiculoides Lam.) in Madagascar
- Authors: Lehavana, Adolphe
- Date: 2021-10-29
- Subjects: Pontederiaceae Madagascar , Water lettuce Madagascar , Salvinia molesta Madagascar , Azolla filiculoides Madagascar , Introduced aquatic organisms , Aquatic weeds Economic aspects , Aquatic weeds Social aspects , Aquatic weeds Geographical distribution
- Language: English
- Type: Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/191225 , vital:45072
- Description: In Madagascar, as in several countries in the world, the invasion by four aquatic weeds (Pontederia crassipes Mart. (Pontederiaceae), Pistia stratiotes L. (Araceae), Salvinia molesta D.S. Mitch Salviniaceae) and Azolla filiculoides Lam. (Azollaceae) are among the drivers of environmental and socio-economic deterioration in aquatic ecosystems. Pistia stratiotes was first recorded on the island in the 19th century, and P. crassipes from the beginning of the 20th century, while S. molesta and A. filiculoides were only documented during in the 21st century. From the 1920s, botanists such as Henri Perrier de la Bathie and Raymond Decary were already aware of the dangers caused, in particular by P. crassipes in other countries, and raised the alarm, but little attention has been paid to these species. The aim of the research conducted for this thesis was to determine the distribution, socio-economic and ecological impacts of these four invasive alien aquatic weeds in Madagascar and to make recommendations for their control. First, the distributions of these four aquatic weeds were mapped. This mapping exercise compiled data from different sources including herbarium records, online data and field visits across Madagascar. The mapping study was undertaken from August 2015 to June 2020. Except for mountainous areas above 1800 m (Tsaratanana Massif, Ankaratra Massif and Andringitra Massif) where no data were available, all of Madagascar's bioclimates were invaded by at least one of the four aquatic weeds. In total, at least one species was recorded in 18 of the 22 Regions. Pontederia crassipes was recorded in 13 Regions, S. molesta in 14 Regions, P. stratiotes in 12 Regions, and A. filiculoides in 13 Regions. Herbarium records revealed the oldest record for P. stratiotes to be 1847, 1931 for P. crassipes, 1995 for S. molesta and there were no herbarium specimens for A. filiculoides prior to the start of the current study in 2015. We now know where these four weeds occur and how abundant they are. An objective of this research was to assess the impacts of the four invasive aquatic plants on the socio-economy of the island, mainly on rice production and fishing. Between 2016 and 2019, 102 households in three regions, Soanierana Ivongo, Foulpointe and Antananarivo, were randomly selected and questioned on the impact of these weeds in their aquatic ecosystems and their livelihoods such as fishing and rice growing. Surveys revealed that the four aquatic weeds significantly threatened household activities. On the east coast of Madagascar, the invasions of these four invasive species decreased fish and freshwater shrimp production by 82%. On the high plateau of Madagascar, they reduced rice yield by 30% despite requiring an additional expense of US$ 1,107/ha for control. Although farmers surveyed only used manual control to manage these weeds, they were receptive to other control methods, including integrated control using herbicides and biological control. Another objective of this research was to determine the ecological impacts of the four weeds and specifically if freshwater ecosystem functioning would return after control. To assess the ecological impact, between February 2017 to August 2019, on Lake Antsokafina, the following abiotic and biotic factors were considered: physico-chemistry of water, succession of macrophyte community and animal diversity. With the exception of turbidity, the values of the physico-chemical parameters of the water (pH, electrical conductivity, water temperature and turbidity), were similar between the infested zone and cleared zone. A study on the invasion process of aquatic weeds showed that the plant community succession of the lake changed over time in the areas that had been cleared. The submerged species Ceratophyllum demersum was the pioneer, followed by creeping species such as Echinochloa colona and Ipomoea aquatica, before the area was recolonized by aquatic weeds. Among the aquatic weeds, S. molesta was the most aggressive, covering 92% of the area one year after the start of the experiment. For