Ecological consequences of non-native fish invasion in Eastern Cape headwater streams
- Authors: Ellender, Bruce Robert
- Date: 2014
- Subjects: Fishery management -- South Africa -- Eastern Cape Fishes -- Conservation -- South Africa Introduced fishes -- South Africa -- Eastern Cape Introduced organisms
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/69065 , vital:29380
- Description: The introduction, spread and concomitant impacts of non-native species are a global problem. Fish are among the most widely introduced vertebrate groups, with their impacts affecting multiple levels of organisation- from individuals, populations and communities, to entire ecosystems. In South Africa, the largest perceived threat to range-restricted endemic headwater stream fishes is said to be invasion by non-native fishes, however, as is the case elsewhere, invasive impacts are often a case of risk perception rather than actual risk analysis. Two range-restricted headwater species, the Eastern Cape redfin Pseudobarbus afer and the Border barb Barbus trevelyani are redlisted by the International Union for the Conservation of Nature (IUCN) as ‘Endangered’, primarily due to invasion by non-native fishes. To investigate invasions in South Africa, and provide a quantitative estimate of the impact of non-native fishes on the two imperilled endemics, P. afer and B. trevelyani, the overall aims of this thesis were to: (A) Provide a literature review on non-native fish invasions in South Africa; (B) Using two case studies on the headwaters of the perennial Keiskamma and episodic Swartkops River systems, investigate the naturalisation-invasion continuum to provide a holistic view of the invasion process in these variable environments. The specific thesis objectives were: (1) Reviewing current knowledge of invasive impacts of non-native fishes in South Africa; (2) Investigating invasibility of headwater stream environments by non-native fishes; (3) Determining the establishment success of non-native fishes, (4) Assessing the spatial and temporal impacts of invasion; (5) Understanding mechanisms responsible for non-native fish impacts; (6) Investigating the threat of non-native fish invasion on the genetic diversity of two the two headwater fishes, P. afer and B. trevelyani. Results from the literature review of fish invasions (Chapter 1) showed that South Africa has a long history of non-native fish introductions, spanning two and a half centuries. Currently, 55 species have been introduced or translocated. Many of these introduced species have become fully invasive (36%). Their impacts also span multiple levels of biological organisation. There was a general paucity of studies on fish invasions (38 studies), however, of those conducted, reviewed studies placed emphases on invasive impacts (25 studies) and the transport, introduction, establishment and spread stages of the invasion process were largely ignored. The two study systems, the Swartkops and Keiskamma Rivers, were heavily invaded and numbers of introduced species surpassed that of natives (Chapter 2, 3 and 5). Headwater streams had varying invasibility and a number of non-native species were successfully established (Chapter 2, 3, 5 and 6). The remainder of the invasions were casual incursions into headwater streams from source populations in mainstream and impoundment environments which were invasion hotspots. Irrespective of establishment, four predatory invaders (largemouth bass Micropterus salmoides, smallmouth bass M. dolomieu, brown trout Salmo trutta and rainbow trout Oncorhynchus mykiss) impacted heavily on native fish communities (Chapter 3, 4 and 5). Two broad types of invasion were documented, top down invasion by non-native O. mykiss and S. trutta and upstream invasion by M. salmoides and M. dolomieu (Chapter 3 and 5). Their impacts included changes in community structure, extirpation from invaded stream reaches resulting in contracted distribution, and isolation and fragmentation of native fish populations. The impacts of non-native predatory fishes were particularly acute for P. afer and B. trevelyani. Where non-native predatory fish occurred, P. afer and B. trevelyani had been extirpated (Chapter 3 and 5). As a result both native species exhibited contracted distributions (>20% habitat loss due to invasion). Upstream invasion by centrarchids isolated and fragmented P. afer populations into headwater refugia, while top down invasion by salmonids excluded B. trevelyani from invaded, more pristine stream reaches, by forcing the species into degraded unsuitable lower stream reaches. Predation also disrupted population processes such as adult dispersal for P. afer, and centrarchid-invaded zones acted as demographic sinks, where adults dispersing through invaded reaches were rapidly depleted. While the Mandela lineage of P. afer exhibited little within or between drainage genetic structuring, B. trevelyani was >4% divergent between drainages, and up to 2% divergent between streams within the Keiskamma River system (Chapter 7). The distribution of genetic diversity for B. trevelyani also indicated that the loss of diversity was imminent without immediate conservation interventions. This thesis has provided conclusive evidence that native fishes are vulnerable to invasion and that non-native predatory fishes have significant impacts on native fishes in Eastern Cape headwater streams. If management and conservation measures are implemented, the unwanted introduction and spread of non-native fishes may be restricted, allowing native fishes opportunities for recovery.
