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Bicuar National Park (BNP) is a protected area in southwestern Angola where biodiversity has been poorly studied. BNP is located on the Angolan plateau on Kalahari sands, in a transition zone between the Angolan Miombo Woodland and the Zambezian Baikiaea Woodland ecoregions. Herpetological surveys were conducted in BNP and surrounding areas, through visual encounter surveys, trapping, and opportunistic collecting of specimens from 2015 to 2018. The regional herpetofauna is described here based on these surveys, literature records, and additional unpublished records. In total, 16 amphibian, 15 lizard, 18 snake, two testudine, and one crocodilian species were observed from the recent surveys, and in combination with historical records the species counts are 21, 36, 32, four, and one species for these herpetofauna groups, respectively. Important observations include the first record of Xenocalamus bicolor bicolor (Günther, 1868), the second records of Sclerophrys poweri (Hewitt, 1935) and of Amblyodipsas ventrimaculata (Roux, 1907), and the fourth record of Monopeltis infuscata (Broadley, 1997) for Angola. Additionally, the type locality of Hyperolius benguellensis (Bocage, 1893) is discussed. A part of the material could not be confidently identified to species level, reflecting the taxonomic uncertainty associated with the Angolan herpetofauna. Fossorial herpetofauna was well represented, reflecting adaptation to sandy soils, the dominant substrate in the area. The likely presence of endemic and poorly known species in BNP reinforces the importance of the park for the conservation of Angolan biodiversity. Further surveys are necessary for a more comprehensive understanding of the park’s fauna and biogeographic affinities, and to improve conservation planning.

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A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.

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Today Ichnotropis is still a poorly understood genus. This is an attempt to initialize some change herein, by providing an extensive bibliographic history of the genus, together with some remarks on the described taxa. Necessary future research is suggested.

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On October 16, 2019 an image of Ichnotropis bivittata, taken on May 12, 2018 in Cassongue county, Cuanza Sul province, Angola, was uploaded on iNaturalist by Rogério Ferreira for determination. This specimen may have some features of Ichnotropis microlepidota, in which case it would be the first time that this species has been observed since the species description by Marx (1956).

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Southeast Angola has remained poorly studied as the extensive wetlands make movement difficult, as do the wide-spread land mines scattered during the long civil war. Due to its biodiversity and scenic beauty the Okavango Delta of Botswana was declared a World Heritage Site in 2014. Its survival as a functioning ecosystem and the world’s only major inland delta depends on the waters that flow south from Angola. These wetlands form the catchments for three of southern central Africa’s major river systems, the Zambezi, Kwanza and Okavango drainages. The herpetological survey formed part of the National Geographic Okavango Wilderness Project (NGOWP), initiated to survey the extraordinary biodiversity of the region, and to assess the current state of these important wetlands and identify any impacts on them. This is a personal overview of the trip, and the scientific details are given elsewhere (Conradie et al. 2016). Material is accessioned into the Port Elizabeth Museum (South Africa) and ISCED, Lubango (Angola) collections. The NGOWP surveys (2015–17) to the watersheds form the most extensive regional herpetological surveys in the history of Angola. The survey documented 110 species from the region, comprising 38 snakes, 32 lizards, five chelonians, a single crocodile and 34 amphibians. Seven species were new for Angola, and others were added during the 2016–17 trips. A recur- rent feature of these surveys was the regular discovery of specimens that could not be assigned confidently to known taxa. Numerous new species have been identified; some have been described, for others research is ongoing.

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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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The amphibian fauna of northwestern Zambia was quite well known, but no systematic collection of reptiles had ever been made. This was rectified when I spent the period 23 September to 12 October 1990 based at Sakeji School near Ikelenge and collected 258 reptiles and 77 amphibians. Five reptile species are new for Zambia, i.e. Adolfus africanus, Causus lichtensteinii, Limnophis bicolor, Thrasops j. jacksonii and Rhamnophis aethiopissa ituriensis.

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A review of the reptiles recorded from southeast Katanga Province in the Democratic Republic of Congo, employing the evolutionary species concept, indicates that there are at least 119 species in the area (7 chelonians, 37 lizards, 4 amphisbaenians, 69 snakes and 2 crocodilians), 15 of which are endemic (12.6 %). Analysis of zoogeographical affinities shows that this reptile fauna is derived from all directions, with forest forms entering from the north accounting for 14.4 % and savanna forms entering from the east contributing 22 %.

