PROCEEDINGS of the (Series 4) 2018 * Volume 65 * Supplement II Instifute for Biodii^ersity Science & Sustainability CALIFORNIA ACADEMY OF SCIENCES Copyright © 2018 by the California Academy of Sciences All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without permission in writing from the publisher. Scientific Publications Publisher: Shannon Bennett, Ph.D. Chief of Science and Research Collections California Academy of Sciences Editorial Board Alan E. Leviton, Ph.D., Editor Katherine Piatek, M.A., Managing Editor Michael T. Ghiselin, Ph.D., Associate Editor Tomio Iwamoto, Ph.D., Associate Editor Gary C. Williams, Ph.D., Associate Editor & Website Coordinator Cover images Front cover: Python anchietae from Omahua, Namibe Province, Angola. Photo by Luis M.P. Ceriaco Cover Design Gary C. Williams & Alan E. Leviton California Academy of Sciences ISSN 0068-547X 'Tfo ‘P’toeeediMf^' tie ^4eadem^ Scieeeee is an international journal that accepts manuscripts for publication in the Natural Sciences and selected areas in the Earth Sciences, such as bios¬ tratigraphy, regional and global tectonics as they relate to biogeography, and paleoclimatology, and topics in astrobiology, anthropology, as well as the history of science as they relate to institutions of natural his¬ tory, to individuals, and to activities, such as expeditions and explorations, in the natural sciences. All manuscripts submitted for publication in any of the Academy's scientific publication series {Pro¬ ceedings, Occasional Papers, Memoirs) are subject to peer review. Peer review includes both internal and external review, internal review by at least one Academy scientist whose interests parallel those of the sub¬ mission, and external review, ordinarily by two individuals who are recognized scholars in the field. Manuscripts accepted for publication are subject to page charges; charges may be waived on a case- by-case basis. Published by the California Academy of Sciences 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118 U.S.A. Printed in the United States of America by Allen Press Inc., Lawrence, Kansas 66044 PROCEEDINGS OF THE CALIFORNIA AC ADEMY OF SCIENCES Series 4, Volume 65, Supplement II, pp. I-50I, 413 figs., 5 tables. Appendix. September 28, 2018 Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola Atlas of Historical and Bibliographic Records (1840-2017) Mariana P. Marques Luis M. P. Ceriaco David C. Blackburn and Aaron M. Bauer 1 Centro de Investigagdo em Biodiversidade e Recursos Geneticos {CIBIO), INBIO, Universidade do Porto, Rua Padre Armando Quintas 7, Vairdo, 4485-661 Porto, Portugal ^ Departamento de Zoologia e Antropologia (Museu Boeage), Museu Naeional de Historia Natural e da Cieneia, Universidade de Lisboa, Rua da Eseola Politeeniea 58, 1269-102 Lisboa, Portugal ^ Department of Biology, Villanova University, 800 Laneaster Avenue, Villanova, Pennsylvania 19085, USA. ^ Department of Natural History, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA. ^ Researeh Assoeiate, California Aeademy of Seienees, 55 Musie Coneourse Drive, San Franeiseo, California, 94118, USA. FOREWORD (English version) The rieh and distinetive Angolan herpetofauna has engendered the interest of eountless sehol- ars around the world. Angola’s unique geographie loeation, at the interseetion of the eentral and southern regions of the Afriean eontinent, has given it two diserete faunas of amphibians and reptiles. Of different eolors, sizes, forms, and behaviors, these speeies have, over the years, entieed biologists, explorers and seientists to boldly and selflessly dedieate themselves to their study. The interest shown by speeialists in the study of Angolan speeimens dates baek more than one hundred years. Sinee the nineteenth eentury, books and papers have been published on the topie as the result of these investigations. The first major bibliographie referenee about the Angolan herpetofauna, Herpetologie d’ Angola et du Congo by Jose Vieente Barbosa du Boeage, dates from 1895. Many researchers and publications followed, greatly enriching the scientific bibliography on the topic. However, the scattered nature of the published works, as well as the dearth of available specimens from earlier times, has made it difficult for researchers, scholars or curious amateurs to access to this information. Because of these difficulties, and to facilitate the access to available data on the Angolan herpetofauna, the Ministry of Environment, through its National Institute of Biodiversity and Conservation Areas, decided to present, in a single volume, all the available scientific records published so far on the diversity of amphibians and reptiles. Such a task, which has its basis in the National Strategy and Action Plan for Biodiversity and Conservation Areas of Angola, was dependent upon the helpful collaboration of important international partners. The present Atlas will be of inestimable assistance to the study of the Natural History of Ango¬ la, not only because of its standardized taxonomic and geographical accounts with distribution maps for all of the amphibians and reptiles known in Angola, but also because of its competent approach to the physiography, climate, biomes, historical herpetological research in Angola, conservation, and other topics. The thoughtful way the book is arranged, with straightforward text, clear tables and interesting figures, makes it easy for even laymen to read and understand. At 501 pages long, profusely documented and richly illustrated, this Atlas provides a grand 1 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II guided tour through the diversity of the Angolan herpetofauna, from the most remote and spectac¬ ular places of the country. Through mountains and escarpments, plains and valleys, savannas and dense forests, rivers and streams, lakes and lagoons, this work encompasses the entire country, aking us on an enlightening tour of knowledge. The methodical labor and persistence of the specialists who produced the present work, ensure that it not only contributes to scientific history, through its compilation of research already completed, but also suggests future directions for researchers and scholars devoted to the topic. Further, this substantial volume will also inform the national authorities with respect to environ¬ ment protection and conservation policies. Given all this, one can see that the present work, reflecting the contributions of the men and women who created it, is destined to become a standard reference to the scientific literature of the country, especially in regard to this part of its fauna. The quality of the scientific information poured into the following pages certainly qualifies the authors of this accurate work as leading herpetologists and places them among the important researchers to have served Angola in the course of its scientific history. (Portuguese version) A riqueza e as caractensticas da herpetofauna angolana, despertaram e continuam a desper- tar, o interesse de inumeros estudiosos em todo o Mundo. A privilegiada localizagdo geogrdfica que coloca Angola nas regides central e meridional do continente africano faz do pais um caso raro, ao acolher no seu regago animals de dois espagos diferentes do continente africano, benefi- ciando por isso de uma deslumbrante variedade de repteis e anfibios, como poucos paises. De cores, tamanhos, formatos e comportamentos diferentes, esses especimes levaram a que bidlogos, exploradores e cientistas se debrugassem sobre o seu estudo, de modo denodadamente abnegado e por longo tempo. O interesse manifestado por especialistas no estudo de especimes angolanos vem de ha mais de uma centena de anos. Assim, em resultado dessas investigagdes, desde o seculo XIX sdo publi- cados livros e opusculos sobre o assunto. A primeira grande referenda bibliogrdfica relativa a herpetofauna angolana data de 1895 e intitula-se Herpetologie d’Angola et du Congo, da autoria de Jose Vicente Barbosa du Bocage. Seguiram-se-lhes vdrios investigadores e obras que enrique- ceram sobremaneira a bibliografia cientifica sobre o assunto, ao longo de mais de um seculo. A natureza esparsa das obras publicadas sobre o tema, assim como a exiguidade de exempla- res de tempos de antanho disponiveis tornam de certo modo dificil o acesso a um conjunto de infor- magdes por parte de investigadores, estudiosos ou simples curiosos. Em razdo desses escolhos e para facilitar o contacto com os elementos informativos jd existentes sobre a herpetofauna ango¬ lana, o Ministerio do Ambiente, atraves do seu Institute Nacional da Biodiversidade e Areas de Conservagdo, decidiu coligir num unico volume todos os dados cientificos ate agora publicados e disponiveis sobre a diversidade de repteis e anfibios. Para tamanha empreitada, que encontra estribo na Estrategia e Plano de Agdo Nacional para a Biodiversidade e Areas de Conservagdo de Angola, socorreu-se da prestimosa colaboragdo cientifica de importantes parceiros internacionais. O presente Atlas representa um inestimdvel auxilio para o estudo da Histdria Natural de Angola ndo so devido d metodica sectorizagdo temdtica em que se destaca a apresentagdo de fichas taxondmicas e geogrdficas com mapas de ocorrencia para todos os anfibios e repteis conhe- cidos em Angola, mas tambem pela habilitada abordagem da fisiografia, do clima, dos biomas, da investigagdo histdrica da herpetofauna angolana, da conservagdo, entre outros. Ndo sera despi- ciendo referir que o modo cuidadoso como foi configurado o livro, com textos arejados, quadros MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 3 claws e gravuras interess antes, mostra a inequlvoca facilidade de leitura e compreensdo sobre a materia ate mesmo para um leigo. Atraves destas quase 500 pdginas, profusamente documentadas com dados numericos e rica- mente ilustradas, o Atlas proporciona-nos uma arrebatante visita guiada pela diversidade da herpetofauna angolana, levando-nos aos mats reconditos e inimagindveis cantos do pals. Por entre montanhas e escarpas, planlcies e vales, savanas e densas florestas, rios e riachos, lagos e lagoas, a obra perpassa o pals de les-a-les, transportando-nos num enleante tour de conhecimento. O labor metddico e a sagacidade dos especialistas que produziram a presente obra fazem com que a mesma cumpra com primores de requinte a dupla fungdo utilitdria de fazer histdria cientlfi- ca, atraves da apresentagdo de diversas obras de investigagdo aqui elencadas, e indicar caminhos para pesquisadores e estudiosos da materia interessados, tal e forte a sua componente documen¬ tal que vem cobrir uma grande lacuna hd muito existente. Este nutrido volume tern ainda o conddo de facilitar as autoridades para a concepgdo de politicas de Protegdo e Conservagdo Ambiental. Por tudo isso, e fdcil inferir que a presente obra, na qual luzem intensamente as prendas dos homens e mulheres que a corporizaram, dispoe dos necessdrios condimentos para ser uma refe¬ renda na literatura cientifica do pais, mormente no que diz respeito a este importante segmento da sua fauna. A qualidade da informagdo cientifica vertida ao longo das pdginas que se seguem eleva certamente os autores deste apurado trabalho d condigdo de obreiros de primeira grandeza no piano da Herpetologia, sendo pressentivel que ndo muito tarde no tempo enfileirar-se-do, segu- ramente, sem desdouro ao lado dos mats sonantes nomes que configuram o escol dos grandes investigadores que serviram Angola ao longo da sua histdria cientifica. Her Excellency Minister of the Environment of the Republic of Angola Sua Excelencia Ministra do Ambiente da Republica de Angola OAtr-" Paula Cristina Francisco Coelho Luanda, April 2018 Luanda, Abril 2018 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II In honor of Jose Vicente Barbosa du Bocage (1823-1907), forefather of Angolan herpetology MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 5 Table of Contents Foreword . 1 Abstract . 9 Introduction . 9 The Angolan landscape . 11 Political and administrative borders of Angola . 11 Physiography . 12 River Basins . 13 Geology . 14 Climate . 14 Biomes and vegetation zones . 14 History of herpetological research in Angola . 23 Diversity, Distribution and Endemism . 37 Conservation . 48 Materials and Methods . 55 Data . 55 Mapping species occurrence . 55 Account standardization . 56 Abbreviations and Acronyms . 56 Taxonomic accounts . 59 Amphibia . 59 Order Anura . 59 Family Pipidae . 59 Genus Xenopus . 59 Family Bufonidae . 66 Genus Mertensophryne . 75 Genus Poyntonophrynus . 76 Genus Schismaderma . 78 Genus Sclerophrys . 66 Family Microhylidae . 79 Genus Phrynomantis . 79 Family Brevicipitidae . 81 Genus Breviceps . 81 Family Hemisotidae . 83 Genus Hemisus . 83 Family Hyperoliidae . 85 GQxms Afrixalus . 85 Genus Cryptothylax . 88 Genus Hyperolius . 89 GQvms Kassina . 116 Family Arthroleptidae . 118 Genus Arthroleptis . 118 Genus Leptopelis . 121 Genus Trichobatrachus . 127 Family Ptychadenidae . 128 Genus Ptychadena . 128 Genus Hildebrandtia . 142 Family Phrynobatrachidae . 144 Genus Phrynobatrachus . 144 Family Pyxicephalidae . 149 Genus Amietia . 149 6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II GQxm?, Aubria . 151 GQwm, Pyxicephalus . 152 Genus Tomopterna . 153 Family Dicroglossidae . 157 Genus Hoplobatrachus . 157 Family Ranidae . 158 GQxms Amnirana . 158 Family Rhaeophoridae . 162 Genus Chiromantis . 162 Reptilia . 162 Order Chelonii . 162 Family Pelomedusidae . 162 Genus Pelomedusa . 162 Genus Pelusios . 164 Family Testudinidae . 169 Genus Kinixys . 169 Genus Stigmochelys . 172 Family Trionyehidae . 173 Genus Cycloderma . 173 Genus Trionyx . 173 Order Croeodylia . 174 Family Croeodylidae . 174 Genus Crocodylus . 174 Genus Mecistops . 176 Genus Osteolaemus . 176 Order Squamata . 177 (Lizards) . Ill Family Gekkonidae . Ill Genus Afroedura . 177 Genus Afrogecko . 178 Genus Chondrodactylus . 179 Genus Hemidactylus . 182 Genus Kolekanos . 188 Genus Lygodactylus . 188 Genus Pachydactylus . 192 Genus Rhoptropus . 199 Family Amphisbaenidae . 205 Genus Dalophia . 205 Genus Monopeltis . 207 Genus Zygaspis . 211 Family Lacertidae . 213 Genus Heliobolus . 213 Genus Holaspis . 214 Genus lehnotropis . 214 Genus Meroles . 218 Genus Nueras . 221 Genus Pedioplanis . 222 Family Cordylidae . 225 Genus Chamaesaura . 225 Genus Cordylus . 227 Family Gerrhosauridae . 229 Genus Cordylosaurus . 229 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 7 Genus Gerrhosaurus . 230 Genus Matobosaurus . 235 Genus Tetradactylus . 236 Family Seineidae . 237 Genus Acontias . 237 Genus Eumecia . 239 Genus Feylinia . 240 Genus Lepidothyris . 242 Genus Leptosiaphos . 243 Genus Lubuya . 244 Genus Melanoseps . 245 Genus Mochlus . 245 Genus Panaspis . 247 Genus Sepsina . 250 Genus Trachylepis . 252 Genus Typhlacontias . 272 Family Varanidae . 275 Genus Varanus . 275 Family Chamaeleonidae . 279 Genus Chamaeleo . 279 Genus Trioceros . 283 Family Agamidae . 284 Genus Acanthocercus . 284 Genus Agama . 287 Serpentes . 292 Family Typhlopidae . 292 Genus Afrotyphlops . 292 Genus Letheobia . 299 Family Leptotyphlopidae . 299 Genus Leptotyphlops . 299 Genus Namibiana . 301 Family Pythonidae . 304 Genus Python . 304 Family Boidae . 307 Genus Calabaria . 307 Family Viperidae . 308 Genus Athens . 308 Genus Bids . 309 Genus Causus . 315 Family Lamprophiidae . 320 Genus Amblyodipsas . 320 Genus Aparallactus . 321 Genus Atractaspis . 323 Genus Boaedon . 327 Germs Bothwphthalmus . 331 Genus Dromophis . 332 Genus Gonionotophis . 332 Genus Hemirhagerrhis . 333 Genus Hypoptophis . 334 Genus Limaformosa . 335 Genus Lycophidion . 338 Genus Mehelya . 340 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Polemon . 341 Genus Prosymna . 342 Genus Psammophis . 345 Genus Psammophylax . 354 Genus Pseudaspis . 357 Genus Pythonodipsas . 358 GQxms, Xenocalamus . 359 Family Elapidae . 361 GQxms Aspidelaps . 361 Genus Dendroaspis . 362 Genus Elapsoidea . 365 Genus Naja . 368 Genus Pseudohaje . 376 Family Colubridae . 376 Genus Chamaelycus . 376 Genus Crotaphopeltis . 377 Genus Dasypeltis . 379 Genus Dipsadoboa . 381 Genus Dispholidus . 381 Genus Grayia . 384 Genus Hapsidophrys . 387 Genus Hormonotus . 387 Genus Lycodonomorphus . 388 Genus Mopaneveldophis . 389 Genus Philothamnus . 389 Genus Rhamnophis . 398 Genus Scaphiophis . 399 Genus Telescopus . 400 Genus Thelotornis . 401 Genus Thrasops . 403 Genus Toxicodryas . 405 Family Natrieidae . 406 Genus Limnophis . 406 Genus Natriciteres . 408 Aeknowledgments . 410 Literature Cited . 412 Addendun . 456 Appendix . 457 Taxonomie Index . 481 MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 9 Abstract The present work eonstitutes an historieal atlas of all known bibliographie reeords of amphib¬ ians and reptiles of Angola. It is the first attempt to eompile in a single doeument all the reeords seattered through hundreds of publieations, published from the first half of the nineteenth eentury to the present day, and provide a eritieal taxonomie revision of the herpetofauna of the eountry. An introduetory text diseusses the Angolan soeio-politieal and physiographie landseape, the history of herpetologieal researeh in the eountry, and the diversity, distribution and endemism of Angolan her¬ petofauna. It also provides a summary of the eonservation eoneems surrounding the herpetologieal fauna. For eaeh taxon noted in the literature we provide a detailed taxonomie aeeount, ineluding data on the original deseription, its lUCN assessment status, global distribution, georefereneed reeords for eaeh known oeeurrenee in Angola (aeeompanied by a point loeality map), and taxo¬ nomie and distributional notes. A synoptie list of all doeumented Angolan amphibian and reptile speeies is provided as are lists of taxa previously erroneously assigned to the Angolan fauna. We reeognize 117 speeies of frogs and 278 speeies of reptiles as oeeurring in Angola, although many of these represent speeies eomplexes or are eurrently reeognized under names of eonvenienee pending taxonomie evaluation. In one instanee we provide a replaeement name, Trachylepis monardi nom. nov., to deal with an instanee of seeondary homonymy. The Atlas is intended to be a working referenee for both eurrent researeh and eonservation planning, as well as a eatalyst for future work. Resumo O presente trabalho eonstitui um atlas historieo referente a todos os registos bibliografieos de anfibios e repteis de Angola. Apresenta-se assim eomo um primeira tentativa de eompilar num unieo doeumento todos os registos dispersos por entre eentenas de publieagoes, desde a primeira metade do seeulo XIX ate ao presente, e provideneia uma revisao taxonomiea critiea da herpeto¬ fauna do pais. Este atlas eontem textos introdutorios sobre Angola, englobando a paisagem soeio- politiea e fisiografiea Angolana, a historia da investiga^ao herpetologiea no pais, a diversidade, distribuigao e endemismo da herpetofauna Angolana, bem eomo uma revisao dos problemas de conserva^ao que afeetam a fauna herpetologiea naeional. Para eada taxa euja oeorreneia esta eonfirmada, sao apresentadas fiehas taxonomieas detalhadas, passando em revista desde os dados referentes a deseri9ao original do taxa, a sua elassifiea9ao de aeordo eom a lUCN, a distribui9ao global, uma detalhada e georefereneiada lista de todos os registos para Angola - todos eles mapea- dos num mapa inteiramente dedieado, bem eomo notas taxonomieas e de distribui9ao. O apendiee eontem listas sinoptieas sobre todas as especies de anfibios e repteis que oeorrem em Angola, bem eom listas dos taxa erradamente referidos para a fauna Angolana. Reconheeemos a oeorreneia em Angola de 117 espeeies de anfibios e 278 de repteis, embora muitas destas representem eomplex- os de espeeies ou estao atualmente refereneiados atraves de nomes de eonvenieneia ate nova avali- a9ao taxonomiea. Num easo provideneiamos um nome de substitui9ao, Trachylepis monardi nom. nov., para resolver uma situa9ao de homonimia seeundaria. O presente Atlas tern eomo objeetivo ser uma refereneia de trabalho para aetuais pesquisas e pianos de eonserva9ao, bem eomo um eatalisador de futures trabalhos. Introduction Angola, with an area of nearly 1,250,000 km^, is one of the largest eountries in Afriea. Due to its geographie position, it is highly heterogeneous in terms of elimate, topography and biomes. Angola’s rieh landseape diversity and geographie position supports an amazingly diverse yet still poorly known biota. Its eomplex and troubled reeent history made the eountry almost inaeeessible 10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II to researchers for many years, effectively preventing modem surveys and studies, with negative implications for the advancement of biodiversity knowledge and conservation in all of sub-Saha¬ ran Africa. Fortunately, the end of the almost 30-year civil war in 2002 and the subsequent social and economic development of the country have created the appropriate climate for the return of sci¬ entific research, with support from both the national and international scientific communities. Biodiversity research and conservation initiatives have flourished in the country in the last decade, with Angola becoming a major player in the area, a fact reflected in Angola’s recent hosting of World Environment Day on 5 June 2016. Numerous studies on Angolan plants (Figueiredo et al. 2009; Romeiras et al. 2014; Gongalves and Goyder 2016), birds (Sinclair et al. 2004; Ryan et al. 2004; Sekercioglu and Riley 2005; Mills and Dean 2007; Mills 2010; Mills et al. 2011; Caceres et al. 2015), mammals (Pitra et al. 2006; Rodrigues et al. 2015; Baptista et al. 2013; Vaz-Pinto et al. 2015, 2016; Themudo et al. 2015), amphibians and reptiles (Fretey et al. 2011; Conradie et al. 2013, 2015, 2016; Ceriaco et al. 2014a,b, 2016a,b; Ernst et al. 2014, 2015; Branch and Conradie 2015) and insects (Serrano and Capela 2013, 2015) including the description of new endemic species (Haacke 2008; Conradie et al. 2012a,b; Carleton et al. 2015; Serrano et al. 2015; Stanley et al. 2016; Svensson et al. 2017) and a new endemic genus (Heinicke et al. 2014), and the publi¬ cation of species atlases and checklists (Pinto 1983; Crawford-Cabral 1987, 1998; Crawford- Cabral and Verissimo 2005; Mendes et al. 2013; Mills and Melo 2013; Kipping et al. 2017) are all clear evidence of the current scientific interest in Angolan biodiversity and its conservation. Angola boasts high herpetofaunal diversity, with 116 amphibian and 273 reptile taxa (278 if marine turtles are taken into account, see Taxonomic Accounts and Appendix Table Al), though this fauna has been neglected, especially when compared to the number of studies on mammals and birds. As an example, there are fewer than 10,000 extant museum specimens of amphibian or rep¬ tile from Angola, contrasting with nearly 40,000 museum herpetological specimens from neigh¬ boring and smaller Namibia, and >130,000 published records for reptiles alone from South Africa (Bates et al. 2014). Only one-third of Angolan amphibian and reptile species have been assessed by the International Union for Conservation of Nature (lUCN), and 29 of these are assessed as Data Deficient. In general, species distributions are poorly known, threats are unassessed, taxonomic problems are rife, and the species richness of Angola is believed to be underestimated. This situa¬ tion is exacerbated by the fact that one of the most important collections of Angolan amphibians and reptiles, containing dozens of type specimens critical to resolving taxonomic problems, was destroyed in the 1978 fire at the Museu Bocage in Lisbon. As part of an international partnership between INBAC, the Institute Nacional de Biodiversi- dade e Areas de Conservagao (Luanda, Angola), the California Academy of Sciences (San Fran¬ cisco, California, USA), the Florida Museum of Natural History at the University of Florida (Gainesville, Florida, USA), Villanova University (Villanova, Pennsylvania, USA), the University of Michigan at Dearborn (Dearborn, Michigan, USA), and the Museu Nacional de Historia Natu¬ ral e da Ciencia (former Museu Bocage, Lisbon, Portugal), new herpetofaunal field surveys and studies have been launched with funding from the United States National Science Foundation. These new surveys aim to re-survey historical type localities to collect topotypical material and provide new data (morphology, DNA, frog calls, etc.) for taxonomic studies currently impeded by the loss of many critical type specimens. Other research is planned to explore previously unsam¬ pled regions in order to better document and understand the diversity and distribution of the Angolan herpetofauna. This partnership has also resulted in other projects and activities related to the amphibians and reptiles of Angola. All museum records of Angolan specimens have been synthesized and georef- erenced through a project funded by the JRS Biodiversity Foundation, and all bibliographic records MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA II were eompiled and reviewed and are presented here. These data provide a starting point and overview of the current knowledge on the Angolan herpetofauna. The present work compiles all available published data regarding the diversity and distribution of the Angolan herpetofauna, span¬ ning more than 150 years of herpetological research in the country. It also highlights taxonomic and distributional issues relating to particular taxa, thus suggesting avenues for future research. We deliberately chose to focus solely on bibliographic records, not including the compiled museum records, as the latter will require further detailed reviews and descriptions that fall out of the scope of this publication. We anticipate that the current phase of active herpetological research by sever¬ al groups will facilitate the eventual production of a comprehensive atlas, along the lines of that recently completed for South Africa, Swaziland and Lesotho (Bates et al. 2014). However, this is not yet feasible and we offer the current work as a tool for future work in both taxonomy and con¬ servation. The Angolan Landscape Political and administrative borders oe Angola. — Mainland Angola is limited by the Atlantic Ocean on the west, by the Democratic Republic of the Congo in the north and northeast, by Zambia in the east, and by Namibia in the south (Fig. 1). The Cabinda enclave, a province of Angola, is separated from mainland Angola by the Democratic Republic of the Congo, which it borders to the south and east. It is bordered by the Republic of the Congo in the north and the Atlantic Ocean in the west. Once part of Portugal (also known as Portuguese West Africa), from which it became independent in 1975, the current borders of mainland Angola were originally defined in the Berlin West African Conference in 1884-1885 and in a few subsequent agreements Figure 1. Map of Angola, showing the provinces and major towns and position of Angola on the African continent. 12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II between European colonial powers. The Cabinda enclave, a Portuguese protectorate since 1885, also known as Portuguese Congo, was added to Angola after independence. Administratively, Angola is currently divided into 18 provinces — Cabinda, Zaire, Uige, Bengo, Luanda, Kwanza Norte, Kwanza Sul, Malanje, Lunda Norte, Lunda Sul, Mexico, Bie, Huambo, Benguela, Namibe, Huila, Cunene and Cuando Cubango — with Mexico being the largest (223,023 km^) and Luanda the smallest (18,826 km^). Angolan provincial delimitations have changed over time. For example, Bengo Province was created in 1980 from areas formerly constituting parts of Luanda and Kwan¬ za Norte provinces, and it has recently decreased in size owing to the expansion of the borders of Luanda. There are plans to divide Mexico and Cuando Cubango provinces into three and two provinces, respectively. Each province is divided into different municipalities (“Municipios”) that are subsequently divided into communes (“Comunas”). Physiography. — The Angolan coastal plain extends from sea level to approximately 250- 12^E 16°E 20°E 24^E 6°S lO^S 14^S 18^S Figure 2. The topography of Angola (after Sayre et al. 2013) MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 13 300 m elevation, or up to about 500 m in the northwest and southwest areas (Fig. 2). A transition zone rises sharply from 500 m to about 1000 m and is assoeiated with sheer eliffs and the Angolan escarpment in the southwest areas of the country (Fig. 2). The majority of the country lies on the Angolan Plateau, with a elevation between 1000-1500 meters (Fig. 2), with the exception of moun¬ tainous areas, mainly in Huambo, Bie and Huila provinces, that rise over 2000 m, reaching 2620 m at Mount Moco, the highest point of Angola. The escarpment in Angola, represented by Serra da Mocaba and Bie Escarpments, is approximately 1,000 km long and is one of the most isolated sec¬ tions of the Afromontane Archipelago (Clark et al. 2011). Due to its isolation, it harbors a rich plant diversity, particularly rich in endemics, as well as the highest number of vertebrate endemics in the subcontinent after South Africa (Clark et al. 2011). River basins. — Angola is extensively irrigated by eight main river drainages and a series of coastal rivers (Fig. 3). While the north, including the provinces of Cabinda, Zaire, Uige, Malanje, \2^E 16^E 20<^E 24^E 6°S lO^S 14^S 18^S Kilomcttn Coastal Rivers - Kwanza Cuito Cuvelai - Kavango Kunene Kwando Zambezi - Congo Figure 3. River basins of Angola (data from https://www.arcgis.com/). 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Luanda Norte, Lunda Sul and the northern parts of Moxico, is drained by the Casai-Congo Basin, the eentral areas of the Angola, and especially the east and southeast, are drained by the Zambezi, Cuando, Cuito, and Cubango rivers. The Cuvelai-Etosha Basin extends from northern Namibia to parts of Cunene and Huila provinces, whereas the Kunene Basin dominates most of Huila Province with the main river bordering Namibia along the southern edges of Namibe and Cunene provinces. Finally, the Kwanza, an exclusively Angolan river, starts in Bie and makes a turn to the west in Malanje Province, passing through Kwanza Norte, Kwanza Sul, Bengo and Luanda provinces where it meets the sea. A large part of the west coast of Angola, including the provinces of Zaire, Bengo, Luanda, Kwanza Sul, Benguela, Namibe and Huila, is drained by smaller, sometimes inter¬ mittent, coastal rivers. Geology. — Angolan geological history is complex, shaped by different events, with rocks from different geological eras scattered across the country, and a considerable diversity of soils (Schluter 2006). While a large part of the metamorphic rocks of the country derive from the Congo- Kasai Craton, there is a considerable diversity of more recent sedimentary and igneous rocks (Schluter 2006). Coastal areas of Angola are divided by three main basins: the Congo, the Kwan¬ za and the Namib marine coastal basins. These first developed during the Lower Cretaceous and are dominated by sedimentary rocks (both carbonate and non-carbonate) with some areas of sand¬ stone and limestone sands in the southwest (Namibe Province), as well as some alluvial sands, such as those at the mouth of the Congo River at the northwestern border of the country (Schluter 2006; Fig. 4). Sedimentary rocks dominate almost all of eastern Angola, which is largely covered by sands and related Aeolian sediments of the Kalahari Group (Schluter 2006). The main river basins and depressions contain alluvial and colluvial deposits in the form of sands, clays, rubbles and gravels, with the exception of the Cuvelai Basin, which contains saline soils (Schluter 2006; Fig. 4). The more central areas of the country are characterized by a mix of metasedimentary, magmat¬ ic, to metamorphic (metaigneous) rocks (Schluter 2006). The Bie Escarpment area is dominated by silica rich soils, originating from volcanic activity, while some intrusive volcanic rocks occur in Huila, Cunene, and Moxico (Schluter 2006; Fig. 4). A considerable area of the escarpment border is limited by metaigneous rocks, possibly from the old Congo-Kasai Craton, which are also pres¬ ent in some areas in the northeast of the country (Lunda Sul and Lunda Norte provinces; Schluter 2006; Fig. 4). Climate. — Most of Angola lies within the zone of intertropical trade winds and has a hot wet summer and a warm dry winter (Huntley 1974). The local climates within Angola reflect combi¬ nations of elevation, latitude, and distance to the coast. Generally, the coastal and southern areas of the country are hotter (although coastal areas in the Namib are cooler than those further north) whereas temperature tends to diminish with higher latitude and elevation, and especially at higher elevations in the central area of the country (Fig. 5). The variation of mean monthly temperatures is more extreme in the southeast and less pronounced in the north and northeast (Huntley 1974). Annual rainfall in Angola increases in a roughly southwest to northeast direction, with coastal areas south of Mo9amedes City, Namibe Province) being extremely dry and most central/northeast areas of the country, as well as Cabinda, having an annual rainfall of 1200-1400 mm (Fig. 6). Most of the southern and coastal areas have a less humid climate (Fig. 6). Rainfall is seasonal through¬ out Angola, with the northeast rainy season being from March to May and that of the southwest being shorter and occurring some time between December and March (Huntley 1974). The north¬ west experiences a short dry period (pequeno cacimbo) in January and February (Huntley 1974). Biomes and Vegetation Zones. — Due to its geographic placement, Angola supports a considerable diversity of biomes and vegetation types, from more tropical evergreen and semi- deciduous rainforests in the north to true desert in the extreme southwest. This enormous habitat MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 15 12^E \6^E 20^E 24^E 14°S 18^S Cnrboniit^ Silicic t ] Non -Carbon ft tc Metftigneous 1 1 Meiasedimeniary 1 1 Aeolian Sediments ■i Alkaline Intrusive Volcanic Alluvium - Fan Deposit Figure 4. The geology of Angola (after Sayre et al. 2013). Alluvium - Fluvial Alluvium - beach, strand, coastal dune Alluvium • Saline Water diversity translates into a high number of plant speeies, some of whieh are endemie. The latest eheeklist of Angolan flora published by Figueiredo et al. (2009) enumerates 7,296 plant taxa in 250 families and 1745 genera, of which nearly 15% are endemic, which gives Angola the second rich¬ est flora in Africa after South Africa. This habitat diversity, combined with the other physiograph¬ ic elements, has an extremely important effect on animal diversity and distribution, including those of amphibians and reptiles. 16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 12°E 16^E 20°E 24^E Figure 5. Annual mean temperatures of Angola (after Hijmans et al. 2005). Huntley (1974) recognized five main biomes in Angola. The Guinea-Forest Biome is an ever¬ green and semi-deciduous rainforest restricted to the interior of Cabinda (Maiombe Forest) and in discontinuous patches of forest in Zaire, Uige, Kwanza Norte, and Kwanza Sul provinces. The Congo Savanna Biome occurs in the north and is an area of tall grasslands interspersed by gallery forests in the valleys and isolated forests patches on the plateau, mostly occurring in the Congo River basin in Zaire, Uige, Malanje and Lunda Norte provinces. The Montane Forest Biome is rep¬ resented in Angola by a few isolated patches of forests on the protected slopes and deep ravines in the mountains of Huambo, Benguela, and Huila districts, at the highest elevations in the country, generally 2000-2500 meters, including Mount Moco. The Brachystegia Biome is dominated by Brachystegia (Miombo) woodlands, but is quite diverse across its 750,000 km^ extent on the cen¬ tral plateau of Angola. The South-West Arid Biome contains the most arid parts of Angola with extended dry seasons and mostly deciduous plants extending from coastal Luanda to Namibe MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 17 12^E 20^E 24''E Figure 6. Annual mean precipitation in Angola (after Hijmans et al. 2005). Province and then following the Angolan-Namibian border eastwards. Last, the Esearpment Zone is represented by a diseontinuous series of moist vegetation types that follow the Angolan Escarp¬ ment from the eoastal areas of Zaire, Bengo, Luanda, and Kwanza Sul Province to inland Huam- bo and Huila. Depending on the location in Angola, the Esearpment Zone shares affinities with neighboring biomes but may also act as a barrier between them. In eontrast to Huntley (1974), other reeent authors have taken a more fine-grained approaeh to characterizing Angolan habitats. This led Olson et al. (2001) to recognize 15 different ecoregions for Angola (Fig. 7): Atlantic Equatorial Coastal Forest in northeastern Cabinda Province (Fig. 8), Western Congolian Forest-Savanna Mosaie in Cabinda, Zaire, Uige and Malanje provinees (Fig. 9); Central African Mangroves in northeast Zaire Province, following the Zaire River mouth; Southern Congolian Savanna Mosaic in parts of Lunda Norte, Lunda Sul and Malanje; an exten¬ sive Angolan Scarp Savanna and Woodland area, forming a continuous distribution from the 18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II coastal areas of Zaire, Bengo, Luanda, and Kwanza Sul provinces to inland Huambo and Huila (Fig. 10); an Angolan Montane Forest-Grassland mosaic in the higher areas of Kwanza Sul, Huam¬ bo, Bie and Huila (also known as the Bie Escarpment as noted above, and supporting one of the most critically endangered and most biodiversity-rich areas in the narrow band known as “cloud forest” or “Angolan Escarpment Woodlands” (Clark et al. 2011); areas of Central Zambezian Miombo Woodlands (Fig. 11), Western Zambezian Grasslands, and Zambezian Cryptosepalum Dry Forest in the eastern parts of Mexico; an immense area of Angolan Miombo Woodland (Fig. 12), covering most of the central plateau, but extending to the coast in some areas of Kwanza Sul Province; most of the southern and southeast borders of the country are charactized by Zambezian Baikiaea Woodlands (Fig. 13) and some Zambezian Flooded Grasslands occur in the extreme I2°E 16'^E 20^E 24°E I I .\nnaliiiii Mipmbg Woaillantls C'cntnl Atrican An^Dlnn Mopanc Waadlainila C’cnlral Zambc/iiin Miamha 'n'ooation status (lUCN): Least Concern. Global distribution: The species is known from few localities in southern Cameroon, north¬ eastern Gabon and western Central African Republic. It was recently found in northwestern Ango¬ la and it has been suggested to be more widespread, extending into Equatorial Guinea, the Congo and the Democratic Republic of Congo. Ocurrences in .Angola (Map 1): Uige: “Serra do Pingano” [-7.67297, 14.93825] (Ernst et al. 2015:148). Taxonomic and distributional notes: The Angolan record (Ernst et al. 2015) pro¬ vides documentation of the southernmost range of this species. Wagner et al. (2013) and Ernst et al. (2015) included the species in the X. fraseri subgroup, which comprised several morphologically similar species that mainly occur in Central Africa. Ernst et al. (2015) clar¬ ified the taxonomic status of the Angolan pop¬ ulations and reviewed the distribution patterns oi Xenopus fraseri Boulenger, 1905 in Central Africa. Evans et al. (2015) regarded the mem¬ bers of the former X. fraseri subgroup, exclu¬ sive of X. fraseri itself (i.e., Xenopus amieti Kobel, Du Pasquicr, Fischberg and Gloor, Map I. DistribuHon of in Angola. 1980; X. andrei Loumont, 1983; X. boumbaensis Loumont, 1983; X. itombwensis Evans, Carter, Tobias, Kelley, Manner and Tinsley, 2008; X. lenduensis Evans, Greenbaum, Kusamba, Carter, Tobias, Mendel and Kelley, 201 1; A. longipes Loumont and Kobel, 1991; A. pygmaeus Loumont, 1986; A. ruwenzoriensis Tymov/ska and Fischberg, 1973; X. vestitus Laurent, 1972; and X. wittei Tinsley, Kobel and Fischberg, 1979) as members of the newly proposed Xenopus amieti group, to which they added two new species described in their publication — Xenopus allofraseri and Xeno¬ pus eysoole. 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Xenopus cf. epitropicalis Fischberg, Colombelli and Picard, 1982 Congolese Clawed Frog Xenopus epitropicalis Fischberg, Colombelli and Pieard 1982:53. Holotype: BMNH 1982.462 (eolleetors not stated, possibly M. Fischberg and J.J. Pieard). Type locality: “confluent de la Funa et de la Kemi, a 8 km au sud du centre de Kinshasa,” Democratic Republic of Congo. Xenopus calcaratus: Peters (1877a:618). Xenopus tropicalis: Laurent (1950a:13; 1954a; 70), Frade (1963:254), Cei (1977:16), Kobel (1981:120). Xenopus epitropicalis'. Loumont (1983:176), Ruas (1996:20), Channing (2001:239), Blaekbum et al. (2014), Evans et al. (2015:2), Frost (2016). Xenopus (Silurana) epitropicalis'. Fretey et al. (2011:22). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Democratic Republic of Congo and north¬ east along the Congo River to near the confluence of the Kwa River as well as from near Point Noire (Jackson and Blackburn 2010). It was was previously considered to be more widespread with populations extending across Central Africa, however many of those other populations are now treated as separate species. Ocurrences in Angola (Map 2): The species occurs in Cabinda enclave and in the northeast of the country. Cabinda: “Chincho- xo” [-5.10000, 12.10000] (Peters 1877a:618; Kobel 1981:120). Lunda Norte: “environs de Dundo galerie forestiere de la Luachimo (Dundo, Luachimo forest gallery)” [-7.38333, 20.83333] (Laurent 1950a: 13, 1954a:70; Kobel 1981:120; Ruas 1996:20). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: Publications previous to 1982 treated this species as Xenopus tropicalis (Gray, 1864) but subsequent studies have reveald that X. tropi¬ calis is restricted to tropical West Africa only from Senegal to Cameroon, and that records from the south and east Africa belong to Xenopus epitropicalis (Fischberg et al., 1982) (Loumont 1983; Ruas 1996; Evans et al. 2015). However, the distribution of both species requires further study and for now the Angolan population is best assigned to Xenopus cf epitropicalis (Ruas 1996; Channing 2001; Fretey et al. 2011; Frost 2016). Genetic data currently confirm the distribution of X. epitropicalis only for Congo and the Democratic Republic of Congo (Evans et al. 2015). It remains unclear whether populations from Angola represent nominotypical epitropicalis or one of the recently described species in subgenus Silurana, likely X. mellotropicalis. Map 2. Distribution of Xenopus epitropicalis in Angola. Xenopus sp. Xenopus fraseri: Laurent (1950a: 13, 1954a:70), Cei (1977:16), Kobel (1981:120), Ruas (1996:20), Channing (2001:240), Channing et al. (2012:294), Wagner et al. (2013:206), Ernst et al. (2015:147). Xenopus {Xenopus) fraseri'. Fretey et al. (2011:22). Global distribution: The taxon is presently known from northeastern Angola, although it like- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 61 ly represents a species that is more widespread and ranging from northern Angola to Cameroon and Central African Republic. Ocurrences in Angola (Map 3): Occurs in the extreme northeast of the country. Lunda Norte: “Muita, Luembe E (Muita, Luachimo)” [-7.80000, 21.45000] (Laurent 1950a:13; Ruas 1996:20); “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1954a:70; Ruas 1996:20); “Lunda Norte (probably Dundo or Muita)” (Kobel 1981 : 120). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomy and natural history notes: The Xenopus amieti subgroup {sensu Evans et al., 2015) contains several morphologically similar species that mainly occur in Central Africa (Wagner et al. 2013; Ernst et al. 2015). Previously this group contained X. fraseri, but Evans et al. (2015) demonstrated that Central African species referred to this taxon instead Map 3. Distribution oi Xenopus sp. in Angola. represent either X. allofraseri or X. parafraseri. The identity of the Angolan populations of Xenopus sp. remains uncertain, but they most likely represent a species within the amieti subgroup. Only two species, X. pygmaeus and X. fraseri, have previously been reported from south of the Congo Basin (Laurent 1950a, 1954a; Ruas 1996; Wagner et al. 2013). The species X. fraseri has frequently been confused withX andrei, a closely related species previously believed to be restrict¬ ed to Cameroon, Gabon, and the Central African Republic, but which was recently found in Ango¬ la (Ernst et al. 2015). Laurent (1950, 1954a) reported many specimens (~133) from Angola, collected in Lunda Norte Province, in “Dundo” and “Muita” regions in northeastern Angola, an area bordering the Democratic Republic of Congo. Ruas (1996) referred these previously published records from the northeastern Angola to X. fraseri although she also provided a complementary map (Ruas 1996) with other records for X. fraseri in central and southern Angola, with no further detail, on the basis of these records. It is likely that all previous records of X. fraseri in Angola actually refer to other species, including X. andrei or recently described species (Evans et al. 2015). Because of the lack of corresponding vouchers and tissue samples for genetic analyses, it cannot at this time be deter¬ mined to which species these records might be best assigned. Xenopus muelleri (Peters, 1844) Muller’s Clawed Frog Dactylethra Muelleri Peters 1844:37. Syntypes: ZMB 3556 [3 tadpoles], 3557 [3 speeimens], 6164 [2 speei- mens] and MZUT An264 fide Bauer et al. (1995:48) (eolleetor W.C.H. Peters). Type loeality: “Mozam¬ bique” (Peters 1844:37), restrieted to “Tete, Zambezi River” [Mozambique] by Loveridege (1953:308). Xenopus muelleri'. Conradie et al. (2016:17). Global conservation status (lUCN): Least Concern. Global distribution: The species is known to occur along the East African coastal belt from extreme southern Kenya through Tanzania to the northwestern border of South Africa. There is an apparently allopatric population from Burkina Faso eastward across Sudan-Guinea to northeastern Democratic Republic of Congo and Angola. 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 4): Occurs in the eastern regions of Angola. Cuando Cubango: “Cubango basin (29)” [-17.8729, 19.83333] (Conradie et al. 2016:17); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie etal. 2016:9-10,17). Taxonomic and distributional notes: Given the known distribution for X. muelleri and the fresh material collected by Conradie et al. (2016), it is expected that the species is more widespread in the eastern regions of Angola, and probably occurs in sympatry with X. poweri. Map 4. Distribution of Xenopus muelleri in Angola. Xenopus petersii Bocage, 1895 Peters’ Clawed Frog Xenopus Petersii Bocage 1895a: 187. Syntypes: MBL 3.476 (S. Salvador do Congo, collector Antdnio Barro- so), 3.473 (Quimbundo), 3.474-477 (Caconda), 3.478 (Dombe), 3.479 (Quibula), 3.481-483 (Huila), 3.485 (Cassange), 3.486-488 (Dondo) (collector J.A. d’Anchieta) [15 specimens] T/t/e Perret (1976a: 16). All destroyed by fire 18 March 1978. Bauer et al. (1996:271) reported one syntype from “Catumbella” ZMB 5831, as a surviving specimen from exchanged from the MBL. Type locality: “S. Salvador do Congo,” “Dondo,” “Caconda,” “Dombe,” “Quibula,” “Huila,” “Cassange” and “Quindumbo” (Bocage 1895a: 187), [= M’Banza Congo, Dondo, Caconda, Dombe, Huila, Cassange and Quindumbo], Angola. Daetylethra laevis: Gunther (1865a:480). Daetylethra miilleri: Bocage (1867b:227). Daetylethra Multeri: Bocage (1879b:89, 1879c:96). Xenopus muelleri-. Boulenger (1882:457), Loveridge (1957:308). Xenopus petersii-. Bocage (1896a:113, 1897a:206), Ferreira (1906:166), Schmidt (1936:128), Bauer et al. (1996:271), Channing (2001:248), Measey and Channing (2003:325), Furman et al. (2015:910), Frost (2016), Ceriaco et al. (2016b:41). Xenopus laevis-. Boulenger (1905:107), Monard (1937a:25, 1938:55, 76), Hellmich (1957a:22), Inger (1959:540), Kobel (1981:120), Gavetti and Andreone (1993:41). Xenopus laevis poweri: Laurent (1964a: 129), Cei (1977:17). Xenopus laevis petersi: Loveridge (1957:308), Schmidt and Inger (1959:8), Laurent (1964a: 130), Perret (1976a: 17), Cei (1977:17), Poynton and Broadley (1985a: 508), Poynton and Haacke (1993:13), Ruas (1996:20, 2002:141). Xenopus {Xenopus) laevis: Fretey et al. (2011:22). Xenopus cf petersii: Ceriaco et al. (2014b:669). Xenopus petersii: Furman et al. (2015:910), Frost (2016), Conradie et al. (2016:17). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Gabon, southwards to the western Democratic Republic of Congo, Angola, and northern Namibia. Ocurrences in Angola (Map 5): The species likely is widely distributed across western and central Angola. Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a: 187, 1897a:206; Loveridge 1957:308; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20). Kwanza 12°E Ifi-E 20^ E 24“E MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 63 Norte: “Cambondo” [-9.15963, 14.65771] (Ferreira 1906:166; Monard 1938:55); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:166); “Ambaea” [-9.26667, 15.18333] (Bocage 1 895a: 1 87, I897a:206; Monard 1938:55; Loveridge 1957:308; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Dondo” [-9.68333, 14.43333] (Boeage I895a:I87, I897a:206; Monard 1938:55; Loveridge 1957:308; Perret 1976a: 17; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20). Malanje: “Duque de Braganga” [-9.13333, 16.06667] (Sehmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 20 14b: 669); “Reserva da Palan- ca Preta (margens do Rio Cuanza)” [- 11.11667, 17.46667] (Ruas 2002:141), Map 5. Distribution of in Angola. “Pungo Andongo” [-9.66667, 15.58333] (Boulenger 1905:107; Monard 1938:55; Ruas 1996:20); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:41). Kwanza Sul: “Cas- songue” [-11.85000, 15.05000] (Loumont 1984:728; Ruas 1996:20). Lunda Norte: “Cassange” [-9.58333, 17.86667] (Boeage 1895a:187, 1897a:206; Monard 1938:55; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20, 2002:141). Lunda Sul: “mare Tehifuka, Alto Cuilo, Lunda (= Chifu- ea)” [-10.00000, 19.58333] (Laurent 1964a:130; Loumont 1984:728; Ruas 1996:20, 2002:141); “Alto Chieapa, Lunda” [-10.88333, 19.23333] (Laurent 1964a:130; Ruas 1996:20, 2002:141); “sourees du Cuilo, Alto Chieapa, rives sans foret, Lunda (= Rio Cuilo)” [-10.86667, 19.40000] (Laurent 1964a:130; Loumont 1984:728; Ruas 1996:20, 2002:141); “Dala” [-11.03333, 20.20000] (Monard 1937a:25, 1938:76; Ruas 1996:20, 2002:141); “mare des rives du Kutele, affl. droit du Cuango, Alto Chieapa, Lunda (= Rio Cutele)” [-11.06667, 18.86667] (Laurent 1964a:130; Ruas 1996:20, 2002:141). Bie: “Gauea” [-11.18333, 17.45000] (Sehmidt 1936:128; Monard 1938:55; Sehmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Chitau” [-11.43333, 17.15000] (Sehmidt 1936:128; Monard 1938:55; Sehmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Nequilo” [-12.58500, 17.07000] (Furman et al. 2015 - see Table SI); “Cubango basin (10)” [-13.71616, 17.09661] (Conradie et al. 2016:8-9, 17). Benguela: “Quibula” [-12.28333, 14.68333] (Boeage 1895a:187; Monard 1938:55; Loveridge 1957:308; Perret 1976a:17; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Catumbella (= Catumbela)” [-12.43333, 13.55000] (Boeage 1895a: 187; Monard 1938:55; Loveridge 1957:308; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a: 187; Monard 1938:55; Loveridge 1957:308; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Benguella (= Benguela)” [-12.58333, 13.41667] (Boeage 1895a:187, 1897a:206; Boulenger 1882:457, 1905:107; Monard 1938:55; Loveridge 1957:308; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Dombe” [-12.95000, 13.10000] (Boeage 1879b:89; 1895a:187, 1897a:206; Monard 1938:55; Perret 1976a:17; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Caimbambo” [-13.01667, 14.01667] (Loumont 1894:728; Ruas 2002:141); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:22); “Catengue” [-13.03333, 13.73333] (Sehmidt and Inger 1959:8; Loumont 1984:728; Ruas 1996:20, 2002:141); “Cubal da Ganda (Mareo de Canavezes)” [-13.08333, 14.33333] (Laurent 1964a:129; Ruas 1996:20, 2002:141); 12'^E l(i*E 2()°£ 24*E 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Hanha” [-13.30000, 14.20000] (Bocage I896a:II3; Monard 1938:55; Ruas 1996:20, 2002:141). “Rio Coporolo (= Coporola)” [-13.93333, 12.96667] (Loumont 1894:728; Ruas 2002:141). Hufla: “Caeonda” [-13.73333, 15.06667] (Boeage I895a:I87; Monard 1938:55; Loveridge 1957:308; Ferret 1976a: 17; Loumont 1984:728; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Kalukem- be (= Caluquembe)” [-13.78333, 14.68333] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Kuvangu (= Cubango)” [-14.46667, 16.30000] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Huilla (=Huila)” [-15.05000, 13.55000] (Gunther I865a:480; Boeage I895a:I87, I897a:206; Monard 1938:55; Schmidt and Inger 1959:9; Ferret I976a:I7; Bauer et al. 1996:271; Ruas 1996:20, 2002:141); “Osi (= Osse)” [-15.08333, 15.41667] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Lagoa Nuntechite” [-15.13333, 13.41667] (Poynton and Haacke 1993:13); “Kampulu (= Campulu-Cambissa)” [-15.21667, I6.II667] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Kului (= Cului)” [-15.41667, 15.73333] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141). Namibe: “Mossamedes” [-15.20000, 12.15000] (Bocage I967b:227; Loumont 1984:728; Ruas 1996:20, 2002:141). Cuando Cubango: “Kandin- gu” [-14.66667, 17.70000] (Monard I937a:25, 1938:55, 76; Ruas 1996:20, 2002:141); “Cubango basin (3)” [-14.94277, 17.71863] (Conradie et al. 2016:8-9, 17); “Cubango basin (5)” [-14.74628, 17.66844] (Conradie et al. 2016:8-9, 17); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 17); “Cubango basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:8-9, 17); “Cuito basin (24)” [-14.60622, 18.46722] (Conradie et al. 2016:8-9, 17). Undetermined Locality: without precise locality (Bocage 1879b:95); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17); “arid territories along the coast” (Cei 1977:17). Taxonomic and distributional notes: See notes below for Xenopus poweri. Xenopus poweri Hewitt, 1927 Power’s Clawed Frog Xenopus poweri Hewit 1927a:413, pi. 24. fig. 3. Holotype: MMK/F/898 (collector J.H. Power). Type locali¬ ty: “Victoria Falls,” Zambia and Zimbabwe border. Xenopus laevis poweri'. Schmidt and Inger (1959:8), Laurent (1964a: 129). Xenopus laevis: Kobel (1981:120), Channing (2001:243). Xenopus laevis petersii: Ruas (1996:20, 2002:141). Xenopus {Xenopus) laevis: Fretey et al. (2011:22). Xenopus poweri: Furman et al. (2015:910), Frost (2016), Conradie et al. (2016:17). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from central and eastern Nigeria to eastern Cameroon and western Central Africa Republic, south to the Democratic Republic of Congo, east¬ ern and southeastern Angola, the Okavango Region of Namibia, Zambia, and western Zimbabwe to northern Botswana. Ocurrences in Angola (Map 6): The species occurs in central and western Angola, while X. poweri is more restricted to the east and southeastern regions. Moxico: “Reserva da Palanca Preta (Rio Calombe)” [-11.83333, 19.93333] (Ruas 1996:20; Ruas 2002:141); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:130; Ruas 1996:20, 2002:141); “Lucusse” [-12.51667, 20.81667] (Ruas 1996:20, 2002:141). Cuando Cubango: “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 17); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie etal. 2016:9-10, 17). Taxonomic and distributional notes: The African clawed frog Xenopus laevis (Daudin, 1903) has a large native distribution over much of sub-Saharan Africa, but despite its important role in biological research, its phylogeography and evolutionary history remains poorly known. The MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 65 historic data for Xenopus laevis in Angola begins with Gunther (1865a), who eited the speeies for “Huila,” followed by several authors ineluding Boulenger (1905), Monard (1937a, 1938), and Hellmieh (1957a). After Gunther’s eitation, Boeage (1895a) deseribed a new speeies, Xenopus petersii, based on numerous speeimens deposited in Museu Boeage, Lisboa (see above), all eonsidered syntypes by Perret (1976a). Unfortunately, the syntypes were destroyed in the fire in the Museu Boeage in 1978. Bauer et al. (1996) reported a surviving syntype from “Catumbel- la” deposited in Museum fur Naturkunde, Berlin. Parker (1936) deseribed four raees or groups of X. laevis, redueing X. petersii to a subspeeies of X. laevis and suggesting that Map 6. Distribution of Xenopus poweri in Angola. Xenopus poweri Hewitt, 1927 is a synonym of X. petersii (Loveridge 1957). When deseribing X. petersii, Boeage (1895a) listed three “varieties” based on differenees in ventral markings. He gave no indieation that this variation was geographieal, but Sehmidt and Inger (1959) reported that 12 FMNH speeimens from northern Angola represent Boeage ’s “var. A” pattern, whereas five southern Angolan speeimens exhibit the “var. B” pattern. They refered X. laevis petersii to “var. A,” whieh represents the northern form distributed from the Upper Cuanza River northward. They assigned Xenopus laevis poweri to “var. B” reaehing from the Lower Cuanza River aeross all of southern Angola to Zambia (Sehmidt and Inger 1959; Poynton and Broadley 1985a). Mertens (1971), however, rejeeted this view. Channing (2001) eonsidered X. petersii as a full speeies, removing it from synonymy with Xenopus laevis. Measey and Channing (2003) provided molecu¬ lar evidence for the distinetiveness of X. petersii from X. laevis and eonsidered X. poweri to be a synonym of X. petersii, although other authors, ineluding Fretey et al. (2011) nonetheless later ineluded X. petersii and X. poweri as synonyms of Xenopus laevis. Reeently, Furman et al. (2015) provided an analysis of moleeular variation in the X. laevis group, whieh elarified the distribution of these lineages and supported the reeognition of X. laevis sensu stricto, X. petersii Boeage, 1895, and X. victorianus Ahl, 1924 and revalidated X. poweri Hewitt, 1927 as a separate speeies. These authors restrieted A laevis to South Afriea and the pop¬ ulation from Western Central Afriea to X. petersii, whieh indieates that the eentral and western Angolan population should be referred to the latter. They also proposed that portions of the eurrently reeognized distributions ofX laevis north of the Congo Basin andX. petersii south of the Congo Basin should be referred to X. poweri. This suggests that populations from eastern and southeastern Angola should be referred to X. poweri (Furman et al. 2015). Based on the available studies and in some new material reeently eolleeted by Conradie et al. (2016), we refer reeords from Moxieo Provinee to X. poweri, ineluding those from “Lueusse,” “Reserva da Palanea Preta,” “Rio Calombe” (Ruas 1996, 2002), and “Cazombo” (Laurent 1964a). 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family Bufonidae Gray, 1825 Genus Mertensophryne Tihen, 1960 Mertensophryne melanopleura (Schmidt and Inger, 1959) Dark-Sided Toad Bufo melanopleura Schmidt and Inger 1959:23, fig. 8, pi. 4, fig. 2. Holotype: IRSNB 1.265, formerly Inst. Parc. Nat. Congo Beige 807 (eollector G.F. de Witte). Type locality: “Kankunda, Parc National de I’U- pemba, Belgian Congo” (Schmidt and Inger 1959 Bufo melanopleura: Poynton and Broadley (1988:481), Ruas (1996:22, 2002:142), Chan- ning (2001:86). Mertensophryne melanopleura: Fretey et al. (2011:24), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the southern Democratic Republic of Congo, eastern Angola, and Zambia. Ocurrences in Angola (Map 7): There is a single record from eastern Angola. Moxico: “Lago Cameia” [-11.71667, 20.80000] (Ruas 1996:22, 2002:142; Channing 2001:86). Taxonomic and distributional notes: The species is presently known only from one published locality in Angola but presumably occurs more widely in woodlands and forests (Poynton and Broadley 1988; Ruas 1996). Mertensophryne aff. mocquardi (Angel, 1924) Bufo Moequardi Angel 1924:270. Syntypes: MNHP R. Jeannel). Type loeality: “mont Kinangop et des forets inferieures du mont Kenya” (Angel 1924:270), [= Kinangop and Kenya Plateau], Kenya. Bufo moequardi: Monard (1937a:28, 1938:56, 80), Cei (1977:17). Bufo moequardi: Channing and Howell (2006:86). Mertensophryne moequardi: Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from Kenya. Ocurrences in Angola (Map 8): There is one record of this species from “Mulondo” in Huila Province. Hufla: “Mulondo” [-15.63333, 15.20000] (Monard 1937a:28, 1938:56, 80). Undetermined Locality: “arid territories along the coasf’ (Cei 1977:17). Taxonomic and distributional notes: :23), Demoeratie Republic of Congo. 12*E l(i*E 2I)°E 24*E Map 7. Distribution of Mertensophryne melanopleura in Angola. Mocquard’s Toad 1924.49-55 [7 specimens] (collectors Ch. Alluaud and 12°E lO'E 20°E 24"E. Map 8. Distribution of Mertensophryne moequardi in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 67 Monard (1937a) identified an individual examined by de Witte from “Molundo” as Bufo mocquardi (Angel, 1924). de Witte considered this specimen close to Bufo taitanus Peters, 1878 but believed that it represented Bufo mocquardi. Monard considered the discovery of this species in Angola to be remarkable, since the type specimen is from Kinangop on Mount Kenya. Currently the species Mertensophryne mocquardi is only recognized from its type locality (Channing and Howell 2006; Frost 2016). Further studies are needed to confirm the identity of Monard’s specimens. Genus Poyntonophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sa, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green and Wheeler, 2006 Poyntonophrynus dombensis (Bocage, 1895) Dombe Toad Bufo dombensis Bocage 1895b:51. Syntypes: BMNH 1947.2.21.3-4 (formerly 96.2.28.3-4), MBL T.4-366 and T.4-367 (eollector J.A. d’Anehieta), destroyed by fire 18 Mareh 1978. Type loeality: “Dombe, sur le littoral, au sud de Benguella” [= Dombe Grande, Benguela] Benguela Provinee, Angola. Bufo dombensis-. Boeage (1897a:206), Monard (1938:55), Inger (1959:540), Ferret (1976a: 16), Cei (1977:17), Frost (1985:43), Poynton and Haaeke (1993:12), Ruas (1996:22), Channing (2001:65). Poyntonophrynus dombensis'. du Preez and Carmthers (2009:162), Fretey et al. (2011:24), Channing et al. (2012:150), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola and northern Namibia. Ocurrences in Angola (Map 9): The species occurs in southwestern Angola. Benguela: “Dombe” [-12.95000, 13.10000] (Bocage 1895b:51; Monard 1938:55; Ferret 1976a: 16; Frost 1985:43, 2016; Ruas 1996:22). Namibe: “Assungao” [-14.86667, 13.10000] (Poynton and Haaeke 1993:12; Ruas 1996:22). Undetermined Locality: “arid territories along the coast” (Cei 1977:17). Taxonomic and distributional notes: Bocage (1895b) described Bufo dombensis (Bocage, 1895) based on eighteen specimens, the whereabouts of only some are now known. Poynton (1964 in Poynton and Broadley 1988) considered B. dombensis to be subspecies of Bufo vertebralis Smith 1842 (Frost 1985), an interpretation that was not followed in later works (Poynton and Broadley 1988). Ferret (1976a) studied the type specimens in Museu Map 9. Distribution of Poyntonophrynus dombensis in Angola. Bocage and found two of the syntypes, which were later destroyed in the fire in Lisbon in 1978. Poyntonophrynus grandisonae (Poynton and Haaeke, 1993) Grandison’s Toad (Endemic) Bufo grandisonae Poynton and Haaeke 1993:11. Holotype: TM 40150 (collector W. D. Haaeke). Type local¬ ity: “5 km E of Assuncao, Mossamedes District” (Poynton and Haaeke 1993:11), [= 5 km E of Assun^ao, Namibe] Namibe Province, Angola. Bufo grandisonae: Ruas (1996:22), Channing (2001:73). Poyntonophrynus grandisonae: Fretey et al. (2011:24), Frost (2016). 68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 10): The species is known only from Namibe Province from coastal regions on granite inselbergs (Poynton and Haacke 1993; Channing 2001; Frost 2016). Namibe: “5 km E of Assun^ao” [-14.86667, 13.10000] (Poynton and Haacke 1993:11; Ruas 1996:22); “Caraculo” [-15.01667, 12.66667] (Poynton and Haacke 1993:11; Ruas 1996:22); “Salona river, 2 km N of Cainde” [-15.48333, 13.36667] (Poynton and Haacke 1993:11; Ruas 1996:22); “20 km W of Virei” [-15.66667, 12.76667] (Poynton and Haacke 1993:11; Ruas 1996:22). Taxonomic and distributional notes: This species exhibits the features of the Poyntonophrynus “vertebralis group,” and while resembling P. dombensis (Bocage, 1895a), it differs in several morphological Mav \0. 'DistnhuMon of Poyntonophrynus grandisonae traits (Poynton and Haacke 1993; Channing m Angola. 2001). Poyntonophrynus kavangensis (Poynton and Broadley, 1988) Kavango Toad Bufo kavangensis Poynton and Broadley 1988:472, fig. 1. Holotype: NMZB 19074 (collector H.D. Jackson). Type locality: “Khwai River, Botswana.” Bufo kavangensis: Poynton and Haacke (1993:13), Ruas (1996:22, 2002:142), Channing (2001:80). Poyntonophrynus kavangensis: Fretey et al. (2011:24), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southwestern Angola to northern Namibia and adjacent Botswana and Zimbab¬ we. It is likely to occur in adjacent Zambia. Ocurrences in Angola (Map 11): The species occurs in southern Angola near the Namibian border, and there is one pub¬ lished record further north. Moxico: “Calom- be, Luso” [-11.83333, 19.93333] (Ruas 1996:22, 2002:142). Cunene: “23 km NW of Pereira de Ega - Rogadas” [-16.95000, 15.56667] (Poynton and Haacke 1993: 13; Ruas 1996:22). Taxonomic and distributional notes: When describing Bufo kavangensis, Poynton and Broadley (1988) mentioned that its distri¬ bution extends across northern Zimbabwe, irE I6®E 20°E :4'>E Map 1 1 . Distribution of Poyntonophrynus kavangensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 69 northern Botswana, northern Namibia, and neighboring southern Angola. The record by Ruas (1996) is from further north in “Calombe,” which calls into question the identification of the spec¬ imen. However, Ruas (1996) noted that this northern extension might relate to the historical distri¬ bution of the Kalahari sands. This species is known from temporary pools and flooded grasslands (Poynton and Broadley 1985a; Poynton and Haacke 1993; Channing 2001). Genus Schismaderma Smith, 1849 Schismaderma carens (Smith, 1848) African Red Toad Bufo carens Smith 1848: pi. 68. Syntypes: BMNH 65.5.11.124-126, 58.11.25.91-93 (collector A. Smith). Type locality: “Interior of Southern Afriea”^(ie Frost (1985:76). Bufo carens: Laurent (1964a: 131), Poynton (1964:60), Cei (1977:16), Poynton and Broadley (1988:84). Schismaderma earens: Frost (1895:76, 2016), Ruas (1996:22), Channing (2001:103), Fretey et al. (2011:24). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southwestern Kenya to eastern Angola, southeast through Zimbabwe to eastern South Africa and along Botswana bor¬ der. Ocurrences in Angola (Map 12): The species is only known from “Cazombo,” in Moxico Province. Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:131; Ruas 1996:22; Channing 2001:104). Undeter¬ mined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). Taxonomic and distributional notes: This monotypic genus was reviewed by Poyn¬ ton (1964) and Poynton and Broadley (1988). It is a widely distributed species found in savannas and common around human settle¬ ments (Channing 2001). 1I°E I6®E 2(}°E :4'*E Map 12. Distribution of Schismaderma carens in Angola. Genus Sclerophrys Tschudi, 1838 Sclerophrys buchneri (Peters, 1882) Buchner’s Toad Bufo Buehneri Peters 1882b: 147. Holotype: ZMB 10103 (eolleetor M. Buehner). Type loeality: “Lunda (Afriea oeeidentalis),” Angola. Bufo buehneri: Bauer et al. (1995:40). Bufofunereus: Tandy and Keith (1972:158), Ruas (1996:22, 2002:142). Amietophrynus buehneri: Fretey et al. (2011:22). Selerophrys buehneri: Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from eastern Angola, but it may also occur in Republic of the Congo and western Democratic Republic of Congo (Frost 2016). Ocurrences in Angola (Map 13): Within Angola, this species is known only from the type 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II locality “Lunda” in eastern Angola. Lunda Sul: “Lunda” [-10.96667, 20.06667] (Peters 18826:146; Bauer et al. 1995:40; Ruas 1996:22, 2002:142). Taxonomic and distributional notes: Various authors, including Tandy and Keith (1972), have suggested treating S. buchneri as a synonym of S.funerea (Bocage, 1866), which was followed by Ruas (1996, 2002). Both the lack of recent data and the uncertainty regard¬ ing the taxonomic status of this species indicate that more work is needed on this species. Boulenger (1904) recognized Bufo decorsei Mocquard 1903, described from Brazzaville in Republic of Congo, as a junior synonym of S. buchneri. If true, this suggests that this species is more broadly distributed in Central Africa. We follow Ohler and Dubois (2016) in using the senior synonym Sclerophrys Tschudi, 1838 for all African bufonids recently assigned to Amietophrynus Frost et al. 2006. Map 13. Distribution of Sclerophrys buchneri in Angola. Sclerophrys funerea (Bocage, 1866) Angolan Toad Bufo funereus Bocage 1866a:56 1866b:77. Holotype: MBL 7.5-363 (collector F. A. P. Bayao) fide Perret (1976a: 17), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” [= Calandula] Malan- je Province, Angola. Bufo benguelensis Boulenger (1882:299). Syntypes: BMNH 1872.2.15.3. Type locality: “Benguela” [Ango¬ la], “Fernando P6” [= Bioko, Guinea] and “West Afrika.” Bufo funereus-. Bocage (1882a:304, 1895a: 186, 1897a:205), Ferreira (1904:114, 1906:166), Monard (1937a:27, 1938:55, 79), Mertens (1937a: 18), Themido (1941:2), Laurent (1950a: 13, 1954a:71), Loveridge (1957:311), Schmidt and Inger (1959:20), Perret (1976a: 17), Cei (1977:16, 17), Frost (1985:45), Ruas (1996:21, 2002:142), Channing (2001:69). Bufo funereus funereus'. Laurent (1964a: 131). Amietophrynus funereus'. Fretey et al. (2011:23), Channing et al. (2012:129). Selerophrys funerea'. Frost (2016), Conradie et al. (2016:12). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from West-Central Africa from Gabon to Uganda, Burundi and Rwanda, southward to Democratic Republic of Congo and Angola. Ocurrences in Angola (Map 14): Reports for this species range across much of Angola, extending from the southwestern to northeast¬ ern regions. Bengo: “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:114; Monard Ma? lA.Ttistxfbution of Sclerophrys funerea 'm Angola. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 71 1938:55). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent I950a:I3; Ruas 1996:21); “Muita, Luembe E (Multa)” [-7.80000, 21.45000] (Laurent I950a:I3; Ruas 1996:21, 2002:142). Lunda Sul: “Alto Cuilo, rives du Cuilo, Lunda” [-10.01667, 19.55000] (Laurent 1964a:131; Ruas 1996:22, 2002:142); “Dala” [-11.03333, 20.20000] (Monard 1938:55, 79; Ruas 1996:21, 2002:142); “Alto Chicapa, Lunda” [-10.88333, 19.23333] (Laurent 1964a:131; Ruas 1996:22, 2002:142^ Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142); “Gumba” [-11.26667, 14.28333] (Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142); “Lembu (Lembe)” [-12.86667, 14.11667] (Ferreira 1904:114; Monard 1938:55; Ruas 1996:21, 2002:142). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:56, 1866b:77, 1882a:304, 1895a:186, 1897a:205; Loveridge 1957:311; Perret 1976a:17; Frost 1985:45, 2016; Ruas 1996:21, 2002:142). Mexico: “Calombe, Luso” [-11.83333, 19.93333] (Ruas 1996:22, 2002:142); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:131; Ruas 1996:22). Bie: “General Machado” [-12.03333, 17.46667] (Mertens 1937a:18). Huambo: “Serra do Moco (Luimbale)” [-12.50000, 15.16667] (Laurent 1954a:71; Ruas 1996:21, 2002:142). Benguela: “Benguella” [-12.58333, 13.41667] (Boulenger 1882:299; Loveridge 1957:311). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1882a:304, 1895a:186, 1897a:205; Monard 1938:55; Themido 1941:2; Perret 1976a:17; Ruas 1996:21, 2002:142); “Mukoti” [-14.20000, 15.80000] (Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142). Cunene: “Ruisseau Mbale (Rio Bale)” [-15.16667, 16.75000] (Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142). Cuando Cubango: “Kakindo (Caquindo)” [-15.45000, 17.05000] (Monard 1937a:27, 1938:55, 79; Ruas 1996:21, 2002:142); “Cubango system around the villages of Kakindo [= Caiundo] and Mbale” [-12.85000, 18.29028] (Conradie et al. 2016:12). Undetermined Locality: “without precise local¬ ity” (Ferreira 1906:116); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: The species was noted as new by Bocage in two dif¬ ferent papers (1866a,b), both of which appeared in the November 1866 issue of the same journal. Unlike many of the nomina in 1866a, however, that of Bufo funereus is a nomen nudum as there is no description. Boulenger (1882) described a new species, Bufo benguellensis, based on one indi¬ vidual from “Benguela” sent to him by Bocage as well as two other specimens from “Fernando Po” and “W. Africa”; Bocage (1895a) later synonymized B. benguellensis with B. funereus. The species Sclerophrys buchneri (Peters, 1882) is also sometimes treated as conspecific (Tandy and Keith 1972). Schmidt and Inger (1959) provided a map with the distribution of Bufo funereus in Central Africa, although they did not provide information about specific localities. Sclerophrys garmani (Meek, 1897) Carman’s Toad Bufo garmani Meek 1897:176. Syntypes: FMNH 415, MCZ 19082 [2 speeimens] (eolleetor Field Museum East Afriean Expedition). Type loeality: Haili [= Haileh], southeast of Berbera, Somalia. Bufo regularis humbensis Monard 1937a:26. Syntypes: MNHC 90.00098-99 [2 speeimens] (eolleetor A. Monard) and MNHB 23712. Type locality: “Mulondo” (Monard 1937a:26), Angola. Bufo regularis humbensis: Monard (1938:78), Barbour and Loveridge (1946:84), Frade (1963:254). Bufo garmani: Tandy and Keith (1972:142, 159), Frost (1985:46), Largen (2001:315). Bufo regularis sensu strieto: Ruas (1996:21). Bufo gutturalis: Ruas (2002:141). Amietophrynus garmani: Fretey et al. (2011:23). Selerophrys garmani: Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: Sclerophrys garmani is known from disjunct populations in East Africa 72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Monard (1937a) originally described a new subspecies ofBufo regularis Reuss, 1 833 based ‘5- Distribution of Sderophrys garmam in Angola. on specimens from “Molundo,” giving it the name Bufo regularis humbensis Monard, 1937. These specimens were deposited in the Musee d’histoire naturelle de La Chaux-de-Fonds, Switzerland, though Barbour and Loveridge (1946) also reported one cotype (MCZ A-23712) from “Molundo.” Ruas (1996) identified B. r humbensis as Bufo regularis sensu stricto, however she noted that some records identified as B. regularis may instead correspond to other Bufo (= Sclerophrys) species (following Tandy and Keith, 1972). Ruas (1996) recognized S. humbensis as a synonym of S. gar- mani rather than S. regularis. If S. garmani is restricted to northeastern Africa (Largen 2001), then Angolan specimens would either represent a relatively isolated population located near the Namib¬ ian border (Tandy and Keith 1972), correspond to another species such as S. poweri, or represent an undescribed species. Channing (2001) did not include A. garmani among the species of Ango¬ la, in contrast to both Fretey et al. (2011) and Frost (2016). from Ethiopia and northern Somalia, Angola, south to northeastern parts of South Africa, and the eastern Caprivi Strip of Namibia. Ocurrences in Angola (Map 15): The species occurs in southern and eastern Angola. Mexico: “Rio Calombe (Reserva da Palanca Negra)” [-11.83333, 19.93333] (Ruas 2002:141). As Sclerophrys garmani humbensis (plotted as triangles): Huila: “Mulondo” [-15.63333, 15.20000] (Monard 1937a:26, 1938:78; Barbour and Loveridge 1946:84; Ruas 1996:21). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937a:26, 1938:78; Ruas 1996:21); “Humbi (Humbe)” [-16.68333, 14.90000] (Monard 1937a:26, 1938:78; Ruas 1996:21). Undetermined Locality: “western subregion and Angolan highlands (Angola coastal or watersheds)” (Frade 1963:254). Taxonomic and distribntional notes: Sclerophrys gutturalis (Power, 1927) Guttural Toad Bufo spinosus Bocage 1867a:845. Holotype: MBL 324 (collector J.A. d’Anchieta), not located by Perret (1976a: 18), destroyed by fire 18 March 1978. Type locality: “Benguella,” later corrected to “Dombe, Benguella, Angola,” by Perret (1976a: 18). Preoccupied by Bufo spinosus Daudin, 1803. Bufo regularis gutturalis Power 1927:416, PI. 21, fig. 2. Syntypes: MCZ A-15403 and MMK [originally 4 specimens but now presumably 3] (collector J.H. Power). Type locality: “Lobatsi” and “Kumman” Botswana-Northern Cape Province border, South Africa. Bufo spinosus: Bocage (1867b:227), Loveridge (1936a: 82). Bufo regularis: Boulenger (1882:299), Bocage (1895a:185), Barbour (1911:135). Bufo regularis gutturalis: Power (1927:416). Bufo regularis regularis: Loveridge (1957:310). Bufo gutturalis: Tandy and Keith (1972:159), Perret (1976a:18), Poynton and Broadley (1988:452), Poynton and Haacke (1993:13), Ruas (1996:21, 2002:141), Channing (2001:74). Amietophrynus gutturalis: Fretey et al. (2011:23), Channing et al. (2012:130). Selerophrys gutturalis: Frost (2016), Conradie et al. (2016:12). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 73 Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southeastern Uganda eastward to coastal Kenya and Tanzania, south through Mozambique to KwaZulu-Natal. Also present in Swaziland, Botswana, northern Namibia, and Angola. Ocurrences in Angola (Map 16): The species is distributed across the middle of Angola, from the coast to the country’s eastern border with Zambia and the Democratic Republic of Congo. Kwanza Norte: “Dondo, bank of Quanza River” [-9.68333, 14.43333] (Poynton and Haacke 1993:13; Ruas 1996:21, Ruas 2002:142). Malanje: “Reserva da Palan- ca Preta (Rio Caluando)” [-11.46667, 17.70000] (Ruas 1996:21, 2002:141). Moxico: “Lago Dilolo” [-11.50000, 22.01667] (Ruas 1996:21, 2002:141); “Luso” [-11.78333, 19.91667] (Ruas 2002:141); “Reserva da Palanca Preta (Rio Calombe)” [-11.83333, 19.93333] (Ruas 1996:21, 2002:141); “Luvuei” [-13.06667, 21.16667] (Ruas 1996:21, 2002:141). Benguela: “Benguella (Benguela)” [-12.58333, 13.41667] (Bocage Distribution of Sderophrys gutturalis in 1867a:845; Loveridge 1936a:82, 1957:310; Ruas 1996:21, 2002:142); “Dombe” [-12.95000, 13.10000] (Bocage 1895a:185; Perret 1976a:18). Huila: “Sa da Bandeira” [-14.91667, 13.50000] (Ruas 1996:21, 2002:141); “Humpata” [-15.03333, 13.40000] (Poynton and Haacke 1993:13; Ruas 1996:21, 2002:142). Namibe: “Mos- samedes” [-15.20000, 12.15000] (Bocage 18676:277); “Cima/Saco do Giraul” [-15.06667, 12.15000] (Poynton and Haacke 1993:13; Ruas 1996:21; 2002:142); “Rio Coroca” [-15.78333, 12.06667] (Poynton and Haacke 1993:13; Ruas 1996:21, 2002:142). Cuando Cubango: “Cuban¬ go basin (2)” [-15.70452, 17.45897] (Conradie et al. 2017:8-9, 12); “Cubango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2017:9-10, 12); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2017:9-10, 12); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2017:9- 10, 12). “Cuito basin (32)” [-17.04880, 19.05333] (Conradie et al. 2017:9-10, 12); “Cuando basin (44)” (Conradie et al. 2017:12); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2017:9-10, 12). Taxonomic and distributional notes: The nomen Bufo spinosus Bocage, 1867 was preoccu¬ pied by the Iberian Spiny-toad (Daudin 1803) (Loveridge 1936). Prior to this recognition, it had been considered a synonym of Bufo regularis Reuss, 1833 by Boulenger (1882) and Bocage (1895a). Power (1927) described this taxon as the subspecies Bufo regularis gutturalis based on material from the border of Botswana and the Northen Cape. This taxon was later elevated to full species by Tandy and Keith (1972), and is currently uniformly accepted as a valid species (Chan- ning 2001; du Preez and Carruthers 2009; Frost 2016; Conradie et al. 2016). Sderophrys lemairii (Boulenger, 1901) Lemaire’s Toad Bufo lemairii-. Boulenger 1901:1, PI. 1, fig. 1. Holotype: not stated, but presumably originally in MRAC (eol- leetor C. Lemaire). Type loeality: “Pweto, sur le lae Moero” [= Pweto, Moero Lake], Demoeratie Repub- lie of Congo. 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Bufo lemairei: Laurent (1950a: 13, 1964a: 131), Cei (1977:16). Bufo lemairii: Poynton and Broadley (1988:464), Frost (1985:51), Haacke (1982a: 11), Ruas (1996:21), Chan- ning (2001:82). Amietophrynus lemairii'. du Preez and Carmthers (2009:144), Fretey et al. (2011:23). Sclerophrys lemairii'. Frost (2016), Conradie et al. (2016:12). gauehe du Cuango-Muque, Lunda (Rio Chim- boma)” [-10.76667, 19.20000] (Laurent 1964a:131; Ruas 1996:21); “Alto Chieapa, sources du Cuilo, patie sans foret riveraine, Lunda (Rio Cuilo)” [-10.88333, 19.23333] (Laurent 1964a: 131; Ruas 1996:21). Mexico: “Rive du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a:131; Ruas 1996:21); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:131; Haacke 1982a: 11; Ruas 1996:21). Cuando Cubango: “Cubango basin (7a)” [-14.42966, 17.82658] (Conradie et al. 2016:8-9, 12); “ Cubango basin (7b)” [-14.43377, 17.82957] (Conradie et al. 2016:8-9, 12); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9-10, 12); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2016:9-10, 12); “Cuito basin (31)” [-17.46777, 20.03333] (Conradie et al. 2016:9-10, 12); “Cuito basin (63)” [-17.93611, 21.10269] (Conradie et al. 2016:9, 12). Undetermined Locality: “areas of forest and savanna in north and northeast Ango¬ la” (Cei 1977:169). Map 17. Distribution of Sclerophrys lemairii in Angola. Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the Okavango Delta of northern Botswana, western Zambia and adjacent areas of eastern and northeastern Angola. It also occurs in the Caprivi Strip of Namibia. The Democratic Republic of Congo and Republic of Congo appear to be the northern limits of the species. Ocurrences in Angola (Map 17): The species occurs in northeastern Angola. Lunda Norte: “Muita, Luembe E (Muita)” [-7.80000, 21.45000] (Laurent 1950a:13; Ruas 1996:21). Lunda Sul: “Alto Chieapa, humidi- herbosa des sources du Tchimboma affl. Rive Taxonomic and distributional notes: None. Sclerophrys poweri (Hewitt, 1935) Power’s Toad Bufo regularis poweri Hewitt 1935:293. Syntypes: AMG? [5 specimens] (collector J.H. Power), probably lost, fide Conradie et al. (2015:66). Type locality: “Kimberley,” Northern Cape Province, Republic of South Africa, Conradie et al. (2015:66). Bufo garmani'. Tandy and Keith (1972:159). Bufo poweri'. Channing (2001:91), Pickersgill (2007a:521). Amietophrynus poweri'. Fretey et al. (2011:23), Channing et al. (2012:135), Channing et al. (2013a). Selerophrys poweri'. Frost (2016), Conradie et al. (2016:14). Global conservation status (lUCN): Least Concern. Global distribution: The species is found in arid southwestern Africa, from extreme southern MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 75 Angola, through northwestern southern Africa, including Namibia, Botswana and western Zim¬ babwe. Ocurrences in Angola (Map 18): The species occurs in the extreme southern Angola. Cuando Cubango: “Angola side of the Cuban¬ go River near Calai (29)” [-17.87291, 19.83333] (Conradie et al. 2016:9-10, 14). Taxonomic and distributional notes: Tandy and Keith (1972) considered this species to be a member of Bufo regularis group and a synonym of what is now recognized as Scle- rophrys garmani (Meek, 1897). Due to its sim¬ ilarities with S. garmani, Poynton and Broadley (1988) suggested that poweri might be conspecific with Bufo pseudogarmani Hulselmans, 1969, although they chose to retain it as a synonym of garmani. Pickergill (2007a) considered S. pseudogarmani a syn¬ onym of S. poweri and suggested that southern African populations should be referred to pow¬ eri. According to M. Largen, J. Poynton, L. Mazibuko and M. Tandy (pers. comm, in Channing et al. 2013a), the eastern limit of the distribution of S. poweri is uncertain due to remaining confusion with S. garmani. Whereas Channing (2001) did not consider S. garmani to occur in Angola, he did recognize populations from southernmost Angola as S. poweri. Recently, Conradie et al. (2016) cited a single specimen collected in Cuando Cubango Province, which represents the first record for Angola, although it is expected to be more widely distributed in southern and eastern Angola. Map 18. Distribution of Sclerophrys poweri in Angola. Sclerophrys pusilla (Mertens, 1937) Mertens’ Striped Toad Bufo cinereus Hallowell 1844:169. Name preoccupied by Bufo cinereus Schneider, 1799 = syn. Bufo bufo (Linnaeus, 1758). Bufo maculatus Hallowell 1854:101. Neotype: BMNH 1984.163 fide Poynton et al. (2016:83). Replacement name for Bufo cinereus Hallowell, 1844. Type locality: “Monrovia, Liberia” Poynton et al. (2016:83). Bufo regularis pusillus Mertens 1937:17. Holotype: SMF 22247 (collected by F. Haas). Type locality: “Leta- ba Camp,” Kruger National Park, South Africa fide Poynton et al. (2016:84). Bufo maculatus: Tandy and Keith (1972:159), Poynton and Broadley (1988:460), Channing (1989:1, 2001:84), Poynton and Haacke (1993:13), Pickersgill (2007a:541). Bufo funereus: Monard (1937a:26, 1938:55, 77, 78). Bufo pusilus: Cei (1977:17), Lambiris (1988:51). Bufo maculatus: Ruas (1996:21, 2002:141). Bufo funereus: Ruas (1996:21, 2002:142). Amietophrynus maculatus: Fretey et al. (2011:23), Sclerophrys maculata: Frost (2016), Ceriaco et al. (2016a: 19). Sclerophrys pusilla: Poynton et al. (2016:84), Conradie et al. (2016:14). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from eastern, central and southern Africa from southern and western Cameroon, eastern through the Central African Republic and Sudan south to Mozambique and northern regions of South Africa and Swaziland. The species also occurs in west¬ ern Zimbabwe, Zambia, and northern Namibia, with a possible wide range in Angola. 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 19): The species occurs in western Angola. Malanje: “Duque de Braganga falls (Duque de Bragan- 9a)” [-9.13333, 16.06667] (Poynton and Haac- ke 1993:13; Ruas 1996:21, 2002:141). Bie: “Caccuchi River” [-13.593333, 16.879861] (Poynton et al. 2016:89). Huambo: “Santo Amaro” [-12.70000, 15.85000] (Monard 1937a:26, 1938:77); “Huambo” [-12.737167, 15.81825] (Poynton et al. 2016:89); “Cubango basin (12a)” [-13.59333, 16.87986] (Conradie etal. 2016:14). Benguela: “31 kmNE of Sousa Lara-Chila (Bocoio)” [-12.46667, 14.13333] (Poynton and Haacke 1993:13; Ruas 1996:21); “Ebanga” [-12.73333, 14.73333] (Monard 1937a:26, 1938:78); “Monguavalo Farm” [-13.45000, 14.61667] (Poynton and Haacke 1993:13; Ruas 1996:21). Hufla: “Kalukembe (Caluquembe)” [-13.78333, 14.68333] (Monard 1938:55, 76; Ruas 1996:21, 2002:142); “Kuvan- gu” [-14.46667, 16.30000] (Monard 1937a:26, 1938:77). Namibe: “Leba pass, between river and highway” [-15.07033, 13.24381] (Ceriaco et al. 2016a:19); “Cainde” [-15.48333, 13.36667] (Poynton and Haacke 1993:13; Ruas 1996:21; Ceriaco et al. 2016a:19); “16 km W of Vila Nova” (Poynton and Haacke 1993:13; Ruas 1996:21; Ceriaco et al. 2016a:19). Cuando Cubango: “Cubango basin (3)” [-14.94277, 17.71863] (Conradie et al. 2016:14); “Cubango basin (6a)” [-14.67155, 17.73525] (Conradie et al. 2016:14); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:14); “Cubango basin (20)” [-14.67175, 17.15331] (Conradie et al. 2016:14); “Cubango basin (22d)” [-14.64991, 16.90739] (Conradie et al. 2016:14); “Cuito basin (24)” [-14.60622, 18.46722] (Conradie et al. 2016:14); “Cubango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:14); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:14); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:14); “Cuando basin (41)” (Conra¬ die et al. 2016:14); “Cubango basin (47)” [-14.70214, 17.37806] (Conradie et al. 2016:14); “Cuito basin (62)” [-17.50875, 20.06608] (Conradie et al. 2016:14). Taxonomic and distributional notes: Tandy and Keith (1972) placed Bufo pusillus {sensu Pienaar, 1963) in the synonymy of Sclerophrys maculata (Hallo well, 1854) due the similar mor¬ phology and mating calls. While Hulselmans (1969) questioned this synonymy, immunological evidence (Maxson 1981; Pickergill 2007a) suggested that it is not distinct from S. maculata. A recent study conducted by Poynton et al. (2016) established that S. maculata is now restricted to West Africa from the Gulf of Guinea eastwards to Cameroon, while S. pusilla is found in eastern and southern Africa, including Angola. Conradie et al. (2016) collected some new material from Cuando Cubango Province identified as S. pusilla. Monard (1937a, 1938) identified specimens from “Kalukembe” as Bufo funereus (Bocage) [= Sclerophrys funera (Bocage, 1866)] but we recently determined these to be S. maculata (Ceriaco et al. 2016b), which now should be refered to S. pusilla. 12°E Ifir'E 20'°:E I4'*E Map 19. Distribution of Sclerophrys pusilla in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 77 Sclerophrys regularis (Reuss, 1833) African Common Toad Bufo regularis Reuss 1833:60. Syntypes: SMF 3429, formerly 1298. le and SMF 1298. d [2 speeimens] (eol- leetor probably E. Ruppell), the speeimens represented by two figures of Grenouille Ponctuee from Saint- Hilaire 1809: p. 4, fig. 1, 2 are also syntypes. Type loeality: “Aegypten” [= Egypt]. Bufo guineensis: Peters (1877a:618), Boeage (1879b:89). Bufo pantherinus: Boeage (1866a: 56). Bufo regularis-. Boulenger (1882:298, 1905:107), Boeage (1887a:192, 1887b:208, 1895a:185, 1896a:113), Ferreira (1903:114), Monard (1937a:26, 1938:55, 77), Parker (1939:145), Themido (1941:2), Laurent (1954a:70), Inger (1959:540), Cei (1977:16, 17), Largen (2001:322). Bufo regularis regularis'. Sehmidt (1936:128), Loveridge (1936a:81), Monard (1938:78), Mertens (1937a:17, 1938a:429), Laurent (1950a: 13^ Hellmieh (1957a:23), Laurent (1964a: 130). Bufo regularis sensu lato'. Ruas (1996:21). Amietophrynus regularis'. Fretey et al. (2011:23), Channing et al. (2012:137), Ceriaco et al. (2014b:669), lUCN SSC Amphibian Speeialist Group (2016), Ceriaeo et al. (2016b:27). Selerophrys regularis: Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: This species has among the broadest distributions of all African amphib¬ ian species. It is known from northern Africa, western Eritrea and Ethiopia (also included Uganda and southern Kenya), southwards along the western coast from Cameroon to the Demoratic Repub¬ lic of Congo to Angola. Ocurrences in Angola (Map 20): The species is widespread in Angola, except in the southeast. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877:618). Zaire; “S. Salva¬ dor do Congo” [-6.26667, 14.23333] (Boeage 1887a;192; Ruas 1996:21). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 13, 1954a:70, 1964a; 130; Ruas 1996:21); “Muita, Luembe E (Muita)” [-7.80000, 21.45000] (Laurent 1950a:13, 1954a:70; Ruas 1996:21). Lunda Sul: “Poste de Luangue, humidiherbosa du ruisseau Katcheleke affl. W. du Luangue, entre le Lun- guena et la Tcha-Pemba, Lunda (Posto do Luangue)” [-9.33333, 19.23333] (Laurent 1964a: 130; Ruas 1996:21); “Riv. Cuvuemba Alto Cuilo, Lunda (Rio Cavemba)” [-10.01667, 19.55000] (Laurent 1964a:130; Map 20. Distribution of SctempA™ in Angola. Ruas 1996:21); “Lunda” [-10.96667, 20.06667] (Monard 1938:78); “Alto Chicapa, humidiherbosa des sources du Tchimboma affl. Rive gauche du Cuango-Muque, Lunda (Rio Chimboma)” [-10.76667, 19.20000] (Laurent 1964a:130; Ruas 1996:21); “Dala” [-11.03333, 20.20000] (Monard 1937a:26, 1938:77; Ruas 1996:21). Kwanza Norte: “N’dalla Tando” [-9.30000, 14.91667] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); “Cazengo” [-9.33333, 14.76667] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] (Parker 1936:145; Ruas 1996:21); “Novo Redondo” [-11.20000, 13.85000] (Ferrreira 1904:113; Monard 1938:55; Ruas 1996:21); “Fuma s/riv. N’Gunza, Novo Redondo (N’Gunza)” [-11.20000, 13.93333] (Laurent 1954a:70; Ruas 1996:21). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Boulenger 1882:298, 1905:107; Boeage 1866a:56; 1895a:185; 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ruas 1996:21); “Pungo-Andongo (Pungo Andongo)” [-9.66667, 15.58333] (Boeage 1895a:185; Boulenger 1905:107; Ruas 1996:21); “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 20146:669); “Marimba” [-8.36667, 17.03333] (Boulenger 1905:107; Ruas 1996:21) “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:27). Mexico: “Teixeira de Sousa, Distr. Villa Luso” [-10.70000, 22.23333] (Mertens 1937a: 17); “environs du lae Calundo, rives de la Lumeje, Moxieo (Rio Lumege)” [-11.51667, 20.76667] (Laurent 1964a: 130; Ruas 1996:21); “Rives du lae Calundo Moxieo” [-11.80000, 20.86667] (Laurent 1964a: 130; Ruas 1996:21); “Reserve de Chasse de Cameia, 120 km a Test de Luso (Cameia)” [-11.83333, 21.00000] (Laurent 1964a:130; Ruas 1996:21); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:130; Ruas 1996:21). Bie: “Bihe (Bie)” [-12.38333, 16.95000] (Boeage 1879b:78, 1895a:185; Ruas 1996:21); “Gauea” [-11.18333, 17.45000] (Sehmidt 1936:128; Monard 1938:55; Ruas 1996:21); “Chitau” [-11.43333, 17.15000] (Sehmidt 1936:128; Monard 1938:55; Ruas 1996:21); “General Maehado” [-12.03333, 17.46667] (Mertens 1937a:17). Huambo: “Bimbi (Bimbe)” [-11.81667, 15.83333] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Cuito, Mt. Moeo” [-12.45000, 15.26667] (Parker 1936:145; Ruas 1996:21); “Santo Amaro” [-12.70000, 15.85000] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Cuma” [-12.86667, 15.06667] (Loveridge 1936a:81). Benguela: “Benguella (Benguela)” [-12.58333, 13.41667] (Boeage 1895a:185; Ruas 1996:21); “Dombe” [-12.95000, 13.1000] (Boea¬ ge 1895a:185); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:23); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:426; Ruas 1996:21); “Cubal da Ganda, Mareo de Canavezes (Mareo de Canavezes)” [-13.08333, 14.33333] (Laurent 1964a:130; Ruas 1996:21); “Hanha” [-13.30000, 14.20000] (Boeage 1896a:113). HmTa: “Caeonda” [-13.73333, 15.06667] (Boeage 1895a:185); “Kalukembe (Caluquembe)” [-13.78333, 14.68333] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Sangeve (Sangueve)” [-13.88333, 15.83333] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Qui- lengues” [-14.10000, 14.08333] (Laurent 1954a:70; Ruas 1996:21); “Kuvangu (Cubango)” [-14.46667, 16.30000] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Kapelongo (Capelongo)” [-14.88333, 15.08333] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Osi (Osse)” [-15.08333, 15.41667] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Kangela (Cangela)” [-15.41667, 15.73333] (Monard 1937a:26, 1938:77; Ruas 1996:21). Nambie: “Mossamedes” [-15.20000, 12.15000] (Boeage 1895a:185; Ruas 1996:21). Cunene: “Kuvelai (Cuvelai)” [-15.65000, 15.80000] (Monard 1937a:26, 1938:77; Ruas 1996:21); “Mupa” [-16.18333, 15.75000] (Monard 1937a:26, 1938:77; Ruas 1996:21). Cuando Cubango: “Kakindo (Caquindo)” [-15.45000, 17.05000] (Monard 1937a:26, 1938:77; Ruas 1996:21). Undetermined Locality: “from almost all localities that he [J. d’ Anchieta] visited” (Boeage 1895a: 185); “Locomi” (Boulenger 1905:107; Ruas 1996:21); without precise locality (Laurent 1954:70); “Carangigo” (Boulenger 1882:298); “Between Benguela and Bihe” (Boulenger 1905:107). “areas of forest and savanna in the north and northeast of Angola” (Cei 1977: 16); “arid territories along the coast” (Cei 1977:17). Taxonomic and distributional notes: Sclerophrys regularis (Reuss, 1833) remains a taxo¬ nomic challenge. While many specimens from Angola have been identified as S. regularis, Ruas (1996) noted that the Angolan records are not representative of Bufo regularis sensu stricto and may represent Bufo maculatus (Hallowell, 1854) (currently Sclerophrys pusilla (Mertens, 1937), see S. pusilla account) and/or Bufo guturallis (Power, 1927). Sclerophrys regularis is a widely distributed in Africa ((lUCN SSC Amphibian Specialist Group 2016), but the boundaries between S. regularis and S. gutturalis in Angola, the Democratic Republic of Congo, Kenya, Uganda, and Tanzania remain poorly understood. Further studies are needed to clarify the distribution and the known range should be regarded as provisional. Ruas (1996) cited several localities (e.g., “Huila,” “Bibala,” “Quibula,” “Quindumbo,” “Cahata,” “Dondo,” “Ambaca,” “Catumbela,” “Galanga,” MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 79 “Humbe,” “Rio Quando,” “Gambos,” “Capagombe,” and “Massabi”) referring to Bocage (1895a) as a souree, although these localities are not cited in Bocage ’s work. Family Microhylidae Gunther, 1858 (1843) Genus Phrynomantis Peters, 1867 Phrynomantis affinis Boulenger, 1901 Spotted Rubber Frog Phrynomantis affinis Boulenger 1901:6, pi. 2, fig. 5. Holotype: MRAC (eolleetor C. Lemaire). Type loeality: “Pweto, Lake Mweru” (Boulenger 1901:6), Demoeratie Republie of Congo. Phrynomerus affinis: Laurent (1964a: 156), Cei (1977:17), Poynton and Broadley (1985a:514), Ruas (1996:22). Phrynomantis affinis: Channing (2001:231), du Preez and Carruthers (2009:284), Fretey et al. (2011:37), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Tanzania and the southern Demo¬ cratic Republic of Congo (former Katanga Province), south to eastern Angola, adjacent Zambia and northern Namiba. Ocurrences in Angola (Map 21): The species occurs in eastern Angola. Moxico: “Rives du lac Calundo, Moxico (Lago Calun- do)” [-11.80000, 20.86667] (Laurent 1964a: 156; Ruas 1996:22). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). Taxonomic and distributional notes: While Boulenger (1901) separated P. affinis from P. bifasciatus based on its smaller eyes and less dilated digits, Poynton and Broadley (1985a) suggested that these were not useful characters for diagnosing these two taxa. Nonetheless, more recent authors, including Channing (2001) and du Preez and Carruthers (2009) have continued to recognize them as distinct. irE I6*E 20°E 24*E Phrynomantis annectens Werner, 1910 Marbled Rubber Frog Phrynomantis annectens Werner 1910:294. Holotype: ZMB 24829 (eolleetor L. Sehultze). “Aar-Rivier” (Werner 1910:294) Aar or Aare River, Namibia. Phrynomerus annectens: Parker (1936:143), Inger (1959:542), Cei (1977:17), Poynton and Haaeke (1993:13), Ruas (1996:22), Channing (2001:232). Phrynomantis annectens: Frost (1985:392, 2016), du Preez and Carmthers (2009:286), Fretey et al. (2011:37), Channing et al. (2012:251), Ceriaeo et al. (2016a:20). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs in arid regions extending from the Namib Desert in Angola, through the rocky mountains of central Namibia, and into the Augrabies area of western Northern Cape Province in South Africa. 80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 22): The species occurs in southwestern Angola. Kwan¬ za Sul: “8 km NE of Novo Redondo, Gabela (Novo Redondo)” [-10.85000, 14.36667] (Poynton and Haacke 1993:13; Ruas 1996:22). Benguela: “Morro de Pondo” [-12.38333, 13.88333] (Parker 1936:143; Ruas 1996:22). Namibe: “Mutiambo River (Mutiambo)” [-14.93333, 12.46667] (Poynton and Haacke 1993:13; Ruas 1996:22; Ceriaco et al. 2016a:53); “Caraculo” [-15.01667, 12.66667] (Poynton and Haacke 1993:13; Ruas 1996:22; Ceriaco et al. 2016a:53); “Omauha lodge” [-16.19872, 12.40008] (Ceriaco et al. 2016a:20). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). Taxonomic and distributional notes: None. irE l(J®E 20°E :4'>E Map 22. Distribution of Phrynomantis annectens in Angola. Phrynomantis bifasciatus (Smith, 1847) Banded Rubber Frog Brachymerus bifasciatus Smith 1847b: pi. 63 and two pages of accompanying unnumbered text. Syntypes: BMNH (4 specimens) fide Poynton and Broadley (1985a: 5 13) (collector A. Smith). Type locality: “coun¬ try to the east and north-east of the Cape Colony,” South Africa. Phrynomantis bifasciata: Boulenger (1882:173), Bocage (1895a: 181), Monard (1937a:30, 1938:56, 83), Frade (1963:254). Phrynomantis bifasciatus: Ferreira (1904:113); Barbour (1911:135), Channing (2001:234), du Preez and Carmthers (2009:288), Fretey et al. (2011:37), Channing et al. (2012:254), Frost (2016). Phrynomerus bifasciatus: Schmidt and Inger (1959:187), Phrynomerus bifasciatus bifasciatus: Loveridge (1957:355), Poynton and Broadley (1985a:513), Ruas (1996:22). Global conservation status (lUCN): Least Concern. Global distribution: The species has a broad distribution, extending from Somalia to the Democratic Republic of Congo, south to South Africa. Ocurrences in Angola (Map 23): The species occurs in southwestern Angola. Kwan¬ za Sul: “Chingo (Novo Redondo)” [-11.20000, 13.85000] (Ferreira 1904:113; Monard 1938:56; Ruas 1996:22). Benguela: “Quissan- ge” [-12.43333, 14.05000] (Bocage 1895a:181; Ruas 1996:22); “Benguella (Benguela)” [-12.58333, 13.41667] (Bocage 1895a:181; Monard 1938:56; Ruas 1996:22). Huila: “Mol- Inger (1959:542), Cei (1977:17, 18). 12°E I6“E 20°E :4»E Map 23. Distribution of Phrynomantis bifasciatus in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 81 undo” [-15.63333, 15.20000] (Monard I937a:30, 1938:56, 83; Ruas 1996:22). Undetermined Locality: without precise locality (Bocage 1 895a: 181); “Western subregion and Angolan highlands (Angolan coastal or watersheds)” (Frade 1963:254); “plateaus regions” (Cei 1977:17); “arid terri¬ tories along the coast” (Cei 1977:18). Taxonomic and distributional notes: Poynton and Broadley (1985a) regarded this species as conspecific with Phrynomantis microps Peters, 1875, which is known from the savannas of West and Central Africa, and is now accepted as a valid species. Ruas (1996) suggested that Phryno¬ mantis bifasciatus (Smith, 1 847) is likely widely distributed in Angola. Family Brevicipitidae Bonaparte, 1850 Genus Breviceps Merrem, 1820 Breviceps sp. Breviceps gibosus Bocage (1870:68). Breviceps gibbosus Bocage (1873b:227). Breviceps mossambicus: Bocage (1895a:182), Parker (1934:194), Monard (1937a:29, 1938:56, 81), Loveridge (1957:357), Inger (1959:532), Laurent (1964a: 156), Cei (1977:17, 18), Gavetti and Andreone (1993:114), Ruas (1996:22). Rana mossambicus: Hellmich (1957a:30). Breviceps mossambicus/adspersus: Poynton (1982:67, 1992:68), Frost (1985:356, 2016), Channing (2001:213), du Preez and Carmthers (2009:108), Fretey et al. (2011:27). Breviceps adspersus: Poynton and Broadley (1985a:523), Conradie et al. (2016:11). Breviceps mossambicus-adspersus: Ruas (2002:142). Global distribution: What appears to be a single taxon, though of uncertain identity, ranges across west-central Angola. This may belong to a more widely distributed species in southern Africa, although phylogenetic support for this is not yet available. Ocurrences in Angola (Map 24): Records are reported from across western and eastern Angola. Lunda Sul: “Alto Chicapa, Lunda” [-10.88333, 19.23333] (Laurent 1964a: 156; Ruas 1996:22, 2002:142); “Riv. Luhemba, Alto Chicapa (Rio Luemba)” (Lau¬ rent 1964a: 156; Ruas 1996:22, 2002:142). Moxico: “Rives du lac Calundo (Lago Calun- do)” [-11.80000, 20.86667] (Laurent 1964a:156; Ruas 1996:22, 2002:142); “Calom- be, Luso” [-11.83333, 16.25000] (Ruas 1996:22; 2002:142); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:156; Ruas 1996:22, 2002:142). Bengo: “R. Donda, 300 miles inland from St. Paul de Loanda (Rio Donda)” [-9.68333, 14.43333] (Parker 1934:195; Ruas 1996:22, 2002:142). Huambo: “Bimbi (Bimbe)” [-11.81667, 15.83333] (Monard Map 24. Distribution of Sr., sp. in Angola. 1937a:29, 1938:81; Monard 1938:56; Ruas 1996:22, 2002:142); “Galanga” [-12.06667, 15.15000] (Monard 1938:56; Ruas 1996:22, 2002:142). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1895a: 182; Parker 1934:195; Monard 1938:56; Ruas 1996:22, 2002:142); “Quindumbo” 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-12.46667, 14.93333] (Bocage 1895a:182; Monard 1938:56; Ruas 1996:22, 2002:142); “Bengue- la” [-12.58333, 13.41667] (Parker 1934:195; Gavetti and Andreone 1993:114; Ruas 1996:22, 2002:142); “Ebanga” [-12.73333, 14.73333] (Monard 1937a:29, 1938:56, 81; Ruas 1996:22, 2002:142); “Chimbassi” [-13.18526, 14.20061] (Hellmich 1957a:30). Hulla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:182; Monard 1938:56; Ruas 1996:22, 2002:142); “Vila-da- Ponte (Kuvangu)” [-14.46667, 16.30000] (Monard 1937a:29, 1938:56, 81; Ruas 1996:22, 2002:142). Namibe: “Biballa (Bibala)” [-14.76667, 13.36667] (Bocage 1895a:182; Ruas 1996:22); “Chiyaka District” [vie. -15.08333, 12.73333] (Parker 1934:195); “Mossamedes” [-15.20000, 12.15000] (Bocage 1873b:227). Undetermined Locality: without precise locality (Bocage 1870:68); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “arid territories along the coasf’ (Cei 1977:18). Taxonomic and distributional notes: Identification of Angolan populations of this Breviceps species remains unclear. It may be conspecific with Breviceps adspersus Peters, 1882 or B. mossambicus Peters, 1 854. Most authors have considered the Angolan species to be the former. There is a high degree of similarity between the species, and they are easily confused. According to Poynton and Broadley (1985a) and Poynton (1992) this situation appears to be an example of “widespread sympatric hybridization” characterized by Woodruff (1973). If true, it may be difficult to be confident in the identification of preserved specimens, including those with morphological characteristics that are intermediate between the parental species. These authors suggested that Angolan populations referred to B. mossambicus are probably an undescribed species requiring further investigation. We consider the Angolan records of Breviceps gibbosus (Linnaeus, 1758) cited by Bocage (1870; 1873b) to refer to this species. Breviceps poweri Parker, 1934 Power’s Rain Frog Breviceps poweri Parker 1934:195. Holotype: BMNH 1947.2.14.89 (collector C. Pitman). Type locality: “Broken Hill (i.e., Kabwe), N. Rhodesia” (Parker 1934:195) [= Kabwe, Zambia] fide Poynton and Broadley (1985a:525) and Frost (2016). Breviceps poweri: Channing (2001:223), du Preez and Carmthers (2009:124), Fretey et al. (2011:27), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Democratic Republic of Congo (former Katanga Province) and adjacent northeastern Angola, east through Zambia and the southern Tanzania south to Mozambique. Ocurrences in Angola: The species occurs in eastern Angola (see notes below). Taxonomic and distributional notes: Channing (2001) provided a map with Angolan records from Lunda Sul and Moxico Province, near the Zambian border, though the source of these records remains unclear. Fretey et al. (2011) and Frost (2016) cited B. poweri from northeastern regions of the country, though without specific locality data, probably on the basis of Channing (2001). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 83 Family Hemisotidae Cope, 1867 Genus Hemisus Wagler, 1827 Hemisus guineensis Cope, 1865 Guinea Snout-Burrower Hemisus guineensis Cope 1865: 100. Holotype: NHMW 1095 (collector unknown) fide Haupl and Tiedemann (1978:18). Type locality: Not stated, later restricted to “Guinea,” by inference according to Haupl and Tiedemann (1978:18). Hemisus guineensis mierops Laurent 1972:55, fig. 27. Holotype: MRAC 851 (collector H. Schouteden). Type locality: “Kidada, Lower-Congo,” Democratic Republic of Congo. Hemisus guttatum: Bocage (1895a:184), Monard (1938:56), Frade (1963:254). Hemisus marmoratum: Monard (1938:56). Hemisus marmoratus: Bocage (1887b:208). Hemisus marmoratus guineensis: Laurent (1950a: 15), Hellmich (1957a:28), Loveridge (1957:355). Hemisus guineensis: Laurent (1964a: 147), Haupl and Tiedemann (1978:18), Channing (2001:121), Rddel and Ernst (2003:27), Channing and Howell (2006:126), Onadeko and Rodel (2009:5), Fretey et al. (2011:27), Frost (2016), Ceriaco et al. (2016b:29). Hemisus guineensis mierops: Cei (1977:17, 18), Poynton and Broadley (1985a:533), Poynton (1992:71), Ruas (1996:28), Conradie et al. (2016:11). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola, south to the Caprivi Strip in Namib¬ ia, northern Botswana, and Mozambique to the extreme northeast border of South Africa, and also occurs from Senegal to Uganda and Kenya. Ocurrences in Angola (Map 25): The species is found in western and eastern Angola. Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a;183; Monard 1938:56; Laurent 1972:55). Kwanza Norte: “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:28). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:29). Lunda Norte: “Muita, Luembe E (Muita)” [-7.80000, 21.45000] (Laurent 1950a: 15; Laurent 1972:55; Ruas 1996:28). Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a: 147; Laurent 1972:55; Poynton 1992:71; Ruas 1996:28). Namibe: “Interieur de Mossamedes (Mossamedes)” [-15.20000, 12.15000] (Bocage 1887b:208; 1895a:184; Monard 1938:56; Ruas 1996:28). Undeter¬ mined Locality: “arid subregion, Angola” (Frade 1963:254); “plateaus regions, Angola” (Cei 1977:17); “arid territories along the coast, Angola” (Cei 1977:18). Taxonomic and distributional notes: See notes below for Hemisus marmoratus. Hemisus marmoratus (Peters, 1854) Marbled Snout-Burrower Engystoma marmoratum Peters 1854:628. Syntypes: ZMB 3548 [2 specimens] and ZMB \Q\{)l,fiide Bauer et al. (1995:41, 1996:260) (collector W.C.H. Peters). Type locality: “Cabaceira” (Peters 1954:628), Mozambique. 12*E l(i*E 2i>°E 24*E Map 25. Distribution of Hemisus guineensis in Angola. 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Hemisus marmoratus: Inger (1959:541), Schmidt and Inger (1959:170), Channing (2001:124), Rodel and Ernst (2003:27), Onadeko and Rodel (2009:5), Fretey et al. (2011:27), Frost (2016). Hemisus marmoratum: Bocage (1887a:183, 1895a:183), Boulenger (1905:107), Monard (1937a:29, 1938:56, 82), Frade (1963:254). Hemisus sudanense: Boulenger (1882:178). Hemisus marmoratus (?) angolensis: Hellmieh (1957a:28). Hemisus guineensis mierops: Laurent (1972:55), Ruas (1996:28). Global conservation status (lUCN): Least Concern. Global distribution: The species is widespread in sub-saharan Africa (excluding rainforest areas), extending from Senegal to Ethiopia and Somalia, south into northern South Africa. Ocurrences in Angola (Map 26): The species is found in western and eastern Angola. Kwanza Norte: “Dondo” [-9.68333, 14.43333] (Bocage 1895a:183; Monard 1938:56; Laurent 1972:55; Ruas 1996:28). Kwanza Sul: “Semba Acendu” [-9.85000, 16.15000] (Boulenger 1905:107; Monard 1938:56; Laurent 1972:55; Ruas 1996:28). Bie: “Chitau” [-11.43333, 17.15000] (Laurent 1972:55; Poynton 1992:71). Benguela: “Catumbella (Catumbela)” [-12.43333, 13.55000] (Bocage 1895a:183; Monard 1938:56; Laurent 1972:55; Ruas 1996:28); “Benguella (Benguela)” [-12.58333, 13.41667] (Boulenger 1882:179; Laurent 1972:55, Poyn¬ ton 1992:71). Huila: “Kalukembe (Caluquem- be)” [-13.78333, 14.68333] (Monard 1937a:29, 1938:56, 82; Laurent 1972:55; Ruas 1996:28). Map 26. Distribution of Hemisus marmoratus in Angola. Cuando Cubango: “Kakindo (Caquindo)” [-15.45000, 17.05000] (Monard 1937a:29, 1938:56, 82; Laurent 1972:55; Ruas 1996:28). Undetermined Locality: “western subregion and Angolan high¬ lands (Angolan coastal or watersheds)” (Cei 1977:18). Taxonomic and distributional notes: The delimitation of Hemisus guineensis (Cope, 1865) and Hemisus marmoratus (Peters, 1854) is more complicated than reflected by the current taxono¬ my. Both are likely part of a larger complex of morphologically similar species (Rodel and Ernst 2003; Onadeko and Rodel 2009). Both species have been reported to occur in Angola. However, Laurent (1972), in the only synoptic taxonomic review of the genus, explicitly considered Angolan populations to represent what he described as a subspecies of H. guineensis {H. guineensis mierops) that might even represent a distinct species. In West Africa, H. guineensis occurs in the forest or near to forests, and is common in grasslands and open woodlands, whereas H. marmora¬ tus is usually associated with savannas (Channing 2001; Channing and Broadley 2002; Rodel and Ernst 2003; Channing and Howell 2006; Onadeko and Rodel 2009; Frost 2016). Channing (2001) provided several records for both species in Angola but without information about the specimens corresponding to these records. Vouchers sampled from diverse habitat types and with associated genetic data are needed to evaluate which taxa are present in Angola as well as their distributions. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 85 Family Hyperoliidae Laurent, 1943 Genus Afrixalus Laurent, 1944 Afrixalus dorsalis (Peters, 1875) Striped Spiny Reed Frog Hyperolius dorsalis Peters 1875:206, pi. 1, fig. 2. Syntype: ZMB 4488 (eolleetor R. Buehholz) fide Bauer et al. (1995:44). Type locality: “Boutry” and “Victoria in einem Wassertumpel” restricted to “Boutry” by Mertens (1938b:24), [= mouth of River Butre], Ghana. Afrixalus dorsalis regularis: Laurent (1964a: 149) Cei (1977:17). Afrixalus dorsalis: Schiotz (1999:47), Channing (2001:136), Fretey et al. (2011:28), Channing et al. (2012:187), Frost (2016). Schiotz (1999) suggested that this species might extend from eoastal Cameroon to Ango¬ la. However, these populations vary in pheno¬ type and Angolan populations are somewhat different with some populations having a pale dorsal surfaee to the erus. For Angola, Schiotz (1999) considered two valid subspeeies, Afrix¬ alus dorsalis regularis Laurent, 1951 and Afrixalus dorsalis leptosomus (Peters, 1877). The latter was described by Peters (1877a) Map 27. Distribution of Afrixalus dorsalis in Angola. based on one individual from “Chinchoxo, Cabinda enclave,” whieh was originally placed in Hyperolius (Amiet 2012). Both, Sehiotz (1999) and Amiet (2012) discussed the nomenclatural confusion regarding leptosomus as a member of the fluvovittatus eomplex and Fretey et al. (2011) ineluded Afrixalus dorsalis regularis and A. d. leptosomus as synonyms of the nominate form. In contrast. Frost (2016) considered A. d. leptosomus to be a synonym of Afrixalus quadrivittatus (Werner, 1908) and^. d. regularis to be a synonym of^. dorsalis. Channing (2001) provided three records for Afrixalus dorsalis in Angola in Benguela Province without detailed information, which probably correspond to previously unpublished museum specimens. Global conservation status (lUCN): Least Coneem. Global distribution: The species is known from West Africa, from eastern Sierra Leone to extreme northern coastal Angola, but it has not been recorded from Benin (Gilles et al. 2006). Ocurrences in Angola (Map 27): The species oecurs in coastal Angola. Benguela: “Cubal da Ganda (Marco de Canavezes)” [-13.08333, 14.33333] (Laurent 1964a:149). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). Taxonomic and distributional notes: Afrixalus fulvovittatus (Cope, “1860” 1861) Four-Lined Spiny Reed Frog Hyperolius fulvovittatus Cope “1860” 1861:517. Holotype: ANSP 3219 fide Malnate (1971:350) (collector possibly Dr. Goheen). Type locality: “Liberia” (Cope “1860” 1861:517). Rappia fulvo-vittata: Gunther (1868 “1869”:479). Hyperolius fulvovittatus: Noble (1924:252). Rappia fulvovittata: Boulenger (1882:121), Ferreira (1904:112). Afrixalus fulvovittatus: Cei (1977:17), Schiotz et al. (2013), Frost (2016). 86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Guinea to western Ivory Coast, south to north-central Cameroon, Republic of Congo and northern Angola. Ocurrences in Angola (Map 28): The species occurs in northern Angola. Bengo: “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:112). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Boulenger 1882:121). Undetermined Locality: “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: The taxonomy of Afrixalus fulvovittatus (Cope, 1861) is still unresolved. Some authors, includ¬ ing Fretey et al. (2011), consider it a synonym of the broadly distributed Afrixalus quadrivit- Distribution Afrimlus fulvovittatus in Angola. tatus Werner, 1908, whereas others, such as Schiotz et al. (2013) and Frost (2016), consider this species to have a limited distribution in western Africa, including eastern Sierra Leone, southern Guinea, northern Liberia, and western Ivory Coast. This confusion, as well as the existence of other available names such as Afrixalus leptosomus (Peters, 1877), makes unclear the taxonomic status of specimens from Angola. A thorough taxonomic review incorporating genetic data from many populations across Central Africa is sorely needed. Afrixalus osorioi (Ferreira, 1906) Osorio’s Spiny Reed Frog Global conservation status (lUCN): Least Concern. Global distribution: The species is dis¬ tributed from west-central Angola northwards across much of the Congo Basin, western Kenya, and eastern Uganda. Ocurrences in Angola (Map 29): The species has only been recorded from its type locality “Quilombo” in Kwanza Norte Province, although is expected to be more widespread in the west-central plateaus of the country. Kwanza Norte: “Quilombo” Rappia osorioi Ferreira 1906:162, pi. 1. Syntypes: MHNFCP 017307 [3 speeimens] Ceriaeo et al. (2014a:22) (eolleetor F. Newton). Type loeality: “Quilombo” (Ferreira 1906:162) Kwanza Norte Provinee, Angola. Hyperolius osorioi: Noble (1924:153). Afrixalus osorioi: Perret (1976b:27), Cei (1977:17), Laurent (1982:24), Frost (1985:222, 2016), Sehiotz (1999:49), Channing (2001:140), Kohler et al. (2005:130), Fretey et al. (2011:28), Ceriaeo et al. (2014a:22). 10'^ IS'-S Map 29. Distribution of Afrixalus osorioi in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 87 [-9.33333, 14.90000] (Ferreira 1906:162; Perret I976b:27; Laurent 1982:24; Frost 1985:222, 2016; Ceriaco et al. 20I4a:22; Frost 2016). Taxonomic and distributional notes: Ferreira (1906) described Rappia osorioi based on three specimens, from “Quilombo” collected by Francisco Newton. Laurent (1982) discussed pat¬ terns of morphological variation in A. osorioi and the minor phenotypic differences from two other Congo Basin species, Afrixalus equatorialis (Laurent, 1941) Afrixalus leucostictus (Laurent, 1950). In his discussion of the variation in color and pattern of A. osorioi, Laurent noted that “la phenotype represente par I’holotype” has an elongate rectangular and dark scapular spot. Howev¬ er, the citation provided in this discussion, as well as the list of specimens examined, indicates Lau¬ rent was, in fact, referring to the holotype of Megalixalus fornasinii congicus Laurent, 1941 and not the type material described by Ferreira (1906). Perret (1976b) listed three type specimens in Museu do Porto lacking catalog numbers (one holotype and two paratypes) and followed Laurent in recognizing this taxon as conspecific withM / congicus (Ceriaco et al. 2014a). Channing (2001) provided two records without specific information that are probably based on museum specimens. Afrixalus quadrivittatus (Werner, 1908) Four-Lined Spiny Reed Frog Hyperolius leptosomus Peters 1877a:619, pi., figs. 5-5a. Holotype: ZMB 9175 (eolleetor Afrieanisehe Gesellsehaft) fide Bauer et al. (1995:44). Type loeality: “Chinehoxo (Westafrika)” (Peters 1877a:611), [= Chinehoxo] Cabinda Provinee, Angola. Synonymy with A. fulvovittatus (Werner, 1908) by Sehiotz (1975:78) and assigned by implieation to A. ''quadrivittatus'" by Piekersgill (2007b :23). Megalixus leptosomus quadrivittatus Werner 1908 “1907”: 1900, pi. 4, fig. 13. Syntypes: NHMW 3723 (eol¬ leetor F. Werner) fide Haupl et al. (1994:29). Type loeality: “On the Nile near Khor Attar, Sudan” (Wern¬ er 1908 “1907”: 1900), Sudan. Megalixus leptosomus: Boulenger (1882:129). Hyperolius leptosomus quadrivittatus: Werner (1908 “1907”: 1900). Afrixalus fiuvovittatus leptosomus: Loveridge (1957:322). Afrixalus dorsalis leptosomus: Perret (1976b:19), Laurent (1982:31), Amiet (2012:71). Afrixalus quadrivittatus: Largen (2001:357), Kolher et al. (2005:131), Frost (2016). Afrixalus “quadrivittatus”: Piekersgill (2007b:23), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from eastern Nigeria to western Ethiopia, southwards through western Tanzania and into northern Angola. Ocurrences in Angola (Map 30): The species occurs in “Chinehoxo” in the Cabinda enclave, but it is also presumed to occur in northern Angola. Cabinda: “Chinehoxo” [-5.10000, 12.10000] (Peters 1877a:619; Boulenger 1882:129; Loveridge 1957:322; Perret 1976b:19; Laurent 1982:31; Kolher et al. 2005:132; Amiet 2012:71). Taxonomic and distributional notes: The taxonomy and distribution of^. quadrivit¬ tatus remains controversial. It was removed from synonymy of Afrixalus fulvovittatus (Cope, 1861) by Sehiotz (1975) and Kolher et 12°E I6'E 20°E :4'>E Map 30. Distribution of Afrixalus quadrivittatus in Angola. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II al. (2005), where it had been placed by Laurent (1982) and recently placed again by Fretey et al. (2011). Pickersgill (2007b) suggested that the validity of this species is questionable, which leaves the Angolan records for this species in need of further study. Afrixalus wittei (Laurent, 1941) de Witte’s Spiny Reed Frog Megalixus wittei Laurent 1941:127. Holotype: MRAC 11500. Type loeality: “Lukafu” (Laurent 1941:127), Demoeratie Republie of Congo. Hyperolius fulvovittatus: Boeage (1866a:55). Rappia fulvovittata: Boeage (1895a: 175). Afrixalus wittei: Ferret (1976b:24), Frost (1985:224, 2016), Poynton and Broadley (1987:191), Sehiotz (1999:64), Channing (2001:143), Fretey et al. (2011:27), Ceriaeo et al. (2016b:30). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from tropical lowland savannas from the southern Democratic Republic of Congo, northeastern Angola and Zambia. Ocurrences in Angola (Map 31): The species was from some time only known from a single record from “Duque de Braganga” [= Calandula], Malanje Province. It was recent¬ ly found in Cangandala National Park, in the same region. Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Boeage 1866a;55, 1895a: 175; Ferret 1976b:24); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo etal. 2016b:30). Taxonomic and distributional notes: Ferret (1976b) considered the specimen from “Duque de Braganga” identified by Boeage as Hyperolius fulvovittatus (Boeage 1866a), and later as Rappia fulvovittata (Boeage 1895a), to be referable to Afrixalus wittei (Laurent, 1941). Through analysis of the morphological diversi¬ ty in several species of Afrixalus, Laurent 3 1 . Distribution of Afrixalus wittei in Angola. (1982) argued that A. wittei is closely related to A. osorioi Ferreira, 1906 (Poynton and Broadley 1987). Genus Cryptothylax Laurent and Combaz, 1950 Cryptothylax greshoffii (Schilthuis, 1889) Greshoee’s Wax Frog Hylambates Greshoffii Schilthuis 1889:286, unnumbered fig. Type: UZ by original designation, now possibly deposited in RMNH fide Hoogomoed in Frost (1985:205) (eolleetor A. GreshofiE). Type loeality: “Boma (Congo, W. Afriea)” (Sehilthuis 1889:284), [= Boma], Demoeratie Republie of Congo. Hylambates greshoffii: Frost (1985:205). Cryptothylax greshoffii: Sehiotz (1999:90), Blaekbum and Jaekson (2006:358), Fretey et al. (2011:29), Chan¬ ning et al. (2012:200), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the forests of southern Cameroon extending into the Democratic Republic of Congo, and south to northern Angola. Ocurrences in Angola: The species occurs in northern Angola. Uige: without precise location (Sehiotz 1999:91). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 89 Taxonomic and distributional notes: This forest edge species has been reported from sever¬ al parts of the Congo Basin, where wetlands are very widespread, and probably represents the center of its dispersion (Amiet 2012). Schiotz (1999) provided a distribution map of the species with a record in Uige Province, in northern Angola, although without further detail. Genus Hyperolius Rapp, 1842 Hyperolius adspersus Peters, 1877 Sprinkled Long Reed Frog Hyperolius adspersus Peters 1877a:619, pi., figs. 6-6a. Holotype: ZMB 9176 fide Laurent (1961:92). Type loeality: “Chinehoxo” Cabinda Provinee, Angola. Rappia nobrei Ferreira 1904:112. Syntypes: MHNFCP 017292 [2 specimens] (collector F. Newton). Type locality: “Cabiri” Bengo Province, Angola. Considered a junior synonym ofH. adspersus by Ceriaco et al. (2014a:21). Hyperolius nasutus adspersus: Laurent (1961:92). Hyperolius granulatus: Laurent (1964a: 155), Cei (1977:17). Hyperolius adspersus: Amiet (2005:274), Fretey et al. (2011:29), Channing et al. (2013b:314), Frost (2016). Hyperolius cf adspersus: Ceriaco et al. (2014a:21). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Cameroon to the lower Congo Basin in Democratic Republic of Congo to northern Angola, including the Cabinda encalve. Ocurrences in Angola (Map 32): The species is known from the type locality “Chin- choxo” and from scattered localities in north¬ eastern Angola. Cabinda: “Chinehoxo” [-5.10000, 12.10000] (Peters 1877a:619; Lau¬ rent 1961:92; Amiet 2005:275; Channing et al. 2013b:317; Frost 2016). Bengo; “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:112; Ceriaco et al. 2014a:21). Lunda Sul: “Alto Chicapa galerie forestiere des sources du ruis- seau Cuilo” [-10.88333, 19.23333] (Laurent 1964a: 155). Moxico: “Reserve de chasse de Cameia, 120 km a Test de Luso” [-11.83333, 21.00000] (Laurent 1964a: 155); “Rives du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a: 155). Taxonomic and distributional notes: This species was described by Peters (1877a) as Hyperolius adspersus based on a specimen collected in “Chinehoxo,” in the Cabinda enclave. Ferreira (1904) described Rappia nobrei from the type locality “Cabiri” collected by Francisco Newton. This taxon was previously considered as incertae sedis in part because of the long lack of study of the type specimens (Frost 2016). A recent study by Ceriaco et al. (2014a) confirmed that this taxon is clearly a member of the genus Hyperolius and is plausibly a junior synonym of H. adspersus. Previously, H. adspersus was considered to be a synonym of H. nasutus Gunther 1865, but Amiet (2005) removed it from synonymy. This species is part of the taxonomically prob¬ lematic Hyperolius nasutus group, which currently contains 1 6 recognized species (Channing et al. 2013b; Ceriaco et al. 2014a). Of these 16 species, only three are known to occur near the type local¬ ity of Rappia nobrei: Hyperolius adspersus, Hyperolius nasutus Gunther, 1865, and H. benguel- Map 32. Distribution of Hyperolius adspersus in Angola. 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II lensis (Bocage, 1893) (Channing et al. 2013b; Ceriaco et al. 2014a; see H. benguellensis and H. nasutus accounts). The syntypes of R. nobrei are similar to both H. adspersus and H. benguel¬ lensis, which makes it difficult to determine to which species Rappia nobrei should be referred. We here follow Ceriaco et al. (2014a) by considering R. nobrei to be a junior synonym of H. adsper¬ sus. Channing et al. (2013b) comidQrQd Hyperolius granulatus (Boulenger, 1901), previously asso¬ ciated with Hyperolius benguellensis (Bocage, 1 893), to be a synonym of H. adspersus (Ferret 1976a; Schiotz and Van Daele 2003). Hyperolius angolensis Steindachner, 1867 Angolan Reed Frog [Note: Because of the complexity of the taxonomy of this species we provide additional commentary on its synonymy/chresonymy] Hyperolius marmoratus Rapp 1842:289, pi. 6, figs. 1-2. Type: Not designated (eolleetor unknown). Type loeality: “Natal” [= KwaZulu-Natal], South Afriea. Hyperolius parallelus Gunther 1858a:326. Syntypes: BMNH 1947.2.9.41-43 [3 speeimens] (eolleetor unknown, probably “Mr. Rieh”). Speeimens BMNH 1947.2.9.41 and 1947.2.9.42 are depleted in Gun¬ ther “1858b” 1859: 86, pi. 8, fig. A. Type loeality: “Sud-Afrika” later ehanged to “South Afriea” and “Angola” (Gunther “1858b” 1859:86) and to “Cape of Good Hope” and “Ambris, Angola” [= Ambriz] by Boulenger (1882:121). Synonymized with H. marmoratus by Boulenger (1882:121) and treated as a subspeeies by Laurent (1951:38). Hyperolius marmoratus var. angolensis Steindaehner 1867:50. Syntypes: NHMW 13487.1-2 fide Haupl and Tiedemann (1978:22) (eolleetor F.A.P. Bayao). Type locality: “Angola” (Steindachner 1867:50). Restricted to “Duque de Bragan^a” [= Calandula] (Ceriaco et al. 2014b:669). Hyperolius insignis Bocage 1867a:844, fig. 2. Syntypes: MBL T.21-164, 27-167 [2 specimens] (collector J. A. d’ Anchieta [Benguella] and P. Barroso [St. Salvador du Congo]), destroyed by fire 18 March 1978. Type locality: “Benguella,” Angola. Corrected by Ferret (1976a: 28) to “St. Salvador du Congo” [= M’Banza Congo] and “Novo Redondo” [= Sumbe], Angola. Synonymized with H. marmoratus by Boulenger (1882:121) and Ferret (1976a:27) and treated as a subspecies by Laurent (1951 “1952”:39). Synonymized with H. parallelus by Loveridge (1953a:329). Hyperolius Toulsonii Bocage 1867a:845, fig. 3. Holotype: MBL T.27-275 (collector M. Toulson), destroyed by fire 18 March 1978. Type locality: “Loanda” [= Luanda], Angola. Synonymized with H. marmoratus by Boulenger (1882: 121), recognized as valid by Ahl (1931b:388), and considered a synonym ofH. m. insignis by Laurent (1952 “1951”:391), and later synonymized with H. parallelus (Ferret 1976a:27). Hyperolius huillensis Bocage 1873b:225. Syntypes: MBL 2-166 [4 specimens] (collector J. A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Huilla,” [Angola] (Bocage 1873b:225). Considered a subspecies ofH. marmoratus by Laurent (1952 “1951”:390). Hyperolius vermieulatus Peters 1882a:8. Lectotype: ZMB 10050, designated by Laurent (1961:88) (collector von F.W.A. von Mechow). Type locality: “Malange” [=Malanje] (Peters 1882a: 8). Considered a sub¬ species ofH. marmoratus by Laurent (1952 “1951”: 88). Rappia plieifera Bocage 1893:118. Syntypes: MBL T.22-209 [2 specimens] (collector J. A. d’Anchieta [Caconda] and F. A. P. Bayao [Duque de Braganga]), destroyed by fire 18 March 1978. Haupl and Tiede¬ mann (1978:29) considered NHMW 22895 [2 specimens] as part of the type series. Type locality: “Caconda,” “Duque de Braganga,” Angola (Bocage 1893:118). Synonymized with /£ m. vermieulatus hy Laurent (1952 “1951”:390) and later with H. parallelus by Ferret (1976a:27). Rappia marmorata var. marginata: Bocage 1895a: 164: Type: MBL, specimen number(s) not known (collec¬ tor F. A. P. Bayao), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” [= Calandu¬ la], Angola. Rappia marmorata var. taeniolata Bocage 1895a: 164: Syntypes: MBL, specimen numbers not known (col¬ lector F. A. P. Bayao [Duque de Braganga], Graga and J. A. d’Anchieta [Huilla] and J. A. d’Anchieta [Caconda and Cahata]), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga,” “Huil¬ la,” “Caconda” and “Cahata” [= Calandula, Huila, Caconda and Caota], Angola. Rappia marmorata var. variegata Bocage 1895a: 164: Syntypes: MBL (collector J. A. d’Anchieta). Type local- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 91 ity: “Cahata” and “Quindumbo” [= Caota and Quindumbo], Benguela Province, Angola. Hyperolius marungaensis 1931b:351, fig. 226. Holotype: ZMB uncatalogued (collector Bohm). Type locali¬ ty: “Marunga,” Angola. Considered a subspecies ofH. marmoratus by Laurent (1961:88) Hyperolius decoratus 1931b:352, fig. 227. Holotype: ZMB uncatalogued, MCZ A-17632 (collectors Bruhl and Gleim) - on exchange from ZMB fide Barbour and Loveridge (1946:127). Type locality: “Longa” (Ahl 1931a:78, 1931b:352), Angola. Considered a subspecies of H. marmoratus by Laurent (1952 ‘T951”:392). Hyperolius mierotietus 1931b:353, fig. 228. Syntypes: ZMB [2 specimens] uncatalogued, ZMB 36100 [2 specimens] (collector unknown). Type locality: “Longa” (Ahl 1931a:80, 1931b:352), Angola. Consid¬ ered a synonym of H. m. deeoratus by Laurent (1961:89). Hyperolius erythromelanus Monard 1937a:36. Syntypes: LMCFM [2 specimens] (collector A. Monard). Type locality: “Sangeve” (Monard 1937a:36), [= Sangueve], Angola. Synonymized with H. m. huillensis by Laurent (1952 “1951”:390). Hyperolius marmoratus alborufus: Laurent 1964a: 153, fig. 40. Holotype: MD 5679 (collector unknown). Type locality: “Cazombo, Alto Zambeze, Moxico, Angola” (Laurent 1964a: 153). Hyperolius huillensis'. Bocage (1879c:89). Hyperolius insignis: Bocage (1867a:844, 1887a: 191). Rappia insignis'. Gunther (1869:479). Hyperolius marmoratus'. Gunther (1864a:480), Bocage (1866a:55, 1886b:74), Peters (1881:150), Noble (1924:253), Schmidt (1936:131), Mertens (1938a:427), Inger (1959:541), Gavetti and Andreone (1993:103), Ceriaco et al. (2014b:669). Hyperolius eitrinus: Bocage (1879c:89). Hyperolius Toulsinii: Bocage (1895a: 166, 1897a:203), Ferreira (1906:161). Rappia marmorata: Boulenger (1882:121, 1905:109), Bocage (1895a: 164, 1896a: 113, 1897b:211), Ferreira (1904:112, 1906:160) Rappia plieifera'. Bocage (1895a:167, 1897a:203), Ferreira (1897b:241, 1904:112, 1906:161), Haupl and Tie- demann (1978:29), Haupl et al. (1994:34). Rana marmorata var. huillensis'. Ferreira (1897b:241). Hyperolius deeoratus'. Loveridge (1936a: 106), Monard (1937a:35, 1938:89); Barbour and Loveridge (1946:127). Hyperolius vermieulatus: Monard (1938:88). Hyperolius mierostietus: Monard (1937a:35, 1938:90). Hyperolius sp.II (ap. angolensis)'. Monard (1937a:38, 1938:92). Hyperolius sp.III (ap. angolensis)'. Monard (1937a:38, 1938:93). Hyperolius sp. I (ap. deeoratus)'. Monard (1937a:37, 1938:92). Hyperolius angolensis'. Monard (1937a:36, 1938:90), Channing (2001:148), Conradie et al. (2012a:2), Ceria¬ co et al. (2016b:31), Conradie et al. (2016:14). Hyperolius graueri'. Mertens (1937a:20). Hyperolius parallelus: Peters (1877a:618), Laurent (1943a:14), Ceriaco et al. (2014b:669), Frost (2016). Hyperolius marmoratus angolensis'. Laurent (1950a: 17, 1954a:80), Frade (1963:254), Laurent (1964a: 152), Cei (1977:17), Broadley (1965a:26), Haupl and Tiedemann (1978:22). Hyperolius marmoratus albofaseiatus'. Loveridge (1953a:350). Hyperolius marmoratus parallelus'. Laurent (1961:89), Cei (1977:17). Hyperolius marmoratus huillensis'. Laurent (1961:88), Cei (1977:17), Poynton and Haacke (1993:15). Hyperolius marmoratus vermieulatus'. Laurent (1961:88). Hyperolius marmoratus marungaensis'. Laurent (1961:88), Cei (1977:17). Hyperolius marmoratus insignis'. Laurent (1961:89, 1964a: 151) Cei (1977:17). Hyperolius marmoratus alborufus'. Laurent (1964a: 153), Cei (1977:17). Hyperolius parallelus alborufus'. Schiotz (1975:185). Hyperolius parallelus plieiferus'. Perret (1976a:27). Hyperolius parallelus toulsoni'. Perret (1976a:27). Hyperolius parallelus insignis'. Perret (1976a:29). 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Hyperolius parallelus huillensis: Ferret (1976a:29). Hyperolius erythromelanus: Frost (1985:210). Hyperolius parallelus-marginatus subgroup: Sehiotz (1999:217). Hyperolius {Hyperolius) marmoratus: Fretey et al. (2011:32). Hyperolius {Hyperolius) parallelus: Fretey et al. (2011:32). Global conservation status (lUCN): Least Coneem. Global distribution: The species is known from Angola and the adjacent countries of Nami- ba, Democratic Republic of Congo, and Zambia. Ocurrences in Angola (Map 33): The species is very widespread for almost the entire territory. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a;618; Bocage 1895a: 164; Laurent 1961:89). “Landana” [-5.21667, 12.15000] (Laurent 1943a:14). Zaire: “St. Salvador du Congo” [-6.26667, 14.23333] (Bocage 1887a:191, 1895a;164; Ferret 1976a:29). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1867a:845, 1895a:166, 1897b:203; Loveridge 1936a:106, 1953a:352; Laurent 1961:88, Ferret 1976a;27); “Longa, Loanda” (Ahl 1931a:78, 1931b:352; Barbour and Loveridge 1946:127). Bengo: “Ambriz” [-7.844312, 13.106493] (Boulenger 1882:121; Loveridge 1953a:352). Malanje: “Tembo Aluma” [-7.70000, 17.28333] (Bou¬ lenger 1905:109); “Bange N’gola” [-8.43333, Distribution of Hyperolius angolensis in 16.56667] (Boulenger 1905:109); “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:55, 1866b:74, 1893:118, 1895a;164, 167, 1897a:203; Boulenger 1882:121); “Malanje” [-9.55000, 16.35000] (Peters 1882a:8, Laurent 1961:88); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:669); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:31). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 17, 1954a;80, 1964a; 152); “Matalarive gauche de laThshiumbe, 40 km a Lest de Dundo” [-7.43333, 21.16667] (Laurent 1950a:17); “Carumbo lagoon” [-7.74422, 19.95467] (Conradie et al 2012a;2); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:17, 1954a:80); “Andrada (Luembe O)” [-7.68729, 21.37003] (Laurent 1954a:80); “Caluan- go, Riv. Camaloa, affl. Caluango” [-8.33333, 19.65000] (Laurent 1964a:152); “Sombo, riv. Melan- da, affl. Chiumbe” [-8.68333, 20.95000] (Laurent 1954a:80, 1964a: 152). Lunda Sul: “Alto Cuilo, mare Tchifuka” [-10.00000, 19.58333] (Laurent 1964a: 152). Mexico: “rives de Lumeje, pres du lac Calundo” [-11.51667, 20.76667] (Laurent 1964a:152); “rives du lac Calundo, 105 km a Lest de Luso” [-11.80000, 20.86667] (Laurent 1964a:152); “Reserve de chasse de Cameia, 120 km a Lest de Luso” [-11.83333, 21.00000] (Laurent 1964a:152); “Cazombo (High Zambeze)” [-11.88333, 22.91667] (Laurent 1964a;153; Sehiotz 1975:185). Kwanza Norte: “N’golla Bumba” [-9.03333, 14.60000] (Ferreira 1906:161); “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:109; Fer¬ reira 1906:161); “Cambondo” [-9.15963, 14.65827] (Ferreira 1906:160); “Canhoca” [-9.25000, 14.68333] (Boulenger 1905:109); “Cazengo” [-9.33333, 14.76667] (Ferreira 1904:112); “Quilom- bo” [-9.33333, 14.90000] (Ferreira 1906:160). Kwanza Sul: “Novo Redondo Sumbe” [-11.20000, 13.85000] (Bocage 1895a:164; Ferret 1976a:29); “Gumba” [-11.26667, 14.28333] (Ferreira MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 93 1904:112). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard I937a:35-36, 38; 1938:89, 90, 92). Bie: “Gauca” [-II. 18333, 17.45000] (Schmidt 1936:131); “Chitau” [-11.43333, 17.15000] (Schmidt 1936:131); “Goedecke Farm, Conjo, 25 km north Genera Maehado (Camaeopa)” [-12.01667, 17.06667] (Mertens I937a:20); “Bihe” [-12.38333, 16.95000] (Boeage I879e:89, 1895a: 164); “Cubango basin (10)” [-I3.7I6I6, 17.09661] (Conradie et al. 2016:8-9, 14); “Cuando basin (I2a)” [-13.59333, 16.87986] (Conradie et al. 2016:8-9, 14); “Cuando basin (16)” (Conradie et al. 2016:8-9, 14). Benguela: “Cahata” [-12.35000, 14.81667] (Boeage 1895a: 164); “Quindum- bo” [-12.46667, 14.93333] (Boeage 1895a: 164); “Benguella” [-12.58333, 13.41667] (Boeage 1867a:844, 1887a:191; Gunther 1869:479; Boulenger 1882:121; Loveridge 1936a:106, 1953a:352; Laurent 1961:89); “Ebanga” [-12.73333, 14.73333] (Monard 1937a:35, 37; 1938:89, 92); “Dombe” [-12.95000, 13.10000] (Boeage 1895a:164; Laurent 1961:89; Gevetti and Andreone 1993:104); “Cubal” [-13.03333, 14.25000] (Mertens 1938:427); “Mareo de Canavezes (Cubal da Ganda)” [-13.08333, 14.33333] (Laurent 1964a:151); “Hanha” [-13.30000, 14.20000] (Boeage 1867a:844, 1896a:113, 1897b:211; Boulenger 1882:121); “Tongrube neben dem Jamba-Flup, Entre Rios” [-13.60000, 16.60000] (Hellmieh 1957a:28). Hufla: “12 km W of Bela Vista” [-12.56667, 16.21667] (Poynton and Haaeke 1993:15); “Rio Cuee” [-13.51667, 15.20000] (Ferrei¬ ra 1897b:241); “Caeonda” [-13.73333, 15.06667] (Boeage 1893:118, 1895a:164, 167, 1897a:203; Perret 1976a:27; Haupl and Tiedemann 1978:29; Haupl et al. 1994:34; Gevetti and Andreone 1993:104); “Kalukembe” [-13.78333, 14.68333] (Monard 1937a:35, 38; 1938:92); “Sangeve” [-13.88333, 15.83333] (Monard 1937a:35; 1938:89, 91; Frost 1985:210); “Kuvangu” [-14.46667, 16.30000] (Monard 1937a:35-38; 1938:90, 93); “Kapelongo” [-14.88333, 15.08333] (Monard 1937a:36, 1938:90); “Humpata” [-14.93333, 13.51667] (Conradie et al. 2012a:2); “Huilla” [- 15.05000, 13.55000] (Gunther 1865a:480; Boeage 1873b:225, 1895a:164; Boulenger 1882:121, Laurent 1961:88; Perret 1976a:29); “Nunteehite lagoon” [-15.13333, 13.41667] (Poynton and Haaeke 1993:15). Cuando Cubango: “Marunga oder Kawende” [-17.45000, 20.03333] (Ahl 1931a:77, 1931b:351; Laurent 1961:88); “Kuandu” [-16.74487, 19.10136] (Monard 1937a:36; 1938:90); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 14); “(7e)” [-14.43916, 17.81491] (Conradie et al. 2016:8-9, 14); “Cuando basin (19)” [-14.70213, 17.37772] (Conradie et al. 2016:8-9, 14); “Cuando basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:8- 9, 14); “Cuando basin (22a)” [-14.66622, 16.97842] (Conradie et al. 2016:8-9, 14); “Cuito basin (23)” [-14.58972, 18.17083] (Conradie et al. 2016:8-9, 14); “Cuito basin (24)” [-14.60622, 18.46722] (Conradie et al. 2016:8-9, 14); “Cuito basin (24a)” (Conradie et al. 2016:8-9, 14); “Cuito basin (26)” [-15.08686, 19.14872] (Conradie et al. 2016:8-9, 14); “ Cuito basin (27)” [-15.17127, 19.19433] (Conradie et al. 2016:8-9, 14); “Cuito basin (28)” [-15.13486, 19.19636] (Conradie et al. 2016:8-9, 14); “Cubango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:9-10, 14); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9-10, 14); “Cuito basin (30b)” [-17.51194, 20.04305] (Conradie et al. 2016:9-10, 14); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2016:9-10, 14); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9- 10, 14); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9-10, 14); “Cuando basin (41b)” [-17.46777, 23.07944] (Conradie et al. 2016:9-10, 14); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 14); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie etal. 2016:9-10, 14); “Cubango basin (45)” [-16.88350, 18.01180] (Conradie etal. 2016:9, 12, 14); “Cuito basin (51)” [-14.58970, 18.1711] (Conradie et al. 2016:9, 12, 14); “Cuito basin (53b)” [-14.56322, 18.44394] (Conradie et al. 2016:9, 12, 14); “Cuito basin (54)” [-14.46810, 18.35488] (Conradie et al. 2016:9, 12, 14); “Cuito basin (55)” [-14.68478, 18.67369] (Conradie et al. 2016:9, 12, 14); “Cuito basin (57)” [-15.45969, 18.76833] (Conradie et al. 2016:9, 12, 14); “Cuito basin (58)” [-15.38206, 19.06375] (Conradie et al. 2016:9, 12, 14); “Cuito basin (62)” [-17.50875, 94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 20.06608] (Conradie et al. 2016:9, 12, 14); “Cuito basin (63)” [-17.93611, 21.10269] (Conradie et al. 2016:9, 12, 14). Undetermined Locality: “Without precise location” (Bocage 1887a:191, Lau¬ rent 1961:88); “north of Quanza” (Bocage 1895a: 164); “Between Benguela and Bihe” (Boulenger 1905:109); “Locomi” (Boulenger 1905:109); “plateaus regions” (Cei 1977:17); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17) Taxonomic and distributional notes: This species was first described as Hyperolius angolen- sis Steindachner, 1 867 based on a single specimen from “Angola,” now corrected to “Duque de Brangan^a” [= Calandula] (Ceriaco et al. 2014b); note, however, that Haupl and Tiedemann (1978) cited two syntypes from “Angola” deposited in the Naturhistorsches Museum in Wein, Austria. This species has been reported from Angola by many previous authors under a variety of names. This taxon is considered to be a synonym of Hyperolius parallelus Gunther, 1858 by some authors such as Wieczorek et al. (2000), Fretey et al. (2011), Ceriaco et al. (2014b), and Frost (2016). Other authors, including Schiotz (1999), considered it part of the Hyperolius parallelus-marginatus subgroup — corresponding to one of three taxa, parallelus, insignis and angolensis — or instead a subspecies of Hyperolius marmoratus (Rapp, 1842) as was suggested by Loveridge (1953a) and Poynton and Broadley (1987). Channing (2001) and Conradie et al. (2012a) comidQVQdi Hyperolius angolensis to be a valid species. Both H. parallelus and H. marmoratus are taxonomically prob¬ lematic. Poynton and Broadley (1987) regarded all the forms in southern Africa as subspecies of H. marmoratus, while Channing (1999) regarded angolensis and possibly other forms from Ango¬ la as specifically distinct from the eastern marmoratus-forms (Schiotz 1999). Hyperolius parallelus is represented in southern and southwestern Africa, including Angola (Schiotz 1999) whereas, H. marmoratus, is widespread across southern and eastern Africa and appears to be absent from Angola (Channing 2001; Frost 2016). The nomina parallelus and marmoratus were previously reported from the country under various names associated with many different species and subspecies names (Ferreira 1906; Loveridge 1953a; Monard 1937a, 1938; Mertens 1938; Laurent 1964a; Perret 1976a; Fretey et al. 2011). This is explained partly by the considerable intraspecific and interspecific variation in coloration and pattern across populations (Ceriaco et al. 2014b). There are several studies in progress on the genus Hyperolius that will help to clarify species boundaries among these challenging taxa. Given the current state of knowledge, we refer all of these records to Hyperolius angolensis, though we recognize that this may contain multiple biological species. Hyperolius benguellensis (Bocage, 1893) Benguela Long Reed Frog Rappia benguellensis Bocage 1893:119. Syntypes: MBL 27.220-223 (collector J. A. d’ Anchieta), destroyed by fire 18 March 1978. Type locality: “Cahata dans I’interieur de Benguella” (Bocage 1893:119), [= Caota] Benguela Province, Angola. Rappia benguellensis: Bocage (1895a:169, 1897a:204, 1897h:211), Ferreira (1906:161). Hyperolius nasutus: Monard (1937a:39, 1938:94). Hyperolius benguellensis: Noble (1924:252), Monard (1937a:34; 1938:87), Frade (1963:254), Perret (1976a:27), Frost (1985:207, 2016), Poynton and Broadley (1987:208), Amiet (2005:292), Schiotz and Van Daele (2003:128), Amiet (2005:275) Schiotz (2006:62), Conradie et al. (2012a:2), Channing et al. (2013:317). Hyperolius oxyrhynehus: Laurent (1950a: 17). Hyperolius nasutus: Schiotz (1999:97), Channing et al. (2002:96). Hyperolius {Hyperolius) benguellensis: Fretey et al. (2011:29). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Democratic Republic of Congo (former MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 95 Katanga Province), southern Angola and adja¬ cent northern Nambia and Botswana. Ocurrences in Angola (Map 34): The species is known from western Angola from the type locality “Cahata, Benguella,” although there are some records further north. Kwanza Norte: “Quilombo” [-9.33333, 14.90000] (Fer¬ reira 1906:161). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:87). Bie: “Cubango basin (11)” [-13.69413, 17.06177] (Conradie et al. 2016:8-9, 15); “Cubango basin (12a)” [-16.59333, 16.87986] (Conradie et al. 2016:8-9, 15); Benguela: “Cahata (Caota)” [-12.35000, 14.81667] (Boeage 1893:119 1895a:169, 1897a:204; Perret 1976a:27; Frost 1985:207, 2016; Poynton and Broadley 1987:208; Channing et al. 2002:96; Amiet 2005:275; Channing et al. 2013:318); “Eban- ga” [-12.73333, 14.73333] (Monard 1938:87); “Hanha” [-13.30000, 14.20000] (Boeage 1897b:211). Huila: “Indungu” [-14.81667, 16.26667] (Monard 1938:87); “Zootecnica Plateau, Humpata” [-14.96581, 13.34458] (Channing et al. 2013:317); “Humpata” [-14.23814, 13.43331] (Channing et al. 2013b:317); “Humpata” [-15.03333, 13.40000] (Conradie et al. 2012a:2). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937a:39, 1938:94). Cuando Cubango: “Cubango basin (5)” [-14.74628, 17.66844] (Conradie et al. 2016:8-9, 15); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 15); “ Cubango basin (7b)” [-14.43377, 17.82957] (Conradie et al. 2016:8-9, 15); “Cubango basin (7c)” [-14.43916, 17.81491] (Conradie et al. 2016:8-9, 15); “Cubango basin (8)” [-14.25705, 17.77852] (Conradie et al. 2016:8-9, 15); “Cubango basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:8-9, 15); “Cubango basin (22b)” [-14.66278, 16.96081] (Conradie et al. 2016:8-9, 15); “Cuito basin (27)” [-15.17127, 19.19433] (Conradie et al. 2016:8-9, 15); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9-10, 15); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2016:9-10, 15); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9-10, 15); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016: 9-10, 15); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 15); “Cubango basin (45)” [-16.88350, 18.01180] (Conradie et al. 2016:9, 12, 15). Undetermined Locality: “Western subregion and Angolan highlands (Angolan coastal or watersheds)” (Frade 1963:254). Taxonomic and distributional notes: In the description, Boeage notes that the type speci¬ mens are similar to Rappia nasuta (Gunther, 1865). In their detailed diseussion of the benguelen- sis-nasutus complex, Poynton and Broadley (1987) had little doubt that the characters used at that time were inadequate to separate all available specimens into nasutus and benguellensis. Schiotz (1999) concluded that it is difficult to separate H. nasutus and H. benguellensis based on mor¬ phology and color patterns. Channing et al. (2002) proposed formal ehanges in nomenelature and gave detailed lists of synonyms, but Schiotz (2006) questioned several of these changes. Schiotz and Van Daele (2003) provided a discussion about the differenees in the advertisement calls of nasutus and benguellensis as well as differences in external morphology. Hyperolius benguellensis (Boeage, 1893) was questionably removed from the synonymy of H. nasutus Gunther, 1865 by Amiet (2005). Channing et al. (2013) used molecular data to argue that H. benguellensis is a valid 12°E \e>E 20°E Map 34. Distribution of Hyperolius benguellensis in Angola. 96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II species. The species has only been confirmed from southern Angola, northern Namibia, and north¬ ern Botswana where it was found in open grassy habitats, along streams or man-made structures with emergent vegetation (Channing et al. 2013). We follow Poynton and Broadley (1987), Schiotz and Daele (2003), and Channing et al. (2013) in regarding H. benguellensis as distinct from the sympatric H. nasutus. Hyperolius oxyrhynchus is regarded as a synonym of H. benguellensis (Channing et al. 2013b), though a record from “Muita” (Laurent 1950) is probably a misidentifi- cation as it is far to the north of the rest of the distribution of H. benguellensis. Laurent’s (1950a) record should probably be referred to another species of the nasutus complex, such as H. adsper- sus or H. nasutus. Hyperolius bicolor Ahl, 1931 Two-Colored Reed Frog (Endemic) Hyperolius bicolor Ahl 193 la: 129. Syntypes: ZMB, lost (collector K. May). Type locality: “Farenda Bango, Loanda” (Ahl 193 la: 129), [= Fazenda Bango, Luanda], Angola. Hyperolius bicolor. Frost (1985:208, 2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 35): The species is known only from the type locality “Fazenda Bango, Loanda”. Luanda; “Farenda Bango” [-9.81767, 14.78916] (Ahl 1931a:129; Frost 1985:208, 2016). Taxonomic and distributional notes: According to Laurent {in Frost 1985:208, 2016) the species is probably a synonym of Hyperolius marmoratus insignis, a member of parallelus-marginatus subgroup (Schiotz 1999) [see Hyperolius angolensis account]. The validity of this taxon is questionable and difficult to address due to loss of the type spec¬ imen. 12*E I6®E 20°E 24*E Map 35. Distribution of Hyperolius bicolor in Angola. Hyperolius bocagei Steindachner, 1867 Bocage’s Reed Frog Hyperolius bocagei Steidachner 1867:51, pi. 5, fig. 11. Holotype: NHMW 14846 fide Haupl and Tiedemann (1978:22) (Collector F.P. Bayao). Type locality: “Angola” (Steindachner 1897:51). Rappia bocagei var. maculata Ferreira 1906:160. Type: Not stated (collector F. Newton), see Ceriaco et al. (2014a:25-26) discussion. Type locality: “Golungo Alto”, Angola. Rappia seabrai Ferreira 1906:163. Holotype: (collector F. Newton), not located, probably lost. Paratype: MNHFCP 018587 (collector F. A. P. Bayao). Type locality: “Quilombo, Rio Luinha” [= Gonguembo] Kwanza Norte, Angola. Hyperolius bocagei: Bocage (1873b:225), Noble (1924:252), Parker (1936:143), Laurent (1950a: 16, 1954a:79, 1964a: 150), Cei (1977: 17); Poynton and Broadley (1987:212), Haupl and Tiedemann (1978:22), Frost (1985:208, 2016), Haupl et al. (1994:26), Schiotz (1999:188), Channing (2001:151), Schiotz and Van Daele (2003:145), Ceriaco et al. (2014a:24). Rappia bocagii: Boulenger (1882:126, 1905:109), Bocage (1895a: 165, 1897a:203). Rappia cinctiventris: Bocage (1895a: 168), Boulenger (1905:110). Rappia bocagei: Ferreira (1904:112, 1906:160). MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 97 Hyperolius seabrai: Noble (1924:253), Schmidt (1936:132), Monard (1937a:39, 1938:95), Cei (1977:17), Frost (1895:118). Hyperolius cinctiventris: Noble (1924:252), Monard (1938:93), Inger (1959:541), Cei (1977:17). Hyperolius {Hyperolius) boeagei: Fretey et al. (2011:29). Global conservation status (lUCN): Least Concern. Global distribution: The species is known only from the Democratic Republic of the Congo, to Tanzania, Angola and Zambia. Ocurrences in Angola (Map 36): The species has been reported from scattered local¬ ities mainly in the westerns regions of the country. Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:165, 1897a:203). Luanda: “Cacuaco?” [-8.78333, 13.36667] (Ferreira 1904:112). Kwanza Norte: “Canhoca” [-9.25000, 14.68333] (Bou- lenger 1905:109); “Golungo Alto” [-9.13333, 14.76667] (Ferreira 1906:160); “Quilombo, Rio Luinha” [-9.33333, 14.90000] (Ferreira 1906:163; Ceriaco et al. 2014a:24). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:143). Malanje: “Bange N’gola” [-8.43333, 16.56667] (Boulenger 1905:110); “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1873b:225, 1895a:165, 1897a:203; Ferreira 1906:163; Ceriaco et al. 2014a:24). Map 36. Disnibutionof//y;.ero//.. Wge/m Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 16, 1954a:79, 1964a: 150); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:16). Mexico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:150). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:40 1938:93, 96). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1936:132); “Bihe” [-12.38333, 16.95000] (Bocage 1895a:168). Benguela: “Ebanga” [-12.73333, 14.73333] (Monard 1938:93); “Dombe” [-12.95000, 13.10000] (Bocage 1895a:168). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:165); “Kuvangu” [-14.46667, 16.30000] (Monard 1937a:40, 1938:93, 95); “Osi” [-15.08333, 15.41667] (Monard 1938:93). Cunene: “ruisseau Mbale” [-15.16667, 16.75000] (Monard 1938:93); “Kamba” [-16.28333, 15.23333] (Monard 1938:93). Cuando Cubango: “Kuandu” [-16.74487, 19.10136] (Monard 1937a:40, 1938:93, 95). Undetermined localities: without precise location (Boulenger 1882:165); “Nana Meya” (Boulenger 1905:110). Taxonomic and distributional notes: Hyperolius boeagei Steindacher, 1867 was described based on a specimen from “Angola”. However, the type locality of H. boeagei should be consid¬ ered “Duque de Braganga [= Calandula], Malanje,” as this is the locality from which the specimen collected by Pinheiro Bayao was sent by Bocage to Steidachner (Ceriaco et al. 2014a). Ferreira (1906) described Rappia seabrai based on a single individual deposited in Museu do Porto, col¬ lected by Francisco Newton. The holotype is believed to be lost (Ceriaco et al. 2014a) and several authors consider it to be a synonym of//, boeagei (Laurent, pers. comm, in Frost, 1985; Fretey et al., 2011; Ceriaco et al. 2014a; Frost 2016). In the original description, Ferreira (1906) also noted that one specimen referable to H. seabrai from “Duque de Braganga” [= Calandula] collected by Bayao was found in Museu Bocage, Lisboa, Portugal, and this was considered a paratype by Ceria¬ co et al. (2014a). The type locality of “Quilombo,” currently Gonguembo in Kwanza Norte 98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Province, is approximately 140 km west of Caladula Falls in Malanje Province, formerly “Duque de Branganga,” the type locality of H. bocagei and the locality from which the paratype of R. seabrai was collected (Ceriaco et al. 2014a). Schiotz (1999) and Schiotz and Van Daele (2003) suggested that H. bocagei might be a junior synonym of a species in the Hyperolius viridiflavus complex. Poynton (1986) considered Hyperolius cinctiventris Cope, 1862 to be a synonym of Hyperolius argus Peters, 1854, and as this species is restricted to East Africa it is unlikely to correspond to Angolan specimens. The specimens identified by Monard (1938) as cinctiventris deposited in the Musee d’Histoire Naturelle, La-Chaux-de Fond, Switzerland were recently stud¬ ied and identificated as Hyperolius bocagei (Ceriaco et al. in prep.). Hyperolius chelaensis Conradie, Branch, Measy and Tolley, 2012 Chela Mountain Reed Frog (Endemic) Hyperolius chelaensis Conradie et al. 2012a:5, figs 5-6. Holotype: PEM A9223 (eolleetor W. Conradie). Type loeality: “a small pateh (< 2.5ha.) of Afromontane forest in a small gorge draining from Serra da Chela above the Estaeao Zooteeniea, near Humpata, Lubango” (Conradie et al. 20 12a: 5), Angola. Hyperolius chelaensis: Conradie et al. (2013:203), Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 37): The species is known only from the type locality “Serra de Chela,” Lubango. Huila: “Serra de Chela gorge, above the Estagao Zooteeniea near Humapata, Lubango” [-14.88944, 13.27417] (Conradie et al. 20 12a: 5; Conradie et al. 2013:203). Taxonomic and distributional notes: This species was recently described by Con¬ radie et al. (2012a). According to the original publication, H. chelaensis is the sister species of H. cinereus Monard, 1937. The type locality is along a small stream that drains from the Serra de Chela escarpment into the ephemeral water courses (e.g., Giraul River) draining west to the Atlantic Ocean (Conradie et al. 2013). It is expected to be more widespread in the Serra de Chela and Leba mountain ranges. Further Map 37. Distribution of Hyperolius chelaensis in surveys are needed to establish a full distribu- tion of the species and its conservation status. Hyperolius cinereus Monard, 1937 Ashy Reed Frog (Endemic) Hyperolius cinereus Monard 1937a:32. Syntypes: MHNC 90.0856-7 (collector A. Monard). Type locality: “Kalukembe” and “Bimbi” (Monard 1937a:32), [= Caluquembe and Bimbe], Angola. Hyperolius cinereus: Monard (1938:85), Cei (1977:17), Frost (1985:209, 2016), Schiotz (1999:192), Chan- ning (2001:152), Conradie et al. (2012a:17), Conradie et al. (2013:222). Hyperolius {Hyperolius) cinereus: Fretey et al. (2011:30). Global conservation status (lUCN): Least Concern. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 99 Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 38): The species is known from southwestern and cen¬ tral Angola. Bie: “western dambo of Cacuchi River, Bie Province” [-13.58333, 16.86667] (Conradie et al. 2013:222); “Cubango basin (11)” [-13.69413, 17.06177] (Conradie et al. 2016:8-9, 16). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:32, 1938:85, Chan- ning 2001:153; Conradie et al. 2013:222); “small stream at Huambo Agriculture Institute, Huambo Province” [-12.71667, 15.80000] (Conradie et al. 2013:222); “1st stream cross¬ ing east of Caccuchi River, Bie Province” [-13.68333, 17.05000] (Conradie et al. 2013:222). Benguela: “Zamba-Seidlung bei Entre Rois, Benguella Province” [-13.01667, 14.63333] (Conradie et al. 2013:222). Huila: “Kalukembe” [-13.78333, 14.68333] (Monard 1937a:32, 1938:85; Frost 1985:209, 2016; Channing 2001:153; Conradie et al. 2013:201, 222); “stream under road before Estacao Zootecnica near Humpata, Lubango” [-14.90400, 13.32556] (Conradie et al. 2012a: 17; Conradie et al. 2013:222); “waterfall below dams, on plateau above Estacao Zootecnica near Humpata, Lubango” [-14.91425, 13.31386] (Conradie et al. 2012a: 17, Conradie et al. 2013:222). Cuando Cubango: “5 km south of Soba Matios Military base, Cuan- do” [-14.25706, 17.77861] (Conradie et al. 2013:222); “small stream after Muvange river, near Menongue” [-14.73333, 17.66667] (Conradie et al. 2013:222); “Muvange river crossing, near Menongue” [-14.81667, 17.66667] (Conradie et al. 2013:222); “Cubango basin (5)” [-14.74628, 17.66844] (Conradie et al. 2016:8-9, 16); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 16); “Cubango basin (7c)” [-14.43916, 17.81491] (Conradie et al. 2016:8-9, 16); “Cubango basin (8)” [-14.25705, 17.77852] (Conradie et al. 2016:8-9, 16); “Cuban¬ go basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:8-9, 16); “Cubango basin (22b)” [-14.66278, 16.96081] (Conradie et al. 2016:8-9, 16); “Cuito basin (27)” [-15.17127, 19.19433] (Conradie et al. 2016:8-9, 16); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9- 10, 16); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2016:9-10, 16); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9-10, 16); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9-10, 16); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie etal. 2016:9-10, 16); “Cubango basin (45)” [-16.88350, 18.01180] (Conradie etal. 2016:9, 12, 16). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Hyperolius cinereus Monard, 1937 was described based on two syntype specimens from “Kalukembe” in Huila Province and “Bimbi” in Huambo Province (Monard 1937a). Laurent (1964a) later assigned a small collection of specimens from “Dundo” in Lunda Norte Province to H. cinereus, althought his characterization of the coloration of this species differs from that of Monard. Both Monard (1937a) and Laurent (1964a) examined only preserved material and could only speculate on the coloration in life (Conradie et al. 2013). Conradie et al. (2013) considered that populations from southern and central Angola, including the localities for Monard’s two syntypes, correspond to H. cinereus. However, the northern populations 100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II identified by Laurent (1964a) correspond to a closely related species that they described as Hyper- olius raymondi Conradie, Branch and Tolley, 2013 (see H. raymondi). Both species occur in flood¬ ed grasslands called “dambos” where water levels reach 30 (northern population) to 50 (southern population) cm deep (Conradie et al. 2013). Monard (1937a) stated that the syntype from “Bimbi” was a juvenile and differed in several characters. Due to that, Conradie et al. (2013a) argued that this specimen should not be considered part of the type series, instead assigning it only provision¬ ally to H. cinereus, which was followed by Frost (2016). We recently studied the type material in the Musee d’Histoire Naturelle, La-Chaux-de Fond, Switzerland and believe that the “Bimbi” spec¬ imen is identifiable as Hyperolius cinereus, and as it was used by Monard in his description it should be recognized as a part of the type series (Ceriaco et al. in prep.). Hyperolius cinnamomeoventris Bocage, 1866 Cinnamon-Bellied Reed Frog Hyperolius cinnamomeo-ventris Bocage 1866a:55, 1866b:75. Holotype: MBL (collector F.A.P. Bayao), not localted by Ferret (1976a:24), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” (Bocage 1866a: 5 5), [= Calandula] Malanje Province, Angola. Rappia tristis Bocage 1866a:56, 1866b:76. Holotype: MBL (collector F.A.P. Bayao), not localted by Ferret (1976a:25), destroyed by fire 18 March 1978. Type locality: “Duque de Bragan^a” (Bocage 1866a:56), [= Calandula] Malanje Province, Angola. Rappia cinnamomeiventris: Bocage (1895a:172, 1897a:204), Ferreira (1906:163). Rappia cinnamomei-ventris: Bocage (1897b:211), Noble (1924:252, 253), Mertens (1938:429). Hyperolius tristis: Bocage (1895a:171, 1897a:204), Boulenger (1882:121), Ferret (1976a:25). Rappia bivittata: Ferreira (1906:161). Hyperolius einnamomeoventris: Parker (1936:144), Laurent (1950a: 16, 1954a:78, 1964a: 149), Cei (1977:17), Ferret (1976a:25), Frost (1985:209, 2016), Schiotz (1999:129), Channing (2001:153), Conradie et al. (2013:206), Ceriaco et al. (2014a: 19), Ceriaco et al. (2016b:37). Hyperolius einnamomeoventris einnamomeoventris: Laurent (1961:79), Schiotz (1975:123). Hyperolius {Hyperolius) einnamomeoventris: Fretey et al. (2011:30). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from central and southern Cameroon south to Angola, extending across the Democratic Republic of Congo to northwestern Zambia, and east to Uganda and western Kenya. Ocurrences in Angola (Map 39): The species is found in western areas along the coast as well as northeastern regions of the country. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:78); “environs de Dundo, foret des sources de la Dundundo” [-7.36667, 20.83333] (Laurent 1964a: 149); “Matala, rive gauche de la Tshihumbwe, 40 km a Lest de Dundo” [-7.43333, 21.16667] (Lau¬ rent 1950a: 16); “Andrada (Luembe O)” [-7.68729, 21.37003] (Laurent 1954a:78); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a; 16); “riv. Kakuje, affl. gauche Luembe pres du village du “soba”, Santana” [-8.56667, 20.56667] (Laurent 1954a:78). Malanje: “Duque de Braganqa” [-9.10000, Map 39. Distribution of Hyperolius cinnamomeoventris in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA lOI 15.95000] (Bocage I866a:55, 56, 1866b:76, I895a:I7I, 172, I897a:204; Laurent 1961:79; Schiotz 1975:123, Perret I976a:25; Frost 1985:209, 2016); “Kalandula” [-9.10000, 15.95000] (Conradie et al. 2013:206), “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:37). Kwan¬ za Norte: “N’golla Bumba” [-9.03333, 14.60000] (Ferreira 1906:161; Ceriaco et al. 2014a: 19); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:163); “Quilombo” [-9.33333, 14.90000] (Fer¬ reira 1906:163). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Ferreira 1906:163); “Con- gulu” [-10.86667, 14.28333] (Parker 1936:144). Benguela: “Hanha” [-13.30000, 14.20000] (Bocage 1897a:204, 1897b:211; Perret 1976a:25); “Cubal” [-13.03333, 14.25000] (Mertens 1938:429). Undetermined Locality: “Mupepe” (Ferreira 1906:163); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Bocage (1866b) described two new species, Hyperolius cinnamomeo-ventris Bocage, 1866 and Rappia tristis (Bocage, 1866) from “Duque de Braganga [= Calandula].” The nomen tristis was synonymized with H. cinnamomeoventris by Laurent (1943b, 1947) and has subsequently been followed by other authors including Perret (1976a). Before the Lisbon fire Perret (1976a) visited the Museu Bocage but did not find the holotype of either of these species. One syntype of Rappia bivittata designated by Ferreira (1906) from “N’Golla Bumba,” Kwanza Sul province seems to represent Hyperolius cinnamomeoventris Bocage, 1 866 (Ceriaco et al. 2014a). Currently this species is accepted and recognized as Hyperolius cinnamomeoventris (Schiotz 1999; Channing 2001; Fretey et al. 2011; Conradie et al. 2013; Frost 2016). This wide¬ spread species represents a complex of species found in Central Africa and several offshore islands and is awaiting taxonomic revision. Hyperolius concolor (Hallo well, 1844) Variable Reed Frog Ixalus concolor Hallowell 1844:60. Holotype: ANSP 3216 (Malnate 1971:350) (collector unknown). Type locality: “Liberia, W. Africa” (Hallowell 1844:60), Liberia. Hyperolius modestus: Bocage (1866a:55, 1866b:74). Rappia concolor. Bocage (1895a:173), (1897b:211). Hyperolius eoneolor Noble (1924:252), Schiotz (1999:104), Amiet (2012:264), Frost (2016). Cameroon, and along the Atlantic coast south to Angola. Ocurrences in Angola (Map 40): The species is recorded mostly from western Ango¬ la. Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:55, 1866b:74). Bie: “Bihe” [-12.38333, 16.95000] (Bocage 1895a:173). Benguela: “Hanha” [-13.30000, 14.20000] (Bocage 1897b:211). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:173); “Huila” [-15.05000, 13.55000] (Bocage 1895a: 173). Undetermined Locality: “Rio Quando” (Bocage 1895a: 173). Taxonomic and distributional notes: The Angolan records of Hyperolius concolor (Hallowell, 1844) probably represent a Map 40. Distribution of Hyperolius concolor in Angola. Global conservation status (lUCN): Least Concern. Global distribution: The species is irE known from eastern Sierra Leone to western I6“E 20*E 102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II misidentification, although Fretey et al. (2011) considered the species present in Angola. Unfortu¬ nately, the specimens used by Bocage (1866a, 1866b, 1895a, 1897b) were destroyed in the 1978 fire at the Museu Bocage, Lisboa. Noble (1924) cited one specimen of//, concolor from Angola at the American Museum of Natural History, although he did not provide more precise locality infor¬ mation. Hyperolius dartevellei Laurent, 1943 Dartevelle’s Reed Frog Hyperolius dartevellei Laurent 1943:71, fig. 3. Holotype: MRAC 38385 (collector E. Dartevelle). Type local¬ ity: “Zambi, Bas-Congo”, Democratic Republic of Congo. Hyperolius nasutus adspersus: Laurent (1961:92). Hyperolius dartevellei'. Channing et al. (2002:96), Amiet (2015:275), Channing et al. (2013:319), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Cameroon to the lower Congo Basin, northern Angola and the highlands of northwestern Zambia. Ocurrences in Angola: The species is known from the most northern parts of Angola as plot¬ ted by Channing (2001). Those occurrences are not mapped here. Taxonomic and distributional notes: Hyperolius dartevellei Laurent, 1943 was recently removed by Channing et al. (2013) from the synonymy of Hyperolius adspersus Peters, 1877, in which it had been placed by Laurent (1961). Channing et al. (2002) and Amiet (2005) considered it a member of nasutus complex. The species is presently only confirmed from northern Angola, including the Cabinda enclave (Channing et al. 2013). Hyperolius fuscigula Bocage, 1866 Brown-Throated Reed Frog (Endemic) Hyperolius fuseigula Bocage 1866a:56, 1866b:76. Syntypes: MBL 19-189 [2 specimens] (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Duque de Bragan^a” (Bocage 1866a:56), [= Calandula], Angola. Rappia fuseigula'. Bocage (1895a: 170, 1897a:204). Hyperolius fuseigula'. Noble (1924:253), Perret (1976a:26), Frost (2016). Hyperolius einnamomeoventris'. Laurent (1943:67). Hyperolius {Hyperolius) fuseigula'. Fretey et al. (2011:30). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 41): The species is known only from the type locality in Malanje. Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a:56, 1866b:76, 1895a:170, 1897a:204; Perret 1976a:26). Taxonomic and distributional notes: Laurent (1943b) tentatively considered //yper- olius fuscigula Bocage, 1866 to be a synonym of Hyperolius einnamomeoventris Bocage. 1866. Perret (1976a) noted the poor condition Map 4 1. Distribution of in Angola. n°E lU'E 20^ E 24“E MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 103 of the type specimens that were later destroyed in the 1978 fire. Because of the lack of records and loss of the type material, the validity of this taxon is uncertain. Hyperolius glandicolor Peters, 1878 Peters’ Reed Frog Hyperolius glandicolor Peters 1878:208, pi. 2, fig. 9. Syntypes: ZMB 9299 [2 speeimens] (eolleetor J.M. Hildebrandt). Type loeality: “Taita”, Kenya. Hyperolius glandicolor. Sehietz (1999:212), Wieezorek et al. (2000: 1333), Channing and Howell (2006: 162), Conradie et al. (2012a:6). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Kenya, southern Somalia, and Tanzania, Angola and adjacent Democratic Republic of Congo and northern Mozambique. Ocurrences in Angola (Map 42): The only Angolan record is from the northwest of the country. Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1866a:56). Taxonomic and distributional notes: This species was removed from the synonymy of Hyperolius marmoratus Rapp, 1842 by Wieezorek et al. (2000), and Schiotz (1999) has argued it is a member of the viridiflavus sub¬ group in the viridiflavus complex. The recog¬ nized distribution of this species is limited to the eastern coast of equatorial Africa and the associated interior, where it is often associated with higher elevations (Wiezezorek et al. 2000). The Angolan record from “Duque de Braganga” [= Calandula] likely should be referred to another species of Hyperolius. Map 42. Distribution of Hyperolius glandicolor in Angola. Hyperolius gularis Ahl, 1931 Luanda Reed Frog (Endemic) Hyperolius gularis Ah\ 1931b:408, fig. 281. Holotype: ZMB uncatalogued (collector K. May), probably lost. Type locality: “Loanda” [= Luanda] Luanda Province, Angola. Hyperolius gularis: Frost (1985:211, 2014). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 43): The species is known only from the type locality “Loanda”. Luanda: “Loanda” [-8.83333, 13.26667] (Ahl 1931a:125; Frost 1985:211, 2016). Taxonomic and distributional notes: The taxonomic validity of this species is doubtful and it has not been discussed in recent synopses of Hyperolius by Schiotz (1999), Channing (2001), or Fretey et al. (2011). However, Laurent {in Frost 1985; Frost 2016) suggested that this species is probably a synonym of Hyperolius marmoratus Rapp, 1842, which we recognize above as Hyper¬ olius angolensis Steindachner, 1867 (see H. angolensis account). 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ifi'E 20°E 24*E Map 43 . Distribution of Hyperolius gularis in Angola. Hyperolius kivuensis Ahl, 1931 Kivu Reed Frog Hyperolius kivuensis Ahl 1931b:280., fig. 151. Holotype: ZMB 36098 (collector Kandt). Type locality: “Kivu- See” (Ahl 1931a:280), [= Lake ICivu], Democratic Republic of Congo. Hyperolius multifaseiatus: Monard (1937a:33, 1938:87). Hyperolius kivuensis kivuensis: Laurent (1950:16, 1954a:78, 1964a: 149), Poynton and Broadley (1976:196). Hyperolius quinquevittatus: Laurent (1950a: 16, 1954a:79). Hyperolius kivuensis: Cei (1977:17), Schiotz (1999:163), Channing (2001:157), Frost (2016). Hyperolius kivuensis multifaseiatus: Pickersgill (2007a:325). Hyperolius {Hyperolius) kivuensis: Fretey et al. (2011:31). Global conservation status (lUCN): Least Concern. Global distribution: The species is widespread in savannas of southwestern Ethiopia, south¬ eastern Sudan, western Kenya, Tanzania, Uganda, Rwanda, and Burundi, eastern and southern Democratic Rebublic of Congo, Angola, Zambia, Malawi, and northwestern Mozambique. Ocurrences in Angola (Map 44): The species is well documented from northeastern Angola, though may also occur in southern regions. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:16, 1954a:78-79); “Matala, rive gauche de la Tshihumbwe, 40 km a Lest de Dundo” [-7.43333, 21.16667] (Lau¬ rent 1950a: 16); “Andrada (Luembe O)” [-7.70000, 21.38333] (Laurent 1954a:78); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 16, 1954a:78); “Kossa” [-7.90000, 21.36667] (Laurent 1950:16). Lunda Sul: “Alto Chicapa” [-10.88333, 12*E I6®E 20°E 24‘E MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 105 19.23333] (Laurent I964a:I49). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:87). Undetermined Locality: “with no precise identification” (Laurent 1954a: 78), “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). Taxonomic and distributional notes: Poynton and Broadley (1987) recognized that Hyper- olius kivuensis Ah\, 1931 is easily confused with quinquevittatusBocsLgQ, 1866, though they stated that the latter is generally smaller and slimmer. Whereas Laurent (1954a) treated H. multifasciatus (Ahl, 1931) as a junior synonym of//, quinquevittatus, Pickersgill (2007a) con¬ sidered it to be a subspecies of Hyperolius kivuensis. Schiotz (1999) and Channing (2001) provid¬ ed some additional records, though without information about the source. Hyperolius langi Noble, 1924 Lang’s Reed Frog Hyperolius langi Noble 1924:266, pi. 39, fig. 1. Holotype: AMNH 9983 (collector Mr. Lang). Type locality: “Niapu” Democratic Republic of Congo. Hyperolius langi: Monard (1937a:40, 1938:95). Hyperolius platyeephalus langi: Cei (1977:17). Hyperolius langi: Schiotz (1999:149), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola, the northeastern Democratic Repub¬ lic of Congo and Uganda. Ocurrences in Angola (Map 45): The species has been recorded from Benguela Province. Benguela: “Ebanga” [-12.73333, 14.73333] (Monard 1937a:40, 1938:95). Undetermined Locality: “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Monard (1937a, 1938) reported H. langi from “Ebanga” in southwestern Angola. There are also two other species of Hyperolius that occur at this location, H. benguellensis (Bocage, 1893) and H. bocagei Steindachner, 1867. The species that Monard (1937a, 1938) identi¬ fied as H. langi probably corresponds to H. bocagei, especially as it does not agree with the original description for H. benguellensis. However, examination of the type material of H. langi is required to confirm this. Distribution of Hyperolius langi in Angola. Hyperolius maestus Rochebrune, 1885 Cabinda Reed Frog (Endemic) Hyperolius Maestus Rochcbmnc 1885:91. Type: Presumably in MNHN (collector unknown), unrecognized as a type, if still extant, formerly in Museo Bouvieri. Type locality: “Locum Landana dictum, in foliis Musamm” [= Landana, Cabinda enclave], Angola. Hyperolius maestus: Frost (1985:212, 2014). Hyperolius {Hyperolius) marmoratus: Fretey et al. (2011:32). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from Angola. 12*E Ifi'E 20°E 2-t*E 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 46): The species is only known from the type locality “Landana” in the Cabinda enclave. Cabinda: “Landana” [-5.21667, 12.15000] (Rochebrune 1885:91; Frost 1985:212, 2016). Taxonomic and distributional notes: The species was described by Rochenbrune (1885) from “Locum Landana dictum, in foliis Musarum” in the Cabinda enclave. Laurent {in Frost 1985), and latter Fretey et al. (2011) con¬ sidered Hyperolius maestus Rochebrune, 1885 to be a possible synonym of Hyperolius marmoratus Rapp, 1842. The type specimen was deposited in Museo Bouvier and is now presumably in Museum national d’Histoire naturelle, Paris and unrecognized as a type if it still exists (Frost 2016). The species is not mentioned in Schiotz (1999) or Channing (2001). Hyperolius nasutus Gunther, 1865 Large-Nosed Long Reed Frog Hyperolius nasutus Gunther 1865a:482, PI XXXIII, fig. 3. Holotype: BMNH 1947.2.9.68 (collector J.F. Bayao). Type locality: “Duque de Bragan^a” [= Calandula], Malanje Province, Angola. Rappia punctulata Bocage 1895a: 168. Holotype: MBL T.23-227 (collector Banyures) destroyed by fire 18 March 1978. Type locality: “sur les bords du Quanza” [= Quanza river], Angola. Hyperolius nasutus'. Bocage (1866a:55), Noble (1924:253), Schmidt (1936:132), Loveridge (1936a: 109, 1953a:362, 1953:362, 1957:333), Monard (1937a:39, 1938:94), Mertens (1938:429), Frade (1963:254), Haacke (1970:279), Schiotz (1975:97), Poynton and Broadley (1976:206), Cei (1977:17, 18), Frost (1985:214, 2016), Poynton and Haacke (1993:14), Schiotz (1999:97, 2006:65), Channing (2001:166), Largen (2001:359), Channing et al. (2002:96), Schiotz and Van Daele (2003:138), Amiet (2005:275f Conradie et al. (2012a:3), Ceriaco et al. (2016b:35), Conradie et al. (2016:11). Rappia nasuta: Boulenger (1882:127), Bocage (1895a: 169, 1897a:204). Rappia punetulata: Bocage (1897a:204), Ferreira (1904:112), Amiet (2005:275). Hyperolius punetulatus: Noble (1924:253), Loveridge (1936a: 107, 1936b:405), Mertens (1937:20), Monard (1938:86), Frade (1963:254), Perret (1976a:27), Frost (1985:216). Hyperolius nasutus nasutus'. Laurent (1950a: 17, 1954a:84, 1964a: 154), Hellmich (1957a:29). Hyperolius {Hyperolius) nasutus'. Fretey et al. (2011:32). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola south to the Okavango Delta of Botswana and adjacent northern Namibia. Ocurrences in Angola (Map 47): The species is known from its type locality “Duque de Bra- ganqa, Malanje Province” and is very widespread across almost the whole of the country. Bengo: “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:112). Kwanza Norte: “3 km W of Salazar” [-9.30000, 14.91667] (Poynton and Haacke 1993:14); “Dondo” [-9.68333, 14.43333] (Loveridge 1936a:107). Kwanza Sul: “Namba” [-11.91667, 14.85000] (Poynton and Haacke 1993:14). Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Gunther 1865a:482; Bocage 1866a:55, 1895a:169, 1897a:204; Boulenger 1882:127; Loveridge 1936a:109, 1936b:405, 1953a:362, 1957:333; Schiotz 1975:97; Frost 1985:214, 2016; Amiet 2005:275); “Calandula” [-9.07917, MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 107 15.79583] (Channing et al. 2013:334); “Kan- gandala” [-9.825II, 16.91225] (Channing etal. 2013:334); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco etal. 2016b:35); Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a: 84); “environs de Dundo, rives de la Luaehimo” [-7.38333, 20.85000] (Lau¬ rent 1950a: 17); “Carumbo lagoon” [-7.74422, 19.95467] (Conradie et al. 2012a:3); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 17); “Kossa a 100 km au sud-est de Dundo” [-7.90000, 21.36667] (Laurent 1950a: 17); “rive de la Tehihumbwe 40 km a Test de Dundo” [-8.01667, 19.31667] (Laurent 1950a:17); “Village Capaia” [-8.33847, 20.24250] (Conradie et al. 2012a:3). Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:39, 1938:94); “Alto Chieapa,sourees du Cuilo” [-10.88333, 19.23333] (Laurent 1964a:154); “Alto Cuilo, rives du Cuilo “ [-10.01667, 19.55000] (Laurent 1964a:154). Mexico: “region du lae Calundo” [-11.80000, 20.86667] (Laurent 1964a:154); “reserve de ehasse de Cameia” [-11.83333, 21.00000] (Laurent 1964a:154). Bie: “Chi- tau” [-11.43333, 17.15000] (Sehmidt 1936:132); “General Maehado” [-12.01667, 17.06667] Mer- tens (1937:20). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:39, 1938:94); “Tongru- be am Jamba Fliipehen” [-13.60000, 16.60000] (Hellmieh 1957a:29). Benguela: “Cubal” [-13.03333, 14.25000] (Mertens 1938a:429); “Mareo de Canavezes (Cubal da Ganda)” [-13.08333, 14.33333] (Laurent 1964a:154). Hufla: “Huilla” [-15.05000, 13.55000] (Boeage 1895a:169, 1897a:204); “Caeonda” [-13.73333, 15.06667] (Boeage 1895a:169, 1897a:204); “Kalukembe” [-15.05000, 13.55000] (Monard 1938:86); “Nunteehite lagoon” [-15.13333, 13.41667] (Poynton andHaaeke 1993:14). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937a:39, 1938:94). Cuando Cubango: “Kuandu” [-16.74487, 19.10136] (Monard 1937a:39, 1938:94). Undeter¬ mined Locality: “Margens do Cuanza” (Boeage 1895a: 168, 1897a:204; Loveridge 1936a: 107, 1957:333; Perret 1976a:27; Frost 1985:216; Amiet 2005:275); “without preeise loeation” (Monard 1937a:39, 1938:94); “Western subregion and Angolan highlands (Angolan eoastal or watersheds)” (Frade 1963:254); “Zambezian highlands” (Frade 1963:254); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17); “arid territories along the eoast” (Cei 1977:18). Taxonomic and distributional notes: While the type speeimen of R. punctulata is now lost, Channing et al. (2013) ineluded it as a synonym of H. nasutus, for whieh there are many junior synonyms. To date, fifteen names have been used for members of the nasutus group, most of whieh have been synonymized with the nominotypieal form (Sehiotz 1999); a relatively reeent list of these synonyms is provided by Amiet (2005). Poynton and Broadley (1987) reeognized three speeies in the southern Afriean savanna, Hyperolius viridis Sehiotz, 1975, Hyperolius nasutus Gunther, 1865 and Hyperolius benguellensis Boeage, 1893. Sehiotz (1999) ineluding//, benguel- lensis as a synonym of H. nasutus after arguing that differenees in morphology and eolor pattern are not suffieient for distinguishing these two speeies. Later, Channing et al. (2002) retained H. benguellensis as a synonym of nasutus, but also divided the group into three eryptie speeies differing in advertisement eall: Hyperolius acuticeps Ahl, 1931, Hyperolius viridis Sehiotz, 1975, 1,2°E 16°E 20°E H°E Map 47. Distribution of Hyperolius nasutus in Angola. 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II and H. nasutus (Schiotz and Van Daele 2003; Schiotz 2006). Schiotz (2006) distinguished two speeies, H. benguellensis and H. nasutus, in the nasutus group for Angola based on the strueture of the advertisement call and color pattern. This widespread species is common in savannas and exhibits a preference for humid areas such as reeds growing in deep water, pools, streams, and large rivers (Schiotz 1999; Channing 2001). HyperoUus nitidulus Peters, 1875 Plain Reed Frog Hyperolius nitidulus Peters 1875:209, pi. 3, figs. 4, 4a. Holotype: ZMB 7729 fide Bauer et al. (1995:44) (eol- leetor R. Buehholz). Type loeality: “Yomba (Lagos)” (Peters 1875:209), Nigeria. Hyperolius nitidulus: Peters (1877a:619), Sehiotz (1999:206), Wieezorek et al. (2000:1238-1241), Rodel et al. (2010:178), Amiet (2012:194), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from West African extending from Guinea east into Nigeria, Angola and Cameroon. Ocurrences in Angola (Map 48): The species was cited from “Chinchoxo” in the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:619). Taxonomic and distributional notes: According to Wieezorek et al. (2000), the species Hyperolius nitidulus Peters, 1875 is a West African species associated with savanna and woodlands (Schiotz 1999; Wieezorek et al. 2000; Rodel et al. 2010; Amiet 2012). Based on the distribution of the species delimited by Wieezorek et al. (2000), the record from north¬ western Angola is likely a misidentification. Hyperolius ocellatus (Gunther, 1858) Hyperolius oeellatus Gunther 1858a:326. Lectotype: BMNH 1947.2.9.22, (collector L. Fraser), formerly BMNH 51.11.13.48, by implication of being referred to as “holotype” by Perret (1975a:27), Syntypes: BMNH [2 specimens] (collector Mr. Fraser, for the “Fernando Po” specimen) (Gunther 1858a:326). Type locality: “Fernando Po” [= Bioko, Guinea] fide Perret (1975a:24), who considered “Angola” an error. Originally given as “Fernando Po” and “Angola” by Gunther (1858a:326). Hyperolius Lueani Rochebmne 1885:91. Type: Presumably in MNHP (collector unknown), uncatalogued as a type, if still extant, formerly in Museo Bouvieri. Type locality: “Locam Landana dictum, in frondibus Borassomm” [= Landana, Cabinda enclave] Cabinda enclave, Angola. Hyperolius oeellatus: Gunther CT858b” 1859:88), Perret (1975a:24), Cei (1977:17), Frost (1985:214, 2016), Schiotz (1999:141), Channing (2001:168), Channing et al. (2012:217). Rappia oeellata: Boulenger (1882:123); Bocage (1895a: 165). Hyperolius lueani: Frost (1985:212, 2016). Hyperolius {Hyperolius) oeellatus: Fretey et al. (2011:32). Global conservation status (lUCN): Least Concern. Global distribution: The species is from tropical Central Africa. It occurs in the Cabinda irE I6®E 20°E :4»E Map 48. Distribution of Hyperolius nitidulus in Angola. Golden-Eyed Reed Frog MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 109 enclave in Angola, extending north into south¬ eastern Nigeria and Bioko Island, east across Democratic Republic of Congo, with possibly occurences in western Uganda and Rwanda. Ocurrences in Angola (Map 49): The species is recorded from northwestern Angola in the Cabinda encalve. Cabinda: “Landana” [-5.21667, 12.15000] (Rochebrune 1885:91; Frost 1985:212, 2016); “Cabinda” (Schiotz 1999:141; Channing 2001:169). Taxonomic and distributional notes: Gunther (1858a) described Hyperolius ocella- tus from “Fernando P6” and “Angola,” the lat¬ ter without clear indication of the locality based on two specimens deposited in British Museum. Perret (1975a) considered the type locality “Angola” to be in error and designated the lectotype as being from “Fernando P6.” Following Gunther’s (1858a) description, Rochebrune (1885) described a new species from “Locam Landana dictum, in frondibus Borassorum” in the Cabinda enclave as Hyperolius lucani. This may be a synonym of Hyperolius ocellatus (Laurent in Frost 1985, 2016; Fretey et al. 2011), although it is not mentioned by Schiotz (1999) or Channing (2001). Hyperolius platyceps (Boulenger, 1900) Benito River Reed Frog Rappia platyceps Boulenger 1900:444 , pi. 27, fig. 4. Syntypes: BMNH 1947.2.9.57-58 and BMNH 1900.2.17.89-90 [2 speeimens] (eolleetor G.L. Bates, formerly BMNH 1900.2.17.89-90). Type loeality: “Benito River, north of the Gaboon River between 20 and 30 miles inland from the eoast, Gaboon” [= Ben¬ ito River], Equatorial Guinea. Rappia bivittata Ferreira 1906:161, pi. 1. Syntypes: MHNFCP 229 017291, 017296 and 017302, [3 speei¬ mens] (eolleetor F. Newton). Type loeality: “N’Golla Bumba”, “Quilombo” [= Gonguembo] and “Rio Luinha,” Kwanza Norte Provinee, Angola. Rappia platyceps var. angolensis Ferreira 1906:161, pi. 1. Syntype: MHNFCP 017303 (Quilombo) (eolleetor F. Newton), the other syntype from “N’golla Bumba” was not loeated. Type loeality: “N’Golla Bumba” and “Quilombo” [= Gonguembo], Kwanza Norte Provinee, Angola. Rappia fasciata Ferreira 1906:164, pi. 1. Holotype: MHNFCP 017294 (eolleetor F. Newton). Type loeality: “Quilombo” [= Gonguembo] Kwanza Norte Provinee, Angola. Hyperolius angolanus Ahl 1931b:271. Replaeement name for Rappia platyceps var. angolensis. Rappia platyceps: Ferreira (1906:161). Hyperolius platyceps: Noble (1924:253), Ceriaeo et al. (2014a:24), Frost (2016). Hyperolius angolanus Laurent (1950a: 15), Frost (1985:207). Hyperolius platyceps angolanus: Laurent (1954a:77), Cei (1977:17). Hyperolius fasciatus: Noble (1924:252), Cei (1977:17), Frost (1985:210). Hyperolius ferreirai: Frost (1985:210). Hyperolius {Hyperolius) platyceps: Fretey et al. (2011:33). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Cameroon to the southwestern Central African Republic, south to the Democratic Republic of Congo and northwestern Angola. Ocurrences in Angola (Map 50): The species occurs in the north of the country. Lunda irE I6®E 20°E :4'>E Map 49. Distribution oi Hyperolius ocellatus in Angola. 110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Norte: “Dundo” [-7.53333, 21.08333] (Lau¬ rent 1950a: 15); “Luachimo” [-7.36667, 20.83333] (Laurent 1954a:77). Kwanza Norte: “N’golla Bumba” [-9.03333, 14.60000] (Ferreira 1906:161; Frost 1985:207, 210; Ceriaco et al. 2014a:23); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:161; Frost 1985:210); “Quilombo” [-9.33333, 14.90000] (Ferreira 1906:161, 164; Frost 1985:210, 2016; Ceriaeo et al. 2014a:22). Taxonomic and distributional notes: Ahl (1931a) elevated Rappia platyceps var. angolensis Ferreira, 1906. to full species and proposed the replacement name Hyperolius angolanus because of the the name was preoc¬ cupied by Hyperolius marmoratus var. angolensis Steindachner, 1862 (Ceriaco et al. 1 -1 1 Map 50. Distribution of in Angola. 2014a). Ferreira (1906) also described as new Rappia fasciata from “Quilombo” and Rappia bivitatta from “N’golla Bumba”, “Quilombo,” and “Rio Luinha.” Recently Fretey et al. (2011) and Ceriaco et al. (2014a) rQcognizQd Hyperolius platy¬ ceps angolensis, Rappia fasciata, and Rappia bivittata as synonyms of H. platyceps. 12°E I6'E 20°E 24'*E Hyperolius polli Laurent, 1943 Tshimbulu Reed Frog Hyperolius polli Laurent 1943b:96, fig. 20. Holotype: MRAC 656 (collector Achten). Type locality: “Tshim¬ bulu s/Luebi (Kasai)” Democratic Republic of Congo. Hyperolius polli'. Laurent (1954a:78). Hyperolius polli'. Schiotz (1999:153), Frost (2016) Hyperolius {Hyperolius) polli'. Fretey et al. (2011:33). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from the Democratic Republic of Congo and adjacent Angola. Ocurrences in Angola (Map 51): The single species record is in Lunda Norte Province near the border with Congo. Lunda Norte: “Tshinguvu, Riv. Tshikapa” [-7.61667, 20.50000] (Laurent 1954a:78). Taxonomic and distributional notes: According to Laurent (1954a), the specimen from “Tshinguvu (Tshikapa)” is similar to Hyperolius cinnamomeoventris Bocage, 1866, but differs in some morphological characters. Currently it remains poorly known and its validity is uncertain. 12*E l(i*E 2I)°E 24*E Map 5 1 . Distribution of Hyperolius polli in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA III Hyperolius protchei Rochebrune, 1885 Rochebrune’s Reed Frog (Endemic) Hyperolius Protchei Rochebrune 1885:92. Type: Presumably in MNHP (eolleetor unknown), uneatalogued as a type, if still extant, formerly in Museo Bouvieri. Not eited by Guibe (1950). Type loeality: “Loeum Lan- dana dietum, in foliis Musarum” [= Landana, Cabinda enelave], Angola. Hyperolius protehei: Frost (1985:215, 2016). Hyperolius {Hyperolius) marmoratus: Fretey et al. (2011:32). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 52): The species is only known from the type locality “Landana” in the Cabinda enclave. Cabinda: “Landana” [-5.21667, 12.15000] (Rochebrune 1885:92; Frost 1985:215, 2016). Taxonomic and distributional notes: Laurent {in Frost 1985) considered Hyperolius protchei as a possible synonym of Hyperolius marmoratus Rapp, 1842, which in Angola should be considered Hyperolius angolensis Steindachner, 1867 (see Hyperolius angolensis account). The species is not mentioned by Schiotz (1999) or Channing (2001). ^2. Distribution oi Hyperolius protchei in Angola. Hyperolius pusillus (Cope, 1862) Water Lilly Reed Frog Global conservation status (lUCN): Least Concern. Global distribution: The species is rec- tricted to the eastern Africa coastal lowlands, from Somalia to KwaZulu-Natal. Ocurrences in Angola (Map 53): Speci¬ men records are from Malanje and Benguela Province. Malanje: “Fort Don Carlos” [-7.65000, 17.30000]; (Boulenger 1905:110); “Marimba” [-8.36667, 17.03333] (Boulenger 1905:110); “Bange N’gola” [-8.43333, 16.56667] (Boulenger 1905:110); “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:55, 1866b:75, 1895a:173, 1897a:204). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a: 173, 1897a:204). Undetermi- Crumenifera pusilla Cope 1862:343. Holotype: ANSP 11323 (collector unknown) fide Malnate (1971:352). Type locality: “Umvoti,” KwaZulu-Natal, South Africa. Hyperolius mierops: Bocage (1866a:55, 1866b:75), Frade (1963:254), Noble (1924:253). Rappia mierops'. Bocage (1895a: 173, 1897a:204), Boulenger (1882:127; 1905:110). Hyperolius pusillus'. Cei (1977:17), Schiotz (1999:185), Channing (2001:176). Map 53. Distribution of Hyperolius pusillus in Angola. 112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II ned Locality: “bords du Quanza” (Bocage 1895a: 173, 1897:204); “Rio Quando” (Bocage 1895a:173, 1897a:204); “Zambezian highlands” (Frade 1963:254); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Boulenger (1882) noted that this is an East African species, although he recognized (Bocage 1866b) Hyperolius microps as a valid name. Based on this information, the Angolan records possibly belong to another Hyperolius species and it would be important to review this case. Unfortunately, the specimens from Museu Bocage (Bocage 1 866b, 1895a, 1897a) were destroyed in the 1978 fire, although the specimens studied by Boulenger (1905) still exist in the British Museum. Noble (1924) also cited specimens of H. pusillus and H. microps in the collections of the American Museum of Natural History from Angola, though without precise locality information. Hyperolius quinquevittatus Bocage, 1866 Five-Striped Reed Frog Hyperolius quinquevittatus Bocage 1866a:56. Syntypes: MBL 24-214, 24-215 [2 specimens] (collector F.A.P. Bayao). Type locality: “Duque de Braganga” [= Calandula], Malanje Province, Angola. Hyperolius quinquevittatus: Bocage (1866h:77), Frade (1963:254), Noble (1924:253), Laurent (1950A:16, 1954a:79), Perret (1976a:25), Cei (1977:17), Frost (1985:216, 2016), Schiotz (1999:101), Channing (2001:178), Channing and Howell (2006:180). Rappia quinquevittata: Bocage (1895a:174). Hyperolius quinquevittatus quinquevittatus: Schiotz (1975:102), Poynton and Broadley (1987:197), Pickers- gill (2007a:360). Hyperolius {Hyperolius) quinquevittatus: Fretey et al. (2011:33). According to Perret (1976a), the syntypes described by Bocage (1866a), which were destroyed in the 1978 fire in Lisbon, were stud¬ ied by Schiotz (1975) when he concluded that H. quinquevittatus is similar in morphology to H. nasutus Gunther, 1865. Poynton and Broadley (1987) noted, however, that Hyper¬ olius quinquevittatus is more similar to Hyper¬ olius kivuensis even if H. quinquevittatus is in general much smaller and slimmer. Pickersgill Global conservation status (lUCN): Least Concern. Global distribution: The species is known from open savannas at higher elevations in north¬ ern Angola, the southern Democratic Republic of Congo, southern Tanzania, and northern Zambia and Malawi, and Mt. Namuli in northern Mozambique. Ocurrences in Angola (Map 54): The species occurs especially in northern Angola. Malan¬ je: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a:56, 1866b:77, 1895a:174; Schiotz 1975:102; Perret 1976a:25; Frost 1985:216, 2016; Poynton and Broadley 1987:197; Pickersgill 2007a:360). Undeter¬ mined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Map 54. Distribution of Hyperolius quinquevittatus in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA II3 (2007a) suspected that the species belongs to the multifasciatus group, while Fretey et al. (2011) included the H. multifasciatus in the synonym of quinquevittatus. Hyperolius raymondi Conradie, Branch and Tolley, 2013 Raymond’s Reed Frog (Endemic) Hyperolius raymondi Conradie et al. 2013:203, fig. 7. Holotype: PEM A10049 (eollectors W. Conradie, W. Branch, P. Vaz Pinto, S. Batista and N. Batista). Type locality: “Lagoa Cammbo from a dambo near the expedition base camp,” Lunda Norte Province, Angola. Hyperolius cinereus: Laurent (1964a: 149), Schiotz (1999:192), Channing (2001:152). Hyperolius {Hyperolius) einereus: Fretey et al. (2011:30). Hyperolius raymondi'. Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 55): The species is known from the type locality “Lagoa Cammbo” and from surrounding localities in Lunda Norte Province. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a: 149; Channing 2001:153; Conradie et al. 2013:203); “margins Lagoa Samokaza east of Lagoa Cammbo” [-7.73833, 19.98694] (Conradie et al. 2013:203); “Lagoa Cammbo from a dambo near the expedition base camp” [-7.74422, 19.95522] (Conradie et al. 2013:203; Frost 2016); “headwaters of Lovua, north of village Capaia” [-8.33847, 20.24250] (Conradie et al. 2013:203). Taxonomic and distributional notes: Populations that Laurent (1964a) referred to H. cinereus were recognized by Conradie et al. (2013a) as a distinct species and described as Map 55. Distribution of Hyperolius raymondi in Angola. H. raymondi (see further discussion above in account for H. cinereus). This newly described species is so far endemic to Angola, but may also occur in southernmost Democratic Republic of Congo and northwest Zambia. It is found in grass-covered “dambo” (water levels less than 30 cm) in the flood plains of the Lulele River, a southern tributary draining north to the larger Congo River (Conradie et al. 2013). Hyperolius rhizophilus Rochebrune, 1885 African Reed Frog (Endemic) Hyperolius Rhizophilus Rochebmne 1885:92. Type: Presumably in MNHN (eolleetor unknown), uneata- logued as a type, if still extant, formerly in Museo Bouvieri. Not eited by Guibe (1950). Type locality: “Locum Landana dictum, in foliis rhizophoramm” [= Landana, Cabinda enclave], Cabinda Province, Angola. Hyperolius rhizophilus Rochebmne, 1885: Frost (1985:215, 2016). Hyperolius {Hyperolius) rhizophilus Rochebmne, 1885: Fretey et al. (2011:33). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 56): The species is only known from the type locality “Landana” 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II in the Cabinda enclave. Cabinda: “Landana” [-5.21667, 12.15000] (Rochebrune 1885:92; Frost 1985:215, 2016). Taxonomic and distributional notes: The species was described by Rochebrune (1885) from “Locum Landana dictum, in foliis rhizophorarum” in the Cabinda enclave. The type specimen was deposited in the Museo Bouvier, now presumably in the Museum national d’Histoire naturelle, Paris (Frost 2016). The species is not mentioned in Schiotz (1999) or Channing (2001). 12“E I6*E 20°E :4'>E Map 56. Distribution of Hyperolius rhizophilus in Angola. Hyperolius steindachneri Bocage, 1866 Steindachner’s Reed Frog Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the southern Democratic Republic of Congo to northern central-east of Angola and Zambia. Ocurrences in Angola (Map 57): The species occurs in northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1950a:16, 1954a:79, 80, 1964a:151); “Lagoa Carumbo” [-7.74422, 19.95522] (Con- radie et al. 2012a:3); “Camissombo” [-8.15000, 20.65000] (Laurent 1954a:80). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1866a:55, 1866b:75, 1895a: 171, 1897a:204; Loveridge 1936a: 103; Map 57. Distribution of Hyperolius steindachneri in Angola. Hyperolius steindachneri Bocage 1866a:55. Holotype: MBL T.25-226 (collector F.A.P. Bayao). Type locali¬ ty: “Duque de Braganga” [= Calandula], Malanje Province, Angola. Hyperolius machadoi Laurent 1954a: 80, figs. 22, 23. Holotype: MD 495 (collector unknown, presumably A.B. Machado). Type locality: “Camissombo” Lunda Norte, Angola. Hyperolius steindachneri-. Bocage (1866b:75), Noble (1924:253), Loveridge (1936a: 103), Perret (1976a:25), Cei (1977:17), Frost (1985:218, 2016), Poynton and Haacke (1993:15), Schiotz (1999:144), Channing (2001:182), Conradie et al. (2012a:3). Rappia steindachneri'. Bocage (1895a: 171, 1897a:204). Hyperolius steindachneri steindachneri'. Laurent (1950a: 16, 1954a:79, 1961:75, 1964a: 151). Hyperolius machadoi'. Frade (1963:254), Cei (1977:17). Hyperolius {Hyperolius) steindachneri'. Fretey et al. (2011:33). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 115 Perret I976a:25; Frost 1985:218, 2016; Poynton and Haacke 1993:15). Undetermined Locality: without precise locality (Laurent 1954a:79); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: The holotype was destroyed in the 1978 fire in the zoo¬ logical collections in Museu Bocage. Laurent (1954a) described Hyperolius machadoi (Laurent, 1954) from “Camissombo” in Lunda Norte Porvince, but then later Laurent (1964a) synonymized it with H. steindachneri. The only recent report of this species was by Conradie et al. (2012a) who reported several specimens referred to as H. cf steindachneri from Lagao Carumbo in Lunda Norte. Hyperolius vilhenai Laurent, 1964 Vilhena’s Reed Frog (Endemic) Hyperolius vilhenai Laurent 1964a: 155. Holotype: MD 6213 (eolleetor A. Barros Maehado). Type loeality: “galerie forestiere de la riviere, Luita, Poste de Cuilo” [= Luita], Lunda Norte Provinee, Angola. Hyperolius vilhenai'. Cei (1977:17), Frost (1985:219, 2016), Sehiotz (1999:155); Channing (2001:184). Hyperolius {Hyperolius) vilhenai'. Fretey et al. (2011:33). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from northernmost Angola. Ocurrences in Angola (Map 58): The species is known only from the type locality (northern Angola). Lunda Norte: “galerie forestiere de la riviere, Luita, Poste de Cuilo” [-8.03333, 19.41667] (Laurent 1964a:155; Frost 1985:219, 2016). Taxonomic and distributional notes: If valid, the species is also likely to be found in the adjacent Democratic Republic of Congo (Frost 1985, 2016, Sehiotz 1999; Channing 2001). Map 58. Distribution of Hyperolius vilhenai in Angola. Genus Kassina Girard, 1853 Kassina kuvangensis (Monard, 1937) Kuvangu Kassina Cassiniopsis kuvangensis Monard 1937a:41, fig. 19. Holotype: MHNC 90.0012 (collector A. Monard). Type locality: Not stated in the original description but given as “Kuvangu,” Angola by Monard (1938:97). Cassiniopsis kuvangensi: Monard (1938:97), Cei (1977:17). Kassina kuvangensis'. Sehiotz (1975:61), Frost (1985:224, 2016), Poynton and Broadley (1987:179), Sehiotz (1999:245), Channing (2001:187), Fretey et al. (2011:35), Channing et al. (2012:233). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola and Zambia. Ocurrences in Angola (Map 59): The species is known for “Kuvangu,” south-central Angola. Bie: “Cubango basin (10)” [-13.71616, 17.09661] (Conradie et al. 2016: 8-9, 15). Huila: “Kuvan¬ gu” [-14.46667, 16.30000] (Monard 1938:97; Frost 1985:224, 2016; Poynton and Broadley 116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1987:179; Schiotz 1975:61, 1999:247; Chan- ning 2001:188). Cuando Cubango: “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 15). Taxonomic and distributional notes: The type loeality was for some time the only known record for K. kuvangensis. Recently, Conradie et al. (2016) cited the species from Cuando Cubango region. Other authors have also identified populations in Zambia as this species (Schiotz 1999; Channing 2001). 12°E I6'E 20°E 24'>E Map 59. Distribution of Kassina kuvangensis in Angola. Kassina maculosa (Sternfeld, 1917) Marbled Running Frog Megalixalus maculosus Sternfeld 1917:501, pi. 24, fig. 11. Leetotype: SMF 7249 designated leetotype by Mertens (1967:48). Original syntypes SMF [2 speeimens] (eolleetor Sehubotz). Type loeality: “Duma, Ubangi”Aisinhuiion of Kassina senegalensis in Angola. 1993:14); “Molundo” [-15.63333, 15.20000] (Monard 1938:96). Cunene: “5 km W of Pereira de Ega” [-17.06667, 15.73333] (Poynton and Haacke 1993:14). Cuando Cubango: “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9, 16); “Cuito basin (24)” [-14.60622, 18.46722] (Conradie et al. 2016:8-9, 16); “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9-10, 16); “Cuando basin (40)” [-17.45786, 22.91191] (Conradie et al. 2016:9-10, 16). Taxonomic and distributional notes: According to Laurent {in Frost 1985), Largen (1975) mentioned that K. senegalensis may comprise several cryptic species. Hi»E 2&°E 24'‘E Family Arthroleptidae Mivart, 1869 Genus Arthroleptis Smith, 1849 Arthroleptis carquejai Ferreira, 1906 Carqueja’s Squeaker (Endemic) Arthroleptis earquejai Ferreira 1906, pi. 1:165. Holotype: MNHFCP 018586 (eolleetor F. Newton). Type loeality: “Cambondo” (Ferreira 1906:165) Kwanza Norte, Angola. 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Arthroleptis carquejai\ Cei (1977:16), Frost (1985:15, 2016), Ruas (1996:28), Channing (2001:42), Fretey et al. (2011:24), Ceriaco et al. (2014a:27). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 62): The species is known only from northern Angola. Luanda: “Roca Novo (Roqa Novo Mundo)” [-8.433333, 14.13036] (Ceriaco et al. 2014a:27). Kwanza Norte: “Cambondo” [-9.15963, 14.65827] (Ferreira, 1906:165; Frost 1985:15, 2016; Ruas 1996:28; Channing 2001:42; Ceriaco et al. 2014a:27). Taxonomic and distributional notes: Ceriaco et al. (2014a) VQCognizQd Arthroleptis carquejai Ferreira, 1906 as a valid species probably related to a group of species including Arthroleptis variabilis Matschie, 1893. To date it is known only from Angola, although it like¬ ly occurs in the southern Democratic Republic of Congo and possibly Gabon. 12°E l(j®E 24'*E Map 62. Distribution of Arthroleptis carquejai in Angola. Arthroleptis lameerei de Witte, 1921 Lameer’s Squeaker Arthroleptis lameerei de Witte 1921:12. Syntypes: MRAC [5 speeimens],A^i^e R. Laurent in Frost (1985:19) (eolleetor Mission Lemaire). Type loeality: “Lofoi (Katanga),” Demoeratie Republie of Congo. Sehoutedenella lameerer. Laurent (1954a:75, 1964a: 146), Frade (1963:254), Cei (1977:16), Frost (1985:19), Channing (2011:50). Arthroleptis lameerer. Ruas (1996:28), Fretey et al. (2011:24), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from northern Angola, the southern Democratic Republic of Congo, and likely occurs in multiple places in the Albertine Rift Mountains. Ocurrences in Angola (Map 63): The species occurs in northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1954a:75, 1964a: 146; Ruas 1996:28); “Dundo, galerie forestiere de la Luachimo” [-7.38333, 20.85000] (Laurent 1954a:75). Lunda Sul: “Alto Cuilo, Poste de Cacolo, ruisseau-cascade Na-Ipanha” [-10.00000, 19.58333] (Laurent 1964a: 146; Ruas 1996:28); “Alto Cuilo, Poste de Cacolo, rives du lac Cavuemba” [-10.01667, 19.55000] (Laurent 12*E l(i*E 2()®E 24*E Map 63. Distribution of Arthroleptis lameerei in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA II9 1964a: 145); “Alto Cullo, Poste de Cacolo, galerie forestiere du Tcha-Muchito, sous-affluent du Cullo” [-10.01667, 19.45000] (Laurent 1964a;145); “Alto Chicapa, galerie forestiere de laNgun- go, sous-affluent du Cuango-Muque (Rio Gungo)” [-10.83333, 19.28333] (Laurent 1964a: 145; Ruas 1996:28); “Alto Chieapa, Tshimboma, affl. rive gauehe du Cuango-Muque” [-10.76667, 19.20000] (Laurent 1964a: 145; Ruas 1996:28); “Alto Cuilo, rives du Cuilo (Rio Cuilo)” [-10.86667, 19.40000] (Laurent 1964a: 145; Ruas 1996:28); “Alto Chieapa, Caseade de la Kamu- tolonga (Rio Camuntongola)” [-10.88333, 19.25000] (Laurent 1964a: 145; Ruas 1996:28); “Alto Chieapa, sourees de la Tshimboma, affl. rive gauehe du Cuango-Muque (Rio Chimboma)” [-10.88333, 19.23333] (Laurent 1964a: 146; Ruas 1996:28). Undetermined Locality: without precise locality (Laurent 1964a: 146), “areas of forest and savanna in the north and northeast of Angola (Cei 1977:16). Taxonomic and distributional notes: Channing (2001) provided two unspecified localities for A. lameerei in northwestern Angola, and he noted that when collected in Angola this species was found in in leaf litter. Arthroleptis spinalis Boulenger, 1919 Tanganyika Screeching Frog Arthroleptis spinalis Boulenger 1919b: 187. Holotype: MRAC (eolleetor Stappers), fide R. Laurent in Frost (1985:20). Type loeality: “laplaine Saint-Louis, auLake Tanganyika” (Boulenger 1919b: 187), [= St. Louis Plain on the western shore of Lake Tanganyika] Demoeratie Republie of Congo. Athroleptis boulengeri: Laurent (1950a: 15). Schoutedenella spinalis: Cei (1977:16). Arthroleptis spinalis: Fretey et al. (2011:22), Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from northern Angola and the eastern Democratic Republic of Congo. Ocurrences in Angola (Map 64): The species has one record in the northeast of the country. Lunda Norte: “Dundo, galerie fores¬ tiere de la Luachimo” [-7.38333, 20.83333] (Laurent 1950a: 15). Taxonomic and distributional notes: Boulenger (1919a) described Arthroleptis spinalis from eastern Democratic Republic of Congo, which is generally recognized as the only known locality for this species (Fretey et al. 2011, Frost 2016). However, Laurent (1950a) cited Arthroleptis boulengeri (de Witte, 1921) from northeastern Angola and then later recognized this species as a junior synonym of A. spinalis (Laurent 1954). If cor¬ rect, this suggests that this species may be more widespread throughout northern Angola and southern Democratic Republic of Congo. ^4. Distribution of Arthroleptis spinalis in Angola. 12*E l(i*E 2I)°E 24*E 120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Arthroleptis stenodactylus Pfeffer, 1893 Common Squeaker Arthroleptis stenodactylus Pfeffer 1 893 :93, pi. 1 , fig. 1 1 . Holotype: ZMH, presumably destroyed in World War II (eolleetor F. Stuhlmann). Type loeality: “Kihengo” (Pfeffer 1893: pi. I), Tanzania. Corachodichus stenodactylus: Frade (1963:254), Cei (1977:16). Corachodichus stenodactylus stenodactylus: Laurent (1964a: 144). Arthroleptis stenodactylus: Ruas (1996:28, 2002:145), Channing (2001:46), Fretey et al. (2011:25), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is broadly distributed from Kenya and Tanzania, west into the Democratic Republic of Congo and Angola, and south through Zambia, Zimbabwe, Malawi, Mozambique, and extreme northeastern South Africa. Ocurrences in Angola (Map 65): The species is known only for Moxico Province. Moxico: “Calombe, Luso” [-11.83333, 19.93333] (Ruas 1996:28, 2002:145; Channing 2001:47); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a; 144; Ruas 1996:28; Channing 2001:47); “Calunda, Alto Zambeze” [-12.11667, 23.46667] (Laurent 1964a:144; Ruas 1996:28; Channing 2001:47). Undeter¬ mined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “Zambezian highlands” (Frade 1963:254). Taxonomic and distributional notes: According to Ruas (1996), Arthroleptis sten¬ odactylus Pfeffer, 1893 in Angola occurs only in the east of the country. Previous authors (e.g., Channing and Howell 2006; Pickersgill 2007a) have suggested that multiple distinct species exist within what is referred to A. stenodactylus, and preliminary molecular genetic data provides some support for this (Blackburn 2008). It is not yet clear to which of these cryptic species these Angolan populations might correspond. Map 65. Distribution of Arthroleptis stenodactylus in Angola. Arthroleptis xenochirus Boulenger, 1905 Plain Squeaker Arthroleptis xenochirus Boulenger 1905:108, pi. 4, figs. 2, 2a. Holotype: BMNH 1947.2.30.54 (eolleetor W.J. Ansorge), formerly 1904.5.2.101, R. Laurent in Frost (1985:20). Type loeality: “Marimba” (Boulenger 1905:109), Angola. Schoutedenella xenochira: Laurent (1954e:36). Schoutedenella xenochirus: Cei (1977:16), Laurent (1964a:145), Frost (1985:20), Channing (2001:52). Arthroleptis xenochirus: Laurent (1950a: 15), Frade (1963:254), Ruas (1996:28, 2002:145), Fretey et al. (2011:25), Frost (2016), Ceriaeo et al. (2016b:23). Global conservation status (lUCN): Least Concern. Global distribution: A wide-ranging species found in northern Angola, the southern Demo¬ cratic Republic of Congo, northern Zambia and possibly southwestemmost Tanzania. Ocurrences in Angola (Map 66): The species occurs in northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950:15). Lunda Sul: “Poste de Cacolo, Alto Cuilo, gale- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I2I rie forestiere du Tcha-Muchito, sous affluent du Cuilo” [-10.01667, 19.45000] (Laurent 1964a: 145); “Alto Chicapa, galerie forestiere de la Ngungo, sous-affluent du Kwango- Muque” [-10.83333, 19.28333] (Laurent 1964a: 145). Malanje: “Marimba” [-8.36667, 17.03333] (Boulenger 1905:108; Laurent 1954c:36; Frost 1985:20, 2016); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:23). Mexico: “Calombe, Luso” [-11.83333, 19.93333] (Ruas 2002:145). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: This species is well documented from north¬ eastern Angola. Given past taxonomic uncer¬ tainties (e.g., Laurent 1964a; Poyton and Broadley 1985a), it is possible that what is referred to as A. xenochirus across its currently recognized range may in fact represent more than one species. Map 66. Distribution of Arthroleptis xenochirus Angola. Genus Leptopelis Gunther, 1859 Leptopelis anchietae (Bocage, 1873) Anchieta’s TreeFrog (Endemic) Global conservation status (lUCN): Least Concern. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 67): The species is known from the type locality “Huila” and from scattered localities throughout much of the western half of Angola. Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a:147). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1936:131); “Cubango basin (11)” [-13.69413, 17.06177] (Conradie et al. 2016:8-9, 10); “Cubango basin (13)” Hylambates anchietae Bocage 1873b:226. Holotype: MBL T. 13-233 (Perret 1976a:22-23) (collector J.A. Anchieta), destroyed 18 March 1978. Type locality: “I’interieur de Mossamedes” (Bocage 1873b:226), [= Namibe] Angola, later corrected to “Huila” and “Caconda,” Huila Province by Perret (1976a:22). Hylambates anehietae: Bocage (1895a: 177, 1897a:205), Boulenger (1882:133, 1905:110). Leptopelis anehietae'. Schmidt (1936:131), Monard (1938:84), Loveridge (1957:317), Frade (1963:254), Lau¬ rent (1964a: 147), Perret (1976a:22), Cei (1977:17), Frost (1985:227, 2016), Schiotz (1999:306), Channing (2001:196), Channing et al. (2012:106), Fretey et al. (2011:35). Leptopelis cf anehietae'. Conradie et al. (2016:10). Map 67. Distribution of Leptopelis anchietae in Angola. 122 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-13.28061, 16.74722] (Conradie et al. 2016:8-9, 10). Benguela: “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a:177, 1897a:205); “Ebanga” [-12.73333, 14.73333] (Monard 1938:84); “Mount Chininga regions, between 1200 and 1700m” (Chaning 2001:196). Hmla: “Caconda” [-13.73333, 15.06667] (Boeage 1897a:205; Ferret 1976a:12); “Kalukembe” [-13.78333, 14.68333] (Monard 1938:84); “near Humpata” [-14.23814, 13.43331] Channing et al. (2012:106); “Kuvan- gu” [-14.46667, 16.30000] (Monard 1938:84); “Huila” [-15.05000, 13.55000] (Boeage 1895a:177, 1897a:205; Ferret 1976a:22). Namibe: “Tinterieur de Mossamedes (Mossamedes)” [-15.20000, 12.15000] (Boeage 1873b:226; Boulenger 1882:133; Loveridge 1957:317); “Chiyaka distriet” [vie. -15.08333, 12.73333] (Ferret 1976:22). Undetermined Locality: “Western subregion and Angolan highlands (Angola coastal or watersheds)” (Frade 1963:254); “areas of forest and savan¬ na in the north and notheast of Angola” (Cei 1977:17). Taxonomic and distributional notes: Ferret (1976a) suggested that Leptopelis anchietae (Boeage, 1873) is closely related to Leptopelis nordequatorialis Ferret, 1966 and L. oryi Inger, 1968. Leptopelis aubryi (Dumeril, 1856) Gabon Forest Treefrog HylaAubryi Dumeril 1856:561. Syntypes: MNHN 833, 1571 and 4603 (collector J. Auhry-Lecomte). Type locality: “Gabon.” Hylambates Aubryi: Peters (1877a:618), Boeage (1895a:181). Leptopelis aubryi: Laurent (1954a:75), Loveridge (1957:318), Cei (1977:17), Schiotz (1999:266), Fretey et al. (2011:35), Channing et al. (2012:107), Forst (2016). Global conservation status (lUCN): Least Concern. Global distribution: A forest-associated species distributed broadly in Central Africa, extend¬ ing from southeastern Nigeria through Cameroon and Gabon, east into the Central African Republic, and south through the Dem¬ ocratic Republic of Congo to northern Angola. Ocurrences in Angola (Map 68): The species occurs in the Cabinda enclave and in the extreme northeast of Lunda Norte Province. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:618; Boeage 1895a: 176); “cote de Loango” (Boeage 1895a: 181); “Cabinda enclave” (Channing et al. 2012:107). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:75). Taxonomic and distributional notes: Many species of Leptopelis in Central Africa have been confused with Leptopelis aubryi, which has a broad distribution and occurs both within forests and along the margins of forests. Map 68. Distribution of Leptopelis aubryi in Angola. Leptopelis bocagii (Gunther, 1865) Bocage’s TreeFrog Cystignathus Boeagii Gunther 1865a:481, pi. 33, fig. 2. Holotype: not stated, destroyed by fire 18 Mareh 1978 (eolleetor F.P. Bayao). Two speeimens reeeived by BMNH from MBL may be types fide Frost (2016). Type loeality: “Duque de Braganee” (Gunther 1865a:482), [= Calandula], Malanje Provinee, Angola; Hylambates angolensis Boeage 1893:119. Syntypes: MBL T.14-242-244 [3 speeimens] (eolleetor J.A. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 123 d’ Anchieta), destroyed by fire 18 Mareh 1978. Type loeality: “Cahata” [= Caota], “Quindumbo”, “Quis- sange”, Benguela Provinee and “Caeonda” Huila Provinee, Angola. Hylambates bocagei var. leucopunctata Ferreira 1904:113. Syntype: MHNFCP 017324 (eolleeted by F. New¬ ton). Type loeality: “Gumba” (Ferreira 1904:113), [= Gumba, Serra de Selles] Kwanza Norte Provinee, Angola. Cystignathus Bocagir. Boeage (1866a:54). Hylambates bocagii: Boulenger (1882:133), Boeage (1895a: 176, 1897a:205). Hylambates angolensis: Boeage (1895a:179, 1896a:113, 1897a:205, 1897b:211), Perret (1976a:23). Hylambates boeagei var. leueopunetata: Ferreira (1904:113). Hylambates boeagei'. Ferreira (1906:164). Leptopelis angolensis: Sehmidt (1936:131), Monard (1938:84), Frade (1963:254). Leptopelis boeagei'. Loveridge (1933:393, 1953a:342, 1957:317), Laurent (1950a: 15, 1954a:76), Hellmieh (1957a:30), Inger (1959:539, 541), Cei (1977:17), Sehiotz (1975:14), Frost (1985:228, 2016), Poynton and Broadley (1987:174), Sehiotz (1999:289), Channing (2001:198), Largen (1977:96, 2001:349), Fretey et al. (2011:36), Ceriaeo et al. (2014a:25), Ceriaeo et al. (2016b:25). Leptopelis parboeagii: Poynton and Broadley (1987:171), Sehiotz and Van Daele (2003:146). Leptopelis ef. boeagii: Amiet (2012:466). Global conservation status (lUCN): Least Concern. Global distribution: This savanna species has a broad distribution extending from Ethiopia in the north, south to Angola, northeastern Namibia, Zambia, Zimbabwe, and Mozambique. Ocurrences in Angola (Map 69): The species occurs from the western regions to northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:76; Ceriaeo et al. 2014a:26); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a;15, 1954a:76; Ceriaeo et al. 2014a:26). Kwanza Norte: “N’Golla Bumba” [-9.03333, 14.60000] (Ferreira 1906:164; Ceriaeo et al. 2014a:26); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:164; Ceriaeo et al. 2014a:26); “Quilombo” [-9.33333, 14.90000] (Ferreira 1906:164; Ceriaeo et al. 2014a:26). Kwanza Sul: “Gumba” [-11.26667, 14.28333] (Ferreira 1904:113; Loveridge 1957:317; Ceriaeo et al. 2014a:25). Malanje: “Duque de Bragance (Duque de Brangan9a)” [-9.10000, 15.95000] (Gunther 1865a;482; Boeage 1866a:54, Map 69. Distribution of Leptopelis bocagei in Angola. 1895a:176, 1897a:205; Loveridge 1933:393, 1953a:342, 1957:317; Sehiotz 1975:14; Perret 1976a:23; Frost 1985:228, 2016; Poynton and Broadley 1987:174; Largen 2001:349; Ceriaeo et al. 2014a:27), “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:25). Bie: “Chi- tau” [-11.43333, 17.15000] (Schmidt 1936:131). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:84). Benguela: “Quibula” [-12.28333, 14.68333] (Boeage 1893:119, 1895a:179, 1897a:205); “Cahata” [-12.35000, 14.81667] (Boeage 1893:119, 1895a:179, 1897a;205; Perret 1976a:23); “Quissange” [-12.43333, 14.05000] (Boeage 1893:119, 1895a:179, 1897a:205; Love¬ ridge 1933:393; Perret 1976a:23); “Quindumbo” [-12.46667, 14.93333] (Boeage 1893:119, 1895a:179, 1897a:205; Perret 1976a:23); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:30); “Hanha” [-13.30000, 14.20000] (Boeage 1896a:113; 1897b:211). Huila; “Caeonda” 124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-13.73333, 15.06667] (Boeage 1893:133, 1895a:179, 1897a:205; Loveridge 1957:317; Ferret 1976a:23). Taxonomic and distributional notes: Ferret (1976a) noted that the synonymy ofZ. angolen- sis with L. bocagei was diffieult to evaluate due to the differenees in maturity between the type speeimens of these two speeies, which were lost in the 1978 fire in Museu Boeage. Poynton and Broadley (1987) described a similar species, Leptopelis parbocagii based on five specimens collected at Mabwe on the eastern shore of Lake Upemba, in what is now the Democratic Repub¬ lic of Congo. Because of their morphological similarity and overlapping distributions, the identifi¬ cation of these two species remains difficult (Schiotz 1999; Schiotz and Van Daele, 2003). Leptopelis bocagii is a widespread species in southern and central Africa (Channing 2001), though it may comprise several cryptic species (Largen 1977; Amiet 2012). Ferreira (1904) described the variety leucopunctata, currently considered a synonym of Leptopelis bocagii (Gunther, 1865). Ceriaco et al. (2014a) provided more details about the taxonomic identity and nomenclature of L. bocagii and located a syntype from “Gumba, Sa [Serra] de Selles = Angola,” collected by Francisco Newton during the 1903-1905 expedition in Angola, in the Museu de Historia Natural da Universidade do Porto, Portugal. Leptopelis cynnamomeus (Boeage, 1893) Angola Forest Treefrog Hylambates cynnamomeus Boeage 1893:120. Holotype: MBLT. 16-250 (collector J. A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Quilengues,” Huila Province, Angola. Hylambates eynnamomeus: Bcoage (1895a: 180, 1897a:205). Leptopelis viridis einnamomeus: Laurent (1964a: 148), Cei (1977:17). Leptopelis eynnamomeus'. Schiotz (1975:24), Frost (1985:229, 2016), Poynton and Broadley (1987:177), Schiotz (1999:299), Channing (2001:200), Schiotz and Van Daele (2003:146), Fretey et al. (2011:36). Leptopelis boeagii: Loveridge (1957:317). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from central Angola, northwestern Zambia, and southern Democratic Republic of Congo. Ocurrences in Angola (Map 70): The species is known from the type locality “Quillengues” in Huila, as well as from central and northeast¬ ern Angola, next to the boundary with the Democratic Republic of Congo and Zambia. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a:148; Channing 2001:201). Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a: 148; Channing 2001:201). Huila: “Quillengues” [-14.06667, 15.08333] (Boeage 1893:120, 1895a:180, 1897a:205; Loveridge 1957:317; Schiotz 1975:24; Ferret 1976a:22; Frost 1985:229, 2016; Channing 2001:201). Taxonomic and distributional notes: Ferret (1976a) examined the holotype from Angola in the collection of Museu Boeage and condisered two paratypes from “Boloma” in Guinee-Bissau to represent Leptopelis viridis (Gunther, “1868” 1869). Distribution of Leptopelis cynnamomeus in MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 125 Leptopelis jordani Parker, 1936 Congulu Forest Treefrog (Endemic) Leptopelis jordani Parker 1936:144. Holotype: BMNH 1947.2.19.95 [formerly BMNH 1936.8.1.226] (eol- leetor K. Jordan). Type loeality: “Congulu (700-800m)” [= Congulo] Kwanza Sul Provinee, Angola. Leptopelis jordani Parker, 1936: Frost (1985:230, 2016), Fretey et al. (2011:36). Global conservation status (lUCN): Data Deficient. Global distribution: This species is known only from Angola. Ocurrences in Angola (Map 71): The species is known only from the type locality “Congulu” in western Angola. Kwanza Sul: “Congulu (700-800m)” [-10.86667, 14.28333] (Parker 1936:144; Frost 1985:230, 2016). Taxonomic and distributional notes: In his description, Parker (1936) noted similari¬ ties to the widespread Leptopelis aubryi (Dumeril, 1856). There have been no recent records since the original description and the validity of this taxon remains in question. This species is not mentioned in Schiotz (1999) or Channing (2001). 12°E ICr'E 20°£ :4'*E Map 71. Distribution of Leptopelis jordani in Angola. Leptopelis marginatus (Bocage, 1895) Quissangue Forest Treefrog (Endemic) Hylambates marginatus Bocage 1895a: 178. Holotype: MBL T.218-241 (colleetor J.A. d’Anehieta), destroyed hy fire 18 March 1978. Type locality: “Quissange, dans Finterieur de Benguella” [= Quissange] Bengue- la Province, Angola. Hylambates marginatus: Boeage (1897a:205), Cei (1977:17). Leptopelis boeagii: Loveridge (1933:393). Leptopelis anehietae: Loveridge (1957:317). Leptopelis marginatus: Perret (1976a:23), Forst (1985:230, 2016), Fretey et al. (2011:36). Global conservation status (lUCN): Data Deficient. Global distribution: This species is known only from Angola. Ocurrences in Angola (Map 72): The species is known only from the type locality in estern Angola. Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1895a:179, 1897a:205; Loveridge 1933:393, 1957:317; Perret 1976a:23, Frost 1985:230, 2016). Taxonomic and distributional notes: Perret (1976a) concluded that L. marginatus was a valid species, in contrast to Loveridge (1933) who recognized it as a synonym of Leptopelis boeagii (Gunther, 1865). There have been no other recent records and the holotype Map 72. Distribution of Leptopelis marginatus in Angola. 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II was destroyed in the 1978 fire in Museu Bocage. This species is not mentioned Schiotz (1999) or Channing (2001). This should be recognized as a nomen dubium. Leptopelis notatus (Peters, 1875) Common Forest Treefrog Hylambates notatus Peters 1875:205, pi. 2, figs. 1, la. Holotype: ZMB 8471 (eollector R. Buehholz) fide Bauer et al. (1995:43). Type loeality: “Camemns” (Peters 1875:206), [= Cameroon]. Leptopelis tessmanni: Laurent (1950a: 15), Frade (1963:254). Leptopelis notatus: Laurent (1964a: 147), Cei (1977:17), Schiotz (1999:264), Fretey et al. (2011:36), Chan¬ ning et al. (2012:116), lUCN SSC Amphibian Specialist Group (2013a), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: This is a widespread species occurring in and near forests extending from Cameroon, Gabon, Republic of Congo, the western Democratic Republic of Congo, and into northernmost Angola. Ocurrences in Angola (Map 73): The species occurs in the northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1964a; 147); “Mussalomuca” [-7.43333, 20.45000] (Laurent 1964a: 147); “rive de la Tchihumbwe, 40 km a Lest de Dundo” [-8.01667, 19.31667] (Laurent 1950a:15). Lunda Sul; “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a: 147). Undeter¬ mined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:17). 24°E Taxonomic and distributional notes: Map 73. Distribution of Leptopelis notatus in Angola. Fretey et al. (2011) questioned the presence of this species in Angola. Leptopelis viridis (Gunther, “1868” 1869) Rusty Forest Treefrog Hylambates viridis Gunther “1868” 1869:487. Holotype: BMNH 1947.2.10.23 (purchased from Mr. Stevens), formerly BMNH 65.5.3.63. Type locality: “West Africa”. Hylambates viridis: Bocage (1873b:226, 1895a: 176, 1897a:205), Boulenger (1882:134). Leptopelis viridis: Fretey et al. (2011:37), lUCN SSC Amphibian Specialist Group (2013b), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: This species is found in savannas across western and Central Africa, extending south into Angola. Ocurrences in Angola (Map 74): The species has been recorded from Malanje Province. Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1873b:206, 1895a: 179, 1897a:205). Taxonomic and distributional notes: Bocage (1873b, 1895a, 1897a) cited a single juvenile from “Duque de Braganga” collected by Bayao whereas Boulenger (1882) cited another individual from Angola collected by J. J. Monteiro, although without locality. Many recent summaries of the distribution of this species, including Schiotz (1999), lUCN SSC Amphibian Specialist Group (2013b), and Frost (2016), have excluded Angola from its distribution though Fretey et al. (2011) MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 127 considered that it was present. Due to the lack of recent data and the limited information sup¬ plied by previous authors for Angolan speci¬ mens, the presence of L. viridis in Angola is questionable. 1I°E It'E. 20°E 24*E Map 74. Distribution of Leptopelis viridis in Angola. Genus Trichobatrachus Boulenger, 1900 Trichobatrachus robustus Boulenger, 1900 Hairy Frog Trichobatrachus robustus Boulenger 1900:443, pi. 30. Syntypes: BMNH 1947.2.30.85-86 (eollector G.L. Bates), formerly BMNH 1900.2.17.59-60. Type locality: “Benito River, Gaboon” (Boulenger 1900:443), Equatorial Guinea. Trichobatrachus robustus'. Amiet and Burger (2004), Frost (2016). Trichobatrachus cf robustus'. Ernst et al. (2014:298, 299). Global conservation status (lUCN): Least Concern. Global distribution: This forest species occurs throughout the Atlantic Coastal Forests, extending from Nigeria and Cameroon south into northern Angola. Ocurrences in Angola (Map 75): This species was only recently reported from Uige Province in the north of the country. Uige; “Serra Pingano (along Mbalage creek)” [-7.66814, 14.93519] (Enst et al. 2014:289); “Serra Pingano, approximatley 40 m away from Mbalage ereek” [-7.68467, 14.93242] (Enst et al. 2014:289); “Mbalage creek, Serra Pingano” [-7.68514, 14.92944] (Enst et al. 2014:289). Taxonomic and distributional notes: Ernst et al. (2014) discovered a distinct lineage of Trichobatrachus in “Serra do Uige,” an iso¬ lated forest remnant in northern Angola. Fur¬ ther studies are needed to clarify whether this population is conspecifie with those from far¬ ther north in Cameron and Gabon (Ernst et al., 2014). There remains little information Map 75. Distribution of Trichobatrachus robustus in Angola. 128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II available on the habitat requirements and natural history of this charismatic species (Amiet and Burger 2004; Ernst et al. 2014). Family Ptychadenidae Dubois, 1987 Genus Hildebrandtia Nieden, 1907 Hildebrandtia ornatissima (Bocage, 1879) Angola Ornate Frog (Endemic) Rana ornatissima Bocage 1879a:98. Holotype: MBL T.78-160 (collectors H.C. Capello and R. Ivens), destroyed by fire 18 March 1978. Type locality: “Bihe” [= Bie] Bie Province, Angola. Hildebrandtia angolensis Nieden 1908:657. Holotype: ZMB 20024 ? (Gleim). Type locality: “Loanda in Angola” (Nieden 1908:657), [= Luanda], Luanda Province, Angola. Rana ornatissima: Bocage (1879c:89, 1895a: 157, 1897a:202), Boulenger (1905:107, 1919a:35), Schmidt and Inger (1959:38), Frade (1963:254). Rana {Hildebrandtia) ornatissima: Monard (1938:105). Rana (Hildebrandtia) myotympanum: Monard (1937a:49, 1938:106). Rana ornata ornatissima: Schmidt and Inger (1959:40). Hildebrandtia myotympanum: Cei (1977:16, 17). Hildebrandtia ornatissima: Perret (1976a: 19), Cei (1977:16, 17). Hildebrandtia ornata ornatissima: Ruas (1996:24). Hildebrandtia ornata: Channing (2001:294). Hildebrandtia ornatissima: Frost (1985:460, 2016); Fretey et al. (2011:40). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 76): This species is distributed in south-central Angola. Luanda: “Loanda” [-8.83333, 13.26667] (Nie¬ den 1908:657). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:105; Ruas 1996:24); “Galanga” [-12.06667, 15.15000] (Bocage 1895a:157, 1897a;202; Perret 1976a;19; Ruas 1996:24). Bie: “Bihe” [-12.38333, 16.95000] (Bocage 1879a;98, 1879c;89, 1897a:202; Bou¬ lenger 19 19a: 3 5; Perret 1976a: 19; Ruas 1996:24; Frost 1985:460, 2016); “Bingondo” [-12.06667, 17.41667] (Boulenger 1905:107; Ruas 1996:24). Huila: “Kuvangu” [-14.46667, 16.30000] (Monard 1937a:49, 1938:106; Ruas 1996:24); “Osi” [-15.08333, 15.41667] (Monard 1937a:49, 1938:106; Ruas 1996:24); “Dongue” [-15.43333, 14.05000] (Poyntonand Haacke 1993:14; Ruas 1996:24); “Molundo” [-15.63333, 15.20000] (Monard 1937a:49, 1938:106; Ruas 1996:24). Undetermined Locality: “Konodoto” (Boulenger 1919a:35); swamp” (Boulenger 1919a:35). Taxonomic and distributional notes: The putative holotype of Hildebrandtia angolensis Nieden, 1908 is not marked as such in the ZMB catalogue, but it appears to be the only candidate specimen in the Berlin collection that is consistent with the type locality of “Loanda.” See also taxonomic account for Hildebrandtia ornata. Map 76. Distribution of Hildebrandtia ornatissima in Angola. Cafita MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 129 Hildebrandtia ornata (Peters, 1878) Ornate Frog Pyxicephalus ornatus Peters 1878:207, pi. 2, fig. 7. Holotype: ZMB 9297 (eolleetor J.M. Hildebrandt) fide Bauer et al. (1995:49). Type loeality: “Taita” (Peters 1878:207), [= Teita], Kenya. Rana ornatissima: Boulenger (1919a:35). Rana ornata: Inger (1959: 541). Hildebrandtia ornata ornata: Poynton and Broadley (1985b: 141), Poynton and Haaeke (1993:14), Ruas (1996:24). Hildebrandtia ornata: Channing (2001:294), Fretey et al. (2011:40). Global conservation status (lUCN): Least Concern. Global distribution: The species is broadly distributed in the savannas of sub-Saharan Africa, extending from the Sahel of West Africa, throughout eastern and south-central Africa, and into northeastern South Africa. Ocurrences in Angola (Map 77): This species is distributed in south-central Angola. HmTa: “Dongue” [-15.43333, 14.05000] (Poynton and Haaeke 1993:14). Cunene: “2 km NW of Calequero” [-16.28333, 14.30000] (Poynton and Haaeke 1993:14; Ruas 1996:24); “23 km NW of Pereira de Ega - Rogadas” [-16.95000, 15.56667] (Poynton and Haaeke 1993:14; Ruas 1996:24); “Ponang Kuma” [-17.05000, 14.65000] (Boulenger 1919a:35). Taxonomic and distributional notes: Hildebrandtia ornatissima was described by Bocage (1879c) based on a single specimen from the type locality “Bihe.” Later, Nieden (1908) described Hildebrandtia angolensis from “Loanda” but this nomen was preoccu¬ pied by Rana angolensis Bocage, 1866. Boulenger (1919) proposed Rana miotympa- Map 77. Distribution in Angola. num as a replacement name for Hildebrandtia angolensis, and Schmidt and Inger (1959) later recognized this as a junior synonym of Rana ornata ornatissima (Schmidt and Inger 1959). Perret (1976a) removed//, ornatissima from synonymy With Hildebrandtia ornata (Peters, 1878), though this has not been accepted by all subsequent authors and whether these represent one or two species remains unclear. Channing (2001) referred all specimens of Hildebrandtia from southern and central Africa, including from Angola, to H. ornata. In contrast, Poynton and Haaeke (1993), Ruas (1996), and Fretey et al. (2011) recognized both H. ornata and H. ornatissima in Angola. The delimitation of these two species requires further review. Genus Ptychadena Boulenger, 1917 Ptychadena anchietae (Bocage, 1867) Anchieta’s Grass Frog Rana anehietae Bocage 1867a:843, fig. 1. Syntypes: MBL T.9.134 [3 specimens] (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Benguella” [= Benguela] Benugela Province Angola. Rana anehietae: Peters (1877a:618). Rana maseareniensis: Boulenger (1882:52), Bocage (1895a:160). Rana oxyrhynehus oxyrhrynehus: Loveridge (1957:340). Ptyehadena anehietae: Perret (1976a:19, 1979:18), Poynton and Broadley (1985b:146), Frost (1985:471, 2016), Poynton and Haaeke (1993:14), Ruas (1996:25, 2002:144), Largen (2001:335), Channing 130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (2001:319), Channing et al. (2012:305), Ceriaco et al. (2016b:43). Ptychadena {Ptychadena) anchietae: Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: This broadly distributed species is known from Ethiopia south to Ango¬ la and southeast to South Africa. Ocurrences in Angola (Map 78): The species occurs in both western regions and eastern regions near the Zambian border. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a: 6 18). Kwanza Norte: “Dondo” [-9.68333, 14.43333] (Poynton and Haacke 1993:14). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:43). Moxico: “Diolo lake” [-11.50000, 22.01667] (Ruas 1996:25, 2002:144). Bengue- la: “Benguella” [-12.58333, 13.41667] (Boca- ge 1867a:843; Loveridge 1957:340; Perret 1976a: 19; Poynton and Broadley 1985b: 146; Frost 1985:471, 2016; Largen 2001:335); “Dombe” [-12.95000, 13.10000] (Bocage 1895a: 160; Ruas 1996:25). Undetermined Locality: “savanna from Ethiopia to Natal, A 1 55 • 1 nnn i Map 78. Distribution of Ptychadena anchietae in across to Angola (Cei 1977:16). Angola Taxonomic and distributional notes: Perret (1979) erroneously referred to “Huila” as the type locality. Bocage (1895a) and Boulenger (1882) considered R. anchietae to be a synonym of Rana mascareniensis (Dumeril and Bibron, 1841). The latter species was also confused with Rana oxyrhynchus (Smith, 1849) because of its morphological similarity (Perret 1976a). Poynton (1964a) recognized P. anchietae as valid, but later (Poynton 1970) considered it to be a synonym of P. superciliaris (Gunther, 1858). Perret (1976a), however, disagreed noting that P. superciliaris is a forest species distinct from P. anchi¬ etae, and this interpretation was followed by Poynton and Broadley (1985b). Ptychadena ansorgii (Boulenger, 1905) Ansorge’s Grass Frog Rana Ansorgii Boulenger 1905:107, pi. 4, fig. 1. Holotype: BMNH 1947.2.29.48 (eolleetor W.J. Ansorge). Type loeality: “Between Benguella and Bihe” [= between Benguela and Bie], Angola. Rana ansorgii-. Loveridge (1933:371, 1936a:95, 1936b:419, 1953a:373, 1957:343). Rana {Ptychadena) ansorgii'. Monard (1937a:52, 1938:110), Parker (1939:142). Ptychadena ansorgei: Laurent (1950:14). Rana ansorgei: Inger (1959:541), Sehmidt and Inger (1959:61). Ptychadena ansorgi: Cei (1977:16, 17). Ptychadena ansorgii: Loveridge (1936a:95), Poynton and Broadley (1985b: 152), Ruas (1996:26), Largen (2001:177), Channing (2001:321), Frost (1985:471, 2016). Ptychadena {Ptychadena) ansorgii: Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola, southern Democractic Republic of Congo, northern Zambia, and Malawi. Ocurrences in Angola (Map 79): The species occurs in the extreme northeast and in the west- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I3I em regions of the country. Lunda Norte: “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 14; Ruas 1996:26). Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:52, 1938:110; Ruas 1996:26). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:142; Ruas 1996:26). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:52, 1938:110; Ruas 1996:26); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:14; Ruas 1996:26). Benguela: “Benguela” [-12.58333, 13.41667] (Loveridge 1936a:95). HuiTa: “Kalukembe” [-13.78333, 14.68333] (Monard 1937a:52, 1938:110; Ruas 1996:26). Cuando Cubango: “Kandingu (Kului)” [-14.66667, 17.70000] (Monard 1937a:52, 1938:110; Ruas 1996:26). Undetermined Locality: “Between Benguella and Bihe” (Boulenger 1905:107; Map 79. Distribution of Ptychadena ansorgii in Angola. Loveridge 1933:371, 1936b:419, 1953a:373, 1957:343; Schmidt and Inger 1959:61; Poynton and Broadley 1985b: 152; Frost 1985:471, 2016; Ruas 1996:26), “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Relatively little is known of the biology of this species, which is found in the forests and savannas of south-central Africa. Ptychadena bunoderma (Boulenger, 1907) Rough Grass Frog Rana bunoderma Boulenger 1907a:214. Holotype: BMNH 1947.2.2.86 (eolleetor W.J. Ansorge). Type loeal- ity: “Caeonda,” Huila Provinee, Angola. Rana (Ptychadena) buneli: Monard (1937a:55, 1938: A. Monard). Type loeality: “environs de Dala, sur le Tyiumbwe,” Lunda Sul Provinee, Angola. Rana bunoderma'. Sehmidt (1936:129). Ptychadena bunoderma'. Laurent (1964a: 142), Cei (1977:16), Poynton and Broadley (1985b: 152), Frost (1985:471, 2016), Ruas (1996:26), Largen (2001:177), Channing (2001:324). Ptychadena (Ptychadena) bunoderma'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola and northwestern Zambia. Ocurrences in Angola (Map 80): The species occurs in central and eastern Angola. Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a: 142; Ruas 1996:26); “Alto Chicapa, humidiherbosa des sources de la Kamassaka (Rio Camassaca)” [-10.88333, 114). Syntypes: MNHNC [three speeimens] (eolleetor 12°E l(i®E 20°E I4'*E Map 80 Distribution of Ptychadena bunoderma in Angola. 132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 19.25000] (Laurent 1964a: 142; Ruas 1996:26); “Alto Chicapa, humidiherbosa des sources de la Kamutongola (Rio Camuntongola)” [-10.53000, 19.15000] (Laurent 1964a: 142; Ruas 1996:26); “environs de Dala, sur le Tyiumbwe (Rio Chiumbe)” [-11.03333, 20.20000] (Monard 1937a:55, 1938:114; Ruas 1996:26). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1936:129; Ruas 1996:26). Hmla: “Caconda” [-13.73333, 15.06667] (Boulenger 1907a:214; Poynton and Broadley 1985b: 152; Frost 1985:471, 2016; Ruas 1996:26). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: Gaston de Witte initially identified Monard ’s types of R. buneli as ''Rana n. sp., voisine de Rana mascareniensis D.B.,” although based on Monard’s (1937a) description it is closer to P. bibronii or P. bunoderma. Laurent (1964a) provided some new records for Ptychadena bunoderma in Angola, including from near the type locality of P. buneli which he placed in the synonymy of P. bunoderma. Channing (2001) provided several records further north for the species without detailed information. Ptychadena grandisonae Laurent, 1954 Grandison’s Grass Frog Ptychadena grandisonae Laurent 1954b: 11, pi. 1, figs. 2, 5, pi. 3, figs. 1-2, pi. 4, figs. 1, 9. Holotype: pre¬ sumably MD 506? (eolleetor M. de Petehkowsky). Type loeality: “Muita, Luembe E” [= Luembe] Lunda Norte, Angola. Rana (Ptyehadena) bibroni: Monard (1937a:51, 1938:109). Ptyehadena bibronii'. Laurent (1950a: 14). Ptyehadena grandisonae'. Laurent (1964a: 139), Cei (1977:16), Poynton and Broadley (1985b: 150), Frost (1985:472, 2016), Poynton and Haaeke (1993:14), Channing (2001:325). Ruas (2002:144). Rana grandisonae'. Sehmidt and Inger (1959:70). Ptyehadena (Ptyehadena) grandisonae'. Fretey et al. (2011:41). Ptyehadena ef grandisonae'. Conradie et al. (2016:17). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the Democratic Republic of Congo, Angola, and Zambia. Ocurrences in Angola (Map 61): The species occurs in the northeast of the country, although there are some records further south. Lunda Norte: “environs du Dundo” [-7.36667, 20.83333] (Laurent 1964a: 139); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 14, 1954b: 11; Schmidt and Inger 1959:70; Poynton and Broadley 1985b: 150; Frost 1985:472, 2016). Lunda Sul: “Alto Chicapa, mare Tcha-Mutuka, pres des sources du Cuango-Muque” [-10.76667, 19.20000] (Laurent 1964a:139); “Alto Chica¬ pa, sources du Cuilo” [-10.86667, 19.40000] (Laurent 1964a: 139); “sources de la Kamuton¬ gola, Alto Chicapa” [-10.53000, 19.15000] (Laurent 1964a:139); “Lunda” [-10.96667, 20.06667] (Monard 1937a:51, 1938:109). Mexico: “Luso” [-11.78333, 19.91667] (Ruas 2002:144). Benguela: “Ebanga” [-12.73333, 12°E l(j®E 2()°E I4"E Map 81. Distribution of Ptychadena grandisonae in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 133 14.73333] (Monard I937a:5I, 1938:109). Hulla: “Kalukembe” [-13.78333, 14.68333] (Monard I937a:5I, 1938:109); “5 km W of Calucembe” [-13.80000, 14.68333] (Poynton and Haacke 1993:14); “Nuntechite lagoon” [-15.13333, 13.41667] (Poynton and Haacke 1993:14). Cuando Cubango: “on the edge of the upper Longa River floodplain (53b)” [-14.56322, 18.44394] (Con- radie et al. 2016:9, 12, 17). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: Laurent (1954b) considered Monard’s usage of the name Ptychadena bibroni {non Hallowell) to refer to Ptychadena grandisonae Laurent, 1954. We agree with that conclusion and we also considQrRana {Ptychadena) bibroni (Monard 1937a, 1938) a synonym of grandisonae, since the distribution range for the real Ptychadena bibroni (Hallow¬ ell, 1845) is limited to the north of Gulf of Guinea, from the Gambia and Mauritania to the nort¬ heastern Demoeratie Republic of Congo and presumably to South Sudan (Frost, 2016). Ptychadena guibei Laurent, 1954 Guibe’s Grass Frog Ptychadena chrysogaster Laurent 1954b:18, pi. 2, fig. 2, pi. 1, fig. 3, pi. 3, figs. 5-6, pi. 4, figs. 3, 5. Holo- type: MRAC 109096 fide Lang (1990:12). Type loeality: “Lae Karago, Terr, de Kisenyi, Ruanda.” Ptyehadena ehrysogaster guibei Laurent 1954b:23. Holotype: MD 2097 (eolleetor M. de Petehkowsky). Type loeality: “Muita, Luembe E” [= Luembe] Lunda Norte, Angola. Ptyehadena bibroni: Laurent (1950a: 14). Rana ehrysogaster guibei: Sehmidt and Inger (1959:65). Ptyehadena ehrysogaster guibei: Laurent (1964a: 136), Cei (1977:16). Ptyehadena guibei: Poynton and Broadley (1985b: 154), Ruas (1996:26), Largen (2001:177), Channing (2001:326), Frost (2016), Ceriaeo et al. (2016b:47), Conradie et al. (2016:18). Ptyehadena ehrysogaster: Frost (1985:471). Ptyehadena {Ptyehadena) guibei: Fretey et al. (2011:41). Ptyehadena {Ptyehadena) ehrysogaster: Fretey et al. (2011:41). Global conservation status (lUCN): Least Coneem. Global distribution: The speeies is known from the Democratic Republic of Congo and east¬ ern Angola, through Zambia, Botswana, Zimbabwe, Malawi, and northern Mozambique. Ocurrences in Angola (Map 82): The speeies occurs especially in eastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1964a: 136; Ruas 1996:26); “Muita (Luem¬ be E)” [-7.80000, 21.45000] (Laurent 1950:14, 1954b:23; Schmidt and Inger 1959:65; Poyn¬ ton and Broadley 1985b: 154; Ruas 1996:26). Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a: 136; Ruas 1996:26). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:47). Cuando Cubango: “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9-10, 18); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 18); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). 12°E Id'E 20°E UeE Map 82. Distribution of Ptychadena guibei in Angola. 134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: Poynton and Broadley (1985b) elevated P. guibei to a full species. Fretey et al. (2011:41) cited Ptychadena (Ptychadena) chrysogaster Laurent, 1954 for Angola though we believe that this should be referred to P. guibei (Frost 2016). New material cited by Conradie et al. (2016) extends the species distribution into extreme southeastern Angola. Ptychadena keilingi Monard, 1937 Keiling’s Grass Frog Rana {Ptychadena) keilingi Monard 1937a:53, figs. 14-16. Syntypes: MNHC 90.0829 and 90.0830 (collector A. Monard). Type locality: “Dala,” Lunda Sul, Angola. Rana (Ptyehadena) keilingi'. Monard (1938:112), Marx (1959:442), Laurent (1964a: 141). Ptyehadena keilingi'. Forcart (1946:126), Cei (1977:16), Poynton and Broadley (1985b: 154), Frost (1985:472, 2016), Ruas (1996:26), Largen (2001:177), Channing (2001:327). Ptyehadena {Ptyehadena) keilingi'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the Democratic Republic of Congo and adja¬ cent northeastern Angola to extreme northwestern Zambia. Ocurrences in Angola (Map 83): The species occurs in northeastern Angola. Lunda Norte: “Riviere Luita river, Poste de Cuilo (Rio Luita)” [-8.03333, 19.41667] (Laurent 1964a: 141; Ruas 1996:27); “Poste de Luangue, humidiherbosa du risseau Katcheleka, affl. ouest Luangue, entre la Lunguena et le Tcha- Pemba (Posto do Luangue)” [-9.08333, 19.71667] (Laurent 1964a:141; Ruas 1996:27). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:141; Ruas 1996:27); “Dala” [-11.03333, 20.20000] (Monard 1937a:53, 1938:112; Forcart 1946:126; Marx 1959:442; Poynton and Broadley 1895b: 154; Frost 1985:472, 2016; Ruas 1996:26). Moxico: “rives du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a:141; Ruas 1996:27). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977: 16). Distribution of Ptychadena keilingi in Angola. Taxonomic and distributional notes: Two of the five type specimens noted by Monard (1937a) remain in the Musee d’Histoire Naturelle, La-Chaux-de Fond, Switzerland. Little is known of this frog species. Ptychadena cf. mascareniensis (Dumeril and Bibron, 1841) Mascarene Grass Frog Rana maseareniensis Dumeril and Bibron 1841:350. Syntypes: MNHN 4379-4381 fide Guibe (1950), MCZ A- 1044 (exchange from MNHN) fiide Barbour and Loveridge (1929), USNM 10975 (2 specimens) fiide Cochran (1961) (collectors J.R.C. Quoy and J.P. Gaimard, J.-J. Dussumier, J.-B. Leschenault, Nivoy). Type locality: “lies Mascareignes, ou les Sechelles, Maurice et Bourbon.” Restricted to “Island of Reu¬ nion, Mascarene Islands (Indian Ocean)” by Cochran (1961) and “He Bourbon” [= Reunion] by Blom- mers-Schlosser and Blanc (1991). Ptyehadena maseareniensis hylaea Schmidt and Inger 1959:83). Holotype: FMNH 57965 (collector H.A. Beatty). Type locality: “Mount Nimba, Liberia” Liberia. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 135 Rana mascareniensis: Boulenger (1905:107). Rana (Ptychadena) mascareniensis'. Monard (1937a:50, 1938:108). Rana maseareniensis maseareniensis: Loveridge (1957:342), Ruas (1996:25, 2002:144). Ptyehadena maseareniensis'. Cei (1977:16, 17), Frost (1985:473), Charming (2001 329), Pickersgill (2007a: 128), Frost (2016), Conradie et al. (2016:18). Ptyehadena maseareniensis bibroni: Perret (1979:6). Ptyehadena hylaea: Lamotte (1967:647), Pickersgill (2007a: 134). Ptyehadena (Ptyehadena) maseareniensis'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: Populations from throughout sub-Saharan Africa currently referred to this species likely represent multiple undescribed species or should be referred to other described species in this genus. Ocurrences in Angola (Map 84): The species is widespread across the country. Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:50, 1938:108, Ruas 1996:25). Moxico: “Rio Caluando” [-11.46667, 17.70000] (Ruas 1996:25, 2002:144); “Lago Dilolo” [-11.50000, 22.01667] (Ruas 1996:25, 2002:144); “Lago Cameia” [-11.71667, 20.80000] (Ruas 1996:25, 2002:144). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt and Inger 1959:84; Ruas 1996:25). Huila: “Huila” [-11.71667, 20.80000] (Schmidt and Inger 1959:84). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937a:50, 1938:108; Ruas 1996:25). Cuando Cubango: “Cuabango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:9-10,18); “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9- . ^4. Distribution of Ptyehadena cf. mascareniensis 10,18); “Cubango basin (41a)” [-17.46777, 23.06667] (Conradie et al. 2016:9-10,18); “Cubango basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10,18); “Cubango basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10,18); “Cuito basin (63)” [-17.93611, 21.10269] (Conradie et al. 2016:9,12,18). Undetermined Locali¬ ty: “Between Benguella and Bihe (Entre Benguela e Bie)” (Boulenger 1905:107; Ruas 1996:25); “Angola Quest?” (Perret 1979:9); “areas of forest and savanna in the north and northeast of Ango¬ la” (Cei 1977:16); “arid territories along the coast” (Cei 1977:17); “plateaus regions” (Cei 1977:17). Taxonomic and distributional notes: Ptychadena mascareniensis is a complex of multiple cryptic forms, possibly representing as many as ten separate species (Vences et al. 2004; Zimkus et al. 2017) with the nominotypical form likely being restricted to Madagascar, the Seychelles, and the Mascarenes. While Zimkus et al. (2017) did not include Angolan populations in their molecu¬ lar phylogenetic study, it is likely that Angolan populations correspond to both P. nilotica and one undescribed species (their OTU 6) that occurs in Gabon and Democratic Republic of Congo. Schmidt and Inger (1959) described Ptychadena mascareniensis hylaea from Mount Nimba, Liberia and remarked on an individual (CNHM 74214) from “Huila in the highlands of southern Angola,” deposited in the Chicago Natural History Museum (now the Field Museum of Natural 136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II History), that they considered to belong to hylaea. Lamotte (1967) subsequently considered hylaea as a distinct species, whereas Ferret (1979) considered it to be a synonym of P. macare- niensis bibroni (Hallowell, 1845) (= Ptychadena bibroni) (Frost 1985). Fretey et al. (2011) synonymized hylaea with P. mascareniensis, while authors such as Pickersgill (2007a) continue to recognize the species. A taxonomic revision to determine the status of this species in the country is being conducted by R. Ernst (Conradie et al. 2016). Ptychadena cf. mossambica (Peters, 1854) Mozambique Grass Frog Rana mossambica Peters 1854:626. Syntypes: ZMB 4418, 4419a-b, 37555 (eolleetor W.C.H. Peters). Type loeality: “Cabeeeira, Queillimane, Tette, Boror” [= Cabeeeira, Quelimane (Companhia do Boror)] Mozam¬ bique. Ptychadena ef mossambica: Conradie et al. (2016:18). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Kenya south to Mozambique and KwaZulu-Natal, eastwards to Botswana, the Caprivi Strip of Namibia and adjacent south¬ eastern Angola. Ocurrences in Angola (Map 85): Possi¬ bly limited to the southeastern region of the country. Cuando Cubango: “lower Cuando River near the village of Jamba (38)” [-17.58830, 22.65694] (Conradie et al. 2016:9- 10.18) ; “lower Cuando River near the village of Jamba (39)” [-17.46333, 22.86638] (Con¬ radie et al. 2016:9-10,18); “lower Cuando River near the village of Jamba (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9- 10.18) . Taxonomic and distributional notes: Conradie et al. (2016) recently collected some individuals from Cuando Cubango Province Map 85. Distribution of Ptychadena cf mossambica in Angola. and provisionally assigned them to the Ptychadena mossambica complex. Ptychadena oxyrhynchus (Smith, 1849) Sharp-Nosed Grass Frog Rana oxyrhynchus Smith 1849a: pi. 77, figs. 2, 2a, 2b, 2c and two accompanying unumbered pages. Leeto- type: BMNH 58-11-25-97 designated by Guibe and Lamotte (1961:382) (eolleetor A. Smith). Type loeal¬ ity: “Cape of Good Hope” fide Guibe and Lamotte (1961’T960”:382), South Africa. According to Boulenger (1882:52) the specimens noted are from “Cape of Good Hope” and “Natal”, as also reported by Guibe and Lamotte (1961 “1960”:382). The original type locality is “Kaffir land and the country about Port Natal.” Rana oxyrhyneha: Bocage (1866a:53, 1870:68, 1895a: 159, 1897b:210), Ferreira (1900a:53, 1904:112, 1906:160), Schmidt (1936:129), Schmidt and Inger (1959:91). Rana oxyrhynehus: Boulenger (1882:51, 1905:108), Bocage (1887e:211), Rana (Ptyehadena) oxyrhynehus: Monard (1937a:49, 1938:107), Parker (1936:142). Rana oxyrhynehus oxyrhynehus: Mertens (1937a: 19, 1938a:427), Laurent (1950a: 14, 1954a:73). Rana {Ptyehadena) oxyrhynehus oxyrhynehus: Hellmieh (1957a:27). MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 137 Ptychadena oxyrhynchus: Guibe and Lamotte (1960 ‘T961”:382), Laurent (1964a: 133), Cei (1977:17), Frost (1985:474, 2016), Ruas (1996:25), Channing (2001:334), Cerlaco et al. (2016b:46). Ptychadena {Ptychadena) oxyrhynchus: Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is widely distributed in savannas across western, Central, and into southern Africa. Ocurrences in Angola (Map 86): The species occurs from western Angola to the northeastern regions. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 14, 1954a:73, 1964a;133; Ruas 1996:25); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a; 14, 1954a:73; Ruas 1996:25). Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:49, 1938:107; Ruas 1996:25). Moxico; “Teixeira de Sousa” [-10.70000, 22.23333] (Mertens 1937a: 19). Malanje: “Duque de Bra- ganga” [-9.10000, 15.95000] (Bocage 1866a:53, 1895a:159; Boulenger 1882:51, 1905:108; Ruas 1996:25); “Pungo-Andongo” [-9.66667, 15.58333] (Bocage 1895a;159; Ruas 1996:25), “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b;46). Kwanza Norte: “Rio Luinha” [-9.26667, Map 86. Distribution of Ptychadena oxyrhynchus in 14.53333] (Ferreira 1906:160; Ruas 1996:25); “N’dalla Tando (Dala Tando)” [-9.30000, 14.91667] (Ferreira 1904:112; Ruas 1996:25); “Cazen- go” [-9.33333, 14.76667] (Ferreira 1904:112; Ruas 1996:25); “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:27). Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] (Parker 1936:142; Ruas 1996:25): “Chingo” [-11.20000, 13.85000] (Ferreira 1904:112; Ruas 1996:25); “Novo Redondo, grotte de Fuma, fleuve N’Gunza (Rio N’Guanza)” [-11.20000, 13.93333] (Lau¬ rent 1954a:73; Ruas 1996:25). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1936:129; Ruas 1996:25); “Chitau” [-11.43333, 17.15000] (Schmidt 1936:129; Ruas 1996:25). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:107; Ruas 1996:25); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:142; Ruas 1996:25). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:159; Ruas 1996:25); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:159; Ruas 1996:25); “Quin- dumbo” [-12.46667, 14.93333] (Bocage 1895a:159; Ruas 1996:25); “Benguella” [-12.58333, 13.41667] (Bocage 1895a:159; Ruas 1996:25); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:27); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:427; Ruas 1996:25); “Hanha” [-13.30000, 14.20000] (Bocage 1897b:210; Ruas 1996:25). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:159; Ruas 1996:25); “Kalukembe” [-13.78333, 14.68333] (Monard 1937a:49, 1938:107; Ruas 1996:25); “Sangeve” [-13.88333, 15.83333] (Monard 1937a:49, 1938:107; Ruas 1996:25); “Indungu” [-14.81667, 16.26667] (Monard 1937a:49, 1938:107; Ruas 1996:25); “Osi” [-15.08333, 15.41667] (Monard 1937a:49, 1938:107; Ruas 1996:25); “Kakulaka- ze (Kului)” [-15.41667, 15.73333] (Monard 1937a:49, 1938:107; Ruas 1996:25). Cunene: “Rius- seau Mbale (Bale)” [-15.16667, 16.75000] (Monard 1937a:49, 1938:107; Ruas 1996:25). Cuando Cubango: “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9,18); “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9-10,18); “Cuando basin (41a)” 138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-17.46777, 23.06667] (Conradie et al. 2016:9-10,18); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10,18). Undetermined Locality: without precise locality (Bocage 1870:68; Boulenger 1900:53; Laurent 1954:73); “Rio Quando” (Bocage 1895a: 160; Ruas 1996:25); “Rio Quanza” (Boulenger 1905:108); “Carangigo” (Boulenger 1882:51); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:17); “arid territories along the coast” (Cei 1977:17). Taxonomic and distributional notes: This species occurs throughout much of sub-Saharan Africa in areas of moist savanna, near forest edges, in secondary vegetation with tall herbaceous vegetation, and in marshy and agricultura areas (Schmidt and Inger 1959; Ruas 1996; Rodel et al. 2004; Frost 2016). It appears to be relatively common in Angola, except in the arid and semiarid areas in southern regions of the country (Ruas 1996). According to Channing (2001) this species often occurs alongside angolensis (Bocage, 1866). Ptychadena perpUcata Laurent, 1964 Many-Ridged Grass Frog Ptychadena perpUcata Laurent 1964a: 136, fig. 36. Holotype: MD 5513 (collectors A. Barros Machado and E. Luna de Carvalho). Type locality: “Alto Chicapa, humidiherhosa des souces du Cuilo” [= Cuilo River] Lunda Sul Province, Angola. Ptychadena ansorgei: Laurent (1954a:74). Ptychadena perpUcata'. Cei (1977:16), Frost (1985:474, 2016), Ruas (1996:27), Largen (2000:177), Channing (2001:336). Ptychadena {Ptychadena) perpUcata'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola and Zambia. Ocurrences in Angola (Map 87): Pub¬ lished records are limited to the far north of the country. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:74, 1964a:136; Ruas 1996:27). Lunda Sul: “Alto Chicapa, humidi- herbosa des souces du Cuilo (Rio Cuilo, nascente)” [-10.88333, 19.23333] (Laurent 1964a: 136; Frost 1985:474, 2016; Ruas 1996:27, Largen 2001:177). Taxonomic and distributional notes: According to Ruas (1996) and Largen (2000), P. perpUcata is only known from forest gal¬ leries near the type locality. However, Chan¬ ning (2001) provided several records without detailed information, presumably based on museum specimens, that suggest this species is more widespread in the country. Ptychadena porosissima (Steindachner, 1867) Rana porosissima Steindachner 1867:18, pi. 1, figs. 9-13. Holotype: NHMW 14772 (collector F.A.P. Bayao) fide Haupl and Tiedemann (1978:28). Type locality: “Angola”. Ptychadena loveridgei Laurent 1954b: 14. Holotype: MRAC 109038. Type locality: “Tare, Busanza, region d’Astrida, env. 1. 800 m., Ruanda.” i:°E Ifi'E 20°E :4“E Map 87. Distribution of Ptychadena perpUcata in Angola. Striped Grass Frog MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 139 Rana porosissima: Bocage (1887a: 191, 1897b:211), Schmidt and Inger (1959:96), Haupl and Tiedemann (1978:28), Haupl et al. (1994:32). Rana mascareniensis: Bocage (1866a:53), Boulenger (1882:52). Rana mascareniensis vw. porosissima'. Bocage (1895a: 160), Ferreira (1897b:240). Rana subpunetata: Loveridge (1953a:329). Rana (Ptyehadena) maseareniensis: Guibe and Lamotte (1957:978). Rana loveridgei: Loveridge (1957:343). Ptyehadena porosissima'. Poynton and Broadley (1985b: 149), Frost (1985:474, 2016), Ruas (1996:25), Largen (2001:342), Channing (2001:337), Channing et al. (2012:311). Ptyehadena {Ptyehadena) porosissima'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is widely distributed in southern and eastern Africa. Ocurrences in Angola (Map 88): The species occurs from central-west regions to northeastern Angola. Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a:191, Bocage 1895a;160; Ruas 1996:25). Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Bocage 1895a:160; Ruas 1996:25). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1866a:53, 1895a: 160; Bou¬ lenger 1882:52; Ruas 1996:25). Lunda Norte: “Sombo” [-8.68333, 20.95000] (Laurent 1954b:14). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:160; Ruas 1996:25); “Hanha” [-13.30000, 14.20000] (Bocage 1897b:211; Ruas 1996:25). Huila: “Rio Cuce (Rio Cusse)” [-13.51667, 15.20000] (Ferreira 1897b:240; Ruas 1996:25); “Caconda” [-13.73333, 15.06667] (Bocage 1895a: 160; Ferreira 1897b:240; Ruas 1996:25). Taxonomic and distributional notes: Ptyehadena porosissima (Steindachner, 1867) was considered a variety of Rana mascareniensis (Dumeril and Bibron, 1841) by Bocage (1895a) and Ferreira (1897b) and later by Guibe and Lamotte (1957). Ptyehadena loveridgei Laurent, 1954 was synonymized with P. porosissima by Schmidt and Inger (1959). Loveridge (1953a) mistakenly considered P. porosissima a synonym of P. subpunetata, a position refuted by Schmidt and Inger (1959) who considered it a distinct species. Because of the similar general appearance of many species of Ptyehadena and the use of characters of limited diagnostic utility in some descriptions, many names have been misapplied to more than one species. In this case, some records allocated to the P. mascareniensis complex may belong to P. porosissima. Map Angola. Distribution of Ptyehadena porosissima in Ptyehadena subpunetata (Bocage, 1866) Spotted Grass Frog Rana subpunetata Bocage 1866b:73. Holotype: MBL (collector F.A.P. Bayao) not located by Perret (1967a:21), possibly lost or destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” [= Calandula] Malanje Province, Angola. Rana subpunetata'. Bocage (1866a:54, 1895a: 161, 1897a:203), Inger (1959:541). Rana maseareniensis subpunetata'. Mertens (1938a:427). Rana subpunetata'. Schmit and Inger (1959:102). 140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ptychadena subpunctata: Laurent (1964a: 134), Ferret (1976a:21), Cei (1977:16, 17), Poynton and Broadley (19856:143), Frost (1985:476, 2016), Ruas (1996:25, 2002:143), Channing (2001:340), Piekersgill (2007a: 136), Conradie et al. (2016:18). Ptychadena (Ptychadena) subpunctata'. Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola, extending east into the southern Democratic Republic of Congo and Zambia, and south to northern Botswana and northern Namibia. Ocurrences in Angola (Map 89): The species occurs from central-west to eastern areas of Angola. Malanje: “Duque de Bra- ganga” [-9.10000, 15.95000] (Bocage 1866a:54, 1866a:73, 1895a:161, 1897a:203; Ferret 1976a;21; Schmidt and Inger 1959:102; Poynton and Broadley 1985b: 143; Frost 1985:476, 2016; Ruas 1996:25). Moxico: “Lago Dilolo” [-11.50000, 22.01667] (Ruas 1996:25, 2002:143); “Lago Cameia” [-11.71667, 20.80000] (Ruas 1996:25, 2002:143); “Rives du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a: 134; Ruas 1996:25); “Chites de la Lusiavo, affl. du Zambeze, Calunda” [-11.86667, 23.58333] (Laurent 1964a: 134; Ruas 1996:25); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:134; Ruas 1996:25). Benguela: “Cubal” of Ptychadena subpunctata in [-13.03333, 14.25000] (Mertens 1938a:427; Ruas 1996:25). Cuando Cubango: “Cubango basin(6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9,18); “Cubango basin (27)” [-15.17127, 19.19433] (Conradie et al. 2016:8-9,18); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9-10,18); “Cuito basin (30f)” [-17.52638, 20.05825] (Conradie et al. 2016:9-10,18); “Cuito basin (33a)” [-16.90980, 19.30769] (Conradie et al. 2016:9-10,18); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9-10,18); “Cuando basin (41a)” [-17.46777, 23.06667] (Conradie et al. 2016:9-10,18); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10,18); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10,18); “Cuito basin (56)” [-15.06275, 19.14322] (Conradie et al. 2016:9,12,18); “Cuito basin (63)” [-17.93611, 21.10269] (Conradie et al. 2016:9,12,18). Unde¬ termined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:16); “Angola” (Poynton and Broadley 1985b: 143); “north Angola” (Schmidt and Inger 1959:140). Taxonomic and distributional notes: Rana subpunctata was used by Bocage (1866a) as a nomen nudum, but in the same issue of the journal, Bocage (1866b) published a valid description of the new species. According to Ferret (1976a), the holotype cited by Bocage from “Duque de Bragan^a” was not present in the Museu Bocage when he examined that collection. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I4I Ptychadena taenioscelis Laurent, 1954 Dwarf Grass Frog Ptychadena taenioscelis Laurent 1954b:25, pi. 5, fig. 1, pi. 4, fig. 6. Holotype: MRAC 13122 (collector G.F. de Witte). Type locality: “Lukula, pres de Kiambi, Tanganyika,” Democratic Republic of Congo. Ptychadena taenioscelis: Laurent (1964a: 140), Cei (1977:16), Poynton and Broadley (1985b: 153), Poynton and Haacke (1993:14), Ruas (1996:26), Channing (2001:341), Frost (2016), Conradie et al. (2016:18). Ptychadena pumilio: Largen (2001:342). Ptychadena {Ptychadena) pumilio: Fretey et al. (2011:41). Global conservation status (lUCN): Least Concern. Global distribution: The species is broadly distributed across southern, Central, and eastern Africa. Ocurrences in Angola (Map 90): The species occurs in the east of the country. Lunda Norte: “Riviere Luita, Poste de Cuilo (Rio Luita)” [-8.03333, 19.41667] (Laurent 1964a: 140; Ruas 1996:26). Lunda Sul: “Lua- chimo, 120 km au sud de Vila Henrique de Car¬ valho (Luachimo)” [-10.68333, 20.26667] (Laurent 1964a: 140; Ruas 1996:26); “Alto Chicapa, partie superieure des chutes du Cuan- go-Muque (Rio Cuango-Muque)” [-10.76667, 19.20000] (Laurent 1964a: 140; Ruas 1996:26); “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a: 140; Ruas 1996:26). Malanje; “20 km NW of Quimbango (Quimbango)” [-10.76667, 17.43333] (Poynton and Haacke 1993:14; Ruas 1996:26). Moxico: “Reserve de chasse de Cameia, fosse inonde pres de la route” [-11.83333, 21.00000] (Laurent 1964a: 140; Ruas 1996:26). Cuando Cubango: “Cubango basin (3)” [-14.94277, 17.71863] (Conradie et al. 2016:8-9,18); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9,18); “Cubango basin (19)” [-14.70213, 17.37772] (Conradie et al. 2016:8-9,18); “Cuito basin (30d)” [-17.51327, 20.06111] (Conradie et al. 2016: 9-10,18); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:19-10,18); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10,18); “Cuando basin (43b)” (Conra¬ die et al. 2016:9, 18); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10,18); “Cuito basin (62)” [-17.50875, 20.06608] (Conradie et al. 2016:9,12,18). Undetermined Locali¬ ty: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: Ptychadena taenioscelis Laurent, 1954 has occasional¬ ly been treated as a synonym of Ptychadena pumilio (Boulenger 1920). Perret (1979) suggested that P. taenioscelis is probably a subspecies of P. pumilio and referred records of P. taenioscelis from western and Central Africa to this species. Largen (2001) noted the occurrence of P. pumilio in Angola, as did Fretey et al. (2011) who included P. taenioscelis and P. smithi Guibe, 1960 as synonyms of P. pumilio. Further studies are needed to understand whether these are two distinct species or in fact are one and the same. However, we choose to treat this as a species distinct from Ptychadena pumilio, following Poynton and Broadley (1985b), Channing (2001), Pickersgill (2007a), and Frost (2016). Map 90. Distribution of Ptychadena taenioscelis Angola. 142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ptychadena upembae (Schmidt and Inger, 1959) Upemba Grass Frog Rana upembae Schmidt and Inger 1959: 111, fig. 50. Holotype: MRAC 1228 (collector Mission G.F. de Witte), formerly Inst. Parcs Natl. Congo Beige fide Lang (1990:13). Type locality: “Kaswabilenga, Parc National de I’Upemba, Province Katanga, Belgian Congo” (Schmidt and Inger 1959:111), Democratic Republic of Congo. Ptyehadena upembae maehadoi Laurent 1964a: 134. Holotype: MD 5364 (unnamed local collector). Type locality: “Alto Chicapa, Lunda, ’’Angola. Ptyehadena upembae maehadoi'. Cei (1977:16). Ptyehadena upembae'. Poynton and Broadley (1985b: 150), Frost (1985:477, 2016), Ruas (1996:26, 2002:144), Channing (2001:342). Ptyehadena (Ptyehadena) upembae'. Fretey et al. (2011:42). Global conservation status (lUCN): Least Concern. Global distribution: The species extends across south-central African from Angola, the southern Democratic Republic of Congo and Zambia to Mozambique. Ocurrences in Angola (Map 91): The species occurs in eastern Angola. Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a: 134; Ruas 1996:25). Moxico: “Lago Cameia” [-11.71667, 20.80000] (Ruas 1996:25, 2002:144). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Channing (2001) additionnally plotted unspecific locali¬ ties in a 2” map. Taxonomic and distributional notes: Currently P. upembae maehadoi is recognized as a synonym of Ptychadena upembae (Channing irE I6"E 2(}°E I4'>E Map 91. Distribution of Ptychadena upembae in Angola. 2001; Fretey et al. 2011, Frost 2016). Ptychadena uzungwensis (Loveridge, 1932) Udzungwa Grass Frog Rana maseareniensis uzungwensis Loveridge 1932a:384. Holotype: MCZ A-16626 (eolleetor A. Loveridge). Type loeality: “Dabaga, Uzungwe Mountains, Tanganyika Territory” [Tanzania]. Rana maseareniensis subpunetata'. Schmidt (1936:129). Ptyehadena uzungwensis'. Laurent (1954b:10, 1964a:139), Cei (1977:16), Poynton and Broadley (1985b:151), Poynton and Haacke (1993:14), Frost (1985:477, 2016); Ruas (1996:26, 2002:145), Channing (2001:344^ Conradie et al. (2016:18). Rana maseareniensis uzungwensis: Loveridge (1953a:372, 1957:342). Rana uzungwensis: Schmidt and Inger (1959:117). Ptyehadena subpunetata: Ruas (1996:25). Ptyehadena (Ptyehadena) uzungwensis: Fretey et al. (2011:42). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from central eastern Angola and the southeastern Democratic Republic of Congo, Zambia and Tanzania south to northern Zimbabwe. Ocurrences in Angola (Map 92): The species occurs chiefly in eastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954b: 10). Lunda Sul: “Alto Chicapa, hemidi- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 143 herbosa pres de la cascade du Cuango-Muque” [-10.76667, 19.20000] (Laurent 1964a:139); “Alto Chicapa, Lunda hemidiherbosa des sour¬ ces du Cullo” [-10.86667, 19.40000] (Laurent 1964a: 139); “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:139); “Dala” [-11.03333, 20.20000] (Laurent 1964a:139). Malanje: “20 km NW of Quimbango” [-10.76667, 17.43333] (Poynton and Haacke 1993:14). Mexico: “Calombe” [-11.83333, 19.93333] (Ruas 2002:145); “Reserve de chas- se de Cameia dans la steppe de Narikumbi” [-11.83333, 21.00000] (Laurent 1964a:139). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1936:129; Laurent 1954b:9; Schmidt and Inger 1959:118); “Chitau” [-11.43333, 17.15000] (Schmidt 1936:129; Laurent 1954b:9; Schmidt and Inger 1959:118). Cuando Cubango: “Cuito basin (27)” [-15.17127, 19.19433] (Conradie et al. 2016:8-9,18). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola”(Cei 1977:16). Taxonomic and distributional notes: Loveridge (1953a) believed thaX Rana mascareniensis uzungwensis Loveridge, 1932 was erroneously synonymized with Rana subpunctata (Bocage, 1866) by Schmidt (1936) and cited specimens from “Gauca” and “Chitau” in Angola as belonging to P. uzungwensis. This species is considered valid by the majority of modem authors (e.g., Poyn¬ ton and Broadley 1985b; Channing 2001; Frost 2016) and according to Poynton and Broadley (1985b) it inhabits grasslands and dambos. Poynton and Haacke (1993) reported one individual from “20 km NW of Quimbango” collected by G. G. A. Voigt in a water-logged grassland. Chan¬ ning (2001) provided several records in the western regions of the country based on previously unpublished museum records. Family Phrynobatrachidae Laurent, 1941 Genus Phrynobatrachus Gunther, 1862 Phrynobatrachus brevipalmatus (Ahl, 1923) Ahl’s Screeching Frog (Endemic) Hylarthroleptis brevipalmatus Ahl “1923” 1925:102. Holotype: ZMB 26689 (collector K. May). Type local¬ ity: “Loanda” [= Luanda] Luanda Province, Angola. Phrynobatrachus brevipalmatus'. Fretey et al. (2011:38), Frost (2016). Global conservation status (lUCN): Data Deficient. Global distribution: The species is only known from Angola. Ocurrences in Angola (Map 93): The species is known from Luanda. Luanda: “Loanda” [-8.83333, 13.26667] (Ahl “1923” 1925:102; Frost 2016). Taxonomic and distributional notes: According to the original description, this species is closely related to Hylarthroleptis graueri (Nieden, 1911) from eastern Africa. This species was not discussed by Zimkus et al. (2010) and no further details are available for its biology and systematics. 1I°E I6'E irE 24°E Map 92. Distribution of Ptychadena uzungwensis in Angola. 144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 12°E lO^E 20°E 24®E. Map 93 . Distribution of Phrynobatrachus brevipalmatus in Angola. Phrynobatrachus cryptotis group Given the considerable nomenclatural and taxonomic confusion regarding P. cryptotis, P mababiensis, P minutus and P. parvulus, we have opted to combine all of their distributional and taxonomic notes into the same account, although, specific accounts for each one, with the list of chresonyms and type specimen data still follow below. Records attributed to each species are provided below separately, indicated by the name and noting the different symbols used to plot the records on the combined map. Ocurrences in Angola (Map 94): Phrynobatrachus cryptotis (plotted as cir¬ cles). Lnnda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a: 144; Ruas 1996:27); “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a: 144; Ruas 1996:27). Hulla: “Boca de Humpata” [-14.93333, 13.51667] (Laurent 1964a: 144; Ruas 1996:27). Undetermined Locality: “arid territories along the coast” (Cei 1977:18). Phrynobatrachus mababiensis (plotted as triangles). Bie: “Cuando basin (11)” [-13.69413, 17.06177] (Conradie et al. 2016:8-9,16); “Cuando basin (12a)” [-13.59333, 16.87986] (Conradie et al. 2016:8- 9,16). Huambo: “Cuando basin (18)” [-12.67105, 16.11111] (Conradie et al. 2016:8- 9,16). Hulla: “Nuntechite lagoon” [-15.13333, 13.41667] (Poynton and Haacke 1993: 14; Ruas 1996:27). Cuando Cubango: “Cuando basin (3)” [-14.94277, 17.71863] (Conradie et al. 12°E lO'E 20°E 24''E Map 94. Distribution of Phrynobatrachus cryptotis group in Angola. 2016:8-9,16); “Cuando basin (6b)” [-14.67458, MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 145 17.73544] (Conradie et al. 2016:8-9,16); “Cuando basin (19)” [-14.70213, 17.37772] (Conradie et al. 2016:8-9,16); “Cuando basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:8-9,16); “Cuan¬ do basin (22e)” [-14.65386, 16.93547] (Conradie et al. 2016:8-9,16); “Cuando basin (22d)” [-14.64991, 16.90739] (Conradie et al. 2016:8-9,16); “Cuando basin (29)” [-17.87291, 19.833333] (Conradie et al. 2016:9-10,16); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016: 9-10,16); “Cuando basin (38)” [-17.58830, 22.65694] (Conradie et al. 2016:9-10,16); “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9-10,16); “Cuando basin (40)” [-17.45786, 22.91191] (Conradie et al. 2016:16); “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10,16); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10,16). Phrynobatrachus minutus (plotted as pentagons). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:74; Ruas 1996:27); “galerie forestiere de la Luaehimo (Lua- chimo)” [-7.38333, 20.85000] (Laurent 1950a: 15; Ruas 1996:27); “Marrura rive droite de la Tshi- kapa, 50 km sud-oest de Dundo (Marrora)” [- 7.60000, 20.51667] (Laurent 1954a:74; Ruas 1996:27). Cunene: “Donguena (Cunene)” [-17.01667, 14.71667] (Laurent 1954a:74; Ruas 1996:27). Undetermined Locality: “Station Quanza (Esta^ao do Quanza)” (Laurent 1954a:74; Ruas 1996:27); with no preeise loeality (Laurent 1954a:74); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:17). Phrynobatrachus parvulus (plotted as diamonds). Lunda Sul: “Alto Cuilo, Poste de Caeolo, ruisseau cascade Na- Ipanha” [-10.00000, 19.58333] (Laurent 1964a:144); “Alto Cuilo, rives du Cuilo (Cuilo)” [-10.01667, 19.55000] (Laurent 1964a: 144); “Alto Chicapa, humidiherbosa des sources de la Kamutongola” [-10.88333, 19.25000] (Laurent 1964a: 144). Moxico: “Teixeira de Sousa” [-10.70000, 22.23333] Mertens (1937:20). Malanje: “Range N’golla (Dange)” [-8.43333, 16.56667] (Boulenger 1905:109; Loveridge 1933:386; Marx 1958:425; Schmidt and Inger 1959:160; Poynton and Broadley 1985b:169; Frost 1985:449, 2016; Ruas 1996:27). Kwanza Sul: “Calaongo below Congulu (Carlaongo)” [-10.78333, 14.26667] (Parker 1936:142; Ruas 1996:27). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1936:131; Ruas 1996:27); “Chitau” [-11.43333, 17.15000] (Schmidt 1936:131, Ruas 1996:27); “General Machado” [-12.03333, 17.46667] Mertens (1937:20). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:58, 1938:118, Ruas 1996:27). Benguela: “Cubal” [-13.03333, 14.25000] Mertens (1937:20). Cuando Cubango: “Cuando basin (43)” [-17.53500, 23.189916] (Conradie et al. 2016:9-10,17). Undetermined Locality: “without precise locality” (Schmidt 1936:131). Taxonomic and distributional notes: Opinions in the literature vary as to whether three commonly reported species from Angola are conspecific: P. mababiensis FitzSimons, 1932, P. parvulus (Boulenger, 1905) and P. minutus (Boulenger, 1895). Loveridge (1953) assigned a specimen previously identified as P. parvulus to P. mababiensis, suggesting that P. parvulus was a western relative of P. mababiensis or perhaps a synonym of Phrynobatrachus minutus. Poynton and Broadley (1985b:165, 170) discussed difficulties in separating P. parvulus and P. mababiensis based on external morphology. Zimkus et al. (2010) recently used molecular phylogenetic analy¬ ses to reveal that there are probably three different species currently recognized as P. mababiensis. If P. minutus is known only from Ethiopia, it is highly likely that the Angolan records assigned to this species are misidentified and instead belong to one of the other three species. With some reser¬ vations, Laurent (1964a) cited P. cryptotis for Angola, though he recognized that these might represent P. parvulus instead, which is a similar species (Drewes and Vindum 1994). It is likely that the Angolan specimens mapped here represent a combination of P. mababiensis and P. parvulus. 146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Phrynobatrachus cryptotis Schmidt and Inger, 1959 Cryptic River Frog Phrynobatrachus cryptotis Schmidt and Inger 1959:143, fig. 60, pi. 5, fig. 5. Holotype: IRSNB 1.493 (col¬ lector G.F. de Witte), formerly Institut des Parcs Nationaux du Congo Beige n. 1926. Type locality: “upper Bwalo River (an affluent from the left in on the left of the Muye, which is an affluent from the right of the Lufira)” (Schmidt and Inger 1959:143), [= Parc National de PUpemba, upper Katanga], Democratic of Republic of Congo. Phrynobatrachus cryptotis'. Laurent (1964a: 144), Cei (1977:18), Frost (1985:445, 2016), Ruas (1996:27), Fretey et al. (2011:39). Global conservation status (lUCN): Data Deficient. Global distribution: It is known with certainty only from the Democratic Republic of Congo. Phrynobatrachus mababiensis FitzSimons, 1932 Mababe Puddle Frog Phrynobatrachus mababiensis FitzSimons 1932:40. Holotype: TM 14838, formerly Vemay-Lang Khalari Expedition no. 989 (collector V.F. FitzSimons). Type locality: “Tsotsoroga Pan, Mababe Flats” (FitzSi¬ mons 1932:40), South Africa. Phrynobatrachus mababiensis'. Poynton and Broadley (1985b: 165), Forst (1985:447, 2016), Poynton (1992:67), Poynton and Haacke (1993:14), Ruas (1996:27), Channing (2001:305), Channing and Howell (2006:284), Zimkus et al. (2010:42), Fretey et al. (2011:39), Channing et al. (2012:278), Conradie et al. (2016:16). Global conservation status (lUCN): Least Concern. Global distribution: This species occurs in savannas across southern and central Africa. Phrynobatrachus minutus (Boulenger, 1895) Ethiopian Dware Puddle Frog Arthroleptis minutus Boulenger 1895a:539, pi. 30, fig. 4. Holotype: BMNH 1947.2.30.51 (collector A.D. Smith), formerly 95.6.11.8 fide Grandison in Frost (1985:448). Type locality: “Durro” (Boulenger 1895a: 539), [= Duro], Ethiopia. Arthroleptis minutus'. Monard (1938:118). Phrynobatraehus minutus: Laurent (1950a: 15, 1954a:74), Loveridge (1957:349), Inger (1959:540), Cei (1977:16, 17), Ruas (1996:27), Fretey et al. (2011:39), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: This species is likely limited to Ethiopia. Phrynobatrachus parvulus (Boulenger, 1905) Dware River Frog Arthroleptis parvulus Boulenger 1905:109, pi. 4, figs; 3, 3b. Syntypes: BMNH 1947.2.6.93-95 [3 specimens], formerly 1904.5.2.97-99 and FMNH 75381 (collector W.J. Ansorge) fide Grandison in Frost (1985:449). Type locality: “Bange N’golla” (Boulenger 1905:119), [= Dange] Malanje Province, Angola. Arthroleptis parvulus: Loveridge (1933:386), Schmidt (1936:131), Parker (1936:142), Mertens (1937:20). Arthroleptis minutus: Monard (1937a:58, 1938:118). Phrynobatraehus parvulus: Marx (1958:425), Schmidt and Inger (1959:160), Laurent (1964a: 144), Poynton and Broadley (1985b: 169), Frost (1985:449, 2016), Poynton (1992:67), Ruas (1996:27), Channing (2001:308), Channing and Howell (2006:289), Fretey et al. (2011:40). Phrynobatraehus cf. parvulus: Conradie et al. (2016:17). Global conservation status (lUCN): Least Concern. Global distribution: This species occurs in savannas across southern and central Africa. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 147 Phrynobatrachus natalensis (Smith, 1849) Natal Dwarf Puddle Frog Stenhorhynchus natalensis Smith 1849b:24. Holotype: BMNH 1947.2.5.13 (formerly BMNH 62.3.14.20) (collector A. Smith) fide Grandison in Frost (1985:448). Type locality: “the country around Port Natal” [= Durban, KwaZulu-Natal], South Africa. Phrynobatraehus natalensis: Gunther (1865a:481), Bocage (1866a:54, 1870:68, 1879c:89, 1895a:162, 1897b:211), Boulenger (1882:112, 1905:108), Ferreira (1904:113, 1906:166), de Witte (1919:223), Schmidt (1936:130), Monard (1937a:57, 1938:117), Mertens (1937a:20), Parker (1939:142), Laurent (1950a: 15, 1954a:74, 1964a: 143), Loveridge (1957:348), Hellmich (1957a:24), Schmidt and Inger (1959:155), Inger (1959:540), Cei (1977:16, 17, 18), Poynton and Haacke (1993:14), Ruas (1996:27, 2002:145), Largen (2001:348), Channing (2001:307), Zimkus et al. (2010:886), Fretey et al. (2011:39), Frost (2016), Ceriaco et al. (2016b:39), Conradie et al. (2016:17). Phrynobatraehus cf natalensis: Ceriaco et al. (2014b:669). Global conservation status (lUCN): Least Concern. Global distribution: Populations referred to P. natalensis occur across sub-Saharan Africa, including in Angola, though this taxon likely contains multiple undescribed species. Ocurrences in Angola (Map 95): The species is widespread across nearly all of Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a; 15, 1954a:74; Ruas 1996:27); “Andrada (Luembe O)” [-7.70000, 21.38333] (Laurent 1954a;74; Ruas 1996:27); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 15, 1954a:74; Ruas 1996:27). Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Alto Cuilo, rives du Cuilo (rio Cuilo)” [-10.01667, 19.55000] (Laurent 1964a:143; Ruas 1996:27). Malanje: “Bange N’golla (Dange)” [-8.43333, 16.56667] (Boulenger 1905:108; Ruas 1996:27); “Duque de Bragan- ga” [-9.10000, 15.95000] (Bocage 1866a:54, 1895a: 162; Gunther 1865a:481; Boulenger 1882:112; de Witte 1919:223; Poynton and Haacke 1993:14; Ruas 1996:27); “16 km SE of Quissol - Garibo” [-9.63333, 16.41667] (Poynton and Haacke 1993:14; Ruas 1996:27); “Cangan- dala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:39); “Capanda” [-11.58333, 17.61667] (Ceriaco et al. 2014b:669); “ Reserva da Palanca Preta (nascentes do rio Cuanza)” [-11.11667, 17.46667] (Ruas 2002:145); “Rio Caluando (nascente)” [-11.11667, 17.46667] (Ruas 1996:27, 2002:145). Mexico: “Teixeira de Sousa” [-10.70000, 22.23333] (Mertens 1937:20); “Posto Bussaco” [-11.43333, 20.16667] (Ruas 2002:145; Ruas 1996:27); “Rives du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a: 143; Ruas 1996:27); “Rio Calombe (Reserva da Palanca Preta)” [-11.83333, 19.93333] (Ruas 1996:27, 2002:145); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:143; Ruas 1996:27). Kwanza Norte: “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:24). Bengo: “Catete” [-9.11667, 13.70000] (Ferreira 1904:113; Ruas 1996:27). Kwanza Sul: “Congulu (Congulo)” [-10.86667, 14.28333] (Parker 1936:142; Ruas 1996:27). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1936:130; Ruas 1996:27); “Chitau” [-11.43333, 17.15000] (Schmidt 1936:130; Ruas 1996:27); “General Map 95. Distribution of Phrynobatrachus natalensis in Angola. 148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Maehado” [-12.03333, 17.46667] (Mertens 1937a:20); “Bihe” [-12.38333, 16.95000] (Bocage 1879a:89, 1895a: 162; de Witte 1919:223; Ruas 1996:27); “Cubango basin (10)” [-13.71616, 17.09661] (Conradie et al. 2016:8-9,17); “Cubango basin (12a)” [-13.59333, 16.87986] (Conradie et al. 2016:17); “Cubango basin (17)” [-12.57008, 16.49111] (Conradie et al. 2016:8-9,17). Huam- bo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:57, 1938:117; Ruas 1996:27). Benguela: “Quissange” [-12.43333, 14.05000] (Boeage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Ebanga” [-12.73333, 14.73333] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:24); “Hanha” [-13.30000, 14.20000] (Bocage 1897b:211; Ruas 1996:27). Hufla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:162; de Witte 1919:223; Ruas 1996:27); “Kalukembe” [-13.78333, 14.68333] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Sangeve” [-13.88333, 15.83333] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Muko- ti” [-14.20000, 15.80000] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Kuvangu” [-14.46667, 16.30000] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Boca de Humpata” [-14.93333, 13.51667] (Laurent 1964a:143; Ruas 1996:27); “Kangela (Kului)” [-15.41667, 15.73333] (Monard 1937a:57; 1938:117; Ruas 1996:27). Namibe: “Cunene mouth (Foz do Cune- ne)” [-17.28333, 11.80000] (Poynton and Haacke 1993:14; Ruas 1996:27). Cunene: “ruisseau Mbale (Bale)” [-15.16667, 16.75000] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Kuvelai” [-15.65000, 15.80000] (Monard 1937a:57, 1938:117; Ruas 1996:27); “Donguena” [-17.01667, 14.71667] (Laurent 1954a:74; Ruas 1996:27). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1938:117; Ruas 1996:27); “Cubango basin (3)” [-14.94277, 17.71863] (Conradie et al. 2016:8-9,17); “Cubango basin (6b)” [-14.67458, 17.73544] (Conradie et al. 2016:8-9,17); “Cubango basin (19)” [-14.70213, 17.37772] (Conradie et al. 2016:8-9,17); “Cubango basin (21)” [-14.66586, 17.07661] (Conradie et al. 2016:17); “Cubango basin (22d)” [-14.66278, 16.96081] (Conradie et al. 2016:17). Undetermined Locality: without precise locali¬ ty (Bocage 1870:68; Ferreira 1906:166; Schmidt 1936:130); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:17); “arid territories along the coast” (Cei 1977:18). Taxonomic and distributional notes: Schmidt and Inger (1959) provided a distribution map for Phrynobatrachus natalensis (Smith, 1849) with some records in Angola, though without addi¬ tional information. The molecular phylogenetic analysis by Zimkus et al. (2010) VQYQdlQd Phryno¬ batrachus natalensis to be a species complex, with Angolan records likely corresponding to their lineage ''natalensis A.” Phrynobatrachus plicatus (Gunther, 1858) Coast River Frog Hyperolius plicatus Gunther 1858a:326. Holotype: BMNH 1947.2.29.97 [wrongly given as 1947.2.29.47 by Grandison in Frost (1985)], formerly BMNH 49.10.9.1 fide Grandison in Frost (1985:449) (eolleetor unknown). Holotype is depleted in Gunther ‘T858b” 1859:8, pi. 7, fig. C. Type loeality: “Guinea” (Giin- ther 1858a:326), later modified to “Coast of Guinea” (Gunther ‘T858b” 1859:88). Arthroleptis plicatus: Peters (1877a:618). Phrynobatrachus plicatus: Zimkus et al. (2010:896), lUCN SSC Amphibian Speeialist Group (2013e), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: This is a widespread species from western Africa. Ocurrences in Angola (Map 96): The only record of this species in Angola is from the Cabin¬ da enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:618). Taxonomic and distributional notes: The record of this species in Angola is almost certain- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 149 ly a misidentification as today the recognized distribution of this species is restricted to west¬ ern Africa. It is possible that the Cabinda frog may be referable to Phrynobatrachus auritus Boulenger, 1900. i:°E l(i®E 20°:E I4"E Map 96. Distribution of Phrynobatrachus plicatus in Angola. Family Pyxicephalidae Bonaparte, 1850 Genus Amietia Dubois, 1987 Amietia angolensis (Bocage, 1866) Angola River Frog Rana angolensis Bocage 1866b:73. Lectotype: MBL T.8-94 designated by implieation by Ferret (1976a: 18), having listed syntype as holotype. Destroyed by fire 18 Mareh 1978. Type loeality: “Duque de Braganga” (Boeage 1866b:73), [= Calandula] Malanje Provinee, Angola. Rana angolensis: Boeage (1866a:54; 1866b:73, 1887e:211, 1895a:158, 1897a:203) Ferreira (1903:111), Boulenger (1882:50, 1905:108), Themido (1941:2), Laurent (1964a: 132), Ferret (1976a: 18), Cei (1977:16, 17), Poynton and Broadley (1985b: 131), Frost (1985:480), Gavetti and Andreone (1993:91), Poynton and Haaeke (1993:14), Drewes and Vindum (1994:64), Ruas (1996:23, 2002:143), Largen (2001:331). Rana Delalandi: Boeage (1870:68). Rana ehapini: Noble (1924:214). Rana {Rana) angolensis: Monard (1937a: 43, 1938:99). Rana fuseigula angolensis: Loveridge (1933:362, 1936b:410, 1953a:365, 1957:339), Sehmidt (1936:128), Mertens (1937a: 19, 1938a:426), Parker (1939:141), Laurent (1950a: 14, 1954a:71), Hellmieh (1957a:25). Rana fuseigula: Inger (1959:540), Sehmidt and Inger (1959:48). Afrana angolensis: Channing (2001:255), Channing and Howell (2009:250), Piekersgill (2007a:98), Fretey et al. (2011:42). Amietia angolensis: Channing et al. (2012:318), Channing and Baptista (2013:501), Frost (2016), Channing et al. (2016:15). Amietia ef angolensis: Ceriaeo et al. (2016a:20). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from western central Africa, extending from south¬ ern Congo and the Democratic Republic of Congo south through Angola and possibly in the Caprivi Strip of Namibia. 150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 97): The speeies is found throughout Angola, especially in the northern and western regions. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1950a:14, 1954a:71; Ruas 1996:23); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1954a:71; Ruas 1996:23); “rive de la Tshimumbwe, 40 km a Test de Dundo (rio Chiumbe)” [-8.01667, 19.31667] (Laurent 1950a: 14; Ruas 1996:23); “Tshinguvu gallerie forestiere de la Tshikapa, 50 km au sud-ouest de Dundo (Chingufo)” [-8.01667, 19.31667] (Laurent 1950a: 14; Ruas 1996:23). Lunda Sul: “Alto Cuilo, rives du Cuilo, (rio Cuilo)” [-10.01667, 19.55000] (Laurent 1964a: 132; Ruas 1996:23); “Alto Chicapa, partie superieur des chutes du Cuango-Muque (rio Cuango- Muque)” [-10.76667, 19.20000] (Laurent Map 97. Distribution of in An^ 1964a:132; Ruas 1996:23); “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:132; Ruas 1996:23); “Dala” [-11.03333, 20.20000] (Monard 1937a:43, 1938:99; Ruas 1996:23). Mexico: “Calombe (Luso)” [-11.83333, 19.93333] (Ruas 1996:24, 2002:143); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:132 Ruas 1996:23); “Calunda, Alto Zambeze)” [-12.11667, 23.46667] (Laurent 1964a: 132; Ruas 1996:23). Uige: “Serra do Pingano” [-7.68528, 14.92956] (Channing et al. 2016:16, 66); “Quilomosso” [-7.6599, 149821] (Channing et al. 2016:16, 66). Malanje: “Duque de Bragan^a (Calandula)” [-9.06667, 17.90000] (Bocage 1866a:54, 1866b:73, 1895a:158, 1897a:203; Loveridge 1933:362, 1936b:410; 1953a:365, 1957:339; Perret 1976a:18; Poynton and Broadley 1985b: 131; Frost 1985:480, 2016; Ruas 1996:23; Largen 2001:331; Pickersgill2007a:98, Channing and Baptista 2013:508; Channing et al. 2016:16); “Pungo-Andongo” [-9.66667, 15.58333] (Bocage 1895a:158; Boulenger 1905:108; Ruas 1996:23). Kwanza Norte: “N’dalla Tando (DalaTando)” [-9.30000, 14.91667] (Ferreira 1903:111; Ruas 1996:23); “Ndalatando Bota¬ nical Garden” [-9.33422, 14.89895] (Channing et al. 2016:16, 66); “Zembe” [-9.350000, 14.683333] (Ferreira 1903:111; Ruas 1996:23). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:141; Ruas 1996:23); “Congulu” [-10.86667, 14.28333] (Parker 1936:141; Ruas 1996:23). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:99 Ruas 1996:23); “Galanga” [-12.06667, 15.15000] (Bocage 1895a:158; Ruas 1996:23); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:141; Ruas 1996:23); “Santo-Amaro” [-12.70000, 15.85000] (Monard 1937a:43, 1938:99; Ruas 1996:23); “Cubango basin (18)” [-12.67105, 16.11111] (Conradie et al. 2016:18). Bie: “Gauca” [-11.18333, 17.45000] (Schimdt 1936:128; Ruas 1996:23), “Chitau” [-11.43333, 17.15000] (Schimdt 1936:128; Ruas 1996:23; Channing et al. 2016:16); “General Machado” [-12.03333, 17.46667] (Mertens 1937a:19); “Cubango basin (11)” [-13.69413, 17.06177] (Con¬ radie et al. 2016:18); “Cubango basin (12a)” [-13.59333, 16.87986] (Conradie et al. 2016:18.19); “Cubango basin (13)” [-13.28061, 16.74722] (Conradie et al. 2016:18); “Cubango basin (14)” [-12.87242, 16.76742] (Conradie et al. 2016:19). Benguela: “Benguela” [-12.58333, 13.41667] (Loveridge 1936:90; Channing et al. 2016:16); “Quibula” [-12.28333, 14.68333] (Bocage 1895a:158; Ruas 1996:23); “Quissange” [-12.43333, 14.05000] (Bocage 1887c:211, 1895a:158; Perret 1976a:18; Ruas 1996:23); “Entre-Rios” [-13.01667, 14.63333] (Hellmich 1957a:25); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:426; Ruas 1996:23). Huila: “Monguaval farm” MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I5I [-13.45000, 14.61667] (Poynton and Haacke 1993:14; Ruas 1996:23); “Tongrube am Jamba-FluP” [-13.60000, 16.60000] (Hellmich I957a:25); “Caconda” [-13.73333, 15.06667] (Bocage I895a:I58; Perret 1976:18; Gavetti and Andreone 1993:91; Ruas 1996:23); “Kalukembe” [-13.78333, 14.68333] (Monard I937a:43, 1938:99; Ruas 1996:23); “Sangeve” [-13.88333, 15.83333] (Monard I937a:43, 1938:99; Ruas 1996:23); “Kuvangu” [-14.46667, 16.30000] (Monard I937a:43, 1938:99; Ruas 1996:23); “Boca de Humpata” [-15.01667, 13.38333] (Laurent 1964a: 132; Ruas 1996:23; Channing et al. 2012:318; Channing and Baptista 2013:508); “Zootec- nica Station, Humapata” [-I4.9I4I7, 13.31653] (Channing et al. 2016:16, 66); “Humapata eamp” [-14.95000, 13.26667] (Channing et al. 2016:16, 66); “Huila” [-15.05000, 13.55000] (Boeage 1895a:158; Themido 1941:2; Perret 1976a:18; Ruas 1996:23). Namibe: “Lebapass, between river and highway” [-15.07006, 13.24414] (Ceriaeo et al. 2016a:20). Cuando Cubango: “Cubango basin (4)” [-14.81913, 17.67450] (Conradie et al. 2016:18); “Cubango basin (22d)” [-14.64991, 16.90739] (Conradie et al. 2016:18); “Cuando Cubango Provinee” [-14.64991, 18.4672] (Chan¬ ning et al. 2016:16). Undetermined Locality: “without precise locality” (Bocage 1870:78); “Rio Quando” (Bocage 1895a: 158; Ruas 1996:23); “areas of forest and savanna in the north and north¬ east of Angola” (Cei 1977:16); “plateaus regions” (Cei 1977:16); “arid territories along the coast” (Cei 1977:16); “Mombola” (Channing et al. 2016:16). Taxonomic and distributional notes: Many authors, including Poynton (1964), Drewes and Vindum (1994), and Channing and Howell (2006) recognized that there are probably several unde¬ scribed cryptic species within the enormous range of this taxon. Pickersgill (2007a) named three new species from Eastern Africa populations that were previously included within Amietia angolensis. Channing and Baptista (2013) and Channing et al. (2016) restricted nominal A. angolensis to Angolan populations, though recognized that it may occur more broadly, espe¬ cially in neighboring countries for which they lacked specimens with genetic data. Genus Boulenger, 1917 Aubria sp. Rana subsigillata Dumeril 1856:560. Holotype: MNHN 1566 (collector J. Aubry-Lecomte). Type locality: “Gabon” (Dumeril 1856:560). Aubria masako Ohler and Kazadi 1990:29, figs. 4, 8, 10, 12, 24. Holotype: MNHN 1989.2775 (collector M. Kazadi). Type locality: “Foret de Masako pres du village Batiabongena a 15 km du centre- ville de Kisangani sur I’ancienne route Buta (Zaire)” (Ohler and Kazadi 1990:29), [= Masako forest, near Batiabongena village, 15 km from Kisangani], Democratic Republic of Congo. Rana {Aubria) subsigillata: Monard (1937a:47, 1938:104). Aubria subsigillata: Cei (1977:17), Perret (1996:96). Aubria masako: Channing (2011:284), Fretey et al. (2011:42), Channing et al. (2012:329), Frost (2016). Global distribution: Aubria masako and A. subsigillata are known from the tropical forests of central Africa, with ranges that together extend from the Atlantic coast of Cameroon and Gabon and into the Congo Basin of Democratic Republic of Congo. Ocurrences in Angola (Map 98): Both Aubria subsigillata and A. masako have only been reported from “Kakindo (Kuvangu)” in southern Angola. Cuando Cubango: “Kakindo (Kuvan¬ gu)” [-14.46667, 16.30000] (Monard 1937a:47, 1938:104; Channing 2001:285). Taxonomy and natural history notes: Monard (1937a, 1938) cited one specimen of Aubria subsigillata (Dumeril, 1856) from “Kakindo (Kuvangu),” Angola that was identified by Gaston de Witte. This first and only record for the genus in Angola is unlikely due to the habitat, a dry savan¬ na, and its long distance from congeneric populations in central or western Africa. Perret (1996) revisited this specimen and cited morphological differences from A. subsigillata in Cameroon to 152 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II suggest that it might represent a new species or even a new genus. It remains unclear whether this record indeed corresponds to Aubria and, if so, to which species it should be referred. The specimen cited by Monard (1937a, 1938) was recently located in the collections of the Musee d’Histoire Naturelle, La-Chaux-de-Fond, Switzerland and is presently being studied (Ceriaco et al. in prep.). 12°E l(j®E 2()°E 24'*E Map 98. Distribution oi Aubria sp. in Angola. Genus Pyxicephalus Tschudi, 1838 Pyxicephalus edulis Peters, 1854 Edible Bullfrog Pyxicephalus edulis Peters 1854:626. Leetotype: ZMB 50301 (formerly part of ZMB 10056) (eolleetor: W.C.H. Peters) designated by Seott et al. (2013:201). Type loeality: restrieted to “Tete,” Mozambique by Loveridge (1953:375). The original syntype series ineluded ZMB 3349, 3350 [4 speeimens from Mozam¬ bique], 10056 [2 speeimens (Tete)], and 10057 [2 speeimens (Sena)]. Bauer et al. (1995:49) mentioned ZMB 10058 (not loeated), 50260, 50290, and 50301-02, formerly ineluded under the earlier numbers, one of these may be the BMNH speeimen from “Mossambique” mentioned by Boulenger (1882:34) as “typi- eal of P. edulis"' and reeeived from W. Peters. Original type locality: “Mosambique, Boror, Tette” (Peters 1854:626), [= Mozambique Island and adjacent mainland, Boror, Tete] Rana adspersa: Bocage (1895a: 157), Inger (1959:541). Rana (Pyxieephalus) adspersus: Monard (1937a:46, 1938:103), Frade (1963:254). Pyxieephalus adspersus: Cei (1977:17), Frost (1985:477), Channing (2001:346), du Preez and Camthers (2009:414), Channing et al. (2012:123). Pyxieephalus adspersus edulis: Loveridge (1950:255, 1953a:375), Parry (1982:286), Poynton and Broadley (1895b: 123), Poynton and Haacke (1993:13), Ruas (1996:23). Pyxieephalus edulis: Channing et al. (1994:154), Fretey et al. (2011:42), Scott et al. (2013:201), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is broadly distributed in sub-Saharan Africa, extending from Senegal to Kenya, and south through Mozambique, Malawi, Angola, into easternmost Namibia and northeastern South Africa. Ocurrences in Angola (Map 99): The species is known from southern Angola. Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a:157; Ruas 1996:23); “23 km NW of Pereira de Eqa (Roqadas)” [-16.95000, 15.56667] (Parry 1982:286; Poynton and Haacke 1993:13; Ruas 1996:23); “Pereira de Eqa” [-17.06667, 15.73333] (Poynton and Haacke 1993:13; Ruas 1996:23); “Mupanda” [-17.13333, 15.76667] (Monard 1937a:46, 1938:103; Ruas 1996:23). Taxonomic and distributional notes: There has been on-going confusion about the taxono¬ my of Pyxicephalus edulis Peters 1854 andR adspersus Tschudi 1838. The former was recognized as a subspecies of the latter by Boulenger (1882) and many subsequent authors (Loveridge 1950, MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 153 1953a; Parry 1982; Poynton and Broadley 1895b; Poynton and Haacke 1993; Ruas 1996), though Channing et al. (1994) elevated P. edulis based on differenees in the ealls and breeding biology, as well as some morphologi- eal differenees (Seott et al. 2013). Channing et al. (1994) also suggested that much of the liter¬ ature regarding Pyxicephalus adspersus should instead apply to P. edulis. Several authors including Channing (2001), du Preez and Caruthers (2009), and Channing et al. (2012) considered that P. edulis does not occur in Angola, whereas Fretey et al. (2011) consid¬ ered it present there. Ceriaco et al. (in prep.) recently examined Monard’s (1937a, 1938) specimens from “Mupanda” identified as Rana {Pyxicephalus) adspersus in the Musee d’his- toire naturelle de La Chaux-de-Fonds, Switzer- Distribution of Pyxicephalus edulis in Angola. land and considered that these should be referred to P. edulis based on distinctive morphological characters. i:°E lO'E 2'0°E 24*E Genus Tomopterna Dumeril and Bibron, 1841 Tomopterna cryptotis (Boulenger, 1907) Tremelo Sand Frog Rana cryptotis 'QouXengex 1907b:109. Syntypes: BMNH 1947.2.1.73-78, 1947.2.28.48-5, and MCZ A- 19268, exchanged from BMNH fide Barbour and Loveridge (1946:182) (collector Dr. W.J. Ansorge), formerly BMNH 1907.6.29.82-96. Type locality: “Catequero,” “Ponang Kuma (Dongwenna)” and “Kafitu swamps” (Boulenger 1907b: 109) Namibe Province, Angola. Rana cryptotis'. Barbour and Loveridge (1946:182). Tomopterna cryptotis: Poynton and Broadley (1985b: 125), Poynton and Haacke (1993:13), Frost (1985:528, 2016), Ruas (1996:23), Channing (2001:365), Largen (2001:330), Channing and Howell (2006:329), Pick- ersgill (2007a: 11 8). Tomopterna {Tomopterna) cryptotis: Fretey et al. (2011:43). Tomopterna cryptotis: Conradie et al. (2016:19). Global conservation status (lUCN): Least Concern. Global distribution: This widespread species is found in xeric regions across much of sub- Saharan Africa. Ocurrences in Angola (Map 100): The species occurs in southwestern Angola. Namibe: “25 km W of Virei” [-15.66667, 12.76667] (Poynton and Haacke 1993:13; Ruas 1996:23); “Miranda” (Boulenger 1907b: 109; Poynton and Haacke 1993:13; Ruas 1996:23). Cunene: “2 km NW of Calequero (Cahama)” [-16.28333, 14.30000] (Poynton and Haacke 1993:13); “Catequero” [-16.56667, 14.90000] (Boulenger 1907b:109; Poynton and Broadley 1985b:125; Frost 2016, 1985:528; Ruas 1996:23; Largen 2001:330); “Ponang Kuma (Dongwenna)” [-17.05000, 14.65000] (Boulenger 1907b: 109; Barbour and Loveridge 1946:182; Poynton and Broadley 1985b: 125; Frost 2016, 1985:528; Ruas 1996:23); “Kafitu swamps” (Boulenger 1907b: 109; Poynton and Broadley 1985b: 125; Frost 2016, 1985:528; Largen 2001:330). Cuando Cubango: “Cubango basin (22c)” [-14.65386, 16.93547] (Conradie et al. 2016:8-9,19); “Cubango basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:9-10,19). 154 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: Originally described based on specimens from Angola, many subsequent authors have identi¬ fied populations across much of Africa as T. cryptotis. Recent molecular genetic analyses suggest at least some populations found far from Angola, including Somalia and Kenya, represent other distinct species (Zimkus and Larson 2011; Wasonga and Channing 2013). Because of the difficulty in identifying some species of Tomopterna based on morphology alone, the extent of the distribution of this species remains uncertain (Channing 2001; Channing and Howell 2006). Some records of T. cryptotis might refer instead to T. tandyi. Channing (2001) provided a map for T. tandyi with records from southern Angola (Pickersgill 2007a; Channing et al. 2012), between Namibe and Cunene provinces. 12°E l(j®E 2i>°E 24'*E Map too. Distribution of Tomopterna cryptotis in Angola. Tomopterna damarensis Dawood and Channing, 2002 Damaraland Sand Frog Tomopterna damarensis Dawood and Channing 2002:130, figs 1, 2, 3. Holotype: TM 83913 (collector A. Channing). Type locality: “Khorixas” (Dawood and Channing 2002:133), Kunene Region, Namibia. Tomopterna damarensis: Frost (2016), Ceriaco et al. (2016a:20). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from southwestern Angola and northern Namibia. Ocurrences in Angola (Map 101): The species occurs in southwestern Angola. Namibe: “Pediva Hot springs” [-16.28462, 12.56329] (Ceriaco et al. 2016a;20). Taxonomic and distributional notes: Dawood and Channing (2002) suggested that this species might occur more broadly. Recent¬ ly Ceriaco et al. (2016a) reported the first record for Angola from Iona Natinoal Park. Previous records of some other species of Tomopterna, including T. tandyi, might refer to this species (Heinicke et al. 2017). 1I°E It'E 20°E 24”E Map 101. Distribution of Tomopterna damarensis in Angola. Tomopterna krugerensis Passmore and Carruthers, 1975 Knocking Sand Frog Tomopterna krugerensis Passmore and Carmthers 1975:32, figs. 4, 5, 6, 7, 8. Holotype: TM 44670 (collectors N.I. Passmore and V.C. Carmthers). Type locality: “Machayipan, Kmger National Park,” South Africa. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 155 Tomopterna krugerensis: Poynton and Broadley (1985b: 127), Channing (2001:368). Tomopterna {Tomopterna) krugerensis: Fretey et al. (2011:43). Global conservation status (lUCN): Least Concern. Global distribution: The species is occurs in savannas extending from Angola and Namibia, east through Botswana into northern South African and southern Mozambique. Ocurrences in Angola: The species occurs in southwestern Angola. Undetermined Locali¬ ty: without precise locality (Channing 2001:369). Taxonomic and distributional notes: This species is similar morphologically to T. cryptotis but has a distinct call (Poynton and Broadley 1985b) and is phylogenetically distinct (Zimkus and Larson 2011). Because of the morphological similarity, some previous reports from Angola for T. cryptotis might be referable to this species or T. tandyi. Channing (2001) presented the only records in Angola for T. krugerensis with mapped points in Namibe and Cunene provinces, though without specific locality details. Tomopterna tandyi Channing and Bogart, 1996 Tandy’s Sand Frog Tomopterna tandyi Channing and Bogart 1996:80, figs. 2, 3. Holotype: PEM A-2283 (collector M. Snyman). Type locality: “Bedford in the Eastern Cape” (Channing and Bogart 1996:80), South Africa. Tomopterna tandyi: Channing (2001:372), Pickersgill (2007a: 117), Fretey et al. (2011:43), Channing et al. (2012:360), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs in South Africa, Namibia, and southwestern Angola. Ocurrences in Angola: The species occurs in southwestern Angola. Undetermined Locali¬ ty: “without precise locality” (Channing 2001:369). Taxonomic and distributional notes: A tetraploid species, likely of hybrid origin between Tomopterna cryptotis (Boulenger, 1907) and Tomopterna delalandii (Tschudi, 1838) (Channing and Bogart 1996, Frost 2016). This species is morphologically indistinguishable from T. crypto¬ tis and T. delalandii and many records of Tomopterna cryptotis, including in Angola, may refer to T. tandyi. Channing (2001) provided a map for T. tandyi with records from southwestern Angola between Namibe and Cunene provinces (followed by Pickersgill 2007a, Channing et al. 2012, and Frost 2016), though without specific information. Tomopterna tuberculosa (Boulenger, 1882) Rough Sand Frog Pyxieephalus rugosus Gunther 1865a:479, pi. 33, fig. 1. Syntypes: BMNH 1947.2.29.10-11 (collector F.M.J. Welwitsch), formerly BMNH 64.7.13.5-6. Type locality: “Pungo Andongo” and “W. Africa” (Gunther 1865a:479), Angola. Preoccupied by Rana rugosa Temminck and Schlegel, 1838. Rana tuberculosa Boulenger 1882:30. Replacement name for Pyxieephalus rugosus Gunther, 1865. Rana {Tomopterna) signata A\\\ “1923” 1925:43. Syntypes: ZMB 6458, 7765 [2 specimens] (donor J.V.B. Bocage). Type locality: “Huilla” [= Huila], Malanje Province, Angola. Synonymy by Laurent (1954a: 72). Rana {Tomopterna) eaeondana Ahl “1923” 1925:43. Holotype: ZMB 15477 (purchased Linnea). Type local¬ ity: “Caconda,” Huila Province, Angola. Synonymy by Laurent (1954a:72). Pyxieephalus rugosus: Bocage (1870:68, 1873b:226, 1887c:211). Rana tuberculosa Bocage (1895a: 156, 1897b:211), Ferreira (1904:111), Schmidt (1936:130), Mertens (1937a: 19), Schmidt and Inger (1959:36). Rana {Tomopterna) tuberculosa: Monard (1938:102). Pyxieephalus tuberculosus: Parker (1939:142), Haacke (1970:276). Tomopterna signata: Cei (1997:17). Tomopterna rugosa: Laurent (1954a:72). Tomopterna tuberculosa: Loveridge (1957:345), Laurent (1964a: 133), Cei (1997:17), Poynton and Broadley 156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (19856:130), Poynton and Haacke (1993:13), (1996:23), Channing (2001:372), Frost (2016). Tomopterna (Tomopterna) tuberculosa: Fretey et al. (2011:43). Global conservation status (lUCN): Least Coneem. Global distribution: This is a widespread species occurring in savannas extending from Angola and northwestern Namibia across southern Democratic Republic of Congo to Tanzania, Zambia, and Zimbabwe. Ocurrences in Angola (Map 102): The species occurs throughout much of Angola. Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Gunther 1865a:480, 1869:479; Boulenger 1882:30; Bocage 1895a: 156; Loveridge 1957:345; Schmidt and Inger 1959:36; Poynton and Broadley 19856:130; Ruas 1996:23). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:142; Ruas 1996:23); “Novo Redondo” [-11.20000, 13.85000] (Ferreira 1904:111; Schmidt and Inger 1959:37; Ruas 1996:23). Moxico: “Reserve de Cameia (Cameia)” [-11.83333, 21.00000] (Laurent 1964a:133; Ruas 1996:23). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1936:130; Ruas 1996:23); “General Machado” [-12.03333, 17.46667] (Mertens 1937a:19). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1938:102; Ruas 1996:23); “Galanga” [-12.06667, 15.15000] (Bocage 1895a:156; Ruas 1996:23); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:142; Ruas 1996:23). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1887c:211, 1895a:156; Ruas 1996:23); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:156; Ruas 1996:23); “Ebanga” [-12.73333, 14.73333] (Monard 1938:102; Ruas 1996:23); “Hanha” [-13.30000, 14.20000] (Bocage 1897b:211; Ruas 1996:23). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:156; Ahl “1923” 1925:45; Ruas 1996:23); “Kalukembe (Caluquembe)” [-13.78333, 14.68333] (Monard 1938:102; Ruas 1996:23); “13 km N of Cutenda (Cutenda 13 km N)” [-14.25000, 15.05000] (Poynton and Haacke 1993:13; Ruas 1996:23); “7 km N of Cutenda (Cutenda 7 kmN)” [-14.36667, 15.08333] (Poynton and Haacke 1993:13; Ruas 1996:23); “Kuvan- gu” [-14.46667, 16.30000] (Monard 1938:102; Ruas 1996:23); “Huilla” [-15.05000, 13.55000] (Bocage 1873b:226, 1895a:156; Ahl “1923” 1925:45; Ruas 1996:23); “7 km SE of Jau (Jau 7 km)” [-15.21667, 13.51667] (Poynton and Haacke 1993:13); “Cascade de Ongueria (Chibia)” [-15.30000, 13.51667] (Laurent 1954a:72; Ruas 1996:23); “Christo Rei (Cristo Rei)” [-14.93333, 13.51667] (Poynton and Haacke 1993:13; Ruas 1996:23). Cunene: “riusseau Mbale (Bale)” [-15.16667, 16.75000] (Monard 1938:102; Ruas 1996:23). Namibe: “Biballa (Bibalay’ [-14.76667, 13.36667] (Bocage 1895a:156; Ruas 1996:23); “Bottom of Leba Pass (Leba)” [-15.06667, 13.23333] (Poynton and Haacke 1993:13; Ruas 1996:23). Undetermined Locality: without precise locality (Bocage 1870:68). Taxonomic and distributional notes: The nomen Pyxicephalus rugosus was preoccupied and was replaced by Rana tuberculosa Boulenger, 1882. Ahl (1925 “1923”) described two new species of Tomopterna based on Angolan types, which Laurent (1954a) subsequently treated as junior synonyms of what is now T. tuberculosa. Map 102. Distribution of Tomopterna tuberculosa in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 157 Family Dicroglossidae Anderson, 1871 Genus Hoplobatrachus Peters, 1863 Hoplobatrachus occipitalis (Gunther, 1858) African Crowned Bullfrog Rana occipitalis Gunther 1858a:320. Syntypes: BMNH 1853.2.21.4, 1858.11.25.99-102, -112-113, 1932.11.1.1-2 [12 specimens] (collector unknown). One of the type specimens is depicted in Gunther ‘T858b” 1859:130, pi. 11. Type locality: “West-Afrika” (Gunther 1858a:320). Rana bragantina Bocage (1864:254). Holotype: MBL (collector F.A.P. Bayao), not located by Ferret (1967a: 19), possibly lost or destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” (Boca¬ ge 1864:254) [= Calandula], Malanje Province, Angola. Rana occipitalis: Bocage (1866a:53, 1870:68, 1895a: 155), Boulenger (1882:27), Mertens (1938a:426), Themido (1941:2), Loveridge (1957:344), Gavetti and Andreone (1993:98), Frost (1985:508). Dicroglossus occipitalis: Laurent (1950a: 14, 1954a:71), Hellmich (1957a:24), Perret (1976a: 19), Cei (1977:16). Euphlyctis occipitalis: Poynton and Broadley (1985b: 125), (1996:23). Hoplobatrachus occipitalis: Channing (2001:295), Fretey et al. (2011:37), Channing et al. (2012:166), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from most of sub-Saharan Africa. Ocurrences in Angola (Map 103): The species known from northern and western Angola, though probably occurs more widely. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a: 71); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:14, 1954a:71; Ruas 1996:23); “Muita, Luembe E, dans la vallee marecageuse de la Kasseke (afluente de la Muita)” [-7.80000, 21.45000] (Laurent 1950:14; Ruas 1996:23). Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1864:254, 1866a:53, 1895a:155; Per¬ ret 1976a: 19; Ruas 1996:23). Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Bocage 1895a:155; Ruas 1996:23); “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:24); “Dondo, sur le bord droit du Quanza” [-9.68333, 14.43333] (Bocage 1895a:155; Ruas 1996:23). Kwanza Sul: “Novo Redon¬ do” [-11.20000, 13.85000] (Bocage 1895a:155; Map 103. Distribution oi Hoplobatrachus occipitalis in Angola. Ruas 1996:23). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage, 1895a:155; Themido 1941:2; Gavetti and Andreone 1993:98; Ruas 1996:23); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:426; Ruas 1996:23). Undeter¬ mined Locality: without precise locality (Bocage 1870:68; Laurent 1954a:71); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: When publishing the description of R. bragantina, Bocage (1864) was unaware of Gunther’s (1858) R. occipitalis, but quickly recognized this and synonymized the two shortly thereafter (Bocage 1866a). 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family Ranidae Batsch, 1796 Genus Amnirana Dubois, 1992 Amnirana albolabris (Hallowell, 1856) White-Lipped Frog Rana albolabris Hallowell 1856:153. Syntypes: ANSP [4 specimens], lost fide J.L. Ferret in Frost (1985:479) (collector P. du Chaillu). Type locality: “W. Africa” (Hallowell 1856:153), restricted to “Gabon” by Ferret (1977:843). Limnodytes albolabris: Peters (1877a:618). Rana albolabris: Bocage (1895a: 162). Hylarana albolabris albolaris: Laurent (1950a: 14, 1954a:74), Loveridge (1957:338). Hylarana albolabris: Ferret (1977:843), Ruas (1996:24). Amnirana albolabris: Channing and Howell (2006:256), Channing et al. (2012:360), Frost (2016), Jongsma et al. (2018:275). Rana (Amnirana) albolabris: Fretey et al. (2011:43). [-5.10000, 12.10000] (Peters 1877a:618; Boca¬ ge 1895a; 162). Uige: “Uige, University Kimpa Vita campus” [-8.607967, 20.968804] (Jongs¬ ma et al. 2018:275); “Serra do Pingano” [-7.68528, 14.92956] (Jongsma et al. 2018:275); “Negage” [-7.7625, 15.285] (Jongsma et al. 2018:275); “Uige fish farm, Kibokolo” [-6.274417, 15.067472] (Jongsma et al. 2018:275). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 14, 1954a: 74; Ruas 1996:24); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:14; Ruas 1996:24); “rive de la Tshihumbe, 40 km a Lest de Dundo (Rio Chiumbe)” [-8.01667, 19.31667] (Laurent 1950a: 14; Ruas 1996:24); “Tshinguvu, galerie forestiere de la Tshikapa, 50 km au sub-ouest de Dundo (Chingufo)” [-8.01667, 19.31667] (Laurent 1950a: 14; Ruas 1996:24); “Sombo” [-8.68333, 20.95000] (Laurent 1954a:74; Ruas 1996:24). Taxonomic and distributional notes: Ferret (1977) restricted the distribution of this species to the forested area of Cameroon and Congo (Channing and Howell 2006). Laurent (1964a) was uncertain whether his Angolan material corresponded to A. albolabris or instead to A. lemairei (Laurent 1950a, 1954a). Jongsma et al. (2018) included Angolan populations in their molecular phylogenetic analyses and found that these fell within a large complex of likely cryptic species all currently referred to as A. albolabris. Map 104. Distribution of Amnirana albolabris in Angola. Global conservation status (lUCN): Least Concern. Global distribution: The wide-ranging forest species found across western and Central Africa likely represents multiple species, each with more restricted distributions. Ocurrences in Angola (Map 104): The species occurs in northern Angola and in the Cabinda enclave. Cabinda: “Chinchoxo” Amnirana darlingi (Boulenger, 1902) Darling’s White-Lipped Frog Rana darlingi Boulenger 1902:15. Syntypes: BMNH 1947.2.2.67-68 (eolleetor J. ffolliott Darling), formerly MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 159 BMNH 1902.2.12.114-115" before period. Type loeality: “Mazoe and between Umtali and Marandellas” [= Mashonaland], Zimbabwe. Rana albolabris adiscifera Sehmidt and Inger (1959:48). Holotype: FMNH 21171 (colleetor R. Boulton). Type loeality: “Chitau”, Angola. Rana {Rana) darlingi: Monard (1937a:44, 1938:100). Rana albolabris: Schmidt (1936:130). Rana albolabris adiscifera: Marx (1976:62). Rana darlingi: Frade (1963:254), Laurent (1964a: 132), Cei (1977:16). Hylarana darlingi: Poynton and Broadley (1985b: 139), Ruas (1996:24, 2002:143), du Preez and Carruthers (2009:452). Amnirana darlingi: Channing (2001:266), Channing et al. (2012:364), Frost (2016), Conradie et al. (2016:19), Jongsma et al. (2018:275). Rana (Amnirana) darlingi: Fretey et al. (2011:43). Hylarana cf darlingii: Ceriaco et al. (2016b:49). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs in south-central Africa, extending from Angola through the Democratic Republic of Congo, Zambia, Malawi, Zimbabwe, and parts of Botswana and Mozambique. Ocurrences in Angola (Map 105): The species occurs especially in the eastern regions of Angola. Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:25). Lunda Norte: “Poste de Luangue, humidiherbosa du ruisseau Katcheleka, affl. W. Luangue, entre le Lunguena et le Tcha-Pemba (Posto do Luangue)” [-9.08333, 19.71667] (Laurent 1964a: 132; Ruas 1996:24); “Lagoa Carumbo” [-7.738333, 19.9833] (Jongsma et al. 2018; 275)". Lunda Sul: “Dala” [-11.03333, 20.20000] (Monard 1937a:44, 1938:100; Ruas 1996:24); “Alto Chicapa, sour¬ ces du Cuilo (Rio Cuilo)” [-10.86667, 19.40000] (Laurent 1964a; 132; Ruas 1996:24); “mare des rives Kutele, affl. rive droite du Cuango, Alto Chicapa (Rio Cutele)” [-11.06667, 18.86667] (Laurent 1964a:132; Map 105. Distribution of in Angola. Ruas 1996:24); “sources du Tchimboma, Alto Chicapa (Rio Chimboma)” [-10.93333, 19.18333] (Laurent 1964a: 132; Ruas 1996:24). Mexico: “Rives du lac Calundo (Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a: 132; Ruas 1996:24); “Rio, Calombe, Reserva da Palanca Preta” [-11.83333, 19.93333] (Ruas 2002:143); “Calombe, Luso” [-11.83333, 19.93333] (Ruas 1996:24, 2002:143). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1936:130; Schmidt and Inger 1959:48; Marx 1976:62; Ruas 1996:24). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937a:44, 1938:100; Ruas 1996:24). Huila: “Sangeve” [-13.88333, 15.83333] (Monard 1937a:44, 1938:100; Ruas 1996:24); “Kuvangu” [-14.46667, 16.30000] (Monard 1937a:44, 1938:100; Ruas 1996:24); “Kambisa” [-15.31667, 16.21667] (Monard 1937a;44, 1938:100; Ruas 1996:24). Cunene: “ruis¬ seau Mbale (Bale)” [-15.16667, 16.75000] (Monard 1937a:44, 1938:100 Ruas 1996:24). Cuando Cubango: “Cubango basin (22a)” [-14.66622, 16.97842] (Conradie et al. 2016:19); “Cuito basin (55)” [-14.68478, 18.67369] (Conradie et al. 2016:19). Undetermined Locality: “Zambezian 160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II highlands” (Frade 1963:254); “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: Laurent (1964a) synonymized Rana albolabris adis- cifera with Rana darlingi. Amnirana lemairei (de Witte, 1921) Lemaire’s White-Lipped Frog Rana Lemairei de Witte 1 92 1 : 1 , pi. 1 , fig. 1 -4. Holotype: MRAC 1 72 (eolleetor Mission Lemaire). Type loeal- ity: “Lofoi (Katanga)” (de Witte 1921:3), Demoeratie Republie of Congo. Hylarana albolabris lemairei: Laurent (1964a: 133), Cei (1977:16) Hylarana lemairei: Poynton and Broadley (1985b: 140), Ruas (1996:24, 2002:143), Channing et al. (2012:365). Amnirana lemairei: Channing (2001:269), Frost (2016). Rana {Amnirana) lemairei: Fretey et al. (2011:43). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from northeastern Angola, the southern Demo¬ cratic Republic of Congo, and western Zambia. Ocurrences in Angola (Map 106): The species occurs in northeastern Angola. Lunda Norte: “rives du Luachimo, Dundo (Luachi- mo)” [-7.38333, 20.85000] (Laurent 1964a: 133; Ruas 1996:24); “Riv. Chitato, Dundo (Rio Chitado)” (Laurent 1964a: 133; Ruas 1996:24). Lunda Sul: “Alto Cuilo, dans I’eau, tres basse, des sources d’un ruisseau (Alto Cuilo)” [-10.01667, 19.55000] (Laurent 1964a: 133; Ruas 1996:24); “Chutes du Cuan- go-Muque, Alto Chicapa (Rio Cuango-Muque, quedas)” [-10.76667, 19.20000] (Laurent 1964a: 133; Ruas 1996:24); “sources de la Tchimboma, affl. gauche du Cuango-Muque, Alto Chicapa (Rio Chimboma, nascente)” [-10.93333, 19.18333] (Laurent 1964a:133; Ruas 1996:24). Mexico: “Lumeje I, galerie forestiere isolee a cote de la route, pres de I’en¬ Map 106. Distribution of Amnirana lemairei in Angola. tree de la reserve de chasse de Cameia (Rio Lumege)” [-11.58333, 21.00000] (Laurent 1964a: 133; Ruas 1996:24); “Lago Cameia” [-11.71667, 20.80000] (Ruas 1996:24). Undetermined Locality: “areas of forest and savanna in the north and northeast of Angola” (Cei 1977:16). Taxonomic and distributional notes: This taxon was discussed by Schmidt and Inger (1959) as a subspecies of Rana albolabris (Hallowell, 1856). Laurent (1964a) identified some individuals as Rana albolabris lemairei, and Perret (1977) later recognized this as a full species, although he expressed some doubts about it occurence in Angola. Amnirana lepus (Andersson, 1903) Andersson’s White-Lipped Frog Chiromantis lepus Andersson 1903: 142. Holotype: NHR ? (eolleetor Y. Sjostedt). Type loeality: “Kamemn,” [=Cameroon]. Amnirana lepus: Jongsma et al. (2018:275). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I6I Global conservation status (lUCN): Least Concern. Global distribution: The species occurs from Cameroon south to northern Angola and east through the central Democratic Republic of Congo. Ocurrences in Angola (Map 107): The species occurs in the northern regions of the country. Uige: “Serra Pingano” [-7.68528, 14.92956] (Jongsma et al. 2018:275); “Quilo- moQo” [-7.642333, 15.005722]. Lunda Norte: “Carumbo” [-7.752722, 19.95642] (Jongsma et al. 2018:275). Taxonomic and distributional notes: Jongsma et al. (2018) recently published the only reported records for this species in Ango¬ la based on comparisons of genetic data to specimens of A. lepus from throughout its range. The relationship between A. lepus and the similar Angolan endemic species A. parkeriana requires further research. Map Angola. 107. Distribution of Amnirana lepus in n”}: i6.“t 2-i'>E Amnirana parkeriana (Mertens, 1938) Parker’s White-Lipped Frog (Endemic) Rana albolabris acutirostris Parker (1936:141). Holotype: BMNH 1936.8.1.1, currently BMNH 1947.2.3.69. (collector K. Jordan). Type locality: “Congulu” [= Congulo] Kwanza Sul, Angola. Preoccupied by Rana fusca acutirostris Fatio, 1872. Rana albolabris parkeriana: Mertens (1938b: 14), Loverdige (1941a). Rana albolabris acutirostris: Barbour and Loveridge (1946:181). Rana parkeriana: Perret (1977:844), Frost (1985:509). Hylarana parkeriana: Ruas (1996:24). Amnirana parkeriana: Channing (2001:270), Frost (2016). Rana {Amnirana) parkeriana: Fretey et al. (2011:43). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 108): The species is known only from a small area on the western escarpment of Angola. Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:141; Barbour and Loveridge 1946:181; Ruas 1996:24); “Congulu” [-10.86667, 14.28333] (Parker 1936:141; Frost 1985:509; Ruas 1996:24). Taxonomic and distributional notes: Parker (1936) described this species as Rana I(J®E 2(}°:E I4'*E Map 108. Distribution of Amnirana parkeriana in Angola. 162 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II albolabris acutirostris based on a single specimen from “Congulu” and four from “Quirimbo.” As the name was preoccupied, Mertens (1938b) provided the replacement name Rana albolabris park- eriana (Loverdige 1941a; Barbour and Loveridge 1946; Ferret 1977). There have been no new records since the description of this species and its distinction from other large-bodied species of Amnirana, such as A. lepus, remains in doubt. Family Rhacophoridae Hoffman, 1932 (1858) Genus Chiromantis Peters, 1854 Chiromantis xerampelina Peters, 1854 Grey Foam-Nest Treeerog Chiromantis xerampelina Peters (1854:627). Syntypes: ZMB 6593, 6594 [4 speeimens] (eolleetor W.C.H. Peters) fide Bauer et al. (1995:50). Type loeality: “Tette und Sena” Mozambique. Chiromantis xerampelina: ^ch\0tz {\999-3K), C\m\- ning (2001:375), Piekersgill (2007a:296), Fretey et al. (2011:43), Frost (2016). Global conservation status (lUCN): Least Concern. Global distribution: The widespread species occurs in savannas of eastern and cen¬ tral Africa, extending into northeastern South Africa. Ocurrences in Angola (Map 109): The species is known from southwestern Angola. Benguela: “Benguela” (Schiotz 1999:39; Channing 2001:376). Taxonomic and distributional notes: Schiotz (1999) and Channing (2001) provided maps with a record of this species in Benguel- la Province, though without information about the locality; Fretey et al. (2011) also listed this species as occurring in Angola. Map 109. Distribution of Chiromantis xerampelina in Angola. REPTILIA Order CHELONII Brongniart, 1800 Family Pelomedusidae Cope, 1868 Genus Pelomedusa Wagler, 1830 Pelomedusa subrufa (Bonnaterre, 1789) Helmeted Terrapin Testudo Subrufa Bonnaterre 1789:28; pi. 6, fig. 5. Holotype: MNHN 7970 (eolleetor P. Commerson, see Notes below) fide Bour (1982:533). Type loeality: “Les grandes Indes” [= India] (Bonnaterre 1789:28), in error, eorreeted by Mertens (1937b: 141) to “Kap der Guten Hoffnung,” South Afriea, and later by Bour (1982:535) to “Taolanaro (Fort-Dauphin), Republie Malagasy (Madagasear)” [= Tolanaro], Madagasear. Pentonyx Gehafie: Boeage (1870:68). Pelomedusa galeata: Boeage (1887b:202, 1895a:5), Mertens (1937b: 152), Sehimdt (1933:3), Monard (1937b: 147). Pelomedusa subrufa subrufa: Loveridge (1941b:470), Bour (1982:533). MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 163 Pelomedusa subrufa: Iverson (1986:236, 1992:52), Boycott and Bourquin (2008:007.1), Turtle Taxonomy Working Group (2014:435), Petzold et al. (2014:542), Ceriaco et al. (2016a:54), Conradie et al. (2016:26). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is distributed from the subtropical savannas and semi-desert regions in Angola, Namibia, Botswana, Zimbabwe, Zambia, Malawi, the southeastern Democratic Republic of Congo, parts of Mozambique, and portions of northeastern South Africa. It has also been introduced to Madagascar. Ocurrences in Angola (Map 110): This species has been confirmed from numerous localities across southern and central Angola. Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1895a:5; Monard 1937b: 146; Loveridge 1941b:479). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:3; Loveridge 1941b;479); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:3; Love¬ ridge 1941b;479). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1895a:5; Monard 1937b: 146; Loveridge 1941b: 479); “Catumbella-Miindung, Benguella” [-12.33333, 13.55000] (Mertens 1926:152). Huila: “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:147; Loveridge 1941b:479); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:5; 1937b:146; Loveridge 1941b:479); Map 110. Distribution of Pelomedusa subrufa in Angola. “sur les hordes du Kuvangu, au sud de Vila-da-Ponte (Kuvangu river S. Vila da Ponte)” [-14.46667, 16.30000] (Monard 1937b: 147; Loveridge 1941b:479); “Huila District” [-15.24179, 14.52210] (Petzold et al. 2014, supplemental information); “Kahuihui” [Undetermined Locality] (Monard 1937b: 147; Loveridge 1941b:479); “Kalundunga” [Undetermined Locality] (Monard 1937b: 147; Loveridge 1941b:479). Namibe: “Mucungu” [-14.78333, 12.48333] (Schmidt 1933:3; Loveridge 194bl:479; Ceriaco et al. 2016a:54); “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:5; Monard 1937b:146; Loveridge 1941b:479; Ceriaco et al. 2016a:54); “Capangombe” [-15.10000, 13.15000] (Bocage 1887b:202; 1895a:5; Monard 1937b:146; Loveridge 1941b:479; Ceriaco et al. 2016a:54); “Mossamedes” [-15.20000, 12.15000] (Bocage 1887b:202; 1895:5; Loveridge 1941b:479; Ceria¬ co et al. 2016b:54). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a:5; 1937b:146; Loveridge 1941b:479). Cuando Cubango: “23 km E Jamba” [-17.48690, 22.98790] (Petzold et al. 2014, supplemental information); “Cuando basin (40)” [-17.45786, 22.91191] (Conradie et al. 2016:9-10, 26). Taxonomic and distributional notes: This species was first described by Lacepede (1788) as Testudo subrufa, but following the ICZN ruling that this was an unavailable non-binominal work (ICZN 2005, Opinion 2104, Case 3226), authorship has been attributed to Bonnaterre (1789). Mertens (1937b) and Loveridge (1941b) demonstrated that T. subrufa pre-dated T. galeata, until then widely employed as the name for the helmeted terrapin. As the collector named by Lacepede, Pierre Sonnerat, was known to have collected in the southwestern Cape region of South Africa, Mertens (1937b) restricted the type locality to this area. However, Bour (1982) presented evidence that the type was probably actually collected by Philibert Commerson in Madagascar in 1770 (see review in Fritz et al. 2014). Loveridge (1941b) cited several species and subspecies for 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Pelomedusa, but subsequently P. subrufa was long treated as monotypic (e.g., Gasperetii et al. 1993; Fritz and Havas 2007; Turtle Taxonomy Working Group 2007; Boyeott and Bourquin 2008). However, Vargas-Ramirez et al. (2010) identified several species-level lineages within P subrufa. Petzold et al. (2014) have formally resurrected several species from synonymy and have described additional species corresponding to the lineages identified genetically. The populations in Angola belong to the same taxon as the Madagascan type of P. subrufa, which represents a population introduced from the mainland of Africa (Vargas-Ramirez et al. 2010; Wong et al. 2010). Iverson (1986, 1992) and The Turtle Taxonomy Working Group (2007, 2014, 2017) presented point local¬ ity distribution maps for the species. Genus Pelusios Wagler, 1830 Pelusios bechuanicus FitzSimons, 1932 Okavango Mud Turtle Pelusios bechuanicus FitzSimons 1932:37. Holotype: TM 14688 fide Broadley (1981a:664) and Mashinini and Mahlangu (2014:184) (collector V.F. FitzSimons). Type locality: “Thamalakane River at Maun, Ngamiland, Bechuanaland,” [Botswana]. Sternothaerus nigricans'. Monard (1931:109, 1937b: 148). Pelusios bechuanicus'. Laurent (1964a:27), Iverson (1986:239, 1992:57), Fritz and Havas 2007:346, Turtle Taxonomy Working Group (2014:436), Conradie et al. (2016:27). Pelusios bechuanicus bechuanicus'. Broadley (1981a:664). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is limited to the greater Okavango basin, from the Cuban¬ go/Okavango River in the west to the Kafue Flats in Zambia and south to northeastern Namibia and northern Botswana. Ocurrences in Angola (Map 111): This species occurs in central- southeastern Angola. Moxico: “Riv. Chonga, affl. de la Lumeje, 100 km a Test de Luso” [-11.75000, 20.75000] (Laurent 1964a:27; Broadley 1981a:666). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1931:109, 1937b: 148; Broadley 198 la: 666). Cuando Cubango: “flood waters of the lower Cuando River (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:90-10, 27). Taxonomic and distributional notes: FitzSimons (1932) short description was sup¬ plemented by a more extensive and illustrated version (FitzSimons 1935). Laurent (1964a) resurrected Pelusios bechuanicus from the syn¬ onymy of Pelusios subniger (Bonnaterre, 1789) based on a single Angolan specimen from Moxico Province, and subsequently Broadley (1981a) allocated Sternothaerus nigricans (non Donndoff, 1798) from “Chimporo” (Monard 1931, 1937b) to this species as well. Iverson (1986, 1992) provided a distribution map with two records from Angola, which correspond to the “Chimporo” and “Chonga River” records listed above. Map Angola. 111. Distribution of Pelusios bechuanicus in MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 165 Pelusios castaneus (Schweigger, 1812) West African Mud Turtle Emys castanea Schweigger 1812:314. Holotype: MNHN speeimen lost fide Bour (1978 “1979”) (eolleetor unknown); Neotype: MNHN 2008.0303 (eolleetor Fouehard-Togamin), designated by Bour (2008). Type loeality: “Patria ignota” [unknown]; Neotype loeality: “vicinity of Koutchatcha (7°20’ N, 1°18’ E), a vil¬ lage elose to the Amou River (ea. 30 km East of Glei), Ogou Prefeeture, Plateaux Region, Togo.” Sternothaerus derbianus: Peters (1877a:611), Frade (1963:252). Sternothaerus Derbianus: Boeage (1895a:3). Pelusios subniger Loveridge (1941b:491, 1957:175). Pelusios eastaneus: Iverson (1986:241, 1992:60), Fritz and Havas 2007:346, Kindler et al. (2016:305), Bour et al. (2016:095.1). Pelusios eastaneus eastaneus: Turtle Taxonomy Working Group (2014:436); Bour et al. (2016:095.3). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from West Africa, from Senegal east through extreme southern Chad and the adjacent Central African Republic. A southern population occurring in near-coastal localities from Gabon to northern Angola is disjunct from the main distribution. Introduced populations occur on Sao Tome and on Guadeloupe (but not on the Cape Verde Islands as previously noted; see Bour et al. 2016). Ocurrences in Angola (Map 112): This species occurs along the coast in the northwest from southern Bengo Province north to Cabin¬ da. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611; Loveridge 1941b:501). Luanda: “Loanda” [-8.83333, 13.26667] (Boeage 1895a;3; Loveridge 1941b:501); “Cacuaco, 10 km NE Luanda” [-8.78333, 13.36667] (Bour et al. 2016:095.3). Taxonomic and distributional notes: One non-nominate subspecies has been recog¬ nized, the extinct P. c. seychellensis (Sieben- rock, 1906) (Turtle Taxonomy Working Group 2014; Bour et al. 2016), although molecular evidence suggests that it should be considered a junior synonym of the nominate form (Stuckas et al. 2013; Kindler et al. 2016). Loveridge (1941b) considered Sternothaerus Map 112. Distribution of Pelusios castaneus in Angola. derbianus as used by Peters (1877a) and Boeage (1895a) as a synonym of Pelusios subniger (Lacepede, 1789). Laurent (1956) revived/] derbianus and P. rhodesianus Hewitt, 1927 as races of Pelusios castaneus (Schweigger, 1812) although Bour (1978 “1979”) applied the name P. castaneus castaneus to the West African form, with derbianus as a synonym (Broadley 1981a). Bour et al. (2016) mapped several additional localities in Angola, all from near-coastal sites in Luanda and Bengo provinces, although place names were not given. Pelusios chapini Laurent, 1965 Central African Mud Turtle Pelusios eastaneus ehapini Laurent 1965:21, pi. 3, figs. 1-4. Holotype: MRAC 20937 (eolleetor R.F. Lau¬ rent). Type loeality: “Kasenyi, Lake Albert, Bunia Terr., Ituri, Congo” [= Lake Albert, Kasenyi, Ituri Provinee], Demoeratie Republie of Congo. Pelusios ehapini: Turtle Taxonomy Working Group (2014:437), Kindler et al. (2016:305). Global conservation status (lUCN): Not Evaluated. 