Thesis (MSc)--University of Stellenbosch, 2006. / ENGLISH ABSTRACT: The present study aims to determine the phylogenetic relationships among the sand lizards, Pedioplanis. In addition, a single mitochondrial gene is used to investigate the geographic genetic structure of the widey distributed P. burchelli. With 11 species, Pedioplanis is the most speciose genus among the southern African genera of the family Lacertidae. All the species are restricted to the subcontinent with the exception of three (P. namaquensis, P. undata and P. benguellensis), which extend their range northwards into Angola. A total of 2200 nucleotide positions derived from two mitochondrial markers (ND2 and 16S rRNA) and one nuclear gene (RAG-1) are used to determine the phylogenetic relationships among ten of the eleven Pedioplanis species. The first well resolved gene tree for the genus, drawn from 100 individuals, is presented and this is largely congruent with a phylogeny derived from morphology. Contrary to some previous suggestions, Pedioplanis forms a monophyletic assemblage with Heliobolus and Nucras. The genus Pedioplanis is monophyletic with P. burchelli/P. laticeps forming a sister clade to all the remaining congeners. Two distinct geographic lineages can be identified within the widespread P. namaquensis; one occurs in Namibia, while the other occurs in South Africa. The “P. undata” species complex is monophyletic, but one of its constituent species, P. inornata, is paraphyletic. Relationships among the subspecies of P. lineoocellata are much more complex than previously documented. An isolated population previously assigned to P. l. pulchella is paraphyletic and sister to the three named subspecies. The phylogeny identifies two biogeographical groupings that probably diverged during the mid-Miocene. The development of the Benguella Current could have initiated isolation mechanisms associated with changes in habitat that could have generated barriers and played a role in the evolution of this group. At the lower taxonomic level, the mtDNA phylogeographic structure of the wide spread P. burchelli in South Africa reveal at least six distinct clades that are geographically partitioned. The first one is restricted to the eastern mountains along the Great Escarpment (GE). The next three are found along the Cape Fold Mountains (CFM): the north-west CFM, central CFM and eastern CFM. The fifth one shares samples from central CFM and GE. The last clade is restricted to the eastern central mountains of the GE. These six geographic groupings are genetically divergent from each other and they started separating in the early Pliocene period. Phylogeographic studies on other taxa in the region have found different levels of genetic structuring among or within taxa. The fact that P. burchelli is restricted to high altitude areas could have resulted in limited dispersal and consequently contributed to its geographic structure. However, the exact cause of the pattern obtained is not readily apparent. Habitat fragmentation in the past is probably one of the most influential factors shaping the genetic distribution of the species across South Africa. The inclusion of nuclear markers will shed more light on the evolutionary history of P. burchelli in South Africa. / AFRIKAANSE OPSOMMING: Die huidige studie stel ten doel om ‘n filogenie daar te stel vir die Sand akkedisse, Pedioplanis. ‘n Enkele mitochondriale geen is ook gebruik om die geografiese genetiese struktuur van die wydverspreide P. burchelli vas te stel. Met 11 spesies is Pedioplanis die mees spesieryke genus onder die suidelike Afrika genera wat aan die Lacertidae familie behoort. Al die spesies is beperk tot die subkontinent met die uitsondering van drie (P. namaquensis, P. undata en P. benguellensis), wat ‘n uitgebreide verspreiding het noordwaarts tot in Angola. ‘n Totaal van 2200 nukleotied posisies wat afkomstig is van twee mitochondriale merkers (ND2 en 16S rRNA) en een nukluêre geen (RAG-1) is gebruik om die filogenetiese verwantskappe tussen 10 van die 11 Pedioplanis spesies vas te stel. Die eerste goed geondersteunde geen boom vir die genus, gebasseer op 100 individue, is verkry en dit is meestal ooreenstemmend met ‘n filogenie gebasseer op morfologie. In teenstelling met sekere voorstelle van die verlede vorm Pedioplanis ‘n monofiletiese groep tesame met Heliobolus en Nucras. Die genus Pedioplanis is monofileties met P. burchelli/P. laticeps wat ‘n suster groep vorm van al die oorblywende lede van die genus. Twee herkenbare geografiese lyne kan geidentifiseer word in die wydverspreide P. namaquensis; een kom in Namibia voor, terwyl die ander een in Suid Afrika voorkom. Die “P. undata” spesies kompleks is monofileties, maar een van die spesies wat deel uitmaak van die groep, P. inornata, is parafileties. Verwantskappe tussen die subspesies van P. lineoocellata is meer kompleks as wat aanvanklik aanvaar is. ‘n Geisoleerde bevolkimg wat voorheen toegesê is aan P. l. pulchella is parafileties en verteenwoordig ‘n suster groep van die benaamde subspesies. Die filogenie identifiseer twee biogeografiese groeperings wat moontlik gedivergeer het gedurende die middel-Miocene. Die ontwikkeling van die Benguella stroom het dalk versperrings geinisiëer as gevolg van die gesamentlike veranderinge in habitat wat dalk ook ‘n rol gespeel het in die evolusie van die groep. Op die laer taksonomiese vlak het die mtDNA filogeografiese struktuur van die wydverspreide P. burchelli in Suid Afrika ten minste ses groepe aangetoon wat geografies van mekaar geskei is. Die eerste een is beperk tot die oostelike berge wat aan die Groot Eskarpement (GE) behoort. Die volgende drie word gevind in die Kaapse Vouberge (KVB): die noord-westelike KVB, sentrale KVB en oostelike KVB. Die vyfde een deel eksemplare van beide die GE en die KVB. Die laaste groep is beperk tot die oostelike en sentrale berge van die GE. Hierdie ses geografiese groepe is geneties geskei van mekaar en hulle het begin om apart te ontwikkel gedurende die vroë Pliocene periode. Ander filogeografiese studies in die area het verskillende vlakke van genetiese struktuur vertoon tussen en binne taksa. Die feit dat P. burchelli beperk is tot hoogliggende dele kon moontlik bygedrae het tot die geografiese struktuur. Die presiese oorsaak van die patroon wat verkry is, is nie ooglopend nie. Habitat fragmentasie in die verlede is moontlik een van die mees invloedrykste faktore wat die genetiese verspreiding van die spesie in Suid Afrika beinvloed het. Die insluiting van nukluêre merkers sal meer lig warp op die evolusionêre geskiedenis van P. burchelli in Suid Afrika.

Thesis (MSc)--University of Stellenbosch, 2006. / ENGLISH ABSTRACT: Marion Island is situated approximately 2600 km southeast of Cape Town and is the larger of the two islands that comprise the Prince Edward Island group. During past glaciation events, Marion was partially covered by ice with fauna and flora confined to isolated refugia across the island. As a result of these long-term isolation events, it is postulated that species predating these glaciation events might show geographic partitioning of genetic variation. Subsequently, the majority of Southern Oceanic islands have a history of sealing activities with vessels frequenting various islands in their hunt for seals. These combined visits to several islands could have facilitated the introduction of alien species, or the reintroduction of species already present on islands. These incidents, combined with more recent scientific voyages, have led to the establishment of several exotic species on Marion Island. Two indigenous Collembola species (Cryptopygus antarcticus (Willem, 1901) and Tullbergia bisetosa (Börner, 1903)) were chosen as model species to investigate the possible effects of glaciation on the spatial distribution of genetic variation on indigenous species. Given the negative impact that alien taxa have on indigenous species and ecosystems, the phylogeographic population structure of the recently introduced Isotomurus cf. palustris (Müller, 1876) was determined and compared to those described for the two indigenous species. To address these questions, two mitochondrial DNA genes were targeted: cytochrome oxidase subunit I (COI) and cytochrome oxidase subunit II (COII). These genes were sequenced (GENBANK accession numbers DQ147289-DQ147558) and analysed using, amongst others, AMOVA (Analysis of Molecular Variance), SAMOVA (Spatial Analysis of Molecular Variance) and NCA (Nested Clade Analysis). As expected, the two indigenous species showed distinct signs of population expansion, whilst the recently introduced species exhibits little genetic variance across its range. The life histories of species have an influence on their resultant genetic signature; therefore life history parameters were incorporated to explain differences in the phylogeographic patterns observed for these three species. / AFRIKAANSE OPSOMMING: Marion Eiland is die groter van twee eilande wat die Prins Edward eilandgroep vorm, ongeveer 2600 km suid-oos vanaf Kaapstad. Gedurende glasiasie episodes in die verlede, was Marion gedeeltelik met ys bedek en fauna en flora was geisoleer in klein gedeeltes van die eiland wat beskut was. As gevolg van hierdie langtermyn isolasie word daar voorgestel dat spesies wat op die eiland was voor hierdie gebeurtenisse plaasgevind het, ‘n geografiese verdeling van genetiese variasie sal toon. Sedertdien het die meerderheid van die eilande in die Suidelike Oseaan ook ‘n geskiedenis van skepe wat op een of meer van die eilande aangedoen het in die jag op robbe. Hierdie besoeke deur die robjagters aan die eilande het meegebring dat daar eksotiese spesies op die eiland aangekom het of dat spesies wat reeds op die eiland voorgekom het, opnuut daar geland het. Hierdie, tesame met die onlangse wetenskaplike reise na Marion Eiland, het daartoe gelei dat baie eksotiese spesies ingevoer is na die eiland. Twee inheemse spesies, naamlik Cryptopygus antarcticus (Willem, 1901) en Tullbergia bisetosa (Börner, 1903) is gekies om as model spesies te dien om die moontlike gevolge van glasiasie op die geografiese verspreiding van genetiese variasie te toets. In ag geneem die moontlike negatiewe impak wat eksotiese spesies op die inheemse spesies mag hê, is die filogeografie van ‘n uitheemse spesie, Isotomurus cf. palustris (Müller, 1876) ook bepaal en vergelyk met díe van die bogenoemde inheemse spesies. Dit is gedoen deur twee gene, sitokroom oksidase I (COI) en sitokroom oksidase II (COII) te analiseer. Hierdie gene se basispaarvolgordes is bepaal (GENBANK toetreenommers DQ147289-DQ147558), en hulle is ge-analiseer deur gebruik te maak van AMOVA (Analise van Molekulêre Variasie), SAMOVA (Ruimtelike Analises van Molekulêre Variasie) en NCA (geseteldegroepsanalises). Net soos verwag, het die twee inheemse spesies albei duidelike tekens getoon van ‘n bevolkingstoename terwyl die uitheemse spesie amper geen variasie oor sy verspreidingsgebied getoon het nie. Die lewensstyle van spesies het ‘n beduidende invloed op hulle genetiese bevolkingstruktuur. As sulks is díe in ag geneem om die verskille tussen hulle genetiese strukture te verduidelik.