- Full Text:
- Date Issued: 2014
Petrogenesis and metallogenesis of the Panzhihua Fe-Ti oxide ore-bearing mafic layered intrusion, SW China
- Authors: Howarth, Geoffrey Hamilton
- Date: 2013
- Subjects: Petrogenesis Metallogeny Intrusions (Geology) -- China -- Panzhihua Metallogenic provinces -- China -- Panzhihua Igneous rocks -- China -- Panzhihua Geochemistry Iron Titanium Oxides Ores Magmas
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4915 , http://hdl.handle.net/10962/d1001810
- Description: The Panzhihua intrusion is one of several large Fe-Ti oxide ore bearing intrusions related to the major flood volcanism of the Emeishan Large Igneous Province (ELIP), SW China. The Panzhihua intrusion in particular has recently become the focus of numerous studies owing to the excellent exposure in large open pit mining operations. The formation of Fe-Ti oxide ore layers has been the focus of these studies and has become a somewhat controversial topic with three separate models currently proposed for ore formation. The gabbroic Panzhihua intrusion extends for ± 19 km along strike, has a maximum thickness of 3000 m and hosts extensive (up to 60 m thick) Fe-Ti oxide ore layers in the lower portions of the intrusion. The intrusion has been divided into five zones: marginal zone (MGZ), lower zone (LZ), middle zone A (MZa), middle zone b (MZb) and the upper zone (UZ). The gabbroic rocks are comprised of plagioclase, clinopyroxene and interstitial Fe-Ti oxides with minor olivine. Apatite is present within the MZb only and shows no correlation with Fe-Ti oxide ore layers. Fe-Ti oxides are present throughout the stratigraphy of the intrusion. This is unlike typical layered intrusions where significant Fe-enrichment through fractionation of Fe-poor silicate phases (i.e. plagioclase) is required before Fe-Ti oxide saturation. There are no oxide-free cumulate rocks at the Panzhihua intrusion, implying either an evolved parent magma or very high Fe content of the source rocks. I present here new mineral composition data, whole-rock major and trace element geochemistry along with whole-rock Sr-Nd isotopes and PGEs in order to constrain the evolution of the Panzhihua parent magma en route from source to chamber and the formation of Fe-Ti oxide ore layers. Furthermore an initial pilot study using O-isotope data is conducted on Ti-magnetite and plagioclase separates from gabbroic vs. ore rocks. Results are coupled with detailed thermodynamic modeling using the software PELE in order to further constrain Fe-Ti oxide ore layer formation. The intrusion is characterised by extreme depletion of PGEs relative to the coeval flood basalts and picrites. High Cu/Pd and Pd/Pt imply two separate stages of S-saturated and S-undersaturated depletion of PGEs. Pd is highly compatible in sulphide and is quickly scavenged by sulphide liquids resulting in an increase in Cu/Pd of the residual liquid. Furthermore decoupling of Pd and Pt can be achieved by either late stage hydrothermal alteration or through S-undersaturated stage of PGE depletion where Pt is scavenged by Pt-rich alloys or oxide minerals. I show that the latter is more likely. Fractionation modeling suggests that the Panzhihua parent magma formed at depth from original picritic magma. This is consistent with several other recent studies on other layered intrusions of the ELIP. Sr-Nd isotopic ratios indicate very little crustal contamination has occurred en route to the current chamber. Sr and Nd concentrations of footwall rocks are too low to produce any significant change in initial Sr and Nd isotopic ratios of the intruding basaltic magmas, indicating that crustal contamination will not be indicated by Sr-Nd isotopic ratios. Gradational change in the Sr-Nd isotope ratios across the MGZ provides strong evidence for formation in an open system by multiple replenishments of progressively less contaminated magmas from depth. Contamination is difficult to constrain but must be occurring prior to emplacement at the current level (low Sr and Nd contents of footwall). A gradational upward decrease in highly incompatible element across the MGZ can then be explained by continuous magma flow, which effectively removes the evolved intercumulus liquids from the growing cumulate pile at the base of the chamber. The initial stages of formation of the Panzhihua intrusion are interpreted to result from prolonged low volume pulses of magma into a slowly opening chamber. The timing of Fe-Ti oxide crystallisation is fundamental in the understanding of the petrogenesis of ore layers. Distinct geochemical variation in whole-rock Fe2O3/TiO2 and Zr/Nb indicates that Timagnetite is the dominant oxide within the lower ± 270 m of the intrusion whereas above this level both Ti-magnetite and ilmenite are present as cumulus phases. This is interpreted to indicate a variation in the fO2 where the lower intrusion crystallises at higher fO2 relative to that above this level. Silicates within the ore layers, in particular plagioclase, are highly embayed and resorped where in contact with Fe-Ti oxides. This characteristic of the silicate grains implies early crystallisation prior to Fe-Ti oxides with subsequent disequilibrium conditions resulting in resorption. Furthermore distinct reaction rims of kaersutite amphibole, Fo-enriched olivine, An-enriched plagioclase and pleonaste are observed. The abundance of amphibole suggests H2O involvement in this reaction and consumption of silicates. A model for parent magma crystallisation at various H2O contents indicates that plagioclase crystallisation temperature is very sensitive to H2O content of the parent magma. Plagioclase crystallises early for “dry” compositions but significantly later for “wet” compositions. Fe-Ti oxide ore layers are generally well layered, contain gabbroic xenoliths and are observed raversing/cross-cutting the cumulate stratigraphy. I present here a new model for ore layer formation in order to account for these distinct features of the ore layers. A model invoking multiple replenishments of magma with variable oxide microphenocryst content, H2O content and volume is proposed. Magma evolving in the plumbing system and fed to the Panzhihua chamber is variably enriched in H2O, which results in significantly different crystallisation paths. High H2O magmas (> 2 wt %) crystallise Fe-Ti oxides early whereas low H2O magmas (< 1 wt %) crystallise oxides late. Early pulses of H2O-poor magma crystallise a sequence of plag+cpx+Fe-Ti oxide (±ol). Later pulses of H2Orich magma subsequently intrude the partially crystallised cumulate sequence incorporating and consuming previously crystallised silicates with subsequent early crystallisation of Ti-magnetite and formation of ore layers. H2O-rich magmas likely have suspended Ti-magnetite microphenocrysts as well, which crystallise at depth in the plumbing system. This model can account for the various characteristic features of the Fe-Ti oxide ore layers at the Panzhihua intrusion as well as other Fe-Ti oxide ore bearing intrusions in the region.
- Full Text:
- Date Issued: 2013
The pyrrolizidine alkaloids of Senecio chrysocoma and Senecio paniculatus
- Authors: Logie, Catherine Gwynedd
- Date: 1996
- Subjects: Senecio -- Analysis Pyrrolizidines Botanical chemistry
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4339 , http://hdl.handle.net/10962/d1005000
- Description: In order to compare the pyrrolizidine alkaloid content of two closely related species, Senecio chrysocoma and S. paniculatus, nine populations of plants distributed between the two species, were examined. Three novel pyrrolizidine alkaloids, 7ß-angelyl-l-methylene-8∝-pyrrolizidine, 7ß-angelyl-l-methylene-8∝-pyrrolizidine and 7ß-angelyl-l-methyleneSO!-pyrrolizidine-4-oxide, as well as eight known pyrrolizidine alkaloids, 7-angelylhastanecine, 9-angelylhastanecine, 7-angelylplatynecine, 9-angelylplatynecine, 9-angelylplatynecine-4-oxide, sarracine, neosarracine and retrorsine, were isolated and identified by NMR and GC-MS techniques. Traces of five tiglyl isomers, 9-tiglylplatynecine, 9-tigl ylplatynecine-4-oxide, 7ß-tiglyl-l-methylene-8∝-pyrrolizidine, sarranicine and neosarranicine, were also isolated and tentatively identified; however, these compounds could have been artefacts of the extraction and analytical procedures. While both species of plant investigated, S. chrysocoma and S. paniculatus, were found to be morphologically different, their pyrrolizidine alkaloid content was, in fact, very similar. The presence of retrorsine in S. paniculatus plant extracts, but not in those from S. chrysocoma plants, was the only major chemical difference observed. It is perhaps significant that retrorsine was the only macrocyclic pyrrolizidine to be identified. A comprehensive, computerised database of physical data for pyrrolizidine alkaloids has been compiled, which has facilitated the identification of new pyrrolizidines and the examination of trends in proton and carbon-13 NMR data for pyrrolizidine alkaloids. A stereospecific synthesis of 7ß-angelyl-l-methylene-8∝-pyrrolizidine was undertaken toconfirm the absolute stereochemistry of the product isolated from S. chrysocoma and S. paniculatus. An inseparable 5:2 mixture of 7ß-angelyl-l-methylene-8∝-pyrrolizidine and 7ß-angelyl-l,2-didehydro-l-methyl-8∝-pyrrolizidine, together with a small amount of tiglyl isomer, was finally synthesised. The application of various chiral differentiating chromatographic and spectroscopic techniques confirmed that both the natural and synthetic products had the same stereochemistry, permitting the natural alkaloid to be identified as 7ß-angelyl-methylene-8∝-pyrrol izidine