animal diversity, bird, shrimp and fish community were assessed. The cleaning of the plots in the lake allowed the resumption of fishing activity providing 50 to 200g/catch for shrimp and from 0.25 to 0.5kg/catch for fish per person per day, while no catch was obtained in the areas infested by aquatic weeds were fishermen still attempting to harvest fish/shrimp from the aquatic weed infested areas. Three species of birds, Humblot’s Heron (Ardea humbloti), the white-faced whistling duck (Dendrocygna viduata) and red-billed teal (Anas erythrorhyncha) returned once the areas had been cleared. A manipulated outdoor as descriptor for laboratory experiment was conducted to determine the level and nature of competition of four aquatic weeds species against the indigenous floating fern, Salvinia hastata Desv. (Salviniaceae), using an additive series density model. It was shown that all four invasive species outcompeted S. hastata, with P. crassipes being 24 times more dominant, followed by P. stratiotes at 12 times, S. molesta at 8 times, and finally A. filiculoides at 1.2 times more dominant. This study provided direct evidence of the biodiversity impact of these four species and thus also provided an environmental argument for their control. Based on the findings of this study, a series of recommendations was formulated to manage the invasions of alien species in Madagascar with particular attention to invasive aquatic weeds. These recommendations mainly concern the establishment of management structures and legal instruments such as the creation of a lead government agency at national level and a cross-sectorial invasive species advisory committee, which should review legislation and regulations related to invasive species. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2021
- Full Text:
- Date Issued: 2021-10-29
Quantifying the impacts of cactus biological control agents in South Africa
- Authors: Mnqeta, Zezethu
- Date: 2022-04-08
- Subjects: Cactus Biological control South Africa , Noxious weeds South Africa , Invasive plants Biological control South Africa , Insects as biological pest control agents South Africa , Cochineal insect South Africa , Mealybugs South Africa , Agricultural productivity South Africa
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/294631 , vital:57239 , DOI 10.21504/10962/294631
- Description: Invasive alien cacti are prominent weeds that threaten indigenous biodiversity and have a negative impact on agricultural productivity in South Africa. These plants are problematic because they form dense thickets that reduce the carrying capacity of rangelands; restrict the movement of livestock and wildlife thus reducing access to shade and water sources; and are directly harmful to livestock, wildlife and people due to their sharp spines. Biological control is the most effective, affordable and environmentally friendly method to control invasive alien cacti and minimize their negative impacts. Cochineal insects (Dactylopius spp.: Dactylopiidae) and the gall-forming mealybug, Hypogeococcus sp. (Pseudococcidae) are used as biological control agents for cacti. The agents are however poor dispersers, so mass-rearing and augmentative releases are required in order to establish the agents at sites where they are needed. This study aimed to evaluate the mass-rearing and release efforts of cactus biological control in South Africa, quantify the impact of the biological control agents on cactus plant populations through long-term monitoring, and assess the benefits accrued due to the biological control agents through the perceptions of land-users. An assessment of the effectiveness of the release effort for cactus biological control agents was conducted by comparing where biological control agents have been released with the known distribution of the target weeds. Only 26% of the quarter degree squares that are known to be occupied by invasive alien cacti have had biological control agents released in them. This indicated that the mass-rearing and release efforts in South Africa are inadequate and should be increased, especially in areas where few releases have been made and many cacti are present, such as the Limpopo Province. The impact of the biological control agents on cactus plant populations was assessed by monitoring agent densities, target plant densities and target plant reproductive outputs before and after releases. Plant biomass and reproductive output were reduced by biological control agents for three of the target weeds that were assessed, while