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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).

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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.

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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).

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A striking new sandveld lizard of the Nucras tessellata group is described from the Lambert’s Bay Strandveld of the Western Cape Province, South Africa. It is sister to the clade N. livida + N. tessellata, and is phenetically most similar to N. tessellata, from which it differs in its more elongate body and possibly increased number of presacral vertebrae and patternless orange dorsal coloration. The form elegans, described as a species by Andrew Smith (1838), but treated as an infrasubspecific variant by Broadley (1972), also exhibits weak patterning, but is likely a regional color variant. Nucras aurantiaca sp. nov. is the ninth member of the genus found in southern Africa. Its discovery in the well-collected coastal Western Cape suggests that further herpetofaunal surveys are needed in this region, which is threatened by agricultural activity and tourism-related development.

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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.

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

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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.

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This review summarises the current status of our knowledge of Angolan reptile diversity, and places it into a historical context of understanding and growth. It is compared and contrasted with known diversity in adjacent regions to allow insight into taxonomic status and biogeographic patterns. Over 67% of Angolan reptiles were described by the end of the nineteenth century. Studies stagnated dur- ing the twentieth century but have increased in the last decade. At least 278 reptiles are currently known, but numerous new discoveries have been made during recent surveys, and many novelties await description. Although lizard and snake diversity is currently almost equal, most new discoveries occur in lizards, particularly geckos and lacertids. Poorly known Angolan reptiles and others from adjacent regions that may occur in the country are highlighted. Most endemic Angolan reptiles are lizards and are associated with the escarpment and southwest arid region. Identification of reptile diversity hotspots are resolving but require targeted surveys for their delimitation and to enable protection. These include the Kaokoveld Centre of Endemism, Angolan Escarpment and the Congo forests of the north. The fauna of Angola remains poorly known and under-appreciated, but it is already evident that it forms an important centre of African reptile diversity and endemism.

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A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.

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A collection of reptiles and new locality records are reported from the littoral Southern Namib Desert, Northwestern Cape Province, Republic of South Africa. The new localities, by species, are: Chersina angulata, Swartlintjies River; Chondrodactylus angulifer, Swartlintjies River and Kleinsee; Agama atra, Papendorp, Swartlintjies River, and Kleinsee; Chamaeleo pumilus, MacDougall`s Bay; Mabuya occidentalis, Hondeklipbaal, Swartlintjies River, and Kleinsee; Acontias lineatus, Hondeklipbaal; Nucras tessellata, Kleinsee, and MacDougall`s Bay; Eremias capensis, Papendorp, Hondeklipbaal, and Kleinsee; Meroles knoxii, Elandsbaal; Meroles ctenodactyla, Papendorp, and Kleinsee; Psammophylax rhombeatus, Elandsbaal; Psammophis crucifer, Elandsbaal; and Naja nivea, Kreefbaal, and Kleinsee, 1978.

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A collection of reptiles and new locality records are reported from the littoral Southern Namib Desert, Northwestern Cape Province, Republic of South Africa. The new localities, by species, are: Chersina angulata, Swartlintjies River; Chondrodactylus angulifer, Swartlintjies River and Kleinsee; Agama atra, Papendorp, Swartlintjies River, and Kleinsee; Chamaeleo pumilus, MacDougall`s Bay; Mabuya occidentalis, Hondeklipbaal, Swartlintjies River, and Kleinsee; Acontias lineatus, Hondeklipbaal; Nucras tessellata, Kleinsee, and MacDougall`s Bay; Eremias capensis, Papendorp, Hondeklipbaal, and Kleinsee; Meroles knoxii, Elandsbaal; Meroles ctenodactyla, Papendorp, and Kleinsee; Psammophylax rhombeatus, Elandsbaal; Psammophis crucifer, Elandsbaal; and Naja nivea, Kreefbaal, and Kleinsee, 1978.