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- Date Issued: 1996
Synthesis, characterisation and biological activity of 2-(methylthiomethyl)anilines, 2-(methylthio)anilines, their Schiff-base derivatives and metal(II) (Co, Ni, Cu) complexes
- Authors: Olalekan, Temitope Elizabeth
- Date: 2013
- Subjects: Aniline , Schiff bases , Ligands , Nuclear magnetic resonance spectroscopy , Chelates , X-ray crystallography , Antimalarials
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:4562 , http://hdl.handle.net/10962/d1020868
- Description: A series of 31 sulfur-nitrogen donor ligands and 64 metal(II) complexes have been investigated. The thiomethylated aniline ligands 2–(methylthiomethyl)aniline 2MT and 2–(methylthio)aniline 2MA were synthesized with their substituted derivatives (-Me, -MeO, -Cl, -Br, -NO2) to serve as chelating agents. These ligands behave as bidentate ligands with SN donor group with Co(II), Ni(II) and Cu(II). The Co(II) and Ni(II) complexes have the ML2Cl2 molecular formula while the Cu(II) complexes formed with MLCl2 stoichiometry where L is the bidentate ligand. The ligands and their metal(II) complexes have been characterized by elemental analysis and with spectroscopic techniques. The trend observed in the NMR spectra and IR frequencies of the thiomethylated compounds shows there is a significant difference between the 2MT and 2MA series as a result of sulfur lone pairs extending the conjugation of the aromatic ring in the case of the latter. The effect of the position and electronic nature of ring substituent on the NMR shifts of the amine protons is discussed. The 6- and 5-membered chelate complexes formed by the 2MT and 2MA ligands respectively do not show significant diversity in their spectroscopic properties. From the elemental analysis for the Co(II) and Ni(II) complexes, their compositions reveal 1:2 M:L stoichiometry with 2 chlorine atoms from the respective metal salts. In addition, the spectroscopic data are largely indicative of tetragonally distorted structures for these solid complexes. The X-ray crystallography data reveal the Cu(II) complexes exist as square pyramidal dimers and with long Cu–Cl equitorial bonds fit into the tetragonally distorted octahedral structure. The electrolytic nature of Co(II) and Cu(II) complexes in DMF were found to be similar, they behave as non electrolytes in contrast to Ni(II) complexes which are 1:1 electrolytes. The electronic spectra of these metal(II) complexes were found to be different for both their solid forms and in solutions of DMF and DMSO and this has been discussed. The thiomethylated aniline ligands possess the amine and thioether groups which are present in many known biologically active compounds, hence the biological activity of the ligands and their metal complexes were tested against three strains of bacteria and one fungus. The methoxy-substituted derivatives were found to possess better inhibitory activity and this was similarly reflected in the metal(II) complexes. The activity of the complexes can be said to be in the order, Cu(II) > Co(II) > Ni(II). The Schiff-base derivatives were prepared from the ligands and para-methoxysalicylaldehyde and their Cu(II) complexes were synthesized in order to determine their biological activity. The Schiff-base ligands were found to be less active than their parent ligands. The Cu(II) complexes are not soluble in water, DMSO or DMF, as a result and could not be evaluated for their biological activity. Based on the good results from the antimicrobial evaluation, the antiplasmodial activity of some of the Co(II), Ni(II) and Cu(II) complexes of the thiomethylated ligands against Plasmodium falciparum (FCR-3) was determined. At 50 μM concentration level, the Cu(II) complexes show activity equal or better than the prophylactic chloroquine. The Cu(II) complexes with the methoxy-substituted demonstrated exceptional activity but their Co(II) and Ni(II) analogues did not show any activity. The cytotoxicity of the active Cu(II) complexes at 50 μM concentration was determined against the breast cancer cell line (MDA-MB-231). The compounds destroyed the cancer cell in the range of 28–40%, thus showing their preferred activity against the parasitic cell instead of the cancer cell. The selectivity demonstrated by these compounds have shown them to be potential antimalarial agents and this could be further investigated.