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We present an annotated catalog of the type specimens of amphibians and reptiles in the collections of the Museu de História Natural da Universidade do Porto in Portugal. These specimens, all from present-day Angola, formed the basis of taxonomic descriptions by both José Júlio Bethencourt Ferreira and José Vicente Barbosa du Bocage in the latest 19th and early 20th century. We provide details for all type specimens and summarize the history and taxonomy for each species. Specimens of Rappia bocagei var. maculata and Typhlops bocagei could not be located during our survey, and we believe these to be lost. The collections at the University of Porto contain type specimens of one snake, Typhlops boulengeri, and eight frogs, Arthroleptis carquejai, Hylambates bocagei var. leucopunctata, Rappia platyceps var. angolensis, Rappia bivittata, Rappia fasciata, Rappia nobrei, Rappia osorioi, and Rappia seabrai. Of these, only two are currently recognized: Afrixalus osorioi and Arthroleptis carquejai.

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The herpetological collections of the Museu do Dundo in Lunda Norte Province, northeast Angola, are among the most important in southern Africa and represent one of the largest collections of Angolan amphibians and reptiles in the world. The collection comprises more than 2,750 preserved specimens, including type specimens of taxa described by Raymond F. Laurent during the 1950s and 1960s,when he was affiliated with the Musee royal del’Afrique centrale (RMCA) in Tervuren, Belgium, and the Museum of Comparative Zoology (MCZ) at Harvard University, Cambridge, Massachusetts, where portions of these type series were also deposited. We provide details for all type specimens and summarize the history and taxonomy for each species represented in the type collection. The collections contain type specimens of 28 amphibian and reptile species, including seven snakes: Typhlops praeocularis lundensis, Dispholidus typus punctatus, Lycodonomorphus subtaeniatus, Lycophidion hellmichi, Gonionotophis brussauxi prigoginei, Prosymna ambıgua brevis, and Elapsoidea decosteri huilensis; 13 lizards: Rhoptropus boultoni montanus, Rhoptropus taeniostictus, Hemidactylus nzingae, Lygodactylus tchokwe, Cordylus vittifer machadoi, Chamaesaura anguina oligopholis, Gerrhosaurus bulsi, Nucras scalaris, Ichnotropis bivittata pallida, Mabuya bayonii huilensis, Mabuya ivensii septemlineata, Trachylepis raymondlaurenti, and Eumecia anchietae major; one amphisbaenian: Monopeltis vanderysti vilhenai; and seven frogs: Hyperolius machadoi,Hyperolius marmoratus alborufus, Hyperolius vilhenai, Ptychadena grandisonae, Ptychadena loveridgei, Ptychadena perplicata, and Ptychadena upembae machadoi. The typespecimens of the snake Xenocalamus bicolor machadoiwere not found in the collections. A brief history of the museum and remarks on the overall herpetological collections are also provided. We also note additional information about the related type material of these taxa at the MCZ.

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Angola`s herpetofauna has been neglected for many years, but recent surveys have revealed unknown diversity and a consequent increase in the number of species recorded for the country. Most historical Angola surveys focused on the northeastern and southwestern parts of the country, with the southeast , now comprising the Kuando-Kubango Province, neglected. To address this gap a series of rapid biodiversity surveys of the upper Cubango-Okavango basin were conducted from 2012‒2015. This report presents the results of these surveys, together with a herpetological checklist of current and historical records for the Angolan drainage of the Cubango, Cuito, and Cuando Rivers. In summary 111 species are known from the region, comprising 38 snakes, 32 lizards, five chelonians, a single crocodile and 34 amphibians. The Cubango is the most western catchment and has the greatest herpetofaunal diversity (54 species). This is a reflection of both its easier access, and thus greatest number of historical records, and also the greater habitat and topographical diversity associated with the rocky headwaters. As a result of these surveys, five new species records were added to the Cubango catchment, 17 to the Cuito catchment, and nine to the Cuando catchment. Seven of the records for the eastern catchments, including three for the Cuito and four for the Cuando, were also new for Angola. Resumo.—A herpetofauna de Angola foi negligenciada durante muitos anos, mas vários levantamentos realizados recentemente revelaram uma diversidade desconhecia e um consequente aumento no número de espécies registadas para o país. A maior parte dos levantamentos históricos realizados em Angola focaram-se no nordeste e sudoeste do país, sendo o sudeste, que agora abrange a província do Kuando-Kubango, bastante negligenciado. Para preencher esta lacuna, foram realizados vários levantamentos de biodiversidade na bacia superior do Cubango-Okavango entre 2012 e 2016. Neste relatório são apresentados os resultados destes levantamentos, bem como uma lista de registos herpetológicos recentes e históricos para as bacias dos rios Cubango, Cuito, e Cuando. Resumidamente, conhecem-se 111 espécies para a região, incluindo 39 espécies de cobras, 32 espécies de lagartos, 5 espécies de quelónios, apenas um crocodilo, e 34 espécies de anfíbios. A bacia hidrográfica do Cubango é a que está localizada mais a oeste, e tem a maior diversidade de herpetofauna (54 espécies). Isto reflecte por um lado a sua maior acessibilidade, e consequentemente um maior número de registos históricos, e ainda a maior diversidade topográfica e de habitat, associada às nascentes rochosas. Como resultado destes levantamentos, foram adicionadas cinco novas espécies à bacia do Cubango, 17 à do Cuito, e nove à do Cuando. Sete dos registos para as bacias a este, incluindo trois para o Cuito e quatro para o Cuando, são também novos registos para Angola.

