×

The basic anatomy of the lacertid hemipenis (intromittent organ) and methods for its investigation are described. In many members of the Lacertidae, the hemipenis has a structure quite unlike that of other squamate reptiles: the distal lobes of the retracted organ are complexly folded and there is a well-defined supporting structure of dense connective tissue, the armature. This incorporates blood sinuses and has an intramuscular portion embedded in the m. retractor penis magnus and two club-shaped bodies, the clavulae, that support the lobes in the erect organ. Unarmatured hemipenes occur in some lacertids and, like those of other squamates, possess sac-like lobes in the retracted state, but they are singular in having the lobes invested by the m. retractor penis magnus. It is argued that many of these apparently primitive hemipenes are in fact secondary derivatives of the armatured type. There is considerable inter-specific variation in hemipenial structure which is described systematically. In some cases this involves differences in size, asymmetry and simplification, which may arise as physical isolating mechanisms and is useful in distinguishing otherwise very similar species, particularly in the genus Mesalina (p. 1253). Other shared derived hemipenial features provide useful information about relationships between species and higher taxa and a summary of the hypotheses that they support is given (p. 1254).

×

Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.

×

The six floristic biomes in the Western Cape Province (W.C.P.), namely the Fynbos, Afromontane Forest, Thicket, Grassland, Nama and Succulent Karoo Biomes (Low and Rebelo, 1996), are not only diverse with regard to the variety of plant species and communities occurring there, but also contain a wide diversity of animal species, biogeographical zones, landscapes and natural features, both within the terrestrial and aquatic (freshwater and marine) context. In addition to the topographical diversity of the Cape Fold Mountains, the coastal zone and lowlands, and their transition into surrounding habitats, the W.C.P. experiences a wide climatic diversity too. These features have resulted in an extensive and complex diversity of habitat types which partly explain the rich biological diversity within the W.C.P. Past climatic changes on a global scale have also influenced ecological systems and processes within the W.C.P. to the extent where it is believed that vicariant speciation processes and events during global climatic changes have resulted in evolutionary driving forces that have had significant impacts on the biodiversity within the biogeographical boundaries of the W.C.P. (Vrba, 1985).

×

New and additional distribution records are given for 12 amphibian, 25 lizard, one amphisbaenian, and five snake taxa from the Cape Province, South Africa. Some of the new records represent large range extensions, whereas others fill in gaps or confirm the occurrence of taxa in the region.

×

The taxonomic status, distribution and ecology of the herpetofauna of the Orange Free State are discussed, based on the examination of 10 096 specimens housed in various southern African museums. The majority of specimens were collected during the years 1972 - 1978 and 1983 - 1992, and are housed at the National Museum, Bloemfontein. A total of 25 amphibian and 95 reptilian (one translocated species) forms have been determined as occurring in the Orange Free State, 12 (three amphibians, nine reptiles) of which are new (or recently published) records for the province. An additional 10 amphibian and 28 reptilian forms have been determined as occurring nearby but extralimitally to the O.F.S., and several of these forms are expected to occur in the province. New distributional records have resulted in the amendment of the ranges of several species. Detailed taxonomic data on new material, including rare species, have been given. Some problem areas in the taxonomy of O.F.S. taxa have been higlighted. An analysis of habit utilization indicated that 84,0% of amphibians and 61,1% of reptiles are terricolous, whereas up to 21,5% of reptiles are rupicolous. The majority of snakes (72,2%) are terrestrial in habits, but 13,9% are fossorial. More than half of all lizards (52,8%) are terrestrial, although 34,6% are rupicolous. A total of six amphibian and 27 reptile forms utilize inactive termitaria as a microhabitat, including several basically terrestrial forms. Snake forms were particularly well represented in termitaria, 60,6% of all forms known from the O.F.S. having been recorded from inactive termitaria. General features of the ecology of O.F.S. amphibians and reptiles have also been discussed. The biogeographical analysis indicated that O.F.S. amphibian forms can be classified into one of nine range clusters (common patterns of distribution), and reptiles into 13 such cluster groups. By testing these classifications by means of a transect through the northern O.F.S. (from eastern to western borders), it was determined that a fairly distinct east-west subtraction of amphibian and reptilian species and subspecies occurs in the O.F.S. Clustering of range boundaries and high species and subspecies diversity at the western and eastern ends of the transect zone suggest dynamic biogeographical situations occurring in those areas - the western group being associated with the transition from grassland to bushveld, and the eastern group associated with the transition from Highveld Grassland to Drakensberg Mountains. The general eastern and western groupings of taxa appear to be associated with the cooler, wetter and mountainous east vs the warmer, drier and lowerlying west, respectively. Despite a great deal of collecting having been conducted in the O.F.S. from 1972 to 1992, an analysis of the number of taxa collected in each quarter-degree unit in `the O.F.S. indicated that additional collecting would be required in order to conduct effectively a mathematically-based biogeographical analysis.