the duration of the study was too short to measure reductions for the fourth target weed. Benefits to land-users were then quantified through a questionnaire survey. Land-users perceived biological control as an effective management option, with 81% of the land-users reporting that there was less invasive alien cactus after releasing biological control agents on their land. Forty-nine percent of the land-users believed that the negative impacts of the cactus had been reduced and that they benefited more from their land since control was achieved. Since land-users were only interviewed within four years of the releases being conducted, it is expected that the percentage of land-users who gained benefits from biological control will increase in future. Ninety-seven percent of the land-users stated that the agents were safe and had not fed on any other plants or had any detrimental impacts. These perceptions indicated that land-users regarded biological control as a safe and effective method of controlling invasive alien cacti. This study confirms that biological control is an effective and safe way of controlling invasive alien Cactaceae. It is also the first to assess some of the benefits that land-users have accrued due to biological control of cactus weeds. It is however evident that a greater mass-rearing and release effort is required for South Africa to get the maximum benefits possible from the use of the biological control agents for cactus weeds that are available in the country. , Izityalo ze-cactus ezisuka kwamanye amazwe zilukhula olubalaseleyo, olwenza ingxaki kwintlobo ngeentlobo zezityalo nezilwanyana kwaye ezizityalo zinefuthe neziphumo ezingalunganga kwimveliso yezolimo eMzantsi Afrika. Ezi zizityalo ziyingxaki kuba zenza amatyholo ashinyeneyo athi anciphise umthamo wokusebenzisa umhlaba: zinqanda ukuhamba nogkukhululekileyo kwemfuyo nezilwanyana zasendle ngokwenza oko zingakwazi ukufikelela emthuzini xa kutshisa nakwimithombo yamanzi yokusela; zikwayiyo nengozi kwimfuyo, izilwanyana zasendle kunye nabantu ngenxa yamave abukhali afumaneka kwezizityalo. Ukulawula nokwehliza ubunzini nezinga lemigcipheko yezizityalo, kusetyenziswa indlela ekuthiwa yi-biological control. Lendlela yeyona ndlela isebenza ngokuphucukileyo, efikelelayo, nengenabungozi kokusingqongileyo. Izinambuzane ze- cochineal (i-Dactylopius spp.: Dactylopiidae) kunye ne-mealybug, i-Hypogeococcus sp. (Pseudococcidae) zisetyenziswe njengezixhobo ze-biological control ezinceda ukulawula ezizityalo ze-cactus zingafunekiyo. Ingxaki yazo ezizinambuzane zizixhobo ze-biological control zingentla kukuba azikwazi kuzisasaza ngokwazo ukuba zifikelele nakwizityalo ezikude ngoko ke ukukhuliswa nokukhutshwa ngobuninzi bazo kuyafuneka ukukhawulelana nalengxaki kunye nokwandisa amathuba wokuba zifikelele kuzozonke izityalo ze-cactus ekufuneka zizilawule. Olu phononongo lujolise ekuvavanyeni iinzame zokukhulisa ngobuninzi nokukhupha ezezinambuzane zizixhobo ze-bioloigical control zokulawula izityalo ze-cactus eMzantsi Afrika, ukujonga ubungakanani befuthe notshintsho elenziwa zezinambuzane zizixhobo ze-bioloigical control kwizityalo ze-cactus emva kwexesha elide lokuzijonga, kunye nokuvavanya inzuzo efunyenwe ngenxa yokulawula ezizityalo ze-cactus ngokwemibono yabasebenzisi bomhlaba. Uvavanyo lweenzame zokukhutshwa kwezixhobo zezixhobo zezinambuzane ze-bioloigical control kwizityalo ze-cactus lwenziwa ngokuthelekisa iindawo apho izixhobo zezinambuzane ze-biological control zikhutshwe khona kunye neendawo apho izityalo ze-cactus kwaziwayo ukuba ziyafumaneka khona. Yi-26% kuphela yesikwere sekota eyaziwayo ukuba kukhona izityalo ze-cactus nalapho kukhutshwe khona kwezinambuzane ezizixhobo ze-biological control. Oku kubonisa ukuba iinzame zokukhulisa nokukhutshwa kwezinambuzane ezizixhobo ze-biological control eMzantsi Afrika azanelanga kwaye kufuneka zandiswe, ngakumbi kwiindawo apho kukho ukhutsho olumbalwa olwenziweyo kunye nezityalo ze-cactus ezininzi ezifumaneka khona, njengePhondo laseLimpopo. Iimpembelelo yokukhutshwa kwezinambuzane ezizixhobo ze-biological control kwizityalo ze-cactus zavavanywa ngokujonga ubunizi bezinambuzane ezizixhobo ze-biological control, ukuxinana kwezityalo ze-cactus ekujoliswe kuzo kunye nemveliso yokuzala kwezityalo ze-cactus phambi nasemva kokuba kukhutshwe izinambuzane ezizixhobo ze-biological control. Ubungakanani bezityalo ze-cactus kunye nemveliso yokuzala ziye zacutheka emva kokukhutshwa kwezinambuzane ezizixhobo ze-biological control kwinzityalo ezintathu ze-cactus