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Here we provide the first phylogenetic analysis that include Afrogecko ansorgii and a detailed morphological comparison with other species of leaf-toed geckos. For this purpose, we used two mitochondrial (16S, ND2) and four nuclear (RAG1, RAG2, CMOS, PDC) genes to produce a robust phylogenetic reconstruction. This allowed us to show that A. ansorgii is not related as previously believed to circum-Indian Ocean leaf-toed geckos and is rather more closely related to other Malagasy leaf-toed geckos. Additionally, we explore and compare osteological variation in A. ansorgii skulls through High Resolution X-ray Computed Tomography with previously published material. This allowed us to describe herein a new genus, Bauerius gen. nov., and additionally provide a detailed redescription of the species (including the first description of male material), supplementing the limited original description and type series, which consisted of only two females.

Four new species of flat geckos in the Afroedura bogerti Loveridge, 1944 group are described from south-western and west-central Angola. The description of these new species significantly restricts the distribution range of typical A. bogerti, a morphologically very similar species, from which they differ genetically by 5.9–12% divergence for the mitochondrial 16S ribosomal RNA gene. Morphologically and genetically, Angolan Afroedura are divided into two main groups: a mostly south-western coastal group and a west-central inland/highland group. These two groups are further divisible into three and two subgroups respectively, all geographically isolated, differing by a combination of the following features: colouration, average adult size, number of mid-body scale rows, number of scale rows on dorsal and ventral surface of each tail verticil and if nostril scales are in contact or not. All five Angolan species are morphologically distinguishable and in agreement with the molecular results. An updated dichotomous key to the Afroedura transvaalica group is provided. The new discovery adds to a growing number of endemic Pro-Namib reptiles described from Angola in recent years.

The Scaled Sandveld Lizard Nucras scalaris Laurent, 1964 is a poorly known lacertid endemic to north-eastern Angola and is only known from the type series collected more than half a century ago. The original description provided a comprehensive morphological description, but there was a lack of information regarding its evolutionary relationships and habitat associations. A recent discovery of N. scalaris from Cusseque, Bié Province, on the central Angolan plateau provided the opportunity to address some knowledge gaps of this species. A phylogenetic analysis confirmed its distinctiveness at the species level. Despite a limited dataset, the analysis suggests that N. scalaris is sister to N. broadleyi rather than N. lalandii, the latter having been assumed to be the sister species, because of morphological similarity between them. The new record of N. scalaris represents a range extension of more than 350 km to the southwest and extends the elevation range to 1 570 m above sea level, compared to previous known localities at about 1 300 m above sea level (Alto Cuílo and Alto Chicapa, in Lunda Sul Province). The specimen also provides new information on live coloration, as well as the habitat association of grassy shrubland that is dominated by geoxylic suffrutices (the so-called ‘underground forests’) characteristic of the Angolan Miombo Woodlands. This new information raises the question whether poor survey data have led to an under-estimation of faunal diversity in this peculiar and overlooked vegetation type, and reinforces the need for further surveys which could highlight the importance of this habitat.

Background The African continent is comprised of several different biomes, although savanna is the most prevalent. The current heterogeneous landscape was formed through long-term vegetation shifts as a result of the global cooling trend since the Oligocene epoch. The overwhelming trend was a shift from primarily forest, to primarily savanna. As such, faunal groups that emerged during the Paleogene/Neogene period and have species distributed in both forest and savanna habitat should show a genetic signature of the possible evolutionary impact of these biome developments. Crotaphopeltis and Philothamnus (Colubridae) are excellent taxa to investigate the evolutionary impact of these biome developments on widespread African colubrid snakes, and whether timing and patterns of radiation are synchronous with biome reorganisation. Methods A phylogenetic framework was used to investigate timing of lineage diversification. Phylogenetic analysis included both genera as well as other Colubridae to construct a temporal framework in order to estimate radiation times for Crotaphopeltis and Philothamnus. Lineage diversification was estimated in Bayesian Evolutionary Analysis Sampling Trees (BEAST), using two mitochondrial markers (cyt–b, ND4), one nuclear marker (c–mos), and incorporating one fossil and two biogeographical calibration points. Vegetation layers were used to classify and confirm species association with broad biome types (‘closed’ = forest, ‘open’ = savanna/other), and the ancestral habitat state for each genus was estimated. Results Philothamnus showed an ancestral state of closed habitat, but the ancestral habitat type for Crotaphopeltis was equivocal. Both genera showed similar timing of lineage diversification diverging from their sister genera during the Oligocene/Miocene transition (ca. 25 Mya), with subsequent species radiation in the Mid-Miocene. Philothamnus appeared to have undergone allopatric speciation during Mid-Miocene forest fragmentation. Habitat generalist and open habitat specialist species emerged as savanna became more prevalent, while at least two forest associated lineages within Crotaphopeltis moved into Afromontane forest habitat secondarily and independently. Discussion With similar diversification times, but contrasting ancestral habitat reconstructions, we show that these genera have responded very differently to the same broad biome shifts. Differences in biogeographical patterns for the two African colubrid genera is likely an effect of distinct life-history traits, such as the arboreous habits of Philothamnus compared to the terrestrial lifestyle of Crotaphopeltis.

Background The African continent is comprised of several different biomes, although savanna is the most prevalent. The current heterogeneous landscape was formed through long-term vegetation shifts as a result of the global cooling trend since the Oligocene epoch. The overwhelming trend was a shift from primarily forest, to primarily savanna. As such, faunal groups that emerged during the Paleogene/Neogene period and have species distributed in both forest and savanna habitat should show a genetic signature of the possible evolutionary impact of these biome developments. Crotaphopeltis and Philothamnus (Colubridae) are excellent taxa to investigate the evolutionary impact of these biome developments on widespread African colubrid snakes, and whether timing and patterns of radiation are synchronous with biome reorganisation. Methods A phylogenetic framework was used to investigate timing of lineage diversification. Phylogenetic analysis included both genera as well as other Colubridae to construct a temporal framework in order to estimate radiation times for Crotaphopeltis and Philothamnus. Lineage diversification was estimated in Bayesian Evolutionary Analysis Sampling Trees (BEAST), using two mitochondrial markers (cyt–b, ND4), one nuclear marker (c–mos), and incorporating one fossil and two biogeographical calibration points. Vegetation layers were used to classify and confirm species association with broad biome types (‘closed’ = forest, ‘open’ = savanna/other), and the ancestral habitat state for each genus was estimated. Results Philothamnus showed an ancestral state of closed habitat, but the ancestral habitat type for Crotaphopeltis was equivocal. Both genera showed similar timing of lineage diversification diverging from their sister genera during the Oligocene/Miocene transition (ca. 25 Mya), with subsequent species radiation in the Mid-Miocene. Philothamnus appeared to have undergone allopatric speciation during Mid-Miocene forest fragmentation. Habitat generalist and open habitat specialist species emerged as savanna became more prevalent, while at least two forest associated lineages within Crotaphopeltis moved into Afromontane forest habitat secondarily and independently. Discussion With similar diversification times, but contrasting ancestral habitat reconstructions, we show that these genera have responded very differently to the same broad biome shifts. Differences in biogeographical patterns for the two African colubrid genera is likely an effect of distinct life-history traits, such as the arboreous habits of Philothamnus compared to the terrestrial lifestyle of Crotaphopeltis.

