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A prominent scenario for the evolution of reptilian placentation infers that placentotrophy arose by gradual modification of a simple vascular chorioallantoic placenta to a complex structure with a specialized region for nutrient transfer. The structure of the chorioallantoic placenta of Niveoscincus ocellatus, apparently described originally from a single embryonic stage, was interpreted as a transitional evolutionary type that provided support for the model. Recently, N. ocellatus has been found to be as placentotrophic as species with complex chorioallantoic placentae containing a specialized region called a placentome. We studied placental development in N. ocellatus and confirmed that the chorioallantoic placenta lacks specializations found in species with a placentome. We also found that this species has a specialized omphaloplacenta. The chorioallantoic placenta is confined to the region adjacent to the embryo by a membrane, similar to that found in some other viviparous skinks, that divides the egg into embryonic and abembryonic hemispheres. We term this structure the "inter-omphalopleuric" membrane. The position of this membrane in N. ocellatus is closer to the embryonic pole of the egg than to the abembryonic pole and thus the surface area of the omphaloplacenta is greater than that of the chorioallantoic placenta. In addition, the omphaloplacenta is regionally diversified and more complex histologically than the chorioallantoic placenta. An impressive and unusual feature of the omphaloplacenta of N. ocellatus is the development of extensive overlapping folds in the embryonic component of midgestation embryos. The histological complexity and extensive folding of the omphaloplacenta make this a likely site of placental transfer of nutrients in this species.

I evaluated possible constraints on the evolution of viviparity in the lizard genus Sceloporus by experimentally extending the length of egg retention past the normal time of oviposition for a number of oviparous species. Observations also included a representative of the genus Urosaurus, the sister genus to Sceloporus. I determined the effects of retention on embryonic development, hatchlings, and gravid females. Results indicated that the proximate constraints on longer retention times and viviparity are 1) embryonic development becomes arrested or severely retarded in utero, and 2) the ability to maintain gravidity past the normal time of oviposition is limited in some species. Observations on Urosaurus further showed that extended egg retention results in hatchlings with traits that are associated with lower fitness. I also tested the hypothesis that reproductive Sceloporus lower their body temperatures during activity because their normal body temperatures are detrimental to embryos. Observations on a viviparous species of Sceloporus indicated that the normal body temperature of the female was detrimental to embryonic development. This result is indicates that viviparity would be constrained in some squamate lineages if maternal body temperatures are too high for successful embryonic development. I also evaluated the hypothesis that the evolutionary transition from oviparity to viviparity involves a "reduction" of the eggshell concurrent with longer durations of egg retention. If this hypothesis is correct, then attributes of eggshells that should enhance exchange of respiratory gases (i.e., thickness, density, permeability to water vapor) would be correlated with the maximal developmental stage that embryos are able to attain in the oviducts (i.e., stage of developmental arrest). The results of this study indicated that these features of shells do not determine the stage at which development becomes arrested. Thus, the results do not support the hypothesis that shell reduction occurs concurrently with longer periods of egg retention. The results are consistent with the alternative hypothesis that reduction of the eggshell occurs after viviparity has evolved. / Ph. D.

Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: As ectotherms, lizards are particularly vulnerable to changes in the thermal landscape and face extinction risk if they lack the capacity to rapidly adapt or behaviourally mitigate increasingly altered thermal environments. Theoretical models that predict lizards‟ response to climate change often fail to take into account the thermal characteristics of the microenvironment, the ability of lizards to behaviourally buffer climate variation in the habitat and the plastic nature of both behaviour and physiology over ecologically relevant time-scales. Here, I address this major knowledge gap using two separate research chapters in an experimental physiology approach. In Chapter 1, I investigated the temperature-dependence and plasticity of resting metabolic rate, water-loss rate and preferred body temperature of Cordylus oelofseni at several temporal scales (within and between seasons) and incorporated field observations to acquire a better understanding of this species‟ adaptive potential to buffer thermal changes in the habitat. Cordylus oelofseni showed plasticity of both behaviour and physiology in response to thermal acclimation, but relied on distinct strategies depending on the time-scale investigated. These results highlighted the complexity of underlying mechanisms used by these organisms to buffer temperature variation. In Chapter 2, I used an experimental approach to examine the energetic costs of thermoregulation in C. oelofseni and test the cost-benefit model of thermoregulation. This model‟s primary prediction states that lizards should thermoregulate carefully only when the associated costs are low. Using four enclosures that simulated different thermal qualities (temporal and spatial distributions of operative temperatures) in the habitat, I found limited support for the cost-benefit model. Lizards in the low-quality heterogeneous enclosures invested the same energetic effort and thermoregulated with similar overall accuracy as lizards in the high-quality heterogeneous enclosure. The costs incurred were not necessarily energetic, but reflected missed opportunities (e.g. less time to forage), something that, along with important interaction effects with body mass, deserves further attention when testing this model. Together, these results illustrate the importance of incorporating ecological reality at various time and spatial scales in order to make relevant predictions regarding the fate of lizards with projected climate change. / AFRIKAANSE OPSOMMING: As ektotermiese diere, is akkedisse veral sensitief vir veranderinge in die termiese landskap en staar uitsterwingsrisiko in die gesig as hulle nie die vermoë het om vinnig aan te pas of gedragsveranderinge te maak in omgewings wat toenemend verwarm nie. Teoretiese modelle wat akkedisse se reaksie op klimaatsverandering voorspel, neem dikwels nie die termiese eienskappe van die mikro-omgewing, die vermoë van akkedisse om met gedragsveranderinge klimaat variasie in die habitat te buffer en die plastieke aard van beide gedrag en fisiologie oor ekologies relevante tydskale in ag nie. Hier bespreek ek hierdie groot kennisgaping met behulp van twee afsonderlike navorsingshoofstukke in 'n eksperimentele fisiologie benadering. In Hoofstuk 1 het ek ondersoek ingestel na die temperatuur-afhanklikheid en plastisiteit van rustende metaboliese tempo, waterverlies tempo en voorkeur liggaamstemperatuur van Cordylus oelofseni by verskeie tydskale (binne en tussen seisoene) en inkorporeer veld waarnemings om 'n beter begrip te verkry van hierdie spesie se aanpasbare potensiaal om termiese veranderinge in die habitat te buffer. Cordylus oelofseni het plastisiteit van beide gedrag en fisiologie in reaksie op hitte-akklimatisering getoon, maar staatgemaak op verskillende strategieë, afhangende van die tyd-skaal wat ondersoek is. Hierdie resultate beklemtoon die kompleksiteit van die onderliggende meganismes wat gebruik word deur hierdie organisme om temperatuur verandering te buffer. In Hoofstuk 2 het ek 'n eksperimentele benadering gebruik om die energiekoste van termoregulering in C. oelofseni te ondersoek en die kostevoordeel model van termoregulering te toets. Hierdie model se primêre voorspelling verklaar dat akkedisse slegs versigtig moet termoreguleer wanneer die gepaardgaande koste laag is. Deur gebruik te maak van vier afskortings wat verskillende termiese eienskappe gesimuleer het (tyd en ruimtelike verspreiding van operatiewe temperature) in die habitat, het ek beperkte ondersteuning gevind vir die koste-voordeel model. Akkedisse in die lae-gehalte heterogene afskortings het dieselfde energieke moeite belê en getermoreguleer met soortgelyke algehele akkuraatheid as akkedisse in die hoë-gehalte heterogene kamp. Die kostes wat aangegaan is, is nie noodwendig energiek nie, maar weerspieël geleenthede wat gemis is (bv. minder tyd om kos te soek), iets wat, saam met belangrike interaksie effekte met liggaamsmassa, verdere aandag verdien wanneer hierdie model getoets word. Tesame illustreer hierdie resultate die belangrikheid van die integrasie van ekologiese werklikheid op verskillende tyd en ruimtelike skale, om relevante voorspellings oor die lot van akkedisse met geprojekteerde klimaatsverandering te kan maak.

Male growth is best described by a von Bertalanffy growth model and female growth by a logistic growth model. This sexual dimorphism is correlated with greater relative surface activity and lower survivorship of juvenile males when compared to juvenile females. Early-hatching (first clutch) offspring exhibit greater body size and survivorship compared to late-hatching (second clutch) offspring. (Abstract shortened with permission of author.)

The thermoregulation of lizards depends highly on the environment they inhabit. Thermoregulation is crucial for physiological processes and affects therefore the life history of the lizard. High altitudes are characterized by a colder climate than on lower altitudes, and these environmental factors affects the thermoregulation of lizards. Lizards can however adapt to colder climates by adjusting their physiology and/or behavior. In this study two populations of Liolaemus aparicioi, an endemic lizard inhabiting the valleys close to La Paz, were observed at two different altitudes to investigate any differences in behavior or habitat use that could compensate for the high altitude. The behavioral displays associated to thermoregulation and the use of habitat were recorded in order to assess how they allocated their time and how they used the habitat available. The mean values of most parameters measured indicated that the population on the high altitude do compensate for the high altitude by behavior and habitat use.