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We discuss three aspects of the thermal biology of lacertid lizards. First, we provide an overview of the available data on field body temperatures (Tb), the thermal sensitivity of various performance functions and selected body temperatures in different species of lacertid lizards. We also briefly summarise information on the mechanisms of thermoregulation. Second, we discuss recent developments to estimate the »precision« of thermoregulation, and the contribution of distinct behavioural mechanisms. Finally, we revise available evidence for the existence of evolutionary adjustments of thermal characteristics in lacertid lizards. Existing studies have mainly dealt with within- and among-species differences in thermoregulatory behaviour (selected temperatures) and thermal physiology of adults (optimal temperatures, heating rates). Available data provide only limited evidence for clear-cut evolutionary shifts in thermal physiology characteristics along climatic gradients.

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Namibe Province is the southernmost province of Angola and, as the result of sever- al expeditions from the nineteenth century to the present, it is one of the most her- petofaunally well-known areas of the country. The Province harbors a high diversi- ty of amphibians and reptiles, including roughly one-third of the reptile taxa report- ed for Angola as a whole. In this paper we present the results of a joint herpetologi- cal expedition to Namibe Province in 2013 by the California Academy of Sciences and the Instituto Nacionalda Biodiversidade e Áreas de Conservação, as well as a synoptic list of all the herpetological bibliographic records for the taxa known from the Province. A total of 37 herpetological taxa was collected, including at least three (then) undescribed species, two new country records, and new records for rarely cited taxa in Angola. These taxa belong to four amphibian genera and 15 reptile gen- era. Species accounts are provided for each of the species collected. We also highlight biogeographic patterns, conservation issues, and possible future paths for the explo- ration and knowledge of the herpetofauna of Namibe.

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Most lacertids are active foragers, but intrafamilial variation in foraging mode is greater than in most lizard families. We collected data on eight species of African lacertids to assess this variation. Both active and ambush foraging occurred within Pedioplanis and Meroles. Meroles ctenodactylus had a proportion of time moving and proportion of attacked prey detected while moving intermediate to those for actively foraging and ambushing Pedioplanis, but its number of movements per minute was exceptionally high. This species has a unique mixed foraging mode. Like active foragers, it seeks food by tongue-flicking while moving and spends a high percentage of the time moving. Like ambush foragers, it searches visually for prey during pauses between movements. Our findings confirm published data on four Kalahari lacertids. We discuss the history of foraging modes in advanced lacertids.

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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.

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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.

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The available material of the collection of reptiles and amphibians collected and described from 1831 by Andrew Smith, is re-examined to determine in how far it may be regarded as typical. A complete list of the new species described by Smith, together with notes and comments pertaining thereto, is given. Illustrated with a plate on the type specimen of Pelusios sinuatis.

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An annotated checklist of indigenous and potentially indigenous Namibian terrestrial, aquatic and marine reptiles is presented. The purpose is to serve as an interim description of Namibian reptile diversity, to establish a taxonomic and biogeographical baseline, and as a preliminary review of the conservation status of Namibian reptiles. Two hundred and forty species of indigenous reptiles are presently known to occur in Namibia. These species comprise an array of approximately 265 described (but not always recognized) taxa, several of which are probably unwarranted. Species accounts are presented for all these species. Four accounts are for new species currently being described. Nineteen species have not yet been recorded from Namibia, but are expected to (accounts given) and another 6 species are less likely to occur (no accounts given). Full accounts are given for the 17 species which have been formally recorded in the past, but the lack of recent evidence suggests that the species is now locally extinct, the original report erroneous, or the species’ occurred as vagrants. Four additional species had been included on various published lists in the past, but have never been formally documented, no specimens are known to exist, and it is unlikely that the species would occur today even as vagrants (no accounts given). In total, 276 species-accounts are presented. Each account cites the original reference and type locality for each taxon, and a short description of the Namibian distribution. Emphasis is placed on Namibian and international legal and conservation status. Eighty-five species (33%) were found to be of local conservation concern. Gaps in knowledge (e.g. taxonomy, biogeography, and conservation status), where future research should be directed, are noted.

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The herpetofauna of the southern Kalahari has mixed affinities, as this area lies on a rainfall gradient in a critical area where a transition between the arid south-west and the moister north-east takes place. As the variation in substrate type is relatively limited, the effect of the rainfall gradient appears to influence and determine the range limits of many taxa in both directions, resulting in an area in which of 55 recorded reptiles, 11 western taxa overlap or form a parapatric zone with 25 eastern taxa, while the remaining taxa are endemic or wideranging.

