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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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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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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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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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The lacertid genus Pedioplanis is a moderately speciose group of small-bodied, cryptically-colored lizards found in arid habitats throughout southern Africa. Previous phylogenetic work on Pedioplanis has determined its placement within the broader context of the Lacertidae, but interspecific relations within the genus remain unsettled, particularly within the P. undata species complex, a group largely endemic to Namibia. We greatly expanded taxon sampling for members of the P. undata complex and other Pedioplanis, and generated molecular sequence data from 1,937 bp of mtDNA (ND2 and cyt b) and 2,015 bp of nDNA (KIF24, PRLR, RAG-1) which were combined with sequences from GenBank resulting in a final dataset of 455 individuals. Both maximum likelihood and Bayesian analyses recover similar phylogenetic results and reveal the polyphyly of P. undata and P. inornata as presently construed. We con- firm that P. husabensis is sister to the group comprising the P. undata complex plus the Angolan sister species P. huntleyi + P. haackei and demonstrate that P. benguelensis lies outside of this clade in its entirety. The complex itself comprises six species including P. undata, P. inornata, P. rubens, P. gaerdesi and two previously undescribed entities. Based on divergence date estimates, the P. undata species complex began diversifying in the late Miocene (5.3 ± 1.6 MYA) with the most recent cladogenetic events dating to the Plio- cene (2.6 ± 1.0 MYA), making this assemblage relatively young compared to the genus Pedioplanis as a whole, the origin of which dates back to the mid-Miocene (13.5 ± 1.8 MYA). Using an integrative approach, we here describe Pedioplanis branchi sp. nov. and Pedioplanis mayeri sp. nov. representing northern populations previously assigned to P. inornata and P. undata, respectively. These entities were first flagged as possible new species by Berger-Dell’mour and Mayer over thirty years ago but were never formally described. The new species are supported chiefly by differences in coloration and by unique amino acid substitutions. We provide comprehensive maps depicting historical records based on museum specimens plus new records from this study for all members of the P. undata complex and P. husabensis. We suggest that climatic oscillations of the Upper Miocene and Pliocene-Pleistocene era in concert with the formation of biogeographic barriers have led to population isolation, gene flow restrictions and ultimately cladogenesis in the P. undata complex.

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Although reptile diversity in Africa is high, it is poorly represented in Angola, with just 257 species known. Despite its greater surface area and habitat diversity Angola has significantly lower lacertid lizard diversity than adjacent Namibia. This is particularly notable in African sand lizards (Pedioplanis), where 10 species (two endemic) are known from Namibia but only two are recorded from adjacent Angola. Pedioplanis benguelensis was described from Angola, but its taxonomic status is problematic and it was previously synonymised with P. namaquensis. All other Angolan Pedioplanis were referred to Namibian P. undata, although this taxon is now known to comprise a complex of at least five different species and the relationship of Angolan material to this complex has not been assessed. In this study, we investigated the phylogenetic placement of Angolan Pedioplanis using two mitochondrial (ND2 and 16S) and one nuclear (RAG-1) markers. A Bayesian analysis was conducted on 21 samples from Angola, combined with existing data for 45 individuals from GenBank and three additional samples from central Namibia. The phylogeny demonstrates that P. benguelensis is a valid species and that it is not the sister taxon to P. namaquensis with which it has been morphologically confused. In addition, Angolan lacertids previously referred to P. undata are not conspecific with any of the Namibian or South African species in that complex. Rather, there is strong support for the presence in Angola of additional species of Pedioplanis, which form a wellsupported sister clade to the P. undata complex (sensu stricto) of Namibia and two ofwhich are described herein. These discoveries highlight the need for further biodiversity surveys in Angola, as similar increases in species diversity in other Angolan taxa might be found given sufficient investment in biodiversity surveys.