ebekujoliswe uvavanyo kuzo, ngelixa ixesha lophononongo lalifutshane kakhulu ukuvavanya unciphiso kwisityalo se-cactus sesine. Izibonelelo zabasebenzisi bomhlaba zaye zavavanywa kusetyenziswa uhlobo lwemibuzo. Abasebenzisi bomhlaba balubona ukusetyenziswa kwezinambuzane ezizixhobo ze-biological control njengendlela yolawulo olusebenzayo, yi81% yabasebenzisi bomhlaba echaze ukuba izityalo ze-cactus zecuthekile emva kokukhutshwa kwezinambuzane ezizixhobo ze-biological control emhlabeni wabo. Amashumi amane anesithoba eepesenti zabasebenzisi bomhlaba bakholelwa ukuba impembelelo ezingalunganga ze-cactus zincitshisiwe kwaye baxhamle kakhulu kumhlaba wabo okoko lwaphunyenzwa. Oludliwano-ndlebe belwenziwe kubasebenzisi bomhlaba kwisithuba seminyaka emine emva kokuba kukhutshwe izinambuzane ezizixhobo ze-biological control, kulindeleke ukuba ipesenti yabasebenzisi bomhlaba abathe bafumana izibonelelo kwi-biological control lonyuke kwixesha elizayo. Amashumi alithoba anesixhenxe ekhulwini abasebenzisi bomhlaba bachaze ukuba izinambuzane ezizixhobo ze-biological control zikhuselekile kwaye khange zidle kuzo naziphi na ezinye izityalo ezingezi eze-cactus ebekujoliswe kuzo okanye zineempembelelo eyingozi. Ezi mbono ziyabonisa ukuba abasebenzi bomhlaba bayithatha i-biological control njengendlela ekhuselekileyo nesebenzayo yokulawula izityalo ze-cactus zamanye amazwe. Olu phononongo luqinisekisa ukuba ulawulo olusebenzisa izinambuzane ezizixhobo ze-biological control yindlela esebenzayo nekhuselekileyo yokulawula iCactaceae yamanye amazwe. Ikwangolokuqala ukuvavanya ezinye zezibonelelo ezizuzwe ngabasebenzisi bomhlaba ngenxa ye-biological control yezityalo ze-cactus. Nangona kunjalo kucacile ukuba ukwandisa inzame zokukhulisa nokukhupha ngobuninzi izinambuzane ezizixhobo ze-biological control kuyafuneka ukuze uMzantsi Afrika ufumane izibonelelo eziphakamileyo enokubakho ngokusetyenziswa kwe zinambuzane ezizixhobo ze-biological control zokulawula ukhula lwe-cactus olukhoyo kweli lizwe. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2022
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- Date Issued: 2022-04-08
Effect of Helicosporidium sp. (Chlorophyta; Trebouxiophyceae) infection on Cyrtobagous salviniae Calder and Sands (Coleoptera: Curculionidae), a biological control agent for the invasive Salvinia molesta D.S. Mitchell (Salviniaceae) in South
- Authors: Mphephu, Tshililo Emmanuel
- Date: 2022-10-14
- Subjects: Salvinia molesta South Africa , Weeds Biological control , Cyrtobagous salviniae , Ketoconazole
- Language: English
- Type: Academic theses , Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/365815 , vital:65792 , DOI https://doi.org/10.21504/10962/365815
- Description: The effectiveness of established biological control agents depends on biotic and abiotic interactions in the introduced range. The weevil, Cyrtobagous salviniae Calder and Sands (Coleoptera: Curculionidae), was released as a biological control against Salvinia molesta D.S. Mitchell (Salviniaceae) in South Africa in 1985. This agent has been highly successful against S. molesta and has significantly reduced the weed’s populations around the country. However, in 2007, the parasitic alga, Helicosporidium sp. (an undescribed species), was detected in field-collected C. salviniae adults in South Africa. The distribution and impacts of this disease on the weevil and its efficacy as a control agent were not known. In this thesis, the prevalence, infection load, and impact of Helicosporidium sp. on C. salviniae was determined. In 2019, adult weevils were collected from 10 sites across the Eastern Cape, KwaZulu-Natal, Limpopo, and Western Cape provinces and screened to determine the occurrence, infection load, and geographic distribution of Helicosporidium sp. Transmission mechanisms of this disease in C. salviniae were then evaluated. The possible impact of Helicosporidium sp. was assessed by comparing the feeding rates and the reproductive output of the diseased and healthy adults of C. salviniae. An attempt was then made to eliminate the disease in C. salviniae through the application of the antibiotic, ketoconazole. Further, the role of temperature on infection load in C. salviniae was also assessed. Finally, recommendations for the long-term biological control programme against S. molesta in South Africa were made. The disease covers the entire distribution range of C. salviniae in South Africa, with the disease occurrence rate ranging from 92.15% to 100% insects infected per site. Helicosporidium sp. was found