Since 1957, all padlopers were considered Homopus, which was an anomaly, because only two species have four claws on the front and hind limbs. The revival of the genus Chersobius for the five-toed species (signatus, boulengeri and solus) now limits Homopus to the four-toed species (areolatus and femoralis). Molecular data indicate that Homopus is paraphyletic with two well-supported clades: the four-toed Homopus in one clade and the five-toed Chersobius together with Chersina angulata in a second clade. There are numerous distinct morphological and ecological differences between the genera. All Chersobius species have a single, large inguinal scute, whereas the number is variable in Homopus. The midline pectoral scute of Chersobius is consistently longer, and the nuchal scute shorter, than in Homopus. The fourth to fifth vertebral suture is narrow in Homopus and broad in Chersobius. Forelimb scales also differ with 3–4 large rows and 5–6 smaller rows, respectively, in Homopus and Chersobius. Chersobius males have a plastral concavity and Homopus males not. Two Chersobius species (C. signatus and C. boulengeri) are distinguishable from Homopus by having 12 or more marginal scutes, whereas Homopus species have 11 or fewer. The glans penes are respectively V-shaped and heart-shaped in Homopus and Chersobius. Ecological differences include that Chersobius species are rupicolous and inhabit arid regions, whereas Homopus species inhabit more mesic habitats. Homopus females produce multi-egg clutches and tend to be bigger compared to single-egg clutching Chersobius females. Females of all species are larger than conspecific males, and all males have larger shell openings than females have. ZooBank—urn:lsid:zoobank.org:pub: 18229454-6662-46F3-8B15-836BF2AC38EB

English-born Donald George Broadley spent most of his adult life working as a herpetologist at two museums in Zimbabwe. His many taxonomic reviews and other publications made him one of the most re...

Donald G. Broadley was one of Africa’s most prolific recent authors. He produced over 410 articles, including numerous comprehensive and detailed taxonomic reviews of lizard and snake genera. A rev...

A new species of tree snake Dipsadoboa montisilva Branch, Conradie & Tolley sp. nov. (Serpentes: Colubridae) is described from the ‘sky islands’ of Mount Mabu and Mount Ribáuè in northern Mozambique. Features of scalation, colour, body form and habitat distinguish the new species from other Dipsadoboa. This is supported by a phylogenetic analysis using one mitochondrial marker (cytochrome b) that shows the new Mozambican species is divergent from other sampled Dipsadoboa, including D. flavida and D. aulica, the only congeners known to occur in Mozambique. Morphologically, the new Dipsadoboa forms part of the D. werneri-shrevei complex from east and southeast Africa, but differs in having higher subcaudal counts, a different temporal pattern and only two supralabials entering the orbit. Phylogenetically, it occurs in a clade with D. shrevei and D. werneri. The status of D. shrevei in East Africa is reassessed, particularly in terms of the poorly-known Dipsadoboa shrevei kageleri from northern Tanzania. It is morphologically well defined from D. shrevei shrevei and utilises a different habitat. Although based on limited genetic data, it appears to be well-defined from typical D. shrevei and is accordingly raised to specific status. The only Tanzanian record for typical D. shrevei from Mtene, Rondo Plateau in southeast Tanzania is well isolated from the species’ range to the west (e.g. Zambia, Angola) and the published scalation features, particularly ventral counts, do not fully accord with D. shrevei. The Rondo Plateau population is treated as Dipsadoboa incerta sedis, and because we return D. shrevei to its binomial status, we can no longer consider D. shrevei as occurring in Tanzania. Biogeographically, the Rondo Plateau population may have a stronger affinity to the new Mozambican species. The discovery of isolated populations of the new species in mid-altitude forest remnants on Mt Mabu and Mt Ribáuè emphasizes the high conservation importance of the Mozambique forest ‘sky islands’ from which numerous other endemic new species have been recently discovered. These species are impacted by ongoing habitat destruction through slash and burn clearing for subsistence agriculture.

Aim: The expansion of open habitats during the mid-Miocene has been hypothesized as a driver of allopatric speciation for many African taxa. This habitat-dependent mode of diversification has been implicated in the shift from C3 (e.g. forest/woodland) to C4 dominated systems (i.e. open savanna, grasslands) in a number of African squamates. We examined this hypothesis using a genus of African viperid snakes (Bitis) with both open habitat and forest-dwelling representatives. \ud \ud Location: Africa. \ud \ud Methods: A comprehensive multilocus dataset was used to generate a calibrated species tree using a multispecies coalescent model. Individual gene trees and patterns of nuclear allele sharing were used to assess species monophyly and isolation. To test the habitat-dependent evolution hypothesis, we generated an ancestral character state reconstruction for open and closed habitats using the dated phylogeny. This was related to the timing of open habitat expansion and forest/woodland contraction in Africa. \ud \ud Results: The genus Bitis originated in the Oligocene, with species level diversification in the late Miocene/Pliocene. Four well-supported clades correspond to the recognized subgenera Bitis, Keniabitis, Macrocerastes and Calechidna. Several previously unrecognized lineages potentially represent cryptic species. \ud \ud Main conclusions Habitat-dependent evolution does not appear to have been a main driver for generic level viperine diversification: the ancestral state for Bitis was open habitat and at least one clade moved into forest in the Miocene, long after forest had contracted and fragmented. Forest-dependent species diversified only in the late Miocene, presumably as forest became further reduced in extent, fitting an allopatric model of speciation. Although our results do not favour a general pattern of habitat-dependent diversification in Bitis, cladogenesis within the subgenus Calechidna for “arenicolous” species (Bitis caudalis complex) and “rupicolous” species (B. Atropos-cornuta complex), corresponds to the aridification of southwest Africa. This suggests there are subtleties not captured in the broad open habitat category, which are relevant for understanding the role of habitat-dependent evolution.

Two rare and endemic psammophines (Serpentes: Psammophiinae) occur in Angola. The taxonomic status of Psammophylax rhombeatus ocellatus Bocage, 1873 and Psammophis ansorgii Boulenger, 1905 have long remained problematic, with both having varied past and present taxonomic assignments, and whose distributions may therefore present zoogeographic anomalies. Little was known of their biology, habitat associations, or phylogenetic relationships. New material was collected during biodiversity surveys of the Humpata Plateau, near Lubango, Angola. It allowed fuller descriptions of scalation and live coloration for both species, and resolution of their taxonomic status. Genetic analysis confirms that both are distinct at the specific level. In addition, within Psammophis, Jalla’s Sand Snake (Psammophis jallae Peracca, 1896), of which P. rohani Angel, 1925, remains a synonym, is sister to P. ansorgii, and Boulenger’s comment on similarities with P. crucifer are not supported. The status of an unusual skaapsteker from Calueque, Cunene Province, Angola, is discussed and its assignment to Ps. ocellatus is provisional and requires additional material for taxonomic resolution. The new P. ansorgii records from Tundavala represent a range (+400 km southwest) and altitude (1800 m to 2286 m a.s.l) extension from the previous only known precise locality of Bela Vista (= Catchiungo), Huambo Province, whilst that for Ps. ocellatus doubles the known altitude from 1108 m to 2286 m a.s.l and extends the range about 122 km to the northwest from historical material from the plateau of Huíla and Cunene provinces.

The Port Elizabeth Museum herpetology collection contains 407 type specimens, representing 70 primary and 55 secondary squamate types. The type series comprise 93 African taxa (84 lizards and 9 snakes), of which 75 are still regarded as valid. It is the third largest primary reptile type collection in Africa. This is the first catalogue of this important African squamate type collection. It provides the original name, original publication date, journal volume number and pagination, reference to illustrations, current name, museum collection number, type locality, and notes on the status of all types and important additional non-type material mentioned in historical descriptions. Photographs of all primary types, as well as original illustrated material are provided.

Angolan amphibians have been studied since the mid-nineteenth century by explorers and scientists from all over the western world, and collections have been deposited in around 20 museums and institutions in Europe, Northern America, and Africa. A significant interruption of this study occurred during Angola’s liberation struggle and civil war for nearly four decades and, as a consequence, knowledge about the country’s biodiversity became outdated with critical gaps. Since 2009, a new era in Angolan biodiversity studies started as expeditions scattered in southwestern, northeastern, southeastern, and northwestern Angola lead to exciting discoveries, including new records for the country, descriptions of new species, range extensions and taxonomical updates. Currently 111 amphibian species are listed for the country (of which 21 are endemic), but this number is an underestimate and the various unresolved taxonomical issues challenge the study of every other aspect of this group. The Angolan amphibian fauna remains one of the most poorly known in Africa and much still has to be done in order to understand its diversity, evolution and conservation needs. An overview of existing knowledge of Angolan amphibians is presented, including an updated checklist for the country, comments on problematic groups, endemic species, biogeography, recent findings, and priority research topics.

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 during 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.

