An island in a sea of sand: a first checklist of the herpetofauna of the Serra da Neve inselberg, southwestern Angola.

The Serra da Neve inselberg in Namibe Province, southwestern Angola is the second highest peak of Angola with an elevation of 2489 m. It remains one of the least explored regions in the country, despite several endemic species having been recently described from this inselberg. Here we provide an inventory of the amphibian and reptile species ocurring in Serra da Neve and compare its fauna with that of the surrounding habitats at lower elevations. We also examine the phylogenetic affinities of the inselberg taxa. A total of 59 herpetological taxa were recorded for the Serra da Neve inselberg and its immediate surroundings. These include 11 species of amphibians, belonging to nine genera and seven different families, and 48 species of reptiles, belonging to 32 genera and 12 families. Of these, one amphibian and seven reptiles from seven different genera are strictly endemic, making the inselberg the richest region in southwestern Africa with respect to strict endemics, with one endemic reptile taxa per 127 km2. Not surprisingly, most of the recorded taxa belong to clades that are endemic, or at least strongly associated, with southern Africa, but two are representatives of central African clades, and another two are more closely related to eastern African highland taxa. We also provide comments on the threats to the conservation of this endemic-rich inselberg.

Lack of country-wide systematic herpetology collections in Portugal jeopardizes future research and conservation.

Natural History Collections (NHCs) represent the world's largest repositories of long-term biodiversity datasets. Specimen collection and voucher deposition has been the backbone of NHCs since their inception, but recent decades have seen a drastic decline in rates of growth via active collecting. Amphibians and reptiles are amongst the most threatened zoological groups on the planet and are historically underrepresented in most worldwide NHCs. As part of an ongoing project to review the Portuguese zoological collections in the country's NHCs, herpetological data from its three major museums and smaller collections was gathered and used to examine the coverage and representation of the different taxa extant in Portugal. These collections are not taxonomically, geographically, or temporally complete. Approximately 90% of the Portuguese herpetological taxa are represented in the country's NHCs, and around half of the taxa are represented by less than 50 specimens. Geographically, the collections cover less than 30% of the country's territory and almost all of the occurring taxa have less than 10% of their known distribution represented in the collections. A discussion on the implications for science of such incomplete collections and a review of the current status of Portuguese NHCs is presented.

Historical natural history collections allow the description of a new and presumably extinct species of dwarf gecko (Squamata: Gekkonidae: Lygodactylus Gray, 1864) from Fernando de Noronha Island, Brazil.

A series of specimens of an unidentified species of the genus Lygodactylus from Fernando de Noronha Island were found in the herpetological collections of the Museu Nacional, Rio de Janeiro (Brazil). No species of this genus were known to occur in the archipelago. A review of the historical reports regarding the herpetofauna of the island failed to provide evidence regarding the presence of the species in the last centuries. Morphological comparisons with the two other species of the genus occurring in South America, L. klugei and L. wetzeli, allowed us to confidently confirm that the Fernando de Noronha population belonged to a putatively new species. Here we describe this population as a new species and discuss its possible extinction causes in the archipelago. We also debate the importance of historical natural history collections to the study of biodiversity.

Tightening the requirements for species diagnoses would help integrate DNA-based descriptions in taxonomic practice.

Modern advances in DNA sequencing hold the promise of facilitating descriptions of new organisms at ever finer precision but have come with challenges as the major Codes of bionomenclature contain poorly defined requirements for species and subspecies diagnoses (henceforth, species diagnoses), which is particularly problematic for DNA-based taxonomy. We, the commissioners of the International Commission on Zoological Nomenclature, advocate a tightening of the definition of "species diagnosis" in future editions of Codes of bionomenclature, for example, through the introduction of requirements for specific information on the character states of differentiating traits in comparison with similar species. Such new provisions would enhance taxonomic standards and ensure that all diagnoses, including DNA-based ones, contain adequate taxonomic context. Our recommendations are intended to spur discussion among biologists, as broad community consensus is critical ahead of the implementation of new editions of the International Code of Zoological Nomenclature and other Codes of bionomenclature.