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In recent years there has been an exponential increase in the amount of published DNA sequences, as it has become one of the most popular sources of data for reconstructing phylogenies. A number of studies have attempted to resolve the relationships of groups of lacertids using mitochondrial DNA sequence data, in particular Podarcis and Gallotia (e.g. THORPE et al., 1994; 1996; GONZALEZ et al., 1996; FU et al., 1997; HARRIS, 1997; HARRIS et al., 1998a; 1999; CASTILLA et al., 1999a,b). Reassessment of these data suggest some alternative phylogenies to those previously published, and also highlight possible problems. Two studies have attempted to unravel the phylogeny of the whole family using mitochondrial DNA sequences (FU, 1998; HARRIS et al., 1998b). By combining these essentially congruent data sets, and reanalysing levels of support for the phylogenies produced, a more complete estimate of relationships across the family is produced. This is then compared to alternative sources of evidence, primarily morphological (ARNOLD, 1989), biochemical (MAYER & BENYR, 1994) and karyological (OLMO et al., 1993). Reasons for discrepencies are discussed, as are the relative advantages and disadvantages of the different sources of phylogenetic inference.

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DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.

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Namibia is mostly an arid and semi-arid country with a high number of reptile and fewer amphibian species. We review the herpetological literature dealing with Namibian species over the past fifty years, and provide up-to-date amphibian and reptile accounts using a widely accepted taxonomy and nomenclature. We critically discuss species accounts, draw attention to the historical development of species inventories for the country, and indicate species endemism for Namibia and the Namib Desert. In Namibia, the lizard families Gekkonidae, Lacertidae, and Scincidae have undergone adaptive radiations and are species-rich. This also applies to the snake family Psammophiidae. Areas of herpetological research that have received most attention are systematics (with its disciplines faunistics (area inventories), taxonomy, and phylogeny), ecology, and physiology. The former is indicative of early stages of herpetological research such as area inventories and the subsequent analyzes of the collections. The latter two were largely enabled by (1) species highly adapted to life in the hyper-arid Namib Desert, and (2) by the accessibility of these species in the Namib Desert through the infrastructure provided by the Gobabeb Research and Training Center. The majority of the eco-physiological research has focused on three highly psammophilus, diurnal lizard species; Meroles anchietae, M. cuneirostris, and Gerrhosaurus skoogi, whilst diverse geckos form the basis of eco-morphological studies. The concentration of research localities around cities and the Gobabeb Research and Training Center is characteristic for opportunistic research. Geographic centers of herpetological research have been the central Namib Desert (i.e. Gobabeb), and areas around Swakopmund and Windhoek. Extensive parts of Namibia remain barely touched. Herpetological publication frequency has been approximately the same since its beginning in the early 1800`s until the 1970`s. The period between 1986 and 2003 experienced a remarkable increase of publication activity that has slightly subsided around 2004 and picked up again in recent years. Recent conservation related studies investigate the impact of overgrazing with land degradation and water related issues such as canals and hydroelectric dam projects on herpetological communities. In the near future the impact of mining, especially Uranium mining in the Namib Desert, and the effects of climate change with the predicted drying and warming will demand increased attention. Advances in biotechnology with ever-increasing amounts of data and decreasing cost have and will progressively enable advances in traditional disciplines like taxonomy, phylogeny, and systematics. Additionally, these technologies will increasingly empower the newer disciplines of molecular ecology and conservation biology in Namibia. Annotated, updated species checklists highlight Namibian and Namib diversity and endemicity, and also direct researchers to the numerous taxonomic problems that still confound full understanding of the region`s herpetofauna.

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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.

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Background: Although current molecular clock methods offer greater flexibility in modelling evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate the evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates. Results: According to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic. Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges.

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Closely related lacertid lizards (Eremias, Nucras) in the Kalahari desert differ in patterns of foraging behaviour. Some species are relatively sedentary (‘sit-and-wait’) whereas others are more active (‘widely-foraging’) predators. We determined whether whole-animal locomotor capacities (cruising endurance on a treadmill, initial speed and maximum burst speed in a racetrack, and sprint endurance in a torus-shaped track) correlated with interspecific differences in foraging behaviour. Two of three widely-foraging species had greater cruising endurance, graater sprint endurance, but lower burst speed than did a sit-and-wait species. However, the two species that sprinted quickly also had limited endurance, and vice versa. Pre-feeding negatively influenced endurance but not sprint capacity. Theoretical models of foraging behaviour should recognize that ectotherms have limited endurance, that there can be a trade-off between speed and endurance, and that pre-feeding can reduce some aspects of locomotor capacity.