Thesis (MSc (Botany and Zoology))--University of Stellenbosch, 2006. / The role of the southern harvester termite, Microhodotermes viator, and several climatic parameters in the distribution of the group-living lizard, Cordylus cataphractus, was investigated. Microhodotermes viator is considered the most important prey item of C. cataphractus and termitophagy as the causative agent in the evolution of group-living in this species. One would therefore expect a high degree of correspondence in the ranges of C. cataphractus and M. viator. As climate will also play a role in the distribution of any species, various climatic variables were investigated to determine their influence on the distribution of C. cataphractus. Species distributions were visualized using the minimum polygon technique and the degree of overlap was determined using standard geographic information systems (GIS) techniques. A total of 53 C. cataphractus localities were investigated for the presence of termites. The climatic limits of the geographical distribution of C. cataphractus were investigated by means of three models, namely Classification Trees, General Discriminant Analysis and Logistic Regression. The range of C. cataphractus was completely included within the range of M. viator Microhodotermes viator was included in the diet of C. cataphractus at 73 % of the localities sampled within the lizard’s range. The current geographical range of C. cataphractus is mainly correlated with two climatic factors, namely the low summer rainfall and high monthly solar radiation. The restricting role of both these factors can be directly linked to the group-living nature of C. cataphractus. If termitophagy were the overarching cause of group-living in C. cataphractus, then one would expect a close relationship between termite density and lizard density and termite density and lizard group size. I investigated these relationships at both a local and regional scale. For the local scale study, 25 quadrats of 25 × 25 m were plotted at a selected site, and for the regional scale study, ten 35 × 35 m quadrats at sites throughout the lizard’s range were used. In each quadrat, a range of variables were recorded, the most important of which were lizard density, lizard group sizes, termite foraging port density, distance to nearest termite foraging ports, vegetation height and vegetation cover. I found that the density of termite foraging ports determines C. cataphractus density. Vegetation height and cover affects crevice selection by C. cataphractus groups, probably because an unobstructed view is necessary to locate termite activity at foraging ports. I also investigated possible differences in the use of termites by different sized groups of C. cataphractus during different times of the year. Faecal samples, collected once a month at Eland’s Bay from small, medium and large groups from January 2005 to December 2005, were analysed for the presence of termite head material. I found that large groups fed on termites to a greater extent than small groups during certain times of the year and there was a general tendency for this phenomenon throughout the year. The results collected in this study indicate that the southern harvester termite, M. viator, plays a central role in the ecology of the group-living lizard, C. cataphractus.

Thesis (MSc)--Stellenbosch University, 2002. / ENGLISH ABSTRACT: Food demands for the ever-increasing human population is increasing the pressure on the agricultural sector to produce more food. In order to satisfy these demands, farmers are turning to chemical biocides for the control of pest species to produce greater crop yields. All pesticides must be toxic or poisonous to the target species they intend to control. Unfortunately, most pesticides are toxic or poisonous to non-target organisms as well, with detrimental effects on their health. Organic farming was developed to enhance the overall health of the farm's natural soilmicrobe- plant-animal biodiversity. No synthetic fertilisers and/or pesticides are used when farming organically. Life in the soil consists of intricate food webs and interactions between the soil dwelling invertebrates. The soil-organisms are divided into three main groups, viz., Micro-organisms (e.g. protozoa, bacteria and fungi) mesofauna (nematodes, Collembola and Acari) and macro-fauna (e.g. millipedes, isopods, insects, molluscs and earthworms). The invertebrates are very susceptible to chemical contamination by chemical biocides in natural and agro-ecosystems. The soil invertebrate communities are responsible for the decomposition of organic material in soil, thereby remineralising the soil. The decomposition processes start with comminution of the large pieces of organic material by meso- and macro-fauna and ends with the micro-fauna and microbial organisms that complete these processes by returning the nutrients in an inorganic form to the soil. The aim of this study was to investigate whether, and to what extent the soil organisms are influenced by different management practices viz., organic management practices versus conventional management practices. A vineyard on the farm Plaisir de Merle, in Simondium, Western Cape was used for the present study. One half of a one hectare vineyard was managed organically and the other half conventionally. Within each vineyard block six different treatments were performed. Three of the treatments were strictly organic and the other three were strictly conventional. Four replicates of each management treatment were performed. The bait-lamina technique was used to assess the feeding activity of the soil organisms exposed to the different management treatments. In addition to the bait-lamina trials in the vineyard itself, bait-lamina tests were performed in microcosm studies with soil from the organically and conventionally managed vineyard blocks under controlled conditions. In order to assess the impact of the various pesticides that are used in the vineyards in the conventional way, on the soil fauna, standard acute toxicity tests and behavioural tests were performed on Eisenia fetida, the compost worm. The bait-lamina tests in the vineyard revealed that the moisture content of the soil plays an important role in the biological activity of soil fauna. The different management treatments did affect the biological activity of the soil fauna, but seasonal changes also proved to be one of the important factors governing biological processes in the soil. The acute toxicity tests showed that the active ingredients (mancozeb, penconazole and trifloxystrobin) of three of the pesticides that were tested in this study, had negatively affected E. fetida at their recommended application concentrations. The remaining two pesticides' active ingredients (glyphosate and N-acetyl salicylic acid) did not affect the earthworms negatively at the recommended application concentrations. The preference behavioural trials showed that E. fetida could detect and avoid contaminated substrates at the LCso-concentrations of the different pesticides. All the earthworms were influenced positively in the preference behaviour experiments. Because of certain limitations of the bait-lamina technique, it was difficult to formulate conclusions on what happens in the soil. A possible explanation for the differences in feeding activity of soil fauna could be attributed to the migration of the soil fauna to more habitable soil horizons during the dry summer conditions, when most of the pesticides are applied. The ecological relevance of the acute toxicity tests conducted need to be investigated further. It is clear that the acute toxicity tests provided important information that should be considered, but care should be taken and the necessary safety factors be determined and considered when doing risk assessment studies. The results of the preference behaviour studies showed that for certain pesticides E. fetida can be a sensitive bioindicator of acute and/or sub-acute lethal toxicity testing but this might not necessarily be the case for other pesticides. The goal of doing laboratory studies is to gain as much information to make reliable extrapolations to field situations from laboratory data. Laboratory-to-field extrapolations are very complicated because of the physico-chemical composition of soil, the unpredictable way pesticides behave within soil and the reaction of soil organisms to the soil and to the chemical biocides that are used. Further studies need to be done in order to fully understand to what extent the soil fauna were affected by the different management practices applied to the vineyard at Plaisir de Merle. / AFRIKAANSE OPSOMMING: Voedsel aanvraag vir die toenemende menslike bevolking plaas groot druk op die landbou sektor om meer kos te produseer. Om aan die voedsel eise te voldoen, gebruik boere al meer chemiese biosiede om pesspesies te bestry. Alle pestisiede moet toksies of giftig wees vir die teiken organisme waarvoor dit bestem is. Ongelukkig is die meeste pestisiede ook toksies of giftig vir nie-teiken organismes, wat tot nadelige effekte op hul gesonheid kan lei. Organiese boerdery is ontwikkel om die algehele gesondheid van die plaas se natuurlike grond-mikrobe-plant-dier biodiversiteit te bevorder. Geen sintetiese bemestingstowwe en/of pestisiede mag gebruik word wanneer daar organies geboer word nie. Die lewe in die grond bestaan uit ingewikkelde voedselwebbe en interaksies tussen die grondlewende invertebrate. Die grond invertebrate word verdeel in drie hoof groepe, nl. mikro-organismes (bv. Protozoa, bakterieë en fungi) meso fauna (Nematoda, Collembole en Acari) en makrofauna (bv. Millipoda, Isopoda, Insecta, Mollusca en erdwurms). Die Invertebrata is die mees vatbaarste vir chemiese kontaminasie deur chemiese biosiedes in natuurlike en landbou ekosisteme. Die grond invertebraat gemeenskappe is verantwoordelik vir die afbreek van alle organiese materiaal in die grond en dus vir remineralisering van die grond. Die afbreekproses begin by die komminusie van groter stukke organiese materiaal deur die meso- en makrofauna en eindig met die mikrofauna en mikrobes wat die prosesse voltooi deur die nutriente terug te plaas in die vorm van anorganiese produkte in die grond. Die doel van hierdie studie was om te ondersoek of, en tot watter mate, grond organismes geraak word deur verskillende grondbestuurspraktyke, nl. Organiese grondbestuurspraktyke teenoor die konvensionele grondbestuurspraktyke. 