Range change is a common species response to global change. Comparing historical species distribution data with recent biological surveys has the potential to quantify changes to species geographic ranges. However, the broad-scale sampling strategies typically employed to acquire primary species distribution data are prone to errors of omission. The aim of this study was to evaluate the South African Frog Atlas Project (SAFAP) as a means for detecting changes in amphibian species distributions and to relate observed range changes to extrinsic environmental factors and intrinsic species characteristics. The SAFAP provided historical (1905 – 1995) and recent (1996 – 2003) species distributions of the amphibians of South Africa. Geographic sampling bias in the dataset was assessed by relating collection density and species richness to hypothesised sources of bias. Several methods for managing differing sampling intensity were tested on hypothetical ranges. The best methods were applied to the South African species to investigate range dynamics. Changes to the size of species ranges and shifts in mean range centre were assessed. An Ecological Niche Factor Analysis provided comparative measures of climate and habitat niche breadth for each species. SAFAP sampling was concentrated around cities, roads and protected areas, resulting in relatively overestimated species richness and range sizes near to these features. Large parts of the arid northwestern regions were under-sampled. An increase in sampling intensity over time resulted in the false detection of range expansions. The most reliable method to correct for increased sampling was a mathematical correction factor, according to which, 60.2% of South African frog species have undergone range contractions. Upslope shifts of 47.6 m were found for South African species and species of the Bushveld region shifted towards an area of Savanna Biome resilience. While several of the observed changes to species ranges were consistent with global change predictions, southern hemisphere amphibians may show a differing response to global change to that which is commonly predicted. Small range size, habitat specialisation and climate specialisation were significant predictors of range contractions for all species. Contracting habitat specialists were concentrated within two areas of endemism that also had high levels of land transformation. The use of methods that correct for sampling variation has allowed the SAFAP to be valuable in investigating species range change.

Thesis (PhD)--Stellenbosch University, 2010. / ENGLISH ABSTRACT: Biological invasions are a growing threat to biodiversity, trade and agriculture in South Africa. Though alien reptiles and amphibians (herpetofauna) are not currently a major issue, escalating problems worldwide and increased trade in South Africa suggest a possible increase in future problems. In this thesis I explore practical measures for risk assessment implementable under national legislation. I began by documenting record-keeping and legislative differences between provinces in South Africa. This revealed some serious deficiencies, complicating attempts to compile accurate inventories and discern import trends. International trade data, however, revealed an exponential increase in the number of imports to South Africa over the last 30 years. Characterising the abundance of species in this trade is important as species introduced in large numbers pose a higher establishment risk. In South Africa, I found a tendency for venomous and expensive species to be traded in low numbers, whereas species that are easy to breed and handle, or that are colourful or patterned are traded in higher numbers. Unlike South Africa, California and Florida have had a large number of well-documented herpetofaunal introductions. These introductions were used to verify the role of several key predictors in species establishment. I first evaluated the role of each variable separately. I examined different approaches for bioclimatic modelling, the predictive power of different sources of distribution data, and methods of assigning a climate-match score. I also present the first test of Darwin’s naturalization hypothesis for land vertebrates using two new phylogenies inferred for native and introduced reptiles in California and Florida. I then used boosted regression trees (BRT) to infer the relative contribution of each factor to species establishment success. Results from the BRTs were incorporated into a user-friendly spreadsheet model for use by assessors inexperienced in complex modelling techniques. Introduction effort was found to be the strongest contributor to establishment success. Furthermore, species with short juvenile periods were more likely to establish than