×

Data relating to the Sandveld lizards (Nucras) occurring in Namibia, southwest Africa are reviewed. In particular, we investigated records of N. holubi, a chiefly southeastern African species, and attempted to identify recently collected material that could not be assigned to any species currently recognized in Namibia. A phylogenetic analysis of Nucras based on three mitochondrial markers revealed a deep divergence between Namibian Nucras holubi and two presumably conspecific clades from Limpopo Province, South Africa. In addition, the coloration pattern and scalation of the Namibian material differ from those of the eastern forms, supporting its recognition as a separate species. The name Nucras damarana Parker, 1936, long relegated to the synonymy of N. holubi, is here resurrected for this apparently endemic northern Namibian species. Nucras damarana is restricted to the Kunene, Omusati, Oshikati, Kavango, and Otjozondjupa regions of north-central Namibia. A distinctive specimen of Nucras from near Ruacana in the Kunene Region was identified as allied to Nucras broadleyi, a species recently described from southwestern Angola, on the basis of genetic data, although it differs substantially in color pattern. With the addition of N. aff. broadleyi and the resurrected N. damarana to its fauna, as well as the removal of N. holubi from the nation’s species list, four species of Nucras are confirmed to be present in Namibia. Although the conservation status of N. damarana, N. tessellata, and N. intertexta is Least Concern, the uncertain taxonomic status of N. aff. broadleyi precludes a meaningful threat assessment.

×

New informations concerning systematical questions within the family Lacertidae, which were published during the last year, are presented.

×

A checklist of the lizards of the Cape Province, South Africa is presented. A total of 143 species and subspecies is recognized. The taxonomic status of no less than 84 (59,1%) of these has changed since the last review of the Cape lizards by FitzSimons (1943). Comments on the distribution and taxonomic status of problematic groups are included.

×

Numerous new distribution records are given for 19 amphibian, 3 tortoise, 32 lizard, and 15 snake species and subspecies in the Cape Province, particularly in the eastern and central karroid regions. Many species are shown to have much wider distributions than previously believed. The importance of the mountains of the inland escarpment as a refugium for temperate species, the presence of isolated populations of western arid species in the karroid valleys of the northern Cape Fold Mountains, and the possible vicariant effects caused by sea level fluctuations on species inhabiting the southern Cape coastal plain, are briefly discussed.

×

The phenotype of an individual has often been used as the descriminating factor in distinguishing species. However, with the advent of more precise molecular techniques, the genotype of species is increasingly being used as the preferred method in taxonomic classifications. Many taxa have recently been demonstrated to be incongruent in terms of their genetic and morphological groupings, and this may due to the influence that the environment may have on the morphological and functional aspects of a species. Selective pressures often act upon the performance of a species within a particular habitat first, and then selection for the morphological characters that allow for optimal performance occurs. Should genetically disparate species inhabit a particular environment, convergence in morphologies and performance may evolve. Historically, lizard species descriptions were based primarily on external morphologies, and thus misclassfication of species may have occurred due to mistakenly grouping species with convergent morphologies together. In the current dissertation, the links between morphology, performance capacities, diet and behaviour is explored in comparison to the environment and genetic relationships of southern African lacertid lizards. The performance capacities and associated morphological traits were expected to be more closely linked with the environment, and not closely linked with genetic relationships. To investigate these expectations, a multidisciplinary approach was taken, and genetic, morphological and performance analyses were done and compared with dietary behavioural and environmental analyses. In the first chapter, the link between habitat openness and the lizard bauplans is investigated and the presence of convergent morphologies within this group of lizards is uncovered. These convergences are shown to have resulted in misclassification of two lacertid species, and taxonomic revisions within the family are discussed. The second chapter explores the link between performance and associated morphological traits, and the dietary composition of the members of the Nucras genus. The third chapter identifies the link between the predator escape strategies employed by the members of the Meroles genus, and their morphologies and performance capacities. The fourth chapter explores the intraspecific, inter-population differences in morphologies and investigates the link between the morphological groupings and the population genetic groupings within Pedioplanis lineoocellata. The final chapter identifies whether adaptation to a novel habitat can occur over a relatively short period of time, and the morphological traits, functional aspects, and population genetic structure is investigated in conjunction with environmental analyses of vegetation and substrate between the populations of Meroles knoxii. It was concluded that the morphological and functional aspects of the southern African lacertid lizards are more closely related to the environment, particularly the microhabitat structure, than to their genetic relationships, and that future work using this group of lizards should involve a multidisplinary approach as different selective pressures are playing a role in shaping the morphologies and performance capacities of these lizards, compared to those that are acting upon the genotypes of the lizards.