'n Wingerd op die plaas Plaisir de Merle, in Simondium, Wes-Kaap, was gebruik vir die huidige studie. Een helfte van 'n een hektaar wingerd is organies bestuur en die ander helfte is op die konvensionele manier bestuur. Op elk van die twee wingerd blokke is ses verskillende behandelings toegepas. Drie van die behandelings was streng organies en die ander drie was streng konvensioneel van aard. Vier replikate van elke behandeling is toegepas op elk van die twee wingerdblokke. Die bait-lamina metode is gebruik om die voedingsaktiwiteit van die grond organismes te asses seer. As toevoeging tot die bait-lamina proewe in die wingerd self, is bait-lamina toetse ook in mikro-kosmosse in die laboratorium gedoen met grond afkomstig vanaf die twee wingerdblokke. Om die impak van die verskillende pestisiede op die grondorganismes te ondersoek, is standaard akute toksisteitstoestse en gedragstoetse uitgevoer met die komposerdwurm, Eisenia fetida. Die bait-lamina resultate in die wingerd het getoon dat die voginhoud van die grond die belangrikste rol speel wat die biologiese aktiwiteit van die grondorganismes beïnvloed. Die verskillende behandelings het die biologiese aktiwiteit van die grond fauna geaffekteer, maar seisoenale veranderings is ook uitgesonder as een van die bepalende faktore wat die biologiese prosesse in die grond stuur. Die akute toksisiteitstoetse het getoon dat die aktiewe bestandeie van drie van die pestisiede (mancozeb, penconazole en trifloxystrobin), E. fetida negatief beïnvloed het teen die aanbeveelde konsentrasies wat toegedien is. Die aktiewe bestandeie van die ander twee pestiede (glyphosate en N-asetiel sallisiel suur) het nie die erdwurms nadelig beïnvloed teen die aanbeveelde konsentrasies wat toegedien is nie. Die gedragsproewe het getoon dat E. fetida die LCso-konsentrasies van al die verskillende pestisiede kan waarneem en vermy. Al die erdwurms is positief beïnvloed in die gedragseksperimente met die verskillende pestisiede. Omdat die bait-laminametode sekere beperkings het, was dit moeilik om tot gevolgtrekkings te kom oor wat presies in die grond gebeur. 'n Moontlike verklaring vir die verskillende voedingsaktiwiteite van die grond fauna kan toegereken word aan die migrasie van die grondorganismes na meer leefbare grondhorisonne gedurende die droë somer toestande, wat toevallig met die spuit van die meeste pestisiede ooreenstem. Die ekologiese relevansie van die akute toksisiteitstoetse wat uitgevoer is, moet meer deeglik ondersoek word. Die belangrikheid van die akute toksisteitstoetse is duidelik en het waardevolle informasie gelewer, maar sorg moet geneem word, en die nodige veiligheids faktore moet bepaal word en in ag geneem word, wanneer riskobepalingstudies gedoen word. Die gedragsproewe het getoon dat vir sekere pestisiede E. fetida 'n sensitiewe bioindikator van akute en/of sub-akute letale toksisiteits toetse kan wees, maar nie noodwendig vir ander pestisiede nie. Die doel van laboratoriumstudies is om so veel as moontlik inligting te versamelom vertroubare ekstrapolasie te kan maak na situasies in die veld vanaf laboratorium data. Laboratorium-na-veld ekstrapolasies is dikwels baie gekompliseerd as gevolg van die fisies-chemiese samestelling van die grond, die onvoorspelbare manier waarop chemiese pestisiede met die grond reageer en die reaksie van die grondorganismes op chemiese biosiede in die grond. Verdere studies moet gedoen word om so deeglik maanlik die mate van die impak wat die verskillende bestuurspraktyke op die grond fauna het, te verstaan op Plaisir de Merle.

Thesis (MSc)--Stellenbosch University, 2003. / ENGLISH ABSTRACT: The geometric tortoise, Psammobates geometricus, found in the Western Cape, South Africa, is one of the world's rarest tortoises. At present, it is only known from five isolated geographical areas, one being the Worcester-Tulbagh Valley. The primary aims of my study were to determine the status of geometric tortoise subpopulations along the Nl highway in the Worcester Valley, to determine the population structure of these subpopulations and, if necessary, to provide recommendations for the mitigation ofNI highway-induced impacts on geometric tortoise subpopulations along the road. All Renosterveld remnants along the Nl between Du Toitskloof and Worcester were surveyed for the presence of geometric tortoises, and tortoise density and population structure were determined for each remnant that supports tortoises. Two geometric tortoise subpopulations on opposite sides of the Nl were identified. In the larger remnant of the two, females dominated both the subadult and adult classes, whereas in the other remnant male frequency was slightly higher than female frequency for the adult class only. In both remnants, adult females were significantly larger than adult males in mean carapace length. Mean body mass for adult females and males also differed significantly in both remnants. Analysis of habitat quality showed that both remnants supporting the geometric tortoise are seriously infested by encroaching indigenous and alien plants. The barrier effect of the Nl highway can be mitigated by constructing underpasses, linking the two remnants on opposite sides of the road. Furthermore, road mortality of tortoises can be avoided by erecting tortoise proof fencing along appropriate sections of the highway in the Worcester Valley. By linking the remnants, the largest single area available to the geometric tortoise in the Worcester Valley will also be formed. An additional aim of my study was to investigate the correspondence of geometric tortoise density in individual habitat remnants to the West-to-East change in vegetation composition/characteristics occurring along the Nl in the Worcester Valley. Vegetation composition/characteristics were thus determined in all remnants also surveyed for tortoises. The West-to-East change in vegetation composition/characteristics of the study area indicates that there is a transition from Renosterveld in the West to karroid veld in the East. / AFRIKAANSE OPSOMMING: Die geometriese skilpad, Psammobates geometrieus, wat in die Wes-Kaap, Suid- Afrika, voorkom, is een van die wêreld se skaarste skilpaaie. Vandag word hierdie skilpad net nog in vyf geïsoleerde geografiese streke gekry, een van hulle die Worcester-Tulbagh Vallei. Die primêre doel van my studie was om die status van geometriese skilpad subpopulasies langs die Nl snelweg in die Worcester Vallei te ondersoek en om die populasiestruktuur van hierdie subpopulasies te bepaal. Verder was die doelook, om aanbevelings te maak oor hoe die impakte van die pad op die geometriese skilpad subpopulasies langs die Nl verminder kan word. Al die Renosterveld oorblyfsels langs die Nl tussen Du Toitskloof en Worcester, en binne die bekende gebied van die geometriese skilpad, is deursoek vir geometriese skilpaaie. Populasiedigtheid en populasiestruktuur van die skilpaaie is bepaal vir elke oorblyfsel wat skilpaaie bevat. Twee geometriese skilpad subpopulasies is gekryaan teenoorgestelde kante van die Nl. In die groter oorblyfsel, het wyfies albei die onvolwasse en volwasse klasse gedomineer. In die ander oorblyfsel is gevind dat die frekwensie van mannetjies ietwat hoër was as die frekwensie van wyfies, maar vir slegs die volwasse kategorie. Binne albei oorblyfsels is gevind dat volwassse wyfies betekenisvol groter in gemiddelde doplengte is as volwasse mannetjies. Gemiddelde liggaamsgewig het ook betekenisvol verskil tussen volwasse wyfies en mannetjies in albei oorblyfsels. 'n Analise van habitatkwaliteit het gewys dat beide oorblyfsels wat die geometriese skilpad ondersteun grootliks ingeneem is deur inheemse indringers, sowel as uitheemse plante. Die versperrings effek van die Nl snelweg kan verminder word deur duikwege te skep wat die twee ooblyfsels, op teenoorgestelde kante van die pad, verbind. Bowendien, deur 'n ondeurdringbare heining langs passende dele van die snelweg in die Worcester Vallei op te rig, sal skilpadmortaliteit vermy word. Deur oorblyfsels te verbind, salook die grootste enkele area, beskibaar aan die geometriese skilpad in die Worcester Vallei, gevorm word. 'n Addisionele doel van hierdie studie was om veranderinge in vegetasie langs 'n wes-na-oos gradient binne die studie area te korreleer met die teenwoordigheid van skilpaaie langs hierdie gradient. Vegetasie samestelling/eienskappe is dus bepaal vir elke oorblyfsel wat deursoek is vir skilpaaie. Die wes-na-oos verandering in vegetasie samestelling/eienskappe binne die studie area wys dat daar 'n oorgang van Renosterveld in die weste na 'n karooagtige veld in die ooste is.