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- Date Issued: 2013
Understanding the reading practices of Fort Hare students
- Authors: O’Shea, Cathy
- Date: 2017
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/6886 , vital:21197
- Description: Universities world-wide are battling to offer access to far greater numbers than ever before. The University of Fort Hare, specifically, is also part of a troubled South African education system and is located in a disadvantaged, rural area. The main aim of this study was to understand Fort Hare students’ reading practices, as reported by the students themselves. This thesis used a framework of New Literacy Studies, which views student learning as a process of mastering discipline-specific, socially constructed norms and values, and sees the adopting of a literacy as including the adoption of an identity. Since discourse, in the NLS tradition, has been found to be a mediating mechanism in the social construction of identity, a critical discourse analysis was adopted to begin understanding aspects of Fort Hare students’ reading practices and the links between these and their identities. Critical realism is the ontological underpinning of this thesis. This means that the study aimed to identify the tendencies of certain mechanisms - in this case, Discourses - to affect students’ reading practices, by analysing interview transcripts of focus group discussions held with 30 students. Frameworks and tools provided by Fairclough and Gee were applied to interview data analysis. The ‘We blacks’ Discourse was one of one of the prominent Discourses that interviewees drew on when talking about their reading practices. It was closely allied to the ‘Resistance to reading’ Discourse, as participants explained that they tended to disregard books and did not enjoy reading for leisure. The ‘We blacks’ Discourse in this way homogenised class and other differences between black students, and indicated the ways in which their experiences were outside of academic Discourses. This Discourse served as a constraining mechanism for some, and indicated that those who used it tended not to identify with the academy. There was an evident link between the ‘We blacks’ Discourse, the ‘Resistance to reading’ Discourse and the ‘Better than us’ Discourse, in which students who enjoyed reading were called names for supposedly being conceited. Two opposing discourses (with a small ‘d’) emerged when students talked about literacy sponsors like parents and lecturers. Some used the ‘Our parents don’t chase us’ discourse to depict family members who were not encouraging, overlapping with the ‘We blacks’ Discourse. The contrasting ‘Go read anything’ discourse described more encouraging teachers and relatives. This discourse was also used to describe educators who had forced them to read, with several interviewees describing corporal punishment as being a necessary part of school-based literacy practices. It also became clear that Fort Hare’s institutional identity played a role in some interviewees’ self-identities, as the ‘Resistance to reading’ Discourse was linked to the ‘Why bother?’ Discourse. The latter seemed part of a defensive positionality that arose partly because some students see Fort Hare as a university with relatively low academic standards. However, the implication is that lecturers and others can work towards changing Discourses and so endeavour to enable reading practices. Educators could also take steps to address resistant attitudes and encourage reading.