Saving collections: taxonomic revision of the herpetological collection of the Instituto de Investigação Científica Tropical, Lisbon (Portugal) with a protocol to rescue abandoned collections.

The herpetological collections of the Instituto de Investigação Científica Tropical, Lisbon, are amongst the most important collections from the former Portuguese territories in Africa and Asia. The collection comprises more than 5000 preserved specimens, including type specimens of nine taxa, Trachylepis adamastor, Trachypelis thomensis, Panaspis thomensis, Naja peroescobari, Dalophia angolensis, Hemidactylus nzingae, Boaedon fradei, Platysaurus maculatus maculatus, and Platysaurus maculatus lineicauda. The collection was abandoned in the early years of 2000s and was at risk of being lost. In this paper the entire collection is reviewed, a catalogue provided of the extant specimens, and a brief account of the history of herpetological research at IICT given. Details are also provided on the recovery of the collection and a protocol to rescue abandoned collections.

How the African house gecko (Hemidactylus mabouia) conquered the world.

Alien species are among the greatest threats to biodiversity, but the evolutionary origins of invasiveness remain obscure. We conducted the first range-wide sampling of Hemidactylus mabouia from more than 120 localities across Africa, Madagascar and the Neotropics to understand the evolutionary history of one of the most widely distributed, invasive vertebrates in the world. We used a multi-locus phylogeny, species delimitation, fossil-calibrated timetree, ancestral area reconstruction and species distribution models (SDMs) to determine how many putative species-level lineages are contained within H. mabouia, the timing and tempo of diversification, and the origins of commensality-providing insights into the evolutionary origins of invasiveness. Our analyses suggest 'H. mabouia' originated in the Miocene in the Zambezian biogeographic region and includes as many as 20 putative species-level lineages, of which only Hemidactylus mabouia sensu stricto is invasive and widely distributed, including all Neotropical records. Zambezia is the hotspot for diversity within the group with 14 species in southeastern Zambezia. SDMs suggest that H. mabouia was able to establish in the Neotropics due to habitat suitability, and globalization and the slave trade probably allowed it to cross the Atlantic. Distribution models for the H. mabouia complex overpredict the range of the invasive H. mabouia sensu stricto-highlighting the importance of taxonomy in invasive species management.

A new species of Rain Frog (Brevicipitidae, Breviceps) endemic to Angola.

Recent molecular phylogenetic work has found that Breviceps Merrem, 1820 comprises two major clades, one of which, the B. mossambicus group, is widely distributed across southern sub-Saharan Africa. This group is notable for harboring abundant cryptic diversity. Of the four most recently described Breviceps species, three are members of this group, and at least five additional lineages await formal description. Although Breviceps has long been known to occur in Angola, no contemporary material has been collected until recently. The three most widespread taxa, B. adspersus, B. mossambicus, and B. poweri, may all occur in Angola, but accurate species assignment remains challenging given the rampant morphological similarity between these taxa, and, until recently, the lack of genetic resources. Phylogenetic, morphological, and acoustic analyses of recently collected samples from disparate localities within Angola provide evidence for an undescribed species that is sister to B. poweri. The new species can be diagnosed from its sister taxon by lacking pale spots along the flanks, a pale patch above the vent, and a short, dark band below the nares (all present in B. poweri). Additionally, the male advertisement call differs from the three other Breviceps that might occur in Angola in having both a longer interval between consecutive calls and a higher average dominant frequency. We here describe this lineage as a distinct species, currently only known from Angola, and discuss the presence of other Breviceps taxa within Angola.