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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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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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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 diversity of lacertid lizards in Africa is highest in the southern African subcontinent, where over two-thirds of the species are endemic. With eleven currently recognized species, Pedioplanis is the most diverse among the southern African genera. In this study we use 2200 nucleotide positions derived from two mitochondrial markers (ND2 and 16S rRNA) and one nuclear gene (RAG-1) to (i) assess the phylogeny of Pedioplanis and (ii) estimate divergence time among lineages using the relaxed molecular clock method. Individual analyses of each gene separately supported different nodes in the phylogeny and the combined analysis yielded more well supported relationships. We present the first, well-resolved gene tree for the genus Pedioplanis and this is largely congruent with a phylogeny derived from morphology. Contrary to previous suggestions Heliobolus/Nucras are sister to Pedioplanis. The genus Pedioplanis is monophyletic, with P. burchelli/P. laticeps forming a clade that is sister to all the remaining congeners. Two distinct geographic lineages can be identified within the widespread P. namaquensis; one occurs in Namibia, while the other occurs in South Africa. The P. undata species complex is monophyletic, but one of its constituent species, P. inornata, is paraphyletic. Relationships among the subspecies of P. lineoocellata are much more complex than previously documented. An isolated population previously assigned to P. l. pulchella is paraphyletic and sister to the three named subspecies. The phylogeny identifies two biogeographical clades that probably diverged during the mid-Miocene, after the development of the Benguella Current. This probably led to habitat changes associated with climate and, in conjunction with physical barriers (Great Escarpment), contributed towards speciation within the genus Pedioplanis.

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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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Lizards are a diverse clade in which one radiation consists entirely of sit-and-wait foragers and another consists of wide foragers. Lizards utilizing these two foraging modes are known to differ in diet, but little is known about how feeding morphology relates to diet and/or foraging mode. This study tested the hypothesis that skull morphology and biting performance are related to diet preference, and consequently, coevolve with foraging mode. Four species of lacertid lizard were studied because they vary in foraging mode, their phylogenetic relationships are known and they are well studied ecologically. Using an ‘ecomorphological’ approach, skull morphology and biting performance were quantified and mapped on to the phylogeny for the species. The results indicate that sit-and-wait species have shorter, wider skulls than the wide foraging species, and that all are significantly different in overall head shape. The sit-and-wait species had similar values for biting performance; however, clear phylogenetic patterns of covariation were not present between sit-and-wait and wide foraging species for either biting performance or skull morphology. Thus, skull morphology and performance have little influence on diet and foraging mode in these species. Instead it is likely that other factors such as seasonal prey availability and/or life history strategy shape foraging mode decisions.

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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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The genus Pedioplanis reaches its northernmost limit in western Angola, where it is represented by three species, Pedioplanis benguelensis, P. haackei and P. huntleyi. The taxonomic status of P. benguelensis remains problematic, mainly due to the vague original description and the loss of the original type material. Here we provide a revision of the Angolan representatives of the genus, with the description of a new species, Pedioplanis serodioi sp. nov., from the lowlands of southwestern Angola. Phylogenetic analyses using a combination of mitochondrial (16S and ND2) and nuclear (RAG-1) markers, as well as morphological data, support the recognition of the new species. For purposes of nomenclatural stability, we designate a neotype for P. benguelensis and provide motivation to correct the spelling of the specific epithet to “benguelensis”. The clarification of the status of P. benguelensis and the description of a new species contribute to a better understanding of the taxonomy and biogeography of the genus Pedioplanis, as well as the general biogeographic context of southwestern Angola, adding to the growing evidence in favor of the recognition of this region as a hotspot of lizard diversity and endemism. An updated key to the genus is also provided.

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The present paper of N. N. SZCZERBAK was originally published in Russian as `Katalog afrikanskih Jascurok` by the Academy of Sciences, Institute of Zoology, Museum of Zoology, USSR, Kiev (83 pp., 30 maps) in 1975. Lists of synonyms, bibliography, maps and table of contents - all being parts of the original paper - have not been included in this translation which was carried out with the consent of the author by R. GÜNTHER (Berlin) and H. GRILLITSCH (Vienna). The English summary was taken over as provided in the original version. As a SHORT NOTE in this issue of HERPETOZOA subsequent to the translation comments and updated addenda by W. MAYER are provided indicated by [aa* bb* etc.] in the text.