×

The functional characteristics of prey items (such as hardness and evasiveness) have been linked with cranial morphology and performance in vertebrates. In lizards particularly, species with more robust crania generally feed on harder prey items and possess a greater bite force, whereas those that prey on evasive prey typically have longer snouts. However, the link between dietary niche breadth, morphology, and performance has not been explicitly investigated in lizards. The southern African genus Nucras was used to investigate this link because the species exhibit differing niche breadth values and dietary compositions. A phylogeny for the genus was established using mitochondrial and nuclear markers, and morphological clusters were identified. Dietary data of five Nucras species, as reported previously, were used in correlation analyses between cranial shape (quantified using geometric morphometrics) and dietary niche breadth, and the proportion of hard prey taken and bite force capacity. Dietary niche breadth and the proportion of hard prey eaten were significantly related to cranial shape, although not once phylogeny was accounted for using a phylogenetic generalized least squares regression. The proportion of evasive prey eaten was a significant predictor of forelimb length when phylogeny was taken into account. We conclude that, in Nucras, the percentage of evasive prey taken co-evolves with forelimb morphology, and dietary niche breadth co-evolves with cranial shape. However, although head width is correlated with the proportion of hard prey eaten, this appears to be the result of shared ancestry rather than adaptive evolution.

×

Eremiadinae, one of three subfamilies of Lacertidae, are distributed throughout Asia and Africa. Previous phylogenetic studies suggested that one of the main groups of Eremiadinae (the Ethiopian clade) consist of two clades with predominately East-African and South-African distribution. Yet, especially the latter one, which includes the genera Pedioplanis, Meroles, Ichnotropis, Tropidosaura and Australolacerta, was not well supported in the molecular phylogenetic analysis. In this study, we analysed the phylogenetic relationships among the genera of the ‘South African clade’ to assess whether this group actually forms a highly supported clade and to address questions concerning the monophyly of the genera. We sequenced sections of the widely used mitochondrial genes coding for 16S rRNA, 12S rRNA and cytochrome b (altogether 2045 bp) as well as the nuclear genes c-mos, RAG-1, PRLR, KIF24, EXPH5 and RAG-2 (altogether 4473 bp). The combined data set increased the support values for several nodes considerably. Yet, the relationships among five major lineages within the ‘South African clade’ are not clearly resolved even with this large data set. We interpret this as a ‘hard polytomy’ due to fast radiation within the South African lacertids. The combined tree based on nine marker genes provides strong support for the ‘South African Clade’ and its sister group relationship with the ‘East African Clade’. Our results confirm the genus Tropidosaura as a monophylum, while Ichnotropis is paraphyletic in our trees: Ichnotropis squamulosa appears more closely related to Meroles than to Ichnotropis capensis. Furthermore, the monophyly of Meroles is questionable as well. Based on our results, I. squamulosa should be transferred from Ichnotropis into the genus Meroles. Also, the two species of Australolacerta (A. australis and A. rupicola) are very distantly related and the genus is perhaps paraphyletic, too. Finally we propose a phylogeographical scenario in the context of palaeoclimatic data and compare it with a previously postulated hypothesis.