Thesis (MSc)--Stellenbosch University, 2013. / This study examined the biogeography and taxonomic status of the Cape legless skink, Acontias meleagris species complex using phylogenetic analyses, population genetics, demographic history aspects, time of lineage diversification estimation, environmental statistic analyses and a morphological evaluation. A total of 231 specimens from 55 localities were collected from the entire known distribution range of the A. meleagris complex throughout the Eastern, Northern and Western Cape, South Africa. Partial sequence data were collected from two mitochondrial DNA loci, 16S rRNA and cytochrome oxidase subunit one (COI), and one protein-coding nuclear DNA locus, exophilin 5 (EXPH 5). DNA sequences were analyzed for phylogenetic methods and biogeographical dating, while population genetic analyses were conducted on the COI sequences. Geographical boundaries amongst cryptic lineages were determined and evolutionary drivers of cladogenesis within the species complex were inferred. Marked genetic structure was observed within the A. meleagris complex, and five clades were retrieved, most of which were statistically well supported. These five clades were also evident within the haplotypic analyses and were characterized by demographic stability. Lineage diversification and the current biogeographical patterning observed for lineages within the A. meleagris species complex reflect the impact of sea level oscillations on historical coastal habitat availability. Additional historical evolutionary drivers within this subterranean species complex were inferred and discussed. The five clades within this species complex were considered discrete species, characterised by phylogenetic and biogeographic distinctiveness. While, morphological characters that could be used to identify the five species demonstrated widespread overlap for morphometric and meristic characters as well as colour pattering. Consequently, the phylogenetic species concept was employed for a taxonomical revision of A. meleagris sensu lato. Here, three of the previously recognised subspecies A. m. meleagris, A. m. orientalis and A. m. orientalis–'lineicauda' were elevated to full species, and two new species A. caurinus sp. nov. and A. parilis sp. nov. were described. / National Research Foundation (NRF)

Thesis (PhD (Botany and Zoology))--University of Stellenbosch, 2010. / ENGLISH ABSTRACT: The use of organophosphate pesticides is still an integral part of commercial farming activities and these substances have been implicated as a major source of environmental contamination in South Africa. Evidence exists that many non target animals in and around agricultural areas are at risk of being affected due to the mobile nature of pesticides and the intermittent nature of pesticide application. The extent to which non-target animals are affected by exposure to two organophosphates (azinphos-methyl and chlorpyrifos) was investigated through monitoring selected biomarker responses and life cycle effects under laboratory conditions in two selected test species. A representative species from both the aquatic and terrestrial environment was used as these two compartments of the environment are inevitably linked due to the mobility of pesticides from the area of application to surrounding areas. The earthworm Eisenia fetida was used as test organism in the terrestrial environment while the fish Oreochromis mossambicus served as representative of the aquatic environment. Juvenile life stages of both species were subjected to standard acute toxicity tests which showed that for both species, juvenile life stages were more sensitive to both pesticides than adults. It was also illustrated that azinphos-methyl is more toxic than chlorpyrifos to both species. Both test species were also subjected to an intermittent exposure regime in order to assess the effects of repeated pesticide application on biomarker, life-cycle and behaviour responses. The results indicated that for similar exposure regimes, azinphos-methyl was more toxic to E. fetida than chlorpyrifos and detrimentally affected all endpoints investigated. The present study suggests that exposure concentration may have a more pronounced effect in inducing a toxic response than exposure interval, irrespective of the pesticide used. In addition to this, E. fetida was unable to avoid the presence of these pesticides in soil, even at concentrations as high as 50% of the LC50 value, indicating that the presence of pesticides in the soil pose a realistic threat to earthworms and other soil dwelling organisms. Biomarker responses, morphological effects and feeding behaviour was assessed for O. mossambicus and similar to the terrestrial toxicity experiments, there was evidence to suggest that in the case of an intermittent exposure scenario, azinphos-methyl was more hazardous than chlorpyrifos to this species. For the majority of endpoints that were investigated, it appeared that exposure interval played a more important role in inducing an effect than exposure concentration. At a shorter exposure interval, the majority of endpoints showed no difference between higher and lower exposure concentrations, while at a longer exposure interval the effects of exposure concentration became evident. In addition, feeding behaviour was affected by pesticide exposure in a dose-dependent manner. The present study yielded important results that improve the understanding of biological impacts of pesticide pollution on the environment. This can aid in optimising farming practices such as pesticide application not only in terms of eradicating the pest organisms, but also in terms of mitigating the environmental effects associated with large-scale pesticide use, thereby ensuring sustained biodiversity in these areas. / AFRIKAANSE OPSOMMING: Die gebruik van organofosfaat plaagdoders is ‘n integrale deel van kommersiële landbou aktiwiteite maar hierdie middels is ook ‘n prominente bron van omgewingsbesoedeling in Suid-Afrika. Daar is bewys dat verskeie nie-teiken diere in en om landbouareas geaffekteer word weens die nie-statiese aard van plaagdoders in die omgewing, sowel as die herhalende aard van plaagdodertoediening. Die graad waartoe nie-teiken diere geaffekteer word deur die plaagdoders azinphos-metiel en chlorpyrifos is ondersoek deur die monitering van verskeie biomerkerresponse en lewenssiklus-effekte in geselekteerde toetsspesies binne ‘n beheerde laboratoriumomgewing. ‘n Verteenwoordigende spesie van beide die akwatiese en die terrestriële omgewing is gebruik aangesien hierdie twee dele van die omgewing onlosmaaklik verbind is weens die beweging van plaagdoders vanaf die area van toediening na omringende areas. Die erdwurm Eisenia fetida is gekies as toetsorganisme vir die terrestriële omgewing en die varswatervis Oreochromis mossambicus het gedien as verteenwoordigende spesie vir die akwatiese omgewing. Onvolwasse diere van beide spesies is onderwerp aan standaard akute toksisiteitstoetse en daar is gevind dat, vir beide spesies, onvolwasse diere meer sensitief vir die betrokke plaagdoders is as volwasse diere. Dit is ook gevind dat azinphosmetiel giftiger is as chlorpyrifos vir beide spesies. Beide toetsspesies is ook onderwerp aan ‘n chroniese blootstellingsregime om die effek van herhaalde plaagdodertoediening op biomerker-, lewenssiklus- en gedragsresponse te ondersoek. Die resultate van die herhaalde blootstelling het aangedui dat vir soortgelyke blootstellingsregimes, azinphos-metiel giftiger is as chlorpyrifos vir E. fetida en dat beide middels alle eindpunte wat ondersoek is, nadelig affekteer. Die huidige studie toon ook bewyse dat blootstellingskonsentrasie ‘n meer prominente effek as blootstellingsinterval kan hê in die teweegbringing van ‘n toksiese respons. Verder was E. fetida nie in staat om die teenwoordigheid van die plaagdoders in grond te vermy nie, self nie by konsentrasies so hoog as 50% van die LC50 waarde nie. Laasgenoemde resultaat dui dus aan dat die aanwesigheid van plaagdoders in die grondomgewing ‘n realisitese bedreiging inhou vir erdwurms en ander grondorganismes. Soortgelyk aan die terrestriële toksisiteitseksperimente, was daar getuienis vir die verhoogde toksisiteit van azinphos-metiel relatief tot chlorpyrifos vir O. mossambicus. Dit blyk dat blootstellingsinterval ‘n meer prominente rol as blootstellingskonsentrasie speel in die teweegbringing van effekte vir die meerderheid van die eindpunte wat ondersoek is. In die geval van ‘n korter blootstellingsinterval het die meerderheid van eindpunte wat ondersoek is geen verskille getoon tussen ‘n hoër en ‘n laer konsentrasie nie, terwyl met ‘n langer blootstellingsinterval daar ‘n aanduiding was dat blootstellingskonsentrasie ‘n meer prominente rol gespeel het. Verder is gevind dat voedingsgedrag in O. mossambicus geaffekteer is op ‘n konsentrasie verwante manier. Die huidige studie toon resultate wat ‘n belangrike bydrae kan lewer tot die begrip van die biologiese impakte van organofosfaat plaagdoders op die omgewing. Die resultate kan gebruik word vir die optimisering van boerderypraktyke soos plaagdodertoediening, sodat laasgenoemde effektief is vir die beheer van pes-organismes, maar ook die impakte van grootskaalse plaagdodertoediening kan minimaliseer en sodoende die biodiversiteit binne hierdie areas sal beskerm.