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- Date Issued: 2017
The early proterozoic Makganyene glacial event in South Africa : its implication in sequence stratigraphy interpretations, paleoenvironmental conditions and iron and manganese ore deposition
- Authors: Polteau, Stéphane
- Date: 2005
- Subjects: Geology, Stratigraphic -- South Africa -- Northern Cape Geochemistry -- South Africa -- Northern Cape Paleogeography -- South Africa -- Northern Cape Petrology -- South Africa -- Northern Cape Ore deposits -- South Africa -- Northern Cape
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5039 , http://hdl.handle.net/10962/d1007612
- Description: The Makganyene Formation forms the base of the Postmasburg Group in the Transvaal Supergroup in the Griqualand West Basin. It consists of diamictites, sandstones, banded iron-formations (BIFs), shales, siltstones and carbonates. It is generally accepted that the Makganyene Formation rests on an erosive regional unconformity throughout the Northern Cape Province. However this study demonstrates that this stratigraphic relationship is not universal, and conformable contacts have been observed. One of the principal aims of this study is to identify the nature of the Makganyene basal contact throughout the Griqualand West Basin. Intensive fieldwork was carried out from Prieska in the south, to Danielskuil in the north. In the Sishen and Hotazel areas, only borehole material was available to assess the stratigraphy. The Griquatown Fault Zone delimits the boundary between the deep basin and platform facies. The Koegas Subgroup is only present south of the Griquatown Fault Zone, where it pinches out. However, the transition Griquatown BIFs-Koegas Subgroup occurs in lacustrine deposits on the Ghaap platform (Beukes, 1983). The Griquatown Fault Zone represents the edge of the basin, which corresponds to a hinge rather than a fault zone. The Makganyene Formation rests with a conformable contact on the Koegas Subgroup south of the Griquatown Hinge Zone, and north of it the Makganyene Formation lies unconformably on the Asbestos Hills Subgroup. The Makganyene Formation displays lateral facies changes that reflect the paleogeography of the Griqualand West Basin, and the development of ice sheets/shelves. The Ghaap platform is characterised by coarse immature sand interbedded with the diamictites. The clasts in this area contain local Asbestos Hills material and no dropstones are present. Such settings are typical of sediments that are being deposited below a grounded ice mass. At the Griquatown Hinge Zone, the sandstone lenses are smaller, and the clasts consist of chert, of which a great number are striated and faceted. In the Matsap area, the presence of dropstones is strong evidence for the presence of a floating ice shelf that released its material by basal melting. Further south, the Makganyene Formation contains stromatolitic bioherms that only form if clastic contamination is minimal and therefore the ice that transported the detritus to the basin did not extend far into open sea conditions. The base of the Hotazel Formation also contains diamictite levels. Dropstones have been identified, implying a glacial origin. The Hotazel diamictites are interbedded with hyaloclastites and BIFs. The Makganyene glacial event, therefore, was not restricted to the Makganyene Formation, but also included the Ongeluk Formation, through to the base of the Hotazel Formation. Petrographic studies of the Makganyene Formation and the base of the Hotazel Formation reveal mineral assemblages that are diagnostic of early to late diagenetic crystallisation and of low-grade metamorphism not exceeding the very low green-schist facies. The facies identified display the same sense of basin deepening, from shallow high-energy Hotazel area on the Ghaap platform, to the deep basin in the Matsap area. Whole-rock geochemical analyses reveal that the elemental composition of the Makganyene Formation is very similar to that of the Asbestos Hills BIFs, which were the most important source of clastic detritus for the Makganyene Formation. However, minor amounts of carbonates of the Campbellrand Subgroup, as well as a felsic crustal input from the Archean granitoid basement, made contributions. On the Ghaap platform, the Makganyene diamictite is enriched in iron, calcium, and magnesium, while in the deeper parts of the basin the diamictites are enriched in detrital elements, such as titanium and aluminium, which occur in the fine clay component. The Hotazel diamictite displays a distinct mafic volcanic input, related to the extrusion of the Ongeluk basaltic andesites, which was incorporated in the glacial sediments. Sequence stratigraphy is based on the recognition of contacts separating the different systems tracts that compose a depositional sequence. However, because the basal contact of the Makganyene Formation has not been properly identified in previous work, no correct model has been proposed so far. Therefore correlations between the Griqualand West and the Transvaal basins, based on lithostratigraphic similarities and extrapolations of unconformities, have to be reviewed, especially since the publication of new radiometric ages contradict all previously proposed correlations. It is proposed here that the Transvaal Supergroup in the Griqualand West Basin represents a continuous depositional event that lasted about 200 Ma. The Makganyene glacial event occurred during changing conditions in the chemistries of the atmosphere and ocean, and in the continental configuration. A Snowball Earth event has been proposed as the causative process of such paleoenvironmental changes. However, evidence presented here of less dramatic glacial conditions, with areas of ice-free waters, implies an alternative to the Snowball Earth event. The paleoenvironmental changes are thought to represent a transition from an anaerobic to aerobic atmosphere, that was responsible for the global cooling of the surface of the Earth, Such a glacial event may have aided in the large-scale precipitation of iron and manganese in areas of intense upwellings.