species that started breeding later, as were species with more distant relatives in regional biotas. Average climate match and life form were also important. Of the herpetofaunal groups, frogs and lizards were most likely to establish, while snakes and turtles established at much lower rates, though analysis of all recorded herpetofaunal introductions shows slightly different patterns. Predictions made by the BRT model to independent data were relatively poor, though this is unlikely to be unique to this study and can be partially explained by missing data. Though numerous uncertainties remain in this field, many can be lessened by applying case by case rules rather than generalising across all herpetofaunal groups. The purpose for import and potential trade volume of a species will influence the threat it poses. Considering this in conjunction with a species’ environmental tolerances and previous success of species with similar life histories, should provide a reasonable and defendable estimate of establishment risk. Finally, a brief summary of the potential impacts of introduced alien herpetofauna is provided in the thesis. / AFRIKAANSE OPSOMMING: Indringer spesies hou ‘n al groter bedreiging in vir die biodiversiteit, handel en landbou van Suid- Afrika. Alhoewel uitheemse reptiele en amfibieërs (herpetofauna) tans nie ‘n groot bedreiging in Suid-Afrika is nie, dui groeiende probleme wêreldwyd asook 'n toename in plaaslike handel op moontlike toekomstige probleme. In hierdie tesis, ondersoek ek praktiese metodes vir risikobepaling wat onder nasionale wetgewing toegepas kan word. Ek begin deur die verskille in stoor van rekords en wetgewing tussen provinsies te dokumenteer. Hierdie proses het ernstige tekortkominge uitgewys, wat pogings om akkurate inventarisse saam te stel en invoertendense te bepaal, bemoeilik. Internasionale handelsdata het egter getoon dat daar ‘n eksponensiële toename in die hoeveelheid invoere na Suid-Afrika oor die laaste 30 jaar was. Die hoeveelheid spesies in hierdie handel is belangrik omdat spesies wat in groot hoeveelhede ingevoer word, ‘n hoër vestigingsrisiko het. In Suid-Afrika is ‘n tendens gevind vir handel in giftige en duur spesies teen lae hoeveelhede, terwyl spesies wat maklik teel, maklik hanteer kan word en kleurvol is of mooi patrone het, in groter hoeveelhede mee handel gedryf word. Kalifornië and Florida, in teenstelling met Suid-Afrika, het ‘n hoë aantal goed-gedokumenteerde gevalle van herpetofauna wat in die natuur vrygestel is. Hierdie introduksies was gebruik om die rol van verskeie belangrike faktore in die vestiging van populasies te bepaal. Eerstens het ek die rol van elke faktor apart ondersoek. Ek het verskillende benaderinge vir bioklimatiese model-bou ondersoek, die akuraatheid van verskillende bronne van distribusiedata getoets en drie metodes om ‘n “climate match score” te bereken, voorgestel. Ek bied ook die eerste toets van Darwin se naturalisasie-hipotese vir landwerveldiere aan, deur gebruik te maak van twee nuwe filogenieë wat ek gebou het vir inheemse en ingevoerde reptiele in Kalifornië en Florida. Ek het verder gebruik gemaak van “boosted regression trees” (BRT) om die relatiewe bydrae van elke faktor tot die vestigings-potensiaal van spesies te bepaal. Resultate van hierdie BRTs was ingekorporeerd in ‘n gebruikersvriendelike ontledingstaat wat deur bestuurders, onervare in komplekse modelboutegnieke, gebruik kan word. Invoer-hoeveelheid was die faktor wat die sterktste bygedra het tot suksesvolle vestiging. Verder is spesies met kort jeugperiodes en dié met verlangse familie in streeks-biotas, meer geskik om suksesvol te vestig. Gemiddelde klimaatooreenstemming en lewensvorm was ook belangrik. Paddas en akkedisse was die mees waarskynlikste van die herpetofauna groepe om populasies te vestig, terwyl slange en skilpaaie teen laer tempo’s populasies gevestig het, alhoewel analise van alle gedokumenteerde gevalle van herpetofauna-invoerings wêreldwyd effens verskillende tendense toon. Voorspellings wat deur die BRT-model vir onafhangklike data gemaak is was redelik swak, alhoewel hierdie resultaat onwaarskynlik nie uniek aan die studie is nie, en word gedeeltelik verduidelik deur die gebrek aan data. Alhoewel talle onsekerhede steeds bestaan, kan dié verminder word deur geval-tot-geval reëls toe te pas eerder as om vir herpetofauna as ‘n groep te veralgemeen. Die doel van invoer en potensiële handel-volumes van ‘n spesie, sal die bedreiging wat die spesie toon, bepaal. Hierdie faktore moet saam met omgewingstoleransie en voorafgaande sukses van spesies met soortgelyke lewenswyses oorweeg word, om ‘n aanvaarbare en verdedigbare beraming van vestigingsrisiko te gee. Laastens, word ‘n kort opsomming van die effekte wat uitheemse herpetofauna mag hê, verskaf.