to transmit vertically within the populations of C. salviniae. Infection by the Helicosporidium sp. disease reduced the reproductive output of C. salviniae as well its impact on biomass reduction of S. molesta when a diseased culture was compared to a healthy culture from the USA. 98.44 to 98.55% of Helicosporidium sp. loads were reduced through multiple applications of ketoconazole concentrations under in vitro trials. In vivo treatments resulted in 70% control of Helicosporidium sp. in the adults of C. salviniae that were fed ketoconazole three times over a 21 day period. Adult C. salviniae feeding and survival performances were similar when fed fronds of S. molesta inoculated with ketoconazole and water. The lowest and highest disease loads of Helicosporidium sp. were recorded when the weevils were reared at 30°C and 14°C, respectively. As expected, the highest impact and reproductive output of C. salviniae were at 30°C. The evaluations discussed in this thesis highlight the role of diseases in biological control agents, and gaps in both the pre-release and post-release monitoring that should integrate screening of diseases in these studies. Although the combined application of the antibiotic and temperature will reduce Helicosporidium sp. loads and impact, this technology is most likely only applicable where the weevils are reared in small numbers in a rearing facility and not really applicable to the field situation. It is important to release healthy agents that will cause efficient control of the target weed plant species, therefore, when introducing new biological control agents, the health status of such agents needs to be understood. Therefore, long-term field monitoring and assessment of the impact of C. salviniae on S. molesta should be conducted to track all the changes that may result due to the presence of Helicosporidium sp. This long-term monitoring and assessment will give a more informative role of Helicosporidium sp. in field populations of C. salviniae. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2022
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- Date Issued: 2022-10-14
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
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- Date Issued: 2023-03-31
The biological control of Egeria densa Planch. (Hydrocharitaceae) in South Africa
- Authors: Smith, Rosali
- Date: 2021-10-29
- Subjects: Egeria (Plant genus) Biological control South Africa , Hydrocharitaceae Biological control South Africa , Aquatic weeds Biological control South Africa , Leafminers South Africa , Plant invasions South Africa , Resilience (Ecology) South Africa , Freshwater ecology South Africa , Hydrellia South Africa , Submerged macrophyte
- Language: English
- Type: Doctoral theses , text
- Identifier: http://hdl.handle.net/10962/191102 , vital:45060 , 10.21504/10962/191102
- Description: Over the last thirty years, biological control, the use of host-specific natural enemies, has been a huge asset in the management exotic aquatic macrophytes such as Pistia stratiotes L. (Araceae), Pontederia crassipes Mart. (Solms) (Pontederiaceae), Azolla filiculoides Lam. (Azollaceae), Salvinia molesta D.S. Mitch (Salviniaceae) and Myriophyllum aquaticum (Vell.) Verdc. (Haloragaceae), also known as the “Big Bad Five” in South Africa. Despite these successes, freshwater ecosystems in South Africa have been harder to restore to an invasive macrophyte-free space, due to chronic disturbances such eutrophication, propagule dispersal and hydrological alterations. In the Anthropocene, where human activities have profound effects on their environment, these disturbances weakens ecological resilience and drive aquatic plant invasions. Due to long periods of invasions and the presence of a new suite of exotic aquatic plant propagules, native vegetation recolonization has been slow or even absent. Instead, the release of resources, such as sunlight, nutrient and space through aquatic weed management acts as a catalyst for secondary biological invasion. New invasive aquatic weeds include submerged and rooted emergent growth types, with Egeria densa Planch. (Hydrocharitaceae) the most widely distributed submerged aquatic weed in South Africa. It can quickly form dense monoculture stands that have ecological, economic and social impacts. Because of its ability to regenerate from plant fragments with double nodes, mechanical control is inappropriate. Additionally, mechanical and chemical control not only affects E. densa but have significant