School of Animal, Plant and Environmental Sciences (ETDs)

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    The past, present and future of cactus invasions in South Africa in response to rising atmospheric CO2 and climate change
    (University of the Witwatersrand, Johannesburg, 2023) Venter, Nicolaas; Byrne, Marcus
    Cactaceae originate from the Americas and over the past 250 years have been introduced into South Africa and elsewhere for agricultural and horticultural purposes. Numerous species, including useful taxa, have become important invasive weeds that have serious socio-economic and environmental impacts. Fortunately, management strategies, in particular biological control, have been successful in controlling certain species. However, with rising atmospheric CO2 invasive cacti are likely to pose a renewed threat, whereby evidence shows that cactus species are responsive to CO2 fertilisation, which is likely to increase their vigour, mainly through improved water use efficiency (WUE). Additionally, studies show that plant quality declines with increasing CO2 which in general has negative effects on their arthropod herbivores. This study sought to determine the effect of CO2 on two functionally different invasive cactus species and how they may respond to predicted increases in atmospheric CO2. Opuntia stricta (a succulent shrub) is an obligate CAM photosynthetic species that invades grasslands and savannas across semi-arid to subtropical environments whereas Pereskia aculeata (a scrambling woody vine with well-developed leaves) is a CAM-cycling photosynthetic species that invades forest type habits across subtropical environments. Plants were grown at three CO2 concentrations that represented pre-industrial (sub-ambient - 250 ppm), current (ambient - 400 ppm) and future (elevated - 600 ppm) atmospheric CO2 conditions. Plants were also subject to suboptimal and optimal watering treatments for the duration of the experiments to determine the ameliorative effect of CO2 on productivity in response soil moisture deficits. In addition, an experiment was conducted on O. stricta to test the effects of the three CO2 concentrations on plant quality and the subsequent effect on the fitness and efficacy of its insect biological control agent, Dactylopius opuntiae. Growth and productivity of O. stricta and P. aculeata responded positively to increasing CO2, however the responses varied with CO2 concentration. Increasing atmospheric CO2 concentration from date of introduction to present possibly facilitated O. stricta invasion whereas this was less likely for P. aculeata. In both species WUE increased with increasing CO2. Under suboptimal watering, there was partial amelioration of productivity at 600 ppm CO2, but the plant traits that benefitted varied according to species. Plant quality declined for both species, most notably cladode nitrogen (N) content decreased, and carbon/nitrogen ratios (C/N) increased. When D. opuntiae were exposed to O. stricta grown at elevated CO2 (only tested on well-watered plants), D. opuntiae fitness was reduced which in turn reduced the rates of plant mortality due to the insect damage. Using the WUE results from O. stricta, a mechanistic species distribution model (SDM) created here predicted greater increases in its potential distribution in South Africa under climate change relative to the SDM that did not include WUE as a predictor variable. This suggests that improved WUE under future CO2 concentrations can offset the effect of declining rainfall in certain regions of South Africa. Overall, these results suggest that O. stricta and P. aculeata will show more vigorous growth and are also likely to expand their ranges into regions where rainfall currently limits their distribution. This expanded distribution may be further facilitated by reduced biocontrol agent efficacy as host plant quality declines. These findings suggest that management of these two species and other invasive cacti is likely to become more challenging with increasing atmospheric CO2 and climate change.
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    The reproductive system of Campuloclinium macrocephalum and its implications for biocontrol implementation
    (University of the Witwatersrand, Johannesburg, 2022) Moodley, Saness; Glennon, Kelsey; Goodman-Cron, Glynis
    Invasive species are a threat to biodiversity therefore it is imperative to determine the factors that facilitate the invasion potential of a species. Campuloclinium macrocephalum Less. (DC), the ‘pompom weed’, is an alien invasive species in South Africa and is currently threatening the persistence of the grassland, wetland, and savanna biomes. The species is also significantly contributing to a decline in plant diversity by outcompeting native vegetation in these areas. Various integrated approaches using combinations of chemical, mechanical and biocontrol management programs have been developed to manage the spread of the species in its invaded range, however the species has still been able to persist. The persistence of the species was hypothesised to be a consequence of the co-occurrence of apomixis and polyploidy, however despite the identification of triploid and tetraploid cytotypes in South African populations of the pompom weed, the reproductive strategy of the species has not yet been determined. The aim of this study was therefore to infer whether populations of C. macrocephalum (pompom weed) reproduces via vector-mediated crosses, self-pollination or apomixis (either facultative or obligate) and examine the relationship of the mode of reproduction with ploidy level. Male fertility was also assessed to ascertain if interploidy gene flow was possible. The collated information was then used to infer the potential impact of reproductive strategies and polyploidy on biocontrol. All examined populations were shown to have high mean pollen viability percentages of 90% and 98% with no significant differences in pollen viability amongst the four populations. The high pollen viability percentages were supported by prolific pollen grain germination on the stigmatic surfaces (margins of style at base of style branches) and the sides of the style. This suggested that the pollen grains can fertilize and interploidy mating is likely possible in South African populations of the pompom weed. It is plausible that the high pollen viability is enabling triploids to act as a ‘triploid bridge’. However, the high pollen viability was confounded by the pollen tube analyses revealing that pollen tube growth is being arrested on the stigmatic surface suggesting that overall male fertility is low. The arrested pollen tube growth is typically associated with a ‘triploid block’. Nevertheless, the production of viable gametes can reduce the triploid block and facilitate gene flow between populations. The predominant mode of reproduction was determined by assessing the contribution of insects to pollen transfer, pollinator exclusion experiments, germination trials, pollen tube growth to the ovules and genetic analyses. We found that the African Monarch butterfly (Danaus chrysippus) and the honeybee (Apis mellifera) contributed most to pollen transfer in comparison to the other insects visiting C. macrocephalum. A pollinator exclusion experiment showed that the pompom weed can set seed in the absence of pollinators, albeit at lower quantities than in the open treatments. Nevertheless, germination percentages showed that reproductive success was similar between open and bagged treatments in each population. The Modderfontein population showed lower reproductive success and seedling establishment in comparison to the other populations, presumably due to the severity of the biocontrol infestation on this population. Genetic analyses revealed low genetic variation within and amongst populations. Pollen tube analyses showed no pollen tube growth to the ovules in all samples, which suggests that seed set is independent of fertilization. The lack of pollen tube growth is a strong indicator of obligate autonomous apomixis which is further corroborated by the low genetic differentiation between maternal plants and their respective offspring. The co-occurrence of apomixis and polyploidy made it difficult to discern which factor contributes more to the invasiveness of the species, however, we hypothesise that autonomous apomixis provides the pompom weed with the competitive advantage to persist in its invaded range. However, further studies on the reproductive strategies of tetraploid cytotypes are needed to confirm this hypothesis. The low genetic variation suggests that all populations should be equally susceptible to biocontrol agents, however that this may be affected by other factors such as environmental conditions or phenotypic plasticity. Phenotype plasticity refers to a single genotype producing different phenotypes in response to environmental conditions. This could reduce the efficacy of biocontrol agents as they may exhibit differential responses on different phenotypes