ResumoInvestigações moleculares recentes revelaram que o género Breviceps Merrem, 1820, é composto por duas linhagens principais, uma das quais, o grupo B. mossambicus, é amplamente distribuído na região sul da África subsaariana. Este grupo é notável por albergar uma abundante diversidade críptica. Das quatro espécies de Breviceps recentemente descritas, três pertencem a este grupo, e pelo menos outras cinco linhagens adicionais aguardam a sua descrição formal. Apesar de o género ser conhecido de Angola desde há muito tempo, só muito recentemente foram colhidos novos espécimes. Os três taxa mais amplamente distribuídos, B. adspersus, B. mossambicus e B. poweri podem todos, porventura, ocorrer em Angola, no entanto a correta identificação destas espécies têm sido problemática devido às semelhanças morfológicas extremas entre este taxa, e, até muito recente, a completa ausência de material genético. Análises filogenéticas, morfológicas e acústicas dos espécimes recentemente colhidos em diferentes locais de Angola apontam para a existência de uma espécie nova para a ciência, irmã de B. poweri. A nova espécie pode ser diferenciada do seu táxon irmão pela falta de marcas pálidas nos flancos, mancha pálida acima do ventre e pequena banda negra abaixo do nariz (presentes em B. poweri). Para além destas características, o chamamento dos machos difere das outras três espécies de Breviceps que podem ocorrer em Angola por ter um maior intervalo entre chamamentos consecutivos e uma maior frequência média dominante. Descrevemos aqui esta linhagem como uma espécie distinta, atualmente apenas conhecida de Angola, e discutimos a presença de outras espécies de Breviceps em Angola.

A phylogenetic study of Haemocystidium parasites and other Haemosporida using complete mitochondrial genome sequences.

Haemosporida are diverse vector-borne parasites associated with terrestrial vertebrates. Driven by the interest in species causing malaria (genus Plasmodium), the diversity of avian and mammalian haemosporidian species has been extensively studied, relying mostly on mitochondrial genes, particularly cytochrome b. However, parasites from reptiles have been neglected in biodiversity surveys. Reptilian haemosporidian parasites include Haemocystidium, a genus that shares morphological features with Plasmodium and Haemoproteus. Here, the first complete Haemocystidium mitochondrial DNA (mtDNA) genomes are studied. In particular, three mtDNA genomes from Haemocystidium spp. sampled in Africa, Oceania, and South America, are described. The Haemocystidium mtDNA genomes showed a high A + T content and a gene organization, including an extreme fragmentation of the rRNAs, found in other Haemosporida. These Haemocystidium mtDNA genomes were incorporated in phylogenetic and molecular clock analyses together with a representative sample of haemosporidian parasites from birds, mammals, and reptiles. The recovered phylogeny supported Haemocystidium as a monophyletic group apart from Plasmodium and other Haemosporida. Both the phylogenetic and molecular clock analyses yielded results consistent with a scenario in which haemosporidian parasites radiated with modern birds. Haemocystidium, like mammalian parasite clades, seems to originate from host switches by avian Haemosporida that allowed for the colonization of new vertebrate hosts. This hypothesis can be tested by investigating additional parasite species from all vertebrate hosts, particularly from reptiles. The mtDNA genomes reported here provide baseline data that can be used to scale up studies in haemosporidian parasites of reptiles using barcode approaches.

A correction to a recent review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two additional species.

A correction to the recent revision of the genus Hemidactylus Goldfuss, 1820 from Angola is presented. The previous revision assigned some historical specimens to H. bayonii and H. nzingae, with one specimen even being considered a paratype of the latter taxon. A review of the morphological data of all of the specimens used in the previous study found that two other undescribed taxa, morphologically similar to H. bayonii and H. nzingae, respectively, were overlooked. We here present a re-description of these specimens and provide the diagnostic characters that allow their recognition as two new species, both endemic to Angola. In order to provide an accurate picture of the diversity of Hemidactylus in the country and to correct our previous misidentifications that could lead to some taxonomic instability, we herein describe H. hannasabinae sp. nov. and H. vernayi sp. nov.