Assessments of biodiversity patterns and threats among African reptiles have lagged behind those of other vertebrate groups and regions. We report the first systematic assessment of the distribution, threat status, and climate change vulnerability for the reptiles of Tanzania. A total of 321 reptile species (including 90 Tanzanian endemics) were assessed using the global standard IUCN Red List methodology and 274 species were also assessed using the IUCN guidelines for climate change vulnerability. Patterns of species richness and threat assessment confirm the conservation importance of the Eastern Arc Mountains, as previously demonstrated for birds, mammals and amphibians. Lowland forests and savannah-woodland habitats also support important reptile assemblages. Protected area gap analysis shows that 116 species have less than 20% of their distribution ranges protected, among which 12 are unprotected, eight species are threatened and 54 are vulnerable to climate change. Tanzania's northern margins and drier central corridor support high numbers of climate vulnerable reptile species, together with the eastern African coastal forests and the region between Lake Victoria and Rwanda. This paper fills a major gap in our understanding of the distribution and threats facing Tanzania's reptiles, and demonstrates more broadly that the explicit integration of climate change vulnerability in Red Listing criteria may revise spatial priorities for conservation.

The assimilation of information on taxonomy, distribution, basic ecology and conservation status of Africa's reptiles lags far behind that for most other continents. Many regions of mainland Africa are rarely surveyed, resulting in severe knowledge gaps that currently limit effective conservation of African reptiles. Here, we provide a precis on the knowledge gaps and conservation status of mainland African reptiles, and quantify the main threats based on IUCN Red List of Threatened Species assessments using publicly available distribution data. Our results show that these data are insufficient to confidently identify areas of high biodiversity, with large gaps in knowledge in the Horn of Africa, central Africa and West Africa. There is a strong overall taxonomic bias in extinction risk with 45% of families more threatened than expected by chance. Furthermore, Amphisbaenidae, Chameleonidae, Gerrhosauridae, Testudinidae, Viperidae all have a high percentage of their constituent species at risk. Overall, land transformation for agriculture, particularly subsistence farming, constitutes the primary threat to African reptiles, and our derived Threat Index based on socio-economic traits of African countries show that risk is high in Burundi, Ethiopia, Liberia, Malawi, Rwanda and Sierra Leone. These findings highlight important challenges facing the conservation of African reptiles, and we suggest that conservation priorities in mainland Africa be focussed on areas where the potential for overall loss of biodiversity is high, particularly in regions where knowledge is inadequate.

Africa, inclusive of the West Indian Ocean islands, harbours 11 of the world's 16 extant testudinid genera. Fossil records indicate that testudinids originated in Asia and dispersed first to North America and Europe (Early Eocene) and later to Africa (Late Eocene). We used mitochondrial (1870 bp) and nuclear (1416 bp) DNA sequence data to assess whether molecular data support the late cladogenesis of Southern African testudinid lineages. Our results revealed strong support for the monophyly of a clade consisting of Kinixys, the two Malagasy genera and four Southern African genera (Psammobates, Stigmochelys, Homopus and Chersina). Kinixys diverged from this clade in the Late Palaeocene, suggesting that testudinids occupied Africa at an earlier date than indicated by fossil records. The Southern African tortoises consist of three, strongly supported clades: Psammobates + Stigmochelys; the five-toed Homopus + Chersina; and the four-toed Homopus. Due to the paraphyly of Homopus, we propose the taxonomic resurrection of Chersobius for the five-toed Homopus species (boulengeri, signatus and solus). Cladogenesis at the genus level occurred mainly in the Eocene, with Chersina and Chersobius diverging in the Oligocene. The latter divergence coincided with species-level radiations within Homopus (areolatus and femoralis) and Psammobates (oculifer, geometricus and tentorius). Our phylogeny could not resolve relationships within Psammobates, indicating rapid speciation between the Late Oligocene and Early Miocene. The Chersobius species were the last to diverge in the Early to Mid-Miocene. By the Mid-Miocene, P. tentorius started to differentiate into four lineages instead of the three recognized subspecies: P. t. tentorius, P. t. trimeni and two P. t. verroxii subclades occurring north and south of the Orange River, respectively. Terminal radiations in several taxa suggest the existence of cryptic species and a more diverse tortoise fauna than currently recognized. Factors contributing to this diversity may include the early origin of African testudinids and climatic fluctuations over a heterogeneous landscape.

A recent molecular revision of the snake-eyed skinks of the genus Panaspis Cope, 1868 uncovered extensive cryptic diversity within the P. wahlbergi (Smith, 1849) and P. maculicollis Jacobsen & Broadley, 2000 species complexes. We here describe an unnamed central and northern Namibian lineage of the P. maculicollis group as a new species. We base the description on previously published molecular evidence and newly collected morphological, meristic and coloration data. Members of the group share a conservative morphology and the newly described species is very similar to P. maculicollis, from which it differs mostly on the basis of its coloration. This description raises the number of Panaspis recognized to 17 species and reinforces the already strong evidence that northern Namibia is a hotspot of reptile diversity.

Trachylepis (Mabuyinae) includes ∼80 species of fully-limbed skinks found primarily in Africa and Madagascar, but a robust species-level phylogeny for this genus is lacking and this impedes studies on a wide-range of topics from biogeography to character evolution. Trachylepis and its close relatives (which together form the Mabuya group or Mabuyinae) are notable in that they have undergone multiple transitions and remarkable specializations in their reproductive modes. A Trachylepis phylogeny will be particularly useful for investigating reproductive evolution, because it includes species that exhibit oviparity, viviparity, and bimodal parity (species with both oviparous and viviparous populations). We sequenced DNA at four mitochondrial and five nuclear loci for 67 (∼84% of) Trachylepis species to infer a phylogeny for this genus. We performed stochastic character mapping of parity mode under a variety of parity mode transition models to infer ancestral parity mode states and the number and type of parity mode transitions. We recovered a strongly supported phylogeny of Trachylepis that is generally consistent with earlier phylogenetic studies. The best-fit model of reproductive mode evolution supports an oviparous ancestor for Trachylepis, and supports at least three viviparity to oviparity transitions. We compared parity mode evolution under the overall best-fit model (no constraints on parity mode transitions) to the best-fit model among the subset of models that assume viviparity to oviparity transitions are impossible. Our results support a model of reproductive evolution that allows for reversibility from viviparity to oviparity, a process that is not generally accepted. Alternatively, the best-fit model of evolution among the set of models that eliminate reversals from viviparity to oviparity suggests that bimodal reproduction may have persisted for millions of years within multiple lineages.

Theory predicts that sexually dimorphic traits under strong sexual selection, particularly those involved with intersexual signaling, can accelerate speciation and produce bursts of diversification. Sexual dichromatism (sexual dimorphism in color) is widely used as a proxy for sexual selection and is associated with rapid diversification in several animal groups, yet studies using phylogenetic comparative methods to explicitly test for an association between sexual dichromatism and diversification have produced conflicting results. Sexual dichromatism is rare in frogs, but it is both striking and prevalent in African reed frogs, a major component of the diverse frog radiation termed Afrobatrachia. In contrast to most other vertebrates, reed frogs display female-biased dichromatism in which females undergo color transformation, often resulting in more ornate coloration in females than in males. We produce a robust phylogeny of Afrobatrachia to investigate the evolutionary origins of sexual dichromatism in this radiation and examine whether the presence of dichromatism is associated with increased rates of net diversification. We find that sexual dichromatism evolved once within hyperoliids and was followed by numerous independent reversals to monochromatism. We detect significant diversification rate heterogeneity in Afrobatrachia and find that sexually dichromatic lineages have double the average net diversification rate of monochromatic lineages. By conducting trait simulations on our empirical phylogeny, we demonstrate that our inference of trait-dependent diversification is robust. Although sexual dichromatism in hyperoliid frogs is linked to their rapid diversification and supports macroevolutionary predictions of speciation by sexual selection, the function of dichromatism in reed frogs remains unclear. We propose that reed frogs are a compelling system for studying the roles of natural and sexual selection on the evolution of sexual dichromatism across micro- and macroevolutionary timescales.