Thesis (PhD)--Stellenbosch University, 2011. / ENGLISH ABSTRACT: The terrestrial Oligochaete species Eisenia andrei and Enchytraeus doerjesi were exposed to different concentration series of Cd and Zn, both separately and in mixtures for 28 days in artificial OECD soil at 15, 20 and 25°C. At the end of the four week exposure period, survival and reproduction were assessed in E. doerjesi and survival, reproduction, biomass change, metal uptake and biomarker responses (MTT and comet assays) in E. andrei. Survival results for both E. andrei and E. doerjesi indicated that the lethality of Cd increased at higher temperatures, whereas the opposite was observed for Zn.Cadmium LC50 values were the highest at 15°C and the lowest at 25°C. In the Zn exposures LC50 increased with increasing temperature. Mixture results in both test organisms indicated that mixtures were less lethal than the metals separately. Effects of mixtures on survival, nonetheless, increased with increasing temperature. In the Cd experiment, reproduction in E. andrei was only recorded in the control treatments at the three temperatures investigated. However, the deleterious effect of Zn on the reproduction of both E. andrei and E. doerjesi decreased with increasing temperature and Zn EC50 for reproduction increased with increasing temperature. Results for exposures to mixtures indicated in both test organisms that the interaction between Cd and Zn were antagonistic. In both E. andrei and E. doerjesi, the effect of mixture exposures on reproduction decreased with increasing temperatures. The highest mixture EC50 values for reproduction were found at higher temperature. In E. doerjesi (using reproduction results in MixToxModules) Cd and Zn interactions were dose level dependent at the three temperatures investigated. Antagonism was the predominant interaction at lower mixture concentrations whereas synergism occurred at mixture concentrations equal to or higher than the mixtures’ EC50 values. Biomass loss increased with increasing temperature in the Cd exposures (p 9 0.05) but not in the Zn exposures in E. andrei. In this species mixture results indicated antagonistic interactions between Cd and Zn at all temperatures investigated. The deleterious effect of mixtures on the biomass of E. andrei increased with increasing temperature. When Cd and Zn interactions were further investigated in E. andrei (using biomass results in MixToxModules) it was found that they were dose level dependent at the three temperatures investigated. Antagonism was the predominant interaction at lower mixture concentrations whereas synergism occurred at mixture concentrations higher than the mixtures EC50 values. The assessment of metal uptake in E. andrei revealed a temperature dependent Cd uptake with higher Cd body burdens occurring at higher exposure concentrations and temperatures (p ≤ 0.05). In the case of Zn, although uptake was lower at higher temperature, there was no statistical difference in uptake between exposure concentrations and between temperatures. Mixture results however indicated that in mixture exposures less Cd was accumulated by E. andrei than in single Cd exposures (p ≤ 0.05). Inversely, in mixture exposures more Zn was accumulated by E. andrei than in single Zn exposures (p ≤ 0.05). Biomarker studies revealed that Cd and Zn were both cytotoxic and genotoxic whether in single or mixture exposures. Factorial ANOVA analyses of the effects of temperature and metals on the reduction of MTT by E. andrei indicated that temperature rather than the metals was the most important factor controlling mitochondrial activity (p < 0.001). In both Cd and Zn exposures significant deleterious metal effects on mitochondrial processes were found to increase with temperature (p ≤ 0.01). Mixture exposures indicated decreasing cytotoxicity with increasing temperature (p ≤ 0.05) and possible antagonism between Cd and Zn at cellular level. Results of the comet assay showed that the genotoxic profile of Cd was the opposite of the genotoxic profile of Zn. Cd was less genotoxic at lower temperature and increasingly deleterious at higher temperature while Zn was more genotoxic at lower than higher temperature (p ≤ 0.05). The results of mixture exposures indicated decreasing mixture genotoxicity with increasing temperature and suggested that the interactions between Cd and Zn at molecular level were probably antagonistic. / AFRIKAANSE OPSOMMING: Eisenia andrei en Enchytraeus doerjesi is aan verskillende konsentrasiereekse van Cd en Zn, afsonderlik en in mengsels, vir 28 dae in OECD kunsmatige grond onderskeidelik by 15, 20 en 25°C blootgestel Die volgende eindpunte is aan die einde van die vier weke blootstellingsperiode gemeet: oorlewing en voortplanting (by E. doerjesi) en oorlewing, voortplanting, biomassaverandering, metaalopname, MTT en komeettoetse (by E. andrei). Oorlewingsresultate by beide E. andrei en E. doerjesi het getoon dat toenemende temperatuur die letale toksisiteit van Cd laat toeneem terwyl die teenoorgestelde waar was vir Zn. By die Cd blootstellings was die LK50 waardes die hoogste by 15°C en die laagste by 25°C. By die blootstellings aan Zn het die LK50 waardes toegeneem by hoër temperatuur. Resultate by die mengsels by beide toestspesies het aangetoon dat die mengsels minder letaal was as die afsonderlike metale. Effekte van mengsels op oorlewing het nietemin toegeneem met toenemende temperatuur. By die Cd blootstellings is voortplanting slegs by die kontroles en in die geval van E. andrei by die drie onderskeie temperature ondersoek. Die nadelige uitwerking van Zn op voortplanting by beide E. andrei en E. doerjesi het afgeneem met stygende temperatuur en die Zn LK50 vir voortplanting het toegeneem met toenemende temperatuur. Resultate van die blootstellings aan mengsels het getoon dat die wisselwerking tussen Cd en Zn by beide spesies antagonisties was. By beide spesies het die invloed van die mengsels op voortplanting afgeneem met stygende temperatuur. Die hoogste mengsel LK50 waardes vir voortplanting is by hoër temperature gevind. By E. doerjesi was Cd en Zn wisselwerkings by blootstelling aan mengsels (voortplantingsresultate ondersoek deur van MixToxModules gebruik te maak) dosisvlak verwant by die drie temperature wat ondersoek is. Antagonisme was die oorwegende wisselwerking by laer mengsel konsentrasies terwyl sinergisme voorgekom het by mengsel konsentrasies gelyk aan of hoër as die LK50 waardes van die mengsels. In die geval van die Cd blootstellings by E. andrei het biomassaverlies toegeneem met toenemende temperatuur (p ≤ 0.05) maar nie by die Zn blootstellings nie. Resultate van blootstellings aan mengsels het getoon dat die uitwerking van mengsels op die biomassa van E. fetida toegeneem het met toenemende temperatuur. By die verdere ondersoek van Cd en Zn wisselwerkings, waar gekyk is na dosis verhouding of dosisvlak antagonisme (deur van biomassa resultate in MixToxModules gebruik te maak), is gevind dat Cd en Zn wisselwerkings dosisvlak afhanklik was by die drie temperature wat ondersoek is. Antagonisme was die oorwegende wisselwerking by laer mengselkonsentrasies terwyl sinergisme voorgekom het by mengselkonsentrasies hoër as die mengsel EK50 konsentrasies. Die bepaling van metaalopname deur E. andrei het ’n temperatuurafhanklike opname van Cd getoon met hoër Cd liggaamskonsentrasies by hoër blootstellingskonsentrasies en temperature (p ≤ 0.05). Alhoewel Zn opname laer was by hoër temperatuur was daar geen statisties betekenisvolle verskille in opname tussen blootstellingskonsentrasies of temperature nie. Die bepaling van metaalopname by wurms wat aan mengsels blootgestel is, het getoon dat minder Cd deur E. andrei opgeneem is as waneer die wurms aan Cd as enkelmetaal blootgestel is (p ≤ 0.05), Daarteenoor het die teenoorgestelde gebeur in die geval van Zn, Meer van die metaal is opgeneem wanneer E. andrei aan mengsels blootgestel is as aan die enkelmetaal. Biomerkerstudie het getoon dat Cd en Zn beide sito- en genotoksies kan wees ongeag of dit as enkelmetale of in mengsels toegedien is. Faktoriale ANOVA analises van die effekte van temperatuur en metale op die verlaging van MTT by E. andrei het getoon dat temperatuur ’n belangriker faktor was as metaalbesoedeling by die kontrole van mitochondriale aktiwiteit. (p ≤ 0.001). By beide Cd en Zn blootstellings was daar in elk geval statisties betekenisvolle metaaleffekte op mitochondriale prosesse met toename in temperatuur (P ≤ 0.01). By blootstellings aan mengsels is gevind dat sitotoksisiteit afgeneem het met toenemende temperatuur (p ≤ 0.05) asook ’n moontlike antagonisme tussen Cd en Zn op sellulêre vlak. Resultate van die komeettoets het getoon dat die genotoksiese profiel van Cd die teenoorgestelde was as die van Zn. Cd was minder genotoksies by laer temperature en meer en meer skadelik by hoër temperature terwyl Zn meer genotoksies was by laer as by hoër temperature (p ≤ 0.05). Die resultate van blootstlling aan mengsels het laer genotoksisiteit getoon met toename in temperatuur. Dit dui daarop dat wisselwerkings tussen Cd en Zn op molekulêre vlak moontlik antagonisties was.