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We discuss seasonal variation in thermoregulatory behavior and its consequences on body temperature for 12 species of diurnal lizards in the southern Kalahari semidesert of Africa and also evaluate several methods of attempting to document thermoregulatory behavior using a descriptive data base. Lizards vary time of activity among seasons, which limits the variation in ambient conditions actually experienced. Ground-dwelling lizards and probably arboreal lizards move nonrandomly with respect to sun and shade; thus the percentage of lizards in sun in inversely proportional to air temperature. Arboreal lizards shift to higher perches at midday in summer and to logs or ground in winter thus decreasing and increasing incident heat loads, respectively. Both juveniles and adults of 3 species, only juveniles of 2 species, and only adults in 1 species are active in winter: both adults and juveniles of 6 species brumate [= hibernate]. Mean body temperature (T`b) varies within days and among months and is positively correlated with corresponding mean air temperature (T`a) in almost all species. Nonetheless, correlation and regression analysis suggests that thermoregulatory behaviors reduce the impact of variations in ambient conditions on Kalahari lizards. The mean T`b of different species reflect evolutionary relationships. In summer, mean T`b is proportional to the percentage of lizards in sun and with the tendency of lizards to be active only in summer. Thus, lizards with inferred low optimal temperatures are active during more months of the year.

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Desert lizards are typically either widely foraging or sit-and-wait predators, and these foraging modes are correlated with major differences in ecology. Foraging mode is related to the type of prey eaten by lizards. Widely foraging lizards in the Kalahari desert, the Western Australian desert, and the North American desert generally eat more prey that are sedentary, unpredictably distributed, and clumped (e.g., termites) or that are large and inaccessible (inactive scorpions) than do sit-and-wait lizards. In contrast, sit-and-wait lizards eat more prey that are active. Foraging mode also appears to influence the types of predators that in turn eat the lizards. For example, a sit-and-wait snake eats predominately widely foraging lizards. Crossovers in foraging mode thus exist between trophic levels. Widely foraging lizards may also encounter predators more frequently, as suggested by analyses of relative tail lengths; but tail break frequencies are ambiguous. Daily maintenance energetic expenditures of widely foraging lizards appear to be about 1.3-1.5 times greater than those of sit-and-wait lizards in the same habitats, but gross food gains are about 1.3-2.1 times greater. Widely foraging species also have lower relative clutch volumes, apparently in response to enhanced risks of predation. Foraging mode within one species varies with changes in food availability. Physiology, morphology, and risk of predation might generally restrict the flexibility of foraging mode. Because foraging mode constrains numerous important aspects of ecology, any general model of foraging velocity must be complex.

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The performance of ectotherms integrated over time depends in part on the position and shape of the distribution of body temperatures (Tb) experienced during activity. For several complementary reasons, physiological ecologists have long expected that Tb distributions during activity should have a long left tail (left‐skewed), but only infrequently have they quantified the magnitude and direction of Tb skewness in nature. To evaluate whether left‐skewed Tb distributions are general for diurnal desert lizards, we compiled and analysed Tb (∑ = 9,023 temperatures) from our own prior studies of active desert lizards in three continents (25 species in Western Australia, 10 in the Kalahari Desert of Africa and 10 species in western North America). We gathered these data over several decades, using standardized techniques. Many species showed significantly left‐skewed Tb distributions, even when records were restricted to summer months. However, magnitudes of skewness were always small, such that mean Tb were never more than 1°C lower than median Tb. The significance of Tb skewness was sensitive to sample size, and power tests reinforced this sensitivity. The magnitude of skewness was not obviously related to phylogeny, desert, body size or median body temperature. Moreover, a formal phylogenetic analysis is inappropriate because geography and phylogeny are confounded (i.e. are highly collinear). Skewness might be limited if lizards pre‐warm inside retreats before emerging in the morning, emerge only when operative temperatures are high enough to speed warming to activity Tb, or if cold lizards are especially wary and difficult to spot or catch. Telemetry studies may help evaluate these possibilities.