A revision of the Dwarf Geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with the description of three new species.

At present the genus Lygodactylus is represented by three species in Angola confirmed by voucher specimens-L. angolensis, L. bradfieldi, and L. capensis-and two others believed to be present, but without specimens with precise localities, L. chobiensis and L. lawrencei. We present a detailed taxonomic revision of the group in Angola and describe three new species, Lygodactylus baptistai sp. nov. L. nyaneka sp. nov. and Lygodactylus tchokwe sp. nov. Phylogenetic analysis using the mitochondrial marker ND2, as well as morphological data support the recognition of the new species. In addition, data suggest that specimens historically assigned to L. capensis in Angola represent misidentifications of L. nyaneka sp. nov. and L. tchokwe sp. nov. We revisit the identity of Lygodactylus laurae, a junior synonym of L. angolensis. We also present the first confirmed record of L. lawrencei in the country, using both morphological and molecular data. The description of the new species and the revision of the taxonomic identity of the Angolan populations of the genus, raises the number of species occurring in the country to five. A key to the Angolan species is presented.

The Bush Vipers, genus Atheris Cope, 1862 (Squamata: Viperidae) of Bioko Island, Gulf of Guinea, with the description of a new species.

Two species of Bush Vipers, genus Atheris Cope, 1862, have been reported for the continental island of Bioko, Gulf of Guinea - Atheris squamigera and a putatively undescribed species, morphologically similar to Atheris chlorechis. The latter was only known from one specimen collected in the early 1900s and its taxonomic identity has never been fully resolved. Based on recently collected specimens of Atheris from Bioko Island, we reviewed all the available data and specimens through detailed morphological comparisons against all known species of the genus. Our results confirm that Atheris squamigera is present in Bioko Island, and that the second taxon is a new species, which we describe here as Atheris hetfieldi sp. nov. The newly described species presents a series of morphological characters that clearly differentiate it from all of its congeners. The description of this new species is the first of a new snake species from Bioko in more than 100 years and this species is the only snake currently recognized as endemic to the island.

A review of the African snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species.

The genus Panaspis in Angola is represented by four species, most of them part of taxonomically and nomenclaturally challenging species-complexes. We present a taxonomic revision of the group in the region and describe one new species, Panaspis mocamedensis sp. nov., endemic to the lowland areas of the Namibe province, southwestern Angola. Phylogenetic analysis using a combination of mitochondrial (16S, cytb) and nucleares (RAG1, PDC) markers, as well as morphological and meristic data support the recognition of the new species. In addition, these data support the presence of nominotypical Panaspis cabindae, P. wahlbergi and P. maculicollis in Angola. Reexamination of the Angolan population of P. breviceps was based on morphological analysis, as no molecular data from Angola is available for this species. According to our results, this population likely represents the nominotypical form, but due to its complex taxonomic and nomenclatural history and the lack of molecular data, this population needs to be reconsidered when molecular data become available. The description of a new species and revision of the Angolan Panaspis contributes to a better understanding of the true species richness of the Angolan herpetofauna, as well as to understanding the major biogeographic patterns of the region. A key to Angolan Panaspis species is also presented.

A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species.

The genus Hemidactylus in Angola is represented by six species, all of them part of taxonomically and nomenclaturally challenging species complexes. We present a detailed taxonomic revision of the group in the region and describe two new species, Hemidactylus nzingae sp. nov. and Hemidactylus paivae sp. nov., both occuring in and potentially endemic to the highlands of Angola. Phylogenetic analysis using a combination of mitochondrial (ND2) and nuclear (MXRA5, PDC, RAG1) markers, as well as morphological and scalation data support the recognition of the new species. In addition, data support the revalidation of Hemidactylus bayonii Bocage, 1893, and Hemidactylus benguellensis Bocage, 1893. We also provide a redefinition of Hemidactylus longicephalus Bocage, 1873 with which we synonymize Hemidactylus mabouia molleri Bedriaga, 1892, from São Tomé in the Gulf of Guinea. Given that the type material of H. bayonii, H. benguellensis, H. longicephalus and H. mabouia molleri have all been lost or destroyed, we designate neotypes for all of these nomina for purposes of nomenclatural stability. The description of the new species and the revision and revalidation of the Angolan species already described contributes to a better understanding of the taxonomy and biogeography of West and Central African Hemidactylus, as well as to the general biogeographic and evolutionary patterns of Angolan fauna. A key to the Angolan species is also presented.