Thesis (PhD)-- Stellenbosch University, 2012. / ENGLISH ABSTRACT: Climate change and biological invasions are major threats to biodiversity. In particular, these threats are predicted to influence terrestrial systems in the sub-Antarctic, where significant ecosystem responses to both have already been seen. In this thesis, the sub-Antarctic Prince Edward Island group is used as a model system in which to investigate key questions relating to climate change and invasive species impacts. The island group comprises two islands, Marion (MI) and Prince Edward (PEI), both of which are experiencing rapid warming, yet have different invasive assemblages and in consequence are experiencing different impacts. Variation in the patterns of invasive species richness and abundance and their underlying causes are matters of considerable ecological and conservation significance. While an increase in thermal energy availability typically results in an increase in species richness, the mechanisms underlying these patterns are poorly understood. In Chapter 2 of this thesis, these relationships are explored for springtails, an important component of the soil fauna on Marion Island. Energy explains a large amount of the spatial variation in indigenous and invasive springtail species richness. Disturbance thresholds and stressful temperatures are more important than increased population sizes in determining this variation in species richness. As both indigenous and invasive springtail species richness and abundance are strongly related to temperature, a warming climate could have far-reaching consequences for these organisms. In particular, invasive species are predicted to be at an advantage relative to indigenous species under warming conditions. One species where this seems especially likely, given its physiological responses to experimental warming and drying, is the large invasive tomocerid, Pogonognathellus flavescens. Determining whether this will be the case depends on understanding the factors underlying its range limits and abundance structure. Moreover, few studies have sought to distinguish the causal basis of abundance structure and range limits, particularly for invasive species. Thus, in Chapter 3, local microclimate variables and physiological tolerances of the invasive springtail, P. flavescens (a habitat generalist), are examined. The results suggest that the species should be widely distributed across a range of habitats on MI. However, the springtail is restricted to indigenous Poa cookii tussock grassland habitats in the southeast. The current range limits are set by dispersal limitation (i.e. contingent absences) whilst abundance structure is a function of variation in soil substrate quality. However, over time, the widening distribution of P. cookii, as a consequence of a major management intervention (the eradication of feral cats), may enable P. flavescens to colonise all suitable areas. In Chapter 4, the focus changes to what has been considered the third major response to climate change, along with range and phenological responses - changing animal body sizes. Body size is one of the most significant and obvious features of animals and is of considerable ecological and physiological importance. A prediction of the temperature-size rule (TSR) is that with warming, body size of the weevil species on both MI and PEI should decline. However, predation by mice of the weevils on MI should fundamentally affect the pattern of such change, causing it to differ from neighbouring PEI, indicating synergistic impacts between climate change and invasions. Analysis of a 24-year data set indicates a decline in the body size of all weevil species on PEI with increasing temperature. However, on MI, a negative relationship between mean annual temperature and body size is found only for Palirhoeus eatoni, a species not eaten by mice. A possible explanation for the positive relationships found for the other species could be due to higher metabolic demands imposed on mice in colder years than in warmer ones. Any increase in predation coupled with a preference for larger sizes, which the mice clearly show, would lead to a decline in the mean size of the weevil species. Due to the relationship between body size and metabolic rate and the importance of the weevils in the islands’ food webs, changes to the body size of these organisms could have significant consequences for the island ecosystems’ functioning. The thermal environment experienced by organisms also has a direct effect on survival, growth and reproduction. The physiological response of organisms to rapidly changing climates is therefore a primary concern. Organisms may respond to variable environmental conditions through phenotypic plasticity as well as behaviour. Chapter 5 of this thesis shows that of the weevil species and populations investigated on MI, most display phenotypic plasticity, the form of which is in keeping with the ‘Hotter is Better’ hypothesis. This could be due to rare extreme temperature events and the advantage for the performance curves to incorporate high temperatures experienced in the environment. Mismatches between thermal optima and preferred temperatures displayed by all species could mean that these weevils are well equipped to cope with warming conditions on MI unless the prediction of an increase of rare extreme events such as extreme temperatures is realised. Rapidly changing climates and an increase in the introduction of non-indigenous species are issues of major conservation concern. This has increased the significance of studies on the impacts of these threats. However, this thesis shows that to understand such processes, it is essential that an integration of disciplines be undertaken. This thesis thus adopts a multidisciplinary approach and highlights key issues associated with both climate change and biological invasions. The patterns and predictions of species and community responses to these environmental changes are complex. Moreover, predicting such responses is likely to be problematic, especially as multiple factors will change concurrently and how these factors might change is unclear. This highlights the importance of long-term records for understanding organism responses to such changes. Furthermore, impacts on indigenous species are likely to be exacerbated by the predicted increase in the rate of introductions with climate change. This makes the case for preventing the dispersal of invasive species to new areas all the more important. / AFRIKAANSE OPSOMMING: Klimaatsverandering en indringer spesies is belangrike bedreigings vir biodiversiteit. In besonder word voorspel dat hierdie bedreigings terrestriële sisteme in die sub-Antarktiese sal beïnvloed, waar beduidende ekosisteem reaksie aan beide reeds gesien is. In hierdie tesis word die sub-Antarktiese Prince Edward eiland groep as a model sisteem gebruik om belangrike vrae met betrekking tot klimaatsverandering en die impak van indringer spesies te ondersoek. Die eiland groep bestaan uit twee eilande, Marion (ME) en Prince Edward (PEE), wat beide versnellende verwarming ervaar, maar tog verskillende indringer samestellings het en vervolgens verskillende impakte ervaar. Variasie in die patrone van indringerspesierykheid en vollopheid en hulle onderliggende oorsake is van aansienlike omgewings en bewarings betekenis. Terwyl 'n toename in die beskikbaarheid van energie tipies lei tot 'n toename in spesierykheid, word die onderliggend meganismes van hierdie patrone swak verstaan. In Hoofstuk 2 van hierdie tesis, word hierdie verhoudings vir springsterte ondersoek, 'n belangrike komponent van die grond fauna op ME. Energie verduidelik 'n groot hoeveelheid van die ruimtelike variasie in inheemse en indringende springstert spesierykheid. Versteuringsdrempels en stressvolle temperature is meer belangrik as die toename in bevolking groottes in die bepaling van hierdie variasie in spesierykheid. Aangesien beide inheemse en indringende springstert spesierykheid en vollopheid sterk verwant is aan temperatuur, kan 'n verwarmende klimaat verreikende gevolge vir hierdie organismes hê. In die besonder word voorspel dat indringerspesies bevoordeeld sal wees relatief tot inheemse spesies onder verwarmende toestande. Een spesie waar dit veral blyk om geneig te wees, gegewe sy fisiologiese reaksie tot eksperimentele verhitting en uitdroging, is die groot indringer tomocerid, Pogonognathellus flavescens. Om te bepaal of dit die geval sal wees, hang af van die begrip van die onderliggende faktore van sy voorkomsgrense en vollopheidstruktuur. Daarbenewens is daar min studies wat gepoog het om te onderskei tussen die veroorsakende basis van vollopheidstruktuur en voorkomsgrense, veral vir indringerspesies. Dus, in Hoofstuk 3, word plaaslike mikroklimaat veranderlikes en fisiologiese toleransies van die indringer springstert, P. flavescens ('n habitat generalis), ondersoek. Die resultate stel voor dat die spesie wyd verspreid moet wees oor 'n verskeidenheid van habitatte op ME. Maar, die springstert is beperk tot inheemse Poa cookii polle grasveldhabitatte in die suidooste. Die huidige voorkomsgrense word daar gestel deur verspreidingsbeperking (dws voorwaardelike afwesighede), terwyl vollopheidstruktuur 'n funksie is van die variasie in die grond substraat kwaliteit. Maar, die uitbreidende verspreiding van P. cookii as gevolg van 'n groot bestuursingryping (die uitwissing van wilde huiskatte), kan P. flavescens in staat stel om alle geskikte gebiede te koloniseer met verloop van tyd. In Hoofstuk 4 verander die fokus na wat as die derde groot reaksie op klimaatsverandering beskou word, saam met voorkoms en fenologiese reaksies - veranderende diere liggaamsgroottes. Liggaamsgrootte is een van die beduidendste en mees voor die hand liggende eienskappe van diere en is van aansienlike ekologiese en fisiologiese belang. 'n Voorspelling van die temperatuur-grootte-reël (TGR) is dat met verwarming, liggaamsgrootte van die snuitkewerspesies op beide ME en PEE sal afneem. Hoe ookal, predasie deur muise van die snuitkewers op ME moet fundamenteel die patroon van sodanige verandering op PEE beïnvloed, wat sinergistiese impakte tussen klimaatsverandering en indringings aandui. Die ontleding van 'n 24-jarige datastel dui aan op 'n afname in die liggaamsgrootte van alle snuitkewer spesies op PEI met ‘n toename in temperatuur. Maar, op ME is 'n negatiewe verhouding tussen die gemiddelde jaarlikse temperatuur en liggaamsgrootte net gevind vir Palirhoeus eatoni, 'n spesie wat nie deur die muise geëet word nie. 'n Moontlike verduideliking vir hierdie positiewe verhoudings wat gevind is vir die ander spesies kan wees as gevolg van hoër metaboliese eise op die muise in kouer jare as in warmer jare. Enige toename in predasie, tesame met 'n voorkeur vir groter mates, wat die muise duidelik wys, sou lei tot 'n afname in die gemiddelde grootte van die snuitkewer spesies. As gevolg van die verhouding tussen liggaamsgrootte en metaboliese tempo, sowel as die belangrikheid van die snuitkewers in die eilande se voedselwebbe, kan veranderinge in die liggaamsgrootte van hierdie organismes beduidende gevolge op die eiland ekosisteme se funksionering hê. Die termiese omgewing wat deur organismes ervaar word het ook 'n direkte invloed op oorlewing, groei en voortplanting. Die fisiologiese reaksie van organismes op vinnig veranderende klimate is dus 'n primêre bron van kommer. Organismes kan reageer op veranderlike omgewingstoestande deur fenotipiese plastisiteit sowel as gedrag. Hoofstuk 5 van hierdie tesis toon dat van die snuitkewerspesies en bevolkings wat ondersoek is op ME, die meeste fenotipiese plastisiteit vertoon, die vorm wat in ooreenstemming is met die ‘Warmer is Beter’ hipotese. Dit kan wees as gevolg van seldsame uiterste temperatuur gebeure en die voordeel vir die prestasie kurwes om hoë temperature wat ervaar word in die omgewing in te sluit. Mismatches tussen termiese optima en voorkeur temperature vertoon deur alle spesies kan beteken dat hierdie snuitkewers goed toegerus is om die verhitting op ME te hanteer, tensy die voorspelling van 'n toename van seldsame uiterste gebeure soos uiterste temperature gerealiseer word. Vinnig veranderende klimate en 'n toename in die bekendstelling van nie-inheemse spesies is kwessies van groot bewarings kommer. Dit het die betekenis van studies oor die impak van hierdie bedreigings verhoog. Hierdie tesis toon egter dat om sulke prosesse te verstaan, dit noodsaaklik is dat 'n integrasie van die dissiplines onderneem word. Hierdie tesis aanvaar dus 'n multi-dissiplinêre benadering en beklemtoon die belangrike kwessies wat verband hou met beide klimaatverandering en biologiese indringing. Die patrone en voorspellings van spesies en die gemeenskapsreaksies op hierdie omgewingsveranderinge is kompleks. Verder, die voorspelling van sodanige reaksies sal waarskynlik problematies wees, veral omdat verskeie faktore gelyktydig sal verander en hoe hierdie faktore kan verander is onduidelik. Dit beklemtoon die belangrikheid van lang termyn rekords vir die begrip van organisme reaksies op sulke veranderinge. Verder, die impak van inheemse spesies is geneig om te vererger deur die voorspelde toename in die tempo van bekendstellings met klimaatsverandering. Dit maak die taak vir die bestuur van die voorkoming dat indringerspesies nuwe gebiede bereik al hoe meer belangrik.