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We discuss seasonal variation in thermoregulatory behavior and its consequences on body temperature for 12 species of diurnal lizards in the southern Kalahari semidesert of Africa and also evaluate several methods of attempting to document thermoregulatory behavior using a descriptive data base. Lizards vary time of activity among seasons, which limits the variation in ambient conditions actually experienced. Ground-dwelling lizards and probably arboreal lizards move nonrandomly with respect to sun and shade; thus the percentage of lizards in sun is inversely proportional to air temperature. Arboreal lizards shift to higher perches at midday in summer and to logs or ground in winter thus decreasing and increasing incident heat loads, respectively. Both juveniles and adults of 3 species, only juveniles of 2 species, and only adults in I species are active in winter: both adults and juveniles of 6 species brumate [= hibernate]. Mean body temperature (T,) varies within days and among months and is positively correlated with corresponding mean air temperature (T,) in almost all species. Nonetheless, correlation and regression analysis suggests that thermoregulatory behaviors reduce the impact of variations in ambient conditions on Kalahari lizards. The mean Tb of different species reflect evolutionary relationships. In summer, mean T i s proportional to the percentage of lizards in sun and with the tendency of lizards to be active only in summer. Thus, lizards with inferred low optimal temperatures are active during more months of the year.

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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.

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The present work constitutes an historical atlas of all known bibliographic records of amphib- ians and reptiles of Angola. It is the first attempt to compile in a single document all the records scattered through hundreds of publications, published from the first half of the nineteenth century to the present day, and provide a critical taxonomic revision of the herpetofauna of the country. An introductory text discusses the Angolan socio-political and physiographic landscape, the history of herpetological research in the country, and the diversity, distribution and endemism of Angolan her- petofauna. It also provides a summary of the conservation concerns surrounding the herpetological fauna. For each taxon noted in the literature we provide a detailed taxonomic account, including data on the original description, its IUCN assessment status, global distribution, georeferenced records for each known occurrence in Angola (accompanied by a point locality map), and taxo- nomic and distributional notes. A synoptic list of all documented Angolan amphibian and reptile species is provided as are lists of taxa previously erroneously assigned to the Angolan fauna. We recognize 117 species of frogs and 278 species of reptiles as occurring in Angola, although many of these represent species complexes or are currently recognized under names of convenience pending taxonomic evaluation. In one instance we provide a replacement name, Trachylepis monardi nom. nov., to deal with an instance of secondary homonymy. The Atlas is intended to be a working reference for both current research and conservation planning, as well as a catalyst for future work.The interest shown by specialists in the study of Angolan specimens dates back more than one hundred years. Since the nineteenth century, books and papers have been published on the topic as the result of these investigations. The first major bibliographic reference about the Angolan herpetofauna, Herpétologie d’Angola et du Congo by José Vicente Barbosa du Bocage, dates from 1895. Many researchers and publications followed, greatly enriching the scientific bibliography on the topic. However, the scattered nature of the published works, as well as the dearth of available specimens from earlier times, has made it difficult for researchers, scholars or curious amateurs to access to this information. Because of these difficulties, and to facilitate the access to available data on the Angolan herpetofauna, the Ministry of Environment, through its National Institute of Biodiversity and Conservation Areas, decided to present, in a single volume, all the available scientific records published so far on the diversity of amphibians and reptiles. Such a task, which has its basis in the National Strategy and Action Plan for Biodiversity and Conservation Areas of Angola, was dependent upon the helpful collaboration of important international partners. The present Atlas will be of inestimable assistance to the study of the Natural History of Ango- la, not only because of its standardized taxonomic and geographical accounts with distribution maps for all of the amphibians and reptiles known in Angola, but also because of its competent approach to the physiography, climate, biomes, historical herpetological research in Angola, conservation, and other topics. The thoughtful way the book is arranged, with straightforward text, clear tables and interesting figures, makes it easy for even laymen to read and understand. At 501 pages long, profusely documented and richly illustrated, this Atlas provides a grand guided tour through the diversity of the Angolan herpetofauna, from the most remote and spectac- ular places of the country. Through mountains and escarpments, plains and valleys, savannas and dense forests, rivers and streams, lakes and lagoons, this work encompasses the entire country, aking us on an enlightening tour of knowledge. The methodical labor and persistence of the specialists who produced the present work, ensure that it not only contributes to scientific history, through its compilation of research already completed, but also suggests future directions for researchers and scholars devoted to the topic. Further, this substantial volume will also inform the national authorities with respect to environ- ment protection and conservation policies. Given all this, one can see that the present work, reflecting the contributions of the men and women who created it, is destined to become a standard reference to the scientific literature of the country, especially in regard to this part of its fauna. The quality of the scientific information poured into the following pages certainly qualifies the authors of this accurate work as leading herpetologists and places them among the important researchers to have served Angola in the course of its scientific history.