Description of a new long-tailed skink (Scincidae: Trachylepis) from Angola and the Democratic Republic of the Congo.

Angola and the Democratic Republic of the Congo are relatively unknown in terms of their herpetological diversity. Based on specimens collected in the Congolese region of the Katanga and the northeast of Angola during the first decades of the twentieth century, de Witte and Laurent independently suggested, based on morphological and coloration differences, that populations of T. megalura of these regions could belong a new "race". We compared specimens of T. megalura (including the type specimens of T. megalura and T. massaiana) with Angolan and Katangan museum specimens as well as newly collected specimens from Angola. Coloration pattern and morphological characters, in combination with substantial divergence in the 16S mitochondrial gene, confirm the distinctiveness of the west Central African form, and it is here described as a new species. Data regarding its natural history, ecology and global distribution are presented.

A new species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola.

A new species of rupicolous girdled lizard is described from Serra da Neve Inselberg, Namibe Province, southwestern Angola. We sequenced two mitochondrial gene regions (16S and ND2) for the new species and compared these data with those previously published for other Cordylus species. The new species is genetically divergent from the closely related Cordylus machadoi and C. namakuiyus and morphologically distinguished by head scale arrangement, coloration, and osteological characters. Serra da Neve is the second highest peak in Angola and one of the most understudied areas of the country. The discovery of this new species highlights the importance of this inselberg for regional endemism and emphasizes the high conservation importance of the area.

A new earless species of Poyntonophrynus (Anura, Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola.

African pygmy toads of the genus Poyntonophrynus are some of the least known species of African toads. The genus comprises ten recognized species endemic to sub-Saharan Africa, five of which are restricted to southwestern Africa. Recent field research in Angola provided new material for three species of Poyntonophrynus, including a morphologically distinctive population from the Serra da Neve Inselberg. Based on a combination of external morphology, high-resolution computed tomography scanning, and molecular phylogenetic analysis, the Serra da Neve population is described as new species that is nested within the genus. The most striking character that differentiates the newly described species from its congeners is the lack of a tympanic middle ear, a condition common in the family Bufonidae, but so far not known for Poyntonophrynus. The description of this new species from southwestern Angola reinforces the biogeographic importance of the region and further suggests that southwestern Africa is the cradle of diversity for this genus.

Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of São Tomé, Príncipe and Annobon islands.

Five problematic cases regarding the presence of reptile taxa on the island of São Tomé (Gulf of Guinea) are reviewed. These cases deal with 1) the past presence of crocodilians in the island, 2) the putative type locality of Dipsas pulverulenta Fischer, 1856, 3) the dubious presence of a mamba (genus Dendroaspis) on the island, 4) an erroneous reference to the presence of Gastropyxis smaragdina, and 5) two specimens (Psammophis lineatus and Psammophis elegans) from Carl Weiss collection, currently deposited in the Zoological Museum of Hamburg and wrongly labelled as being from São Tomé island. A provisional checklist to the terrestrial herpetofauna of the three oceanic islands of the Gulf of Guinea-São Tomé, Príncipe and Annobon-is presented in order to clarify the current numbers of the endemics of each island, as well as to review and present current research.

A new species of African snake-eyed skink (Scincidae: Panaspis) from central and northern Namibia.

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.