Dissertation (PhD)--University of Stellenbosch, 2007. / ENGLISH ABSTRACT: The taxonomic status of southern Africa’s rupicolous crag lizards (genus Pseudocordylus) was investigated. As considerable confusion exists in the literature regarding the type specimens and type localities of the various taxa, resolution of these problems were considered the starting point of the study. Examination of museum specimens allowed for the designation of lectotypes, alloparalectotypes and/or paralectotypes. Of particular relevance to this study was the rediscovery of Andrew Smith’s type specimens of P. m. melanotus and P. m. subviridis. Restriction of the type locality of P. m. subviridis, based on entries in Smith’s diary and journal, allowed for the confirmation of previous interpretations and definitions of the two taxa. The geographical distribution of the various taxa and populations was determined using an extensive locality database. Two kinds of molecular markers, namely allozymes and mitochondrial DNA, were used in an attempt to resolve taxon boundaries within the P. melanotus species complex. The allozyme analysis indicated that P. m. melanotus might be polyphyletic and comprised of two unrelated lineages. Furthermore, fixed allelic differences between parapatric populations of P. m. melanotus and P. m. subviridis, and between sympatric populations of P. m. subviridis and P. langi, suggested that all three forms might be considered full species, with the possibility of more cryptic species present in the complex. Pseudocordylus transvaalensis differed from most other populations by 1-3 fixed allelic differences, but was indistinguishable from the Nkandhla district (central KwaZulu-Natal) population of P. m. melanotus. There were no heterozygous individuals in a sample from Monontsha Pass (Qwa-Qwa), a population reportedly comprising P. m. melanotus and P. m. subviridis, as well as intermediates, and all specimens were assignable to P. m. subviridis. The allozyme study was, however, based on phenetic principles and for further taxonomic resolution a cladistic approach was required. An mtDNA analysis (16S rRNA gene) using Maximum Parsimony, Maximum Likelihood and Bayesian analyses was therefore conducted to determine phylogenetic relationships among species and subspecies and to re-assess the taxonomic status of forms in the P. melanotus species complex. The mtDNA analysis corroborated most of the results obtained in the allozyme analysis. Firstly, P. langi was again found to be basal. With the addition of P. microlepidotus and P. spinosus to the ingroup, it is now apparent that P. langi is the basal species in the genus. (Recent studies have indicated that P. capensis and P. nebulosus are not congeneric with Pseudocordylus.) Secondly, the 16S rRNA results confirm that P. m. melanotus, as presently construed, is comprised of two clades that are not sister groups. The northern populations of P. m. melanotus (Sabie and Lochiel) form a fairly deeply divergent clade that may represent a separate species. The Nkandla population was, however, found to cluster with the other southern P. m. melanotus populations and not with P. transvaalensis as was the case in the allozyme electrophoretic analysis. However, the most surprising result of the 16S rRNA analysis was the finding that both P. microlepidotus and P. spinosus are embedded within P. m. subviridis. This suggests that these two species evolved from within P. m. subviridis and may have been separated only recently, with rapid morphological divergence occurring, but with limited genetic differentiation. It is suggested that all of the above three taxa be provisionally treated as full species. There was also morphological support for the uniqueness of all groupings indicated by the mtDNA analysis. Pseudocordylus transvaalensis is characterized by its large size, unique dorsal and gular (black) colour patterns, as many as three horizontal rows of lateral temporal scales, a series of small scales posterior to the interparietal scale, and usually two subocular scales behind the median subocular on either side of the head. The various populations currently classified under the name P. melanotus are more difficult to separate, but P. m. melanotus and P. m. subviridis usually differ as follows: frontonasal divided in P. m. melanotus, undivided in P. m. subviridis (and most Northern melanotus); lateral temporals in two rows, upper more elongate versus single row of much elongated scales; longitudinal rows of dorsolaterals closely-set versus widely separated; femoral pores of females pit-like versus deep with secretory plug. Northern melanotus differs from Southern melanotus in usually having an undivided frontonasal scale and seldom having a small scale present behind the frontonasal. Pseudocordylus langi has unique dorsal and gular colour patterns (including a series of blue spots on the flanks), granular dorsals with 6-9 paravertebral rows of enlarged flat scales, high total numbers of femoral pores (25-34) and usually only five (smooth not keeled or ridged) infralabial scales on either side of the head. Pseudocordylus spinosus also has unique dorsal and gular colour patterns, spinose lateral scales, frontonasal longer than wide and excluded from the loreal scales, low total femoral pore counts (6-9), and females (not only males) have differentiated femoral scales. Both Principal Components Analysis (PCA) and Canonical Discriminant Analysis (CDA) distinguished four groups, namely P. transvaalensis, P. langi, P. spinosus and a P. melanotus/subviridis/microlepidotus cluster. A separate CDA of all P. melanotus populations partly distinguished between Southern melanotus and P. m. subviridis, and largely separated Northern melanotus; whereas a CDA of P. transvaalensis showed that all three allopatric populations are 100% distinguishable in morphological space. A Nested Clade Analysis indicated that fragmentation as well as range expansion played a role in the distribution of the P. melanotus species complex. This may be explained by climatic oscillations (high-low temperatures and wet-dry cycles) during the Cenozoic that caused habitat expansion and contraction. Based on the topology of the mtDNA phylogram it is apparent that the genus Pseudocordylus originated along the eastern escarpment. A P. langi-like ancestor may have had an extensive range along the eastern escarpment, with the Maloti-Drakensberg forming the southern limit of its range. During a subsequent rise in global temperatures, range contraction and fragmentation took place, leaving an isolated population in the south and one in the north. The southern population survived unchanged in the Maloti-Drakensberg refugium, but the northern population was forced to adapt to the warmer conditions. Thereafter, the northern form expanded its range again, but during a subsequent cooler period, range contraction occurred, resulting in an isolated north-eastern population in the Sabie-Lochiel area in Mpumulanga (Northern melanotus) and a western population. Relationships in the latter clade are not sufficiently resolved to allow further reconstruction of biogeographic history, but it is clear that a P. m. subviridis-like form became isolated in the south where it eventually came into contact with P. langi at high elevations. Pseudocordylus m. subviridis eventually extended its range southwestwards into the inland mountains of the Eastern Cape and Cape Fold Mountains to give rise to the P. microlepidotus complex. This cycle of range expansion and contraction may also account for the isolated populations at Suikerbosrand, Nkandhla district, and in the Amatole-Great Winterberg mountain region. Furthermore, it is suggested that P. spinosus originated from a P. m. subviridis-like ancestral population that became isolated on the lower slopes of the Drakensberg where terrestrial predation pressure resulted in a quick shift in morphology from fairly smooth body scales to a more spiny morphology. / AFRIKAANSE OPSOMMING: Die taksonomiese status van suidelike Afrika se rotsbewonende krans-akkedisse (genus Pseudocordylus) is ondersoek. Omdat daar aansienlike verwarring in die literatuur bestaan met betrekking tot die tipe monsters en die tipe lokaliteite van die verskillende taksa, is die oplossing van hierdie probleme as die beginpunt