×

The evolutionary diversification of many terrestrial vertebrate groups is strongly linked to climatic events in the Cenozoic, the period from 65 Million years ago to today when modern animals first appeared. I investigated the effects of Cenozoic climate change on the taxonomic and morphological diversification of the Old World lizard family Lacertidae, with particular emphasis on the African radiation. African lacertids exhibit an unusual pattern of diversification, in which their highest species richness occurs in deserts north and south of the equator, despite being spread throughout the continent. This disparity is particularly surprising given that desert lacertids are thought to be evolutionarily younger than their mesic-dwelling relatives, suggesting increased diversification rates in arid habitats. To identify the evolutionary factors underlying this pattern, I use a combination of phylogenetic, morphological and ecological techniques. In Chapter 1, I apply Bayesian methods and fossil-based calibrations to molecular sequence data to construct a time-calibrated phylogeny for Lacertidae. I estimate that the family arose in the early Cenozoic, with the majority of their African radiation occurring in the Eocene and Oligocene. In Chapter 2, I describe changes in lacertid body shape across biomes and substrates, and find widespread morphological convergence in similar habitat types. I suggest that in addition to foraging demands, fluctuating and extreme climatic conditions, largely driven by precipitation and temperature, contribute to morphological convergence across independent arid-dwelling clades. Finally, I test if ancestral transitions in ecology, morphology, and rates of diversification temporally coincide with paleoclimatic events in the Cenozoic. I use High Resolution X-ray Computed Tomography to characterize changes in the skull related to life in arid habitats, and apply maximum likelihood methods to test if the origins of those traits temporally coincide with significant shifts in habitat, diversification rates and climatic changes. My results show that African lacertids experienced three major peaks in diversification, accompanied by the evolution of suites of arid-adapted morphological traits. These changes coincide with climatic shifts in Africa, including the transition from closed forests to open grasslands and savanna in the late Oligocene, prior to the peak temperatures of the mid-Miocene Climatic Optimum, and following the formation of the Benguela current leading to hyper-aridity in southern Africa. I conclude that deserts are important centers for reptile evolution, but that expected changes in climate due to global warming may outpace the ability of arid-dwelling species to adapt and persist in the future.

×

Several specimens of different genera of Lacertid lizards were collected during a 4-week-trip into the Limpopo Province of South Africa. Depending on their habitat members of the different species occurred within different reptile species assemblages. On the Soutpansberg specimens of Nucras intertexta were recorded in unusual habitat and a genetic study shall resolve their systematic position. No Lacertids were recorded in the high altitudes of the Blouberg and Wolkberg, which is most probably attributed to sampling artifacts.

×

Aim To map and assess the richness patterns of reptiles (and included groups: amphisbaenians, crocodiles, lizards, snakes and turtles) in Africa, quantify the overlap in species richness of reptiles (and included groups) with the other terrestrial vertebrate classes, investigate the environmental correlates underlying these patterns, and evaluate the role of range size on richness patterns. Location Africa. Methods We assembled a data set of distributions of all African reptile species. We tested the spatial congruence of reptile richness with that of amphibians, birds and mammals. We further tested the relative importance of temperature, precipitation, elevation range and net primary productivity for species richness over two spatial scales (ecoregions and 1° grids). We arranged reptile and vertebrate groups into range-size quartiles in order to evaluate the role of range size in producing richness patterns. Results Reptile, amphibian, bird and mammal richness are largely congruent (r = 0.79–0.86) and respond similarly to environmental variables (mainly productivity and precipitation). Ecoregion size accounts for more variation in the richness of reptiles than in that of other groups. Lizard distributions are distinct with several areas of high species richness where other vertebrate groups (including snakes) are species-poor, especially in arid ecoregions. Habitat heterogeneity is the best predictor of narrow-ranging species, but remains relatively important in explaining lizard richness even for species with large range sizes. Main conclusions Reptile richness varies with similar environmental variables as the other vertebrates in Africa, reflecting the disproportionate influence of snakes on reptile richness, a result of their large ranges. Richness gradients of narrow-ranged vertebrates differ from those of widespread taxa, which may demonstrate different centres of endemism for reptile subclades in Africa. Lizard richness varies mostly with habitat heterogeneity independent of range size, which suggests that the difference in response of lizards is due to their ecological characteristics. These results, over two spatial scales and multiple range-size quartiles, allow us to reliably interpret the influence of environmental variables on patterns of reptile richness and congruency.