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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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Molecular phylogenetic analyses of southern African lacertid lizards (Eremiadini) using mitochondrial and nuclear markers revealed two examples of generic assignments incompatible with monophyletic clades. Australolacerta Arnold 1989, a genus endemic to South Africa and to which two isolated species have been referred, is paraphyletic at the generic level. In addition, the species Ichnotropis squamulosa Peters 1854 was found to be embedded within the genus Meroles. To resolve the paraphyly in Australolacerta we erect a new genus, Vhembelacerta Edwards, Branch, Herrel, Vanhooydonck, Measey, & Tolley, gen. nov., to accommodate Lacerta rupicola FitzSimons 1933. To maintain a monophyletic Ichnotropis Peters 1854, Ichnotropis squamulosa Peters 1854 is transferred to Meroles Gray 1838, now named Meroles squamulosus comb. nov. Where necessary the genera affected by these actions are re-characterized.

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The Port Elizabeth Museum collection holds nearly 500 reptile specimens from Zambia and adjacent southeastern Democratic Republic of the Congo. These are reviewed, and biological and distribution data on 5 chelonians, 27 lizards and 38 snake species from Zambia are presented. These include information on 2 chelonians, 11 lizards and 19 snake species recorded from the poorly-known northern lVwinilunga District, Northwestern Province. Among the important findings are: the second largest specimen of and second Zambian locality for the dwarf terrapin, Pelusios nanus, with details on H`e first documented data on reproduction and sperm retention; the close proximity, without intergradation, of Lygodactylus heeneni and L. angularis suppOlting elevation of the former to specific status; a range extension of about 345km and the most northerly record (Chingola) for the gecko Lygodactylus chobiensis; a north-westerly range extension of more than 300km (to Chingola) for the gecko Hemidactylus mabouia (both these records may be translocations); the third and fourth reco]`ds for Zambia, and the most southern to date, for the gecko Pachydactylus tuberculosus; the first detailed biological information on the rare skink, Mabuya ivensii, which was first collected from Zanbia during these collections; asynchronous reproduction in Sakeji populations of both Mabuya maculilabris and M. wahlbergii; a southern range extension to Shimabala for the rare skink Eumecia anchietae; a range extension for the skink Lygosoma afrum to Sampfya town; the first record of lchnotropis capensis in the Copperbelt, and a north-westerly range extension of mon: than 300km to Chingola; the absence of asynchronous reproductive cycling between lchnotropis capensis and I bivittata in July at Sakeji; support for the specific status of Limnophis bangewoliCi,lS based on differences in colouration of the supralabials and subcaudals; new record sizes for both sexes of Dipsadoboa shrevei shrevei; a new record size for female Psammophis brevirostris leopardinus; and probable sympatry between Naja annulifera and its sister species N. anchietae at Livingstone.

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A lizard specimen recently collected from Gabon can not be assigned to any known species for the country. It is identified as Ichnotropis bivittata, a lacertid previouslnot reported for the country. The specimen is described in details and compared to related species. An older specimen examined by Boulenger (1921) was collected in the same area but not identified as from Gabon since the country was not always with the same borders and under the name `Congo français`.

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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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A commented lacertids list for Europe, Africa, the Middle East, including the Arabian Peninsula and Asia is given.

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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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We provide an updated, commented list of the reptiles of Gabon, with their occurrence in the country’s protected areas, national parks and reserves. This list is part of a book describing the current status of knowledge of the biodiversity of Gabon, including detailed descriptions and abundant illustrations of its ecosystems and protected areas.

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Presented is a compilation of new locality records with natural history data for 49 reptile species in Haut-Ogooué Province, southeastern Gabon, Equatorial Africa. One new snake species record is added to Gabon (Colubridae: Philothamnus hughesi), 14 new reptile species records to Haut-Ogooué Province, three new reptile records for Léconi Park, and 28 new reptile records for Lékédi Park. A predation case is reported of Naja melanoleuca preying on Dasypeltis fasciata and of Boaedon perisilvestris on a micromammal.

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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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Lacertidae from page 84.