The African green and bush snakes of the genus Philothamnus currently comprises 21 species and three subspecies and occurs throughout sub-Saharan Africa. The genus has been the subject of previous taxonomic revisions based on traditional morphological characters and limited genetic assessment, and may not reflect their evolutionary history. Indeed, previous findings based on phylogenetics show discordant results of interspecific relationships and question the monophyly of the genus, although taxon sampling has been limited to date. We investigated phylogenetic affinities within Philothamnus with more inclusive genetic and geographical sampling, with the aim of better understanding their evolutionary history, so that future taxonomic revision of Philothamnus can be better informed. Species relationships were examined within a phylogenetic context and sampling included 133 ingroup samples from 16 taxa. Phylogenies were constructed in Bayesian and likelihood frameworks using three mitochondrial (16S, cyt b and ND4) and two nuclear (c-mos and RAG1) markers. Competing hypotheses relating to the monophyly of the genus were tested with a Shimodaira-Hasegawa test. To examine species boundaries, Bayesian General Mixed Yule-Coalescent Model and multi-rate Poisson Tree Processes analyses were conducted. In addition, a barcoding approach was used to further clarify species-level relationships by comparing frequency distributions between intra- and interspecific sequence divergence. The genus was recovered as monophyletic; however, species-delimitation results suggest that the current taxonomy does not reflect the evolutionary history of this group. For example, Philothamnus s. semivariegatus is paraphyletic, with at least four distinct clades. Philothamnus carinatus consists of two cryptic (sister) lineages from Central and West Africa that are deeply divergent, suggesting a long history of isolation between those regions. Furthermore, the subspecies P. n. natalensis and P. n. occidentalis show strong support for species-level divergence, which reflects their morphological and ecological differences. Accordingly, we elevate P. occidentalisnov. comb. to a full species. A fully informed taxonomic revision of these taxa will require additional morphological and ecological data for corroboration, but it seems that the morphological characters (e.g. scalation, dentition) used to describe these species to date are labile within and between species. This most likely has clouded our understanding of the species boundaries within the genus. Our phylogeny and species-delimitation analyses should provide a sounder framework for taxonomy, but may also prove useful toward understanding the morphological adaptations of these species to their respective habitats.

Members of the snake subfamily Aparallactinae occur in various habitats throughout sub-Saharan Africa. The monophyly of aparallactine snakes is well established, but relationships within the subfamily are poorly known. We sampled 158 individuals from six of eight aparallactine genera in sub-Saharan Africa. We employed concatenated gene-tree analyses, divergence dating approaches, and ancestral-area reconstructions to infer phylogenies and biogeographic patterns with a multi-locus data set consisting of three mitochondrial (16S, cyt b, and ND4) and two nuclear genes (c-mos and RAG1). As a result, we uncover several cryptic lineages and elevate a lineage of Polemon to full species status. Diversification occurred predominantly during the Miocene, with a few speciation events occurring subsequently in the Pliocene and Pleistocene. Biogeographic analyses suggested that the Zambezian biogeographic region, comprising grasslands and woodlands, facilitated radiations, vicariance, and dispersal for many aparallactines. Moreover, the geographic distributions of many forest species were fragmented during xeric and cooler conditions, which likely led to diversification events. Biogeographic patterns of aparallactine snakes are consistent with previous studies of other sub-Saharan herpetofauna.

Using up to 2117 bp of mitochondrial DNA and up to 2012 bp of nuclear DNA, we analysed phylogeographic differentiation of six widely distributed species of African hinged terrapins (Pelusios spp.) representing different habitat types. Two taxa each live in savannahs or in forests and mesic savannahs, respectively, and the remaining two species occur in intermediate habitats. The species living in forests and mesic savannahs do not enter dry savannahs, whereas the savannah species may occur in dry and wet savannahs and even in semi-arid steppe regions. We found no obvious correlation between habitat type and phylogeographic pattern: one savannah species (P. rhodesianus) shows phylogeographic structure, i.e. pronounced genetic differences among geographically distinct populations, and the other (P. nanus) not. One species inhabiting forests and mesic savannahs (P. carinatus) has phylogeographic structure, the other (P. gabonensis) not. The same pattern is true for the two ecologically intermediate species, with phylogeographic structure present in P. castaneus and absent in P. chapini. Nuclear evidence suggests that the latter two taxa with abutting and partially overlapping ranges are distinct, while mtDNA is only weakly differentiated. Pelusios castaneus shows pronounced phylogeographic structure, which could reflect Pleistocene range interruptions correlated with the fluctuating forest cover in West and Central Africa. Our results do not support the recognition of an extinct subspecies of P. castaneus for the Seychelles. Pelusios carinatus contains two well supported clades, which are separated by the Congo River. This species is closely related to P. rhodesianus, a taxon consisting of two deeply divergent mitochondrial clades. One of these clades is paraphyletic with respect to P. carinatus, but the two clades of P. rhodesianus are not differentiated in the studied nuclear markers and, again, paraphyletic with respect to P. carinatus. Using mtDNA sequences from the type material of P. rhodesianus, we were able to allocate this name to one of the two clades. However, owing to the confusing relationships of P. rhodesianus and P. carinatus, we refrain from taxonomic decisions.

Key to the Afroedura transvaalica- group 1. Midbody scale rows more than 95........................................................................2 - Midbody scale rows less than 95; Angola.................................................. A. bogerti (sensu lato) 2. Rostral bordering nostril................................................................................3 - Rostral excluded from nostril................................................................... A. loveridgei 3. Anterior nasals in contact (very rarely separated); scales around midbody-South Africa 102-118 (= 109), northern Zimbabwe 108-119 (= 114)........................................................................... A. transvaalica - Anterior nasals separated by 1-3 granules; scales around midbody 99-101 (= 100)................ A. gorongosa sp. nov.