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13 species of the genera Aporosaura, Meroles, Pedioplanis, and Heliobolus horn Namibia and additionally Pedioplanis burcheUi from the Republic of South-Africa have been investigated proteinelectrophoretically concerning 14 genetic loci. Pedioplanis and Meroles turned out to be sister groups, Heliobolus and Aporosaura are standing further apart. The su bgenera of Meroles (Meroles s. str. and Saurites) could not be parted distinctly. The morphological correspondence between Aporosaura and Saurites has been interpreted as convergent adaptions to their habitat.

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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.

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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.

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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.

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Aim Temperature influences most components of animal ecology and life history – but what kind of temperature? Physiologists usually examine the influence of body temperatures, while biogeographers and macroecologists tend to focus on environmental temperatures. We aim to examine the relationship between these two measures, to determine the factors that affect lizard body temperatures and to test the effect of both temperature measures on lizard life history. Location World-wide. Methods We used a large (861 species) global dataset of lizard body temperatures, and the mean annual temperatures across their geographic ranges to examine the relationships between body and mean annual temperatures.We then examined factors influencing body temperatures, and tested for the influence of both on ecological and life-history traits while accounting for the influence of shared ancestry. Results Body temperatures and mean annual temperatures are uncorrelated. However, accounting for activity time (nocturnal species have low body temperatures), use of space (fossorial and semi-aquatic species are ‘colder’), insularity (mainland species are ‘hotter’) and phylogeny, the two temperatures are positively correlated. High body temperatures are only associated with larger hatchlings and increased rates of biomass production. Annual temperatures are positively correlated with clutch frequency and annual longevity, and negatively correlated with clutch size, age at first reproduction and longevity. Main conclusions Lizards with low body temperatures do not seem to have ‘slower’ life-history attributes than species with high body temperatures. The longer seasons prevalent in warm regions, and physiological processes that operate while lizards are inactive (but warm enough), make environmental temperatures better predictors of lizard life-history variation than body temperatures. This surprisingly greater effect of environmental temperatures on lizard life histories hints that global warming may have a profound influence on lizard ecology and evolution.

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Point distribution data were used to evaluate the biogeographical influence of the arid Tankwa Karoo Basin on the distribution of reptiles in the south-western districts of South Africa. Under-representation of the Tankwa Karoo in the dataset required an additional field survey of this region. Prior to the survey, available records from the Tankwa Karoo Basin represented only 13 reptile species. A total of 36 species (24 lizards, nine snakes, three chelonians) was recorded during our survey of this area. Turnover across the Basin is high, species richness is lower than in surrounding mountainous areas and there are no species endemic to the area. The Tankwa Karoo Basin acts as a dispersal barrier for many reptile species occurring in the surrounding more mesic areas. At the same time, the ranges of a number of typical northern, arid-adapted species extend southward along the Tankwa plains. A number of species range extensions in the region are reported.

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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.

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From 12 to 18 species of desert lizards occur in ecologic sympatry on 10 study areas in the Kalahari desert of southern Africa. The number of species of diurnal lizards is similar to the number in the North American deserts, while the nocturnal component of the saurofauna is more comparable to that occurring in the Australian deserts. The between-habitat component of species density is similar in the American and the Kalahari deserts, but is markedly greater in the Australian desert. A variety of climatic and vegetational statistics are examined for the 10 areas, and correlations with lizard species density are discussed.