van hierdie studie geneem. Die bestudering van akkedismonsters in museums het dit moontlik gemaak om lektotipes, alloparalektotipes en/of paralektotipes aan te wys. Van besondere belang vir hierdie studie is die herontdekking van Andrew Smith se tipe monsters van P. m. melanotus en P. m. subviridis. Die beperking van die tipe lokaliteit van P. m. subviridis, gebaseer op inskrywings in Smith se dagboek en joernaal, het dit moontlik gemaak om vorige interpretasies en definisies van die twee taksa te bevestig. Die geografiese verspreiding van die verskillende taksa en bevolkings is bepaal deur middel van ’n omvattende lokaliteit databasis. Twee soorte molekulêre merkers, naamlik allosieme en mitokondriale DNS, is gebruik in ʼn poging om uitsluitsel te verkry oor die takson-grense binne die P. melanotus-spesiekompleks. Die allosiem-analise het daarop gedui dat P. m. melanotus moontlik polifileties mag wees en uit twee onverwante stamboom-vertakkings kan bestaan. Verder het vaste alleliese verskille tussen parapatriese bevolkings van P. m. melanotus en P. m. subviridis, en tussen simpatriese bevolkings van P. m. subviridis en P. langi, daarop gedui dat al drie vorme as volledige spesies beskou kan word, met die moontlikheid dat meer kriptiese spesies in die kompleks teenwoordig kan wees. Pseudocordylus transvaalensis het van die meeste ander bevolkings verskil met 1-3 vaste alleliese verskille, maar was ononderskeibaar van die bevolking van P. m. melanotus van die Nkandhla distrik (sentraal KwaZulu-Natal). Daar was slegs homosigote individue in ʼn steekproef van Monontsha Pas (Qwa-Qwa), ʼn bevolking wat volgens die literatuur P. m. melanotus en P. m. subviridis, sowel as intermediêre omvat, en alle monsters was toekenbaar aan P. m. subviridis. Die allosiemstudie is egter gebaseer op fenetiese beginsels en vir verdere taksonomiese oplossing is ʼn kladistiese benadering vereis. ʼn Mitokondriale DNS-analise (16S rRNS geen) wat gebruik maak van Maksimum Parsimonie-, Maksimum Waarskynlikheids- en Bayes-analises is daarom uitgevoer om die filogenetiese verwantskappe tussen spesies en subspesies te bepaal en om die taksonomiese status van vorme in die P. melanotus-spesiekompleks te herondersoek. Die mtDNS-analise het die meeste van die resultate van die allosiem-analise bevestig. Eerstens, P. langi is weer bevind om basaal te wees. Met die byvoeging van P. microlepidotus en P. spinosus tot die binne-groep het dit nou duidelik geword dat P. langi die basale spesie in die genus is. (Onlangse studies het aangedui dat P. capensis en P. nebulosus nie kongeneries met Pseudocordylus is nie.) Tweedens, die 16S rRNS resultate bevestig dat P. m. melanotus, soos tans vasgestel, saamgestel is uit twee klade wat nie sustergroepe is nie. Die noordelike bevolkings van P. m. melanotus (Sabie en Lochiel) vorm ʼn redelik diep divergente klaad wat ʼn afsonderlike spesie mag verteenwoordig. Dit is egter bevind dat die Nkandla bevolking saamgegroepeer het met die ander suidelike P. m. melanotus-bevolkings en nie met P. transvaalensis soos wat die geval was in die allosiem-elektroforetiese analise nie. Die mees verbasende resultaat van die 16S rRNS-analise was egter die bevinding dat beide P. microlepidotus en P. spinosus genestel was binne P. m. subviridis. Dit dui daarop dat hierdie twee spesies kon ontwikkel het vanuit P. m. subviridis en slegs onlangs van mekaar geskei het, toe vinnige morfologiese splitsing voorgekom het, maar met beperkte genetiese differensiasie. Dit word voorgestel dat al drie die bogenoemde taksa voorlopig as volledige spesies beskou word. Daar was ook morfologiese steun vir die uniekheid van al die groeperings wat die mtDNS-analise uitgewys het. Pseudocordylus transvaalensis kan uitgeken word aan sy bogemiddelde grootte, unieke dorsale en (swart) kleurpatrone op die keel, so veel as drie horisontale rye lateraaltemporale skubbe, ʼn reeks klein skubbe agter die interpariëtale skub, en gewoonlik twee subokulêre skubbe agter die middelste subokulêre skub op beide kante van die kop. Die verskillende bevolkings wat tans geklassifiseer word as P. melanotus is moeiliker om van mekaar te skei, maar P. m. melanotus en P. m. subviridis verskil gewoonlik soos volg: frontonasale skub in twee gedeel in P. m. melanotus, heel in P. m. subviridis (en in die meeste Noordelike melanotus); lateraal-temporale skubbe in twee rye, die boonste ry met verlengde skubbe teenoor ʼn enkele ry verlengde skubbe; longitudinale rye van dorsolaterale skubbe naby aan mekaar teenoor ver uit mekaar; femorale porieë van wyfies klein en vlak teenoor diep met sekreterende proppe. Noordelike melanotus verskil van Suidelike melanotus deurdat hulle gewoonlik ʼn heel frontonasale skub het en daar selde ʼn klein skub teenwoordig is agter die frontonasale skub. Pseudocordylus langi het unieke dorsale en keel-kleurpatrone (wat ʼn reeks blou kolle op die sye insluit), granulêre dorsale skubbe met 6-9 rye vergrote plat skubbe langs die rugsteen, ʼn groot totale aantal femorale porieë (25-34), en gewoonlik net vyf (glad, ongerif) infralabiale skubbe op elke kant van die kop. Pseudocordylus spinosus het ook unieke dorsale en keel-kleurpatrone, skerp laterale skubbe, frontonasale skub langer as wyd en nie in kontak met die loreale skubbe nie, klein totale aantal femorale porieë (6-9), en wyfies (nie net mannetjies nie) het gedifferensieerde femorale skubbe. Die Hoof-komponent Analise (HKA) en die Kanonieke Diskriminant Analise (KDA) het albei vier groepe geïdentifiseer, naamlik P. transvaalensis, P. langi, P. spinosus en ʼn P. melanotus/subviridis/microlepidotus groepering. ʼn Aparte KDA van alle P. melanotus bevolkings het gedeeltelik onderskei tussen Suidelike melanotus en P. m. subviridis, en die Noordelike melanotus is grootliks van die ander onderskei; terwyl ʼn KDA van P. transvaalensis daarop gedui het dat al drie allopatriese bevolkings 100% onderskeibaar in morfologiese ruimte is. ʼn Genestelde Klaad-Analise het aangedui dat fragmentasie, sowel as gebiedsuitbreiding, ʼn rol gespeel het in die verspreiding van die P. melanotus-spesiekompleks. Dit kan moontlik verklaar word deur die klimaatswisselinge (hoë-lae temperature en nat-droë siklusse) gedurende die Senosoikum wat habitat-uitbreiding en –verkleining veroorsaak het. Gebaseer op die topologie van die mtDNS filogram is dit duidelik dat die genus Pseudocordylus al langs die oostelike platorand ontstaan het. ʼn Voorouer soortgelyk aan P. langi kon ʼn uitgebreide gebied al langs die oostelike platorand gehad het, met die Maloti-Drakensberg wat die suidelike limiet van hierdie gebied gevorm het. Gedurende ʼn daaropvolgende toename in globale temperature het gebiedsverkleining en fragmentasie plaasgevind, wat ʼn geïsoleerde bevolking in die suide en een in die noorde tot gevolg gehad het. Die suidelike bevolking het onveranderd oorleef in die Maloti-Drakensberg skuilplek (“refugium”), maar die noordelike bevolking is geforseer om aan te pas in die warmer toestande. Daarna het die noordelike vorm se gebied weer uitgebrei, maar gedurende ʼn daaropvolgende koeler periode het gebiedsverkleining weer plaasgevind, met die gevolg dat daar ʼn geïsoleerde noord-oostelike bevolking in die Sabie-Lochiel-area in Mpumalanga (Noordelike melanotus) en ʼn bevolking in die weste was. Verwantskappe in die laasgenoemde klaad is nie voldoende opgelos om verdere rekonstruksie van die biogeografiese geskiedenis moontlik te maak nie, maar dit is duidelik dat ʼn vorm soortgelyk aan P. m. subviridis geïsoleer geraak het in die suide waar dit eindelik op hoë liggings in kontak gekom het met P. langi. Die gebied van P. m. subviridis is ook later suidweswaarts uitgebrei tot in die binnelandse berge van die Oos-Kaap en Kaapse Plooiberge om tot die ontstaan van die P. microlepidotuskompleks aanleiding te gee. Hierdie siklus van gebiedsuitbreiding en verkleining kan ook ʼn verklaring bied vir die geïsoleerde bevolkings by Suikerbosrand, Nkandhla distrik, en in die Amatole-Groot Winterberg-streek. Verder word voorgestel dat P. spinosus ontstaan het uit ʼn voorouerlike bevolking soortgelyk aan P. m. subviridis wat geïsoleerd geraak het op die laer hange van die Drakensberg waar die druk van aardsbewonende roofdiere tot ʼn vinnige verandering in morfologie vanaf redelik gladde liggaamskubbe tot ʼn meer skerppuntige morfologie gelei het.