Afroedura gorongosa sp. nov. (Figs. 4–5) Afroedura transvaalica (part) Broadley 1966, p. 111. Holotype. ZMB 83293 (GNP 484), adult male, western flank of Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (1038 m a.s.l., 18˚28’04.3”S, 34˚02’53.2”E), collected by M.- O. Rödel and M.F. Barej, 11 December 2015. Paratypes (three specimens). ZMB 83290 (GNP 438), adult female, with incision in left thigh; PEM R22220 (GNP 439, previously ZMB 83291), adult female, with incision in left thigh; ZMB 83292 (GNP 440; will later be deposited in the zoological collection of the E.O. Wilson Laboratory in Gorongosa National Park), adult male, with everted hemipenes and incision in left thigh; collected on 22 May and 21 July 2015 by J.A. Guyton and P. Naskrecki from gallery forest near Murombodzi Waterfall, Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (842 m a.s.l; 18°29'0.1''S, 34°2'34.6''E). Additional material (two specimens). ZMB 83288 (GNP 433), adult male, ZMB 83289 (GNP 434), adult female; both collected in April 2015 by H. Farooq, P. Naskrecki and J.A. Guyton from the top of Bunga Inselberg, Gorongosa National Park, Sofala Province, Mozambique (approx. 200 m a.s.l., 18˚35’58.1”S, 34˚20’34.8”E). Etymology. The specific name refers to Mount Gorongosa and Gorongosa National Park in Central Mozambique, to which the species is endemic. We suggest Gorongosa Flat Gecko is a suitable common name. Afroedura, based on Oedura, is feminine and the specific epithet is treated as a noun in apposition. Diagnosis. A member of the A. transvaalica -complex in possessing two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Differs from other members of the complex by having 97–102 midbody scale rows (less than 95 in A. bogerti- complex, 113–120 (= 117) in A. loveridgei and 102–119 in A. transvaalica (102–118, = 109 in South Africa—Jacobsen et al. 2014; 108–119, = 114 in Zimbabwe adjacent to Mount Gorongosa); by the rostral bordering the nostril (nostril excluded from rostral in A. loveridgei); by the anterior nasals being separated by 1–3 granules (always in broad contact in A. loveridgei; usually in broad contact in A. transvaalica - A. loveridgei; 17–20 in South African A. transvaalica, and 15–19 in Zimbabwe populations adjacent to Mozambique); and in having a higher average number of precloacal pores in males (8–13, = 10; 6–10, = 8 in A. transvaalica; 8–10, = 9 in A. loveridgei). Holotype description. Morphology. Adult male; SVL 50.8 mm; tail 52.2 mm (partly regenerated), with a small incision in the left thigh for the removal of muscle tissue. Head and body dorsoventrally depressed; HL 12.3 mm, HW 9.0 mm, broadest at mid-eye and 1.37x longer than wide. Eye large (3.7 mm wide), pupil vertical with indented margins; circumorbital scales small and granular, becoming elongate, almost ciliate, at upper posterior margin. Snout rounded, 4.9 mm long, longer than distance between eye and ear openings (3.6 mm). Scales on top of snout granular, rounded and convex, the largest being 2/3rds the width of the scales on the back, which are granular and juxtaposed, with no intervening minute granules. Scales on snout almost twice as large as those on crown and throat. Scales on belly flattened, imbricate, more-or-less ovate at mid-ventrum, and twice the size of lateral granules and 1.5x those along backbone. Ear opening deep, oblique and roughly oval, only half as high as wide. Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at posterior margin of limb insertions. All digits with a large pair of distal scansors, separated by a large, curved claw, and followed after a large gap (twice length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; 12 scale rows under 4th toe. Nostril pierced between rostral, 1 st supralabial and three nasal scales; the supranasal being much larger than the subequal postnasals and separated by a granule bordering the rostral, followed behind by two smaller granules. Rostral roughly rectangular but with its upper edges elongated due to extensions into the nostril. Nine supralabials on each side, the labial margin flexing dorsad at the rictus, with 1–2 minute scales proximal to the flexure. Nine infralabials on either side, with a small scale proximad to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only 2/3rds the width of rostral, and in contact with two distinctly elongate postmental scales. Approximately 20 scales across crown at level of front of eye; 13 scales between nostril and front of eye; 23 scales from front of ear to back of eye; 99 scales around midbody. A roughly V-shaped row of 13 precloacal pores occurs eight scale rows anterior to the cloacal lip, with these scales more imbricate and triangular adjacent to the precloacal pores. Original portion of tail slightly dorsoventrally flattened and indistinctly verticillate, without obvious lateral constrictions; each verticil comprising 6–7 imbricate rows of scales dorsally and 5 imbricate scale rows ventrally, and with ventral scales approximately twice the size of those on dorsal surface. Hemipenal pouches well-developed, with an angular, ventro-lateral series of 3 enlarged tubercular scales on each side (largest nearest cloaca), and with two openings of the cloacal sacs on the posterior lip of the cloaca. Colour in life (from photographs, Fig. 4a and unpublished). Dark black-brown above with a series of five white dorsal spots (4–5 scales across) between the fore- and hind-limb insertions, that are surrounded by a chocolate brown zone that encompasses a series of 2–4 laterally arranged smaller pale spots (2–3 scales wide), and posteriorly by a diffuse pale region that forms an irregular band across the mid dorsum. Top of head, flanks and upper surfaces of limbs with small, scattered pale spots. Head with a dark brown band across the posterior edge of crown encompassing a small central pale spot; a vague, thin pale canthal stripe, extends on both sides from the nasal region to front of eye; upper and lower labials grey-brown with diffuse yellowish bars; iris reddish brown with a heavy dark brown reticulation, and pupil with crenulated edge. Original segment of tail mottled in dark brown and pale blotches; regenerated tail grey with vague darker blotches. Ventrum mottled grey-cream. Colour in preservative. Dark black-brown above with irregular markings on back, comprising four equally spaced dark bars, each with a central white spot 4–5 scales wide, and an irregular, paler brown bar behind; top and sides of head, flanks and upper surfaces of limbs with small, irregular pale spots; ventral surface of throat, neck, belly, limbs and tail uniform dark grey. Side of head and labials dark brown, with a few scattered pale spots on upper labials, and vague pale edges to lower labials. Variation in paratypes and additional material (Figs. 4b, 4c, 5; Tables 3–4). Morphology. SVL from 54.6 mm (ZMB 83289) to 61.2 mm (PEM 22220); head length 1.24–1.38 times head width (max. in paratypes 1.38); snout only 1.5 times diameter of eye in one specimen. The supranasals always separated by granules, usually with a single large granule in contact with the rostral between the supranasals, followed by 2–3 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril, and the width of the rostral at the lip margin is always wider than that of the mental (except in ZMB 83290 were they are subequal); always two postmental scales; supralabials more than in holotype (10–11), infralabials (9–10); scales between anterior edge of eye 19–21; scales between nostril and anterior edge of orbit 12–13; scales between anterior edge of ear and rear margin of orbit 21–24; scales around mid-body 97–101; subdigital lamella beneath 4th toe 10–11; dorsal scales per tail verticel 7–8; ventral scales per tail verticel 5. In one female (ZMB 83290) the terminal, medially-constricted subdigital lamella of the 3rd toe of left foot is swollen and scansor-like, bearing small setae. Precloacal pores 8–13 in males, females without dimples in precloacal scales (as in A. transvaalica, but unlike A. loveridgei where five females had 5–9 dimpled precloacal scales, that lacked secretory activity, and only one had none). Colour in life (based on female paratype PEM R22229). Boldly patterned, with six dorsal dark transverse bands, posteriorly edged with cream, the first on the neck, the second and sixth at the fore- and hindlimb insertions, respectively, and with three thin bands equally spaced between them on the body; background colour of body, head and upper surfaces of limbs flecked on light brown and cream with occasional pale spots 2–4 scales wide; a dark canthal stripe runs through the rostral to the front of the eye, with a paler zone above; the scales of the anteriordorsal region of the orbital rim are bright yellow, with a faint yellowish infusion of the crown scales between the eyes; the original tail is boldly barred dorsally with seven, subequally-spaced, dark brown-black bars with a yellow posterior edge; belly uniform cream with scattered yellowish scales, greyer on ventral surfaces of limbs and tail. Colour in preservative. Similar to holotype, but dorsum pale brown to grey; larger dorsal markings usually reticulated, occasionally forming irregular transverse bands (6–7 from occiput to between hindlimbs); smaller dorsal markings vary from vague to distinct. A few specimens have irregular pale blotches along the posterior edges of the darker dorsal patches/bands, and also scattered on the dorsal surfaces of the limbs and back. Dorsal surface of regenerated tails usually with irregular pale and dark blotches that may form wavy bands, and which become flecks or grey blotches below. Size. Largest male (ZMB 83288)—SVL 56.5 mm, tail (partially regenerated) 55.7 mm; largest female (PEM R22220)—SVL 61.2 mm, tail (original) 64.0 mm. It is thus intermediate in size between A. transvaalica (malemax. SVL 72 mm, female—max. SVL 64 mm) and A. loveridgei (male—max. SVL 59 mm, female—max. SVL 58 mm). Only one of six specimens had an original tail (105% SVL). Hemipenis (based on ZMB 83292). Approximately 5 mm long, with a simple, smooth, possibly longitudinally flounced pedicel; capped with two large cups covered with fine calyculate ornamentation; sulcus simple and draining into the conjoined base of the distal cups. Distribution. Known from only three localities in the Mount Gorongosa region (Fig. 2). The holotype and paratypes, plus various non-collected individuals, have been found on the western flanks of Mount Gorongosa between about 900 and 1100 m a.s.l. (Figs. 6a–c). Access to further potential areas on the mountain, i.e. rock faces at the eastern slopes, was not possible due to security concerns. The additional material and various non-collected specimens have been observed at a lower altitude on the Bunga Inselberg (about 212 m a.s.l.) in the north-western corner of the National Park, approximately 40 km from the type locality (Fig. 6d). No tissue samples from the Bunga individuals were available, and the assignment of these individuals to the new species was thus based on morphological similarity. Natural history. The Gorongosa National Park is situated in Central Mozambique approx. 100 km south-west of the Zambezi River, and is in the southern part of the great African Rift Valley. Mount Gorongosa is an isolated granite mountain massif rising to 1863 m a.s.l., north-west of the National Park’s plains. At its highest elevations the massif’s plateau is covered with montane grasslands and forest patches dominated by cypresses (Widdringtonia nodiflora); the upper slopes of the mountain (900–1600 m) support a belt of moist evergreen forest. The type series was collected in an area between 900 and 1100 m on the western flanks of the mountain (Figs. 6a–c, 7). The paratypes were collected in a small alcove beneath a boulder pile in the closed-canopy riverine forest beside the Murombodzi Waterfall, and thus in comparatively cool and moist microclimates (Figs. 6c, 7). Some rocks with lizards present were located within the splash zone of the waterfall. Other specimens were observed but not collected at this site. The holotype was collected from a deep narrow crack in a large granite rock (i.e. 8 m ³) in highly degraded grassland below the edge of the rainforest and almost fully exposed to the sun (Fig. 6b). It was the only specimen seen at that site. At both sites the new species lived in syntopy with the cordylid Smaug mossambicus (FitzSimons, 1958). The additional non-type material comes from the Bunga Inselberg (Fig. 6d), a partly eroded inselberg composed mostly of trachyte in the north-western corner of the national park, covered and surrounded by dense Miombo woodland at a relatively low altitude (approx. 212 m a.s.l.). Here the geckos were observed during the day and at night in several places in deep cracks between very large boulders. No geckos were observed outside of these cracks. Syntopic lizards were Smaug mossambicus, the skink Trachylepis margaritifera (Peters, 1854) and the gecko Hemidactylus platycephalus Peters, 1854. Further sympatric, but not syntopic geckos were: Lygodactylus cf. capensis (Smith, 1849), Pachydactylus punctatus Peters, 1854 and Chondrodactylus turneri (Gray, 1864). The new species was observed regularly between April and December 2015. Like A. transvaalica, it is tolerant of conspecifics and frequently several specimens, and sometimes as many as over 20, were observed sheltering in deep rock crevices. One adult female (ZMB 83290) contained two eggs of approximately 11.5 x 7.5 mm. Conservation Status. The population of the new gecko present on the lower slopes of Mount Gorongosa is threatened by illegal deforestation taking place within the riverine forest adjacent to Murombodzi Waterfall. Slashand-burn removal of tall, old growth trees around the rocks may cause changes to the humidity and water availability in these currently very moist, shaded microhabitats that seem to be preferred by the lizards. Reforestation efforts led by Gorongosa National Park and introduction of shade-grown coffee as an alternative to slash-and-burn agriculture still practiced around the mountain will hopefully slow or halt the loss of the native riverine and evergreen vegetation in that area. On Bunga Inselberg the population of A. gorongosa is currently protected within the core area of the National Park and no immediate threat to its survival exists, though deforestation for agriculture is slowly encroaching on the area. However, based on the existence of only a few known sites and small area of occupancy, i.e. the presumed restriction to a fragmented range on Mount Gorongosa and potentially some rocky areas surrounding the mountain, as well as the ongoing high rate of habitat degradation and conversion on the mountain, the species may be considered highly threatened, potentially justifying an assessment as Endangered or even Critically Endangered. Further surveys and formal IUCN Red List assessment are required to confirm this.