- Full Text:
- Date Issued: 2005
Phylogenetics of the keratin-feeding beetle family Trogidae (Coleoptera: Scarabaeoidea)
- Authors: Strümpher, Werner Petrus
- Date: 2016
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/1232 , vital:20037
- Description: Trogidae constitute a monophyletic and biologically unique family within the Scarabaeoidea, being the only keratinophagous group in its superfamily. Traditionally, the family has been divided into three distinctive genera, Omorgus Erichson, Polynoncus Burmeister, and Trox Fabricius. Although the taxonomy of the group is relatively well studied, recently changes to the existing classification have been proposed without the family as currently constituted being subjected to phylogenetic analyses. In this study I present the first molecular phylogeny for this cosmopolitan family, based on three partially sequenced gene regions: 16SrRNA, 18S rRNA and 28S rRNA domain 2. Analyses resolved relationships between and within genera and subgenera that are largely congruent with existing taxonomic hypotheses based on morphology. I recovered four well-supported radiations: Polynoncus, Omorgus, Holarctic Trox and Afrotropical Phoberus. One of the more surprising results was the monophyly of Phoberus. Based on the evidence, I propose reinstating the genus Phoberus to accommodate all Afrotropical (including Madagascan endemic) species. The results re-emphasise the validity of Scholtz’s (1986a) classification system for the genus Omorgusand its subgeneraand provide evidence of an African origin for Madagascan trogids. Divergence analysis was able to date the major events for the origin of extant lineages of Trogidae. The subsequent diversification of the major lineages is largely attributed to Pangaean and Gondwanan vicariance events in the Mid-Jurassic and early Late Cretaceous, respectively. The separation of the landmasses resulted in the evolution of four distinct groups: Omorgus, Polynoncus, Trox and Phoberus. The monophyletic subgenus Phoberus MacLeay is of particular interest; about one-third (14 of 40) of the species in the Afrotropical region have lost the ability to fly. To gain insights into the evolution of flightlessness in this clade, phylogenetic relationships among species were inferred using molecular sequence data. Four partial sequences of four genes (COI, 16S, 18S and 28S domain 2) were obtained for 27 species, covering all of the recognised species-groups. Estimated times of divergence were based on published mutation rates for COI. The results recovered eight monophyletic lineages, supporting the morphology-based subdivisions of the genus. Flightlessness evolved at least five times within the subgenus. There is no strong support for the re-acquisition of functional wings once they are lost. Diversification, and the present relictual distributions, in African Phoberus can be linked to climatic and geological events associated with the Miocene and Pliocene Climate Optimum in southern Africa. The evolution of flightlessness in Phoberus is consistent with the habitat stability hypothesis. All flightless species occupy refugial areas with relatively long-term environmental stability. Phoberus is a promising model group for studying evolutionary trends relating to flightlessness, speciation and biogeography. Flightlessness is a counterintuitive evolutionary development, given the many advantages of flight. To interpret the diversification in the flightless Phoberus capensis Scholtz, the phylogenetic relationships among several populations of P. capensis are investigated. Phylogenetic relationships among populations were inferred using molecular sequence data that suggested three distinct evolutionary lineages, which was also supported by morphological characters. Divergence time estimates suggest a Pliocene-Pleistocene diversification. Based on these results, it is suggested that P. capensis experienced climatically-driven allopatric speciation with sheltered Afrotemperate forests and high mountain peaks serving as important refugia in response to climatic exacerbations. The P. capensis complex thus represents a speciation process in which flight-restricted populations evolved in close allopatry, possibly as recently as the Pleistocene. Two of the three divergent and geographically distinct lineages are described as novel species. This study represents the most recent comprehensive work on the Trogidae. On the basis of this study, it is proposed that taxonomic changes to the generic classification of the family be made. Trogidae are formally divided into two subfamilies, Omorginae and Troginae. The subgenus Phoberus is restored to genus rank to include all the Afrotropical species, and Afromorgus is confirmed at subgeneric rank. The genus Madagatrox is synonymised with Phoberus.
- Full Text:
- Date Issued: 2016