×

Habitat transfonnation is the primary anthropogenic threat to global biodiversity. Fragmentation of reptile populations following habitat transfonnation within a landscape can lead to the extirpation of species. We investigated the effects of land-use on the species richness and abundance of reptile assemblages in three habitat types (two natural and one modified) in the grasslands of Gauteng, South Africa. Using trap arrays, we surveyed reptiles in primary grassland with little or no rock cover, primary grassland with large quartzite outcrops and scattered rocks, and secondary grasslands that were historically ploughed and cropped. We measured vegetation height and vegetation cover at these same localities. We caught significantly fewer reptile species in the historically cultivated sites than in either of the two natural habitat types. Differences in the reptile assemblage of each habitat type were not explained by either the spatial location or the vegetation structure of our trap sites but were well explained by the sites` habitat type. Estimates of total species richness indicated that we were able to adequately sample the reptile assemblages in the three habitat types, further supporting our observation of reduced species richness in the secondary grasslands. We infer that habitat transfonnation associated with cultivation e.g., rock removal, has had a detectable, negative impact on the species richness and composition of the local reptile assemblages. We recommend that land-use planning in Gauteng emphasise the need for areas of inter-connected, untransfonned habitat in order to mitigate the negative impacts of habitat transfonnation on the local reptile diversity.

×

The family Lacertidae encompasses more than 250 species distributed in the Palearctis, Ethiopis and Orientalis. Lacertids have been subjected in the past to several morphological and molecular studies to establish their phylogeny. However, the problems of convergent adaptation in morphology and of excessively variable molecular markers have hampered the establishment of well supported deeper phylogenetic relationships. Particularly the adaptations to xeric environments have often been used to establish a scenario for the origin and radiation of major lineages within lacertids. Here we present a molecular phylogenetic study based on two nuclear marker genes and representatives of 37 lacertid genera and distinct species groups (as in the case of the collective genus Lacerta). Roughly 1600 bp of the nuclear rag1 and c-mos genes were sequenced and analyzed. While the results provide good support to the hitherto suggested main subfamilies of Gallotiinae (Gallotia and Psammodromus), Eremiainae and Lacertinae [Harris, D.J., Arnold, E.N., Thomas, R.H., 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc. Lond. B 265, 1939–1948], they also suggest unexpected relationships. In particular, the oriental genus Takydromus, previously considered the sister-group to the three subfamilies, is nested within Lacertinae. Moreover, the genera within the Eremiainae are further divided into two groups, roughly corresponding to their respective geographical distributions in the Ethiopian and the Saharo-Eurasian ranges. The results support an independent origin of adaptations to xeric conditions in different subfamilies. The relationships within the subfamily Lacertinae could not be resolved with the markers used. The species groups of the collective genus Lacerta show a bush-like topology in the inferred Bayesian tree, suggesting rapid radiation. The composition of the subfamilies Eremiainae and Lacertinae as well as their phylogeography are discussed.

×

Sandveld lizards (genus Nucras) are widespread in southern Africa, but are generally secretive and poorly known. We examined 385 preserved specimens from five species of Nucras collected over a broad time span (104 years) and a geographic area covering most of South Africa and Swaziland. We had three main objectives: to test for sexual size dimorphism, to quantify male and female reproductive cycles, and to determine diet. In addition, we examined the importance of scorpions in the diet of Nucras based on previous studies reporting an unusually high incidence of scorpions in the diet of Kalahari N. tessellata. Males of all species except N. lalandii had significantly larger heads than females of the same body size, although females had significantly greater snout–vent length than males in three of the five species examined. The general reproductive pattern was for females to contain vitellogenic eggs during late spring and early summer. No females simultaneously contained oviductal and vitellogenic eggs, which suggests that females produce only one clutch per season. Clutch size was unrelated to female body size in all species examined. Female follicle volume generally coincided with male testicular volume, indicating for the most part synchronized reproductive cycles between the sexes. Testis volume was generally highest during spring–early summer, with only N. holubi showing a second peak in autumn. We recorded 15 arthropod orders in the diet of Nucras. All species feed on invertebrates, primarily insects, and, to a lesser degree, spiders and centipedes. Termites, grasshoppers, and beetles both numerically and volumetrically dominated their diet. We found no evidence that scorpions form a major part of the diet of any of these five species of Nucras, including N. tessellata from biomes outside the Kalahari Desert.

×

Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World-wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.

×

Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.