In 1944 Loveridge described two new geckos from Angola Afroedura karroica bogerti and Pachydactylus scutatus angolensis. The descriptions of both species have vague and confusing type localities and refinements are suggested based on early expedition reports historical accounts from the region and a review of cartographic material. Numerous new distribution records are reported for both species from expeditions undertaken from 1956–2016 by the authors or their colleagues. The taxonomic status of both species has changed but new material from diverse habitats altitudes and geological substrates indicates that further taxonomic adjustments are likely in order to reflect additional cryptic diversity.

In this first cytogenetic survey on the lamprophiid snake subfamily Pseudoxyrhophiinae, we studied the karyology of ten snake species belonging to seven genera from Madagascar (Compsophis, Leioheterodon, Liophidium, Lycodryas, Madagascarophis, Phisalixella and Thamnosophis) using standard and banding methods. Our results show a wide range of different karyotypes ranging from 2n = 34 to 2n = 46 elements (FN from 40 to 48), with nucleolus organizer regions (NORs) on one (plesiomorphic) or two (derived/apomorphic) microchromosome pairs, and W chromosome at early or advanced states of diversification from the Z chromosome. The observed W chromosome variations further support the most accepted hypothesis that W differentiation from the Z chromosome occurred by progressive steps. We also propose an evolutionary scenario for the observed high karyotype diversity in this group of snakes, suggesting that it is derived from a putative primitive pseudoxyrhophiine karyotype with 2n = 46, similar to that of Leioheterodon geayi, via a series of centric fusions and inversions among macrochromosomes and translocations of micro- either to micro- or to macrochromosomes. This primitive Pseudoxyrhophiinae karyotype might have derived from a putative Lamprophiidae ancestor with 2n = 48, by means of a translocation of a micro- to a macrochromosome. In turn, the karyotype of this lamprophiid common ancestor may have derived from the assumed primitive snake karyotype (2n = 36 chromosomes, with 16 biarmed macro- and 20 microchromosomes) by a series of centric fissions and one inversion. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112, 450–460.

The montane inselbergs of northern Mozambique have been comparatively little-studied, yet recent surveys have shown they have a rich biodiversity with numerous endemic species. Here we present the main findings from a series of scientific expeditions to one of these inselbergs, Mt Mabu, and discuss the conservation implications. Comprehensive species lists of plants, birds, mammals and butterflies are presented. The most significant result was the discovery of a c. 7,880 ha block of undisturbed rainforest, most of it at medium altitude (900–1,400 m), a forest type that is not well represented elsewhere. It is possibly the largest continuous block of this forest type in southern Africa. To date, 10 new species (plants, mammals, reptiles and butterflies) have been confirmed from Mt Mabu, even though sampling effort for most taxonomic groups has been low. The species assemblages indicate a relatively long period of isolation and many species found are at the southern limit of their range. Conservationists are now faced with the challenge of how best to protect Mt Mabu and similar mountains in northern Mozambique, and various ways that this could be done are discussed.

Phylogenetic reconstruction using the mitochondrial 16S marker shows that geographically separated populations of the poorly known Hyperolius cinereus (Anura: Hyperoliidae) from Angola form two distinct clades. The description of H. cinereus was originally based on only a single preserved adult male. Fresh material of both sexes allowed a detailed redescription of the species, which is restricted mainly to the south-draining Cunene and Cubango river systems. Bioacoustic and morphological characters, in conjunction with colouration differences, allow the description of a cryptic sister species from Lagoa Carumbo in north-eastern Angola, occurring in the Luele and Lovuo river systems of the Congo drainage basin. Tadpoles, for H. cinereus and the new species, are described.