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Four lacertid lizards, Pedioplanis laticeps, P. lineo-ocellata, Meroles knoxii and Nucras tessellata, occur sympatrically on the arid plains of the Tankwa Karoo Basin in South Africa. The aim of this study was to evaluate the significance of foraging strategy in resource partitioning among the four species, allowing them to co-occur in a structurally simple system with a limited number of potential niches. Previous workers already identified P. lineo-ocellata and M. knoxii as sit-and-wait foragers and N. tessellata as an active forager. We recorded data on three foraging variables: movements per minute, proportion of time spent moving, and proportion of attacks on prey whilst moving, for juveniles and adults of P. laticeps. By comparing the foraging data obtained for P. laticeps to those for other lacertid species, we were able to demonstrate that adult P. laticeps are ambush foragers. We also noted a significant ontogenetic shift in foraging behaviour in P. laticeps, and, due to a significantly higher frequency of short brief movements, we classified juveniles as mixed foragers. The sharing of an ambush foraging strategy by at least three of the four lacertid species co-occurring on the Tankwa plains, suggests considerable overlap along the trophic dimension of ecological space. This overlap presumably promotes occupation of separate microhabitats by the three ambush foragers in the Tankwa Karoo Basin.

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Four lacertid lizards, Pedioplanis laticeps, P. lineo-ocellata, Meroles knoxii and Nucras tessellata, occur sympatrically on the arid plains of the Tankwa Karoo Basin in South Africa. The aim of this study was to determine whether the four species are spatially separated in terms of the microhabitat each species occupies, allowing them to occur in sympatry in this structurally simple system with a limited number of potential niches. The habitat preferences of the four species were investigated on the farm Gansfontein in the southwestern parts of the Tankwa Karoo Basin. The area was surveyed by walking transects through different habitat types and recording all active lizards. For each lizard observed, the species was identified and a photograph was taken of its microhabitat. At the end of the sampling period the microhabitats in which lizards were found were sorted into distinct types on the basis of substrate composition. During the survey, only one individual of N. tessellata was recorded, but sample sizes for the other three species were sufficiently large to provide a clear perspective of habitat segregation among the three species. The results showed that the three species are spatially separated at the study site, with P. lineo-ocellata being restricted to rocky areas, P. laticeps preferring gravel substrates and M. knoxii frequenting sandy areas. Although only anecdotal information is available, all indications are that the observed spatial separation of the three species is mainly the result of historical adaptation to the different substrate types.

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Locomotion is important to animals because it has direct implications for fitness through its role in predator escape, prey capture, and territory defence. Despite significant advances in our understanding of animal locomotion, studies exploring how substrate properties affect locomotor performance remain scant. In the present study, we explore how variation in substrate (sand, slate, cork) affects locomotor performance in lacertid lizards that differ in morphology. Moreover, we explore whether substrate effects are the same for different types of locomotor performance (speed, acceleration, and stamina). Our results show that the substrate affected most types of locomotor performance studied but not always in the same way. Although substrate effects were species-dependent for the maximal speed over 50 cm and the distance run to exhaustion, this was not the case for acceleration capacity. These results suggest that substrate texture differentially affects burst performance vs. longer duration measures of locomotor performance. Finally, straightforward relationships between habitat use and the substrate on which performance was maximized were not observed. This suggests that the evolution of locomotor capacity is complex and that animals may show compromise phenotypes allowing them to deal with a variety of substrates in their natural environment.

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Evolutionary changes in foraging style are often believed to require concurrent changes in a complex suite of morphological, physiological, behavioural and life-history traits. In lizards, species from families with a predominantly sit-and-wait foraging style tend to be more stocky and robust, with larger heads and mouths than species belonging to actively foraging families. Here, we test whether morphology and foraging behaviour show similar patterns of association within the family Lacertidae. We also examine the association of bite force abilities with morphology and foraging behaviour. Lacertid lizards exhibit considerable interspecific variation in foraging indices, and we found some evidence for a covariation between foraging style and body shape. However, the observed relationships are not always in line with the predictions. Also, the significance of the relationships varies with the evolutionary model used. Our results challenge the idea that foraging style is evolutionarily conservative and invariably associated with particular morphologies. It appears that the flexibility of foraging mode and its morphological correlates varies among lizard taxa.