non-target effects. In response to its rapid spread over the last 20 years, especially following floating invasive aquatic management, a biological control programme was initiated, and in 2018, the leaf-mining fly, Hydrellia egeriae Rodrigues (Diptera: Ephydridae) was released. This was the first release of a biological control agent against E. densa in the world, and the first agent released against a submerged aquatic weed in South Africa. This thesis comprises the subsequent step of a biological control program when permission for the release of an agent have been obtained. A brief history of macrophyte invasions in South Africa’s unique freshwater systems are given in the literature review. Contributing factors to secondary invasions within the context of ecological resilience are introduced. An argument for the benefit of biological control as nuisance control is given, especially because E. densa and its natural enemy, H. egeriae is the focus species of this thesis. The main goal after permission for the release of an agent have been obtained, is to establish and build-up field populations. Research questions in this thesis aimed to investigate factors that contribute to or negate this goal. Through laboratory and field experiments we investigated the thermal physiology of the agent, and its climatic suitability to its novel range; different release strategies on field establishment and biotic resistance through the acquisition of novel parasitoids. Considering the longevity of this biological control program, we investigated the effects of elevated CO2 on the interaction between E. densa and H. egeriae through open top chamber experiments. Laboratory thermal physiology results showed that the agent is able to survive, develop and proliferate at all E. densa sites throughout the year. This is confirmed with the establishment of the agent at two release sites, the Nahoon River in the Eastern Cape Province and the Midmar Dam in KwaZulu-Natal. Post-release surveys showed that H. egeriae requires augmentative releases to sustain field populations. Without augmentative releases, H. egeriae herbivory levels were almost negligent. However, a contributing factor to low field-populations was parasitism. The biological control agent acquired three parasitoids, which have previously been described from Hydrellia lagarosiphon Deeming (Diptera: Ephydridae), a specific herbivore to Lagarosiphon major (Ridl.) Moss (Hydrocharitaceae). These results provide information on the immediate establishment and effectiveness of the H. egeriae. Results from the elevated CO2 study suggest that E. densa will become less nutritious through a shift in leaf C/N ratio, when ambient 800ppm is bubbled into experimental growth chambers. Hydrellia egeriae feeding was affected by ambient CO2 levels and plant nutrient availability. The set levels of ambient CO2 levels used in this experiment produced dissolved inorganic carbon levels that were lower than dissolved inorganic carbon levels in E. densa invaded sites. This suggests that, submerged aquatic plant-insect interactions may be harder to predict from only laboratory experiments. Further investigations are necessary to establish system-specific characteristics i.e. dissolved inorganic carbon and target plant nutritional quality. The biological control of E. densa in South Africa is still in its infancy. This study presents results from post-release surveys up until two years after the agent was released. From this study, Hydrellia egeriae exhibits the potential to be an effective biological control agent, but release strategies should be adapted to sustain field populations and to limit field parasitism effects. Continued post-release surveys will provide a more comprehensive idea of the seasonal fluctuations of field-populations and parasitism. Surveys at multiple sites will provide information on potential site specific characteristics that contribute to or negate biological effort. Considering the high nutrient status of South African freshwater systems, a more holistic approach to E. densa management is necessary. This will require the strengthening of ecological resilience to prevent systems from shifting into an alternate invasive stable state. In addition, aquatic weed management needs to be addressed by a resilient social network, which ultimately calls for the strengthening of socio-ecological resilience. , Thesis (PhD) -- Faculty of Science, Zoology and Entomology, 2021
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- Date Issued: 2021-10-29