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& Geociencias Noroeste Argentino, Salta, Argentina Inst Venezolano Invest Cient, Caracas, Venezuela Umweltbundesamt, Vienna, Austria Penclen, Plumelec, France CNRS MNHN UPMC, Ctr Rech Paleobiodivers & Paleoenvironm, Paris, France Staatl Museum Nat Kunde, Stuttgart, Germany Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivada, Buenos Aires, DF, Argentina Ctr Biol Gest Populat INRA, Montferrier Sur Lez, France Loyola Univ Chicago, Chicago, IL USA Pontificia Univ Catolica Rio do Sul, Porto Alegre, RS, Brazil Durban Museum Nat Sci, Durban, South Africa Univ Estado Amazonas, Manaus, Amazonas, Brazil Brigham Young Univ, Provo, UT 84602 USA Museo Hist Nat La Salle, Caracas, Venezuela Univ Vet Med Hannover, Hannover, Germany Australian Museum, Sydney, NSW, Australia Texas Tech Univ, Lubbock, TX 79409 USA Francis Marion Univ, Florence, SC USA Univ Washington, Seattle, WA 98195 USA Univ Connecticut, Groton, CT USA Fed Univ ABC, Santo Andre, SP, Brazil Univ Fed Uberlandia, Uberlandia, MG, Brazil Fed Univ Para, Belem, Para, Brazil Yale Peabody Museum, New Haven, CT USA Univ Texas Arlington, Arlington, TX 76019 USA Senckenberg Deutsch Entomol Inst, Muncheberg, Germany CSIRO, Natl Res Collect, Canberra, ACT, Australia Univ Venda, Thohoyandou, South Africa Univ Ulsan, Ulsan, South Korea Univ Estadual Sao Paulo, Botucatu, SP, Brazil Senckenberg Forsch Inst & Nat Museum, Frankfurt, Germany Ocean Univ China, Inst Marine Biodivers & Evolut, Qingdao, Shandong, Peoples R China Univ Fed Para, Altamira, Brazil Univ Estadual Paulista, Assis, Brazil UPMC, Ctr Rech Paleobiodiversite & Paleoenvironm, CNRS, MNHN, Paris, France Tech Univ Kaiserslautern, Kaiserslautern, Germany Charles Univ Prague, Dept Zool, Prague, Czech Republic Univ Nacl Expt los Llanos Occident Ezequiel Zamor, Guanare, Venezuela Long Isl Univ, Brooklyn, NY USA Univ Ryukyus, Okinawa, Japan Pontificia Univ Catolica Ecuador, Museo Zool, Escuela Ciencias Biol, Quito, Ecuador Indiana Univ Penn, Indiana, PA USA IRD, Dakar, Senegal State Museum Nat Hist Karlsruhe, Karlsruhe, Germany Univ Toulouse III Paul Sabatier, Toulouse, France Univ Ghent, Ghent, Belgium Univ Antwerp, Antwerp, Belgium Karlsruhe Inst Technol, Karlsruhe, Germany Ctr Cellular & Mol Biol, Hyderabad, India Univ Fed ABC, Santo Andre, Brazil Yale Peabody Museum Nat Hist, New Haven, CT USA Univ Innsbruck, Mondsee, Austria Bethune Cookman Univ, Daytona Beach, FL USA Natl Museum Nat & Sci, Tokyo, Japan Zool Garten Koln, Cologne, Germany Univ Estadual Paulista, Botucatu, SP, Brazil Univ Estadual Paulista, Sao Vicente, Brazil Univ Estadual Paulista, Rio Claro, Brazil Univ Estadual Paulista, Sao Jose Do Rio Preto, Brazil Univ Estadual Paulista, Assis, Brazil

The girdled lizard genus Cordylus is represented in Angola by two species, Cordylus angolensis and C. machadoi, separated from their nearest congeners by over 700 km. Here we describe a new species, Cordylus namakuiyus sp. nov., endemic to the arid lowlands west of the southern Angolan escarpment. Phylogenetic analysis using three mitochondrial and eight nuclear genes shows that the low-elevation forms and the proximate, high-elevation species C. machadoi are genetically divergent and reciprocally monophyletic, and together form the earliest diverging lineage of the northern Cordylus clade. Morphological data, collected using computed tomography and traditional techniques (scalation and morphology), identify consistent phenotypic differences between these high- and low-elevation species and allows for a detailed description of the osteology and osteodermal arrangements of the new species. A series of 50 specimens, collected during the 1925 Vernay expedition to southwestern Angola and housed at the American Museum of Natural History, are assigned to the new species, although the identity of Cordylus from northern Namibia remains ambiguous and requires further investigation.

Using phylogenetic and haplotype network analyses of 2036 bp of mitochondrial DNA, we compare samples of the two hinged terrapin species Pelusios castanoides and P. subniger from continental Africa, Madagascar and the Seychelles to infer their biogeography. Owing to the long independent history of Madagascar and the Seychelles, the populations from those islands should be deeply divergent from their African conspecifics. Seychellois populations of the two species are currently recognized as Critically Endangered endemic subspecies. However, even though we found within P. subniger evidence for a cryptic species from the Democratic Republic of the Congo, all other samples assigned to this species were undifferentiated. This suggests that Malagasy and Seychellois populations of P. subniger were introduced by humans and that the Seychellois subspecies P. s. parietalis is invalid. This has implications for current conservation strategies for the Critically Endangered Seychellois populations and suggests that measures should rather focus on endemic species. The situation of P. castanoides could be different. Samples from Madagascar and the Seychelles are weakly, but consistently, differentiated from continental African samples, and Malagasy and Seychellois samples are reciprocally monophyletic in maximum likelihood analyses. However, due to a lack of samples from central and northern Mozambique and Tanzania, we cannot exclude that identical continental haplotypes exist there.

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 benguellensis 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 Angol...

We examine the phylogeography, phylogeny and taxonomy of hinge-back tortoises using a comprehensive sampling of all currently recognized Kinixys species and subspecies and sequence data of three mitochondrial DNA fragments (2273 bp: 12S rRNA, ND4 + adjacent DNA coding for tRNAs, cytb) and three nuclear loci (2569 bp: C-mos, ODC, R35). Combined and individual analyses of the two data sets using Bayesian and Maximum Likelihood methods suggest that the savannah species of Kinixys are paraphyletic with respect to the rainforest species K. homeana and K. erosa, and that the rainforest species may be derived from a savannah-living ancestor. The previously recognized savannah species K. belliana was a conglomerate of three deeply divergent clades that we treat here as distinct species. We restrict the name K. belliana (Gray, 1830) to hinge-back tortoises ranging from Angola to Burundi, while five-clawed hinge-back tortoises from the northernmost part of the formerly recognized range of K. belliana, together with four-clawed tortoises from West Africa, are assigned to the species K. nogueyi (Lataste, 1886). These two species are allied to K. spekii, whereas Southeast African and Malagasy hinge-back tortoises formerly lumped together with K. belliana represent the distinct species K. zombensis Hewitt, 1931, which is sister to K. lobatsiana. The latter two species together constitute the sister group of the rainforest species K. homeana and K. erosa. Mitochondrial data suggest that K. natalensis has a basal phylogenetic position in a clade embracing K. belliana sensu stricto, K. nogueyi and K. spekii, while nuclear data and the two data sets combined favour a sister group relationship of K. natalensis to all other hinge-back tortoises. Phylogeographic structure is present in all wide-ranging species and correlates in K. homeana and K. erosa with the Dahomey Gap and former rainforest refugia. The Malagasy population of K. zombensis is weakly differentiated from its South African conspecifics and further sampling is needed to determine whether there is support for the subspecific distinctness of Malagasy tortoises.

Currently, four species of the lacertid lizard genus Adolfus are known from Central and East Africa. We sequenced up to 2,825 bp of two mitochondrial (16S and cyt b) and two nuclear (c-mos and RAG1) genes from 41 samples of Adolfus (representing every species), two species each of Gastropholis and Holaspis, and in separate analyses combined this data with GenBank sequences of all other Eremiadini genera and four Lacertini outgroups. Data from DNA sequences were analyzed with maximum parsimony (PAUP), maximum-likelihood (RAxML) and Bayesian inference (MrBayes) criteria. Results demonstrated that Adolfus is not monophyletic: A. africanus (type species), A. alleni and A. jacksoni are sister taxa, whereas A. vauereselli and a new species from the Itombwe Plateau of Democratic Republic of the Congo are in a separate lineage. Holaspis and Gastropholis were recovered in separate clades. Based on this molecular data, relatively substantial sequence divergence and multiple morphological differences, we describe a new genus of lacertid for the lineage including A. vauereselli and the new Itombwe species. The recognition of this new, endemic genus underscores the conservation importance of the Albertine Rift, especially the Itombwe Plateau, a unique region that is severely threatened by unchecked deforestation, mining and poaching.

A new species of gecko of the Pachydactylus weberi complex is described from the NamibRand Reserve in southern Namibia. It is morphologically well differentiated from all other members of this group, in lacking thigh tubercles, and can further be distinguished by its small size (< 45 mm snout–vent length), participation of the first supralabial in the nostril rim, snout scales much larger than interorbital scales, and three body bands in juveniles and adults. It has a distinctive, bold, contrasting hatchling coloration that superficially resembles that of P. mclachlani, from which it is distinguished both morphologically and genetically. DNA sequence data from two nuclear genes (RAG-1, PDC) and the ND2 mitochondrial gene plus its five flanking tRNAs (2,975 bp) reveal that this new species is most closely related, among the taxa sampled, to P. monicae from the lower Orange River valley.

Aim East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia. Location East Africa. Methods We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal. Results Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre-dates the Pliocene (> 6 Ma). These dates are much older than the Plio-Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages. Main conclusions The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa.

The snake family Lamprophiidae Fitzinger (Serpentes: Elapoidea) is a putatively Late Eocene radiation of nocturnal snakes endemic to the African continent. It incorporates many of the most characteristic and prolific of Africa’s non-venomous snake species, including the widespread type genus Lamprophis Fitzinger, 1843 (house snakes). We used approximately 2500 bases of mitochondrial and nuclear DNA sequence data from 28 (41%) of the approximately 68 recognised lamprophiid species in nine of the eleven genera to investigate phylogenetic structure in the family and to inform taxonomy at the generic level. Cytochrome b, ND4 and tRNA gene sequences (mitochondrial) and c-mos sequences (nuclear) were analysed using Maximum Likelihood, Bayesian Inference and Maximum Parsimony methods. The genus Mehelya Csiki, 1903 was paraphyletic with respect to Gonionotophis Boulenger, 1893. To address this, the concept of Gonionotophis is expanded to include all current Mehelya species. The genus Lamprophis emerged polyphyletic: the enigmatic Lamprophis swazicus was sister to Hormonotus modestus from West Africa, and not closely related to its nominal congeners. It is moved to a new monotypic genus (Inyoka gen. nov.). The remaining Lamprophis species occur in three early-diverging lineages. (1) Lamprophis virgatus and the widely distributed Lamprophis fuliginosus species complex (which also includes Lamprophis lineatus and Lamprophis olivaceus) formed a clade for which the generic name Boaedon Dumeril, Bibron & Dumeril, 1854 is resurrected. (2) The water snakes (Lycodonomorphus) were nested within Lamprophis (sensu lato), sister to Lamprophis inornatus. We transfer this species to the genus Lycodonomorphus Fitzinger, 1843. (3) We restrict Lamprophis (sensu strictissimo) to a small clade of four species endemic to southern Africa: the type species of Lamprophis Fitzinger, 1843 (Lamprophis aurora) plus Lamprophis fiskii, Lamprophis fuscus and Lamprophis guttatus.

Girdled lizards (Cordylidae) are sub-Saharan Africa's only endemic squamate family and contain 80 nominal taxa, traditionally divided into four genera: Cordylus, Pseudocordylus, Chamaesaura and Platysaurus. Previous phylogenetic analysis revealed Chamaesaura and Pseudocordylus to be nested within Cordylus, and the former genera were sunk into the later. This taxonomic revision has received limited support due to the study's poor taxon sampling, weakly supported results and possible temporary nomenclatural instability. Our study analyzes three nuclear and three mitochondrial genes from 111 specimens, representing 51 in-group taxa. Parsimony, likelihood and Bayesian analyses of concatenated and partitioned datasets consistently recovered a comb-like tree with 10, well-supported, monophyletic lineages. Our taxonomic reassessment divides the family into 10 genera, corresponding to these well-supported lineages. Short internodes and low support between the non-platysaur lineages are consistent with a rapid radiation event at the base of the viviparous cordylids.

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.

Jackson's Forest Lizard (Adolfus jacksoni) is widespread throughout the highlands of the Albertine Rift, southern Uganda, western and central Kenya, and northern Tanzania. To understand th...

Nadzikambia is a monotypic genus described to accommodate the Mulanje chameleon, N. mlanjensis. During herpetological surveys of isolated mountains in Mozambique a population of chameleons was discovered on Mount Mabu. It is referable to the genus Nadzikambia, but features of scalation, colouration and genetic divergence distinguish this population from N. mlanjensis. It is described as a new species, Nadzikambia baylissi, and represents the second species in the genus and the first record of the genus in Mozambique. Notes on reproduction in N. mlanjensis are appended. The discovery of the new species in a mid-altitude evergreen forest remnant on Mount Mabu emphasises the high conservation importance of the region, previously signalled by the discovery of a new species of forest viper (Atheris mabuensis), and a number of undescribed butterflies and freshwater crab species in the region.

We investigated the phylogeographic differentiation of the widely distributed African helmeted terrapin Pelomedusa subrufa based on 1503 base pairs of mitochondrial DNA (partial cyt b and ND4 genes with adjacent tRNAs) and 1937 bp of nuclear DNA (partial Rag1, Rag2, R35 genes). Congruent among different analyses, nine strongly divergent mitochondrial clades were found, representing three major geographical groupings: (1) A northern group which includes clades I from Cameroon, II from Ghana and Ivory Coast, III from Benin, Burkina Faso and Niger, IV from the Central African Republic, and V from Kenya, (2) a northeastern group consisting of clades VI from Somalia, and VII from Saudi Arabia and Yemen, and (3) a southern group comprising clade VIII from Botswana, the Democratic Republic of Congo, Madagascar and Malawi, and clade IX from South Africa. Malagasy and continental African populations were not clearly differentiated, indicating very recent arrival or introduction of Pelomedusa in Madagascar. The southern group was in some phylogenetic analyses sister to Pelusios, rendering Pelomedusa paraphyletic with respect to that genus. However, using partitioned Bayesian analyses and sequence data of the three nuclear genes, Pelomedusa was monophyletic, suggesting that its mitochondrial paraphyly is due to either ancient introgressive hybridization or phylogenetic noise. Otherwise, nuclear sequence data recovered a lower level of divergence, but corroborated the general differentiation pattern of Pelomedusa as revealed by mtDNA. This, and the depth of the divergences between clades, indicates ancient differentiation. The divergences observed fall within, and in part exceed considerably, the differentiation typically occurring among chelonian species. To test whether Pelomedusa is best considered a single species composed of deep genealogical lineages, or a complex of up to nine distinct species, we suggest a future taxonomic revision that should (1) extend the geographical sampling of molecular data, specifically focusing on contact zones and the possible sympatric occurrence of lineages without admixture, and (2) evaluate the morphology of the various genealogical lineages using the type specimens or topotypical material of the numerous junior synonyms of P. subrufa.

This study constitutes the first evolutionary investigation of the snake family Psammophiidae--the most widespread, most clearly defined, yet perhaps the taxonomically most problematic of Africa's family-level snake lineages. Little is known of psammophiid evolutionary relationships, and the type genus Psammophis is one of the largest and taxonomically most complex of the African snake genera. Our aims were to reconstruct psammophiid phylogenetic relationships and to improve characterisation of species boundaries in problematic Psammophis species complexes. We used approximately 2500 bases of DNA sequence from the mitochondrial and nuclear genomes, and 114 terminals covering all psammophiid genera and incorporating approximately 75% of recognised species and subspecies. Phylogenetic reconstructions were conducted primarily in a Bayesian framework and we used the Wiens/Penkrot protocol to aid species delimitation. Rhamphiophis is diphyletic, with Rhamphiophis acutus emerging sister to Psammophylax. Consequently we transfer the three subspecies of Rhamphiophis acutus to the genus Psammophylax. The monotypic genus Dipsina is sister to Psammophis. The two species of Dromophis occupy divergent positions deeply nested within Psammophis, and we therefore relegate Dromophis to the synonymy of Psammophis. Our results allow division of the taxonomically problematic Psammophis 'sibilans' species complex into two monophyletic entities, provisionally named the 'phillipsii' and 'subtaeniatus' complexes. Within these two clades we found support for the status of many existing species, but not for a distinction between P.p. phillipsii and P. mossambicus. Additionally, P. cf. phillipsii occidentalis deserves species status as the sister taxon of P. brevirostris.

Homopus bergeri Lindholm 1906 was based on a single partial shell from Namibia. No other Homopus were recorded from the country until a population of Homopus was discovered in the Aus region, southern Namibia, in 1955. It was originally referred to H. boulengeri Duerden 1906, and then inaccurately to H. bergeri. Inspection of the type of H. bergeri Lindholm 1906 shows that it is a junior synonym of Psammobates tentorius verroxii (A. Smith 1839). The Aus Homopus is morphologically distinguishable from all other Homopus and is described as a new species, Homopus solus. Morphological variation, distribution and biology in H. solus are described. The use of the combinations Homopus ‘solos’ by Devaux (2003) and Homopus bergeri (solos) (Lindholm, 1906) by Bonin et al. (2006) have no nomenclatural standing.