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global distribution: The species is known from Central Africa, from the Central African Republic to the northern Democratic Republic of Congo and much of the lower Congo drainage, Congo, Gabon and presumably northern Angola (possibly including the Cabinda enclave). Ocurrences in Angola: This species may occur in the far northwest of the country, although there are no previously published specimen records from Angola. Taxonomic and distributional notes: According to the Turtle Taxonomy Working Group (2014) and the recent study provided by Kindler et al. (2016), it is expected that Pelusios chapini occurs in Cabinda and might reach the extreme northwest border of the country in Zaire and/or Uige povinces. The species is closely related to P. castaneus and some introgression may have occurred between the two taxa (Fritz et al. 2011; Kindler et al. 2016). Pelusios gabonensis (Dumeril, 1856) African Forest Turtle Pentonyx gabonensis Dumeril 1856:373, pi. 13, figs. 2, 2a. Holotype: MNHN 4237 (collector M. Aubry). Type locality: “Gabon.” Sternothaerus gabonensis: Bocage (1866a:40, 1866b:57). Pelusios gabonensis: Loveridge (1941b:485), Iverson (1986:244, 1992:63), Fritz and Havas 2007:348, Turtle Taxonomy Working Group (2014:438), Kindler et al. (2016:305). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is i2°e known from forested areas of Central Africa from Cameroon east across most of the central and northern portions of ther Democratic Republic of Congo to far western Uganda, Tan¬ zania and Burundi and southwest to Angola. A disjunct population occurs in Liberia and the Ivory Coast. Ocurrences in Angola (Map 113): This species occurs in northern Angola. Cabinda: “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:40, 1866b:57). Taxonomic and distributional notes: According to Loveridge (1941b), Iverson (1986, 1992) and The Turtle Taxonomy Work¬ ing Group (2014), the species is only known from Angola in Cabinda and Zaire provinces, although The Turtle Taxonomy Working Group 1 1 3 . Distribution of Pelusios gabonensis in Angola. (2017) provided an additional record from Lunda Norte. Pelusios nanus Laurent, 1956 Aerican Dware Mud Turtle Pelusios nanus Laurent 1956:31, pi. IV, figs. 2-4. Holotype, MRAC 7833 (eolleetor G. de Witte). Type loeal- ity: “Dilolo, Haut Lualaba,” Lualaba Provinee, Demoeratie Republie of Congo. Pelusios derbianus: Sehmidt (1933:3). Sternothaerus nigrieans: Monard (1937b: 148). Pelusios rhodesianus: Loveridge (1941b 491). Pelusios nanus: Laurent (1964a:25, 1965:27), Broadley (1981a:657), Iverson (1986:245, 1992:64), Fritz and Havas 2007:348, Ceriaeo et al. (2014b:670), Turtle Taxonomy Working Group (2014:438), Kindler et al. (2016:305). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 167 Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies has a relatively limited distribution aeross eentral Angola, northern Zambia and the southern Demoeratie Republie of Congo. Ocurrences in Angola (Map 114): This speeies oeeurs in eentral Angola. Malanje: “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 2014b:670). Lunda Sul: “Alto Chieapa” [-10.88333,19.23333] (Laurent 1964a:25, 1965:27; Broadley 1981:661). Moxico: “envi¬ rons du lae Calundo” [-11.80000, 20.86667] (Laurent 1964a:25; Broadley 1981a:661); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:25; Broadley 1981a:661). Bie: “Chitau” [-11.43333, 17.15000] (Sehmidt 1933:3); “Catabola, Rio Cunhinga basin” [-12.144732, 17.280407] (Kindler et al. 2016, supplemental material:5); “river between Kuito asnd Catabola” [-12.267883, 17.114582] (Kindler et al. 2016, supplemental material:5); “Kuito” [-12.383599, 16.928029] (Kindler et al. 2016, supplemental material:5). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:148; Loveridge 1941b 501); “Elende” [-12.73333, 15.15000] (Monard 1937b:148; Loveridge 1941b 501). Huila: “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:148; Broadley 1981a:661); “Osi” [-15.08333, 15.41667] (Monard 1937b:148; Loveridge 1941b 501). Taxonomic and distributional notes: Wermuth and Mertens (1977) considered Pelusios nanus as a subspecies of Pelusios adansoni (Schweigger, 1812). However, according to Broadley (1981a), the two species are highly divergent geographically and phylogenetically. Loveridge (1941b) considered specimens refered to Sternothaerus nigricans by Monard (1937b) to be Pelu¬ sios rhodesianus Hewitt, 1927, however, according to Broadley (1981a) these specimens and those allocated to Pelusios derbianus by Schmidt (1933) are P. nanus. Iverson (1986, 1992) presented a distribution map for the species although he did not provided information about the localities plot¬ ted. Iverson’s map agrees with ours, as well as that of the Turtle Taxonomy Working Group (2014). The recent records from Capanda, Malanje Province (Ceriaeo et al. 2014b) extend the known distribution by more than 100 km. Pelusios rhodesianus Hewitt, 1927 Variable Mud Turtle Pelusios nigricans rhodesianus Hewitt 1927:375, figs, la, Ic, pi. 21, figs. 2-3. Holotype: AMG 5432 (now PEM) (collector E. Knowles Jordan). Type locality: “Mpika district, N.E. Rhodesia” [= Mpika, Muchinga Province, Zambia]. Sternothaerus Adansoni'. Bocage (1867b:217), Frade (1963:252). Sternothaerus gabonensis'. Bocage (1866a:40, 1866b:57). Sternothaerus derbianus'. Bocage (1895a:3). Sternothaerus sinuatus'. Bocage (1895a:4), Monard (1937b: 148). Pelusios sinuatus sinuatus'. Schmidt (1933:3). Pelusios derbianus'. Mertens (1938a:430). Pelusios subniger Loveridge (1941b:489), Hellmich (1957a:33), Laurent (1950a: 13). Pelusios gabonensis: Laurent (1954a:70, 1964a:25). i:°E IG'E 20°E I4"E Map 114. Distribution of Pelusios nanus in Angola. 168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Sternothaerus nigricans: Frade (1963:253). Pelusios castaneus: Laurent (1964a:26). Pelusios castaneus rhodesianus: Laurent (1965:31). Pelusios rhodesianus: Iverson (1986:247, 1992:66), Broadley (1981a:689), Fritz and Havas 2007:349, Broadley and Boyeott (2008:004.1), Fritz et al. (2011:119), Ceriaeo et al. (2014b:670), Turtle Taxonomy Working Group (2014:439), Kindler et al. (2016:305). Global conservation status (lUCN): Least Concern. Global distribution: The species is widely distributed in south-central Africa, from Uganda and western Kenya south to Malawi and northern Zimbabwe and west to northeastern Namibia, Angola and Congo. A disjunct population occurs in Swaziland and adjacent parts of South Africa and southern Mozambique. Ocurrences in Angola (Map 115): This species is very widespread in Angola, except in the arid southwest. Cabinda: “Rio Quilo” [-5.18333, 12.18333] (Bocage 1866a;40, 1895a:3; Loveridge 1941b:501; Broadley 1981a:671). Uige; “7 km WUige” [-7.614250, 14.984306] (Kindler et al. 2016, supplemental material:6) Bengo: “Ambriz” [-7.844312, 13.106493] (Bocage 1867b:217, 1895a:3; Loveridge 1941b:501; Broadley 1981a;670). Kwanza Norte: “Rio Cuanza, Mucoso nahe Dondo” [-9.53333, 14.65000] (Hellmich 1957a:33; Broadley 1981a:671); “Dondo” [-9.68333, 14.43333] (Bocage 1895a;3). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1866a:40, 1866b:57, 1895a:4; Loveridge 1941b:501; Broadley 1981a:671); “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 20 14b: 670). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:70, 1964a;25-26; Broadley 1981a:671); “riviere Tchihumbwe” [-8.01667, 19.31667] (Laurent 1950a:13). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a;26). Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Lau¬ rent 1964a:26; Broadley 1981a:671). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:3; Loveridge 1941b;501; Laurent 1965:31; Broadley 1981a;671); “Catabola, Rio Cunhinga basin” [-12.144732, 17.280407] (Kindler et al. 2016, supplemental material:5); “Kuito” [-12.383599, 16.928029] (Fritz et al. 2011:119; Kindler et al. 2016, supplemental material: 6). Huambo: “Galan- ga” [-12.06667, 15.15000] (Broadley 1981a:671). Benguela: “Rio Cuce” [-12.58333, 13.41667] (Bocage 1895a:4; Loveridge 1941b;501; Broadley 1981:671); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:430; Loveridge 1941b:501; Broadley 1981a:671). Huila: “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:148; Loveridge 1941b:501; Broadley 1981a;671). Taxonomic and distributional notes: This species was described by Hewitt (1927) as a sub¬ species of Pelusios nigricans (= Pelusios subniger [Bonnaterre, 1788]) on the basis of a series of specimens. Broadley (1981a) examined five specimens under the same Albany Museum catalogue number, 5432, one of which he regarded as a holotype. These specimens were subsequently trans¬ ferred to the Port Elizabeth Museum. Loveridge (1941b), Hellmich (1957a) and Laurent (1950a) considered it a synonym of P. nigricans, whereas Laurent (1964a, 1965) treated Angolan material Map Angola. 115. Distribution of Pelusios rhodesianus in MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 169 as P. castaneus (Schweigger, 1812). Raw (1978) elevated P. rhodesianus Hewitt, 1927 to specific rank, an action subsequently confirmed by Broadley (1981a) and accepted by most subsequent authors. Iverson (1987, 1992) presented a point distribution map for the species, although he did not provide information about the localities depicted. Broadley and Boycott (2008) also provided a distribution map of P. rhodesianus based on museum and literature records. Family Testudinidae Batsch, 1788 Genus Kinixys Bell, 1827 Kinixys belliana and Kinixys spekii The precise delineation of these two species of Kinixys in Angola remains unclear and genet¬ ic data will be necessary to establish species boundaries (U. Fritz, pers. comm.). To date only K. belliana has been genetically verified in Angola but it is highly likely, given the distribution of K. spekii elsewhere, that it is also present in the country (see notes below). Collectively the species are widespread, although absent from desert and forest habitats. Ocurrences in Angola (Map 116): The following records combine both K. belliana and K. spekii (see K. spekii account below for the specific reference to its records in the cher- sonyms list). Lunda Norte: “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a;13; Loveridge and Williams 1957:396). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:25); “Lunda” [-10.96667, 20.06667] (Monard 1937b; 146; Loveridge and Williams 1957:396). Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:25). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:40, 1895a:2; Loveridge and Williams 1957:396); “Muata-Yamv, Lui River” [-9.41667, 17.33333] (Loveridge and Williams 1957:396); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:670); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:53). Bie: “Gauca” [-11.18333, 17.45000] (Laurent 1964a:25; Loveridge and Williams 1957:396); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:4; Loveridge and Williams 1957:396); “Cuemba” [-12.15000, 18.08333] (Kindler et al. 2012, supporting information: 4); “Kuito” [-12.383599, 16.928029] (Kindler et al. 2012, supporting infor- mation;4); “Cubando basin (13)” [-13.28061, 16.74722] (Conradie et al. 2016a:8-9, 27). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a:2; Loveridge and Williams 1957:396). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1887b:209, 1895a;2; Loveridge and Williams 1957:396); “Benguella” [-12.58333, 13.41667] (Hellmich 1957a:32); “Ebanga” [-12.73333, 14.73333] (Monard 1937b; 146; Loveridge and Williams 1957:396); “Cubal” [-13.03333, 14.25000] (Mertens 1937a:5, 1938a:430; Loveridge and Williams 1957:396). Hufla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a;2; Loveridge and Williams 1957:395); “Kuvan- gu” [-14.46667, 16.30000] (Monard 1937b:146); “Osi” [-15.08333, 15.41667] (Monard 1937b:146; Loveridge and Williams 1957:396). Cunene: “riv. Mbale” [-15.16667, 16.75000] Map 116. Distribution of Kinixys belliana and Kinixys spekii in Angola. 170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (Monard 1937b:146; Loveridge and Williams 1957:396). Namibe: “Capangombe” [-15.10000, 13.15000] (Boeage 1895a:2; Loveridge and Williams 1957:396; Ceriaco et al. 2016a:54). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b: 146; Loveridge and Williams 1957:396). Taxonomic and distributional notes: There is some eonfusion surrounding the deseription of K. belliana (Bour 2006). Gray (1831a) in his Synopsis Reptilium. Cataphracta, the most frequent¬ ly cited source of the description (e.g.. King and Burke 1989), includes in his section “Additions and Corrections” a relatively detailed description headed "'Kinixys Belliana, Gray, Griffith, Trans. Animal Kingd.” In the other publication (Gray 1831b) a very short description is provided under the heading “Bell’s Tortoise. Test. {Kinixys) Belliana, Gray.” The latter work has a note by Gray dated “Oct. 1830,” whereas the first has a preface dated “Jan. 1831.” The title pages for both works are dated 1831. Thus it appears that the shorter description was written first, although this does not clarify which version was published first. We here cite the more extensive description in the Cat¬ aphracta (Gray 1831a). For some time all southern African hinged-back tortoises were treated as a single species, Kinixys belliana Gray, 1831 (Loveridge and Williams 1957). However, Pritchard (1979) suggested that a more detailed investigation might confirm the validity of some of the form described earlier as Kinixys spekii Gray, 1863. Broadley (1981b) divided K. belliana into two sub¬ species, the typical form being restricted to the coastal plain, with the depressed form K b. spekii replacing it inland. He later recognized the two as specifically distinct (Broadley 1989a, 1993). Broadley (1989a) provided a distribution map for K spekii depicting Angolan records previously assigned to K belliana. However, he later stated that K. belliana extends to Angola (Broadley 1992a), whereas the presence of K. spekii had not yet been confirmed (Broadley 1993). Iverson (1986, 1992) placed K. spekii as a synonym of K. belliana and provided a map with all the known records from the country assigned to the latter. Mifsud and Stapleton (2014) and the Turtle Taxon¬ omy Working Group (2014) suggested that K. belliana occurs wiidely in Angola exclusive of the southeast and the true desert, whereas K. spekii is limited to the southeastern border areas with the Caprivi Strip and Zambia. On the other hand, Fritz and Havas (2007), Branch (2008) and Vetter (2011) considered K. belliana to be extralimital, with K. spekii widespread in Angola. A recent study of phylogeography, phytogeny and taxonomy of hinged-back tortoises (Kindler et al. 2012) found that the previously recognized savanna species K. belliana comprises three deeply divergent clades that are now treated as distinct species. Angolan populations are members of the clade that retains the name K. belliana, whereas West African and southeast African clades have been allocated to K. nogueyi (Lataste, 1886) and W zombensis Hewitt, 1931, respectively. Kindler et al. (2012) did identify K. spekii in Angola near the Namibian border, however, so it is highly like¬ ly that it occurs at least in southeastern Angola, although pending further investigation we list all K. belliana/K. spekii records together. Kinixys belliana Gray, 1831 Bell’s Hinge-Back Tortoise Kinixys Belliana Gray, 1831a:69. Holotype: BMNH 1979.919 (collector unknown). Type locality: “?” [“bought at Humphrey’s, West Africa” according the label on the type specimen fide King and Burke (1989)]. Cinixys belliana'. Boeage (1866a:40, 1895a:2), Boeage (1887c:209), Monard (1937b: 146). Kinixys belliana'. Schmidt (1933:4), Laurent (1950a: 13), Iverson (1986:158, 1992:271), King and Burke (1989:89), Broadley (1992a:12), Kindler et al. (2012:193), Turtle Taxonomy Working Group (2014:402), Ceriaco et al. (20 16b: 54). Kinixys belliana belliana'. Mertens (1937a:5, 1938a:430), Hellmich (1957a:32), Loveridge and Wiliams (1957:384), Loveridge (1957:170), Laurent (1964a:25), Broadley (1981b:208, 1992a: 13, 1993:47). Kinixys cf belliana'. Conradie et al. (2016:27). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I7I Global conservation status (lUCN): Not Evaluated [Least Concern 1996] Global distribution: We here follow the distributions presented by Kindler et al. (2012) in their supporting information. The species ranges from the Red Sea coast of Sudan and the Horn of Africa south to the Kenya-Tanzania border area and thence southwest to central Angola. Kinixys spekii Gray, 1863 Spek’s Hinged-Back Tortoise Kinixys spekii Gray, 1863a:381. Holotype: BMNH 1936.5.3.117 (collector J. H. Speke). Type locality: “Cen¬ tral Africa” probably “northwestern Tanzania” Broadley and Howell (1991). Kinixys belliana spekii: Broadley (19816:211). ''Kinixys spekii”: Broadley (1989a:52). Kinixys spekii: Fritz and Havas (2007:286), Branch (2008:53), Vetter (2011:79), Kindler et al. (2012:193), Ceriaco et al. (2014b:670), Turtle Taxonomy Working Group (2014:403), Ceriaco et al. (2016b:53). Global conservation status (lUCN): Not Evaluated. Global distribution: We here follow the distributions presented by Kindler et al. (2012) in their supporting information as modified in The Turtle Texonomy Working Group 2017). The species is known from southern Central Africa from Kenya and Uganda south to northern KwaZu¬ lu-Natal, South Africa (although not in the coastal plain and lowlands of southeastern Kenya, Tan¬ zania and Mozambique) and west to the Caprivi Strip of Namibia and adjacent Cuando Cubango, Angola. Kinixys erosa (Schweigger, 1812) Forest Hinge-Back Tortoise Testudo erosa Schweigger 1812:321. Syntypes: Two specimens originally in the Mannheim Museum and a specimen described and figured by Shaw (1802:59, pi. 13) (collectors unknown). Type locality: “America septentrionali” [in error]. Cinixys erosa: Bocage (1866a:40, 1895a: 1), Peters (1877a:611). Kinixys erosa: Laurent (1964a:24), Iverson (1986:159, 1992:272), Spawls et al. (2004:52), Fritz and Havas 2007:285, Branch (2008:50), Vetter (2011:75), Kindler et al. (2012:193), Luiselli and Diagne (2014:084.1), Mifsud and Stapleton (2014:26), Turtle Taxonomy Working Group (2014:402). Global conservation status (lUCN): Data Deficient. Global distribution: A widespread silvi- colous species in West Africa, preferring moist areas such as marshes and river banks from Gambia eastward through the Democratic Republic of Congo to Uganda, and south areas adjacent to the mouth of the Congo River. Ocurrences in Angola (Map 117): This species is limited to extreme northern regions of the country, including Cabinda. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a;40, 1895a:l; Mifsud and Sta¬ pleton 2014:31). Lunda Norte: “pres de la R. Camualua (ou Camuali), affl. de la rive gauche du Chicapa, a 1’ Guest de Dundo, Posto de Lovua” [-7.31667, 20.15000] (Laurent 1964a:24; Luiselli and Diagne 2014:084.6; Mifsud and Stapleton 2014:31); “affl. non Map 117. Distribution of Kinixys erosa in Angola. 172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II determine du Chicapa, a peu pres dans la meme region que le precedent” (which represent the locality cited above; “affl. de la rive gauche du Chicapa, a TOuest de Dundo, Posto de Lovua “) (Laurent 1964a:24); “Cambulo, 70 km from the border with Democratic Republic of Congo, Kin¬ shasa” [-7.73333, 21.26667] (Luiselli and Diagne 2014:084.6; Mifsud and Stapleton 2014:31). Taxonomic and distributional notes: Bour (in Iverson 1992) stated that the syntype figured by Shaw might have been in the Leverian Museum. This collection was sold at auction in 1806 and dispersed to a number of private and state-owned museums, but this specimen has not been located. Genus Stigmochelys Gray, 1873 Stigmochelys pardalis (Bell, 1828) Leopard Tortoise Testudo pardalis Bell 1828:420, Suppl. PI. 25. Type: Not loeated, originally in the Bell Colleetion, part of whieh was later donated to the OUM (King and Burke 1989). Type loeality: “in Promont. Bonae Spei” (eollector unknown), [= Cape of Good Hope], Western Cape Province, South Africa. Testudo pardalis-. Bocage (1867b:217, 1870:68, 1895a:3), Monard (1937b:147), Hellmich (1957a:32). Testudo pardalis pardalis'. Mertens (1937a: 5). Geoehelone pardalis babeoeki: Loveridge and Williams (1957:235). Geoehelone pardalis'. Iverson (1986:143, 1992:252), Broadley (1989b:43), Le et al. (2006:524). Stigmoehelys pardalis: Fritz and Bininda-Emonds (2007:305), Fritz and Havas (2007:297), Branch (2008:66), Fritz et al. (2010:348), Vetter (2011:91), Turtle Taxonomy Working Group (2014:406), Ceriaco et al. (2016a:54), Conradie et al. (2016:27). Global conservation status (lUCN): Least Concern. Global distribution: This species is distributed from southern Sudan and Ethiopia southwards through eastern Africa to South Africa, and northwestward to Namibia and southern Angola. Ocurrences in Angola (Map 118): This species occurs in southwestern Angola and eastwards along the Namibian border. Benguela: “Benguella” [-12.58333, 13.41667] (Bocage 1867b:217, 1895a:3; Hellmich 1957a:32, Loveridge and Williams 1957:251); “Cubal” [-13.03333, 14.25000] (Mertens 1937a:5; Loveridge and Williams 1957:251). Namibe: “Capangombe” [-15.10000, 13.15000] (Loveridge and Williams 1957:251; Ceriaco et al. 20 16a: 54); “Mossamedes” [-15.20000, 12.15000] (Bocage 1895a:3; Ceriaco et al. 2016a:54). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:147; Loveridge and Williams 1957:251); “Forte Rogadas” [-16.73333, 14.98333] (Monard 1937b: 147; Loveridge and Williams 1957:251); “Dombodola” [-17.33333, Map 118. Distribution of Stigmochelys pardalis in 14.83333] (Monard 1937b: 147; Loveridge and Williams 1957:251). Cuando Cubango: “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 27). Taxonomic and distributional notes: Bell’s (1828) description notes that the type was in his own collection at the time. The majority of the collection was purchased by F. W. Hope, who donat- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 173 ed it to the Oxford University Museum in 1862. However, a recent catalogue of the types there reveals that the type of S. pardalis is not present (Nowak-Kemp and Fritz 2010). Stigmochelys pardalis (Bell, 1828) was previously included in the genus Testudo and more recently, in the genus Geochelone (Loveridge and Williams 1957; Iverson 1992). Based on data from mitochondrial and nuclear genes, Le et al. (2006:524, 528) indicated a sister relationship between S. pardalis and Psammobates and subsumed it within the latter genus. However, based on morphological differ¬ ences, Fritz and Bininda-Emonds (2007) placed it in a monotypic Stigmochelys, which has been accepted by subsequent authors. Loveridge and Williams (1975) recognized two subspecies for Geochelone (= Stigmichelis) pardalis (Bell, 1 828) G. p. pardalis and G. p. babcocki, also distin¬ guished by Le et al. (2006) and Fritz and Havas (2007). Iverson (1992) formerly questioned the recognition of the two subspecies and recently Fritz et al. (2010) concluded that there is no ration¬ ale for recognizing subspecies within S. pardalis. Broadley (1989b) and Iverson (1986, 1992) pre¬ sented distribution maps for the species with some new Angolan localities but without mentioning the localities by name. Family Trionychidae Fitzinger, 1826 Genus Cycloderma Peters, 1854 Cycloderma aubryi (Dumeril, 1856) Aubry’s Softshell Turtle Cryptopodus Aubryi Dumeril 1856:374, pi. 20. Holotype, MNHN 8006 (collector C.E. Aubry-Lecomte). Type locality: “Gabon.” Cycloderma Aubryi: Peters (1877a:611), Bocage (1895a:8). Cycloderma aubryi: Loveridge and Wiliams (1957:453), Frade (1963:252), Iverson (1992:295), Gramentz (2008:117), Turtle Taxonomy Working Group (2014:411). Global conservation status (lUCN): Vul¬ nerable. Global distribution: Central African species known from the Congo Basin from Gabon to the Democratic Republic of Congo and Cabinda in Angola. Ocurrences in Angola (Map 119): This species occurs in the Cabinda enclave. Cabin¬ da: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611; Bocage 1895a:7; Loveridge and Williams 1957:459; Gramentz 2008:140); “Cabinda” [-5.55000, 12.18333] (Frade 1963:252), Turtle Taxonomy Working Group (2014:411). Taxonomic and distributional notes: Bour et al. (1995) discussed the type specimen. Map 119. Distribution of Cycloderma aubryi in Angola. Genus Trionyx Geoffroy Saint-Hilaire, 1809 Trionyx triunguis (Forskal, 1775) Aerican Soetshell Turtle Testudo triunguis ForskM 1775: ix. Holotype, lost fide Webb in Iverson (1992), (collector P. Forskal). Type locality: “In Nilo” [= the Nile], Egypt. Gymnopus aegyptiacus: Bocage (1867b:218). 174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Trionyx triunguis: Peters (1877a:611), Boeage (1895a:7), Loveridge and Wiliams (1957:423), Loveridge (1957:172), Saldanha (1966:8), Hughes (1979:203), Iverson (1986:196, 1992:318), Braneh (1998:43, 2008 74), Turtle Taxonomy Working Group (2014:420). Amyda triunguis: Mertens (1926:152). Amyda triunguis triunguis: Hellmieh (1957a:33). Global conservation status (lUCN): Vulnerable. Global distribution: This species is widespread in tropical West Africa south of the Sahara as well as through the Nile Valley and around the eastern Mediterranean as far as Turkey. A disjunct population occurs in the Jubba and Scabelle rivers (Somalia), draining the Ethiopian Highlands. Ocurrences in Angola (Map 120): This species occurs in western Angola, mainly in near coastal drainages from Cabinda to the Cunene mouth. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611; Boeage 1895a:7; Loveridge and Wiliams 1957:435). Bengo: “Cunga” [-9.23333, 13.76667] (Loveridge and Wiliams 1957:435). Luanda: “Loanda”[-8.83333, 13.26667] (Loveridge and Wiliams 1957:435). Kwanza Norte: “Cuanza, Mucoso” [-9.53333, 14.65000] (Hellmieh 1957a:33; Loveridge and Wiliams 1957:435). Benguela: “Lobito bay” [-12.35000, 13.55000] (Loveridge and Wil¬ iams 1957:435); “Catumbella” [-12.43333, 13.55000] (Boeage 1867b:218, 1895a:7; Mertens 1926:152); Loveridge and Wiliams 1957:435); “Benguella” [-12.58333, 13.41667] (Boeage 1895a:7; Loveridge and Wiliams 1957:435). Namibe: 11.80000] (Hughes 1979:203 [Fig. 67]). Taxonomic and distributional notes: Loveridge and Wiliams (1957) provided a detailed review of the species. Map 120. Distribution of Trionyx triunguis in Angola. Cunene mouth” [-17.28333, Order CROCODYLIA Gmelin, 1789 Family Crocodylidae Cuvier, 1808 Genus Crocodylus Laurenti, 1768 Crocodylus niloticus Laurenti, 1768 Nile Crocodile Crocodylus niloticus Laurenti 1768:53. Type: not loeated fide King and Burke (1989:11). Type loeality: “Indi- en und Agypten”, later restrieted to “Agypten” [= Egypt] by Fuehs et al. (1974a: 110). Crocodilus vulgaris: Gunther (1865a:480), Boeage (1866a:41, 1867b:218, 1895a:8), Peters (1877a:611), Monard (1937b: 150). Crocodilus niloticus: Ferreira (1903:16), Themido (1941:11), Hellmieh (1957a:31), Saldanha (1966:8), Branch and McCarteney (1992:3), Ceriaco et al. (2014b:669), Branch and Conradie (2015:200), Conradie et al. (2016:26). Crocodylus niloticus chamses: Fuchs et al. (1974a: 111). Global conservation status (lUCN): Least Concern. Global distribution: Widespread across much of eastern sub-Saharan Africa, northward along MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 175 Map Angola. 121. Distribution of Crocodylus niloticus in the Nile Valley into Egypt, and westwards across northern Namibia and southern Angola and thence northwards up the Atlantic coast to Cameroon. Also present in Madagascar. Ocurrences in Angola (Map 121): This species occurs across the country, despite the scarcity of publish data the Angolan popula¬ tion as traditionally been reported very high in almost all of the hydrographic basins. Cabin¬ da: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611). Lunda Norte: “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200). Luanda: “Rio Bengo” [-8.71667, 13.40000] (Bocage 1866a:41); “Loanda” [-8.83333, 13.26667] (Bocage 1866a:41). Malanje: “Pungo Andongo” [-9.66667, 15.58333] (Gunther 1865a:480); “Duque de Braganga” [-9.10000, 15.95000] (Ferreira 1903:16); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 20 14b: 669). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1867b:218). Benguela: “Alto Cubal” [-13.03333, 14.25000] (Hellmich 1857a:31). Huila: “fleuve Kuvangu” [-14.46667, 16.30000] (Monard 1937b:150). Cunene: “etangs duKuve- lai” [-15.65000, 15.80000] (Monard 1937b: 150). Cuando Cubango: “sight below the Mupupa Falls” [-17.51667, 20.05000] (Branch and McCartney 1992:3); “Cuito basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:9-10, 26); Cuito basin (35) observation” [-16.62322, 19.05352] (Conradie et al. 2016:9-10, 27); “Cubango basin (43) observation” [-17.53500, 23.18916] (Con¬ radie et al. 2016:9-10, 26); “Cuando basin (44a) observation” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 27); “Cubango basin (45) observation” [-16.88350, 18.01180] (Conradie et al. 2016:9, 12, 27). Taxonomic and distributional notes: Crocodylus niloticus is a widespread species through¬ out much of sub-Saharan Africa, however, some West African populations long assigned to C. niloticus are recognized as C. suchus (Geoffroy Saint-Hiliare, 1 807), a taxon recently resurrect¬ ed from synonymy (Schmitz et al. 2003). Fuchs et al. (1974a) previously recognized seven sub¬ species in C. niloticus and applied the name C. niloticus chamses to the Angolan population. The actual distributions of C. niloticus sensu stricto and C. suchus are poorly known in the region of central West Africa and Angola in particular (Fergusson 2010), thus it is unclear if Angolan popu¬ lations represent one or both of the species. Grigg and Kirshner (2015) provided a distribution map with the ranges of C. niloticus and C. suchus in Africa, which reflects the lack of data from central regions in Angola, but suggests that C. niloticus is the sole species occurring in the country, an interpretation we here accept. Branch and McCartney (1992) stated that the species is commonly found in all the major permanent river systems, including the Cuito and Cubango rivers. Hellmich (1957a) also refered to a large population in the Kwanza River, while Monard (1937b) mentioned sightings from Cunene near Forte Ro9adas, Capolongo along the Cuvango, and at the mouth of Catumbela. Monard (1932) identified the mythical creature named “Libata” by locals of Chiumbe and Cuilo (northeast of Angola) as C. niloticus. 176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Mecistops Gray, 1844 Mecistops cf. cataphractus (Cuvier, 1825) West African Slender-Snouted Crocodile Crocodilus cataphractus (Cuvier 1825:58). Holotype: RCSM 710 (collector unknown). Type locality: “Sene¬ gal, Fuchs et al. (1974b:3). Crocodilus cataphractus'. Peters (1877a:611), Bocage (1895a:9), Loveridge (1957:176), Frade (1963:252), Laurent (1964a:27). Crocodilus cataphractus cataphractus'. Fuchs et al. (1974b:3). Global conservation status (lUCN): Critically Endangered. Global distribution: A widely distributed species throughout West and Central Africa. Ocurrences in Angola (Map 122): This species occurs in northern Angola, close to the Congo basin. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611); “Zona de Cabinda “ [-5.55000, 12.18333] (Frade 1963:252); “cote de Loango” (Bocage 1895a:9). Lunda Norte: “Dundo, canal de la Mussungue” [-7.25000, 20.50000] (Laurent 1964a:27), “environs de Dundo” [-7.36667, 20.83333] (Laurent 1964a:27); “environs de Dundo, R. Mussungue, affl. du Luachimo” [-7.41667, 20.83333] (Laurent 1964a:27); “Dundo, R. Luachimo, pres de T embouchure du fleuve Dilolo” [-7.60000, 20.86667] (Lau¬ rent 1964a:27); “affl. de la Luembe ± 70 km au SE de Dundo” [-8.37140, 21.59368] (Laurent 1964a;27). Taxonomic and distributional notes: The taxonomy of African crocodiles is still unresolved. According to Schmitz et al. (2003) the species Crocodylus cataphractus (Cuvier, 1825) is not closely related to other Crocodylus species, and they considered C. cataphractus and Osteo- laemus tetraspis Cope, 1861 as sister species. McAliley et al. (2006) recommend the resurrection of the historic Genus Mecistops Gray, 1 844 to accommodate this species, based on recent molecu¬ lar and morphological analyses that support the distinction of M. cataphractus from Crocodylus species. The recent mitochondrial DNA analysis of Feng et al. (2010) corroborates this interpreta¬ tion. Historically, M. cataphractus was widely distributed throughout West and Central Africa but recent studies by Shirley (2013) and Shirley et al. (2013) have found molecular and morphological support for two divergent taxa in this genus, one distributed entirely in West Africa and the other in Central Africa (lUCN 2014). Mecistops from Angola belong to undescribed species, occuring in Cabinda Province and Lunda Norte (Grigg and Kirshner 2015), although Shirley (2010) and Shirley et al. (2014) considered the distribution to exclude northeastern Angola. Map 122. Distribution of Mecistops cf. cataphractus in Angola. Genus Osteolaemus Cope, 1861 Osteolaemus tetraspis Cope, 1861 African Dwarf Crocodile Osteolaemus tetraspis (Cope 1861:549). Syntypes: not located fide King and Burke (1989:14) (collector P. B. Du Chaillu [skin]; donation from Museum of the Pennsylvania University [skull of a half-grown individ¬ ual]). Type locality: “Ogobai River, Western Africa” (Cope 1861:550) [= Ogooue River], Gabon. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 177 Crocodylus fwntatus: Bocage (1866a:41), Peters (1877a:611). Crocodylus tetraspis: Boeage (1895a:9), Frade (1963:252), Grigg and Kirshner (2015:12). Global conservation status (lUCN): Vulnerable. Global distribution: Osteolaemus tetraspis sensu lato is known from across the tropical low¬ land regions of sub-Saharan West Africa and West Central Africa, from Senegal to the southeast¬ ern Democratic Republic of Congo. Osteolaemus tetraspis sensu stricto (see Notes below) occus south of the Cameroon Volcanic Line and primarily west of the Congo Basin. Ocurrences in Angola (Map 123): This species occurs in Cabinda enclave and in the north of the country, although all published records come from Cabinda alone. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:611; Bocage 1895a:252); “Rio Quilo” [-5.18333, 12.18333] (Bocage 1866a:41, 1895a;9); “Zona de Cabinda” [-5.55000, 12.18333] (Frade 1963:252). Taxonomic and distributional notes: The taxonomy of the African dwarf crocodile, Osteolaemus tetraspis has been under debate for many years (Eaton et al. 2008). The species was first described by Cope 1861, from the Ogooue Basin. A second dwarf crocodile genus, Osteoblepharon was proposed by Schmidt (1919) but regarded as a synonym of Osteolaemus by Mertens (1943) and Inger (1948). Eaton et al. (2008) revealed that the genus Osteolaemus contains at least three distinct species; O. tetraspis from the greater Ogooue Basin (including Gabon, portions of Cameroon and southwest Congo), O. osborni (Schmidt, 1919) from the Congo Basin, and a putative new species from West Africa. However, Franke et al. (2013) considered O. osborni as a subspecies of tetraspis. In Angola the species is confirmed only from Cabinda, which agrees with the current known distribution presented by Grigg and Kirshner (2015). Map Angola. 123. Distribution of Osteolaemus tetraspis in Order SQUAMATA Oppel, 1811 LIZARDS Family Gekkonidae Gray, 1825 Gtnm Afroedura Loveridge, 1944 Afroedura bogerti complex Loveridge, 1944 Bogert’s Rock Gecko Afroedura karroica bogerti Loveridge 1944:1, fig. 1. Holotype: AMNH 47841 (eolleetors H. and A. Chap¬ man). Type loeality: “Namba (Mombolo),” Kwanza Sul Provinee, Angola. Aftoedura ef bogerti: Branch (1998:232). Afroedura bogerti: Bates et al. (2014:93), Jacobsen et al. (2014:467), Branch et al. (2017:158). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and Namibia. 178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 124): The species was until recently only known from the type locality “Namba (Mombolo),” Kwanza Sul Province. The current distribution com¬ prise the southwestern regions along the coast from Kwanza Sul to Namibe Province. Kwan¬ za Sul: “Namba (Mombolo)” [-11.91667, 14.85000] (Loveridge 1944:1); “on track from Atome to Cassongue (vicinity of Missao da Namba)” [-11.91719, 14.86086] (Branch et al. 2017:162). Huambo: “10 km west of Soque” [-12.36256, 15.02847] (Branch et al. 2017: 161); “1 km south Luimbale” [-12.25369, 15.31689] (Branch et al. 2017:162); “Candum- bo Rocks, 16 km west of Vila Nova” [-12.73600, 15.97439] (Branch et al. 2017:162); “1 km west Kandumbo on road to BoasAguas” [-12.73614, 15.97442] (Branch et Map 124. Distribution of Afroedura bogerti in Angola. al. 2017:162). Benguela: “3 km west of Bicoio” [-12.48278, 14.10689] (Branch et al. 2017:162); “Praia do Meva (near Santa Maria)” [-13.39667, 12.58972] (Branch et al. 2017:162). Namibe: “Lucira road, 5 km south of Catara River” [-13.60431, 12.64556] (Branch et al. 2017:161); “turn off Morro do Chapeu Armando” [-14.52856, 12.50189] (Branch et al. 2017:161); “small granite outcrops in succulent veld, 52 km N on tar road in road to Lucira from junction with Lubango- Namibe road” [-14.65806, 12.52717] (Branch et al. 2017:162); “1 km east of Farm Mucongo” [-14.78361, 12.49694] (Branch et al. 2017:162); “Granite outcrops in sandy veld, 50 km E Namibe on main tar road to Leba” [-15.01558, 12.55503] (Branch et al. 2017:162); “Caraculo” [-15.02686, 12.65210] (Branch et al. 2017:161); “Tambor” [-16.06669, 12.44997] (Branch et al. 2017:161); “Furnas (? - ploted as nearest rock outcrops to centre of quarter-degree square (QDS) 1612Ab” [-16.39167, 12.14167] (Branch et al. 2017:161); “Omauha Lodge, 15 km south of Tambor” [-16.20061, 12.40183] (Branch et al. 2017:162); “0.5 km south of Tambor” [-16.07414, 12.43328] (Branch et al. 2017:162). Taxonomic and distributional notes: Onderstaal (1984) elevated the species to specific sta¬ tus (Jacobsen et al. 2014). There is deep genetic divergence among Angolan populations, sugges¬ tive of a complex of several species (Branch et al. 2017). Genus Afrogecko Bauer, Good and Branch, 1997 Afrogecko ansorgii (Boulenger, 1907) Ansorge’s Gecko (Endemic) Phyllodactylus Ansorgii Boulenger 1907a:212. Holotype: BMNH 1946.8.24.52-53 [2 speeimens] (eolleetor W.J. Ansorge). Type loeality: “Maeonjo, Benguela” [= Fazenda Mueungo], Namibe Provinee, Angola. Afrogecko ansorgii-. Bauer et al. (1997:476), Bates et al. (2014:100), Heinieke et al. (2014:31), Ceriaeo et al. (20 16a: 54). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 125): The species is known only from the type locality “Macon- jo.” Namibe: “Maeonjo (= Fazenda Mueungo)” [-14.782192, 12.486557] (Boulenger 1907a:212; Bauer et al. 1997:478; Ceriaeo et al. 2016a:54). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 179 Taxonomic and distributional notes: According to the original publication and Bauer et al. (1997) the species is restricted to Angola and known only from the type locality. Heinicke et al. (2014) in a revision of the cir- eum-Indian Ocean clade of leaf-toed geekos concluded that the phylogenetic position of Afrogecko ansorgii (Boulenger, 1907) is unclear, although they tentatively retained this species in Afrogecko pending further data. The species has recently been rediscovered (W.R. Branch, pers. comm., January, 2017) and is now known from several localities. I2”E l<)°E 20” E 24”E Map 125. Distribution of Afrogecko ansorgii in Angola. Genus Chondrodactylus Peters, 1870 Chondrodactylus fitzsimonsi (Loveridge, 1947) Button-Scaled Thick-Toed Gecko Pachydactylus laevigatus tessellatus FitzSimons 1938:172, fig. 6. Holotype: TM 17202 (collector V.F.M. FitzSimons). Type loeality: “Kamanyab” [= Kamanjab, Kunene Region] Namibia. Pachydactylus laevigatus fitzsimonsi Loveridge 1947:400. Nomen substitutum (see Notes below). Paehydaetylus laevigatus: Sehmidt (1933:5). Paehydaetylus laevigatus fitzsimonsi: Laurent (1964a:38). Paehydaetylus fitzsimonsi: Benyr (1995:50), Braneh (1998:255). Chondrodaetylus fitzsimonsi: Bauer and Lamb (2005:117), Ceriaco et al. (2016a:23, 54). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is restricted to central and northern parts of west¬ ern Namibia and adjacent Angola. Ocurrences in Angola (Map 126): The species occurs in southwestern Angola, mainly in Namibe Province. Huila: “Ongueria, 55 km au S.S.E. de Sa da Bandeira” [-15.30000, 13.51667] (Laurent 1964a:38). Namibe: “plage «das Conchas», pres de Mo^amedes” [-15.13333, 12.11667] (Laurent 1964a:38; Ceriaco et al. 2016b;54); “environs de Moga- medes, au bord de lar oute de Sa da Bandeira” (Laurent 1964a:38; Ceriaco et al. 20 16a: 54); “Espinheira” [-16.78731, 12.35817] (Ceriaco et al. 2016a:23); “Iona National Park, north of Tambor” [-15.99636, 12.40667] (Ceriaco et al. 2016a:23); “Pico Azevedo” [-15.53400, 12.49197] (Schmidt 1933:5). Map 126. Distribution of Chondrodactylus fitzsimonsi in Angola. 180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: The genus Chondrodactylus was previously considered a monotypic (C angulifer) but Bauer and Lamb (2005) demonstrated that a clade of large-bodied geckos, the Pachydactylus bibronii group (Lamb and Bauer 2002), is sister to C. angulifer and moved these taxa into Chondrodactylus. Pachydactylus laevigatus fitzsimonsi Loveridge, 1947 replaced the name P. 1. tessellatus FitzSimons, 1938 which was preoccupied by P. tessellatus (Werner, 1910), which is currently regarded as a synonym of Pachydactylus capensis (Smith, 1846). Reference to Schmidt’s (1933) plate image of “R laevigatus'' reveals that his specimens were, in fact, C. fitzsimonsi. Although cited in the chresonymy, specific localities from the unpub¬ lished theses of Benyr (1995) and Heinz (2011) have not been listed above. Chondrodactylus pulitzerae (Schmidt, 1933) Pulitzer’s Thick-Toed Gecko Pachydactylus bibronii pulitzerae Schmidt (1933:6, pi. 1). Holotype: CM 5619 (collector R. and L. Boulton). Type loeality: “Pieo Azevedo,” Namibe Provinee, Angola. Homodaetylus Bibroni: Boeage (1867b:220, 1867e:227, 1895a:15). Paehydaetylus bibronii: Boulenger (1885:201), Boeage (1895a: 15, 1887b:202, 1887e:209), Mertens (1926:152), Monard (1937b:53). Paehydaetylus bibronii pulitzerae: Parker (1936:129), Mertens (1937a:7, 1938a:431), Barbour and Loveridge (1946:164), Loveridge (1947:403), Hellmieh (1957a:36, 1957b:49), Laurent (1964a:37), Marx (1959:466). Paehydaetylus laevigatus laevigatus: Loveridge (1947:398; 1957:191). Paehydaetylus bibronii turneri (part): Loveridge (1947:405). Paehydaetylus laevigatus pulitzerae: Benyr (1995:50). Chondrodaetylus pulitzerae: Heinz (2011:55), Ceriaeo et al. (2014b:670), Ceriaeo et al. (2016a:24, 54), Heinieke et al. (2017:4), Ceriaeo et al. (2017:42). Chondrodaetylus ef pulitzerae (part): Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and far northwestern Namibia. Ocurrences in Angola (Map 127): The species occurs along the coast, mainly in the southern Angola (Fig 135). Luanda; "campus of the Universidade Metodista de Angola, in Caop Velha, Cacuaco Municipality [-8.78713889, 13.46361111] (Ceriaeo et al. 2017:42);" “Loanda” [-8.83333, 13.26667] (Boeage 1895a:15; Loveridge 1947:404). Malanje; “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 2014b:670). Huambo: “Huam- bo, zwischen den Fiipen Catumbella und Caporello, Benguela” [-12.76667, 15.73333] (Mertens 1926:152, 1937b:7; Loveridge 1947:404). Benguela: “Morro de Pundo” [-12.38333, 13.88333] (Parker 1936; 129; Loveridge 1947:404); “Catumbella” [-12.43333, 13.55000] (Boeage 1867b: 220; Loveridge 1947:404); “Quissange” [-12.43333, 14.05000] (Boeage 1887d:209; Loveridge 1947:404); “Benguela” [-12.58333, Map 127. Distribution of Chondrodactylus pulitzerae in Angola. 13.41667] (Boeage 1867b:220, 1895a:15; Boulenger 1885:201; Parker 1936:129; Loveridge 1947:404); “Dombe” [-12.95000, 13.10000] (Boeage 1867b:220; Loveridge 1947:404); “Entre Rios” [-13.01667, 14.63333] (Hellmieh MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I8I I957a:36); “Chivitidi” [-13.01667, 14.63333] (Hellmich I957a:36); “Cubal” [-13.03333, 14.25000] (Mertens I937a:7, I938a:43I; Loveridge 1947:404). Namibe: “Mossamedes” [-15.20000, 12.15000] (Bocage I867c:227, I895a:I5; Loveridge 1947:404; Laurent I964a:37; Ceriaco et al. 20I6a:54); “Pico Azevedo” [-15.55000, 12.51667] (Schmidt 1933:5-6; Barbour and Loveridge 1946:164; Loveridge 1947:398, 404; Marx 1959:466; Ceriaco et al. 20I6a:54); “Rio Coroca” [-15.78333, 12.06667] (Bocage I887b:202; Loveridge 1947:404; Ceriaco etal. 20I6a:54); “Iona National Park, 9.65 km (by air) west-south-west of Espinheira” [-16.81200, 12.27126] (Ceriaco et al. 2016a:24); “Espinheira” [-16.78639, 12.35799] (Ceriaco et al. 2016a:24); “Omauha Lodge” [-16.19861, 12.40087] (Ceriaco et al. 2016a:24); “Iona National Park, Rio Curoca in Pedi- va Hot Springs area” [-16.28359, 12.56106] (Ceriaco et al. 2016b:24); “Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road” [-15.01592, 12.64239] (Ceriaco et al. 2016a:24); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaco et al. 2016a:24). Taxonomic and distributional notes: The genus Chondrodactylus was previously considered monotypic (C. angulifer) but Bauer and Lamb (2005) demonstrated that a clade of large-bodied geckos, the Pachydactylus bibronii group (Lamb and Bauer 2002), is sister to C. angulifer and moved these taxa into Chondrodactylus. Pachydactylus bibronii pulitzerae was originally described by Schmidt (1933) from “Pico Azevedo” in southern Angola and was long considered a synonym of R bibronii (Smith, 1846) [= Chondrodactylus bibronii] (Ceriaco et al. 2014a). Benyr (1995), in an unpublished thesis, clarified the distinction between the more temperate C. bibronii and a more tropical Chondrodactylus laevigatus (Fisher, Chondrodactylus turneri (Gray, 1864) lineage and treated C. pulitzerae as a subspecies of Chondrodactylus laevigatus (see also Lamb and Basuer 2002). Recently, Heinz (2011) demonstrated that Chondrodactylus pulitzerae is a distinct, species-level lineage, occurring in sympatry or at least parapatry with C. laevigatus in southern Angola and in extreme northern Namibia. Most records for this species are in the south¬ west of Angola in Benguela Province (Schmidt 1933; Parker 1936; Monard 1937b; Mertens 1937a; Hellmich 1957a; Laurent 1964a), although Bocage (1895a) reported it to be common south of the Kwanza in general and cited two specimens from “Loanda” collected by Bayao in 1 874. A recent record from Capanda Dam is a northward range extension of the species. It has also recently been recorded Cacuaco, Luanda Province (Ceriaco et al. 2017), suggesting that C. pulitzerae may be more widely distributed throughout the savanna biome of Angola (Ceriaco et al. 2014b). Although cited in the chresonymy, specific localities from the unpublished theses of Benyr (1995) and Heinz (2011) have not been listed above. Chondrodactylus laevigatus (Fischer, 1888) Button-Scaled Gecko Pachydactylus laevigatus (Fischer 1888:15, pi. 2, fig. 3). Syntypes: ZMH (not located) (collector J. Stein- grover). Type locality: “bei Aus und auf dem Wege nach Bethanien,” [= Aus on the way to Bethanien, Karas Region] Namibia. Pachydactylus stellatus: Schmidt (1933:5). Pachydactylus bibronii-. Monard (1937b:53). Pachydactylus bibronii turneri (part): Loveridge (1947:405). Paehydaetylus bibronii pulitzerae'. Hellmich (1957b:49) Paehydaetylus laevigatus laevigatus'. Benyr (1995:7). Paehydaetylus turneri (part): Branch (1998:254). Chondrodaetylus turneri (part): Heinz (2011:30), Bates et al. (2014:104), Ceriaco et al. (2014b:670). Chondrodaetylus cf pulitzerae (part): Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. 182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global distribution: The species is known from the north of the Northern Cape Province, South Africa, throughout Namibia and into southern Angola and north and east as far as southern Kenya. Ocurrences in Angola (Map 128): The species appears to occur in Cunene, Huila, and Cuando Cubango provinces, with a single questionable record fom southern Namibe Province. Huila: “Kampulu (pres Kasinga)” [-15.21667, 16.11667] (Monard 1937b:53; Loveridge 1947:407); “Mulondo” [-15.63333, 15.20000] (Schmidt 1933:5; Loverige 1947:407-409); “Gambos” [-15.76667, 14.10000] (Hellmich 1957b:49). Cunene: “Humbi” [-16.68333, 14.90000] (Monard 1937b:53; Loveridge 1947:407); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:53; Loveridge 1947:407). Cuando Cubango: “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9); “Cuito basin (62)” [- 17.50875, 20.06608] (Conradie et al. 2016:9). Taxonomic and distribntional notes: , 128. Distribution of cWmdacofc in Angola. The genus Chondrodactylus was previously considered a monotypic (C. angulifer) but Bauer and Lamb (2005) demonstrated that a clade of large-bodied geckos, the Pachydactylus bibronii group (Lamb and Bauer 2002), is sister to C. angulifer and moved these taxa into Chondrodactylus. Benyr (1995) clarified the distinction between the more temperate Chondrodactylus bibronii (Smith, 1846) and a more northerly distrib¬ uted Chondrodactylus 1. laevigatus (Fisher 1888) and Chondrodactylus 1. turneri (Gray, 1864). Branch (1998) subsequently used the name turneri as a senior synonym of laevigatus, correcting Benyr’s oversight of the former name’s priority (Lamb and Bauer 2002; Ceriaco et al. 2014b) when the two are treated as conspecific. Heinz (2011) demonstrated that C. turneri and C. laevigatus are deeply divergent from one another, but Heinicke et al. (2017) were the first to employ C. laeviga¬ tus as specifically valid subsequent to its transfer to Chondrodactylus. Schmidt (1933) reported P. stellatus [syn. Chondrodactylus laevigatus^ from Pico Azevedo (the type locality of P. pulitzer- ae), but these specimens are referable to C.fitzsimonsi (see account above). Loveridge (1947) erro¬ neously considered Monard’s (1937b) material of Pachydactylus bibronii from “Humbi,” “Kam- pulo,” and “Mupanda” representative of Chondrodactylus turneri, a species now not believed to occur in Angola. Although cited in the chresonymy, specific localities from the unpublished theses of Benyr (1995) and Heinz (2011) have not been listed above. Genus Hemidactylus Oken, 1817 Hemidactylus angulatus complex Hallowell, 1852 African Tropical Gecko Hemidactylus angulatus Hallowell 1852a:63, fig. Holotype: ANSP 7431 (donated by H. Ford). Type loeality: “West eoast of Afriea” [= Gabon] by implieation fide Loveridge (1947). Hemidactylus brookii angulatus: Laurent (1964a:29). Hemidactylus brooki: Spawls et al. (2004:86). Hemidactylus angulatus: Carranza and Arnold (2006:539), Bauer et al. (2006a:8), Trape et al. (2012:222), Trape et al. (2012:222). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 183 Global conservation status (lUCN): Not Evaluated. Global distribution: The range of this speeies extends from Mauritania aeross mueh of the Sahel to Sudan and the Horn of Africa and south through East Africa. In Central Africa it extends as far south as northeastern Angola, although records in the region are scattered and its true distribution is uncertain. Ocurrences in Angola (Map 129): The species has been recorded only from “Alto Cuilo,” Lunda Sul Province. Lunda Sul; “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a;29). Taxonomic and distributional notes: This species was recently elevated to a full species based on molecular evidence that iden¬ tifies it as a member of a completely different major clade from the Asian Hemidactylus brookii Gray, 1845, under wich it was previously subsumed (Carranza and Arnold 2006; Bauer et al. 2006c). Hallowell’s type specimen was originally thought to have come from Liberia as the donor was “Mr. Henry Ford of Liberia,” however, the locality was subsequently corrected to Gabon (Loveridge 1947). Loveridge (1947, 1957) comidQrQd Hemidactylus bayonii Bocage, 1893 a syn¬ onym of H. angulatus and its type constituted his only Angolan record for this species. However, Laurent (1964a) stated that Loveridge ’s synonymy was incorrect and reported that the specimen that he identified from “Alto Cuilo” was actually the first H. angulatus to be captured in Angola. We recently examined this specimen, but it is in too poor a condition to verify specific identity. Numerous biological species are currently subsumed under the name H. angulatus (I. Agarwal et al., unpublished). Hemidactylus bayonii Bocage, 1893 Bayao’s Gecko (Endemic) Hemidactylus Bayonii Bocage 1893:116. Holotype: MBL specimen number not known (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Dondo, sur les bords du Quanza” [= Dondo, near Quanza river edges], Kwanza Norte, Angola. Hemidactylus bayonii: Bocage (1895a: 13, 1897a: 193), Frade (1963:252), Laurent (1964a:30). Hemidactylus brookii angulatus: Loveridge (1947:139, 1957:184). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 130): The species is only known from western regions of the Kwanza watershed. Kwanza Norte; “Dondo” [-9.68333, 14.43333] (Bocage 1893:116, 1895a:13, 1897a:193). Kwanza Sul: “31 km au nord-est de Novo Redondo” [-11.00000, 14.05000] (Laurent 1964a:30). Taxonomic and distributional notes: Loveridge (1947, 1957) considered the type specimen of Hemidactylus bayonii referable to H. angulatus. For some time this species was only known from the type locality, although Laurent (1964a) identified two individuals as Hemidactylus bayonii from “Kwanza Sul, 3 1 km from Novo Redondo”, not far from the type locality. Laurent (1964a) also provided a discussion about Loveridge’s (1947) mistaken synonymy of H. bayonii I2"E ICi^E 20* E 24*E Map 129. Distribution of Hemidactylus angulatus in Angola. 184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II with Hemidactylus brookii angulatus Hallow- ell, 1852. The species was recently recorded from Quigama National Park in Luanda Province (M. Marques, L. Ceriaco, A. Bauer and D. Blackburn pers. obs. 2015, 2016). Il-E HS“E lO-'E 24“E Map 130. Distribution of Hemidactylus bayonii in Angola. Hemidactylus benguellensis Bocage, 1893 Benguela Gecko (Endemic) This species is generally regarded as a syn¬ onym of Hemidactylus mabouia (Moreau De Jonnes, 1818) (Loveridge 1947, 1957; Uetz and Hosek 2017). However, H. mabouia is itself a complex of species (Bauer et al., unpublished) and we tentatively maintain the validity of H. benguellensis until both it and H. mabouia can be thoroughly revised. Hemidactylus benguellensis Bocage 1893:115. Syntypes: MBL specimen numbers not known (collector J.A. d’ Anchieta)”, destroyed by fire 18 March 1978. Type locality: “Cahata” [= Caota], Benguela Province, Angola. Hemidactylus benguellensis: Bocage (1895a:12, 1897a:193), Monard (1937b:52), Frade (1963:252). Hemidactylus mabouia: Loveridge (1947:170, 1957:185). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is i^-e i«°e 2(pe 24“e known from Angola. Ocurrences in Angola (Map 131): The species is only known from southwestern regions of the country. Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1893:115, 1895a:12, 1897a:115; Loveridge 1947:170, 1957:185); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:52). Taxonomic and distributional notes: Map 131. Distribution of Hemidactylus benguellensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 185 Hemidactylus longicephalus Bocage, 1873 Long-Headed Tropical Gecko Hemidactylus longicephalus Bocage 1873b:210. Syntypes: MBL specimen numbers not known (collector J.A. d’ Anchieta), destroyed by fire 18 March 1978. Type locality: “I’interieur de Mossamedes (Capangombe)” and “Catumbella, pres Benguella” [= Capangombe (Namibe Province) and Catumbella (Benguela Province)], Angola. Hemidactylus platycephalus: Bocage (1866a:42, 1866b:60, 1870:68; 1873b:209). Hemidactylus bocagii: Boulenger (1885:125), Bocage (1895a:ll, 1897a:193), Ferreira (1904:117). Hemidactylus longicephalus: Bocage (1887a: 178), Schmidt (1933:4), Parker (1936:128), Hellmich (1957b:49), Managas (1963:227), Loveridge (1947:187), Laurent (1964a:30), Ceriaco et al. (2014b:671), Ceriaco et al. (2016a:54). Hemidactylus bocagei: Ferreira (1906:170). Hemidactylus mabouia: Loveridge (1947:167). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from West Africa from Cameroon to Angola south into northern Namibia. Ocurrences in Angola (Map 132): The species is known from throughout the western regions of the country. Cabinda; “Landana” [-5.21667, 12.15000] (Loveridge 1947:189); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:42, 1866b:60, 1895a:ll, 1897a:193). Zaire; “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a:178, 1895a:ll; Loveridge 1947:189). Uige; “Fazenda Otilia- Encoge, Quitexe” [-7.55000, 15.03333] (Managas 1963:227). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1866a:42, 1866b:60). Bengo; “Ambriz” [-7.844312, 13.106493] (Boulenger 1885:125; Bocage 1895a; 11; Loveridge 1947:189); “Minas de Quizambil, Dande” [-8.31667, 13.73333] (Managas 1963:227); “Cunga” [-9.23333, 13.76667] (Ferreira 1904:117; Loveridge 1947:189). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:49); “Dondo” [-9.68333, 14.43333] (Bocage 1873b:209). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:128; Loveridge 1947:189); “Con- gulu” [-10.86667, 14.28333] (Parker 1936:128; Loveridge 1947:189); “Gumba” [-11.26667, 14.28333] (Ferreira 1904:117; Loveridge 1947:189). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1895a:ll, 1897a:193; Loveridge 1947:189); “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1885:125; Bocage 1895a:ll; Loveridge 1947:189); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:671). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:4; Loveridge 1947:189). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1873b:209, 1895a: 11, 1897a: 193; Loveridge 1947:189). Huila: “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] (Laurent 1964a:30). Namibe; “Capangombe” [-15.10000, 13.15000] (Bocage 1873b:209, 1895a:ll, 1897a:193; Loveridge 1947:189; Ceriaco et al. 2016a:54); “Rio Coroca” [-15.78333, 12.06667] (Bocage, 1895a:ll, 1897a:193; Loveridge 1947:189; Ceriaco etal. 2016a: 54). Undetermined Locality: “Carangigo” (Boulenger 1885:125; Bocage 1895a: 11; Loveridge 1947:189); “Cuanza River” (Loveridge 1947:189). Map 132. Distribution of Hemidactylus longicephalus in Angola. 186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: In the original description Bocage identified the species as Hemidactylus platycephalus, however, at the end of the description he mentioned that he had provisionally registered the species in the Museum catalogue under the name Hemidactylus longi- cephalus. Ferret (1975) believed that H. longicephalus Bocage, 1873 was a junior synonym of H. muriceus Peters, 1870 (see below), but he had been unable to see the type of the former to confirm this. Hemidactylus bocagii Boulenger 1885 was recognized as valid for Angolan speci¬ mens by Bocage (1895a, 1897a) and Ferreira (1904, 1906), although this name was synonymized by Loveridge (1947). The taxonomy of H. longicephalus is unstable and several names that are currently placed in its synonymy require further investigation. This species is broadly sympatric with the rather similar Hemidactylus mabouia (Moreau De Jonnes, 1818), throughout much of its range in Angola (Ceriaco et al. 2014a). Hemidactylus mabouia (Moreau De Jonnes, 1818) Tropical House Gecko Gecko Mabouia Moreau de Jonnes, 1818:138. Holotype: MHNP 6573. Type loeality: “en Amerique, dans les contrees eontinentales qui avoisinent au midi I’archipel des Antilles, et qu’il est egalement repandu dans les lies meme de I’archipel, depuis la Trinite jusqu’a la Jama'ique. “ Restricted to “St. Vincent” [Lesser Antilles] by Stejneger (1904:600). Hemidactylus mabouia'. Peters (1877a:612), Boulenger (1885:122, 1905:110), Bocage (1895a: 10), Ferreira (1904:117, 1906:170), Parker (1936:128), Loveridge (1947:167, 1957:185), Hellmich (1957a:34), Laurent (1950a: 12, 1954a:63, 1964a:29), Kluge (1969:28), Spawls et al. (2004:88), Bates et al. (2014:114), Branch and Conradie (2015:200), Ceriaco et al. (2016b:59). Hemidactylus mabouia mabouia'. Chirio and LeBreton (2007:118). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from almost all sub-Saharan Africa regions, although it is absent from more arid and temperate areas except under anthropogenic conditions. It is also currently found in North, Central and South America and in the Caribbean, where it is an anthropophilic invasive (Kraus 2009). Ocurrences in Angola (Map 133): The species occurs mainly in northern Angola, but also along the coast, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:612; Bocage 1895a; 10; Loveridge 1947:179); “Cabinda” [-5.55000, 12.18333] (Bocage 1895a:10; Loveridge 1947:179). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12, 1964a;29); “Carumbo, Lucapa” [-8.42278, 20.73917] (Branch and Conradie 2015:200). Luanda; “Loanda” [-8.83333, 13.26667] (Bocage 1895a: 10; Loveridge 1947:179; Hellmich 1957a:34). Bengo: “Cabiri” [-8.91667, 13.66667] (Ferreira 1903:117; Loveridge 1947:179); “Catete” [-9.11667, 13.70000] (Ferreira 1903:117; Loveridge 1947:179). Kwanza Norte: “Cambondo” [-9.15963, 14.65827] (Ferreira; 1906: 170; Loveridge 1947:179); “Mucoso bei Dondo” [-9.53333, 14.65000] (Hellmich 1957a;34); “Dondo” [-9.68333, 14.43333] (Bocage 1895a:10; Love- ll-E 2rE :4“E Map 133. Distribution of Hemidactylus mabouia in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 187 ridge 1947:179). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1895a: 10; Love- ridge 1947:179); “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1905:110; Loveridge 1947:179); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 20I6b:59). Bengue- la: “Lobito” [-12.35000, 13.55000] (Parker 1936:128; Loveridge 1947:179; Laurent I954a:63); “Entre Rios” [-13.01667, 14.63333] (Hellmieh I957a:34). Undetermined Locality: “Carangigo” (Boulenger 1885:122; Loveridge 1947:179); “Serra de Stombe” (Ferreira 1903:117; Loveridge 1947:179). Taxonomic and distributional notes: The type locality of H. mabouia was initially restrict¬ ed to “Antilles” by A. Dumeril in Dumeril and Dumeril (1851) and then more precisely to “St. Vin¬ cent” by Stejneger (1904). However, as noted by Kluge (1969), the holotype, which he believed was actually collected in Catragena, Colombia, is not conspecific with the gecko now typically regarded as H. mabouia, and would appear to be a specimen of H. angulatus (Carranza and Arnold 2006). Kluge recommended the maintenance of prevailing usage of the name. Hemidactylus mabouia as currently recognized includes several synonyms. However, it is known to be part of a complex of many species and the taxonomy of the complex is in flux. In Angola, H. mabouia is broadly sympatric with the rather similar Hemidactylus longicephalus Bocage, 1873 throughout much of its range (Ceriaeo et al. 2014b). Kluge (1969) mapped several Angolan localities, but did not provide specific localities. Hemidactylus cf. muriceus Peters, 1870 Guinean Spiny Tropical Gecko Hemidactylus muriceus Peters 1870:641. Holotype: ZMB 6921 (eolleetor R. Sehomburgk). Type loeality: “Keta (Guinea)” [= Keta, Ghana], Hemidactylus muriceus: Peters (1881:147), Boeage (1895a:13), Henle and Bohme (2003:33), Bauer et al. (2006a: 10). Hemidactylus ef muriceus: Ceriaeo et al. (2016b:59). Global conservation status (lUCN): Not Evaluated. Global distribution: This species occurs in West Africa forests, from Ghana to Cameroon, east to the Central African Republic and south to northern Angola. Ocurrences in Angola (Map 134): The species is known from northcentral Angola. Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:59). Undetermined Locality: “Cuango = Quango” (Peters 1881:147; Bocage 1895a: 13) (Malanje Province impossible to georefer¬ ence: See History Section for more detailed information). Taxonomic and distributional notes: The type was incorrectly cited as ZMB 69201 by Perret (1975). According to Perret (1975), Henle and Bohme (2003) and Bauer et al. (2006a) Hemidactylus muriceus Peters, 1870 has frequently been confused with other species. Perret (1975) attributed the confusion to Tornier’s (1902) use of the name for H. intestinalis Werner, 1897, which he consid¬ ered as senior synonym of H. ansorgii Map 134. Distribution of Hemidactylus cf. muriceus in Angola. 188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Boulenger, 1901. However, Henle and Bohme (2003) have argued that H. intestinalis is indeed a synonym of H. muriceus, and that H. ansorgii is the name applieable to another speeies. Ferret (1975) also believed that//, longicephalus Bocage, 1873 was a junior synonym of//, muriceus, but he had been unable to see the type of the former to confirm this. Because of those frequent misiden- tifications, literature records for the species are in many cases unreliable. The identity of Angolan specimens is unclear but they are probably not referable to H. muriceus sensu stricto. Genus Kolekanos Heinicke, Daza, Greenbaum, Jackman and Bauer, 2014 Kolekanos plumicaudus (Haacke, 2008) Feather-Tailed Gecko (Endemic) Afrogecko plumicaudus Haacke 2008:86, figs. 1-3. Holotype: TM 40527 (collector W.D. Haacke). Type local¬ ity: “Tambor (= turn-off of track to the south towards the Curoca River crossing into the Iona Park and the Kunene River mouth, marked by an empty 200 litre dmm),” Namibe Province, Angola. Afrogecko plumicaudus: Mashinini and Mahlangu (2013:167), Bates et al. (2014:100), Ceriaco et al. (2016a:55). Kolekanos plumieaudus: Heinicke et al. (2014:27), Agarwal et al. 2017:649. Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 135): The species is only known from the type locality “Tambor” and neighboring areas in Namibe Province, Southwest Angola. Namibe; “Tam¬ bor (= turn-off of track to the south towards the Curoca River crossing into the Iona Park and the Kunene River mouth, marked by an empty 200 litre drum” [-16.13333, 12.41667] (Haacke 2008:86; Mashinini and Mahlangu 2013:167; Ceriaco et al. 2016a:55); “Curoca River cross¬ ing (N bank)” [-16.30000, 12.43333] (Haacke 2008:86; Ceriaco et al. 20 16a: 55); “8 km /II km E of Iona, along track towards Oncocua, Parque Nacional do Iona” [-16.85972, 12.61111] (Haacke 2008:86, 90; Heincke et al. 2014:27, 38; Ceriaco et al. 2016a;55); Omahua" [-16.199642, 12.39877] (Agarwal et al. 2017:649). Taxonomic and distributional notes: Map 135. Distribution of Kolekanos plumicaudus in Angola. Heinicke et al. (2014) assigned A. pulmicaudus to a new genera Kolekanus, which is readily dis¬ tinguished from all other genera of African leaf-toed geckos based on its uniquely flattened tail with pointed lateral projections. This species is restricted to granite boulders on grassy, sandy plains with sXxxntQd Acacia mellifera thombush (Haacke 2008; Agarwal et al. 2017). Genus Lygodactylus Gray, 1864 Lygodactylus angolensis Bocage, 1896 Angola Dwarf Gecko Lygodactylus angolensis Bocage 1896a: 110. Syntypes: MBL (2 specimens) specimen numbers not known (collector J.A. d’ Anchieta), destroyed by fire 18 March 1978. Type locality: “Hanha, Benguella,” Benguela Province, Angola. Lygodactylus laurae Schmidt (1933:4). Holotype: CM 5860 (collector R. and L. Boulton). Type locality: “Chitau,” Bie Province, Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 189 Lygodactylus capensis: Bocage (1895a: 15), Laurent (1964a:31). Lygodactylus angolensis: Bocage (1870:68, 1873b:209, 1897a: 193), Loveridge (1947:207, 1957:187), Hellmich (1957a:35), Pasteur (1964:56), Branch (1998:245), Broadley and Cotterill (2004:41), Cerlaco et al. (20 16b: 57). Lygodactylus laurae: Mertens (1937a:6), Barbour and Loveridge (1946:147); Marx (1959:464), McCoy and Richmond (1966:154). Global conservation status (lUCN): Not Evaluated. Global distribution: The distribution oi Lygodactylus angolensis is poorly known in mueh of its range. It oeeurs from southern and eentral Angola east through Zambia, the former Katanga Provinee of the Demoeratie Repuiblic of Congo, northern Zimbabwe, northern and western Mozambique and Malawi to seattered loealities in Tanzania and southern Kenya. Ocurrences in Angola (Map 136): Lygo¬ dactylus angolensis is known from widely seat¬ tered loealities aeross the interior of Angola. Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco etal. 2016b:57). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:31). Moxico: “Falls of Luisavo, Poste de Calunda” [-11.86667, 23.58333] (Laurent 1964a:31). Bie: “Gauca” [-11.18333, 17.45000] (Sehmidt 1933:4; Loveridge 1947:208 Marx 1959:464); “Chitau” [-11.43333, 17.15000] (Sehmidt 1933:4; Bar¬ bour and Loveridge 1946:147; Loveridge 1947:208, 1957:187; Pasteur 1964:56; MeCoy and Richmond 1966:154); “Farm Goedecke on Conjo, 25 km north of General Machado (= Camacupa)” [-12.03333, 17.46667] (Mertens 1937a:6; Loveridge 1947:208). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1897a:193; Loveridge 1947:208). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:15, 1897a:193; Monard 1937b:53; Loveridge 1947:208); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:53; Loveridge 1947:208); “Hanha” [-13.01667, 14.63333] (Bocage 1896a:110, 1897a:193; Loveridge 1947:208); “Entre Rios” [-13.30000, 14.20000] (Hellmich 1957a:35). Taxonomic and distributional notes: Some citations to the name of Lygodactylus capensis (Bocage 1895a; Laurent 1964a) were later transferred to the synonymy of Z. angolensis. Pasteur (1964) provided a map (his fig. 15) depictng records for Angolensis, which indicates its wide range from the inland southwest of Angola, where it may be sympatric with Lygodactylus capensis (Smith, 1849). He also provided data on scale varation in this species. Recently, several individu¬ als of L. angolensis where collected in Cangandala National Park, Malanje Province (Ceriaco et al. 2016b), which confirms its northern extent in the country. Map 136. Distribution of Lygodactylus angolensis in Angola. Lygodactylus bradfieldi Hewitt, 1932 Bradfield’s Dwarf Gecko Lygodactylus bradfieldi Hewitt 1932:126, pi. 6, fig. 10. Syntypes: originaly deposited at Albany Museum, now ineluding BMNH 1946.8.30.69, MCZ R-33443 (eolleetor R.D. Bradfield), PEM R14574-82, R14586- 92. PEM material ineorreetly reported as lost by De Lisle et al. (2013). Type loeality: “farm Quiekbom near Okahandja,” Namibia. 190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Lygodactylus capensis: Bocage (1895a: 15), Loveridge (1947:208). Lygodactylus capensis capensis: Schmidt (1933:4) Lygodactylus bradfieldi: Branch (1998:246), Bates et al. (2014:117), Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Northern Cape through Namibia to southern Angola and northwestern Botswana, with an apparently isolated population in northwestern Limpopo Province (Branch 1998; Bates et al. 2014). Ocurrences in Angola (Map 137): Pre¬ sumably restricted to Namibe Province. Namibe: “Mucungu” [-14.78333, 12.48333] (Schmidt 1933:4; Loveridge 1947:212); “Capangombe” [-15.10000, 13.15000] (Bocage 1895a: 15). Cuando Cubango: “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9, 11, 24); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9,11,24). Taxonomic and distributional notes: Lygodactylus bradfieldi Hewitt, 1932 was for some time treated as a western race of L. capensis, although Pasteur (1964) revived it as a full species. Pasteur (1964) considered Angolan and northeastern Namibian records to be referable to L. capensis Smith, 1849, how¬ ever, records assigned to L. capensis from “Mucungu” (Schmidt 1933; Loveridge 1947) and “Capangombe” (Bocage 1895a), Namibe Province may belong to L. bradfieldi given the species distribution and adaptation to arid savanna habitats. Recently, Conradie et al. (2016) col¬ lected three specimens that correspond to L. bradfieldi from Cuito drainage, in Cuando Cubango Province. Namibian and Angolan specimens assigned to both L. bradfieldi and L. capensis require reexamination to confirm the species boundaries. Map 137. Distribution of Lygodactylus bradfieldi in Angola. Lygodactylus capensis (Smith, 1849) Cape Dwarf Gecko Hemidactylus capensis Smith 1849: pi. 75, fig. 3 and two associated unnumbered text pages. Syntpes: lost fide FitzSimons (1937) (eolleetor A. Smith). Type loeality: “Kaffirland, and the distriets to the north of Cape Colony,” South Affiea. Hemidactylus capensis: Boeage (1867b:219, 1870:68). Lygodactylus capensis: Boeage (1895a: 15), Monard (1937b:53), Loveridge (1947:208, 1957:187), Frade (1963:253), Pasteur (1964:62), Braneh (1998:246). Lygodactylus capensis capensis: Bates et al. (2014:118). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from chiefly savanna habitats of central and south¬ ern Africa, from warmer regions of South Africa north to Tanzania in the east and southern Ango¬ la in the west. Isloated records exist from southern Kenya and the south central Democratic Repub¬ lic of Congo. Ocurrences in Angola (Map 138): Published records for Lygodactylus capensis are from the southwestern provinces of Angola. Benguela: “Dombe” [-12.95000, 13.10000] (Bocage MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA I9I I867b:2I9, I895a:I5; Monard I937b:53; Loveridge 1947:212). Hulla: “Caconda” [-13.73333, 15.06667] (Bocage I895a:I5; Monard I937b;53; Loveridge 1947:212). Cunene: “Kuvelai, Kasinga” [-15.65000, 15.80000] (Monard I937b:53; Loveridge 1947:212). Namibe: “Chiyaka district” [-15.08333, 12.73333] (Loveridge 1947:212). Taxonomic and distributional notes: Pasteur (1964) provided a map (his fig. 15) showing the distribution of this species and provided data on scale variation and compar¬ isons with L. angolensis, which he regarderd as a member of the same species group. The species L. grotei Stemfeld, 1911 was previous¬ ly considered a subspecies of L. capensis (e.g., Pasteur 1964), which it replaces in the low¬ lands of Mozambique and Tanzania. Genetic data indicate that there are multiple unde¬ scribed species within the L. capensis complex (Travers 2012) and extensive taxonomic work will be necessary to delimit species in the group as a whole. Isolated records from southern Kenya and the Democratic Republic of Congo especially require investigation. Lygodactylus chobiensis FitzSimons, 1932 Okavango Dwarf Gecko Lygodactylus picturatus chobiensis FitzSimons 1932:35. Holotype: TM 14580 (collector V.F. FitzSimons). Type locality: “Kabulabula, Chobe River,” Botswana. Lygodactylus chobiensis'. Pasteur (1964:77), Haacke (1970:279), Branch (1998:247), Broadley and Martiz (2010). Global conservation status (lUCN): Least Concern. Global distribution: The species extends from the southeast of Angola and adjacent Namib¬ ia and northern Botswana to western Mozambique, northern Zimbabwe and southeastern Zambia. There is an apparently isolated population in northern Mozambique (Broadley and Maritz 2010a). Ocurrences in Angola: The species occurs in Angola, probably in both Moxico and Cuando Cubango provinces (see notes below). Taxonomic and distributional notes: Despite the lack of precise published locality records for Lygodactylus chobiensis from Angola, the currently accepted distribution for the species includes portions of Angola (Haacke 1970; Branch 1998; Broadley and Martiz 2010). Pasteur (1964) provided a provisional map of the distribution of the species from Lygodactylus picturatus group and noted that the species probably occurs in the most eastern areas of Moxico Province, Angola, adjacent to the border with Zambia. It may also be expected in at least southeastern Cuan¬ do Cubango as there are records nerby in the neighboring Caprivi Strip of Namibia. New surveys and aquisition of new fresh material is absolutely required to clarify L. chobiensis distribution. Lygodactylus lawrencei Hewitt, 1926 Lawrence’s Dware Gecko Lygodactylus lawrencei Hewitt 1926b:478. Holotype: SAM 17289 (eolleetor R. F. Lawrenee). Type loeality: “Otjitambi, Kaokoveld, S.W.A.” Kamanjab Constitueney, Kunene Region, Namibia. Lygodactylus lawrencei'. Pasteur (1964:70), Braneh (1998:247). i:°E lO'E. 2'0°E 24®E Map 138. Distribution of Lygodactylus capensis in Angola. 192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from northwestern Namibia and adjacent south¬ western Angola. Ocurrences in Angola: The species occurs in southwestern regions of Angola, near the border with Namibia, but there are no precise published records. Taxonomic and distributional notes: Lygodactylus lawrencei is a species of rocky, dry savannas from the Kaokoveld regions in northern Namibia, extending into southern Angola (Branch 1998). New surveys and acquisition of new material are absolutely required to clarify the range of Z. lawrencei in Angola, given the lack of precise published locality records for this species in the country. Genus Pachydactylus Wiegmann, 1834 Pachydactylus angolensis Loveridge, 1944 Angolan Thick-Toed Gecko (Endemic) Pachydactylus scutatus angolensis (Loveridge 1944a:3). Holotype: AMNH 47874 (eolleetors A. Vemay, H. Lang and R. Boulton). Type loeality: “Hanha, Benguela Provinee,” Angola. Pachydactylus scutatus angolensis: Barbour and Loveridge (1946:165), Loveridge (1947:357), Laurent (1964a:37). Paehydaetylus angolensis: Bauer et al. (2002a:26), Bauer and Lamb (2005:116), Bauer (2010:265); Heinieke et al. (2017:12), Ceriaeo et al. (2016a:25), Braneh et al. (2017:163). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is only known from Angola. Presumptive specimens from Namibia have been determined to be P. parascutatus Bauer, Lamb and Branch, 2002. Ocurrences in Angola (Map 139): The species occurs broadly in southwestern Angola, below the eascarpment. Benguela: “Lobito bay” [-12.35000, 13.55000] (Loveridge 1944:3; Bauer et al. 2002a:23); “Hanha, Benguela” [-12.24500, 13.70750] (Loveridge 1944:3; Barbour and Loveridge 1946:165; Bauer et al. 2002a:23; Bauer 2010:265; Branch et al. 2017:163); “24 km S Benguela” [-12.69747, 13.25222] (Bauer 2010:265; Branch et al. 2017:163); “30 km N of Dombe Grande” [-12.73097, 13.23024] (Bauer 2010:265; Branch et al. 2017:163). Namibe: “Lucira” [-13.86667, 12.53333] (Bauer 2010:265; Ceriaeo et al. 2016a:55; Branch et al. 2017:163); “San Nicolau” [-14.26667, 12.36667] (Bauer 2010:265; Ceriaeo et al. 2016a:55); “Lungo” [-14.31667, 13.20000] (Bauer 2010:265; Ceriaeo et al. 2016a:55; Branch et al. 2017:163); “Bentiaba” [-14.26667, 12.38333] (Branch et al. 2017:163); “Saco de Giraul” [-15.06833, 12.14222] (Bauer 2010:265; Ceriaeo et al. 2016a:55; Branch et al. 2017:163); “environs de Mogamedes (au bord de la route de Sa da Bandeira et dans le plateau littoral de la Plage das Conchas” [-15.16667, 12.15000] (Laurent 1964a:37; Ceriaeo et al. 2016a:55); “Namibe- Lubango road, 2 km E (by road) of Mangueiras, south side of the road” [-15.04361, 13.16000] (Ceriaeo et al. 2016a:25; Branch et al. 2017:163). Map 139. Distribution of Pachydactylus angolensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 193 Taxonomic and distributional notes: Bauer et al. (2002a) re-examined the types of P. s. angolensis and revised the P. scutatus group, revalidating P angolensis as a full species, although closely allied to P. scutatus. Bauer and Lamb (2005) did not include the species in their phylogenetic analyses of southern African Pachydactylus, but they suggested that it was a member of the “northwestern clade.” This has recently been verified by Heinicke et al. (2011), who subse¬ quently (Heinicke et al. 2017) identified a sister group relationship between P. angolensis and P caraculicus FitzSimons, 1959. Ongoing molecular work has identified putative species-level diversity within the species. Pachydactylus caraculicus FitzSimons, 1959 Angola Banded Thick-Toed Gecko Pachydactylus caraculicus FitzSimons 1959:407. Holotype: TM 22880 (collector C. Koch). Type loeality: “Caraeul, S. Angola” [= Caraeulo, Namibe Province, Angola]. Pachydactylus angolensis'. Laurent (1964a:37), Haaeke (1970:280), Braneh (1998:253), Bauer and Lamb (2005: 1 16), Bauer et al. (2002a: 12), Bauer (2010:266), Heinieke et al. (2017: 12), Mashinini and Mahlangu (2013:174); Ceriaeo et al. (2016a:55). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola and northwestern Namibia. Ocurrences in Angola (Map 140): The species is known from the type locality “Carac- ulo” and nearby areas in Namibe Province. Namibe: “Lungo, Vila Arriaga district, S. Angola” [-14.76667, 13.36667] (FitzSimons 1959:407; Ceriaeo et al. 2016a:55); “36 mi northwest of Mocamedes [Namibe]” [-14.82533, 12.54417] (Bauer 2010:266; Ceriaeo et al. 20 16b: 5 5); “Caracul, S. Angola” [-15.01667, 12.66667] (FitzSimons 1959:407; Haaeke 1970:280; Mashinini and Mahlangu 2013:174; Ceriaeo et al. 20 16a: 55); “Cima, Giraul River district” [-15.06667, 12.15000] (FitzSimons 1959:407; Ceriaeo et al. 2016a:55). Taxonomic and distributional notes: In the original description, FitzSimons (1959) considered Pachydactylus caraculicus to be geographically and phylogenetically intermedi¬ ate between Pachydactylus scutatus angolensis Loveridge, 1944 (= Pachydactylus angolensis) and Pachydactylus scutatus scutatus Hewitt, 1927. However, this interpretation was challenged by Laurent (1964a) who noted that P. s. angolensis occurs in sympatry with P. caraculicus in south¬ ern Angola, and therefore must be specifically distinct (Bauer et al. 2002a). Bauer and Lamb (2005) and Heinicke et al. (2011) provided phylogenetic analyses that demonstrated that P. caraculicus is part of a diverse, primarily rupicolous “northwestern clade” of Pachydactylus, and this has been verified by Heinicke et al. (2017), who identified a sister group relationship of P. caraculicus with P. angolensis Loveridge, 1944. Map 140. Distribution of Pachydactylus caraculicus in Angola. 194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Pachydactylus oreophilus complex McLachlan and Spence, 1967 Kaokoland Rock Gecko Pachydactylus oreophilus McLachlan and Spence 1967:4, figs. 1-2. Holotype: PEM 1503/67 [now PEM R1921] (eolleetors G.R. MeLaehland and J.M. Spenee). Type loeality: “20 miles West of Sesfontein,” Kunene Region, Namibia. Pachydactylus oreophilus'. Branch (1998:258), Bauer and Lamb (2005:116), Bauer (2010:266), Heinieke et al. (2011:12), Herrmann and Braneh (2012:98). Pachydactylus ef oreophilus'. Ceriaeo et al. (2016a:26, 55). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from northwestern Namibia and southwestern Angola. Ocurrences in Angola (Map 141): The species occurs in Namibe Province and in the arid southwest of Benguela Province. Bengue- la: “35 km S Dombe Grande towards Lucira” [-13.20979, 12.98017] (Bauer 2010:266). Namibe: “Assuncao” [-14.86667, 13.10000] (Bauer 2010:266; Ceriaeo et al. 2016a;55); “Caraculo” [-15.01667, 12.66667] (Bauer 2010:266; Ceriaeo et al. 2016b:55); “20 km W Virei” [-15.66667, 12.76667] (Bauer 2010:266; Ceriaeo et al. 2016a:55); “6 km S Rio Coroca towards Iona” (Bauer 2010:266; Ceriaeo et al. 20 16a: 55); “Saiona River, 25 km NW Cainde” [-15.40000, 13.20000] (Bauer 2010:266; Ceriaeo et al. 20 16a: 55); “Mutiambo River on road to Lucira” [-15.78333, 12.46667] (Bauer 2010:266; Ceriaeo et al. 20 16a: 55); “Tambor” [-16.13556, 12.42972] (Bauer 2010:266; Ceriaeo et al. 2016a:55); “7 km from Iona towards Oncocau, Iona Reserve” [-16.85831, 12.61275] (Bauer 2010:266; Ceriaeo et al. 20 16a: 55); “Furnas” (Bauer 2010:266; Ceriaeo et al. 20 16a: 55); “Omauha Lodge” [-16.19861, 12.40087] (Ceriaeo et al. 2016a:26). Taxonomic and distributional notes: Bauer and Lamb (2005) and Heinieke et al. (2011, 2017) demonstrated that P. oreophilus belongs to the diverse, largely rupicolous “northwestern clade” of Pachydactylus. Specimens currently allocated to this taxon, in fact, represent a non- monophyletic group of superficially similar forms (Bauer et al., unpublished), with P. oreophilus sensu stricto limited to Namibia. There are further species level differences among the Angolan populations (Bauer et al., unpublished). Map 141. Distribution of Pachydactylus oreophilus in Angola. Pachydactylus punctatus complex Peters, 1854 Speckled Thick-Toed Gecko Pachydactylus punctatus Peters 1854:615. Leetotype: ZMB 4799, designated by Loveridge (1947:354) by implication (see Bauer and Gunther 1991 :294). Type locality: “Sena und Tette” [= Sena and Tete], Mozam¬ bique, restricted to “Tete” by implication fide Bauer et al. (1995:55). Pachydactylus ocellatus: Bocage (1867b:220, 1895a: 16), Boulenger (1885:205, 1905:110), Frade (1963:253). Pachydactylus serval: Monard (1931:90, 1937b:54). Pachydactylus punctatus brunnthaleri'. Schmidt (1933:5). Pachydactylus punctatus punctatus'. Loveridge (1947:352), Laurent (1954a:63, 1964a:36), Hellmich (1957a:37). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 195 Pachydactylus amoenoides: Laurent (1964a:36). Pachydactylus punctatus: Bauer and Braneh (1995a:70), Braneh (1998:259), Bauer et al. (2006b:646), Bauer (1999:57), Bates et al. (2014:130), Cerlaeo et al. (2016a:26, 55). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies as presently eonstrued is widespread across southern Africa from the northwest and northeast of South Africa north to Malawi, the former Katanga Province of the Democratic Republic of Congo, and southern Angola. Ocurrences in Angola (Map 142): Pub¬ lished species records are from southwestern Angola, but members of this clade also occur in parts of central and southeastern Angola. Benguela: “Hanha” [-12.25000, 13.75000] (Bauer and Branch 1995a; 82), “Lobito” [-12.33333, 13.50000] (Laurent 1954a:63, 1964a:36; Bauer and Branch 1995a;82); “Catumbella” [-12.43333, 13.55000] (Bocage 1895a: 16; Loveridge 1947:356); “Benguella” [-12.58333, 13.41667] (Bocage 18676:220, 1895a; 16; Boulenger 1885:205; Loveridge 1947:356). Huila: “Indungu” [-14.81667, 16.26667] (Monard 1937b;54); “Capelongo” [-14.91667, 15.08333] (Bauer and Branch 1995a; 82); “Humpata, environs de Sa da Ban- deira” [-15.03333, 13.40000] (Laurent 1964a;36); “Kului” [-15.41667, 15.73333] (Monard 193 7b: 5 4). Namibe: “ km 60 sur la route de Mogamedes a Sa da Bandeira” [-15.00000, 12.66667] (Laurent 1964a;36; Ceriaco et al. 2016a:55); “11 miNE ofMocamede” [-15.08783, 12.26833] [Bauer and Branch 1995:82; Ceriaco et al. 2016a;55); “environs de Mogamedes” [-15.16667, 12.15000] (Laurent 1964a;36; Ceriaco et al. 2016a:55); “35 km south of Mogamedes” [-15.50000, 12.16667] (Laurent 1964a:36; Ceriaco et al. 2016a:55); “Pico Azevedo” [-15.55000, 12.51667] (Schmidt 1933:5; Loveridge 1947:356; Ceriaco et al. 2016a:55); “Iona National Park” [-16.65669, 12.43672] (Ceriaco et al. 2016a:26); “Espinheira” [-16.78639, 12.35799] (Ceriaco et al. 2016a;26). Cunene: “Riv. Mbale” [-15.16667, 16.75000] (Monard 1931:90); “Forte Rogadas” [-16.71667, 15.01667] (Laurent 1964a:36). Unde¬ termined Locality: “Cuanza” (Boulenger 1905:110; Loveridge 1947:356); “Arid subregion” (Frade 1963:253). Taxonomic and distributional notes: Members of the Pachydactylus punctatus complex have been confused with a number of other southern African Pachydactylus, including the species P. geitje (Sparrman, 1778) and P. serval Werner, 1910, South African and Namibian endemics, respectively (Loveridge 1947; Bauer and Branch 1995a). Bocage’s (1867b, 1895a), Boulenger’s (1885, 1905) and Trade’s (1963) Angolan specimens of P. ocellatus (= P. geitje) and Monard’s (1931, 1937b) Pachydactylus serval specimens from “Mbale,” “Indungu,” and “Kului” have all been redetermined as P. punctatus (Loveridge 1947; Bauer et al. 2006b). Bauer and Lamb (2005) and Heinicke et al. (2011, 2017) identified P. punctatus as part of the “northwestern clade” of Pachydactylus, which comprises numerous morphologically diverse species that are widely dis¬ tributed in southern Angola and northern Namibia. Laurent (1964a) considered Pachydactylus amoenoides Hewitt, 1935 to be distinct from P. punctatus based on their sympatry in southern Map 142. Distribution of Pachydactylus punctatus in Angola. 196 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Angola, and elevated it to a full species. Although occasionally considered a valid species or sub¬ species of P. punctatus, Bauer and Branch (1995a) tentatively included amoenoides in the syn¬ onymy of P. punctatus. Heinz (2011) identified deep, species-level divergences within P. puncta¬ tus sensu lato, and there are a minimum of four taxa in the complex that occur in Angola. A phy- logeographic study of P punctatus is being undertaken (Bauer et al. unpublished). Pachydactylus rangei (Andersson, 1908) Namib Web-Footed Gecko Palmatogecko rangei Andersson 1908:299, pi. 1, figs. a-c. Holotype: MWNH 460 (collector P. Range). “Luderitzbuch in the German South-West Africa,” Karas Region, Namibia. Palmatogecko rangei: Mertens (1937a:6), Haacke (1976b:22-23), Russel and Bauer (1990:198), Branch (1998:263), Bauer (1999:57). Pachydactylus rangei: Bauer and Lamb (2005:116), Herrman and Branch (2013:98), Bates et al. (2014:140), Ceriaco et al. (20 16a: 5 5). Global conservation status (lUCN): Least Concern. Global distribution: The species is restricted to the Namib Desert and immediate¬ ly adjacent areas, from the Holgat River in the Northern Cape, South Africa to extreme south¬ western Angola. Ocurrences in Angola (Map 143): The species occurs in the Nambi Desert, in south¬ western Angola. Namibe: “Mossamedes” [-15.20000, 12.15000] (Haacke 1976b;22-23; Ceriaco et al. 20 16a: 55); “Coroca River” [-15.78333, 12.06667] (Haacke 1976b;22-23; Ceriaco et al. 20 16a: 55); “Porto Alexandre” [-15.80000, 11.83333] (Haacke 1976b:22-23; Ceriaco et al. 2016a:55); “Cunene mouth” [-17.28333, 11.80000] (Haacke 1976b:22-23); “Lacrau” (Haacke 1976b:22-23; Ceriaco et al. 2016a:55); “Namib desert” (Mertens 1937a:6; Hermann and Branch 2013:98; Ceriaco et al. 2016a;55). Taxonomic and distributional notes: This species was initially described as the type species of the genus Palmatogecko Andersson, 1908 (Bates et al. 2014). Joger (1985) used immunological data to infer its close affinities with Pachydactylus and Bauer and Lamb (2005) synonymized the genus Palmatogecko with Pachydactylus, within which it is most closely allied to P. vanzyli (Steyn and Haacke, 1966) and P austeni Hewitt, 1923 (Heinicke et al. 2017). Pachydactylus rangei occu¬ pies compacted windward dune faces in the Namib sand seas and other areas of loose sand, includ¬ ing dry river beds, penetrating inland as far as the Namib fog belt (Haacke 1976b; Russell and Bauer 1990; Branch 1998; Bauer 1999; Bates et al. 2014). Map Angola. 143. Distribution of Pachydactylus rangei in Pachydactylus scherzi Mertens, 1954 Scherz’s Thick-Toed Gecko Pachydactylus scherzi Mertens 1954:175, fig. 1. Holotype: SMF 45696 (eolleetors R. Mertens and E. Seherz). Type loeality: “Welwitsehia-Flaehe der Namib am Sudwestrand des Brandberg-Massivs, Damaraland, Siidwest-Afrika” [= Welwitsehia area of the Namib on the southwestern slope of the Brand- berg massif, Damaraland], Namibia. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 197 Pachydactylus scherzi: Bauer and Branch (1995a:69), Bauer and Lamb (2005:116), Heinicke et al. (2011:12), Bauer and Branch (2012:84) Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from western Angola and Namibia, chiefly in asso¬ ciation with the Great Western Escarpment. Ocurrences in Angola (Map 144): The species occurs in southwestern Angola. Huila: “10 mi. NE Sa da Bandeira” [-14.81000, 13.61783] (Bauer and Branch 1995a:84). Taxonomic and distributional notes: Pachydactylus scherzi is a member of the P. punctatus complex of small mostly terrestri¬ al geckos distributed throughout the semi-arid and arid regions of southern Africa (Bauer and Branch 1995a; Heinz 2011), which is, in turn a member of the “northwestern clade” of Pachy¬ dactylus (Bauer and Lamb 2005; Heinicke et al. 2011, 2017). Although long overlooked (e.g.. Branch 1988), P. scherzi was treated by Bauer and Branch (1995a) as a full species, with several regional morphs. The single pub¬ lished Angolan record corresponds to Bauer and Branch’s (1995a) northern or “Sanitatas type form.” Heinz (2011) identified deep, species level divergences within P. punctatus sensu lato, and found that P. scherzi was embedded within this group. Aphylogeographic study of the P. punctatus complex is being undertaken (Bauer et al., unpublished). h f I ! ) ^ V — 1 - - 1 - 1 - Ji Map Angola. 144. Distribution of Pachydactylus scherzi in Pachydactylus scutatus Hewitt, 1927 Pachydactylus scutatus Hewitt 1927:395, pi. 23, fig. 2. Holotype: SAM 17471 (collector R.F. Lawrence). Type locality: “Kowaris, S.W.A.” [= Ko wares] Kunene Region, Namibia. Pachydactylus scutatus: Bauer et al. (1993:37), Branch (1998:260), Bauer (1999:56), Bauer (2010:266), Ceriaco et al. (2016a:25). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from northwestern Namibia and adja¬ cent Angola. Ocurrences in Angola (Map 145): The species occurs in southwestern Angola. Nami- be: “6 km S Rio Coroca, Iona” [-15.774892, 11.895472] (Bauer 2010:266); “Espinheira” [-16.79772, 12.35422] (Bauer 2010:266; Ceriaco et al. 2016a:25); “Iona, Iona Reserve” [-16.90000, 12.583333] (Bauer 2010:266). Scaly Thick-Toed Gecko I2''E 20°E 24“ E Map 145. Distribution of Pachydactylus scutatus in Angola. 198 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: Bauer et al. (2002a) re-examined the types of P. s. angolensis and revised the P. scutatus group, revalidating P. angolensis as a full speeies, although closely allied to P. scutatus. Ceriaco et al. (2016a) mistakenly stated that their Espinheira record was the first published for the country. Pachydactylus vanzyli (Steyn and Haacke, 1966) Namib Desert Gecko Kaokogecko vanzyli Steyn and Haacke, 1966:6. Holotype: CR 4000/7 (currently NHMW 1635) (collectors W. Steyn, P.S. Swart, C.G. Coetzee and W.D. Haacke). Type locality: “About 18 miles SW of Ompembe, Kaokoveld, South West Africa,” Kunene Region, Namibia. Kaokogecko vanzyli: Haacke (1976a:23). Palmatogecko vanzyli: Branch (1998:264). Pachydactylus vanzyli: Bauer and Lamb (2005:116), Ceriaco et al. (2016a:55). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from far northwestern Namibia and adjacent southern Angola. Ocurrences in Angola (Map 146): The species occurs in southern areas of Namibe Province. Namibe: “Espinheira” [-16.74477, 12.38186] (Haacke 1976a:25-26; Ceriaco et al. 2016a:55); “Kakolo windmill” [-16.66667, 12.33333] (Haacke 1976a:25-26; Ceriaco et al. 2016a:55). Taxonomic and distributional notes: This species was first described by Steyn and Haacke (1966) in a new genus, Kaokogecko, the affinities of which have long been recog¬ nized to be with Pachydactylus, and P. rangei, in particular (Haacke 1976a, 1976c; Joger 1985; Bauer 1990). Kluge and Nussbaum (1995) synonymized Kaokogecko with Palmatogecko and Bauer and Lamb (2005) subsequently synonymized the latter genus with Pachydactylus. These authors, as well as Heinicke et al. (2011, 2017), confirmed Map Angola. 146. Distribution of Pachydactylus vanzyli in K. vanzyWs sister group relationship to P. rangei. Pachydactylus wahlbergii (Peters, 1869) Wahlberg’s Kalahari Gecko Colopus Wahlbergii (Peters 1869:57, pi. fig. 1). Holotype: NHR 385 (collector J. Wahlberg). Type locality: “Damaralande” [= Damaraland], Namibia. Colopus wahlbergii wahlbergii: Haacke (1976b:30, 1984:173). Colopus wahlbergii: Branch and Lamb (2005:118). Pachydactylus wahlbergii: Heinicke et al. (2017:14). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southern Africa from southeastern Angola, through Namibia and Botswana to western Zimbabwe and Limpopo Province, South Africa. Ocurrences in Angola: The species occurs in southeastern Angola in Cuando Cubango Province, close to the Namibian border, although there appear to be no specific published records. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 199 Taxonomic and distributional notes: This species was originally described in the monotyp- ic genus Colopus Peters, 1869. Bauer and Lamb (2005) retained Colopus as valid despite poor molecular phylogenetic support. Heinicke et al. (2017), however, have conclusively shown that Colopus wahlbergii is embedded within Pachydactylus. Haacke (1976b) cited Colopus wahlbergii from Angola, noting its occurrence in southeastern areas near the Namibian border, but without providing specific localities. Genus Rhoptropus Peters, 1869 Rhoptropus afer Peters, 1869 Namib Day Gecko Rhoptropus afer Peters 1869:59, pi., figs. 2, 2a-e. Leetotype: ZMB 6149A (eolleetor J. Wahlberg), designat¬ ed by Loveridge (1947) by implieation fide Bauer and Gunther (1991) and Bauer et al. (1995a:56). Type loeality: “Damaralande” [northwestern Namibia]. Rhoptropus afer. Boeage (1870:68, 1873b:212, 1887b:203, 1895a: 16, 1897b:210), Boulenger (1885:217, 1910:643), Loveridge (1947:286), Bauer and Good (1996:643), Braneh (1998:267), Lamb and Bauer (2001:71), Ceriaeo et al. (2016a:55). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from northern Namibia and adjacent southern Angola, in the Namib Desert, although absent from the dune seas. Ocurrences in Angola (Map 147): The species is known from southern Namibe Province. Namibe: “Maconjo” [-15.01667, 13.20000] (Boeage 1873b:212; Ceriaeo et al. 2016a:55); “Capangombe” [-15.10000, 13.15000] (Boeage 1873b:212, 1895a:16, 1897b:210; Ceriaeo et al. 2016a:55); “Mossa- medes” [-15.20000, 12.15000] (Boulenger 1885:217; Ceriaeo et al. 2016a:55); “Rio Coro- ca” [-15.78333, 12.06667] (Boeage 1887b:203, 1895a:16, 1897b:210; Ceriaeo et al. 2016a:55). Taxonomic and distributional notes: The genus Rhoptropus is a monophyletic group of rupicolous and diurnal geckos endemic to Namibia and southern Angola (Bauer and Good 1996; Lamb and Bauer 2001). A record from Lake Ngami, northern 147. Distribution of in Angola. Botswana (as R. braconnieri (Thominot, 1878)) has long ago been discredited, with the locality in error and the type specimen (MNHN 294) referable to R. afer (see Bauer and Good 1996). Boeage (1873b, 1887b, 1895a) cited the s^QciQS Rhoptropus Peters, 1869 from Namibe Province (“Rio Coroca” and “Campagombe”), later he also identified one specimen from “Hanha,” Benguela Province (Boeage 1897b) as R. afer. Boeage (1897b) considered that the specimen captured in “Hanha” proved that the species probably extended further north, however the distribution of this species is limited to the Namib Desert only (Bauer and Good 1996; Branch 1998). Loveridge (1947) omitted the records from Boulenger (1885, 1910) for “Mossamedes” and “Angola,” respec¬ tively. 200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Rhoptropus barnardi Hewitt, 1926 Barnard’s Namib Day Gecko Rhoptropus barnardi Hewitt 1926a:413, pi. 35, figs. 1-3. Syntypes: SAM 16639 (6 specimens) (K.H. Barnard and R.F. Lawrence). Type locality: “near Eriksson’s Drift, Kunene River,” Cunene Province, Angola. Rhoptropus barnardi'. Laurent (1964a:35), Branch (1998:267), Ceriaco et al. (2016a:26, 55). Global conservation status (lUCN): Not Evaluated. Global distribution: Occurs mainly inland of the northern Namib Desert, from central Namibia to Southern Angola, reaching its eastern limit in the Otavi Highlands of northern Namibia. Ocurrences in Angola (Map 148): The species is known from Namibe and Cunene provinces in southwestern Angola. Cunene; “near Eriksson’s Drift, Kunene River” [-17.26944, 14.525] (Hewitt 1926a:41). Namibe: “ km 60 de la route de Mo^amedes a Sa da Bandeira” [-15.00000, 12.66667] (Lau¬ rent 1964a:35; Ceriaco et al. 2016a:55); “Approximately 7.35 km north-west (by road) of Pico Azevedo” [-15.47519, 12.46319] (Ceriaco et al. 2016a:26); “Omauha Lodge” [-16.20033, 12.40003] (Ceriaco et al. 2016a:26); “ Iona National Park, Rio Curoca crossing. North side ofthe river” [-16.30189, 12.42028] (Ceriaco et al. 2016a:26); “ Iona National Park, Rio Curoca crossing, south side ofthe river” [-16.30408, 12.41667] (Ceriaco et al. 2016a:26); “ Iona National Park, Rio Curo¬ ca in the Pediva Hot Springs area” [-16.28359, 12.56106] (Ceriaco et al. 2016a:26); “Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the road” [-15.04467, 13.15906] (Ceriaco et al. 2016a;27); “Iona National Park” [-15.77317, 12.33303] (Ceriaco et al. 2016a:27). Taxonomic and distributional notes: Schmidt (1933) reported some specimens identified as Rhoptropus barnardi Hewitt, 1926 but, absent appropriate comparative material, he was unaware that his specimens represented a new taxon (Bauer and Good 1996), Rhoptropus taeniostictus Laurent, 1964. Rhoptrops bradfieldi is broadly distributed in a variety of mainly rocky habitats from below the Escarpment and on the “Huila Plateau” (Haacke and Odendaal 1981). However, the range in Angola is poorly known, so if populations are continuous or fragmented is uncertain. Specific records from Huila have not been published but are relatively numerous. A record from “Novo Redondo” [= Sumbe] (Bauer and Good 1996) is in error, as there is no mention of this locality in the work which these authors cite (Haacke and Odendaal 1981). The discovery of a species that is superficially similar to R. barnardi (see Rhoptropus sp. below) neces¬ sitates the reexamination of specimens, especially those occurring north of Mossamedes (Namibe City) in order to determine the true distribution of both taxa. Map 148. Distribution of Rhoptropus barnardi in Angola. Rhoptropus benguellensis Mertens, 1938 Benguela Namib Day Gecko (Endemic) Rhoptropus boultoni benguellensis Mertens 1938a:431, figs. 4-5. Holotype: SMF 25275 (eolleetor W. Sehaek). Type loeality: “Cubal, 900m H., Prov. Benguela, Angola.” MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 201 Rhoptropus afer. Bocage (1897b:210). Rhoptropus ? boultoni: Parker (1936:127). Rhoptropus afer benguellensis: Mertens (1938a: 430, 431). Rhoptropus boultoni benguellensis: Mertens (1967:59), Hellmieh (1957a:37), Laurent (1964a:33), Bauer and Good (1996:643), Heinieke et al. (2017:6). Global conservation status (lUCN): Not Evaluated. Global distribution: Endemic to western Angola. Ocurrences in Angola (Map 149): The species is endemic to the southwest quadrant of Angola, north of the Namib. Benguela: “Bocoio” [-12.46667, 14.13333] (Parker 1936:127); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:431; Laurent 1964a:33; Mer¬ tens 1967:59; Bauer and Good 1996:643); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957b:37; Laurent 1964a:33); “Marco de Canavezes (Cubal de Ganda)” [-13.08333, 14.33333] (Laurent 1964a:33); “Hanha” [-13.30000, 14.20000] (Bocage 1897b:210). Taxonomic and distributional notes: Three subspecies of Rhoptropus boultoni, including R. b. benguellensis, were long recog¬ nized (Bauer and Good 1996; Bauer and Lamb 2001), but recent molecular phylogenetic 11 1 j .1 .L 11 .Li -jy Map 149. Distribution of Rhoptropus benguellensis in research has revealed that all three are specifi- cally distinct (Kuhn 2016; Heinieke et al. 2017) ^ and do not form a monophyletic group. Loveridge (1947) and Laurent (1964) regarded a number of earlier references to other Rhoptropus to be applicable to this taxon, and we here have followed them. Although formally published localities are from Benguela, recent collections have included material from as far north as the Cuanza River in central Angola (Kuhn 2016). This species occu¬ pies some of the most mesic environments of any member of the genus. In the original description of Mertens (1938), the two images of the type specimen were mistakenly labeld as Rhoptropus afer benguellensis. Rhoptropus biporosus FitzSimons, 1957 FitzSimons’ Namib Day Gecko Rhoptropus biporosus FitzSimons 1957:395. Holotype: TM 24198 (collector Bernard Carp/Transvaal Museum Expedition). Type locality: “Ompemhe, 120 miles west of Ohopoho [= Opuwo], Kaokoveld, Southwest Africa,” Kunene Region, Namibia. Rhoptropus biporosus: Bauer and Good (1996:643), Branch (1998:267), Ceriaco et al. (2016a:27, 55). Global conservation status (lUCN): Not Evaluated. Global distribution: Recorded only from inland of the northern Namib dune fields form extreme northern Namibia and into adjacent Angola. Ocurrences in Angola (Map 150): The species is present in southern Namibe Province below the Escarpment. Namibe: “Pico Azevedo” [-15.53400, 12.49197] (Bauer and Good 1996:643; Ceriaco et al. 2016a:55); “Iona National Park” [-16.53347, 12.44560] (Ceriaco et al. 2016a:27); “Iona National Park, 20 km south-south-west (by air) of Espinheira” [-16.93169, 12.24500] 202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (Ceriaco et al. 20I6a:27); “Espinheira” [-16.78886, 12.35761] (Ceriaco et al. 2016a:28). Taxonomic and distributional notes: Rhoptropus biporosus is closely allied to R. barnardi and one or more undescribed species of small-bodied congeners that occupy similar habitats in arid to semi-arid rocky areas bordering the Namib (Kuhn 2016). I2“E 24°E Map 150. Distribution of Rhoptropus biporosus in Angola. Rhoptropus boultoni Schmidt, 1933 Boulton’s Namib Day Gecko Rhoptropus boultoni Schmidt 1933:7, fig. 2, pi. 1 (far right). Holotype: CM 5634 (collectors R. and L. Boul¬ ton). Type locality: “Pico Azevedo,” Namibe Province, Angola. Rhoptropus boultoni'. Barbour and Loveridge (1946:187); Marx (1959:469); McCoy and Richmond (1966:155); Bauer and Good (1996:643). Rhoptropus boultoni boultoni'. Ceriaco et al. (2016a:28, 55). Global conservation status (lUCN): Not Evaluated. Global distribution: Occurs mainly inland of the northern Namib Desert, from central Namibia to southwestern Angola. Ocurrences in Angola (Map 151): The species is known from southwestern Angola, inland of the Namib and as far north as the desert’s northern limit in southern Namibe. Namibe: “Pico Azevedo” [-15.55000, 12.51667] (Schmidt 1933:7; Barbour and Loveridge 1946:181; Marx 1959:469; McCoy and Richmond 1966:155; Ceriaco et al. 20 16a: 55); “Iona National Park, 3.4 km south¬ west (by air) of Espinheira, vicinity of “Lion Cave” [-16.80375, 12.33978] (Ceriaco et al. 2016a;28); “Approximately 7.35 km north¬ west (by road) of Pico Azevedo” [-15.47589, 12.46269] (Ceriaco et al. 2016a:28); “Espin¬ heira” [-16.79150, 12.35168] (Ceriaco et al. 2016b:28); “Omauha Lodge” [-16.19792, 12.39981] (Ceriaco et al. 2016a;28); “Iona National Park, Rio Curoca crossing, south side of river” [-16.30408, 12.41667] (Ceriaco et al. I2"E l«°E 20° E 24°E Map 151. Distribution of Rhoptropus boultoni in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 203 20I6a:29); “Leba Pass, between river and highway” [-15.07033, 13.24381] (Ceriaeo et al. 20I6a;29); “Namibe-Lubango road, 2.0 km east (by road) of Mangueiras, south side of the road” [-15.04467, 13.15906] (Ceriaeo et al. 2016a:29); “Namibe-Lubango road, road marker 59, 1.8 km west by road of Caraculo” [-15.01531, 12.64244] (Ceriaeo et al. 2016a:29); “Pieo Azevedo” [-15.53400, 12.49197] (Ceriaeo et al. 2016a:29); “Espinheira” [-16.78894, 12.35775] (Ceriaeo et al. 2016a:29). Taxonomic and distributional notes: Three subspeeies of Rhoptropus boultoni have been reeognized (Bauer and Good 1996; Bauer and Lamb 2001), but reeent moleeular phylogenetie research has revealed that all three are specifically distinct (Kuhn 2016; Heinicke et al. 2017) and do not form a monophyletic group. Rhoptropus boultoni generally occupies areas of large boulders or occurs in association with large trees, such as baobab, Adansonia digitata (Bauer and Good 1996). There is deep genetic substructure in R. boultoni sensu stricto (Kuhn 2016) and further investigation may reveal that the taxon as now construed constitutes a species complex of similar, large-bodied Rhoptropus. Rhoptropus montanus Laurent, 1964 Mountain Namib Day Gecko (Endemic) Rhoptropus boultoni montanus Laurent 1964a:31. Holotype: MD 1854 (collector A. Barros Machado). Type locality: “Boca da Humpata, region de Sa da Bandeira” [Boca da Humpata, Lubango], Huila Province, Angola. Rhoptropus boultoni montanus: Bauer and Good (1996:644); Ceriaeo et al. (2016a:29), Heinicke et al. (2017:6). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is Endemic to southwestern Angola. Ocurrences in Angola (Map 152): All known localities are on top of the Great escarp¬ ment. Huila: “Boca de Humpata, Sa da Ban¬ deira” [-14.93333, 13.51667] (Laurent 1964a:31; Bauer and Good 1996:644). Namibe: “Leba Pass overlook” [-15.07700, 13.23292] (Ceriaeo et al. 2016b;29); “Leba Pass, between river and highway” [-15.07033, 13.24369] (Ceriaeo et al. 2016a;29). Taxonomic and distributional notes: Three subspecies of Rhoptropus boultoni, including R. b. montanus, were long recog¬ nized (Bauer and Good 1996; Bauer and Lamb 2001), but recent molecular phylogenetic research has revealed that all three are specifi¬ cally distinct (Kuhn 2016; Heinicke et al. 2017) and do not form a monophyletic group. Additional localities from recent collections (Ceriaeo et al. 2016a) reinforce that this is a high elevation species, occurring in cooler, more mesic environments than most of its congeners. We recently examined the holotype of this taxon, which is in poor condition, having gone com¬ pletely black and brittle after more than 60 years in formalin. Map Angola. 152. Distribution of Rhoptropus montanus in 204 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Rhoptropus taeniostictus Laurent, 1964 Angolan Namib Day Gecko (Endemic) Rhoptropus taeniostictus Laurent 1964a;33, figs. 5a-b. Holotype: MD 1967 (collector A. Barros Machado). Type locality: “ km 60 de la route de Mo^amedes a Sa da Bandeira” [= km 60 road from Nambie to Lubango], Angola. Rhoptropus barnardi: Schmidt (1933:6). Rhoptropus taeniostictus: Bauer and Good (1996:644), Ceriaco et al. (2016a:30, 56), Heinicke et al. (2017:6). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is endemic to southwestern Angola. Ocurrences in Angola (Map 153): All published records are from Namibe Province north of the Namib proper and below the Escarpement. Namibe: “Mucungu” [-14.78333, 12.48333] (Schmidt 1933:6; Ceria¬ co et al. 2016a:56); “60 km of the road of Mogamedes to Sa da Bandeira” [-15.00000, 12.66667] (Laurent 1964a:33; Ceriaco et al. 20 16a: 56); “Namibe-Lubango road, 2 km east (by road) of Mangueiras, south side of the road” [-15.04467, 13.15906] (Ceriaco et al. 2016a:30); “Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road” [-15.01611, 12.64369] (Ceriaco et al. 2016a:30). Taxonomic and distributional notes: Laurent (1964a) in the original description noted in some similarities between the new species R. barnardi Hewitt, 1926 and R. boul- toni Schmidt, 1933, although it is apparent that he regarded its closest affinities with R. barnardi. Kuhn (2016) found its phylogenetic position to be equivocal, with nuclear genes suggesting affini¬ ties with R. montanus, Laurent, 1964 and mitochondrial data placing it as the sister to R. barnardi + R. biporosus + R. benguellensis. Map 153. Distribution of Rhoptropus taeniostictus in Angola. Rhoptropus sp. (Endemic) Rhoptropus sp.: Ceriaco et al. (2016a:31). Global distribution: The species is known from Angola. Ocurrences in Angola (Map 154): The species is known from southwestern Angola. Namibe: “Espinheira” [-16.79242, 12.35400] (Ceriaco et al. 2016a:30); “Omauha Lodge” [-16.20033, 12.40003] (Ceriaco et al. 2016a:30); “Iona National Park, north of Tambor” [-15.99636, 12.40667] (Ceriaco et al. 2016a:30); “Leba Pass, between river and highway” [-15.07003, 13.24347] (Ceriaco et al. 2016a:30); “Namibe-Lubango road, 2 m east (by road) of Mangueiras, south side of the road” [-15.04467, 13.15906] (Ceriaco et al. 2016a:30). Taxonomic and distributional notes: This as yet undescribed taxon is superficially most similar to R. barnardi, but is most closely related to R. biporosus (Kuhn 2016). It has a significant elevational range, extending from near the coast to the Escarpment and appears to be parapatri cally distributed relative to R. biporosus. Recognition that there is a cryptic species in Angola MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 205 suggests that northern specimens of R. barnardi should be reexamined to confirm their specific identity. I’^F. 1i"F; 10" F. Map 154. Distribution of Rhoptropus sp. in Angola. Family Amphisbaenidae Gray, 1825 Genus Dalophia Gray, 1865 [see Monopeltis welwitschii account for comments] Dalophia angolensis Cans, 1976 Angolan Worm Lizard Dalophia angolensis Gans 1976:6, fig. 4. Holotype: IICT 167-1959 [= CZL 167] (collector F. Frade). Type locality: “Calombe, 7 km “west” of Vila Luso - Moxico Road” [= Calombe, 7 km west of Luena], Moxi- co Province, Angola. Monopeltis ellenbergeri: Monard (1931:97, 1937b:67). Monopeltis grand trnasvaalensis: Monard (1937b:67). Dalophia pistillum: Loveridge (1941c:436). Dalophia ellenbergeri-. Loveridge (1941c:433), Broadley (1997a:34), Gans (2005:30). Tomorupeltis eolobura luluae: Laurent (1964a:87). Dalophia angolensis'. Broadley et al. (1976:446), Gans (2005:30). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southeastern Angola and adjacent western Zambia. Ocurrences in Angola (Map 155): The species in known from the type locality and near areas, limited to eastern Angola. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Lau¬ rent 1964a:87, Gans 1976:10; Broadley et al. 1976:446). Moxico: “Lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:87; Gans 1976:10; Broadley et al. 1976:446); “Calombe, 7 km west of Vila Luso - Moxico road” [-11.83333, 19.93333] (Gans 1976:6; 12“E I6"E 2rE 24“E Map 155. Distribution of Dalophia angolensis in Angola. 206 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Broadley et al. 1976:446); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:87; Gans 1976:10; Broadley et al. 1976:446). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b:67; Loveridge 1941c:436; Broadley 1997a:34); “vient du Haut-Zambeze, c’est-a-dire d’une region geographique assez proche du Kubango” (Monard 1931:97). Taxonomic and distributional notes: According to Gans (2005) this species is known only from the type series, which is in conflict with Gans (1976) and Broadley et al. (1976), and is unsup¬ ported by any discussion or data. The interpretation of Gans (2005) would mean that this species is an Angolan endemic, however, in the absence of contradictory evidence we accept the earlier data (Gans 1976) that support at least one occurrence in adjacent northwestern Zambia. Dalophia ellenbergeri (Angel, 1920) Ellenberger’s Worm Lizard Monopeltis Ellenbergeri Angel 1920:615, figs. 1-2. Syntypes: MNHN 20.78-20.80 (collector M. Ellenberg- er). Type locality: “Lealui (Haut-Zambeze),” Zambia. Dalophia ellenbergeri-. Loveridge (1941c:433), Broadley (1997a:34), Gans (2005:30). Dalophia pistillum: Branch and McCartney (1992:2). Global conservation status (lUCN): Not Global distribution: The species is known from Zambia and Angola. Ocurrences in Angola (Map 156): A sin¬ gle published locality from southeastern Ango¬ la, near the Zambia border. Cuando Cubango: “approximately 50 km E of Cuito Cuanavale” [-15.23333, 19.61667] (Branch and McCartney 1992:2; Broadley 1997a:34). Taxonomic and distributional notes: Branch and McCartney (1992) collected one specimen near “Cuito Cuanavale” identified as Dalophia pistillum (Boettger, 1985) which, in accordance with Broadley (1997a) should rep¬ resent the first record of Dalophia ellenbergeri from Angola. Earlier Angolan records attrib¬ uted to this species (e.g., Monard 1931; Loveridge 1941c) are referable to Dalophia angolensis Gans, 1976. Evaluated. 12“E I6'=E 2(PE 24“E Map 156. Distribution of Dalophia ellenbergeri in Angola. Dalophia pistillum (Boettger, 1895) Blunt-Tailed Worm Lizard Monopeltis pistillum Boettger 1895:62. Syntypes: SMF 11833 (3 speeimens), formerly 5455 2a (eolleetor not stated). Type loeality: “Sambesi, Ostaffiea” [= Zambezi], Zambia. Monopelis granti kuanyamarum Monard 1937b:67. Holotype: MHNC 91.0623 (eolleetor A. Monard). Type loeality: “Mupanda,” Cunene Provinee, Angola. Monopeltis granti transvaalensis'. Monard (1937b:67). Dalophia pistillum'. Loveridge (1941e:435); Broadley et al. (1976:467), Haaeke (1984:172), Gans (2005). Global conservation status (lUCN): Not Evaluated. Global distribution: This species has the widest range in the genus, ranging from Angola through Zambia to Mozambique and south to Botswana, Namibia, and the Northern Cape Province of South Africa. Ocurrences in Angola (Map 157): Known from the southern Angola. Huila: “Kuvangu” MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 207 [-14.46667, 16.30000] (Monard I937b:68, Loveridge I94Ic:436); Cunene: “Mupanda” [-17.13333, 15.76667] (Monard I937b:67; Loveridge I94Ie:436; Broadley et al. 1976:467). Taxonomic and distributional notes: Loveridge (1941e) eonsidered the forms D. granti (Boulenger 1907), D. colobura (Boulenger 1910), D. transvaalensis (FitzSi- mons 1933), D. mossambica (Cott, 1934) and D. kaynuamarum (Monard, 1937) all syn¬ onyms of D. pistillum. Laurent (1964a) sug¬ gested that these eonstituted three speeies (Z). pistilum, D. mossambicus, D. colobura) with granti, transvaalensis and kuanyamarum potentially synonyms or valid raees of the first of these. Haaeke (1984) provided a map with some points in Angola, one of whieh matehes Map 157. Distribution of Dalophia pistillum in Angola. “Mupanda”, unlike the others from Cuando Cubango Province, which probably correspond to Dalophia angolensis (Gans, 1976) or possibly QVQn Dalophia ellenbergeri Angel, 1920. Genus Monopeltis Smith, 1848 Monopeltis anchietae (Bocage, 1873) Anchieta’s Worm Lizard Lepidosternon {Phractogonus) Anchietae Bocage 1873a:247, figs. 1-4. Holotype: MBL T46.525 (collector J.A. d’Anchieta) destroyed by fire 18 March 1978. Type locality: “Humbe, dans I’interieur de Mossamedes, pres des bords de la riviere Cunene” [= Humbe, near Cunene River], Cunene Province, Angola. Monopeltis okavangensis Monard 1931:95, fig. 5. Syntypes: MHNC 91.0614-15, NMBA 13331 (collector A. Monard). Type locality: “Vila da Ponte” and “Kakindo” [= Cuvango and Caquindo], Huila and Cuan¬ do Cubango provinces, Angola. Monopeltis devisi (Monard 1937b:69). Holotype: MHNC 91.0620 (collector A. Monard). Type locality: “Mupa,” Cunene Province, Angola. Monopeltis Anehietae: Bocage (1895a:28, 1897a: 194). Monopeltis anehietae: Loveridge (1941c:410), Frade (1963:252), Gans (1967:84, 1976:1, 2005:35), Broadley et al. (1976:377), Broadley (1997b:8), Branch (1998:124), Broadley and Measey (2010). Monopeltis okavangensis: Monard (1937b:68). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Angola, Namibia (Caprivi Strip and Damar- aland) and Botswana. Ocurrences in Angola (Map 158): The species occurs in the southern Angola, exclusive of the arid southwest. Huila: “Kuvangu (= Vila da Ponte)” [-14.46667, 16.30000] (Monard 1931:95, 1937b:69; Loveridge 1941c:410, 412; Gans 1967:84, 2005:35, Broadley et al. 1976:377, 381). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:69; Loveridge 1941c:410, 412; Gans 1967:84, 2005:35; Broadley et al. 1976:377, 381); “Humbe” [-16.68333, 14.90000] (Bocage 1873a:247, 1895a:28, 1897a:194; Loveridge 1941c:410; Gans 1967:84, 2005:35; Broadley et al. 1976:377, 381; Broadley 1997b:8). Cuando Cubango: “Kakindo (= Caquindo)” [-15.45000, 17.05000] (Monard 1931:95, 1937b:68; Loveridge 1941c:410, 412; Broadley et al. 1976:377, 381; Gans 2005:35). 208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: MHNC specimens of the nominal taxa described by Monard (1931, 1937), were orig¬ inally without a formal catalogue numbering system and were listed as “LCFM (unnum¬ bered) by Gans (2005)” and earlier workers. The synonyms Monop eltis okavangensis Monard, 1931 and Monopeltis devisi Monard, 1937 are based on the most northemly records for the species (Loveridge 1941c, Broadley et al. 1976; Gans 2005). Monopeltis anchietae is sympatric with M. capensis Smith, 1848 at some localities in the western part of its range (Broadley et al. 1976), and also with M. infus- cata Broadley, 1997 at “Humbe” in Cunene Province (Broadley 1997b). I1“E i6‘=E 2»*’E 24°E Map 158. Distribution of Monopeltis anchietae in in Angola. Monopeltis infuscata Broadley, 1997 Infuscate Wedge-Snouted Worm Lizard Monopeltis infuscata Broadley 1997b:6. Holotype: NMZB 6072 (collector W. Howells). Type locality: “Not¬ tingham Estates, Beitbridge District, Zimbabwe.” Monopeltis capensis'. Bocage (1873b:216, 1895a:28), Loveridge (1941c:425), Gans (1967:85). Monopeltis capensis capensis'. Loveridge (1941c:425). Monopeltis sphenorhynchus'. Gans (1997:47). Monopeltis capensis capensis (var. B): Broadley et al. (1976:388). Monopeltis infuscata'. Branch (1998:126), Gans (2005:36). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known ia to Northern Cape, southwestern to Botswana. Ocurrences in Angola (Map 159): The species occurs in southern Angola adjacent to the Namibian border, exclusive of the arid southwest. Cunene; “Humbe” [-16.68333, 14.90000] (Bocage 1873b:216, 1895a:28; Loveridge 1941c:425; Gans 1967:86; Broadley et al. 1976:388; Broadley 1997b:8). Cuando Cubango: “Tumbo (= Tumbole River)” [-14.45462, 16.59369] (Broadley et al. 1976:388; Broadley 1997b:8). Undetermined Locality: “Sturuba” (Broadley et al. 1976:388; Broadley 1997b:8). Taxonomic and distributional notes: Broadley et al. (1976) identified three allopatric forms of Monopeltis capensis capen¬ sis (A. Smith, 1848) that differed mainly in from southwestern Angola, south through Namib- 12°E I6‘=E 2»°E 24°E Map 159. Distribution of Monopeltis infuscata in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 209 some differences and tansferred the Angolan records to Group B (= var. B). Later, Broadley (1997b) described Monopeltis infuscata Broadley, 1997 allocating these specimens to the new taxon (Gans 2005). Monopeltis infuscata is sympatric With Monopeltis anchietae Bocage, 1973 at “Humbe,” Cunene Province (Broadley 1997b). Monopeltis luandae Gans, 1976 Luanda Worm Lizard (Endemic) Monopeltis luandae Gans 1976:3, fig. 2. Holotype: AMNH 111338 (collector J.A. Quartan). Type locality: “on the ground in Luanda on the road toward the mouth of the Quanza River, Angola.” Monopeltis luandae: Broadley et al. (1976:437), Gans (2005:36). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 160): The species is known from the type locality “Luan¬ da.” Luanda: “on the ground in Luanda on the road toward the mouth of the Quanza River” (Gans 1976:3, 2005:36; Broadley et al. 1976:437, 440); “Luanda” [-8.83333, 13.26667] (Gans 1976:3, 2005:36; Broadley et al. 1976:437, 440); “Airport, Luanda” [-8.83333, 13.26667] (Gans 1976:3, 2005:36; Broadley et al. 1976:437, 440). Taxonomic and distributional notes: This species is only known from the type series. Monopeltis perplexus Gans, 1976 Monopeltis perplexus Gans 1976:6, fig. 3. Holotype: AMNH 47732 (collector Vemay-Angola Expedition). Type locality: “Hanha or Capelongo,” Angola. Monopeltis perplexus: Broadley et al. (1976:413), Gans (2005:37). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola. Ocurrences in Angola (Map 161): The type locality is one of the two sites “Hanha” or “Cape¬ longo.” Benguela: “Hanha” [-13.30000, 14.20000] (Gans 1976:6, 2005:37; Broadley et al. 1976:413, 416). Huila: “Capelongo” [-14.88333, 15.08333] (Gans 1976:6, 2005:37; Broadley et al. 1976:413, 416). Taxonomic and distributional notes: Monopeltis perplexus is only known from the type locality, which is one of two disparate sites (Broadley et al. 1976c; Gans 2005). This species is allopatric from all other forms, differing from Monopeltis capensis Smith, 1848 and Monopletis anchietae (Bocage, 1873) (Broadley et al. 1976). I2'"H Zlt^K Map 160. Distribution of Monopeltis luandae in Angola. Wedge-Snouted Worm Lizard (Endemic) 210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II I2“E 20“E 24°E 6*S Krs \4°S IS°.S Map 161. Distribution of Monopeltis perplexus in Angola. Monopeltis vanderysti de Witte, 1922 Vanderyst’s Worm Lizard Monopeltis Vanderysti de Witte 1922:66, pi. 1, figs. 1, la-e. Leetotype: MRAC 1872 (collector R.P. Vanderyst) designated by Gans (1967). Type locality: “Wombali, Leverville (dist. du Kwango); Lac Leopold II; Kasai” [Democratic Republic of Congo], restricted to “Wombali (dist. Lac Leopold II), Zaire” by Gans (1967:86-87). Monopeltis vanderysti vilhenai Laurent 1954a:66, figs. 16-18. Holotype: MRAC 17503 (formerly MD 5040) (collector A. Barros Machado). Type locality: “Region de Dundo” [= Dundo], Lunda Norte Province, Angola. Monopeltis vanderysti vilhenai'. Laurent (1964a:84), (Gans 1867:87). Monopeltis vanderysti'. Broadley et al. (1976:431), Gans (2005:38). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from south-central Democratic Repub¬ lic of Congo into adjacent Angola. Ocurrences in Angola (Map 162): The species is known from the far north of Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:66, 1964a:84; Gans 1967:87; Broadley et al. 1976:437; Gans 2005:38). Uige: “Sanza Pombo” [-7.14833, 16.37422] (Broadley et al. 1976:436; Gans 2005:38). Taxonomic and distributional notes: According to Broadley et al. (1976) the description of Monopeltis vanderysti vilhenai by Laurent (1954a) contains various errors, and the purported diagnostic characters do not differentiate it from the nominotypical form (Gans 2005). I1"E Ki°E irE 24°E Map 162. Distribution of Monopeltis vanderysti in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 2II Monopeltis welwitschii (Gray, 1865) Welwitsch’s Worm Lizard (Endemic) Dalophia Welwitschii Gray 1865a:455, figs. 7-8. Syntypes: BMNH 1946.8.20.89-90 (formerly BMNH 64.7.13.34-35) (collector F. Welwitsch). Type locality: “Angola; Pungo-Andongo,” Malanje Province, Angola. Monopeltis Welwitschii: Gray (1865b:377), Bocage (1895a:29, 1897a: 194). Monopeltis welwitschii: Loveridge (1941c:431), Frade (1963:252), Gans (1967:87, 2005:38). Dalophia Welwitschii: Broadley et al. (1976:376). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known only from Angola. Ocurrences in Angola (Map 163): The species is only known from the type locality “Pungo-Andongo” but may extend to the south of the type locality and to other provinces (e.g., Kwanza Sul; Bie). Malanje: “Pungo-Andon¬ go” [-9.66667, 15.58333] (Gray 1865a:455, 18656:377; Bocage 1895a;29, 1897a;194; Loveridge 1941c:432; Gans 1967:87, 2005:38; Broadley et al. 1976:376). Taxonomic and distributional notes: This species description (Gray, 1865a) pub¬ lished in the “Proceedings of the Zoological Society of London” was reprinted (Gray 1865b) in “Annals and Magazine of Natural History” under the heading Proceedings of Learned Societies, creating some subsequent confusion as to the correct citation of the , r^c\nr\ 4. ^ 4.\ 4. Map 163. Distribution of Monopeltis welwitschii in paper. Broadley et al. (1976) noted that Dalophia welwitschii Gray, 1865 was the species of the Dalophia group closest to Monopeltis. In fact, Dalophia welwitschii Gray, 1865 is the type species of the genus Dalophia, yet despite this Gans (2005) both recognized Dalophia as valid and listed this taxon under Monopeltis. This paradox appears to have been an oversight on the part of Gans (2005). Resolution through phylogenetic analysis has been hampered by the fact that M. welwitschii is known only from the type specimens. Should it actually belong in Monopeltis, the next available generic name for remaining Dalophia would be Tomuropeltis Laurent, 1947 (type spQcms Monopeltis giganteus Peracca, 1903). Genus Zygaspis Cope, 1885 Zygaspis nigra Broadley and Gans, 1969 Black Round-Snouted Worm Lizard Zygaspis niger Broadley and Gans 1969:1, figs. 1-2. Holotype: NMZB 6698 (formerly UM 6698) (eolleetor R.G. Japp). Type loeality: “Kalabo, Barotse Provinee,” Zambia. Zygaspis quadrifrons capensis: Laurent (1964a: 84). Zygaspis niger: Broadley and Gans (1975:21). Zygaspis nigra: Broadley and Broadley (1997: 13), Braneh (1998: 124), Gans (2005:39), Broadley and Measey (2016:118). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from eastern Angola, western Zambia, the Caprivi Strip of Namibia and into adjacent northern Botswana. 212 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Zygaspis nigra is sympatric with Z quadri- frons (Peters, 1862) in portions of its range (Broadley and Broadley 1997), although allopatrie in eastern Angola and Namibia. The typical habitats for this species are miombo woodlands and Baikiaea woodlands on sandy s Map 164. Distribution of Zygaspis nigra in Angola. Is (Broadley and Broadley 1997; Branch 1998). Zygaspis quadrifrons (Peters, 1862) Kalahari Round-Snouted Worm Lizard Global conservation status (lUCN): Not Evaluated. Ocurrences in Angola (Map 164): Pub¬ lished records in Angola derived from the northeastern provinces. Lunda Sul: “Alto Chi- capa” [-10.88333, 19.23333] (Laurent 1964a: 84; Broadley and Gans 1969:1; Broadley and Broadley 1997:14). Moxico: “Gago Coutinho District” [-14.101654, 21.435308] (Broadley and Gans 1969:1; Broadley and Broadley 1997:14; Broadley and Measey 2016:118, 122); “Colanda 7 km E of Vila Luso” (Broadley and Gans 1975:21; Broadley and Broadley 1997:14; Broadley and Measey 2016:118). Taxonomic and distributional notes: I3°E 16“E 20'=E 24°E Amphisbaena quadrifrons Peters 1862a:25. Syntypes: ZMB 4202-4204 (eolleetor C. H. Hahn). Type loeali- ty: “Neu-Barmen, im Hereolande, an der Westkiiste von Afrika” [= Gross Barmen, Otjozondjupa Region], Namibia. Amphisbaena ambuellensis Monard 1931:93, figs. 1-4. Syntype: MHNC 91.0621 (eolleetor A. Monard). Type loeality: “Chimporo” and “Caquindo” [= Tehimpolo and Caquindo], Cunene and Cuando Cubango provinees, Angola respeetively. Amphisbaena ambuellensis: Monard (1937b:65). Zygaspis quadrifrons quadrifrons: Loveridge (1941e:385), Gans (1967:88). Zygaspis quadrifrons: Broadley and Broadley (1997:14), Gans etal. (1997:44), Gans (2005:39), Conradie et al. (2016:23). Global distribution: A widespread species distributed from Southern Democratic Republic of Congo, south through Zambia, Zimbabwe, and Botswana to the northern regions of South Africa, west to southwestern Angola and northern Namibia, east to southern Malawi and central Mozambique Ocurrences in Angola (Map 165): The species has been reported from southern Ango¬ la, but may be expected to occur in Moxico Province near the border with Zambia as well. Cunene: “Chimporo” [-16.03333, 17.15000] Map 165. Distribution of Zygaspis quadrifrons in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 213 (Monard 1931:93, I937b:65; Loveridge I94Ic:385, 387; Gans 1967:88, 2005:39; Broadley and Broadley 1997:17). Cuando Cubango: “Caquindo (Kakindo)” [-15.45000, 17.05000] (Monard 1931:93, I937b:65; Loveridge I94Ic:385, 387; Gans 1967:88, 2005:39; Broadley and Broadley 1997:17); “lower Cuando river (41b)” [-17.46777, 23.07944] (Conradie et al. 2016:9, 11, 23); “lower Cuando river (44b)” [-17.56916, 23.27305] (Conradie et al. 2016:9, 11, 23); “Cuito basin (59)” [-16.28392, 18.84744] (Conradie et al. 2016:9, 12, 23); “Cuito basin (60)” [-16.92367, 19.29675] (Conradie et al. 2016:9, 12, 23). Taxonomic and distributional notes: Bauer et al. (1995a) and Gans (2005) mistakenly gave the current name of Neu Barmen as Otjimbingue. This is, in fact, the location of another Rhenish mission station operated in the 19* century by the missionary and collector Carl Hugo Hahn. Amphisbaena ambuellensis (Monard, 1931) was synonymized with Z. quadrifrons with some mis¬ givings by Loveridge (1941c) because he expected to find “racial differentiation” between the fauna of southern Angola and that of South West Africa [Namibia] and Broadley and Broadley (1997) likewise considered that these populations might eventually prove to be distinct. Family Lacertidae Bonaparte, 1831 Genus Heliobolus Fitzinger, 1843 Heliobolus lugubris (Smith, 1838) Bushveld Lizard Lacerta lugubris Smith 1838:93. Syntypes: BMNH 1946.8.6.37-45 (collector A. Smith). Type locality: “Dis¬ trict immediately beyond the northern frontier of the [Cape] Colony,” South Africa. Eremias lugubris: Bocage (1867b:221, 1895a:31), Boulenger (1905:111; 1921:239), Monard (1937b:75), Mertens (1938a:437). Lampreremias lugubris: Szczerbak (1975:33). Heliobolus lugubris: Branch (1998:161), Bates et al. (2014:160), Ceriaco et al. (2016a:37, 56), Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: Widely distributed, from southern Angola over the Kalahari sands through Namibia, Botswana, southwestern Zimbabwe, southern Mozambique and South Africa. Ocurrences in Angola (Map 166): The species occurs broadly in southern Angola. Benguela: “Sighting in Lobito” [-12.35000, 13.55000] (Monard 1937b:75); “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:31); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:31); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:221, 1895a:31; Szczerbak 1975:33); “Dombe” [-12.95000, 13.10000] (Bocage 1895a:31); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:437); “Huxe, Benguella” [-12.71667, 13.20000] (Boulenger 1921:243). Namibe: “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:31; Ceriaco et al. 2016b:56); “Namibe-Lubango road, 2 km E of Mangueiras, south side of the road” [-15.04361, 13.16000] (Ceriaco et al. 20 1 6a:3 7); “Capangombe” [-15.1 0000, I2"E 20°E 24“E Map 166. Distribution of Heliobolus lugubris in Angola. 214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 13.15000] (Bocage 1895a:31; Ceriaco et al. 2016a:56); “Konondoto, Mossamedes” [-15.20000, 12.15000] (Boulenger 1921:243; Ceriaeo et al. 2016a:56); “Namibe-Lubango road, 2 km E of Mangueiras, south side of the road” [-15.03333, 13.16000] (Ceriaco et al. 2016a:37); “Mossamedes” [-15.78333, 12.06667] (Szczerbak 1975:33). Cunene: “PonangKuma” [-17.05000, 14.65000] (Boulenger 1921:243); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:75). Cuan- do Cubango: “Cuito basin (63)” [-17.93611, 21.10269] (Conradie et al. 2016:9, 12, 24). Unde¬ termined Locality: “Cuanza R.” (Boulenger 1905:111). Taxonomic and distributional notes: This widespread species has never been investigated in detail and it is likely that there is regional differentiation and possibly even cryptic taxa within H. lugubris as currently construed. Genus Holaspis Gray, 1863 Holaspis guentheri Gray, 1863 Blue-Tailed Tree Lizard Holaspis guentheri Gray 1863a:152, p. 20, fig. 1. Holotype: BMNH 1946.8.7.31 (donor Sir A. Smith, collec¬ tor unknown). Type locality: None stated, but Gray (1863) considered that it might be a “Tropical- Amer¬ ican” form. Holaspis guentheri: Ferreira (1897b:242), Spawls et al. (2004:163), Chirio and LeBreton (2007:218), Trape et al. (2012:322). Holaspis guentheri guentheri: Loveridge (1957:227), Laurent (1964a:56), Chirio and LeBreton (2007:218), Trape et al. (2012:322). Global conservation status (lUCN): Not Evaluated. Global distribution: This is a lowland forest species, found in open woodland adjacent forest and in some coastal forests, from Uganda and Tanzania in the east to Sierra Leone in the west. In the southwest it reaches Angola. Ocurrences in Angola (Map 167): The few published Angolan records are from the northeast of the country. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:56); “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:56). Taxonomic and distributional notes: Based on the species global distribution record from “Rio Cuce,” Huila Province seems doubt¬ ful and is probably based on a misidentifica- tion. Ferreira’s (1897b) remarks relative to Gray’s (1863) and Boulenger’s (1887) descrip¬ tions of the species suggest that the “Rio Cuce” specimen was probably the small, terrestrial gerrhosaurid, Cordylosaurus subtessellatus (Smith, 1844), which shares a similar color pat¬ tern and does occur in the area. Map 167. Distribution of Holaspis guentheri in Angola. Genus Ichnotropis Peters, 1854 Ichnotropis bivittata bivittata Bocage, 1866 Angolan Rough-Scaled Lizard lehnotropis Dumerilii [lehnotropis bivittatus] Bocage 1866a:43: Syntypes: ZMB 5827 fide Bauer et al. (1995b:41), MBL specimen numbers unknown, destroyed by fire 18 March 1978 (collector F.A.P. Bayao); Type loeality: “Duque de Bragan^a” (Boeage 1866a:43) [= Calandula], Malanje Provinee, Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 215 Ichnotropis capensis: Boulenger (1887:78, 1897:277, 1905:110), Bocage (1895a:30), Ferreira (1897b:243, 1903:15). Ichnotropis bivittata: Boulenger (1921:182), Sehmidt (1933:11), Loveridge (1933:308), Parker (1936:135), Monard (1937b:74), Marx (1956:7), Bauer et al. (1995b:41), Spawls et al. (2004:173), Broadley and Cot- terill (2004:44), Edward et al. (2013a: 110), Ineieh and Le Garff (2015:475), Ceriaeo et al. (2016b:61). Ichnotropis capensis bivittata: Hellmieh (1957b:59), Loveridge (1957:234), Managas (1963:237). Ichnotropis longipes: Frade (1963:253), Monard (1937b:75). Ichnotropis bivittata bivittata: Laurent (1964a:63). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from Angola east through southern Katanga region of the Demoeratic Republie of Congo, northern Zambia and northern Malawi to southern Tanza¬ nia. Ocurrences in Angola (Map 168): The speeies is very widespread in the eountry exclusive of the southwest coastal regions: Cabinda: “Chyiaca” [-4.86667, 12.56667] (Boulenger 1921:185). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:63); “Dala” [-11.03333, 20.20000] (Monard 1937b:75). Malanje: “BangeN’gola” [-8.43333, 16.56667] (Boulenger 1905:110, 1921:185); “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:43, 1895a:30; Bou¬ lenger 1887:78, 1905:110, 1921:185; Ferrei- ra:1903:15; Loveridge 1933:308, 1957:234; Bauer et al. 1995b:41); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:61). Mexico: “Lago Cameia” [-11.71667, 20.80000] (Managas 1963:237); “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] (Managas 1963:237); “Calombe, Luso” [-11.83333, 19.93333] (Managas 1963:237). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:11; Marx 1956:7); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:11; Marx 1956:7). Huam- bo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:74); “Galanga” [-12.06667, 15.15000] (Boca¬ ge 1895a:30); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:135; Marx 1956:7); “Bela-Vista” [-12.36667, 16.20000] (Hellmieh 1957b:59); “Santo-Amaro” [-12.70000, 15.85000] (Monard 1937b:74). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:30); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:30); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:74). Huila: “Rio Cuce” [-13.51667, 15.20000] (Ferreira 1897b:243); “Caconda” [-13.73333, 15.06667] (Bocage 1895a:30; Boulenger 1921:185); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:74); “Sangeve” [-13.88333, 15.83333] (Monard 1937b:74); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:74); “Lobango” [-14.91667, 13.50000 (Bocage 1895a:30); “Kasinga” [-15.13333, 16.08333] (Monard 1937b:74); “Kului” [-15.41667, 15.73333] (Monard 1937b:74). Cunene: “Riv. Mbale” [-15.16667, 16.75000] (Monard 1937b:75); “Chimporo” [-16.03333, 17.15000] (Monard 1937b:75); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:75). Undeter¬ mined locality: “Benguela-Bihe” (Boulenger 1921:185). Taxonomic and distributional notes: Bocage (1866a) coined the name 1. bivittatus, but based Map 168. Distribution of Ichnotropis bivittata bivittata in Angola. 216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II on information from Albert Gunther at the British Museum, he was initially convinced that his material was conspecific with I. dumerilii (Smith, 1849) [= 7. capensis (Smith, 1838)] and under that account heading he listed: "'Tropidosaura DumerUii. Smith. Ichnotropis bivittatus. Nob. Mss.” Peters (1882) argued for the validity of Bocage’s species and suggested that the specimen of 7. dumerUii that Gunther had compared it to might not have been Smith’s type specimen. Because of this confusion 7. bivittata was for some time mentioned in the literature as 7. capensis (e.g., Bocage 1895a; Boulenger 1887, 1897, 1905; Ferreira 1897b, 1903). No modem revision of Ichnotropis has yet been carried out (Edwards et al. 2013 a) and the status of a number of taxa remains equivocal. Ichnotropis bivittata pallida Laurent, 1964 Pale Rough-Scaled Lizard (Endemic) Ichnotropis bivittata pallida Laurent 1964a:64, fig. 19. Holotype: MD 1854 (collector A. Barros Machado). Type locality: “Boca da Humpata, Huila” [Huila Province], Angola. Ichnotropis bivittata pallida'. Laurent (1964a: 64), Edward et al. (20 13a: 110). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 169): The species 7. b. pallida has only been recorded from its type locality on the southwestern plateau. Huila: “Boca da Humpata” [-14.93333, 13.51667] (Laurent 1964a:64). Taxonomic and distributional notes: Laurent (1964a) suggested that some speci¬ mens refered to Ichnotropis bivittata by Monard (1937b) from the Cunene Basin might belong to 7. b. pallida. The status of 7. bivittata pallida remains equivocal, although Ineich and Le Garff (2015) recognized it as a valid sub¬ species. Ichnotropis capensis capensis (Smith, 1838) A[lgyra] capensis Smith 1838:94. Syntypcs: lost fide FitzSimons (1943). Type locality: “sandy deserts around Latakoo” [= Lithako, approx. 27°S, 24°E, i.e. btwn. Kumman and Taungs, N. Cape Province,] South Africa. lehnotropis eapensis: Boulenger (1897:277), Laurent (1964a:63), Branch (1998:162), Broadley and Cotterill (2004:22), Broadley (2004:326), Bates et al. (2014:161). lehnotropis eapensis eapensis'. Branch and McCarteney (1992:1). lehnotropis spp.: Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: The typical form of the species is known from the Democratic Republic of Congo and Angola, south into Namibia and eastwards to Mozambique and South Africa. Ocurrences in Angola (Map 170): Ichnotropis capensis capensis has been recorded from Cuando Cubango Province. Cuando Cubango: “approximately 50 km E of Cuito Cuanavale” [-15.23333, 19.61667] (Branch and McCarteney 1992:1); “Cuito basin (25)” [-15.139194, Map 169. Distribution of Ichnotropis bivittata pallida in Angola. The Cape Rough-Scaled Lizard MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 217 19.14350] (Conradie et al. 2016:8, 9, 24); “Cuito basin (30b)” [-I7.5II94, 20.04305] (Conradie et al. 2016:8-9, 24); “Cuito basin (59)” [-16.28392, 18.84744] (Conradie et al. 2016:9, 12, 24); “Cuito basin (55)” [-14.68478, 18.67369] (Conradie et al. 2016:9, 12, 24). Taxonomic and distributional notes: Many early records from Angola attributed to 1. capensis in fact refer to 1. bivittata Bocage, 1866 (see above). Boulenger (1897) reported the first northern record for the species from “Kuango,” presumably in Malanje Province, which in the light of current knowledge does not correspond to the distribution of the /. capensis (Branch 1998; Broadley and Cotter- ill 2004; Broadley 2004; Bates et al. 2014). Although this and most Angolan records were recognized by Monard (1937b) and Laurent (1964a) as referrable to 1. bivittata, confusion Map 170. Distribution of Ichnotropis capensis capensis in Angola. regarding the application of the name /. capensis in Angola remains today. The record of Branch and McCartney (1993) from Cuando Cubango Province is regarded as the first record of the nominotypical form in Angola (Branch 1998; Broadley 2004; Bates et al. 2014), but extensive fieldwork by Conradie in southeastern Angola has yielded additional records (Conradie et al. 2016). Ichnotropis capensis overlaeti de Witte and Laurent, 1942 Lunda Rough-Scaled Lizard (Endemic) Global conservation status (lUCN): Not Evaluated. Global distribution: Only reported from Angola, although expected to occur in the Katanga region of the Democratic Republic of Congo. Ocurrences in Angola (Map 171): Ich¬ notropis c. overlaeti has recorded from north¬ eastern regions of Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12, 1964a:61); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 12); “Caluango, R. Caquele, affluent Luangue” [-8.33333, 19.88333] (Laurent 1964a:61). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:61). Ichnotropis overlaeti de Witte and Laurent 1942a: 173. Holotype: MRAC 9691 (eolleetor F.G. Overlaet). Type loeality: “Kapanga (Lulua)” [= Lulua Distriet, Kasai-Oeeidental Provinee,] Demoeratie Republie of Congo. lehnotropis eapensis overlaeti'. Laurent (1950a: 12, 1964a:61). Map 171. Distribution of Ichnotropis capensis overlaeti in Angola. 218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: The status of Ichnotropis overlaeti has not been criti¬ cally assessed. Its substantially disjunct distribution relative to the nominotypical form suggests that it is likely valid at either a subspecific or specific level. Ichnotropis microlepidota Marx, 1956 Marx’s Rough-Scaled Lizard (Endemic) Ichnotropis microlepidota Marx 1956:5, fig. 1. Holotype: FMNH 74285 (collector G. Heinrich). Type locali¬ ty: “Mt. Moco” [= Serra do Moco], Huambo Province, Angola. Ichnotropis bivittata: Parker (1936:135). Ichnotropis microlepidota: Marx (1959:461), Edwards et al. (2013a: 110), Ineich and Le Graff (2015:477). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known Ocurrences in Angola (Map 172): The species is known only for the type locality “Mt. Moco” in Huambo Province. Huambo; “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:135; Marx 1956:5, 1959:461). Taxonomic and distributional notes: Parker (1936) identified a specimen from “Mt. Moco” as Ichnotropis bivittata (Bocage, 1966), however he noted that the specimen had some different characters, which might represent racial differentiation. Marx (1956) included this specimen, along with a series of lizards also from “Mt. Moco” as Ichnotropis microlepidota. According to Edwards et al. (2013a) and Ineich and Le Graff (2015) the sta¬ tus of this taxon remains equivocal, until fur¬ ther review of the genus with new material for molecular phylogenetic analyses. from Angola. Il-E 20°E I4”E Map 172. Distribution of Ichnotropis microlepidota in Angola. Genus Meroles Gray, 1838 Meroles anchietae (Bocage, 1867) Anchieta’s Dune Lizard Paehyrhynehus Anehietae Bocage 1867b:227, figs. 1-2. Holotype: MBL speeimen number unknown (eollee- tor J.A. d’Anehieta), destroyed by fire 18 Mareh 1978. Type loeality: “Mossamedes” subsequently ehanged to “Rio Coroea, dans le littoral de Mossamedes,” Namibe Provinee, Angola (Boeage 1895a:33). Paehyrhynehus Anehietae: Boeage (1895a:33, 1897a: 195). Aporosaura anehietae: Boulenger (1887:117, 1921:376), Loveridge (1936a:63), Frade (1963:253), Arnold (1989:213, 1991:784). Meroles anehietae: Braneh (1998:164), Ceriaeo et al. (2016a:36, 56). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the dunes of the Namib Desert in south¬ western Angola and western Namibia. Ocurrences in Angola (Map 173): The species occurs in the desert regions of Namibe Province. Namibe: “Rio Coroea, dans le littoral de Mossamedes” [-15.78333, 12.06667] (Bocage 1867b:227, 1895a;33, 1897a:195; Boulenger 1887:117, 1921:376; Loveridge 1936a:63; Ceriaeo et al. 2016a:56); “Porto Alexandre” [-15.80000, 11.83333] (Ceriaeo etal. 2016a:36); “Praia do Navio MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 219 coastal dunes, ca. 124 km SSW of Namibe” [-16.27475, 11.81806] (Ceriaco et al. 2016a:36); “Foz de Cunene” [-17.28333, 11.80000] (Ceriaco et al. 2016a;36). Taxonomic and distributional notes: Arnold (1989) demonstrated that Aporosaura Boulenger, 1887, the generic name under whieh this species was long known, is a syn¬ onym of Meroles Gray, 1838. Until recently the only published data for Angola was from the type locality, but recently an individual was colleeted in “Praia do Navio eoastal dunes, ea. 124 km SSW of Namibe” (Ceriaco et al. 2016a). In the Museu de Historia Natural da Universidade do Porto, Portugal, the Franeisco Newton eollections inelude uneatalogued spec¬ imens of this species also from “Mossamedes” (M. Marques and L. Ceriaeo, pers. obs.) and Map 173. Distribution of Meroles anchietae in Angola. the Ditsong National Natural History Museum has some material collected by Wulf Haacke from “Porto Alexandre” and “Foz de Cunene” (Ceriaeo et al. 2016a). This species is endemie to the Namib Desert, from the Klinghardt Mountains in southern Namibia to southern Angola (Braneh 1998). Meroles reticulatus (Bocage, 1867) Reticulate Sand Lizard Scapateira (?) reticulata Bocage 1867b:225. Syntypes: MBL specimen number unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Mossamedes, Western Africa” restricted to “Rio Coroca,” Namibe Province, Angola by Bocage (1897a: 195). Scapateira reticulata: Bocage (1870:68, 1895a:32, 1897a:195), Strauch (1867:404), Boulenger (1887:111, 1921:357), Loveridge (1936a:63). Scapteira serripes: Boulenger (1887:111). Meroles reticulatus: Szczerbak (1975:45), Arnold (1989:213), Branch (1998:167), Bauer and Giin- ther (1995:55), Ceriaco et al. (2016a:36, 56). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the gravel plains of the Namib Desert in southwestern Angola and northern Namibia. Ocurrences in Angola (Map 174): The species oeeurs in the southwestern Angola near the coast. Benguela; “Benguella” [-12.58333, 13.41667] (Boulenger 1887:111; Szczerbak 1975:45; Bauer and Gunther 1995:55). Nami¬ be: “Mossamedes” [-15.78333, 12.06667] (Bocage 1867a;225; Boulenger 1887:112; Loverdige 1963:63; Ceriaco et al. 20 16a: 56); Angola. 174. Distribution of Meroles reticulatus in 220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:32, 1897a:195; Boulenger 1887:111; Bauer and Gunther 1995:55; Ceriaco et al. 2016a:56); “Praia do Navio coastal dunes, ca. 124 km SSW of Namibe” [-16.27758, 11.82236] (Ceriaco et al. 2016a:36). Taxonomic and distributional notes: Bocage’s (1867a) doubts about the species’ inclusion in Scapteira were laid to rest by Strauch (1867). Boulenger (1887, 1921) noted that one of the types of S. reticulata from “Benguella” was present in Berlin. Bauer and Gunther (1995) identified this specimen as ZMB 6475, and although not found at the time of their writing, it has since been locat¬ ed. However, although this specimen was collected by d’Anchieta and received from Bocage, it cannot be one of the types, as all five syntypes were from Mossamedes. Similarly to M. anchietae, there are very few published records for the species in Angola. Meroles squamulosus (Peters, 1854) Common Rough-Scaled Lizard Ichnotropis squamulosa Peters 1854:617. Syntypes: ZMB 6614, 6615, 6616 fide Bauer et al. (1995:58). Type loeality: “Tette” [= Tete], Mozambique. Ichnotropis squamulosa'. Monard (1937h:74), Loveridge (1957:234), Braneh (1998:163), Edwards et al. (2012:4), Engleder et al. (2013:132). Meroles squamulosus'. Edwards et al. (2013a: 101), Bates et al. (2014:164), Conradie et al. (2016:25). Global conservation status (lUCN): Not Evaluated. Global distribution: Meroles squamulosus occurs from Angola and Tanzania southwards through Zimbabwe, central Mozambique, Botswana, eastern Namibia, and northeastern South Africa. Ocurrences in Angola (Map 175): The species occurs in southern Angola east of the arid southwestern zone. Huila: “Kapelongo” [-14.88333, 15.08333] (Monard 1937b:74). Cunene: “Pereira de Eqa [= Ondijiva])” [-17.06666667, 15.73333333] (Conradie et al. 2016:25). Cuando Cubango: “Cuito basin (30e)” [-17.51430, 20.05527] (Conradie et al. 2016:9, 10, 25); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9, 12, 25). Taxonomic and distributional notes: A recent analysis using nuclear and mitochondri¬ al sequence data revealed that this species, long known by the name Ichnotropis squamulosa, is strongly supported as a member of the genus Meroles (Edwards et al. 2012; Engleder et al. 2013; Edwards et al. 2013a). Confusion with 1 ^ . 1 7- 7 ^ • j Map 175. Distribution of Meroles squamulosus in members of the genus Ichnotropis is under- standable, as they are very similar morpholog¬ ically (Edwards et al. 2012; Edwards et al. 2013a). The geographic range and habitat of/, squa¬ mulosa overlaps with that of a number of Ichnotropis, but not that of other Meroles (Branch 1998). Il-E [6‘“£ 24''E MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 221 Genus Nucras Gray, 1838 Nucras scalaris Laurent, 1964 Scaled Sandveld Lizard (Endemic) Nucras scalaris Laurent 1964a:58, fig. 17. Holotype: MD 5401. Type loeality: “Alto Chieapa,” Lunda Sul, Angola. Nucras scalaris: Arnold (1989:21), Bisehoff (1991:20), Riehman and Bohm (2013). Global conservation status (lUCN): Data Deficient Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 176): The species is only known from two localities in northeastern Angola. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:58; Riehman and Bohm 2013); “Alto Chieapa” [-10.88333, 19.23333] (Laurent 1964a:58; Riehman and Bohm 2013). Taxonomic and distributional notes: This is a poorly known species and further research is needed in order to assess its taxon¬ omy, distribution and conservation status (Bichoff 1991; Riehman and Bohm 2013). We recently examined the type specimens (holo¬ type and three paratypes), which had not been seen by researchers since Laurent (1964a), owing to the inaccessibility of the Dundo collection. Il-E 16°E 20°E 24''E Map 176. Distribution of Nucras scalaris in Angola. Nucras aff. tessellata (Smith, 1838) Western Sandveld Lizard L[acerta]. tesselata (Smith 1838:92). Lectotype: BM 1946.8.6.3 (formerly BMNH 65.5.4.40, along withpara- lectotype, now BMNH 1946.8.6.2) (collector A. Smith) designated by Broadley (1972:27). Type locality: “Eastern parts of the Cape Colony,” South Africa; “Districts on the western coast of Southern Africa, particulary Little Namqualand”7/t/e Broadley (1972:27). Nucras tessellata var. taeniolata: Bocage (1895a:30). Nueras tessellata var. tceniolata: Boulenger (1910:474). Nueras intertexta var. holubi: Boulenger (1917:208, 1920:20). Nueras tessellata: Monard (1937b:73), Laurent (1964a:56). Nueras ornata: Broadley (1965b:23). Nueras tessellata: Broadley (1972:30). Nueras tessellata: Arnold (1989:213), Edwards et al. (2013b:4), Bates et al. (2014:171), Ceriaco et al. (2016b:56). Global conservation status (lUCN): Not applicable (taxonomic status unclear). Global distribution: The nominal species is distributed from the southwestern Western Cape Province in South Africa, through central Namibia and southwestern Botswana. The taxon occur¬ ring in Angola is certainly a different species. Ocurrences in Angola (Map 177): The species occcurs in the southwestern Angola, from the Namibe desert to the plateau at Caconda. Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:30; Broadley 1972:31); “Kapelongo” [-14.88333, 15.08333] (Monard 1937b:73; Broadley 1972:31). Namibe: “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:30; Broadley 1972:31; 222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ceriaco et al. 2016b:56); “ km 34 de lar oute de Mogamedes a Sa da Bandeira” [-15.50000, 12.16667] (Laurent 1964a:56; Broadley 1972:31; Ceriaeo et al. 20 16b: 5 6). Cunene: “Donguena, Mossamedes” [-17.01667, 14.71667] (Boulenger 1910:472, 1917:210, 1920:20; Broadley 1972:31). Taxonomic and distributional notes: Bocage (1895a) recorded Nucras from “Maconjo” and “Caconda” collected by Anchi¬ eta and identified them as belonging to N. tae- niolata (Smith, 1838). Boulenger (1910) also allocated his specimens from “Dongoena” (= Mossamedes fide Boulenger 1910) to this species, but subsequently (Boulenger 1917, 1920) transferred them to N. holubi (Stein- dachner, 1882). Broadley (1972) revised the Nucras tessellata complex and recognised three subspecies: Nucras tessellata tessellata, Nucras tessellata livida (now a full species) and an unnamed subspecies of Nucras tessellata from Angola. Broadley also noted that the Angolan specimens were similar to N. taeniolata, but believed that more material was needed before the status of Angolan populations could be settled. Edwards et al. (2013b) confirmed the sister species relationship of N. tessellata to N. taeniolata, however, a detailed molecular investigation is still needed to resolve the position and status of Angolan Nucras, which may represent one or more new species (W.R. Branch, pers. comm., January 2017). Genus Pedioplanis Fitzinger, 1843 Pedioplanis benguellensis (Bocage, 1867) Bocage ’s Sand Lizard Eremias benguellensis Bocage 1867h:221, 1867d:229. Holotype: MBL specimen number unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Benguella” [= Benguela], Benguela Province, Angola. Eremias benguelensis: Boulenger (1921:287), Parker (1936:134). Eremias benguellensis: Monard (1937b:73), Laurent (1964a:60). Eremias sp.: Bocage (1887b:203). Eremias namaquensis: Boulenger (1887:91), Bocage (1895a:31). Eremias undata: Boulenger (1921:283). Eremias undata undata: Laurent (1964a:60). Mesalina benguelensis: Szczerbak (1975:24). Pedioplanis undata: Arnold (1989:213), Branch (1998:173), Makokha et al. (2007:623), Haacke (2008:90). Pedioplanis benguellensis: Arnold (1989:213), Bauer and Gunther (1995:55), Branch (1998:173), Makokha et al. (2007:623), Conradie et al. (2012b:93), Ceriaco et al. (2016a:37, 56). Pedioplanis namaquensis: Branch (1998:172), Makokha et al. (2007:623). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and Nambia. Ocurrences in Angola (Map 178): The species occurs in the southwestern Angola. Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:31; Monard 1937b:73); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:221, 1867d:229, 1895a:31; Monard 1937b:73; Szczerbak I2"E 1&“E 20“E Map 177. Distribution of Nucras aff. tessellata in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 223 Map 178. Distribution of Pedioplanis benguellensis in Angola. 1975:24; Bauer and Gunther 1995:55); “Huxe, Benguella” [-12.71667, 13.20000] (Boulenger 1921:286, 288-289); “Catengue” [-13.03333, 13.73333] (Parker 1936:134). Hulla: “Cacon- da” [-13.73333, 15.06667] (Bocage I895a:3I; Monard I937b:73). Namibe: “Namibe-Luban- go road, road marker 59, 1.8 km W (by road) from Caraculo, north side of the road” [-15.01606, 13.64483] (Ceriaeo et al. 20I6a:37); “Maeonjo (=Fazendo Mueungo)” [-14.782192, 12.486557] (Boulenger 1921: 286; Bauer and Gunther 1995:55; Conradie et al. 2012b:93; Ceriaeo et al. 2016a:56); “Capan- gombe” [-15.10000, 13.15000] (Boeage 1895a:31; Monard 1937b:73; Ceriaeo et al. 2016a:56); “Mossamedes” [-15.20000, 12.15000] (Boeage 1887b:203, 1895a:31; Monard 1937b:73; Szezerbak 1975:24; Ceria- co et al. 2016a:56). Cunene: “Ponang Kuma” [-17.05000, 14.65000] (Boulenger 1921:289). Taxonomic and distributional notes: Boeage provided descriptions of this species in two dif¬ ferent papers, which appeared back-to-back in the same issue of the “Jomal de Sciencias Mathe- maticas, Physicas e Naturaes.” In Bocage (1867d) the description is short and benguellensis is spelled with two “l”s, whereas in Bocage (1867b) it is more extensive and the specific epithet is spelled with only one “1.” Although most authors have cited the more extensive description in Bocage (1867b), all except Boulenger (1921), Parker (1936) and Szezerbak (1975), including Bocage himself in later works, have adopted the spelling of the shorter description (Bocage 1867d). We here follow page priority and employ the prevailing spelling, benguellensis. Eremias benguel¬ lensis was later considered by Bocage (1895a) as a synonym of E. namaquensis Dumeril and Bibron, 1839. Boulenger (1921) reinstated Eremias. benguelensis as a full species, diagnosing it from E. namaquensis. This interpretation was subsequently followed by several authors, including Parker (1936), Monard (1937b) and Laurent (1964a), although 7^ benguellensis has frequently been synonymized with P. namaquensis. Based on phylogenetic analysis carried out by Conradie et al. (2012b), material referable to P. benguellensis does not fall within the same lineage as (nor is it sis¬ ter to) P. namaquensis, despite past speculation to the contrary (Bocage 1895a; Branch 1998; Makokha et al. 2007). Previous records of P. namaquensis from southern Angola should be con¬ sidered misidentifications, since there is no support for the occurrence of P. namaquensis in Ango¬ la (Conradie et al. 2012b). Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012 Haacke’s Sand Lizard (Endemic) Pedioplanis haackei Conradie et al. 2012b: 101, figs. 3A, B. Holotype: PEM R18465 (eolleetors W.R. Braneh, W. Conradie, G.J. Measey and K.A. Tolley) Type loeality: “along the road to Tambor, Namibe Provinee,” Namibe Province, Angola. Eremias undata undata (part): Laurent (1964a:60). Pedioplanis haackei'. Ceriaeo et al. (2016b:37, 56). Global conservation status (lUCN): Not Evaluated. 224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 179): Endemic to southwestern Namibe Province, mainly on sandy planis surrounding granite outcrops. Namibe: “5 km NW (by road) of Pico Azevedo” [-15.47600, 12.46150] (Ceriaco et al. 2016a:37); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaco et al. 2016a:37); “Red Canyon at Lake Arco” [-15.74597, 12.13989] (Conradie et al. 20 12b: 101; Ceriaco et al. 2016a:56); “10 km south of Lake Arco” [-15.83044, 12.14125] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56); “Road to Tambor” [-15.87606, 12.20583] (Conradie et al. 2012b: 101; Ceriaco et al. 2016a:56); “Road to Tambor at giant Wei- witchia” [-15.88778, 12.36417] (Conradie et Map 179. Distribution of in Angola. al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56); “Road from Lake Arco to Espinheira” [-15.91356, 12.39522] (Conradie et al. 2012b: Online Supplementary Material; Ceria¬ co et al. 2016a:56); “Iona National Park, north of Tambor” [-15.99539, 12.40647] (Ceriaco et al. 2016a:37); “Omauha Lodge” [-15.99681, 12.40683] (Conradie et al. 2012a: Online Supplementary Material; Ceriaco et al. 2016b:37); “Road to Tambor” [-16.05847, 12.42597] (Conradie et al. 2012a: Online Supplementary Material; Ceriaco et al. 2016a:56); “20 km north of Omauha Lodge” [-16.07414, 12.43328] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56); “Pediva hot-springs, south side of the river” [-16.29381, 13.56033] (Ceriaco et al. 2016b:37). Taxonomic and distributional notes: This recently described species was likely previously mistaken for one of its superficially similar congeners, Pedioplanis undata (Smith, 1838). The lat¬ ter was first signaled in southern Angola by Boulenger (1921), who referred specimens from “Maconjo [=Fazendo Mucungo]” and “Benguella” to it. Laurent (1964a) discussed additional Angolan material and the presence of the species in the country was long widely accepted by the scientific community (Branch 1998; Makokha et al. 2007; Haacke 2008). However, since Laurent (1964a), the status of P. undata has changed considerably. According to Conradie et al. (2012b) P. undata is not present in Angola and, as currently understood, the species is endemic to Namib¬ ia. Laurent’s records were considered by Conradie et al. (2012b) to correspond Pedioplanis haack- ei Conradie, Measey, Branch and Tolley, 2012 and Pedioplanis huntelyi Conradie, Measey, Branch and Tolley, 2012. Pedioplanis “undata'' from outside the accepted range of these two species are likely to represent one or more undescribed taxa (W.R. Branch, pers. comm., January 2017). Pedioplanis huntleyi Conradie, Measey, Branch and Tolley, 2012 Huntley’s Sand Lizard (Endemic) Pedioplanis huntleyi Conradie et al. 2012:105, figs. 4A, B. Type: Holotype, PEM R18479 (eolleetors W.R. Braneh, W. Conradie, G.J. Measey and K.A. Tolley). Type loeality: “road to Oneoeua, 7 km from Iona, Namibe Provinee” Namibe Provinee, Angola. Eremias undata undata (part): Laurent (1964a:60). Pedioplanis huntleyi'. Ceriaeo et al. (2016a:56). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 225 Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is endemie to southwestern Angola. Ocurrences in Angola (Map 180): The species is known from southwestern Angola in Namibe and Cunene provinces near the Namib¬ ian border. Namibe: “40 km south of Omauha lodge” [-16.51164, 12.44761] (Conradie et al. 2012b: Online Supplementary Material; Ceria- co et al. 2016a:56); “14 km west of Moimba” [-16.67947, 12.61275] (Conradie et al. 2012b: 105, Online Supplementary Material; Ceriaco et al. 2016a:56); “23 km West of Moimba” [-16.67947, 12.97397] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 20 16a: 56); “26 km East of Iona” [-16.82928, 12.62103] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56); “16 km East of Iona” [-16.79797, 12.68050] (Conradie et al. 2012b: Online Sup¬ plementary Material; Ceriaco et al. 20 16a: 56); Map 180. Distribution of Pedioplanis huntleyi in Angola. “8 km North East of Iona” [-16.82928, 12.62103] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56); “Road to Onocua 7 km NE from Iona” [-16.85831, 12.61275] (Conradie et al. 2012b:105; Ceriaco et al. 2016a:56); “26 km SE of Oncocua” [-16.86881, 13.52756] (Conradie et al. 2012b: Online Supplementary Material; Ceriaco et al. 2016a:56). Taxonomic and distributional notes: This recently described species was likely previously mistaken for one of its superficially similar congeners, Pedioplanis undata (Smith, 1838). Laurent (1964a) cited Angolan material of the latter species and its presence in the country was long wide¬ ly accepted (e.g.. Branch 1998; Makokha et al. 2007; Haacke 2008). However, based on the phylogenetic analysis by Conradie et al. (2012b) P. undata is not present in Angola and, as currently understood, the species is endemic to Namibia. Laurent’s records were considered by Conradie et al. (2012b) to correspond Pedioplanis haackei Conradie, Measey, Branch and Tolley, 2012 and Pedioplanis huntelyi Conradie, Measey, Branch and Tolley, 2012. Pedioplanis ''undata"' from outside the accepted range of these two species are likely to represent one or more unde¬ scribed taxa (W.R. Branch, pers. comm., January 2017). Family Cordylidae Mertens, 1937 Genus Chamaesaura Schneider, 1801 Chamaesaura anguina oligopholis Laurent, 1964 Angolan Snake Lizard Chamaesaura anguina oligopholis Laurent 1964a: 50. Holotype: MD 6003 (collector A. Serralheiro). Type locality: “Calonda, Lunda,” Lunda Norte Province, Angola. Chamaesaura anguina oligopholis'. Branch (1998:185), Stanley et al. (2011:65), Bates et al. (2014:184). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and the adjacent southern Democratic Republic of Congo. Ocurrences in Angola (Map 181): The species is known only from a single locality in Lunda 226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Norte Province. Lunda Norte: “Calonda” [-8.41667, 20.53333] (Laurent 1964a;50). Taxonomic and distributional notes: Stanley et al. (2011) recognized two subspecies of Chamaesaura anguina (Linnaeus, 1758), the isolated Chamaesaura anguina oligopholis Laurent, 1964 from Angola and eastern Demo¬ cratic Republic of Congo, and C. a. anguina (Linnaeus, 1758) from South Africa and Swazi¬ land (Branch 1998; Bates et al. 2014). The rela¬ tionship between the two subspecies remains problematic (Stanley et al. 2011) and a molec¬ ular assessment is required to determine the extent of divergence (Bates et al. 2014), although it is probable that these highly dis¬ junct forms are specifically distinct. ll^E 20“E 24“E Map 181. Distribution of Chamaesaura anguina oligopholis in Angola. Chamaesaura miopropus Boulenger, 1894 Zambian Snake Lizard Chamaesaura miopropus Boulenger 1895b:732. Holotype: BMNH 1946.8.29.49 (eolleetor A. Carson). Type loeality: “Fwambo, British Central Afriea” [= Zambia]. Chamaesaura maerolepis: Boeage (1895a:25), Monard (1937b:61), Hellmieh (1957b:52). Chamaesaura miopropus: Schmidt (1933:10), Loveridge (1944b:107, 1957:226), Stanley et al. (2011:65), Bates etal. (2014:185). Chamaesaura maerolepis miopropus: Branch (1998:185), Spawls et al. (2004:189), Broadley and Cotterill (2004:43). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is ^o^e i4"e known from Angola east to southern Tanzania. Ocurrences in Angola (Map 182): The species occus in central- west Angola. Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:10; Loveridge 1944b: 108). Huambo: “Galanga” [-12.06667, 15.15000] (Boeage 1895a:25; Monard 1937b:61; Loveridge 1944b: 108); “Bela- Vista” [-12.36667, 16.20000] (Hellmieh 1957b:52). HuiTa: “Caconda” [-13.73333, 15.06667] (Boeage 1895a:25; Monard 1937b:61; Loveridge 1944b: 108). Taxonomic and distributional notes: Schmidt (1933) suggested that the two Angolan specimens from “Caconda” and “Galanga” identified as Chamaesaura maerolepis (Cope, 1862) by Boeage (1895a) are more likely refer¬ able to C. miopropus Boulenger, 1895. For Map 182. Distribution of Chamaesaura miopropus in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 227 some time and until recently, C. miopropus was treated as a northern subspecies of C. macrolepis (Cope, 1862) (Broadley 1971d; Branch 1998; Spawls et al. 2004). Although this species is distin¬ guished by C. macrolepis by the presence of forelimbs (Bocage 1895a, Schmidt 1933; Loverdige 1944b; Broadley 1971a; Broadley and Cotterill 2004) which are absent in the latter taxon. Chamae- saura miopropis is geographically isolated, occurring from Angola east to northern Malawi and southern Tanzania, whereas C. macrolepis is endemic to South Africa, Zimbabwe and Swaziland (Broadley and Cotterill 2004; Bates et al. 2014). Chamaesaura miopropus should therefore be con¬ sidered a valid species (Broadley and Cotterill 2004; Stanley et al. 2011; Bates et al. 2014). Genus Cordylus Laurenti, 1768 Cordylus angolensis (Bocage, 1895) Angolan Girdled Lizard (Endemic) Zonurus angolensis Bocage 1895a:25. Holotype: MBL specimen number unknown (collector J.A. d’ Anchi¬ eta), destroyed by fire 18 March 1978. Type locality: “Caconda,” Huila Province, Angola. Zonurus cordylus: Boulenger (1897:277). Zonurus angolensis: Monard (1937b:61), Frade (1963:252). Cordylus cordylus angolensis: Loveridge (1944b: 3 8). Cordylus tropidosternum tropidosternum: Broadley (1971:22) Cordylus angolensis: Broadley and Branch (2002:10), Branch et al. (2005:131), Stanley et al. (2011:67), Greenbaum et al. (2012:36), Nielsen and Colston (2014:167), Stanley et al. (2016:23). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 183): The species is known from the type locality “Caconda” in Huila Province but its distribu¬ tion range reaches the southern regions of Kwanza Sul Province. Kwanza Sul: “Mombo- lo” [-11.91667, 14.85000] (Greenbaum et al. 2012:36). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:24; Monard 1937b:61; Loveridge 1944b:39). Taxonomic and distributional notes: In the description of Cordylus angolensis Bocage (1895a) expressed some doubts about the true identity of C. angolensis, leading him to name it in a “provisional manner.” The species was not cited in any further Bocage works after the description. With the destruction of the type specimen in the Lisbon museum fire, the taxo¬ nomic identity of the species became even more problematic. Broadley (1971b) consid¬ ered it a synonym of Cordylus tropidosternum (Cope, 1869). Loveridge (1944b) and Branch et al. (2005) noted that Bocage’s type description of C. angolensis lacked detail, although it is now con¬ sidered a valid species endemic to central Angola (Broadley and Branch 2002; Stanley et al. 2011; Nielsen and Colston 2014; Stanley et al. 2016). Greenbaum et al. (2012) found three Cordylus specimens in the collections of the American Museum of Natural History assignable to C. angolensis from “Mombolo”, approximately 190 km north-northwest of the type locality. These specimens agree with the original holotype description in some measurments and coloration. Map Angola. 183. Distribution of Cordylus angolensis 228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II although they differ in the scale counts of longitudinal rows of ventral scales (Greenbaum et al. 2012). Angolan Cordylus are currently being revised and the systematic position of C angolensis will hopefully be resolved. Cordylus angolensis was described within the account of Cordylus cordylus (Bocage 1895a). The two specimens treated as C. cordylus were from the Sociedade de Geografia de Lisboa, Portugal, and had only the locality “Angola.” Monard (1937b) considered these specimens to be attributable to Cordylus vittifer (Reichenow, 1887), although Loveridge (1944b) considered this opinion untenable and considered them assignable to C. angolensis. The identity of these specimens, which have recently been re-discovered remains uncertain, but is under study. Cordylus machadoi Laurent, 1964 Machado’s Girdled Lizard Cordylus vittifer machadoi Laurent 1964a:49, fig. 12. Holotype: MD 1840-1 (collector A. Barros Machado). Type locality: “Leba, Humpata, environs de Sa da Bandeira, Alt. 1800 m, Huila” [= Serra da Leba, Luban- go], Huila Province, Angola. Cordylus vittifer machadoi'. Broadley (1971b:22) Cordylus machadoi'. Branch (1998:195), Stanley et al. (2011:67), Greenbaum et al. (2012:23), Bates et al. (2014:192), Nielsen and Colston (2014:171), Stanley et al. (2016:202). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola and northwestern Namibia. Ocurrences in Angola (Map 184): The species is known from the type locality in Huila Province, however the species distribu¬ tion may extend to Namibe Province. Huila: “Leba, Humpata, environs de Sa da Bandeira” [-15.06667, 13.60000] (Laurent 1964a:49); “Humpata” [-14.96167, 13.34972]; [-15.17750, 13.32139E (Stanley et al. 2011:57; Stanley et al. 2016:209); “Nascente do Tchi- viuguira” [-15.16667, 13.30000] (Stanley et al. 2016:209). Taxonomic and distributional notes: The holotype of this taxon has recently been rediscovered in the collection at Dundo. The single juvenile paratype, was exchanged from this collection and is now MCZ R-74120 (fide Stanley et al. 2016). Branch (1998) elevated C. machadoi to full species, an action recently Map 184. Distribution of Cordylus machadoi in Angola. supported by recently Stanley et al. (2016) who based their conclusions on two C. machadoi spec¬ imens collected at the type locality. These were recovered as sister to all East African species of Cordylus (Stanley et al. 2011; Greenbaum et al. 2012; Nielsen and Colston 2014). A cryptic species allied to C. machadoi, was recently discovered at lower elevation (Stanley et al. 2016). Significant genetic variation across such a constrained geographic area suggests that its northern congener, C. angolensis (Bocage, 1895), which has a comparatively broad geographic distribution, may itself represent a species complex (Stanley et al. 2016). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 229 Cordylus namakuiyus Stanley, Ceriaco, Bandeira, Valerio, Bates and Branch, 2016 Kaokoveld Girdled Lizard (Endemic) Cordylus namakuiyus (Stanley et al. 2016:212, figs. 1, 4, 5). Holotype: CAS 254912 (collectors L. Stanley, S. de Sa, S. Bandeira, H. Valerio, A. Kuhn, J. Vindum and L. Ceriaco). Type locality: “large rock outcrop near Caraculo, road from Lubango and Namibe, Namibe Province,” Angola. Cordylus namakuiyus: Ceriaco et al. (2016a:38, 59). Stanley et al. (2016) recently described this form as a new species endemic to the arid low¬ lands west of the southern Angolan escarpment and sister to the geographically proximate, high-elevation species Cordylus machadoi Laurent, 1964. All phylogenetic analyses recovered the Angolan specimens as monophyletic, form¬ ing the earliest diverging lineage of the northern Cordylus clade (Stanley et al. 2016). These authors analyzed the morphology of AMNH Angolan specimens identified as Cordylus cordylus (Linnaeus, 1758) collected during the Vemay expedition in 1925 and showed that they cluster closely with C. namakuiyus. Although the Vemay specimens lack specific locality information, the expedition field notes mention that significant numbers of unidentified lizards were collected at “Pico Azeve- do” and “100 km east of Mogamedes,” the same areas where the eight specimens of the C. namakuiyus were collected, leading the authors to assign the Vemay material to the new species (Stanley et al. 2016). Map 185. Distribution of Cordylus namakuiyus in Angola. Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 185): The species is known from Huila and Namibe provinces in southwestern Angola. Huila: “large rock outcrop near Caraculo, on the road from Lubango and Namibe” [-15.01650, 12.64203] (Stanley et al. 2016:212; Ceriaco et al. 2016a:38, 59). Namibe: “outskirts of Pico Azevedo” [-15.47589, 12.46269] (Stanley et al. 2016:212; Ceriaco et al. 2016a:38, 59); “low rock outcrop bordering the road between Namibe and Omahua lodge” [-15.99681, 12.40683] (Stanley et al. 2016:212; Ceriaco et al. 2016a:38, 59). Taxonomic and distributional notes: 20“E 24''E Family Gerrhosauridae Fitzinger, 1843 Genus Cordylosaurus Gray, 1865 “1866” Cordylosaurus subtessellatus (Smith 1844) Dwarf Plated Lizard Gerrhosaurus subtessellatus Smith 1844: pi. 41, fig. 2 and first of two accompanying unnumbered text pages, pi. 42, figs. 17-20. Holotype: BMNH 1946.8.19.41 (eolleetor A. Smith). Type loeality: “Great Namaqua- land” [= southern Namibia]. Cordylosaurus trivittatus: Boulenger (1887:126), Boeage (1867b:222, 1895a:37). Cordylosaurus subtessellatus: Ceriaeo et al. (2016a:39, 59). 230 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southwestern Angola through western Namibia and into the western parts of South Africa. Ocurrences in Angola (Map 186): The species occurs in the southwestern regions near the coast, but probably also inland (see Notes below). Benguela; “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:37); “Benguela” [-12.58333, 13.41667] (Boulenger 1887:126; Bocage 1895a:37); “Dombe” [-12.95000, 13.10000] (Bocage 1867b:222, 1895a;37). Huila: “Rio Cuce” [-13.51667, 15.20000] (Fer¬ reira 1897b:242). Namibe; “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:37; Ceriaco et al. 2016a:59); “Iona National Park, MA,\%6.Uisx,ihutionoiCordylosaumssubt^^^^^^ 20 km SSW of Espinheira” [-16.93169, 12.24500] (Bocage 1895a:37; Ceriaco et al. 2016a;39). Taxonomic and distributional notes: FitzSimons (1937) recognized a single BMNH speci¬ men (number not given) corresponding to Smith’s description, but the description, although giving details for only a single animal, does note that two specimens were collected by Smith. Ferreira’s (1897b) record of Holaspis guentheri Gray, 1 863 from “Rio Cuce,” Huila may actually refer to this unrelated, although superficially similar species. ll^E 20“E 24"E Genus Gerrhosaurus Wiegmann, 1828 Gerrhosaurus auritus Boettger, 1887 Kalahari Plated Lizard Gerrhosaurus auritus Boettger 1887:148, pi. V, figs. 3a-e. Holotype: SMF 13947 (formerly SMF 6120a) (eol- leetor H. Sehinz) fide Boettger (1893) and Mertens (1922, 1967). Type loeality: “Ondonga in Ovam- boland” Namibia. Gerrhosaurus auritus: Monard (1937b:78). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola, Botswana, the Democratic Republic of Congo and Namibia. Ocurrences in Angola (Map 187): The species occurs in northeastern Angola. Lunda Sul: “Lunda” [-10.96667, 20.06667] (Monard 1937b:78) Taxonomic and distributional notes: Monard considered one specimen from “Lunda” to be possibly referable to Gerrhosaurus multilineatus Bocage, 1866, although Loveridge (1942) reject¬ ed this interpretation and treated this taxon as a subspecies of Gerrhosaurus nigrolineatus Hal- lowell, 1857. Subsequent authors, including FitzSimons (1943), Mertens (1971) and de Witte (1953) have considered G. auritus a full species. However, de Witte’s specimens were, in fact, referable to G. bulsi Laurent, 1954 (Bates et al. 2013), a species described by Laurent (1954) first as a subspecies of auritus. Broadley (1971) treated auritus as a subspecies of G. multilineatus and this has subsequently been followed by Bates et al. (2014). There has been much confusion regarding the applicability of names in Gerrhosaurus due to the lack of fresh material of Angolan MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 231 G. multilineatus, the type series of which was destroyed by fire in the Museu Bocage (Broadley 2007). Griffin (2003) reassessed and recognized the nomen G. auritus as a valid species (Bates et al. 2014) as did Broadley and Cotterill (2004) and Broadley (2007). Accord¬ ing to Bates et al. (2013), G. auritus appears to be closely related to G. nigrolineatus, but they are morphologically distinguishable (Broadley 2007). Il-E 1&°E 20°E irE Map 187. Distribution of Gerrhosaurus auritus in Angola. Gerrhosaurus bulsi Laurent, 1954 Laurent’s Plated Lizard Gerrhosaurus auritus bulsi Laurent 1954a:64. Holotype: MD Ang. 2127 (collector A. Barros Machado). Type locality: “Dundo,” Lunda Norte Province, Angola. Gerrhosaurus nigrolineatus nigrolineatus'. Laurent (1950a: 12). Gerrhosaurus bulsi'. Laurent (1964a:50), Branch and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola, the southern Democratic Republic of Congo, and Zambia. Ocurrences in Angola (Map 188): The species occurs in northeastern and central regions of Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12, 1954a;64, 1964a:50); “Luachimo village” [-7.38333, 20.83333] (Bates et al. 2013:469 [Table 1]); “Lake Carumbo base camp” [-7.74422, 19.95522] (Bates et al. 2013:469 [Table 1]); “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200). Lunda Sul: “Poste de Cacolo, Alto Cuilo” [-10.15000, 19.28333] (Laurent 1964a:51). Moxico: “envi¬ rons du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a: 50). Taxonomic and distributional notes: de Witte’s (1943) specimens of G. auritus Boettger, 1887, are, in fact, referable to this . . 1 .t- Map 188. Distribution of Gerr^os'awrws' bu A/ in Angola. species (Bates et al. 2013). For some time G. bulsi was treated as a synonym of the poorly known Gerrhosaurus multilineatus Bocage, 1866 (Broadley 1971), although it is currently regarded as valid (Broadley and Cotterill 2004; Broadley 232 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 2007). According to Bates et al. (2013) G. bulsi is a distinct species that is sister to the clade {Ger- rhosaurus intermedius [G. auritus, G. nigrolineatus]). Gerrhosaurus multilineatus Bocage, 1866 Kwanza Keeled Plated Lizard (Endemic) Gerrhosaurus multilineatus Bocage 1866a:44. Holotype: MBL specimen number unknown (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” [= Calandula], Malanje Province, Angola. Gerrhosaurus multilineatus-. Bocage (1866b:61, 1867b:221), Peters (1881:147). Gerrhosaurus bulsi'. Ceriaco et al. (2016b:63). Bocage (1866b) remarked that Gerrhosaurus multilineatus might be a well-characterized variety of G. nigrolineatus Hallowell, 1857, however he considered the two species distinct based on color pattern. Loveridge (1942) and FitzSimons (1943) subsequently relegated G. multilineatus to the synonymy of G. nigro¬ lineatus, the former considering it a color variant (Loveridge 1942). Haagner et al. (2000) stated that Broadley (unpublished) considered G. multilineatus to be based on a hybrid specimen and that the name would, therefore, be unavailable. However, according to Article 17.2 of the Code (ICZN 1999), even if the specimen was a hybrid, the name would still be available (Bates et al. 2013). Currently this species is recognized as valid despite the obvious taxonomic confusion, the loss of the original type series to fire at the Museu Bocage and the lack of museum material, especially from Angola (Bates et al. 2013). The status of Gerrhosaurus multilineatus and the assignment of Angolan populations referred to Gerrhosaurus nigrolineatus (e.g., Boulenger 1887, 1905; Ferreira 1900a, 1903, 1904; Angel 1923; Monard 1937b; Mertens 1938a; Hellmich 1957a, 1957b; Manaqas 1963; Parker 1936; Schmidt 1933; Laurent 1954a) remains problematic and resolution must await the collection of material from the type locality for molecular analysis, and a detailed morpholog¬ ical evaluation of the complex (Bates et al. 2013). Ceriaco et al. (2016b) referred to the specimen collected in Cangandala National Park (Malanje Province) as G. bulsi, but ongoing molecular works reveal that it shall be considered true multilineatus. Map 189. Distribution of Gerrhosaurus multilineatus in Angola. Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 189): Pub¬ lished records exist for this species from cen¬ tral and western Angola. Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a;44, 1866b:61); “Malanje” [-9.55000, 16.35000] (Peters 1881:147); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 20 16b: 63). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1867b;221); “Dombe” [-12.95000, 13.10000] (Bocage 1867b:221). Taxonomic and distributional notes: 20“E 24''E MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 233 Gerrhosaurus cf. nigrolineatus Hallowell, 1857 Black-Lined Plated Lizard Gerrhossaurus nigro-lineatus (Hallowell 1857:49). Syntypes: ANSP 3729, 8825 (collector H.A. Ford). Type locality: “Gaboon” [= Gabon]. Gerrhosaurus nigrolineatus: Bocage (1866a:43; 1870:68, 1887c:210, 1895a:35, 1896a:lll), Peters (1877a:613), Boulenger (1887:122, 1905:111), Ferreira (1900a:49, 1903:15, 1904:117), Angel (1923:159), Monard (1937b:78), Laurent (1954a:64), Grillitsch et al. (1996:26), Largen and Parker (2004:16), Ceria- co et al. (2014b:671), Ceriaco et al. (2016a:59). Gerrhosaurus flavigularis nigrolineatus: Schmidt (1919:523,1933:11), Mertens (1926:152, 1938a:435), Loveridge (1936a:65), Parker (1936:133). Gerrhosaurus nigrolineatus ahlefeldti Hellmich and Schmelcher 1956:203, fig. 1. Holotype: ZMH R02563 (formerly ZMH 806) (collector G.A. von Maydell) fide Hallermann (1998). Type locality: “Piri- Dembos, Roca Nova Douro” [= Piri-Dembos, Roga Novo Douro, Fazenda Novo Minho], Kwanza Norte Province, Angola. Gerrhosaurus nigrolineatus nigrolineatus: Hellmich (1957a:54, 1957b:58), Managas (1963:236). Gerrhosaurus nigrolineatus ahlefeldti: Hallermann (1998:208), Frazen and Glaw (2007:222), Bdhme (2014:152). Gerrhosaurus cf nigrolineatus: Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from a large distribution range, from Gabon and the lower Congo eastwards through southern Democratic Republic of the Congo eastern to Kenya, then southwards as far as northern Namibia, northern Botswana and north-eastern South Africa. Ocurrences in Angola (Map 190): The species is known from northeastern Angola. Cabinda; “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:613); “Molembo” [-5.33333, 12.20000] (Bocage 1895a;35); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:43, 1895a;35). Lunda Norte: “Dundo pres du Musee” [-7.36667, 20.83333] (Laurent 1954a: 64). Zaire: “St. Antonio [do Congo]” [-6.13333, 12.36667] (Schmidt 1919:523-524). Bengo: “Ambriz” [-7.844312, 13.106493] (Boulenger 1887:122; Bocage 1895a:35; Gril¬ litsch et al. 1996:26); “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:117). Luanda: “Luanda” [-8.83333, 13.26667] (Largen and Parker 2004:16). Kwanza Norte: “Piri-Dem- bos” [-8.56667, 14.50000] (Hellmich and L— ^ - 1 - Schmelcher 1956:203; Hellmich 1957b:58; Map 1 90. Distribution of Gerr/?05a™ cf. mgro/meato Hallermann 1998:208; Frazen and Glaw 2007:222; Bohme 2014:152); “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:54); “Dondo” [-9.68333, 14.43333] (Bocage 1895a:35). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] (Hellmich 1957b:58), “Quirimbo” [-10.68333, 14.26667] (Parker 1936:133); “Congulu” [-10.86667, 14.28333] (Parker 1936:133). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1895a:35; Ferreira 1903:15; Boulenger 1905:111); “Malanje” [-9.55000, 16.35000] (Bocage 1895a:35); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:671); “Pungo-Andon- go” [-9.66667, 15.58333] (Boulenger 1905:111). Mexico: “Lago Dilolo” [-11.50000, 22.01667] (Managas 1963:236); “Lago Cameia” [-11.71667, 20.80000] (Managas 1963:236). Bie: “Gauca” 234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-11.18333, 17.45000] (Schmidt 1933:11); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:11). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:78); “Galanga” [-12.06667, 15.15000] (Bocage 1895a:35; “Santo Amaro” [-12.70000, 15.85000] (Monard 1937b:78); “Cuma” [-12.86667, 15.06667] (Loverdige 1936a:65). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1887c:210); “Catumbella” [-12.43333, 13.55000] (Bocage 1887c:210); “Quissange” [-12.43333, 14.05000] (Bocage 1887c:210, 1895a:35); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:35); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:54); “Cubal” [-13.03333, 14.25000] (Mertens 1938:435); “Benguela” [-12.58333, 13.41667] (Boulenger 1887:122; Bocage 1895a:35); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:lll); “Catengue” [-13.03333, 13.73333] (Mertens 1926:152). Hulla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:35); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:78); “Kapelongo” [-14.88333, 15.08333] (Monard 1937b:78); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:35); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:35); “Kampulu” [-15.21667, 16.11667] (Monard 1937b:78); “Mulondo” [-15.63333, 15.20000] (Monard 1937b:78). Namibe: “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:35; Ceriaco et al. 2016a:59). Cunene: “riv. Mbale” [-15.16667, 16.75000] (Monard 1937b:78); “Chimporo” [-16.03333, 17.15000] (Monard 1931:109, 1937b: 148). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b:78); “Kwito region, tributary of Kubango” (Angel 1923:159); “Cubango basin (6a)” [-14.67155, 17.73525] (Conradie et al. 2016:8-9, 24); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9- 10, 24); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al. 2016:9-10, 24); “Cubango basin (46)” [-14.58981, 16.907389] (Conradie et al. 2016:9, 12, 24). Undetermined locality: “Carangi- go” (Bocage 1895a:35); “Between, Bihe and Quilenges” (Boulenger 1905:111); “without precise location” (Bocage 1870:68; Ferreira 1900:49). Taxonomic and distributional notes: Bates et al. (2013) suggested that G. nigrolineatus was not monophyletic and resurrected G. intermedius Lonnberg, 1907 for eastern populations (Kenya, Uganda, Rwanda, Tanzania, Malawi, Mozambique, Zimbabwe and South Africa). However, accu¬ rate determination of geographical boundaries for nigrolineatus and intermedius, especially in Cen¬ tral Africa (Angola, Democratic Republic of Congo, Zambia, northern Botswana and northern Namibia) will require additional sampling, as well as additional morphological analysis of speci¬ mens from throughout their extensive ranges. Bates et al. (2013) questioned the assignment of Angolan Gerrhosaurus multilineatus Bocage, 1866 specimens to G. nigrolineatus (e.g., Boulenger 1887, 1905; Ferreira 1900, 1903, 1904; Angel 1923; Monard 1937b; Mertens 1938; Hellmich 1957a, 1957b; Managas 1963; Parker 1936; Schmidt 1933; Laurent 1954), and their relationship (see G. multilineatus account). The status of the G. nigrolineatus complex in Angola is currently under investigation (M. Bates, pers. comm.) and the records plotted here should be considered as only tentatively assigned to this species. Gerrhosaurus nigrolineatus ahlefeldti (Hellmich and Schmelcher, 1956) is now considered a synonym of G. nigrolineatus, the holotype and four paratypes are present in the Zoological Museum in Hamburg and two paratypes are at the Zoolo- gische Staatssammlung, Miinchen (Hallermann 1998; Franzen and Glaw 2007). Gerrhosaurus skoogi Andersson, 1916 Desert Plated Lizard Gerrhosaurus skoogi Andersson 1916:10, fig. 2. Holotype: GNM 1387 (donor Lt. D. Fortunato). Type loeal- ity: “Port Alexander, Portuguese West Afriea” [= Tombua], Namibe Provinee, Angola. Angolosaurus skoogi'. Loveridge (1942:495), FitzSimons (1953:215), Braneh (1998:177), Ceriaeo et al. (2016a:39, 59). Global conservation status (lUCN): Least Concern. Global distribution: A deserticolous species that may form loosely-structured colonies MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 235 restricted to the Namib Desert, in southwestern Angola and northwestern Namibia. Ocurrences in Angola (Map 191): The species occurs in southwestern Angola and is restricted to the Namib Desert. Namibe: “Porto Alexandre, between Mossamedes and the mouth of the Kunene river” [-15.80000, 11.83333] (Andersson 1916; Loveridge 1942:495; FitzSimons 1953:215; Ceriaco et al. 20 16a: 59); “Praia do Navio coastal dunes, ca 12 km SSW ofNamibe” [-16.27233, 11.83164] (Ceriaco et al. 2016a:39). Taxonomic and distributional notes: The Genus Angolosaurus was established by FitzSimons (1953) to accommodate the species Gerrhosaurus skoogi Andersson, 1916. Lang (1991) considered Angolosaurus skoogi the probable sister taxon to mainland African ger- rhosaurids (Lang et al. 2003) and Nance (2007) supported a position as the most basal member of Gerrhosauridae based on morphology. Howev¬ er, Lamb et al. (2003) showed, using mitochondrial DNA data, that the species is embedded with¬ in Gerrhosaurus and this was confirmed by a multilocus phylogeny by Lamb and Bauer (2013). ConsQqgiQntXy Angolosaurus is a synonym of Gerrhosaurus (Adolphs and Bates 2010). Map Angola. 191. Distribution of Gerrhosaurus skoogi in Genus Matobosaurus Bates and Tolley, 2013 Matobosaurus maltzahni (de Grys, 1938) Western Giant Plated Lizard Gerrhosaurus Maltzahni de Grys 1938:58, figs. 1-2. Holotype: ZMH 5114 (collector A. von Maltzahn), lost ? (not reported by Hallermann 1998). Type locality: “Farm Roidina, D.S.W.-Afrika” [= Roidina Nature Farm], Namibia. Gerrhosaurus robustus: Bocage (1870:68, 1887b:203). Gerrhosaurus validus: Bocage (1895a:36), Boulenger (1905:111), Mertens (1938a:435), Frade (1963:253), Branch (1998:181). Gerrhosaurus validus maltzahni'. Loveridge (1942:497), Hellmich (1957b:58). Matobosaurus maltzahni'. Bates and Tolley (2013:470), Ceriaco et al. (2016b:59). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from central and northwestern regions of Nambia and extends into southern Angola. Ocurrences in Angola (Map 192): The species occurs in the southwestern Angola. Bengue- la: “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957b:56); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:435; Loveridge 1942:497). Huila: “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:36; Loveridge 1942:497). Namibe: “Rio Chimba” [-14.30000, 12.40000] (Bocage 1895a:36; Loveridge 1942:497; Ceriaco et al. 2016a:59); “Mossamedes” [-15.20000, 12.15000] (Bocage 1887b:203, 1895a:36; Ceriaco et al. 2016a:59); “0.5 km S of Tambor, Iona” [-16.13556, 12.42972] (Bates et al. 2013:470; Ceriaco et al. 2016a:59) and “Omuaha Lodge, Iona” [-16.19872, 12.40008] (Bates et al. 2013:470 Ceriaco et al. 2016a:59). Undetermined locality: “Between Benguella and Bihe” (Boulenger 1905:111; Loveridge 1942:497); “Mossamedes to Huil- la” (Bocage 1895a:36; Loveridge 1942:497). 236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: The species was for some time considered a subspecies of Gerrhosaurus validus (Smith, 1849). A combination of molecular, morpho¬ logical and geographical evidence, suggests that the two taxa represent separate evolution¬ ary lineages (Bates et al. 2013) that occur in allopatry, with validus distributed in southeast¬ ern Africa and maltzahni restricted to northern Namibia and southern Angola (Bates et al. 2013; Bates et al. 2014). Thus records of Ger¬ rhosaurus validus (or Gerrhosaurus robustus Peters, 1854) from Angola (Bocage 1870, 1887b, 1895a; Boulenger 1905; Mertens 1938a; Frade 1963) are, in fact, referrable to maltzahni. A new genus, Matobosaurus was erected by Bates and Tolley (2013) to incorpo¬ rate this distinctive clade of gerrhosaurs (Bates et al. 2013). ll^E 16°E 20°E 24'*E Map 192. Distribution of Matobosaurus maltzahni in Angola. Genus Tetradactylus Merrem, 1820 Tetradactylus ellenbergeri (Angel, 1922) Ellenberger’s Plated Snake-Lizard Paratetradactylus Ellenbergeri (Angel 1922:151). Holotype: MNHN 1921-514 (donated by V. Ellenberger). Type loeality: “Pays des Barotses (Rhodesie)” [= Barotseland] Zambia. Tetradaetylus lundensis (Monard 1937b:79). Syntypes: MHNC 91.0455-58 [4 speeimens] (eolleetor A. Monard). Type loeality: “Lunda, sur les bords du Tyihumbwe” [= Lunda, on the edges of Tyihumbue river] Lunda Sul Provinee, Angola. Caitia afrieana\ Boeage (1895a:37). Tetradaetylus lundensis (Monard 1937b:79). Tetradaetylus boulengeri lundensis: de Witte and i2»e i&°e 2o°e 24”e Laurent (1942b: 105). Tetradaetylus ellenbergeri ellenbergi: Laurent (1964a:55). Tetradaetylus ellenbergeri boulengeri: Laurent (1964a:55). Tetradaetylus ellenbergeri: Broadley and Cotterill (2004:43), Wagner et al. (2012:35), Conradie et al. (2016:24). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola, Democratic Republic of Congo, Tanzania and Zambia. Ocurrences in Angola (Map 193): The species occurs in norteastem Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau- rent 1964a:55). Lunda Sul: “Lunda” Ua? m. m.Xnhutkyr, of TetraJactylus ellenbergeri in [-10.96667, 20.06667] (de Witte and Laurent Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 237 I942b:I05; Wagner et al. 2012:37); “Lunda, sur les bords du Tyihumbwe” [-11.00000, 20.00000] (Monard I937b:79); “Lunda” [-10.96667, 20.06667] (de Witte and Laurent I942b:I05; Wagner et al. 2012:37). Mexico: “environs du lac Calundo (Chef Sa-Mussamba Village) 105 km east of Luso” [-11.80000, 20.86667] (Laurent 1964a:55). Cuando Cubango: “along a dammed section of the Luassingua River (23)” [-14.58972, 18.17083] (Conradie et al. 2016:24). Undetermined loca¬ tion: “sur les bords du Quando” (Bocage 1895a:37; Monard 1937b). Taxonomic and distributional notes: Bocage (1895a) cited Angolan material of Catia africana, currently Tetradactylus africanus (Gray, 1838), a species now regarded as restricted to coastal regions of KwaZulu-Natal, South Africa and adjacent southern Mozambique (Bates et al. 2014). It is likely that Bocage ’s record is referable to T. ellenbergeri (Angel, 1922). Monard (1936b) described T. lundensis from Angola, althgouh Laurent (1964a) treated it as a synonym of T. ellenbergeri boulengeri (de Witte, 1933). Wagner et al. (2012) noted only the “Lunda” record from Angola, but other historical records exist (see above). Family Scincidae Gray, 1825 Genm Acontias Cuvier, 1816 “1817” Acontias jappi (Broadley, 1968) Barotseland Blind Legless Skink Global conservation status (lUCN): Not Evalutated Global distribution: The species occurs from western Zambia to eastern Angola, west of the Zambezi River. Ocurrences in Angola (Map 194): The species occurs in Mexico Province, southeast¬ ern Angola. Moxico: “Gago Coutinho district” [-10.88333, 19.23333] (Broadley 1968a: 13; Schneider and Bauer 2009:56). Taxonomic and distributional notes: Schneider and Bauer (2009) recognized A. jappi as a full species, and were followed by Lamb et al. (2010) and Wagner et al. (2012), although molecular data are required to con¬ firm its specific status. Map 194. Distribution oi Acontias jappi in Angola. Typhlosaurus lineatus jappi Broadley 1968a: 13. Holotype: Japp). Type loeality: “Kalaho, Barotseland, Zambia”. Typhlosaurus jappi'. Schneider and Bauer (2009:56). Acontias jappi'. Lamb et al. (2010:42), Wagner et al. (2012:294). NMZB 6757 (formerly UM 6757) (collector R.G. I2“E 20°E I4'*E Acontias kgalagadi Lamb, Biswas and Bauer, 2010 Kalahari Legless Skink Typhlosaurus lineatus Boulenger 1887:432, pi. XXVIII, fig. 3. Holotype: BMNH 69.2.4.1 (presented by South African Museum). Type locality: “Cape of Good Hope,” South Africa. Acontias kgalagadi kgalagadi'. Conradie et al. (2016:25). Global conservation status (lUCN): Not Evalutated Global distribution: The species is knwon from northern Namibia and Botswana and adja¬ cent southeastern Angola. 238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 195): Cuan- do Cubango: “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9, 25). Taxonomic and distributional notes: Conradie et al. (2016) collected a single indi¬ vidual from this species from Cuando Cubango constituting the first record in Angola and the first record north of the Okavango River. Lamb et al. (2010) provided a replacement name, Acontias kgalagadi, for Typhlosaurus lineatus, which they placed in Acontias based on a molecular phylogenetic analysis. Boulenger’s name was preoccupied by Acontias lineatus Peters, 1879, a species occurring in western South Africa and southern Namibia. I2"E 1&“E 20“E 2rE Map 195. Distribution oi Acontias kgalagadi in Angola. Acontias occidentalis FitzSimons, 1941 Savanna Legless Skink Acontias plumbeus occidentalis FitzSimons 1941:275. Lectotype: PEM R5105 (formerly AM 6064) desig¬ nated by Mertens (1955:74). Type loeality: Explieitly restrieted to “Okahandja, Damaraland, Sudwestafri- ka [Namibia]” by Mertens (1955). Acontias plumbeus: Monard (1937b:96). Acontias percivali ocidentalis: Broadley (1969:20), Braneh (1998:135), Daniels et al. (2005:646), Daniels et al. (2006:354). Acontias occidentalis: Lamb et al. (2010:36), Bates et al. (2014:247). Global conservation status (lUCN): Least Concern. Global distribution: This species is endemic to southern Africa and adjacent areas in south¬ ern Angola (Branch 1998; Daniel et al. 2005; Bates et al. 2014). Ocurrences in Angola (196): The species occurs in Cunene Province near the Namibian border. Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:96; Broadley 1969:20). Taxonomic and distributional notes: The original type series included 10 specimens from across the species range, including north¬ ern Namibia, Botswana, and parts of the former Transvaal Province of South Africa. Monard (1937b) misidentified a specimen from “Mupa” southern Angola sts Acontias plumbeus Bianconi, 1849, a species endemic to southern Africa (Zimbabwe, Mozambique, Swazialnd) (Bates et al. 2014). We recently located this individual in the Musee d’Histoire Naturelle, La-Chaux-de Fond, Switzerland and identified Angola. 196. Distribution of Acontias occidentalis in MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 239 it as A. occidentalis FitzSimons, 1941. Acontias occidentalis was previously considered a sub¬ species of A. percivali Loveridge, 1935. Based on molecular phylogenetic studies the two are close¬ ly related (Daniels et al. 2006), but considered specifically distinct (Lamb et al. 2010). Genus Eumecia Bocage, 1870 Eumecia anchietae anchietae Bocage, 1870 Western Serpentieorm Skink Eumecia Anchietae Bocage 1870:66, pi. 1. Holotype: MBL specimen number unnkown (collector J.A. d’Anchieta) destroyed by fire 18 March 1978. Type locality: “le plateau de la Huilla dans I’interieur de Mossamedes” [= Huila plateau] Huila Province, Angola. Lygosoma anchietae'. Boulenger (1887:316), Themido (1941:8), Frade (1963:252). Lygosoma {Eumecia) Anchietae'. Bocage (1895a:50, 1897a: 196). Riopa {Eumecia) anchietae'. Smith (1937:230). Eumecia anchieta: Mittleman (1952:10). Riopa anchietae. Hellmich (1957b:58), Loveridge (1957:216). Eumecia anchietae'. Greer (1967:1), Broadley (1965c: 16), Spawls et al. (2004:147), Ceriaco et al. (2016a:56). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola, Democratic Republic of Congo, Kenya, Tanzania and Zambia. ij.e 2o°e 24''e Ocurrences in Angola (Map 197): The species occurs mainly in central-southwest Angola. Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a:50); “Bela- Vista” [-12.36667, 16.20000] (Hellmich 1957b:58). Benguela: “Alto Cubal” [-13.03333, 14.25000] (Hellmich 1957b:58). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:50; Themido 1941:8); “Huila” [-15.05000, 13.55000] (Bocage 1870:66, 1895a:50, 1897a:196; Boulenger 1887:316; Loveridge 1957:216; Ceriaco et al. 2016a:56); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:95). Cunene: “Riv. Mbale” [-15.16667, 16.75000] (Monard 1937b:95). Taxonomic and distributional notes: Bocage (1870) described the genus Eumecia to emphasize similarity between this species and Map 197. Distribution of Eumecia anchietae anchietae in Angola. Eumeces. However, Boulenger (1887) transferred E. anchietae to Lygosoma (Gray, 1828) where it remained until Smith (1937) revived Eumecia for a subgenus of Riopa. Mittleman (1952) gave it a full generic rank in his classification, as recognized by Greer (1967) and subsequent authors, although Loveridge (1957) continued to use Lygosoma. Metallinou et al. (2016) demonstrated that Eumecia, which shows extreme matrotrophy, is sister to Lubuya ivensii, the only other African mabuyine skink to exhibit such a high prenatal maternal investment. Eumecia anchietae major Laurent, 1964 Lunda Western Serpentieorm Skink (Endemic) Eumecia anchietae major Laurent 1964a: 80. Holotype: MD 6002 (eolleetor A. Serralheiro). Type loeality: “Calonda, steppe, Lunda,” Lunda Norte Provinee, Angola. Lygosoma anchietae'. Monard (1937b:95). 240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Not Evaluated. Global distribution: Only known from Angola. Ocurrences in Angola (Map 198): Lunda Norte: “Calonda” [-8.41667, 20.53333] (Laurent 1964a:80). Lunda Sul: “Lunda, near Dala” [-11.03333, 20.20000] (Monard 1937b:95). Taxonomic and distributional notes: Monard (1937b) first noted that a specimen from Lunda Norte Province was larger and dif¬ fered in some features form the more typical southern forms. Laurent (1964a) subsequently described Eumecia anchietae major for these northern specimens. The validity of the named subspecies of Eumecia anchietae has not been rigorously assessed. I2"E 1&“E 20“E 24’’E Map 198. Distribution of Eumecia anchietae major in Angola. Genus Fey lima Gray, 1845 Feylinia currori Gray, 1845 Western Forest Feylinia Feylinia Currori Gray 1845:129: Lectotype, BMNH 1946.8.13.79 (formerly BMNH xv.2.a) (collector J. Cur- ror) designated by Brygoo and Roux-Esteve (1983). Type locality: “Angola.” The paralectotype from “Coast of Africa” is referable to E polylepis (see Brygoo and Roux-Esteve 1983). Feylinia Currori: Bocage (1873b:214; 1887a:179, 1895a:57), Peters (1877a:614), Boulenger (1905:111). Feylinia eurrori: Boulenger (1887:431), Ferreira (1904:116), Parker (1936:139), Hellmich (1957b:59), Lau¬ rent (1964a:84), Brygoo and Roux-Esteve (1983:312), Spawls et al. (2004:159), Wagner and Schmitz (2006:183), Chirio and LeBreton (2007:232). Global conservation status (lUCN): Not Evaluated. Global distribution: The most wide- ii-e i6“e io^e 24<-e spread species of Feylinia occuring from Cameroon to Central African Republic and the Democratic Republic of Congo, east to Tanza¬ nia and south to Angola, with two isolated records from Sierra Leone (questionable) and Nigeria. Ocurrences in Angola (Map 199): The species occurs in the northern regions of the country including the Cabinda enclave. Cabin¬ da: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877:614); “Molembo” [-5.33333, 12.20000] (Bocage 1895a:57); “Cabinda” [-5.55000, 12.18333] (Bocage 1887a:179, 1895a:57); “Chiloango” [-5.18333, 12.18333] (Brygoo and Roux-Esteve 1983:312; Wagner and Schmitz 2006:184). Lunda Norte: “Cassan- Map199. Distribution of Feylinia currori in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 241 guidi” [-7.48333, 21.31667] (Laurent I964a:84). Bengo: “Bom Jesus (Quanza)” [-9.16667, 13.56667] (Ferreira 1904:116). Kwanza Norte: “Roca Novo Duro, Piri-Dembos” [-8.56667, 14.50000] (Hellmich I957b:59; Brygoo and Roux-Esteve 1983:312; Wagner and Schmitz 2006:184); “Catari” [-9.09548, 15.41905] (Ferreira 1904:116); “Golungo Alto” [-9.13333, 14.76667] (Ferreira 1904:116; Brygoo and Roux-Esteve 1983:312; Wagner and Schmitz 2006:184); “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1904:116); “Zembe” [-9.31667, 14.66667] (Ferreira 1904:116). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:139; Brygoo and Roux-Esteve 1983:312; Wagner and Schmitz 2006:184). Undetermined locality: “Mayumba” (Bocage 1887a: 179); “Schinga” (Brygoo and Roux-Esteve 1983:312). Taxonomic and distributional notes: Boulenger (1887) mistakenly cited both syntype spec¬ imens from “Angola.” Brygoo and Roux-Esteve (1983) reviewed the history of the types and pro¬ vided a point locality map for the species’ global range. Feylinia elegans (Hallowell, 1852) Elegant Feylinia Acontias elegans Hallowell 1852a:64, fig. p. 64. Type: ANSP 9456, 9667 [2 speeimens] (eolleetor H.A. Ford). Type loeality: “Liberia, West Coast of Afriea” (Hallowell 1852a:65) [“probably in error for Gabon” Loveridge (1957)]. Anelytrops elegans: Boeage (1866b:214, 1867e:227, 1870:57). Feylinia Currori: Boeage (1873b:214). Feylinia eurrori: Boulenger (1887:431). Feylinia elegans: Sehmidt (1919:605), de Witte and Laurent (1942b: 108, 1943:36), Brygoo and Roux-Esteve (1983:317), Chirio and LeBreton (2007:234). Feylinia eurrori elegans: Loveridge (1933:328, 1957:223). Global conservation status (lUCN): Not Evaluated. Global distribution: Is a typical Central African forest species, occurring in savannas with western affinities, from Gabon, through Central African Republic, Democratic Republic of Congo to northern Angola. Ocurrences in Angola (Map 200): The species is known from some restricted areas in the north of the country (e.g., extreme north of Zaire Province; northwestern regions of Lunda Norte Province) including the Cabinda enclave. Cabinda: “Cabinda” [-5.55000, 12.18333] (Bocage 1866b:214). Namibe: “Mossamedes” [-15.20000, 12.15000] (Bocage 1867c:227) [in error]. Taxonomic and distributional notes: Feylinia elegans was originally described by Hallowell (1852a) as Acontias elegans with its holotype from “Liberia,” and then redescribed as Sphenorhina elegans (Hallowell, 1857) based on two specimens from “Gaboon.” Schmidt (1919) subsequently identified ANSP 9456 as the holotype and Loveridge (1957) explained the discrepancy m locality, which s & applies to other taxa collected by Ford and reported upon by Hallowell. The species was tentative¬ ly referred to the synonymy of Feylinia currori Gray, 1845 by several early workers (e.g., Bocage 1873b; Boulenger 1887), although Schmidt (1919) recognized F. elegans as specifically distinct. 242 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Loveridge (1933, 1957) considered it as a subspecies of currori, whereas de Witte and Laurent (1942b, 1943), considered a full species, as did Brygoo and Roux-Esteve (1893). The distribution of F. elegans in Angola is probably limited to Cabinda region, and some isolated parts of the Congolian Forest- Savanna in northern regions of the country. The “Mossamedes” record in the extreme southwestern region of the country cited by Bocage (1867c) surely represents a misiden- tification or incorrect locality. Feylinia grandisquamis Muller, 1910 Large-Scaled Feylinia Feylinia Currori Gray 1845:129: Lectotype, BMNH 1946.8.13.79 (collector J. Curror) designated by Brygoo and Roux-Esteve (1983). Type locality: “Angola.” Feylinia eurrori grandisquamis Muller 1910:591. Syntypes, ZSM 413/1909 [5 specimens], probably lost fide Brygoo and Roux-Esteve (1983). Type locality: “Dibongo bei Edea” Cameroon. Feylinia eurrori grandisquamis: Schmidt (1919:607). Feylinia elegans grandisquamis: de Witte and Laurent (1942b: 108), Laurent (1964a: 84). Feylinia grandisquamis: Brygoo and Roux-Esteve (1983:330), Chirio and LeBreton (2007:236). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from lowland areas in dense forests, extending from Cameroon to northern Angola and Central ,2“e i6'e 2o°e 24-e Africa. Ocurrences in Angola (Map 201): The species occurs in the north of the country. Zaire: “Cabra, Kakongo” [-6.7500, 12.6500] (Brygoo and Roux-Esteve 1983:331). Unde¬ termined Locality: “Cuango” (Laurent 1964a: 84) (Malanje Province, impossible to georeferenced. See History Section for more detailed information). Taxonomic and distributional notes: Brygoo and Roux-Esteve (1983) plotted the locality “Cabra, Kakongo” in Zaire Province, however Kakongo could also refer to the river of the same name (or related places) in Cabin¬ da. We are unaware of any definitive informa¬ tion that would confirm which locality was intended, but here follow Brygoo and Roux- Esteve (1983). Map 201. Angola. Distribution of Feylinia grandisquamis in Genus Lepidothyris Cope, 1892 Lepidothyris hinkeli joei Wagner, Bohme, Pauwels and Schmitz, 2009 Joe’s Red-Flanked Skink Lepidothyris hinkeli Wagner et al. 2009:12. Holotype: ZFMK 55701 (collector Pacome and H. Hinkel). Type locality: “Cyamudongo, Nyungwe, Rwanda.” Lepidothyris hinkeli joei Wagner et al. 2009:14, figs. 8-9. Holotype: ZFMK 64410 (collector E. Fischer and H. Hinkel). Type locality: “Oyo, Bokouele, Peoples Rep. Congo.” Moehlus fernandi: Laurent (1964a:78). Lygosoma fernandi: Chirio and LeBreton (2007:266). Global conservation status (lUCN): Not Evaluated. Global distribution: Lepidothyris hinkeli joei is a lowland and submontane forest form, rang- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 243 ing around the edges of the Congo Basin in the Democratie Republie of Congo, People’s Republie of Congo and far northeastern Ango¬ la. Ocurrences in Angola (Map 202): The speeies is only known from “Dundo”, Lunda Norte Provinee. Lunda Norte: “Dundo, foret de la Luachimo” [-7.38333, 20.85000] (Lau¬ rent 1964a: 78). Taxonomic and distributional notes: The specifie epithet fernandi (Burton, 1836) has alternately been assoeiated with several different genera: Lygosoma Hardwieke and Gray, 1827, Lepidothyris Cope, 1892, Riopa Gray, 1839 and Mochlus Gunther, 1864 (Wag¬ ner et al. 2009). Laurent (1964a) signaled the first and only record from Angola. Wagner et al. (2009) VQSunQctQd Lepidothyris Cope, 1892 for the fernandi group. They also provided a morphological and genetic analysis in the L. fernandi complex, resulting in the description of new taxa (Wagner et al. 2009), including a new subspecies, Lepidothyris hinkeli joei Wagner, Bohme, Pauwels and Schmitz, 2009 to accommodate the Angolan specimen cited by Laurent (1964a), which we recently examined in the Museu Dundo, and additional material from the Democratic Republic of Congo and People’s Republic of Congo. Map 202. Distribution of Lepidothyris hinkeli joei in Angola. Genus Leptosiaphos Schmidt, 1943 Leptosiaphos dewittei (Loveridge, 1934) Lygosoma (Siaphos) compressicauda de Witte 1933: Overlaet). Type loeality: “Sandoa” [Lualaba Provinee, Demoeratie Republie of Congo]. Lygosoma dewittei: Parker (1936:139). Leptosiaphos dewittei: Broadley (1989e:445), Broadley and Cotterill (2004:42). Global conservation status (lUCN): Not Evaluated. Global distribution: This skink ranges from western Angola to the Upemba National Park, Democratic Republic of Congo where it is most common in the plateau areas. Ocurrences in Angola (Map 203): The species is only known from northeastern Ango¬ la. Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:139). Taxonomic and distributional notes: The name Siaphos dewittei Loveridge, 1934 is a replacement name for Lygosoma {Siaphos) compressicauda de Witte, 1933, itself a junior DE Witte’s Five-Toed Skink 175, figs. 1-4. Holotype: MRAC 8614 (collector F.G. I2"E l&°i: 20°E 24“E Map 203. Distribution of Leptosiaphos dewittei in Angola. 244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II homonym of Lygosoma (Hinulia) compressicauda Werner, 1897. The disjunct localities for this taxon suggest that further taxonomic work is required. Genus Lubuya Horton, 1972 Lubuya ivensii (Bocage, 1879) Ivens’ Skink Euprepes Ivensi Bocage 1879a:97. Syntypes: MBL (3 specimens) numbers not known (collectors H. Capello and R. Ivens), destroyed by fire 18 March 1978. Type locality: “Bihe, dans I’interieur de Benguella” [= Bie], Bie Province, Angola. Mabuya ivensi septemlineata (Laurent 1964a:77, fig. 23). Holotype: MD 5427 (collector A. Barros Macha¬ do). Type locality: “Alto Chicapa, Lunda” [Lunda Sul Province, Angola]. Euprepes Ivensi'. Bocage (1879b:95). Mabuia ivensii'. Boulenger (1887:197). Lygosoma Ivensii'. Bocage (1895a:48, 1897a:196). Mabuya ivensii'. Monard (1937b:86), Mananas (1963:233), Branch and Haagner (1993). Lubuya ivensii'. Horton (1972:17), Metallinou et al. (2016:4). Traehylepis ivensii'. Wagner et al. (2012:39). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is widely distributed through the drainage systems of Ango¬ la, extending into the adjacent regions of the southern Democratic Republic of Congo and Zambia. Ocurrences in Angola (Map 204): The species is known from central-east Angola. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:77; Branch and Haagner 1993:112; Wagner et al. 2012:37); “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a; 77; Branch and Haagner 1993:112; Wagner et al. 2012:37); “Dala” [-11.03333, 20.20000] (Monard 1937b;86; Laurent 1964a: 78; Branch and Haagner 1993:112; Wagner et al. 2012:37). Malanje: “Rio Loan- do” (R. Luando, pres de Mongoa; Luando River, tributary of the Cuanza River, near Mon¬ goa)” [-11.55000, 18.15000] (Bocage 1879b:95; Laurent 1964a:78; Branch and Haagner 1993:112; Wagner et al. 2012:37); “entre Malanje et Pungo Andongo” (Laurent 1964a:78). Moxico: “source of the Calombe Map 204. Distribution of Lubuya ivensii in Angola. River (Luso)” [-11.83333, 19.93333] (Managas 1963:233; Horton 1972:17); “Luso (=Luena)” [-11.78333, 19.91667] (Horton 1972:17; Branch and Haagner 1993:112; Wagner et al. 2012:37). Bie: “Bihe, dans Tinterieur de Benguella” [-12.38333, 16.95000] (Bocage 1879a:97); “Benguella” (“Bihe” according to the original publication fide Bocage 1879a:97) [-12.38333, 16.95000] (Boulenger 1887:197; Horton 1972:17). Huambo: “Huambo” [-12.76667, 15.73333] (Horton 1972:17; Branch and Haagner 1993:109, 112; Wagner et al. 2012:37); “R. Cuando River, affluent du Cunene” [-13.38333, 15.71667] (Laurent 1964a:78; Branch and Haagner 1993:112; Wagner et al. 2012:37). Undetermined locality: “Quando” (Bocage 1895a:48, 1897a: 196; Branch and Haag¬ ner 1993:112); “sur les bords du Quanza et du Quando (margens do Quanza e do Quando)” (Bocage 1895a:49, 1897a: 196); “R. Cuanza” (Laurent 1964a:78; Branch and Haagner 1993:112; Wagner et al. 2012:37). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 245 Taxonomic and distributional notes: Boulenger (1887) incorrectly restricted Bocage’s type locality to “Benguela,” an action later aeeepted by Horton (1972), Braneh and Haagner (1993), and Wagner et al. (2012). The species was described in Euprepes but later moved by Bocage (1895a) to Lygosoma. Horton (1972) described the monotypie genus Lubuya to aeeommodate M. ivensii, differentiating it from Mabuya as then construed. Greer (1977) suggested thsit Lubuya should return to Mabuya, or by implieation to Trachylepis, the name subsequently alloeated to Afriean mabuyine skinks (Bauer 2003). However, Metallinou et al. (2016) ressurected Lubuya to aeeommodate T. ivensii, whieh they found to be the distantly related sister to Eumecia. Monard (1937b) was the first to reeognize that specimens from northeastern Angola had an extra white line on the lower flank and Laurent (1964a) subsequently ereeted Mabuya ivensi septemlineata to reeognize this form; its validity has not been critically assessed. Genus Melanoseps Boulenger, 1897 Melanoseps occidentalis (Peters, 1877) Western Limbless Skink Herpetosaura occidentalis Peters 1877b:416. Holotype: ZMB 9122. Type loeality: Camemns [= Cameroon], Melanoseps oeeidentalis (Peters): Laurent (1964a:81). Melanoseps oeeidentalis zairensis: Brygoo and Roux-Esteve (1981:1179). Melanoseps oeeidentalis: Chirio and Ineieh (2006:38), Chirio and LeBreton (2007:270), Trape et al. (2012:368). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is limit¬ ed to Central Africa, from Gabon to Angola. Ocurrences in Angola (Map 205): The species is only known from “Camaeonde, Dundo” in Lunda Norte Province. Lunda Norte: “galerie forestiere de la riviere Cama¬ eonde (affl. gauehe de la Luaehimo, Dundo)” [-7.36667, 20.83333] (Laurent 1964a:81; Bry¬ goo and Roux-Esteve 1981:1171). Taxonomic and distributional notes: Brygoo and Roux-Esteve (1981) deseribed a new subspeeies, Melanoseps occidentalis zairensis, from the Demoeratic Republie of Congo, distinguished from the nominotypical form by its ventral scale eount (Chirio and Ine- ich 2006), although subsequent authors have treated the speeies as monotypie (Chirio and LeBreton 2007; Trape et al. 2012). Map 205. Distribution oi Melanoseps oeeidentalis in Angola. Genus Mochlus Giinther, 1864 Mochlus sundevallii (Smith, 1849) Sundevall’s Writhing Skink Eumiees [sie] (Riopa) sunderallii Smith 1849b:ll. Holotype: BMNH 1946.8.7.3 (formerly BMNH 65.5.4.59) (eolleetor A. Smith) (see notes below). Type loeality: “eountry to the eastward of the Cape Colony” [South Afriea; “probably from W. Transvaal” 7/t/e Broadley (1966a:4)]. Moehlus afer: Boeage (1867b:222, 1867e:227). 246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Eumeces reticulatus: Bocage (1879c:88). Lygosoma sundevalii: Boulenger (1887:307), Bocage (1895a:49), Branch (1998:150). Lygosoma modestum: Monard (1937b:94); Frade (1963:253). Lygosoma modesta modesta: Hellmich (1957b:57), Loveridge (1957:215). Mochlus sundevalii sundevalii-. Laurent (1964a:78), Bates et al. (2014:259). Riopa sundevalii-. Haacke (1965:18). Riopa sundevalii sundevalii-. Broadley (1966a:4). Lygosoma sundevalii-. Spawls et al. (2004:146). Moehlus sundevalii-. Wagner et al. (2009:2), Spawls (2017), Ceriaco et al. (2016a:56), Conradie et al. (2016:25). Global conservation status (lUCN): Least Concern. Global distribution: The species is widely distributed mainly in southeastern Africa, from Uganda south to Mozambique and Zimbabwe, and west to Namibia, Zambia, and Angola. Ocurrences in Angola (Map 206): The species occurs in southern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1964a:78). Kwanza Norte; “Piri-Dem- bos” [-8.56667, 14.50000] (Hellmich 1957b:57). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage, 1867b:222, 1895a;49); “Benguela” [-12.58333, 13.41667] (Bocage, 1867b:222, 1895a:49; Boulenger 1887:307; Broadley 1966a;3); “Alto Cubal” [-13.03333, 14.25000] (Hellmich 1957b:57). Huila; “Huila” [-15.05000, 13.55000] (Laurent 1964a:78). Namibe: “10 mis E of Caracul” [-15.01667, 12.66667] (Haacke 1965:18; Broa¬ dley 1966a:4; Ceriaco et al. 2016a:56); “Capangombe” [-15.10000, 13.15000] (Boca¬ ge 1895a:49; Ceriaco et al. 2016a:56); “Mossa- medes” [-15.20000, 12. 15000] (Bocage Distribution of Mochlus sundevalii in Angola. 1867b:227, 1895a:49; Ceriaco et al. 2016a:56); “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a;49; Ceriaco et al. 2016;a 56). Cunene: “Mupanda (Kuanyama)” [-17.13333, 15.76667] (Monard 1937b:94; Broadley 1966a:4). Cuando Cubango: “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9, 10, 12); “Cuando basin (42)” [-17.49611, 23.13444] (Conradie et al. 2016:9, 10, 12); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9, 10, 12). Taxonomic and distributional notes: This species was previously included in Lygosoma, which is now restricted to Asia (Wagner et al. 2009). This species is endemic to Africa and widely distributed mainly in southeastern Africa (Bates et al. 2014). There are two Angolan records from “Dundo” (Laurent 1964a) and “Piri-Dembos” (Hellmich 1957b) that probably correspond to a misidentification, since the distribution range for the species in the country is restricted to the southern regions. Smith’s description appears to refer to a single individual and FitzSimons (1937) identified one potential BMNH specimen matching the description. This corresponds to BMNH 1946.8.7.3 and is entered in the BMNH registers as presented by Sir Andrew Smith but, oddly, as coming via the St. Petersburg Museum (ZIL). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 247 Genus Panaspis Cope, 1868 Panaspis breviceps (Peters, 1873) Peters’ Snake-Eyed Skink Euprepes (Mabuia) breviceps Peters 1873:604. Syntypes: ZMB 6303 (purehased from [H.H. ?] Higgins), 8017 (eolleetor A. Reiehenow). Type loeality: “Gabon” (ZMB 6303) and “Cameruns” [Cameroon] (ZMB 8017). Lygosoma {Panaspis) breviceps: Parker (1936:139). Panaspis breviceps: Mittleman (1952:17), Ferret (1973:595, 1975:8), Fuhn (1972:266), Greer (1974:29), Sehmitz et al. (2005:23), Chirio and LeBreton (2007:272), Trape et al. (2012:376), Medina et al. (2016:410). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Cameroon, Central African Republic, Gabon into the central Democratic Republic of Congo to Angola. Ocurrences in Angola (Map 207): The species is represented by a single west central Angolan record. Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:139; Fuhn 1972:266). Taxonomic and distributional notes: Bauer et al. (2003) provided a detailed expla¬ nation for the recognition of two syntypes, rather than a single holotype. The specific epi¬ thet breviceps (Peters, 1873) has successively been associated with several genera. Fuhn (1972) and Ferret (1973) placed it in Panaspis, which had been revalidated by Mittleman (1952). Broadley (1989), however, treated it as Leptosiaphos {Lacertaspis) breviceps. Most recently Medina et al. (2016) included it in an expanded genus Panaspis in which they *- included species formerly assigned to Afroablepharus based on a molecular phylogenetic Map 207. Distribution of Panaspis breviceps in Angola, revision of the group. Panaspis cabindae (Bocage, 1866) Cabinda Snake-Eyed Skink Ablepharus cabindae Bocage 1866h:64. Syntypes: MBL (3 speeimens) numbers not known (eolleetor J.A. d’Anehieta), destroyed by fire 18 March 1978. Type locality: “Cabinda (Afrique occidental au nord du Zaire)” Cabinda Provinee, Angola. Ablepharus Cabindae: Boeage (1866a:45, 1867b:224, 1887a:179, 1897a:196). Ablepharus cabindae: Peters (1877a:614), Boeage (1895a:51), Boulenger (1887:352), Ferreira (1904:116), Parker (1936:136). Ablepharus aeneus: Boulenger (1887:352). Riopa {Panaspis) cabindae: Smith (1937:229). Panaspis cabindae: Greer (1974:29), Ferret (1975:8), Sehmitz et al. (2005:23), Ineieh and Sehmitz (2010), Ceriaeo et al. (2016b:65), Ceriaeo et al. (2016a:57), Medina et al. (2016:411). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from Angola and the Democratic Republic of Congo. Ocurrences in Angola (Map 208): The species is known chiefly from western Angola, 248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II including several localities along the coast. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:45, 1866b;64, 1867b:224, 1897a:196; Boulenger 1887:352). Zaire: “Soyo” [-6.134903, 12.368935] (Medi¬ na et al. 2016:411); “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a:179, 1895a:51, 1897a:196). Bengo: “Catete” [-9.11667, 13.70000] (Ferreira 1904:116); “Riverine Forest, Bengo” (Medina et al. 2016:411). Uige: “Kimpa Vita Uni Campus” [-8.607967, 20.968804] (Medina et al. 2016:411). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:65). Lunda Norte: “Lagoa Carumbo” [-7,74422, 19,95467] (Branch and Conradie 2015:200; Medina et al. 2016:411). Kwanza Norte: “Lucalla” [-9.40000, 15.03333] (Ferrei¬ ra 1903:116). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:139). Benguela: “Dombe” [-12.95000, 13.10000] (Bocage 1867b:224, 1895a:51, 1897a:196). Huila: “SLebaPass” [-15.07003, 13.24339] (Medina et al. 2016:411); “Benero Campsite near Jamba” (Medina et al. 2016:411). Namibe: “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:51, 1897a:196; Ceria¬ co et al. 2016a:57). Taxonomic and distributional notes: The genus Panaspis was erected by Cope (1868) to accommodate his newly described Panaspis anaeus from Angola (Boulenger 1887), for some time considered a synonym of Ablepharus cabindae Bocage, 1866 (Perret 1975; Schmitz et al., 2005). Boulenger (1887) placed the entire genus Panaspis into the synonymy of Ablepharus, although Smith (1937) regarded it as a subgenus in the genus Riopa Gray, 1839. Mittleman (1952) revised the genera belonging to the subfamily Lygosominae and regarded the genus Panaspis as valid. Greer (1974) erected the new genus Afroablepharus for those species with an ablepharine eye, while all species with a movable lower eyelid, including P cabindae with its preablepharine eye were included in Panaspis. Most recently Medina et al. (2016) included it in an expanded genus Panaspis in which they included species formerly assigned to Afroablepharus based on a molecu¬ lar phylogenetic revision of the group. The currently accepted range for the species in Angola is limited to the northwestern regions (Schmitz et al. 2005; Ineich and Schmitz 2010), and records from “Dombe,” Benguela Province, and “Capangombe,” Namibe Province, certainly correspond to another congener. Unfortunately, the relevant specimens were destroyed in the 1978 fire in the Museu Bocage. Panaspis maculicollis Jacobsen and Broadley, 2000 Speckle-Lipped Snake-Eyed Skink Panaspis maculicollis Jacobsen and Broadley, 2000:65, fig. 3b. Holotype: TM 76922 (eolleetors N.H.G. Jaeobsen and R.E. Newbery). Type loeality: “Klein Tshipise, Mutale Distriet, Northern Provinee [= Limpopo Provinee], South Africa.” Panaspis maculicollis'. Conradie et al. (2016:25), Medina et al. (2016:414). Global conservation status (lUCN): Not Evaluated. Global distribution: Centered on the Kalahari sand areas of northern Botswana and western Map 208. Distribution of Panaspis cabindae in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 249 Zimbabwe, extending eastwards via the Zam¬ bezi and Limpopo valleys into western Mozambique and Umpopo Province of South Africa. Ocurrences in Angola (Map 209): A sin¬ gle record for southern Angola. Cuando Cubango: “Cuando basin (43)” [-17.53500, 23.18916] (Conradie et al. 2016:25; Medina et al. 2016:411). Taxonomic and distributional notes: Greer (1974) erected the new genus Afroable- pharus for species with an ablepharine eye, following a suggestion first made by Smith (1935). Perret (1975) trQSitQd Afroablepharus as a subgenus of Panaspis. However, Medina et al. (2016) demonstrated that Panaspis was made paraphyletic by Afroablepharus and synonymized the latter with the former, which has priority. Conradie et al. (2016) collected a single individual of P. maculicollis from Cuando this species in Angola. ll^E t6°E I4"E Map 209. Distribution of Panaspis maculicollis in Angola. Cubango Province, establishing the first record of Panaspis aff. wahlbergii (Smith, 1849) Wahlberg’s Snake-Eyed Skink Cryptoblepharus walbergii [sic] Smith 1849:10. Syntypes: BMNH 1946.8.18.49-50 (formerly BMNH 65.5.4.112-113). Other unloeated syntypes are implied (fide FitzSimons 1937:269). Type loeality: “eoun- try to the eastward of the Cape Colony” (Smith 1849:10), [= Natal], South Afriea. Ablepharus Wahlbergii'. Boeage (1895a:52). Ablepharus wahlbergii'. Boulenger (1905:111), Loveridge (1957:219). Afroablepharus wahlbergii'. FitzSimons (1937:269), Greer (1974:32), Perret (1975:8), Sehmitz et al. (2005:19), Bates et al. (2014:257). ^ ll“F. 14“ F, Panaspis wahlbergii'. Braneh (1998:159), Jaeohsen and Broadley (2000:63), Braneh and Conradie (2015:200), Medina et al. (2016:410). Global conservation status (lUCN): Not Evaluated. Global distribution: Based on molecular data published by Medina et al. (2016) P. wahlbergii sensu stricto may be limited to southeastern Africa, however, specimens cur¬ rently assigned to P. wahlbergii sensu lato are distributed from Tanzania and the southern Democratic Republic of Congo south to South Africa. Ocurrences in Angola (Map 210): The species is poorly documented in Angola and it might be more widespread in the country. Benguela: “Cahata” [-13.73333, 15.06667] (Boeage 1895a:52). HmTa: “Caconda” Map 210. Distribution of Panaspis aff. wahlbergii in Angola. 250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-12.35000, 14.81667] (Bocage 1895a:52). Undetermined Locality: “Between Benguela and Bihe” (Boulenger 1905:111). Taxonomic and distributional notes: The African snake-eyed skinks have been subject to large scale taxonomic changes both at the generic and subgeneric levels (Fuhn 1969; Ferret 1973, 1975; Greer 1974; Schmitz et al. 2005; Medina et al. 2016; see also notes in above species accounts). Panaspis wahlbergii has long been used to refer to skinks occurring throughout much of southern and eastern Africa. The discovery by Jacobsen (1989) of the species subsequently described as P. maculicollis Jacobsen and Broadley, 2000, in sympatry with P. wahlbergii revealed that older records of Panaspis need to be carefully investigated. The molecular phylogeny of Medina et al. (2016) has identified numerous species level lineages within the range previously attributed to P. wahlbergii. Specimens superficially resembling P. wahlbergii have recently been collected in Quigama National Park, Luanda Province (M. Marques and L. Ceriaco pers. obs.) The misspelling of the specific epithet in the original description was corrected by Smith in an errata slip issued with the original work, thus the corrected version of the name {wahlbergii) is ICZN Code compliant. Genus Sepsina Bocage, 1866 Sepsina angolensis Bocage, 1866 Angolan Reduced-Limb Skink Sepsina angolensis Bocage 1866b:63. Holotype: MBL specimen number unknown (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga, a I’interieur d’Angola” [= Calan- dula,] Malanje Province, Angola. Sepsina angolensis Bocage (1866a:45, 1867b:223, 1870:68, 1895a:53, 1896a: 111, 1897a: 197), Peters (1881:147), Boulenger (1887:421, 1905:111), Ferreira (1904:117), Monard (1937b:95), Mertens (1938a:438), de Witte and Laurent (1943:16), Laurent (1964a:81), Branch and McCartney (1993:1), Branch (1998:1467), Ceriaco et al. (2016a:56). Seelotes angolensis: Hellmich (1957a:68). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the Democratic Republic of Congo, Angola, Zambia and northern Namibia. Ocurrences in Angola (Map 211): Species records are widely distributed across the entire country, except the far northwest and northeastern regions. Bengo: “Catete” -9.11667, 13.70000] (Ferreira 1904:117). Kwanza Norte: “Zembe” [-9.31667, 14.66667] (Ferreira 1904:117). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:45, 1866b:63, 1867b:223, 1895a:53, 1897a:197); “Malanje” [-9.55000, 16.35000] (Bocage 1895a:53); “Pungo-Andon- go” [-9.66667, 15.58333] (Boulenger 1905:111). Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:81). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1867b:223). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:95). Benguela: “Ebanga” [-12.73333, ll-E 16“E 20“E 14“ E Map 211. Distribution of Sepsina angolensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 251 14.73333] (Monard I937b:95); “Hanha” [-13.30000, 14.20000] (Bocage I896a:III); “Cahata” [-12.35000, 14.81667] (Bocage I895a:53); “Quissange” [-12.43333, 14.05000] (Bocage I895a:53); “Quidumbo” [-12.46667, 14.93333] (Bocage I895a:53); “Dombe” [-12.95000, 13.10000] (Bocage I867b;223); “Entre Rios” [-13.01667, 14.63333] (Hellmich I957a:68); “Cubal” [-13.03333, 14.25000] (Mertens I938a:438); “Ganda” [-13.03333, 14.63333] (Hellmich 1957a:68). HmTa: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:53); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:95); “Indungu” [-14.81667, 16.26667] (Monard 1937b:54); “Boca de Humpata, Sa da Bandeira” [-14.93333, 13.51667] (Laurent 1964a;81); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:53); “Chibemba (Gambos), Cunene” [-15.75000, 14.08333] (Laurent 1964a:81). Namibe: “Capangombe” [-15.10000, 13.15000] (Monard 1937b;95; Ceriaco et al. 2016a:56). Cunene: “Kuvelai” [-15.65000, 15.80000] (Monard 1937b:95); “Chimporo” [-16.03333, 17.15000] (Monard 1937b:95). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b:95); “vicinity of Cuito Cuanavale - approximately 75 km W of Mavinga” [-15.78333, 19.70000] (Branch and McCartney 1993:1). Undetermined Locality: “Cuango” (Peters 1881:147; Bocage 1895a:53) (Malanje Province, impossible to georeferenced. See History Section for more detailed information). Taxonomic and distributional notes: None. Sepsina bayonii (Bocage, 1866) Bayao’s Reduced-Limb Skink Dumerilia Bayonii Bocage 1866b:63. Holotype: MBL specimen number unknown (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Loanda” [= Luanda], Luanda Province, Angola. Scelotes bipes: Gunther (1865a:480). Dumerilia Bayonii'. Bocage (1866a:45) Scincodipus congicus: Peters (1877a:614). Dumerilia Bayonii'. Bocage (1882a:299, 1897a:197). Sepsina bayonii'. Boulenger (1887:422), Frade (1963:252). Sepsina (Dumerilia) Bayonii'. Bocage (1895a:55). Dumerilia Bayonii'. de Witte and Laurent (1943:16). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and the Democratic Republic of Congo. Ocurrences in Angola (Map 212): The species occurs in the northwest of the country, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614; Bocage 188ab:299, 1895a:55). Luanda: “Loanda (Forte do Penedo)” [-8.78333, 13.26667] (Bocage 1866a:45); “Loanda” [-8.83333, 13.26667] (Bocage 1866b:63, 1882a:299, 1895a:55, 1897a:197). Bengo: “Ambriz” [-7.844312, 13.106493] (Boulenger 1887:422; Bocage 1895a:55). Kwanza Sul: “Conda” [-11.28333, 14.33333] (Gunther 1865a:480). Undetermined: “Carangigo” (Boulenger 1887:422; Bocage 1895a:55). Map 212. Distribution of Sepsina bayonii in Angola. 252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: The species remains poorly known and may have a broader distribution than currently appreciated. Specimens were recently collected at Quigama National Park, Bengo Province (pers. obs.). Gunther (1865a) mentioned Scelotes bipes (Linnaeus, 1766) collected by Welwitsch in “Condo,” in Kwanza Sul Province. Scelotes bipes is endemic to South Africa and this record is likely based on S. bayonii, which likewise lacks forelimbs. We tentatively include this record here. Sepsina copei Bocage, 1873 Cope’s Reduced-Limb Skink (Endemic) Sepsina Copei Bocage 1873b:212. Syntypes: MBL (3 specimens) catalogue numbers unknown (collector J.A. d’Anchieta [Dombe] F.A.P. Bayao [Novo Redondo]), destroyed by fire 18 March 1978. Type locality: “Dombe,” Benguela Province and “Novo Redondo, au nord de Benguella sur le littoral” [= Sumbe], Kwan¬ za Sul Province, Angola. Sepsina eopii: Boulenger (1887:421). Sepsina Copei'. Bocage (1895a:54, 1897a: 197). Sepsina eopei: de Witte and Laurent (1943:16), Ceriaco et al. (2016a:57). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is h^e known from Angola. Ocurrences in Angola (Map 213): The species occurs in western Angola along the coast. Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1895a:54, 1897a:197). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1873b:212, 1895a:54, 1897a: 197). Benguela: “Benguela” [-12.58333, 13.41667] (Boulenger 1887:421; Bocage 1895a:54); “Dombe” [-12.95000, 13.10000] (Bocage 1873:212, 1895a:54, 1897a:197). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a:54, 1897a:197; Ceriaco et al. 2016a:57). Taxonomic and distributional notes: Sepsina copei is poorly known and like its con¬ geners, its taxonomic status has never been thoroughly investigated. Map 213. Distribution of Sepsina copei in Angola. Genus Trachylepis Fitzinger, 1843 Trachylepis acutilabris (Peters, 1862) Wedge-Snouted Skink Euprepes acutilabris Peters 1862a:19. Syntypes: ZMB 4214, 4215, 4216, 64286-90 (formerly ZMB 4214 part), 64348-49 (formerly ZMB 4216 part) fide Bauer et al. (2003) (collector C.H. Hahn) (see notes below). Type locality: “Neu-Barmen” [= Gross Barmen, Otjozondjupa Region, Namibia]. Euprepes acutilabris'. Bocage (1870:68), Peters (1877a:614). Mabuya acutilabis: Boulenger (1887:208), Bocage (1895a:46), Schmidt (1919:551), Monard (1937b:37), Mertens (1937a: 11), Laurent (1947:8, 1954a:65, 1964a:75), Hellmich (1957a:58, 1957b:53), Branch (1998:151). Trachylepis acutilabris'. Haacke (2008:90), Ceriaco et al. (2016a:34, 57). Global conservation status (lUCN): Not Evaluated. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 253 Global distribution: This species occupies sandy substrates from Cabinda and the western Democratic Republic of Congo, through western Angola continuously through to central Namibia, with several additional scattered populations. Ocurrences in Angola (Map 214): The species occurs in the western Angola, along the coast. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614; Bocage 1895a: 46). Zaire: “Noki” [-5.86667, 13.43333] (Schmidt 1919:551-553), “Antonio [do Congo]” [-6.13333, 12.36667] (Schmidt 1919:551-553). Luanda: “Luanda” [-8.83333, 13.26667] (Hellmich 1957a:58). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1895a:46). Benguela: “Plage de Lobi- to (Restinga)” [-12.33333, 13.50000] (Laurent 1954a:65, Laurent 1964a:75); “Lobito” [-12.35000, 13.55000] (Monard 1937b:94; Mertens 1937a:ll); “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:46); “Benguela” [-12.58333, 13.41667] (Boulenger 1887:208; Bocage 1895a:46); “Baia dos Elefantes” [-13.23333, 12.73333] (Laurent 1947:8); “Mullet Bay” [-13.40000, 12.56667] (Laurent 1947:8); “Baia Farta” [-12.60000, 13.20000] (Lau¬ rent 1947:8); “Baia de St. Bras, pres Lobito” (Laurent 1947:8). Namibe: “Cahinde-Ongueira” [-15.48333, 13.36667] (Hellmich 1957b:53; Ceriaco et al. 2016a:57); “Desert de Mossamedes, 35 km au sud de la ville” [-15.49752, 12.20051] (Laurent 1964a:75; Ceriaco et al. 2016a:57); “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:46; Ceriaco et al. 2016a:57); “Iona National Park, 3.4 km south-west (by air) of Espinheira, vicinities of Lion Cave” [-16.81514, 12.33714] (Ceriaco et al. 2016a:34); “Iona National Park, car wreck 20 km south-west (by air) of Espinheira” [-16.93161, 12.24595] (Ceriaco et al. 2016a:34); “Namibe-Lubango road, road marker 59, 1.8 km (by road) of Caraculo, north side of the road” [-15.01647, 12.64267] (Ceriaco et al. 2016a:34); Pico Azevedo [-15.53400, 12.49197] (Ceriaco et al. 2016a:34). Undetermined Locality: “Carangigo” (Boulenger 1887:208; Bocage 1895a:46). Taxonomic and distributional notes: Previously placed in the genus Mabuya, typical skinks from the Afro-Malagasy region were assigned to Trachylepis Fitzinger, 1843 by Bauer (2003). The species appears to vary little morphologically across its extensive range. Additional possible syntypes include MNHN 1471 (Brygoo 1985), BMNH 67.6.14.2-3, 1935.2.8.9, 1946.8.19.19 (Boulenger 1887), and MCZ R-21037-38. (Barbour and Loveridge 1929). As discussed by Bauer et al. (2003), the type series for this species was large, and it is likely that there are numerous spec¬ imens outside Berlin that may have syntype status. Map 214. Distribution of Trachylepis acutilabris Angola. Trachylepis affinis (Gray, 1838) Senegal Skink Tiliqua affinis Gray 1838:289. Holotype: BMNH 1946.8.18.21 (formerly BMNH xiv.92a) (collector unknown). Type locality: Not stated. Euprepes Blandingii: Bocage (1866a:44), Peters (1877a:614). Mabuia Raddonii: Bocage (1895a:40). Mabuia raddonii'. Boulenger (1887:165). 254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Mabuya radoni: Ferreira (1903:14), Parker (1936:138), Hellmieh (1957b:55). Mabuya affinis'. Hoogmoed (1974:6). Euprepis affinis'. Mausfeld-Lafdhiya et al. (2004:159). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is a widespread speeies eomplex distributed across all of western Africa, from Senegal to Angola. Ocurrences in Angola (Map 215): The species occurs in western Angola. However, the accepted distribution range for the species is limited to the northwestern Angola. Cabin¬ da: “Chinchoxo (cote d’Loango)” [-5.10000, 12.10000] (Peters 1877a:614; Bocage 1895a:40). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:55). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Ferreira 1903:14). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:138). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:40). Taxonomic and distributional notes: The long and complex history of Trachylepis [= Mabuya] affinis (Gray, 1838) has been reviewed by Hoogmoed (1974). At present there are numerous synonyms, of which two have been applied to Angolan material: Euprepis blandingii Hallowell, 1857 and Euprepis raddoni Gray, 1845. Molecular phylogenetic studies (Allen 2015) have revealed some significant structure across the distribution, but a thorough revision is needed in order to determine if any of the names in synonymy may be applicable to Angolan populations. The record from “Caconda,” Huila Province (Bocage 1895a), here plotted with an X, is certainly incorrect, although it is unclear to which other species it may apply. Map 215. Distribution of Trachylepis affinis in Angola. Trachylepis cf. albopunctata (Bocage, 1867) Angolan Variable Skink Euprepes Oliveiri var. albo-punctatus Bocage 1867b:223. Syntypes: MBL specimen numbers unknown (col¬ lector J. A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Benguella, and “Catumbella,” [= Benguela and Catumbela], Benguela Province, Angola. Euprepes angolensis (Bocage 1872:89). Holotype: MBL specimen numbers unknown (collector J. A. d’Anchieta [Biballa] and F. A. P. Bayao [Dondo]), destroyed by fire 18 March 1978. Type locality: “Biballa, dans I’interieur de Mossamedes” and “Dondo” [= Bibala and Dondo], Namibe and Kwanza Norte provinces, respectively, Angola. Euprepes Oliveiri'. Bocage (1867c:227). Euprepes angolensis'. Bocage (1872:78). Mabuia varia: Boulenger (1887:202, 1905:111), Bocage (1895a:43, 1896a:lll). Mabuya varia: Schmidt (1933:12), Monard (1937b:87), Mertens (1938a:437), Parker (1936:138), Laurent (1964a:72), Grillitsch et al. (1996:32), Branch (1998:157), Broadley (2000:99). Mabuya varia varia: Hellmieh (1957a:66, 1957b:57), Loveridge (1957: 212). Traehylepis varia: Ceriaco et al. (2016a:31, 58), Ceriaco et al. (2016b:67), Conradie et al. (2016b:26). Traehylepis varia Clade B: Weinell and Bauer (2018:107). Global conservation status (lUCN): Not Evaluated. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 255 Global distribution: This taxon occupies much of Angola except the southeast, as well as western Zambia and the former Katanga Provinee of the Democratic Republic of Congo. The extent of its extralimital distribution is not known. Ocurrences in Angola (Map 216): The species is widely distributed in Angola, except in the southeast, where it is replaced by T damarana. Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Bocage 1895a;43); “Dondo” [-9.68333, 14.43333] (Bocage 1872:78, 1895a;43; Hellmich 1957b;57); unnamed localities [-10.223, 14.811; -10.839, 14.629; -11.901, 15.45] (Weinell and Bauer 20 18: Supplementary Data 3). Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1905:111); “Cangandala National Park” [-9.81858, 16.65403] (Ceriaco et al. 2016b: 67; Weinell and Bauer 20 18: Supple¬ mentary Data 3); unnamed locality [-9.074, 16] (Weinell and Bauer 20 18: Supplementary Data 3). Lunda Norte: “Capaia” [-8.33333, 20.20000] (Grillitsch et al. 1996:32). Lunda Sul: “Alto Chieapa” [-10.88333, 19.23333] (Laurent 1964a:72). Moxico: “Sandando, 85 km east from Luso” [-11.61667, 20.63333] (Laurent 1964a:72). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:12); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:12). Huambo: “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:138); unnamed locality [-12.503, 15.225] (Weinell and Bauer 2018:Supplementary Data 3). Benguela: “Lobito” [-12.35000, 13.55000] (Parker 1936:138); “Cahata” [-12.35000, 14.81667] (Bocage 1895a:43); “Catumbela” [-12.43333, 13.55000] (Bocage 1867c:223; Loveridge 1957:212); “Quis- sange” [-12.43333, 14.05000] (Bocage 1895a:43); “Benguela” [-12.58333, 13.41667] (Bocage 18676:223; Loveridge 1957:212); “Ebanga” [-12.73333, 14.73333] (Monard 1397b:87); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:66); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:437); “Hanha” [-13.30000, 14.20000] (Bocage 1896:111); unnamed loealities [-8.345, 20.212; -8.346, 20.212; -12.163, 13.877; -12.252, 14.266; -11.901, 15.45] (Weinell and Bauer 20 18: Supplementary Data 3). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:43); “Sangeve” [-13.88333, 15.83333] (Monard 1937b:87); unnamed localities [-15.317, 13.533; -14.741, 15.058; -14.94, 13.512; -14.184, 14.428; -14.638, 13.928; -14.769, 14.808; -13.65, 21.426; -13.786, 14.643] (Weinell and Bauer 20 18: Supplementary Data 3). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1872:78; Ceriaco et al. 2016a:58); “Mossamedes” [-15.20000, 12.15000] (Bocage 1867c:227; Ceriaco et al. 2016a:58); “Leba Pass” [-15.07003, 13.24339] (Ceriaco et al. 2016a:31); unnamed locality [-14.733, 13.345] (Weinell and Bauer 2018:Supple- mentary Data 3). Cuando Cubango: “Cubango basin (46)” [-14.58981, 16.90739] (Conradie et al. 2016:9, 12, 25); “Cubango basin (47)” [-14.70214, 17.37806] (Conradie et al. 2016:9, 12, 25). Undetermined locality: “Between Benguela and Bihe” (Boulenger 1905:111). Taxonomic and distributional notes: Although considered as Trachylepis varia by most authors who have cited these skinks for Angola, a recent molecular and morphological review (Weinell and Bauer 2018) demonstrated that true T. varia is absent from Angola, with the Angolan ""varia'" belonging to a separate clade that extends from Angola to Zambia. As noted by Weinell and Map 216. Distribution of Trachylepis cf albopunctata in Angola. 256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Bauer (2018) it is likely that the names Euprepes olivieri var. albo-punctatus Bocage, 1867 and Euprepes angolensis Bocage, 1872 apply to this clade. Boulenger (1887) and Bocage (1895a, 1896a) incuded both in the synonymy of varia. The original description of albopunctata is consis¬ tent with the morphological characters of members of the Angolan clade, and given its precedence, we tentatively recognize it as the name applicable to varia-XikQ Trachylepis in Angola, exclusive of the southeast. Euprepis angolensis, which we consider as a probable junior synonym of T. albop¬ unctata has been considered by some to refer to the same species as Mabuia striata angolensis Monard, 1937 (see Trachylepis monardi account). Unfortunately, all of the original Bocage speci¬ mens associated with this name were destroyed, exacerbating the task of trying to link this name with particular specimens in the T. varia complex. It is likely that some of the older literature records may refer to other species of Trachylepis outside the T varia complex, so our assessment of the distribution of this group in Angola should be interpreted as a rough estimate at best. A review of the Angolan Trachylepis being prepared to resolve this and other similar problems. Trachylepis bayonii (Bocage, 1872) Bayao’s Skink Euprepes Bayonii (Bocage 1872:75). Syntypes: MBL specimen numbers unknown, BMNH 1946.8.19.13 (for¬ merly BMNH 66.6.11.8), ZMB 6477 (collector F.A.P. Bayao [var. A - Duque de Braganga] and J.A. d’Anchieta [var. B - Huila]), destroyed by fire 18 March 1978. Type locality: “Duque de Bragan^a dans I’interieur d’ Angola” [= Calandula], Malanje Province and “Huilla, plateau assez eleve dans I’interieur de Mossamedes” [= Huila], Huila Provincce, Angola. Mabuya bayonii huilensis (Laurent 1964a:67). Holotype: MD 1886 (collector A. Barros Machado). Type loca¬ lity: “Boca de Humpata, environs de Sa da Bandeira, Huila” [= Humpata], Huila Province, Angola. Euprepes Gravenhorstii: Bocage (1866a:44) Euprepes Bayonii (Bocage 1870:68, 1879b:95, 1887a:179). Euprepes Bayonii var. A\ Bocage (1872:75). Euprepes Bayonii var. B\ Bocage (1872:75). Mabuia bayonii Boulenger (1887:201) Mabuia Bayonii Bocage (1895:38). Mabuya bayonii: Boulenger (1887:201, 1905:111), Bocage (1895a:38, 1897a:195), Schmidt (1933:11), Monard (1937b:87), Hellmich (1957b:54), Managas (1963:234), Spawls et al (2004:130), Bauer et al (2003:270). Mabuya bayonii bayonii: Laurent (1964a:67). Euprepes Bayonii: Bauer et al. (2003:270). Traehylepis bayonii: Menegon and Spawls (2011), Ceriaco et al (2016b:57). Global conservation status (lUCN): Data Deficient. Global distribution: The species has been reported from Angola, the Democratic Republic of Congo, and Kenya and Tanzania, although the East African forms are referable to T b. keniensis (Loveridge, 1956) and are almost certainly specifically distinct from the West African T b. bayonii and T b. huilensis. Ocurrences in Angola (Map 217): The species mainly occurs in central Angola and south¬ western Angola. Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a: 179, 1895a;38). Kwanza Norte; “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:54); “Ambaca” [-9.26667, 15.18333] (Boulenger 1905:111). Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a:44, 1872:75, 1895a;38, 1897a;195; Boulenger 1887:201, 1905:111; Bauer et al 2003:270). Lunda Norte: “Cassange” [-9.58333, 17.86667] (Bocage 1879b:95, 1895a;38; 1897a:195). Moxico; “environs du lac Calundo” [-11.80000, 20.86667] (Manaqas 1963:263); “Calombe, Luso” [-11.83333, 19.93333] (Manaqas 1963:263). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:11; Laurent 1964a:67); “Chitau” [-11.43333, 17.15000] (Schmidt MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 257 1933:11). Huambo: “Santo-Amaro” [-12.70000, 15.85000] (Monard I937b:87). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage I895a;38). Hulla; “Caconda” [-13.73333, 15.06667] (Bocage I895a:38); “Kalukembe” [-13.78333, 14.68333] (Monard I937b:87); “Kuvangu” [-14.46667, 16.30000] (Monard I937b:87); “Boca de Humpata, Sa da Bandeira” [-14.93333, 13.51667] (Laurent I964a;67); “Huilla” [-15.05000, 13.55000] (Bocage 1872:75, I895a:38). Taxonomic and distributional notes: Bocage (1872) distinguished between two vari¬ eties: var. A, which corresponds to specimens with a uniform color collected at “Duque de Braganga” [= Calandula] and var. B from the “Huilla Plateau.” He later (Bocage 1895a) sep¬ arated the uniform specimens from those from throughout the rest of the species range. Laurent (1964a) described bayonii huilensis from “Boca de Humpata, environs de Sa da Bandeira.” This subspecies, based on a single juvenile spec¬ imen, has never been critically assessed, and although it may warrant recognition, we have here lumped all literature records to both it and the nominotypical form pending further research. The record from “S. Salvador do Congo” is out of the expected species range and probably represents a misidentification. Trachylepis binotata (Bocage, 1867) Ovambo Tree Skink Euprepes binotatus Bocage 1867b:223; 1867d:230. Syntypes: MBL specimen numbers unknown, destroyed by fire 18 March 1978, MNHN 1462 fide Brygoo (1985), BMNH 1946.8.15.32 (formerly BMNH 67.7.23.26) fiide Boulenger (1887), ZMB 5830 fide Bauer et al. (2003) (collector J.A. d’Anchieta). Type locality: “Benguella”, “Dombe” and “Catumbella” Benguela and Huila provinces, Angola fide Bocage (1867b:223). Euprepes binotatus: Bocage (1879c:88), Brygoo (1985:13). Mabuya quinquetaeniata: Boulenger (1887:198). Mabuya binotata: Bocage (1895a:46, 1897a: 196), Monard (1937b:91), Hellmich (1957a:59, 1957b:54), Lau¬ rent (1964a:68), Branch (1998:151), Bauer et al. (2003:270). Euprepes binotata: Bauer et al. (2003:270). Mabuya quinquetaeniata binotata: Mertens (1926:152, 1937a: 10, 1938a:437). Traehylepis binotata: Ceriaco et al. (2016a:57). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from southern Angola and northwestern Namibia. Ocurrences in Angola (Map 218): The speeies oeeurs in the southwestern Angola. Benguela: “Catumbella” [-12.35000, 14.81667] (Boeage 1867b:223, 1895a:46, 1897a:196; Bauer et al. 2003:27); “Benguella” [-12.58333, 13.41667] (Boeage 1867b:223, 1867d:230, 1895a:46, 1897a:196; Boulenger 1887:198; Brygoo 1985:13; Bauer et al. 2003:271); “Dombe” [-12.95000, 13.10000] (Boeage 1867b:223, 1895a:46, 1897a:196; Bauer et al. 2003:271); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:437); “Catengue Station, Benguella” [-13.03333, 13.73333] (Mertens 1926:152, 1937a:10). Huila: “Caeonda” [-12.43333, 13.55000] (Boeage 1879e:88). Namibe: “Capangombe” [-15.10000, 13.15000] (Boeage 1895a:46; Ceriaeo et al. 2016a:57); “50 km Mo^a- ll^E ,24'’ E Map 217. Distribution of Trachylepis bayonii in Angola. 258 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II medes road to Sa da Bandeira” [-15.01358, 12.518628] (Laurent I964a;68; Ceriaeo et al. 20I6a:57); “Maconjo” [-15.01667, 13.20000] (Bauer et al. 2003:271; Ceriaeo et al. 2016a:57). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:91). Taxonomic and distributional notes: Bocage (1867b) first named Euprepes binota- tus, from three localities in Benguela Province, but without a description or diagnosis. In the subsequent paper, Bocage (1867d) published a detailed description but noted only the locality “Benguella” by name. The original type series was large and syntypes were sent to Bocage’s correspondents at other major European muse¬ ums. Boulenger (1887) erroneously listed the BMNH syntype under Mabuya quinquetaenia- ta. Trachylepis bocagii (Boulenger, 1887) Bocage’s Skink Euprepes Petersi: Bocage 1872:74. Syntypes (see notes below): MBL speeimen numbers unknown (eolleetor F.A.P. Bayao), destroyed by fire 18 Mareh 1978. Type loeality: “Duque de Bragan^a” [= Calandula] Malanje Provinee, Angola. Mabuia boeagii (Boulenger 1887:203). Replaeement name for Euprepes petersi Boeage, 1872, preoeeupied by E. petersi Steindaehner, 1867 = Eutropis dissimilis (Hallowell, 1857). Euprepes quinquetaeniatus: Boeage (1866a:44). Mabuia Petersi'. Boeage (1895a:42, 1897a:197), Ferreira (1900a:49, 1903:15, 1906:170), Tiedemann and Haupl (1980:43), Tiedemann et al. (1994:53). Mabuia quinquetaniata: Boulenger (1905:111). Mabuya boeagii'. Parker (1936:136), Mertens (1938a:437), Bauer et al. (2003:275). Mabuya boeagii boeagii'. Hellmieh (1957a:60). Mabuya quinquetaeniata quinquetaeniata'. Hellmieh (1957b:54). Mabuya boeagei: Frade (1963:252). Euprepes quinquetaeniata'. Brygoo (1985:90). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Africa, occuring from Angola, through Zambia to Zimbabwe and Malawi. Ocurrences in Angola (Map 219): The species has been recorded at widely scattered sites in Angola, but is absent form the arid far southwest and the forested areas of the northwest and north¬ east. Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:44, 1872:74, 1895a:42; Boulenger 1887:203; Brygoo 1985:90; Bauer et al. 2003:275); “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1887:203, 1905:111; Bocage 1895a:42). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:54); “Cambondo” [-9.15963, 14.65771] (Ferreira 1906:170); “Dondo” [-9.68333, 14.43333] (Bocage 1972:80, 1895a:42, Tiedemann and Haupl 1980:43; Tiedemann et al. 1994:53). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:136). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:42, 1897a:195); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:60); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:437). Without precise locality: “with no precise location” (Ferreira 1900a:49). Il-E 20°E ,24'’ E Map 218. Distribution of Trachylepis binotata in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 259 Taxonomic and distributional notes: Euprepes petersi Bocage, 1872 was preoccu¬ pied by E. petersii Steindachner, 1867 (syn. Eutropis dissimilis (Hallowell, 1857)) and Mabuia bocagii was proposed as a replacement name by Boulenger (1887). As such its type material is identical to that of Bocage (1872). Bocage (1872) applied his name to the species he originally called Euprepes quinquetaeniata (Bocage 1866a). His description of E. petersi gives details for only a single specimen but his earlier mention of E. quinquetaeniata from “Duque de Braganga” reveals that there were originally many specimens in the series in Lis¬ bon. Several of these have been suggested to be potential syntypes of E. petersi, including ZMB 6479 in Berlin (Bauer et al. 2003) BMNH 1946.8.15.30 (formerly 64.7.13.23) in Map 219. Distribution of Trachylepis bocagii in Angola. London, MNHN 1286 and 1286"* in Paris (Bygoo 1985), and NHMW 16677 in Vienna (Tiedemann and Haupl 1980; Tiedemann et al. 1994). The last of these, however, is from “Dondo” and cannot be part of the type series. Those from London and Paris are from the type locality and are almost certainly part of the series of E. quinquetaeniata noted by Bocage (1866a). However, they were received from Lisbon several years before the description of E. petersi and were not among those on hand in Lisbon when Bocage prepared the description of E. petersi. The date of receipt of the Berlin specimen is unknown, but it is likely that it too significantly predated the description of E. petersi. Thus, conservatively, any types of this taxon are likely to have been in Lisbon only, although those specimens in other collections nonetheless provide valuable confirmation of Bocage’s concept of the species. There are several literature records of T. quinquetaeniata from Angola, all of which are highly unlikely. This species has a broad distribution across the Sahel, in the Nile Valley and in East Africa, but the nearest records to Angola are in the northeastern Democratic Republic of Congo and even the closely related T. pulcherrima (de Witte, 1953) extends no closer than -1000 km to the nearest purported Angolan sites (Broadley and Bauer 1998). This is borne out by the examination of the specimens upon which these errant records are based. Thus, Malanje and Kwanza Norte references to T. quinquetaeniata (Boulenger 1905; Hellmich 1957b) are here referred to T. bocagii. Trachylepis bocagii has also recently been collected in Qui^ama National Park, Bengo Province (M. Marques and L. Ceriaco, pers. obs.). Trachylepis chimbana (Boulenger, 1887) Chimba Skink Euprepes affinis Bocage 1872:77. Syntypes: MBL 822 (2 speeimens), 824 (eolleetor not stated). Type loeali- ty: “Rio Chimba, dans I’interieur de Mossamedes” [= Rio Chimba], Namibe Provinee, Angola. Mabuia ehimbana Boulenger 1887:204. Replaeement name for Euprepes affinis Boeage, 1872, preoeeupied hy E. affinis (Gray, 1838). Euprepes affinis'. Boeage (1872:77). Mabuia ehimbana'. Boeage (1895a:45, 1897a: 195), Sehmidt (1933:12), Frade (1963:252). Mabuya striata angolensis: Hellmieh (1957b:56). Mabuya striata ehimbana'. Laurent (1964a:69). Mabuya ehimbana'. Broadley (1975a: 13), Braneh (1998:153). Traehylepis ehimbana'. Portik and Bauer (2012:128), Ceriaeo et al. (2016a:57). 260 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and extreme northern Nambia. Ocurrences in Angola (Map 220): The species occurs in southwest and eastern Ango¬ la. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:69); “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a;69) [in error]. Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:69); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a;69); “Calunda” [-12.11667, 23.46667] (Laurent 1964a:69). Huambo: “Bela-Vista (Sanguengue)” [-12.36667, 16.20000] (Hell- mich 1957b:56). Benguela: “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a;45, Distribution of Trachylepis chimbana in 1897a:195; Broadley 1975a:14); “10 km E of Caembombo” [-13.01667, 14.01667] (Broadley 1975a:14); “Coporolo” [-13.38333, 13.88333] (Broadley 1975a: 14); “5 km S of Catara River on Lucira Road” [-13.55000, 12.55000] (Broadley 1975a:14). Huila: “Jau, around Sa da Bandeira” [-15.20000, 13.51667] (Laurent 1964a:69). Nami- be: “Lucira” [-13.86667, 12.53333] (Broadley 1975a:14; Ceriaco et al. 2016a:57); “Rio Chimba” [-14.30000, 12.40000] (Bocage 1872:77, 1895a:45, 1897a:195; Boulenger 1887:204; Broadley 1975a: 14; Ceriaco et al. 2016a:57); “Chapeau Armando turnoff, Mossamedes” [-14.45000, 12.36667] (Broadley 1975a:14; Ceriaco et al. 2016a:57); “Assuncao” [-14.86667, 13.10000] (Broadley 1975a:14; Ceriaco et al. 2016a:57); “14 kmNE Caraculo” [-14.90214, 12.74229] (Broa¬ dley 1975a: 14; Ceriaco et al. 2016a:57); “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:45, 1897a:195; Broadley 1975a:14; Ceriaco et al. 2016a:57); “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:45, 1897a:195; Broadley 1975a:14; Ceriaco et al. 2016a:57); “Mossamedes” [-15.20000, 12.15000] (Boulenger 1887:204; Ceriaco et al. 2016a:57); “Saiona River, NW of Cain- de” [-15.40000, 13.20000] (Broadley 1975:14; Ceriaco et al. 2016a:57); “Cainde” [-15.48333, 13.36667] (Broadley 1975a:14; Ceriaco et al. 2016a:57); “Rio Coroca” [-15.78333, 12.06667] (Broadley 1975a: 14; Ceriaco et al. 2016a:57). Cunene: “Othinjau” [-17.33333, 14.26667] (Hellmich 1957b:56); “Humbe” [-16.68333, 14.90000] (Schmidt 1933:12). Undetermined locali¬ ty: “12 km W of Humbia” (Broadley 1975a:14); “Huke” (Broadley 1975a:14). Taxonomic and distributional notes: Boulenger (1887) provided E. chimbana as a replace¬ ment name for Bocage’s (1872) Euprepes affinis which was preoccupied. Laurent (1964a) treated M. chimbana as a subspecies of Mabuya striata but Broadley (1975a) treated it as a member of the Trachylepis lacertiformis complex. It seems likely that Laurent’s concept of this species was different than that of other authors and that his localities, far to the east of what is usually regard¬ ed as a western species (Broadley 1975a; Branch 1998; Portik and Bauer 2012), apply to another congener. These specimens have recently been rediscovered and are under study. Trachylepis damarana (Peters, 1870) Kalahari Variable Skink Euprepes damaranus Peters 1870:20. Leetotype: ZMB 6153 (eolleetor W. C. H. Peters). Type loeality: “Damaraland,” [= north eentral Namibia], Namibia. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 261 Trachylepis varia: Conradie et al. (2016b: 26). Trachylepis damarana: Weinell and Bauer (2018:215). Global conservation status (lUCN): Not Evaluated. Global distribution: T. damarana is distributed from southeastern Angola and northeastern Namibia, aeross northern Botswana to Zimbabwe and adjaeent western areas of Mozambique and the northeastern South Afriea. Ocurrences in Angola (Map 221); The species occurs only in the southeastern areas of the country. Cuando Cubango: “Cuito basin (30c)” [-17.51222, 20.06027] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 2018:Sup- plementary Data 3); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie etal. 2016:9- 10, 25; Weinell and Bauer 20 18: Supplementary Data 3); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 20 18: Supplementary Data 3); “Cuando basin (37)” [-17.67833, 22.61475] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 2018:Supplementary Data 3); “Cuando basin (38)” [-17.58830, 22.65694] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 2018;Supplementary Data 3); “Cuando basin (39)” [-17.46333, 22.86638] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 2018:Sup- plementary Data 3); “Cuando basin (42)” [-17.49611, 23.13444] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 20 18: Supplementary Data 3); “Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 25; Weinell and Bauer 20 18: Supplementary Data 3); “Cubango basin (59)” [-16.28392, 18.84744] (Conradie et al. 2016:9, 12, 25; Weinell and Bauer 201 8: Supplemen¬ tary Data 3). Taxonomic and distributional notes: Originally described by Peters (1870) from “Damara- land,” then applicable to much of northern Namibia, this species was recently resurrected from the synonymy of Trachylepis varia. Broadley (1966, 2000) considered T. damarana as a synonym of T. varia, but recently Weinell and Bauer (2018) provided molecular and morphological evidence that supports its validity as a distinct species. Map 221. Distribution of Trachylepis damarana in Angola. 12°E I6"E 20-E 24“ E Trachylepis hoeschi (Mertens, 1954) Hoesch’s Skink Mabuya hoeschi Mertens 1954:178, fig. 4. Holotype: SMF 45681 (eolleetor W. Hoeseh). Type loeality: “Roessing-Berge, ostlieh von Swakopmund, SW-Afrika” [= Rossing Mts., east of Swakopmund, Erongo Region], Namibia. Mabuya hoeschi: Laurent (1964a:68), Braneh (1998:153). Trachylepis hoeschi: Ceriaeo et al. (2016b:32, 57). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is restricted to southwestern Africa, occuring from north¬ western Namibia to southwestern Angola. Ocurrences in Angola (Map 222): The species is limited to the Namibe desert region. Namibe: “Plage «das Conchas», Mogamedes” [-15.13333, 12.11667] (Laurent 1964a;68; Ceriaeo 262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II et al. 20I6a:57); “Rio Curoca in the Pediva Hot Springs area” [-16.28359, 12.56106] (Ceriaco et al. 2016a:32); “Namibe Regional-naNatural Park” [-15.77386, 12.33306] (Ceriaco et al. 2016a:32). Taxonomic and distributional notes: None. Il-E 1&°E 20°E 24”E Map 222. Distribution of Trachylepis hoeschi in Angola. Trachylepis cf. lacertiformis (Peters, 1854) Bronze Rock Skink Euprepes lacertiformis Peters 1854:618. Syntypes: ZMB 5523 (4 speeimens) (eolleetor W.C.H. Peters), lost fide Bauer et al. (2003). Type loeality: “Boror” [Mozambique], eonsidered in error for “Tete,” Mozam¬ bique fide Broadley (1975a). Mabuya laeertiformis: Broadley (1975a:12, 2000:101), Braneh (1998:154). Traehylepis laeertiformis'. Broadley (2004:324), Broadley and Maritz (2010b), Portik and Bauer (2012:128), Ceriaeo et al. (20 16a: 5 7). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs from the southern end of Malawi Lake through the Tete District of Mozambique into eastern Zimbabwe and westwards along the Zambezi. The Angolan population is disjunct. Ocurrences in Angola (Map 223): The species occurs in an isolated population in southwestern Angola in Nambie and Cunene Province, near the Namibian border. Namibe: “Cainde, Mossamedes” [-15.48333, 13.36667] (Broadley 1975a: 12; Ceriaco et al. 2016a:57); “14 km NE of Caraculo, Mossamedes” [-14.90214, 12.74229] (Broadley 1975a: 12; Ceriaco et al. 2016a:57). Cunene: “Otchinjau, Huila” [-16.57903, 13.90516] (Broadley 1975a: 12). Taxonomic and distributional notes: Tra¬ chylepis lacertiformis has frequently been con¬ fused with other species, particularly Tra¬ chylepis varia (Peters, 1867) (Broadley 1975). The status of the Angolan population with respect to supposed conspecifics in southeast Africa requires further investigation. I2''E 2«°E irE Map 223. Distribution of Trachylepis cf lacertiformis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 263 Trachylepis laevis (Boulenger, 1907) Angolan Blue-Tailed Skink Mabuia laevis Boulenger 1907a:212. Holotype: BMNH 1946.8.15.31 (formerly BMNH 06.8.24.71) (eollee- tor W.J. Ansorge). Type loeality: “Maeonjo, Benguela” [= Fazenda Mueungo], Namibe Provinee, Angola. Mabuya laevis: Hellmieh (1957b:54), Laurent (1964a:76), Braneh (1998:155). Traehylepis laevis: Cerlaeo et al. (2016a:33, 57). Global conservation status (lUCN): Not Evaluated. Global distribution: This species inhabits in large rock cracks in granite outcrops, from the Kaokoveld of northern Namibia and Southern Angola. Ocurrences in Angola (Map 224): The species occurs in southern Angola, in desert areas. Kwanza Norte: “Piri-Dembos (record questionable)” [-8.56667, 14.50000] (Hellmieh 1957b:54). Namibe: “Munhino 50 km west Sa da Bandeira” [-14.96667, 12.96667] (Laurent 1964a:76; Ceriaco et al. 2016a:57); “Maeonjo (= Fazenda Mueungo)” [-14.782192, 12.486557] (Boulenger 1907a:212; Ceriaco et al. 2016a: 57); “Iona National Park, north of Tambor” [-15.99642, 12.40711] (Ceriaco et al. 2016a:33). Taxonomic and distributional notes: Hellmieh (1957b) cited this species from “Piri- Dembos,” which is significantly further north than the type locality. Laurent (1964a) cited another specimen from Namibe Province and also expressed doubt about the validty of Hellmich’s specimen. The Map 224. Distribution of Trachylepis laevis in Angola. Piri-Dembos” record is highly improbable and surely represents a misidentification, although it is not clear to which species it might actually belong. Trachylepis maculilabris (Gray, 1845) Speckled-Lipped Skink Euprepes maeulilabris Gray 1845:114. Holotype: BMNH 1946.8.18.17 (formerly BMNH xv.l09a) (eolleetor W. Raddon). Type loeality: “W. Afriea” [= West Afriea]. Euprepes notabilis Peters 1879:36. Syntypes: ZMB 9204 (eolleetor A. von. Homeyer), 8485 (eolleetor J.G. Falkenstein) lost fide Bauer et al. (1995, 2003). Type loeality: “Chinehoxo,” [Cabinda Provinee] (ZMB 8485) and “Pungo Andongo,” [Malanje Provinee, Angola] (ZMB 9204). Euprepes Anehietae Boeage 1886a:44 {nomen nudum), 1866b:62. Holotype: MBL speeimen number unknown, (eolleetor J.A. d’Anehieta) destroyed by fire 18 Mareh 1978. Type loeality: “Zaire” [= Zaire Provinee], Angola. Euprepes (Eupr) Perrotetii: Peters (1877a:614), Boeage (1895a:39). Mabuia maeulilabris: Boulenger (1887:164), Boeage (1895a:40), Ferreira (1906:170). Mabuya maeulilabris: Loveridge (1933:312), Parker (1936:138), Laurent (1950a: 12, 1954a:65), Broadley (2000:94), Bauer et al. (2003:275). Mabuya maeulilabris maeulilabris: Hellmieh (1957a:61), Loveridge (1957:209), Laurent (1964a:65). Euprepes notabilis: Bauer et al. (2003:275). Euprepes maeulialbris: Mausfeld et al. (2004:160). Traehylepis maeulilabris: Trape et al. (2012:398), Ceriaeo et al. (2014b:671), Braneh and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global distribution: This species has a wide distribution in sub-Saharan Africa from Liberia to the Horn of Africa southwards as far as Angola in the west and Mozambique in the east. It is a human commensal and may expand its range as habitats are modified. Ocurrences in Angola (Map 225): The species is widely distributed in the northern regions of the country and is represented by scattered records in the south. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614, 1879:36; Bocage 1895a:39; Love- ridge 1957:209; Bauer et al. 2003:275); “Cabinda” [-5.55000, 12.18333] (Bocage 1895a:40); 1957:209 Bengo: “Ambriz” [-7.844312, 13.106493] (Boulenger 1887:164; Bocage 1895a:40; Loveridge 1933:312). Luanda: “Quifangondo” [-8.76667, 13.43333] (Ceriaco et al. 2014b:671). Kwanza Norte: “Golungo” [-9.13333, 14.76667] (Ferreira 1906:170); “Cambondo” [-9.15963, 14.65771] (Ferreira: 1906: 170; Loveridge 1947:179); “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a:61); “Dondo (Quanza edges)” [-9.68333, 14.43333] (Bocage 1895a:40). Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Peters 1879:36; Bocage 1895a:39, 40; Love¬ ridge 1957:209; Bauer et al. (2003:275); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014:671). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12, 1954a:65, 1964a: 65); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a: 12); “Carumbo, Lucapa” [-8.42278, 20.73917] (Branch and Conradie 2000:200). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:65). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:138); “Congulu” [-10.86667, 14.28333] (Parker 1936:138). Benguela: “Entre Rios, Chiviti- di” [-13.01667, 14.63333] (Hellmich 1957a:61). Taxonomic and distributional notes: Mausfeld et al. (2004) noted that Trachylepis mac- ulilabris (Gray, 1 845) may comprise at least two distinct species, one from West Africa represent¬ ing the nominotypic maculilabris, and other from East Africa representing a cryptic species. Ongo¬ ing research suggests that this is an underestimate and that the complex may contain many species. Bocage (1866a) introduced the name Euprepes anchietae as a nomen nudum, but a valid descrip¬ tion appeared in a second paper (Bocage 1866b) in the the same issue of the journal. Peters (1877a) originally referred one of the types of E. notabilis to E. perottetii Dumeril and Bibron, 1839 and although Peters (1879) revised his identification, this correction was not noted by Bocage (1895a). Map 225. Distribution of Trachylepis maculilabris in Angola. Trachylepis cf. megalura (Peters, 1878) Grass-Top Skink Euprepes (Mabuia) megalurus Peters 1878:204, pi. 2, fig. 4. Syntypes: ZMB 9281 (2 speeimens), 9304 (eol- leetor J. M. Hildebrandt). Type loeality: “Taita” [= Kenya]. Mabuya megalura subsp.: Laurent (1964a:74). Traehylepis ef megalura: Ceriaeo et al. (20 16b: 71). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Ethiopia south to Mozambique and west to the southern Democratic Republic of Congo and northern Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 265 Ocurrences in Angola (Map 226): The species has only been recorded from northeast¬ ern Angola. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:74). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco etal. 2016b:71). Taxonomic and distributional notes: According to Laurent (1964a) the two speci¬ mens from “Alto Cuilo” differ from the speci¬ mens of Mabuya megalura (Peters, 1878) {=Trachylepis megalura) from East Africa and around Lake Kivu and probably represent an undescribed “angolo-katangaise” variety or new species. Recently collected specimens from Cangandala National Park, Malanje Province (Ceriaco et al. 2016b) appear to be conspecific with Laurent’s material and further studies are now in progress. Il-E 1&“E 20“E 2*°E Map 226. Distribution of Trachylepis cf. megalura in Angola. Trachylepis monardi nom. nov. Marques, Ceriaco, Blackburn and Bauer Monard’s Skink (Endemic) Mabuia striata angolensis Monard 1937b:89. Syntypes: MHNC 91.0599-0613 [15 specimens], MHNG 858.95 (collector A. Monard). Type locality: “Kuvangu” [= Cuvango], Huila Province and “Bimbi” [=Bimbe] Huambo Province, Angola. Mabuya striata angolensis: Mahnert (1976:488), Ortiz (1989:56). Mabuya angolensis: Laurent (1964a:72). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known only from Angola. Ocurrences in Angola (Map 227): The species occurs mainly in the central-west Angola. Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:89; Mahnert 1976:488; Ortiz 1989:56). Bie: “Silva Porto” [-12.33333, 16.86667] (Laurent 1964a:72). Huila: “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:89; Mahnert 1976:488; Ortiz 1989:56). Taxonomic and distributional notes: Monard (1937b) described Mabuia striata angolensis perhaps unaware of Bocage’s (1872) Euprepis angolensis, a name that has been largely neglected or sometimes subse¬ quently synonymized with Mabuya striata angolensis, however, its identity remains uncertain. Both Boulenger (1887) and Bocage (1895a) treated it as a synonym of Euprepes varia Peters, 1867, and we include the relevant ll-E 20°E 24“ E Map 227. Distribution of Trachylepis monardi in Angola. 266 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II material under that account. As both names are currently placed in the same genus Monard’s name is a junior secondary homonym of Bocage’s. As such Bocage’s name, despite being a junior sub¬ jective synonym of Trachylepis varia, must retain the epithet angolensis, and Monard’s name requires a replacement (the special conditions of Article 23.9 do not apply as an insufficient number of citations by an insufficient number of authors have used Monard’s name). As no junior synonyms are known to exist for Trachylepis angolensis (Monard, 1937) we propose Trachylepis monardi nom. nov. as a replacement name, in recognition of the contributions of Albert Monard (2 September 1886 - 27 September 1952), Swiss naturalist from La Chaux-de-Fonds, who under¬ took two major expeditions to Angola and left a legacy of specimens and published papers relating to the zoology of not only Angola, but also Cameroon, Guinea-Bissau and North Africa. Monard (1937b) noted 26 syntypes in the type series. Fifteen of these remain in La Chaux- de-Fonds. Ortiz (1989) cited MHNN (Museum d’histoire naturelle de Neuchatel) 2130 as the lectotype, giving both localities “Kuvangu” and “Bimbi”. The MHNC specimens thus are paralec- totypes, as is the single Geneva “paratype” cited by Mahnert (1976) and Schatti and Ferret (1997), for which the locality is also uncertain. Monard typically dispersed specimens to the many Swiss institutions that helped sponsor his field work, including high schools. It is likely thjat the “miss¬ ing” nine specimens were or are in such small collections. Trachylepis occidentalis (Peters, 1867) Western Three-Striped Skink Euprepes vittatus var. australis Peters 1862a: 19. Syntypes: ZMB 4212, 64401-02 (formerly ZMB 4212A-B) (eolleetor C.H. Hahn). Type loeality: “Neu-Barmen” [= Gross Barmen, Otjozondjupa Region] Namibia by implieation fide Bauer et al. (1995, 2003). Euprepes oeeidentalis Peters 1867:20, replaeement name for E. v. australis Peters 1862, preoeeupied by Euprepes australis Gray, 1838 [= Ctenotus australis (Gray, 1838 “1839”)]. Euprepes oeeidentalis: Boeage (1870:68). Mabuia oeeidentalis: Boeage (1895a:42). Mabuya oeeidentalis: Laurent (1964a:73), Braneh (1998:156), Broadley (2000:97). Traehylepis oeeidentalis: Bates et al. (2014:263), Ceriaeo et al. (2016a:33, 57). Global conservation status (lUCN): Not Evaluated. Global distribution: Distributed from arid western South Africa and southwestern Botswana through Namibia to arid areas of southwestern Angola. Ocurrences in Angola (Map 228): The species distribution is limited to the desert regions in Namibe Province. Namibe: “Mossamedes desert, 35 km south from the city” [-15.50575, 12.20768] (Laurent 1964a: 73; Ceriaeo et al. 2016a;57); “Rio Coroca” [-15.78333, 12.06667] (Boeage 1895a:42; Ceriaeo et al. 2016a;57); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaeo et al. 2016a:33). Taxonomic and distributional notes: Bauer et al. (2003) also included ZMB 4213, MNHN 1468 (see Brygoo 1985), and BMNH I2"E 1&°E 20°E 24'E Map 228. Distribution of Trachylepis occidentalis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 267 67.6. 14. 1 among the putative syntypes, although they noted that only three specimens could be types as Peters was explicit about the number of specimens he had. The original ZMB catalogue notes in Peters’ hand both that ZMB 4212 corresponds to the types and that there were three spec¬ imens under this number ZMB 4212A and B reported by Bauer et al. (1995, 2003) have since been renumbered. Thus only ZMB specimens have status as types. Trachylepis punctulata (Bocage, 1872) Speckled Sand Skink Euprepes punctulatus Bocage 1872:76. Syntypes: MBL 742 (2 speeimens) (eolleetor J.A. d’Aneheita), destroyed by fire 18 Mareh 1978. Type loeality: “Rio Coroea, dans le littoral an sud de Mossamedes” [= Rio Coroea, Namibe Provinee, Angola]. Mabuia punctulata: Boulenger (1887:204), Boeage (1895a:44, 1897a: 195). Mabuya punctulata: Sehmidt (1933:12), Frade (1963:253), Broadley (2000:100). Mabuya longiloba longiloba: Laurent (1964a:73). Mabuya variegata punctulata: Broadley (1975a:7), Braneh (1998:158). Trachylepis punctulata: Portik (2009:136), Portik and Bauer (2012:128), Bates et al. (2014:264), Ceriaeo et al. (2016a:31, 57). Global conservation status (lUCN): Not Evaluated. Global distribution: Widely distributed across the Kalahari and adjacent regions of Southern Africa, with several extensions eastwards in central South Africa and Mozambique and reaching its northern extent in southwestern Angola with an islaoted record in western Zambia (Broadley 1975a; Branch 1998). Ocurrences in Angola (Map 229): The species occurs in southwestern Angola. Huila: “15 km north Quilengues” [-13.95256, 14.04704] (Laurent 1964a:73). Namibe: “Ina- mangando River on Lucira Road” [-14.05000, 12.38333] (Broadley 1975a:ll; Ceriaeo et al. 2016a:57); “Mucungu” [-14.78333, 12.48333] (Schmidt 1933:12; Broadley 1975:11; Ceriaeo et al. 2016a;57); “Sao Nicolau, Mossamedes” [-14.26667, 12.36667] (Broadley 1975a:ll; Ceriaeo et al. 2016a:57); “17 km N of Sao Nicolau” [-14.12714, 12.37577] (Broadley 1975a: 11; Ceriaeo et al. 2016a;57); “Caraculo” [-15.01667, 12.66667] (Broadley 1975a:ll; Ceriaeo et al. 2016a:57); “15 km W of Caracu¬ lo” [-15.02039, 12.50489] (Broadley 1975a:ll; Ceriaeo et al. 2016a;57); “Cima” [-15.06667, 12.15000] (Broadley 197a5:ll; Ceriaeo et al. 2016a:57); “Pico Acezevedo” [-15.55000, 12.51667] (Broadley 1975:11); “23 km W of Virei” [-15.66907, 12.74847] (Broadley 1975a:ll; Ceriaeo et al. 2016a:57); “Rio Coroea” [-15.78333, 12.06667] (Bocage 1872:76, 1895a:44, 1897a:195; Boulenger 1887:204; Broadley 1975a:ll, 2000:100; Ceriaeo et al. 2016a:57); “6 km S of Rio Coroea on Iona road” [-15.78333, 12.06667] (Broadley 1975a: 11, 2000:100; Ceriaeo et al. 2016a:57); “Porto Alexandre” [-15.80000, 11.83333] (Broadley 1975a:ll); “30 km N of Tambor” [-16.13556, 12.42972] (Broadley 1975:11; Ceriaeo et al. 2016a:57); “Octchinfengo River on Onocua road, Iona Reserve” [-16.61667, 12.88333] (Broadley 1975a:ll; Ceriaeo et al. 2016a:57); Map 229. Distribution of Trachylepis punctulata in Angola. 268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Foz de Cunene” [-17.28333, 11.80000] (Broadley 1975:11; Ceriaco et al 2016a:57); “Espinheira” [-16.78906, 12.35756] (Ceriaco et al 2016a:31); “Namibe-Lubango road, road marker 59, 1.8 km west (by road) of Caraculo, on the north side of the road” [-15.01531, 12.64244] (Ceriaco et al. 2016a;31); “Praia do Navio coastal dunes, ca 124 km SSW of Namibe” [-16.27233, 11.83164] (Ceriaco et al. 2016a:31). Undetermined Locality: “Mossamedes to Benguela” (Broadley 1975a:ll). Taxonomic and distributional notes: This species was long considered as a subspecies of T. variegata (Peters, 1870) following Broadley (1975a, 2000). Based on morphological differences outlined in these papers and a high level of molecular divergence, these two taxa are now treated as full species (Portik 2009; Portik and Bauer 2012; Bates et al. 2014). The monophyly of T. punctulata is relatively poorly supported and further revisionary work may support the recogni¬ tion of additional taxa within this taxon as currently construed (Portik and Bauer 2012). Broadley (1975a) remarked that populations of T. punctulata from the Northern Cape Province, Botswana, Rhodesia, Zambia and Mozambique differ in several morphological characters from populations in Angola and Namibia, and that the eastern populations may ultimately deserve taxonomic recogni¬ tion. The results obtained by Portik and Bauer (2012) partially support this division, although they found evidence for the Northern Cape Province sample clustering with samples of T. punctulata from Namibia (Portik and Bauer 2012, Bates et al. 2014). The presence of a record from Kalabo, Zambia (Broadley 1971, 1975a) close to the eastern border of Angola suggests that further field¬ work may eventually yield T. punctulata, or a superficially similar taxon, from Moxico or Cuando Cubango. Trachylepis spilogaster (Peters, 1882) Kalahari Tree Skink Euprepes (Euprepis) striatus var. spilogaster Peters 1882:68. Type: Not loeated fide Bauer et al (1995:63, 2003:307). Type loeality: “Bei Otjimbingue im Hererolande (Westafrika)” (Peters 1882:68), [= Otjimb- ingwe, Erongo Region], Namibia. Mabuya striata spilogaster. Laurent (1964a:71). Mabuya spilogaster. Braneh (1998:156). Mabuya spilogaster. Broadley (2000:105). Traehylepis spilogaster. Bates et al. (2014:265). Traehylepis ef spilogaster. Conradie et al. (2016:25). Global conservation status (lUCN): Not Evaluated. Global distribution: This species extends from the Kalahari of northwestern South Africa and southwestern Botswana, through the length of Namibia to southwest Angola. Ocurrences in Angola (Map 230): The only published record for this species is from “Serra do Moco” in Huambo Province, although museum specimens from southeastern provinces exist. These latter are treated separately as Trachylepis aff. spilogaster and marked with a triangle in the map. Trachylepis spilogaster (plotted as circles). Huambo: “Serra do Moco, Luimbale” [-12.53333, 15.18333] (Laurent 1964a:71). Trachylepis aff. spilogaster (plotted as triangles). Bie: “Cubango basin (12b)” [-13.59638, 16.87722] (Conradie et al. 2016:8-9, 25). Cuando Cubango: “Cubango basin (6a)” [-14.67155, 17.73525] (Conradie et al. 2016:25); “Cubango basin (20)” [-14.67175, 17.15331] (Conradie et al. 2016:8-9, 25); “Cuito basin (25)” [-15.13919, 19.14350] (Conradie et al. 2016:8-9, 25); “Cuito basin (30a)” [-16.90980, 19.30769] (Conradie et al. 2016:9-10, 25); “Cuito basin (35)” [-16.62322, 19.05352] (Conradie et al 2016:9-10, 25); “Cuito basin (32)” [-17.04880, 19.53333] (Conradie et al 2016:9-10, 25); “Cubango basin (49)” [-14.66300, 17.66550] (Conradie et al. 2016:9, 12, 25); “Cuito basin (57)” [-15.45969, 18.76833] (Conradie et al. 2016:9, 12, 25); “Cuito basin (62)” [-17.50875, 20.06608] (Conradie et al. 2016:9, MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 269 12, 25); “Cubango basin (47)” [-14.70214, 17.37806] (Conradie et al. 2016:25). Taxonomic and distributional notes: Previously considered as a subspecies of T. striata (Peters, 1844), Branch (1998), Broadley (2000) and all subsequent authors have considered this species specifically dis¬ tinct. Conradie et al. (2016) published numer¬ ous records of T. aff spilogaster from Cuando Cubango. We think it is highly unlikely that these records refer to the same lineage as true T. spilogaster. The true affinities of this popu¬ lation will require a comprehensive revision of the Trachylepis of Angola. I1“E 20°E 24“E Map 230. Distribution of Trachylepis spilogaster (circle) and T. aff spilogaster (triangles) in Angola. Trachylepis sulcata (Peters, 1867) Western Rock Skink Euprepes olivaceus Peters 1862a:21. Syntypes: ZMB 4209, 4210, 64329-30 (formerly ZMB 4210 part), 64346-47 (formerly ZMB 4209), MNHN 1470 (eolleetor C. H. Hahn). Type loeality: “Neu-Barmen” [= Gross Barmen, Otjozondjupa Region] Namibia. Euprepes suleatus Peters 1867a:20, replaeement name for E. olivaeeus Peters, 1862 whieh is preoeeupied by E. olivaeeus Gray, 1839 [= Dasia olivaeea (Gray, 1839)]. MabuiaAnsorgiiBovilQngQr 1907a:213. Holotype: BMNH 1946.8.3.33 (formerly BMNH 1906.8.24.62) (eol¬ leetor W.J. Ansorge). Type loeality: “Caeonda, Benguella,” Huila Provinee, Angola. Euprepes olivaeeus-. Boeage (1970:68). Mabuia suleata: Boeage (1895a:41, 1896a:lll). Mabuia suleata ansorgiv. Monard (1937b:90), Mertens (1938a:438), Hellmieh (1957a:64), Laurent (1964a:74). Mabuia boeagi ansorgei: Laurent (1947:8). Mabuya suleata ansorgiv. Braneh (1998:157). Mabuya suleata'. Broadley (2000:102). Traehylepis suleata'. Portik (2009:23), Portik et al. (2010:147), Protik et al. (2011:1745), Ceriaeo et al. (2016a:34, 57). Traehylepis suleata ansorgiv Portik (2009:23), Portik et al. (2010:147), Protik et al. (2011:1745), Bates et al. (2014:267). Global conservation status (lUCN): Not Evaluated. Global distribution: The species extends from the Western Cape Province of South Africa, through most of Namibia except the northeast to the southwestern quadrant of Angola. Ocurrences in Angola (Map 231): The species occurs in southwestern Angola. Huambo: “Nova Lisboa” [-12.76667, 15.73333] (Hellmieh 1957a:64). Benguela: “Farta bay” [-12.60000, 13.20000] (Laurent 1947:8); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:64); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:438), “Hanha” [-13.30000, 14.20000] (Boeage 1896a:lll). Huila: “Rio Cuce” [-13.51667, 15.20000] (Boeage 1895a:41) “Caeonda” [-13.73333, 15.06667] (Boulenger 1907:213; Laurent 1947:8); “Sangeve” [-13.80000, 16.11667] (Monard 1937b:90; Laurent 1947:8). Namibe: “Munhino 50 km a I’ouest de Sa da Bandeira, 270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II district de Mogamedes” [-14.96667, 12.96667] (Laurent I964a;74; Ceriaco et al. 20I6a;57); “Campangombe” [-15.10000, 13.15000] (Bocage I895a:4I; Ceriaco et al. 2016a;57); “Iona National Park, Rio Curoca in Pediva Hot Springs area” [-16.28359, 12.56106] (Ceriaco et al. 2016a:34); “Leba Pass” [-15.07003, 13.24339] (Ceriaco et al. 2016a:34); “Namibe- Lubango road, 2 km east (by road) of Man- gueiras, south side of the road” [-15.04464, 13.15861] (Ceriaco et al. 2016a:34). Taxonomic and distributional notes: Three subspecies of this species have been rec¬ ognized; the nominate form, Trachylepis s. ansorgii (Boulenger, 1907) and T. s. nigra (Werner, 1915) (Portik 2009; Portik et al. 2010, 2011; Bates et al. 2014). Trachylepis sulcata nigra has been synonymized with the typical form but the taxonomic status of T. s. ansorgii remains uncertain (Portik et al. 2010, 2011). Ostensibly T. s. ansorgii is diagnosable by a larger snout- vent length and larger head size (Branch 1998; Portik 2009; Bates et al. 2014). The geo¬ graphic limit of ansorgii is unclear. Branch (1998) depicted both it and T. s. sulcata as occuring parapatrically in northwestern Namibia, but no concerted effort has been made to determine if there are actually two taxa in Angola and adjacent Namibia or, if there are, where their boundaries lie. Preliminary molecular phylogenetic results suggest that there is significant structure within Angolan popluations, but the taxonomic status of these cannot yet be assessed. We here treat all Angolan material T. sulcata pending further study. Map Angola. 231. Distribution of Trachylepis sulcata in Trachylepis wahlbergii (Peters, “1869” 1870) Wahlberg’s Striped Skink Euprepes Wahlbergii (Peters “1869” 1870:661). Leetotype: ZMB 6155 (eolleetor J.A. Wahlberg) designated by implieation by Boulenger (1887). Type loeality: “Damaraland” Namibia. NHR 2190 is the paraleeto- type and has the same data as the leetotype. Euprepes punctatissimus: Boeage (1866a:44, 1870:68). Mabuia striata: Boeage (1895a:41, 1896a:lll, 1897b:211), Boulenger (1905:111), Angel (1923:160), Schmidt (1933:12), Monard (1937b:88), Themido (1941:8). Mabuya striata: Parker (1936:136). Mabuya striata striata: Mananas (1963:235). Mabuya striata wahlbergi: Branch and McCartney (1992:1), Branch (1998:157), Broadley (2000:106). Traehylepis wahlbergii: Branch and Conradie (2015:200), Conradie et al. (2016:26). Traehylepis striata: Ceriaco et al. (20 16b: 69). Global conservation status (lUCN): Not Evaluated. Global distribution: A widespread species in southern Africa, from Malawi, western Mozam¬ bique, northern and western Zimbabwe and norther Botswana through Zambia and parts of the former Katanga region of the Democratic Republic of Congo to most of Namibia and Angola. Ocurrences in Angola (Map 232): Widley distributed across Angola, although apparently absent from the forested far north and the arid far southwest. Uige; “Fazenda Otilia, Encoge” [-7.55000, 15.03333] (Managas 1963:235). Malanje: “Duque de Bragan9a” [-9.10000, 15.95000] (Bocage 1866a:44, 1895a:41); “Cangandala National Park” [-9.81858, 16.65403] (Ceriaco et al. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 271 20I6b:69). Lunda Norte: “Lucapa” [-8.42278, 20.73917] (Branch and Conradie 2015:200). Lunda Sul: “Lunda” [-10.96667, 20.06667] (Monard I937b:88); “Mutianvo” [-11.45000, 19.33333] (Themido 1941:8). Mexico: “Lago Dilolo” [-11.50000, 22.01667] (Mananas 1963:235); “Lago Cameia” [-11.71667, 20.80000] (Mananas 1963:235); “Fazenda Santa Curz, Luso” [-11.78333, 19.91667] (Managas 1963:235); “Calombe-Luso” [-11.83333, 19.93333] (Mananas 1963:235). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:12); “Silva Porto” [-12.33333, 16.86667] (Managas 1963:235); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:12). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage I895a:4I); “Nova Lisboa” [-12.76667, 15.73333] (Managas 1963:235). Benguela: ^32. Distribution of Trachylepis wahlbergU in “Hanha” [-13.30000, 14.20000] (Bocage I896a:III, I897b:2II). Hufla: “Rio Cuce” [-13.51667, 15.20000] (Bocage I895a:4I); “Caconda” [-13.73333, 15.06667] (Bocage I895a:4I); “Sangeve” [-13.88333, 15.83333] (Monard I937b:88); “Kuvangu” [-14.46667, 16.30000] (Monard I937b:88); “Indungu” [-14.81667, 16.26667] (Monard I937b:88); “Kapelongo” [-14.88333, 15.08333] (Monard I937b:88); “Huilla” [-15.05000, 13.55000] (Bocage I895a:4I; Angel 1923:160); “Kambisa” [-15.31667, 16.21667] (Monard 1937b:88); “Kului” [-15.41667, 15.73333] (Monard 1937b:88); “Molundo” [-15.63333, 15.20000] (Monard 1937b:88); “Gambos” [-15.76667, 14.10000] (Bocage 1895a:41). Cunene: “Kuvelai” [-15.65000, 15.80000] (Monard 1937b:88); “Humbe/Humbi” [-16.68333, 14.90000] (Bocage 1895a:41; Monard 1937b:88); “Mupa” [-16.18333, 15.75000] (Monard 1937b:88); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:88). Cuando Cubango: “Kwito region, tributary of Kuban¬ go” [-15.16667, 19.18333] (Angel 1923:160); “Vicinity of Cuito Cuanavale - approximately 75 km W of Mavinga” [-15.78333, 19.70000] (Branch and McCartney 1992:1); “Cuando basin (43a)” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 26). Undetermined locality: “Rio Quando” [-12.58333, 13.41667] (Bocage 1895a:41); “Between Benguela and Bihe” (Boulenger 1905:111). Taxonomic and distributional notes: Trachylepis wahlbergU was long considered a sub¬ species of T. striata (Peters, 1844) (e.g., Broadley 1977a; Branch 1998), but following Broadley (2000), the former subspecies in this complex have been treated as full species. However, Castig- ilia et al. (2006) provided a molecular data analyses that suggested that T. striata and T. wahlbergU may not be reciprocally monophyletic. All records in the T. striata complex from across Namibia are assignable to T. wahlbergU as are all records from western Zambia. It would seem likely, there¬ fore that members in this group in Angola would also be assignable to this taxon. Color patterns of Angolan members of the group are variable and some specimen do at least superficially resemble true T. striata. However, it is probable on biogeographic grounds that most or all older Angolan lit¬ erature records of ‘T. striata” are, in fact, referable to T. wahlbergU, or perhaps to another species, but not to T. striata sensu stricto. Nonetheless, only in recent years have newly collected specimens been assigned to T. wahlbergU at the outset, e.g., specimens from near Mavinga (Branch and McCarthy 1992), and from Lunda Norte Province (Branch and Conradie 2015), and recently in Malanje Province (M. Marques pers. obs), in the the Laiica Dam area. Further research is needed ll^E 20“E 24''E 272 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II to determine definitively if T. wahlbergi and T. striata are specifically distinct and if all Angolan specimens are referable to the former. Pending the outcome of such research, we here tentatively treat all ''striata''' records as T. wahlbergii. Genus Typhlacontias Bocage, 1873 Typhlacontias johnsonii Andersson, 1916 Johnson’s Burrowing Skink Typhlacontias johnsonii Andersson 1916. Lectotype: GNM Re. Ex. 1388a (colleetor D. F. d’Azinhaes), des¬ ignated hy Haaeke (1997:142). Type loeality: “Port Alexander, Portuguese West Afriea” [= Tombua] Namibe Provinee, Angola. Typhlacontias punctatissimus: Boeage (1895a:56), Monard (1937b:86). Typhlacontias johnsonii'. Haaeke (1997:142), Braneh (1998:146), Ceriaco et al. (20 16a: 5 8). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and Namibia. Ocurrences in Angola (Map 233): The species is restricted to the Namibe desert. Namibe: “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:56; Monard 1937b:86; Haaeke 1997:144; Ceriaco et al. 2016a:58); “Porto Ale¬ xandre (= Tombua)” [-15.80000, 11.83333] (Andersson 1916:19; Haaeke 1997:144; Ceria¬ co et al. 2016a:58); “Lacrau” (Haaeke 1997:144; Ceriaco et al. 2016a:58). Taxonomic and distributional notes: Bocage ’s (1895a) Typhlacontias punctatis- simus Bocage, 1873 from “Coroca” was proba¬ bly referable to this species (Haaeke 1997), but it was destroyed in the fire at the Museu Bocage. 16°E 20“E 24‘^f, Map 233. Distribution of Typhlacontias johnsonii in Angola. Typhlacontias punctatissimus punctatissimus Bocage, 1873 Dotted Blind Dart SkinK Typhlacontias punctatissimus Bocage 1873b:213. Syntypes: MBL (2 specimens) catalogue numbers unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Rio Curoca,” Namibe Province, Angola. Typhlacontias punctatissimus: Bocage (1887b:203, 1897a: 197), Boulenger (1887:429), de Witte and Laurent (1943:34), Frade (1963:252-253), Branch (1998:146). Typhlacontias punctatissimus punctatissimus: Haaeke (1997:146), Ceriaco et al. (2016b:58). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and northern Namibia. Ocurrences in Angola (Map 234): Restricted to the Namibe Province. Namibe: “Mossamedes” [-15.20000, 12.15000] (Boulenger 1887:429; Haaeke 1997:150; Ceriaco et al. 2016a:58); “Mossamedes desert, 35 km south from the city” [-15.50000, 12.16667] (Laurent 1964a:82; Ceriaco et al. 2016a:58); “Rio Coroca” [-15.78333, 12.06667] (Bocage 1873b:213, 1887b:203, 1897a: 197; Ceriaco et al. 2016a:58); “Porto Alexandre (=Tombua)” [-15.80000, 11.83333] (Haaeke 1997:147; Ceriaco et al. 2016a:58). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 273 Taxonomic and distributional notes: The nomenclatural history of T. punctatissimus and its subspecies was reviewed by Haacke (1997), who validated the existence of two sympatric subspecies in southern Angola (thus suggesting that they should both be accorded specific status - see account below). Il-E 1&°E 20°E 24"E Map 234. Distribution of Typhlacontias punctatissimus punctatissimus in Angola. Typhlacontias punctatissimus bogerti Laurent, 1964 Bogert’s Dotted Blind Dart Skink (Endemic) Typhlacontias bogerti (Laurent 1964a: 82). Holotype: MD 1946 (collector A. Barros Machado). Type locality: “Desert de Mo^amedes, 35 km au sud de la ville” [= Namibe desert, 35 km south of the village ofNamibe] Namibe Province, Angola. Typhlacontias bogerti'. Haacke (1965:22). Typhlaeontias punctatissimus bogerti'. Haacke (1997:150). Typhlaeontias punetatissimus bogerti'. Branch (1998:147), Ceriaco et al. (2016a:30, 58). Global conservation status (lUCN): Not Evaluated. Global distribution: Known only from southwestern Angola. Ocurrences in Angola (Map 235): Restricted to the Namibe Province. Namibe: (Map 240) “Mossamedes (= Namibe)” [-15.20000, 12.15000] (Haacke 1965:22, 1997:150; Ceriaco et al. 2016b:58); “10 km S of Mogamedes (=Namibe)” [-15.269404, 12.164081] (Haacke 1997:150; Ceriaco et al. 2016b:58); “34 km S of Mogamedes (=Namibe)” [-15.478951, 12.203563] (Haacke 1997:150; Ceriaco et al. 2016b:58); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaco et al. 2016b:31); “8 km of SE of Pico Azevedo” [-15.55000, 12.51667] (Haacke 1997:150); “Rio Coroca” [-15.78333, 12.06667] (Haacke 1997:150; Ceriaco et al. 2016b:58); Ceriaco et al. 2016b:58; “Kakolo windmill, Iona National Park” [-16.66667, 12.33333] (Haacke I2''E 20“E 24“ E Map 235. Distribution of Typhlacontias punctatissimus bogerti in Angola. 274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1997:150; Ceriaco et al. 2016b:58); “Espinheira” [-16.78528, 12.35468] (Ceriaco et al. 2016b:31). Taxonomic and distributional notes: Typhlacontias p. bogerti appears to be endemic to Angola, restricted to northern Namib Desert in Iona Park west of the Coroca Sand Sea and south of the town of Namibe (Haacke 1965, 1997; Branch 1998). Typhlacontias rohani Angel, 1923 Rohan’s Blind Dart Skink Typhlacontias Rohani (Angel 1923:162, figs. 6-8. Holotype: MHNP 1923.1 (collector J. de Rohan-Chabot). Type locality: “Lwankundu, sous-affluent du Kwando” [= Cuando Cuhango Province, sub-tributary of Cuando River], Angola. Typhlacontias rohani: Monard (1937b:96), de Witte and Laurent (1943:35), Guibe (1954:102), Brygoo (1985:92), Haacke (1997:152), Broadley (2006:557), Conradie et al. (2016:26). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is limit¬ ed to the Kalahari sand regions of southeastern Angola, northwestern Namiba, northern Botswana, western Zimbabwe and southern Zambia. Ocurrences in Angola (Map 236): In Angola known only from the southeastern part of the country. Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937b:96). Cuando Cubango: “Lwankundu River, sub¬ tributary of Kwando” (Angel 1923:164; Bry¬ goo 1985:92; Haacke 1997:152); “Cuando basin (38)” [-17.58830, 22.65694] (Conradie et al. 2016:9-10, 26);”Cuando basin (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9- 10, 26). Taxonomic and distributional notes: Guibe (1954) incorrectly listed “Huilla” as the type locality, without justification (Brygoo 1985). Map Angola. 236. Distribution of Typhlacontias rohani in Il'E l&'E 20“E :4'*E Typhlacontias rudebecki Haacke, 1997 Rudebeck’s Blind Dart Skink (Endemic) Typhlacontias rudebecki Haacke 1997:155, fig. 5. Holotype: TM 25465 (collector G. Rudebeck). Type local¬ ity: “Sao Nicolau, Mogamedes [= Namibe] district, Angola (1412Ad).” Typhlacontias rudebecki: Ceriaco et al. (2016a:58). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 237): The species is only known from the type locality. Nami¬ be: “Sao Nicolau, Mo9amedes [= Namibe]” [-14.26667, 12.36667] (Haacke 1997:155; Ceriaco et al. 2016a:58). Taxonomic and distributional notes: This species was described by Haacke (1997) based on a single specimen from “Sao Nicolau, Mogamedes [= Namibe]” collected by G. Rudebeck. According to the orginal description this species is closer to Typhlacontias rohani Angel, 1923 and Typhlacontias gracillis Roux, 1907, than to the Namib species, however, its general proportions differ and its habitat, although not explicitly recorded, is coastal Namib Desert. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 275 ll-E 16°E lO'E W^E Map 237. Distribution of Typhlacontias rudebecki in Angola. Family Varanidae Hardwicke and Gray, 1824 Genus Varanus Merrem, 1820 Varanus albigularis albigularis (Daudin, 1802) White-Throated Monitor [Although we consider the two Angolan subspecies of V. albigularis to be pontentially valid we believe that it is not possible to meaningfully assign many locality records to one or the other. Thus we provide the occurences in Angola and taxonomic and distributional notes for the two combined.] Tupinambis albigularis Daudin 1802 [AnX]:72, pi. XXXII. Holotype: MNHN 6513 (collector unknown) fide Brygoo (1987). Type locality: “de I’Afrique ou de ITnde.” Varanus ocellatus: Bocage (1867b:220; 1870:68). Varanus albigularis: Bocage (1895a:27), Monard (1931:92), Themido (1941:7), Bayless (2002:1645). Varanus exanthematicus: Ferreira (1903:16), Themido (1941), Managas (“1955” 1957:190), Frade (1963:252). Varanus exanthematicus albigularis: Mertens (1926:152), Monard (1937b:63). Varanus albigularis albigularis: Ceriaco et al. (2016a:58). Global conservation status (lUCN): Least Concern. Global distribution: Southern Africa from Angola and Tanzania southwards. Varanus albigularis angolensis Schmidt, 1933 Angolan White-Throated Monitor Varanus albigularis angolensis (Schmidt 1933:10, pi. II). Holotype: CM 5967 (collectors R. and L. Boulton). Type locality: “Gauca, Bihe,” Bie Province, Angola. Varanus albigularis angolensis: Loveridge (1936a:59), McCoy and Richmond (1966:157), Bayless (2002:1645), Ceriaco et al. (2014b:671), Ceriaco et al. (2016a:39, 58). Varanus exanthematicus angolensis: Mertens (1937a:9), Mertens (1938a:437), Laurent (1964a:48, 1964b:3), McCoy and Richmond (1966:157). Varanus (Empagusa) exanthematicus angolensis: Mertens (1942a: 104), Mertens (1942b: 194), Mertens (1942c:359). Varanus exanthematicus: Mananas ( “1955” 1957:192). 276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Map 238. Distribution of Varanus albigularis sbspp. in Angola. Global conservation status (lUCN): Least Concern. Global distribution: This subspecies is known from northern Angola and adjacent Zambia and the Katanga region of the Democratic Republic of Congo. Ocurrences in Angola (Map 238): i2"e 24'-e Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a;48; Bayless 2002:1661). Kwanza Norte: “Golungo” [-9.13333, 14.76667] (Ferreira 1903:16; Monard 193 7b: 63; Mertens 1942c: 3 59, 360; Mananas “1955” 1957:192). Malanje: “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014a:671). Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:48; Laurent 1946b:48). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:10; Loveridge 1936:59; Mertens 1942c:359; McCoy and Richmond 1966:157; Bayless 2002:1661). Huambo: “High-Huambo” [-12.76667, 15.73333] (Mertens 1926:152, 1937a:9, 1938a:437, 1926:152, 1942a:104, 1942b:194, 210, 216, 1942c:359, 360; Bayless 2002:1661). Benguela: “Cuma” [-12.86667, 15.06667] (Bayless 2002:1661); “Lobito bay” [-12.35000, 13.55000] (Bocage 1895a:27; Mertens 1942c:359; Laurent 1964b:3-4; Bayless 2002:1662); “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:27; Mertens 1926:152, 1942a:104, 1942b:194, 216, 1942c:359, 360; Bayless 2002:1661); “Catumbel- la-Coporello” [-13.93333, 12.9666] (Mertens 1926:152, 1942c:359; Bayless 2002:1661); “Cor- porello” [-13.93333, 12.9666] (Mertens 1926:152; Bayless 2002:1661); “Quissange” [-12.43333, 14.05000] (Frade 1963:253; Mertens 1942c:359; Bayless 2002:1662); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:27; Mertens 1942c:359; Bayless 2002:1662); “Benguella” [-12.58333, 13.41667] (Bocage 1867c:220, 1895:27; Themido 1941:7, 8; Mertens 1942c:359; Managas “1955” 1957:192; Bayless 2002:1661); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1661); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:437, 1942c:360; Bayless 2002:1661). “Hanha” [-13.30000, 14.20000] (Bayless 2002:1661). Huila: “Molundo” [-15.63333, 15.20000] (Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1662). Namibe: “Rio Chimba” [-14.30000, 12.40000] (Bocage 1895a:27; Mertens 1942c:359; Bayless 2002:1661); “Biballa” [-14.76667, 13.36667] (Bocage 1895a:27; Mertens 1926:152, 1942c:359; Bayless 2002:1661); “7.5 km, NW (by road) of Pico Azevedo” [-15.47600, 12.69000] (Ceriaco et al. 2016a:39); “Caraculo” [-15.01667, 12.66667] (Bayless 2002:1661); “Chiyaka district” [vie. -15.08333, 12.73333] (Bayless 2002:1661); “Luciro-Inamangando road” [Undetermined Locality] (Bayless 2002:1662); “7.5 km NW (by road) of Pico Azevedo, adult, found in a rock crevice” [-15.47600, 12.46150] (Ceriaco et al. 2016a:39). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:63; Mertens 1942c:360; Bayless 2002:1662); “Chimporo” [-16.03333, 17.15000] (Monard 1931:92, 1937b:63; Mertens 1942c:359, 360; Bayless 2002:1661). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1931:92, 1937b:63; Mertens 1942c:360; Bayless 2002:1661). “Kubango” (Monard 1931:92; Mertens 1942c:359; Bayless 2002:1662). Undetermined localities: “Xangongo, 30 km SEE” (Bayless 2002:1662); “Cazuindo” (Mertens 1942c:359; Bayless 2002:1661). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 277 Taxonomic and distributional notes: The species was long included within the synonymy of the savanna Monitor, Varanus exanthematicus (Bose 1792), now considered limited to the north of the equatorial African forest belt (Bayless 2002). The subspecies angolensis was rejected by Broadley and Howell (1991), although it has been considered valid by many more recent authors (e.g.. Branch 1998; Bayless 2002; Bates et al. 2014). The allocation of specimens from some Angolan localities to the nominotypical form appears to reflect their referral by earlier authors, rather than a critical assessment. Phillips (2004) regarded V. a. angolensis to be present throughout Angola. Pending a more thorough evaluation of the species in Angola we have not attempted to dis¬ tinguish records of the two subspecies. Varanus niloticus (Linnaeus, 1758) Nile Monitor Lacerta mon/Yor Linnaeus 1758:201, nomen rejectum (ICZN, Opinion 540, 1959). Syntypes: Specimens fig¬ ured in Seba (1734, pi. 94, figs. 1-2; 1735, pi. 82, fig. 2, pi. 105, fig. 1), and described by Linnaeus (1754:41) (collectors unknown), presumed lost. Type locality: “in Indiis. “ [= India]. Lacerta nilotica Linnaeus 1766:369. Holotype: Specimen mentioned by Hasselquist (1757) (collector F. Has- selquist), presumed lost. Type locality: “in ^gypto” [= Egypt]. Varanus niloticus: Bocage (1866a:42, 1867b:220, 1895a:26, 1897b:210), Ferreira (1900a:50), Boulenger (1905:110), Managas (“1955” 1957:192). Monitor niloticus: Gunther (1865a:480). Monitor saurus: Bocage (1879b:95, 1887a: 178), Peters (1877a:613) Varanus niloticus: Bocage (1897b:210), Ferreira (1900a:50, 1903:15), Boulenger (1905:110), Monard (1937b:63), Mertens (1938a:436), Ceriaco et al. (2014b:671), Ceriaco et al. (2016b:87), Conradie et al. (2016:26), Dowell et al. (2016:594). Varanus (Polydaedalus) niloticus: Mertens (1942a:94, 1942c:321). Varanus niloticus niloticus: Laurent (1950a: 12; 1954a:63; 1964a:47), Hellmich (1957a:60; 1957b:69), Man¬ anas (1963:239). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Senegal, west to Somalia and Egypt, and south to South Africa. Ocurrences in Angola (Map 239): The species distribution comprises the entire coun¬ try including Cabinda Province, with exception of the desert regions of far southwestern Ango¬ la. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:613); “Rio Quilo” [-5.18333, 12.18333] (Bocage 1866a;42, 1895a:26; Mertens 1942c:323; Mana9as “1955” 1957:193; Bayless 2002:1676); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a;42, 1895a;26; Mertens 1942c;323; Managas “1955” 1957:193; Bayless 2002: 1675). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1866a:42, 1895a:26; Mer¬ tens 1942c:323; Managas “1955” 1957:193). Bengo: “Cunga River” [-9.23333, 13.76667] (Boulenger 1905:110; Mertens 1942c:321, 323; Bayless 2002:1675 ). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hell- Map 239. Distribution of Varanus niloticus in Angola. 12“E I6'E 20°E 24'*E 278 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II mieh 1957b:60); “Golungo Alto” [-9.13333, 14.76667] (Ferreira 1903:15; Mertens 1942e:321; Mananas “1955” 1957:193); “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:15; Mertens 1942e:321, 323; Bayless 2002:); “Rio Membege River, perto de N’dalla Tando” [-9.43333, 14.76667] (Ferreira 1900a:50; Mertens 1942e:321; Managas “1955” 1957:193; Bayless 2002:1675); “Mueoso” [-9.53333, 14.65000] (Hellmieh 1957a:61); “Dondo” [-9.68333, 14.43333] (Boeage 1895a:26; Mertens 1942c:323; Managas “1955” 1957:193; Bayless 2002:1675). Malan- je: “Duque de Braganga (Duque do Cunene [Bayless])” [-9.10000, 15.95000] (Boeage 1895a:26; Mertens 1942e:323; Managas “1955” 1957:193; Bayless 2002:1675); “Capanda” [-9.72841, 15.34585] (Ceriaeo et al. 2014b:671); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:87); “Rio Loando” [-11.55000, 22.01667] (Boeage 1879a:95; Mertens 1942e:323; Bayless 2002:1675). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:63, 1964a:47; Managas “1955” 1957:193); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950:12). Mexi¬ co: “Lago Dilolo” [-11.50000, 18.10000] (Managas 1963:239). Kwanza Sul: “Condo” [-11.28333, 14.33333] (Gunther 1865a:480; Mertens 1942e:323; Bayless 2002:1675). Benguela: “Catumbella (Catumbinda river [Bayless 2002])” [-12.43333, 13.55000] (Boeage 1867b:220, 1895a:26; Mer¬ tens 1942e:323; Managas “1955” 1957:193; Bayless 2002:1675); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:69); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:436; Mertens 1942a:94, 95; Mertens 1942e:324; Bayless 2002:1675); “Hanha” [-13.30000, 14.20000] (Boeage 1897b:210; Mertens 1942e:323; Managas “1955” 1957; Bayless 2002:1675). Huila: “Rio Cuee” [-13.51667, 15.20000] (Boeage 1895a:26; Mertens 1942e:323; Bayless 2002:1675); “Caeonda” [-13.73333, 15.06667] (Boeage 1895a:26; Mertens 1942e:323; Bayless 2002:1675); “Quillangues” [-14.06667, 15.08333] (Boeage 1895a:26; Mertens 1942e:323; Bayless 2002:1675); “Tyitunda” [-14.36667, 16.48333] (Monard 1937b:63; Mertens 1942e:322, 323; Bayless 2002:1676); “Vila da Ponte (= Kuvangu (Kubangu))” [-14.46667, 16.30000] (Monard 1931:93, 1937b:63; Mertens 1942e:322, 32; Mananas “1955” 1957:193; Bayless 2002:1675); “Kului” [-15.41667, 15.73333] (Monard 1937b:63; Mertens 1942e:323; Bayless 2002:1675). Cunene: “Riv. Mbale” [-15.16667, 16.75000] (Monard 1931:93, 1937b:63; Mertens 1942e:322, 323; Managas 1957 “1955”:193); “Humbe” [-16.68333, 14.90000] (Boeage 1895a:26; Mertens 1942e:323; Bayless 2002:1675); “Foz de Cunene” [-17.28333, 11.80000] (Bayless 2002:1675); “Sighting near Cunene” (Monard 1937b:63; Mertens 1942e:323; Bayless 2002:1675). Cuando Cubango: “Cunga” [-17.30000, 20.80000] (Dowell et al. 2016:594); “Cuito basin (29)” [-17.87291, 19.83333] (Conradie et al. 2016:8-9, 26); “Cuito basin (35) not eolleeted” [-16.62322, 19.05352] (Conradie et al. 2016:9-10, 26); “Cuando basin (43) not eolleeted” [-17.53500, 23.18916] (Conradie et al. 2016:9-10, 26); “Cuando basin (44a) not eolleeted” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 26); “Cubango basin (45) not eolleeted” [-16.88350, 18.01180] (Conradie et al. 2016:9, 12, 26). Unde¬ termined Locality: Rio Quanza” (Boeage 1895a:26; Mertens 1942e:323); “Mayumba” (Managas “1955” 1957:193); “Rio Quango” (Boeage 1895a:26; Mertens 1942e:323; Managas “1955” 1957:193; Bayless 2002:1675); “Kubango Gebiet” (Mertens 1942e:322). Taxonomic and distributional notes: The name Lacerta monitor Linnaeus, 1758 initially assoeiated with this and other speeies of varanids, was rejeted by the ICZN under Opinion 540 and Lacerta nilotica, based on a speeimen deseribed by Hasselquist (1757) is the aeeepted name. Varanus niloticus eomprises three major elades of monitors, with the sole Angolan sample inelud- ed elustering with a very broadly distributed southern elade, also ineluding virtually all samples south of 3°N (Dowell et al. 2016). This ineludes those speeimens in this geographie range that had previously been assigned to Varanus ornatus (Daudin, 1803) whieh had been elevated to full speeifie status by Bohme and Ziegler (1997) based on external features, partieularly eolor. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 279 Family Chamaeleonidae Gray, 1825 Genus Chamaeleo Laurenti, 1768 Chamaeleo anchietae Bocage, 1872 Anchieta’s Chameleon Chamaeleon Anchietae Bocage 1872:72, fig. p. 72. Syntypes: MBL (originally 5 specimens) catalogue num¬ bers unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978; Bauer et al. (2006:270) iden¬ tified ZMB 7756 as a surviving syntype, exchanged from Bocage to Wilhelm Peters. Type locality: “Huil- la, dans finterieur de Mossamedes” (Bocage 1872:72), [= Huila], Huila Province, Angola. Chamaeleo anehietae: Boulenger (1887:452), Tilbury and Tolley (2009:63). Chamaeleon Anehietae: Bocage (1895a:62, 1897a:198), Chamaeleon anehietae: Loveridge (1933:333), Frade (1963:252). Chamaeleo senegalensis anehietae: Loveridge (1957:198). Chamaeleo anehietae: Broadley and Cotterill (2004:40), Bauer et al. (2006:270), Tilbury (2010:447), Ceria- co et al. (2016a:58). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs from the interior plateaus to montane grass¬ lands in southwestern Angola through the for¬ mer Katanga Province of the Democratic Republic of Congo and the Albertine Rift to southwestern Tanzania and Namibe. Ocurrences in Angola (Map 240): The species is known from southwestern Angola. Huila: “Lobango” [-14.91667, 13.50000] (Bocage 1895a:62, 1897a:198); “Huilla” [-15.05000, 13.55000] (Bocage 1872:72, 1895a:62, 1897a:198; Loveridge 1933:333, 1957:198; Bauer et al. 2006:270). Namibe: “Namibe” (Tilbury 2010:451; Ceriaco et al. 2016a:58). Taxonomic and distributional notes: Based on Bocage’s (1872) somewhat ambigu¬ ous locality Boulenger (1887) wrongly consid¬ ered the terra typica to be “Mossamedes, South-west Africa.” Bocage (1895a) restated the locality simply as “Huilla”. Map 240. Angola. Distribution of Chamaeleo anchietae in Chamaeleo dilepis quilensis Bocage, 1886 Quito Flap-Neck Chameleon Chamaeleon Dilepis Leach 1819:493. Holotype: BMNH 1946.8.13.75 (formerly BMNH 25.5a) (presented by T.E. Bowditch). Type locality: Not stated. “French Congo [= CongoY fide Klaver and Bohme (1987). Chamaeleo Capellii (Bocage 1866a:42, 1866c:59). Holotype: MBL specimen number unknown (collector A. Capello), destroyed by fire 18 March 1978. Type locality: “Benguela,” Benguela Province, Angola. Chamaeleo dilepis var. quilensis Bocage 1866c:59. Holotype: MBL specimen number unknown (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Rio Quilo,” Angola. Chamaeleon dilepis: Bocage (1866a:42, 1866c:59, 1867b:219, 1870:68, 1895a:59, 1896a:112), Peters (1887:612), Boulenger (1887:450, 1905:112), Monard (1937b:99), Themido (1941:8). Chamaeleo dilepis: Bocage (1879c:88, 1887a: 178, 1887b:202, 1887c:209). Chamaeleon parvilobus: Boulenger (1887:449). Chamaeleon quilensis: Bocage (1895a:60, 1897a: 198). Chamaeleon quilensis: Boulenger (1905:112), Monard (1937b:99). 280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Chamaeleo dilepis quilensis: Angel (1923:165), Loveridge (1933:331, 1957:199), Glaw (2015:201). Chamaeleo dilepis: Schmidt (1933:12), Mertens (1937a:7, 1938a:435), Parker (1936:141), Managas (1963:231), Branch and Conradie (2015:200). Chamaeleo dilepis dilepis: Loveridge (1936a:76, 1957:199), Laurent (1950a: 12, 1954a:65, 1964a:44), Hellmich (1957a:52, 1957b:53), Chirio and LeBreton (2007:182), Glaw (2015:200). Chamaeleo quilensis: Laurent (1954a:65, 1964a:44), Chirio and LeBreton (2007:198). Chamaeleo dilepis ''quilensis'’": Tilbury (2010:495). Chamaeleo dilepis: Tolley (2014), Ceriaco et al. (2016a:58), Ceriaco et al. (2016b:73), Conradie et al. (2016:24). Global conservation status (lUCN): Least Coneem. Global distribution: The species as a whole is widely distributed throughout southern and eastern Africa, although the limits of its range in some areas are under dispute. There is general agreement that the distribution includes the area from Congo east to Tanzania, Kenya, Somalia and Ethiopia, and south to South Africa. However, records from Gabon and north and from Uganda have been questioned (Chirio and Ineich 2006; Pauwels and Vande weghe 2008), and it may be difficult to distinguish this taxon from its near relative C. gracilis, with which it is sympatric in parts of Central Africa. Ocurrences in Angola (Map 241): Chamaeleo dilepis quilensis is widely distrib¬ uted across the entire country. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a;612); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:42, 1866c:59, 1895a;59); Rio Quilo” [-5.18333, 12.18333] (Bocage 1866a:42, 1866b:59, 1895a;60, 1897a:198; Loveridge 1933:331, 1957:199; Chirio and LeBreton 2007:198). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a:178, 1895a:59; 1897a: 198). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12; 1954a:65); “Muita (Luembe E) “ [-7.80000, 21.45000] (Laurent 1950a:121954a:65); “Carumbo” [-7.74172, 19.95467] (Branch and Conradie 2015:200). Lunda Sul: “Cacolo (Minungo)” [-10.15000, 19.28333] (Mananas 1963:231). Moxico: “Lago Dilolo” [-11.50000, 22.01667] (Mananas 1963:231); “Teixeira de Sousa, distr. Villa Luso” [-10.70000, 22.23333] (Mertens 1937:8); “Lago Cameia” [-11.71667, 20.80000] (Mananas 1963:231); “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] (Mananas 1963:231); “Calombe, Luso” [-11.83333, 19.93333] (Managas 1963:231); “environs du lac Calun- do” [-11.80000, 20.86667] (Laurent 1964a:44); “Calunda” [-12.11667, 23.46667] (Laurent 1964a:44); “Macondo” [-12.55000, 23.76667] (Laurent 1964a:44); “Cazombo” [-11.88333, 22.91667] (Laurent 1954a:65). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Bocage 1895a:59); “Cangandala National Park” [-9.81858, 16.65403] (Ceriaco et al. 2016b:73). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1867b:219, 1895a:59). Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:12); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:12); “General Machado (= Camacupa)” [-12.03333, 17.46667] (Mertens 1937a: 7); “Farm Goedecke on Conjo, 25 km north of General Machado (= Camacupa)” [-11.798563, 17.5114] (Mertens 1937a:6; Map 241. Distribution of Chamaeleo dilepis quilensis in Angola. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 281 Loveridge 1947:208). Huambo: “Bela- Vista (= Sanguengue)” [-12.36667, 16.20000] (Hellmich 1957b:53); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:141); “Cuma” [-12.86667, 15.06667] (Loveridge 1936a:76). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:59); “Cahata” [-12.35000, 14.81667] (Bocage 1895a:59); “Catumbella” [-12.43333, 13.55000] (Bocage 1867b:219); “Quissange” [-12.43333, 14.05000] (Bocage 1887b:209, 1895:59); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:59); “Benguella” [-12.58333, 13.41667] (Bocage 1866a:42, 1866c:59, 1867b:219, 1895a:59-60; Loveridge 1957:200); “Dombe” [-12.95000, 13.10000] (Boca¬ ge 1867b:219, 1895a:60); “Entre Rios/Chitidi” [-13.01667, 14.63333] (Mertens 1938a:435, Hell¬ mich 1957a:52); “Cubal” [-13.03333, 14.25000] (Mertens 1937a:7, 1938a:435; Hellmich 1957b:53); “Marco de Canavezes, perto da barragem do rio Cubal” [-13.30000, 14.20000] (Boca¬ ge 1896a:112); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:112); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:99). Hulla: “Caconda” [-13.73333, 15.06667] (Bocage 1879c:88; Loveridge 1936a:76); “Lubango” [-14.91667, 13.50000] (Bocage 1895a:59); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:59); “Mutundo” [-15.23333, 13.36667] (Frade 1963:231); “Gambos” [-15.76667, 14.10000] (Bocage 1895a:59); “Kampulu, environs de Kasinga” [-15.21667, 16.11667] (Monard 1937b:99); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:99); “Kului” [-15.41667, 15.73333] (Monard 1937b:99). Namibe: “Mossamedes” [-15.20000, 12.15000] (Bocage 1867b:219, 1887b:202, 1895a:59; Boulenger 1887:451; Ceriaco et al. 2016a:58); “Chim- ba” [-15.20000, 13.68333] (Hellmich 1957b:53; Ceriaco et al. 2016a:58). Cunene: “Mupanda” [-17.13333, 15.76667] (Monard 1937b:99). Cuando Cubango: “Lwankundu, sub-tributary of Kwando” [Undetermined Locality] (Angel 1923:165); “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9-10, 24). Undetermined locality: “Between Bihe and Quilenges” (Boulenger 1905:112); “Mayumba/Maiumba” (Bocage 1887a:178, 1895a:60, 1897a:198); “Cuan- za r.” (Boulenger 1905:112). Without precise locality: “with no precise locality” (Laurent 1964a:44). Taxonomic and distributional notes: Bocage (1866a) noted some distinct characters in a specimen collected by Anchieta in “Rio Quilo” relative to the typical form, Chamaeleo dilepis Leach, 1819, however, variation across the C. dilepis species complex as a whole suggests that there are few if any consistent differences among these forms and the taxonomy of the C. dilepis complex remains unresolved (Glaw 2015). The issue is exacerbated by the vague type locality of C. dilepis, “Congo,”which, in the BMNH register is given as “Gaboon, French Congo” (although Pauwels and Vande weghe 2008) stated that the species is absent from Gabon). Loveridge (1936a) refered all Angolan specimens from the Field Museum to C. d. quilensis, with the exception of one specimen from “Cuma,” which he regarded as typical dilepis (FMNH 15365). Broadley (1966) and Tilbury (2010) considered quilensis a variant of the polymorphic C. dilepis, although Klaver and Bohme (1986) and Ullenbruch et al. (2007) identified differences in hemipenial morphology and regarded quilensis as subspecifically (Klaver and Bohme 1987) or even specifically distinct from nominotypical C. dilepis (Klaver and Bohme 1986). Tilbury (2010) provided an extended discus¬ sion of the taxonomic and distributional issues surrounding the C. dilepis complex. Regardless if quilensis is taxonomically distinct from C. dilepis sensu stricto, it seems likely that all members of the complex form Angola are referable to the same form (although some authors have assigned some Angolan material to another doubtful form, C. d. roperi) and we here treat all Angolan spec¬ imens as C. d. quilensis pending a more detailed phylogeographic study of the group. 282 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Chamaeleo gracilis etiennei Schmidt, 1919 Etienne’s Graceful ChamaeleoN Chamaeleo gracilis Hallowell 1842:324, pi. 18. Holotype: ANSP 7325 (“purchased by the Rev. Charles Eden of Monrovia, of one of the Affiean natives”). Type loeality: “Liberia, Western Affiea.” Chamaeleon etiennei Sehmidt 1919:574. Holotype: AMNH 11370 (eolleetor H. Lang). Type loeality: “hills near Banana at the mouth of the Congo,” Demoeratie Republie of Congo. Chamaeleon graeilis: Boeage (1866a:41, 1867b:219, 1895a:61), Ferreira (1900a:50, 1903:16, 1904:117), Boulenger (1887:448, 1905:111), Monard (1937b:98), Themido (1941:8) Chamaeleon senegalensis: Boeage (1870:68), Peters (1881:147). Chamaeleon graeilis: Boulenger (1887:448, 1905:111), Boeage (1895a:61). Chamaeleo senegalensis: Gunther (1865a:480). Chamaeleon etiennei: Parker (1936:140), Laurent (1964a:42). Chamaeleo graeilis: Hellmieh (1957a:53, 1957b:53), Laurent (1964a:44). Chamaeleo graeilis graeilis: Loveridge (1957:198). Chamaeleo graeilis etiennei: Loveridge (1957:197). Chamaeleo senegalensis senegalensis: Loveridge (1957:197). Chamaeleon graeilis etiennei: Tilbury (2010:507), Ceriaeo et al. (2014b:670), Glaw (2015:202) Global conservation status (lUCN): Least Concern. Global distribution: Known from the sounthem Congo to northern Angola Ocurrences in Angola (Map 242): Limit¬ ed to areas from Kwanza Sul and Lunda Sul northwards, including Cabinda. Although we located no published historical records from Lunda Norte, Tilbury (2010) plotted them there, and it is likely that they occur in this province as well as the immediately adjacent Democratic Republic of Congo. Cabinda: “Chinchoxo” [-5.10000, 12,10000] (Peters 1877a:612; 1881:147; Boeage 1895a:61); “Cabinda” [-5.55000, 12.18333] (Boeage 1895a:61). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:42); “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a;42). Luanda: “Loanda” [-8.83333, 13.26667] (Boeage 1867b:219). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:53); “Ambaca” [-9.26667, 15.18333] (Boeage 1895a;61); “N’Dalla Tando” [-9.30000, 14.91667] (Ferreira 1904:111); “Cazengo” [-9.33333, 14.76667] (Ferreira 1903:16); “Mucoso” [-9.53333, 14.65000] (Hellmieh 1957a;53); “Dondo” [-9.68333, 14.43333] (Boeage 1895a:61); “Canhoca” [-9.25000, 14.68333] (Ferreira 1904:111; Parker 1936:140). Lunda Sul: “Lunda” [-10.96667, 20.06667] (Monard 1937b:98); “Mutianvo” [-11.45000, 19.33333] (Themido 1941:8). Mexico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:42). Kwanza Sul: “Lembu, Serra de Selles” [-12.86667, 14.11667] (Ferreira 1904:117); “Congulu” [-10.86667, 14.28333] (Parker 1936:140); “Condo” [-11.28333, 14.33333] (Gunther 1865a:480; Boulenger 1887:448; Parker 1936:140). Benguela: “Ebanga” [-12.73333, 14.73333] (Monard 1937b:98). Undetermined locality: “Quango = Cuango” (Peters 1877a:612, 1881:147; Boeage 1895a:61) (Malanje Province, impossible to georeferenced. See History Section for more detailed information); “Carangigo” (Boulenger 1887:448). Map 242. Distribution of Chamaeleo gracilis etiennei in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 283 Taxonomic and distributional notes: Records of Chamaeleo senegalensis Daudin, 1802 from Angola are referable to C g. etiennei (see Peters Bocage 1895a). Both Tilbury (2010) and Glaw (2015) recognized Chamaeleo gracilis etiennei as a subspecifically distinct forest-dwelling form. The more southerly records listed above require confirmation and may refer to C. dilepis. Chamaeleo namaquensis Smith, 1831 Namaqua Chameleon Chcemelio Namaquensis Smith 1831:17. Syntypes: BMNH 65.5.4.61-62, MNHP 8017 (collector A. Smith) (see notes helow). Type locality: “western coast of South Africa, near to the mouth of the Orange river” [Namaqualand]. Chamaeleon namaquensis: Bocage (1867c:227, 1870:68, 1895a:62), Boulenger (1887:462), Frade (1963:253). Chamaeleo tubereuliferus: Gunther (1865a:480). Chamaeleo namaquensis = C. tubereuliferus: Bocage (1872:72). Chamaeleo namaquensis: Branch (1998:228), Tilbury (2010:519), Glaw (2015:203), Ceriaco et al. (2016a:58). Global conservation status (lUCN): Least Concern (Carpenter 2011). Global distribution: The species ranges from western South Africa through Namaqualand and the Namib Desert to southern Angola. Ocurrences in Angola (Map 243): The species occurs only in the arid portion of south¬ western Angola mainly in Namibe Province. Namibe: “Mossamedes” [-15.20000, 12.15000] (Gunther 1865a:480; Bocage 1867c;227, 1872:72, 1895a:62; Boulenger 1887:462; Ceriaco et al. 2016a:58). Taxonomic and distributional notes: Smith (1831) did not specify the number of the type specimens, nor their disposition. There is a long history of one stuffed specimen from Andrew Smith being present in Paris (Dumeril and Dumeril 1851; Dumeril 1852; Brygoo 1983, although not mentioned by Guibe 1954) and this specimen has generally been regarded as a syntype and is, in fact, the only syntype mentioned by Klaver and Bohme (1987) and Glaw (2015). However, FitzSimons (1937) mentioned two specimens in the BMNH that he regarded as types. These specimens correspond to BMNH 65.5.4.61-62, which although not marked as types in the BMNH registers, were obtained from Andrew Smith and have the explicit locality, “mouth of Orange River.” We regard these specimens as part of the original type series. Klaver and Bohme (1997) also noted the possibility that additional types might yet be found in the collection of the National Museums of Scotland. This species represents a distinct phylogenetic lineage within Chamaeleo and some authors have suggested that this taxon could be recognized as generically distinct (Townsend and Larson 2002), however it is strongly supported as a member of the Chamaeleo clade (Bates et al. 2014). Map 243. Distribution of Chamaeleo namaquensis in Angola. 284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Trioceros Swainson, 1839 Trioceros oweni (Gray, 1831) Owen’s Chameleon Global conservation status (lUCN): Least Concern. Global distribution: Widely distributed in Central Africa, from southeastern Nigeria to Cabinda and across the northern half of the Democratic Republic of Congo. Ocurrences in Angola (Map 244): The species has been recorded only from the north¬ ern portion of the Cabinda enclave. Cabinda: “Cabinda” [-5.55000, 12.18333] (Tilbury 2010:703, 705). Taxonomic and distributional notes: This species is the type species of the genus Trioceros Swainson 1831, which Tilbury and Tolley (2009) and subsequent authors have treted as a distinct genus. Map 244. Distribution of Trioceros oweni in Angola. Chamaeleo Oweni Gray 1831b:7. Syntypes: BMNH 1946.8.21.48-49 (formerly BMNH xxiv.l7.b and xxiv.l7a) (collector Capt. W. Owen). Type locality: “Fernando Poo” [Bioko, Equatorial Guinea]. Chamaeleo oweni: Chirio and LeBreton (2007:190). Trioeeros oweni: Tilbury and Tolley (2009:65-66), Tilbury (2010:700), Trape et al. (2012:200), Glaw (2015:229). Family Agamidae Gray, 1827 Genus Acanthocercus Fitzinger, 1843 Acanthocercus cyanocephalus (Falk, 1925) Angolan Tree Agama Agama eyanoeephala Falk 1925:83. Neotype: ZFMK 88492 (collector P. Wagner) designated by Wagner et al. (2018:27). Neotype locality: “garden in Ikelenge (-11.241592, 24.273256), northern Mwinilunga Dis¬ trict,” Zambia. Original type locality: “Angola” without further precision. Stellio nigrieollis: Bocage (1866a:43). Stellio atrieollis: Bocage (1879b:95, 1895a:22), Peters (1881:147), Boulenger (1885:358, 1905:110), Ferreira (1900a:49), Angel (1923:158), Schmidt (1933:9), Parker (1936:132), Monard (1937b:60), Themido (1941:7). Agama atrieollis: Boulenger (1885:356, 1905:110), Laurent (1950a: 12, 1964a:38), Ferreira (1903:15), Schmidt (1933:9), Monard (1937b:58), Parker (1936:132), Themido (1941:7), Frade (1963: 253), Man¬ anas (1963:228). Agama atrieollis atrieollis: Klausewitz (1957:161). Agama eyanogaster: Loveridge (1957:195). Aeanthoeereus atrieollis: Spawls (2010). Aeanthoeereus eyanoeephalus: Ceriaco et al. (2014b:670), Ceriaco et al. (2016b:75), Wagner et al. (2018:27). Aeanthoeereus eyanogaster: Branch and Conradie (2015:200). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from the southern Democratic Republic of Congo, through Angola and western Zambia to far northern Namibia. Occurrences in Angola (Map 245): The species occurs in the entire country with exception MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 285 of the southwest and the northwestern regions. Malanje: “Bange N’gola” [-8.43333, 16.56667] (Boulenger 1905:110); “Duque de Braganga” [-9.10000, 15.95000] (Boeage I866a:43, I895a:22; Boulenger 1885:359, 1905:110; Ferreira 1903:15; Monard 1937b:57; Loveridge 1957:196); “Malanje” [-9.55000, 16.35000] (Peters 1881:147; Boca- ge 1895a:22; Monard 1937b:57); “Pungo- Andongo” [-9.66667, 15.58333] (Boulenger 1905:110); “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 20146:670); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:75). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 12); “Cassange” [-9.58333, 17.86667] (Boeage 1895a:22; Monard 193 7b: 57; Wagner et al. 2018:47); “Carumbo, Lucapa” [-8.42278, 20.73917] (Branch and Conradie 2015:200); “Cacolo (Minungo)” [-10.15000, 19.28333] (Mananas 1963:228). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:38); “Village Sa-Tchisseke, pres des sources du Cuilo, Alto Chicapa” [-10.86667, 19.38333] (Laurent 1964a:39); “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a:38); “Lunda” [-10.96667, 20.06667] (Monard 1937b:57, 58; Wagner et al. 2018:47). Mexico: “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] (Managas 1963:228); “environs du lac Calundo “ [-11.80000, 20.86667] (Laurent 1964a:39); “Calombe, Luso” [-11.83333, 19.93333] (Managas 1963:228); “Cazombo, Haut Zambeze” [-11.88333, 22.91667] (Laurent 1964a:39); “Calunda, Haut Zambeze” [-12.11667, 23.46667] (Laurent 1964a:39); “Cassamba” [-13.10000, 20.35000] (Managas 1963:228); “Sessa (Luchazes)” [-13.91667, 20.80000] (Mananas 1963:228); “Vila Gago Coutinho (Bundas)” [-14.10000, 21.43333] (Mananas 1963:228). Luanda: “Dande” [-8.46667, 13.38333] (Wagner et al. 2018:47). Kwanza Sul: “Mombolo” [-12.16667, 14.83333] (Wagner et al. 2018:47). Bie: “Chitau “ [-11.43333, 17.15000] (Schmidt 1933:9; Wagner et al. 2018:47); “Silva Porto” [-12.38333, 16.95000] (Managas 1963:228); “Cachingues” [-13.06667, 16.75000] (Mana9as 1963:228). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:57, 58); “Galanga (Galange)” [-12.06667, 15.15000] (Boeage 1895a:22; Monard 1937b:57); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:132); “Huambo” [-12.76667, 15.73333] (Wagner etal. 2018:47); “Cuma” [-12.86667, 13.06667] (Wagner et al. 2018:47). Benguela: “Quissange” [-12.43333, 14.05000] (Boeage 1895a:22; Monard 1937b:57); “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a:22; Monard 1937b:57). Huila: “Rio Cuce” [-13.51667, 15.20000] (Boeage 1895a:22; Monard 1937b:57); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:57, 58); “Caconda” [-13.73333, 15.06667] (Boeage 1895a:22; Monard 1937b:57; Themido 1941:7). LFudetermined Locality: “with no precise location” (Boeage 1879b:95), “Quando” (Monard 1937b:57); “Cuanzar.” (Boulenger 1905:110). Taxonomic and distributional notes: Acanthocercus species outside of Arabia and the Horn of Africa, including those in Angola, have long been referred to A. atricollis (Smith, 1849). A ref¬ erence to Stellio nigricolis (Boeage 1866a) appears to be a lapsus for^. atricollis as well (Ceriaco et al. 2014b; Wagner et al. 2018). Klausewitz (1957) divided this species into several subspecies, but Angolan material fell into the nominotypical form (Branch 1998, Ceriaco et al. 2014b). Subse- KFi'; Map 245. Distribution of Acanthocercus cyanocephalus in Angola. 286 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II quent revisionary work on the genus (Wagner et al., 2018) revealed that some Angolan Acantho- cercus are referrable to A. cyanocephalus, a long-forgotten name proposed by Falk (1925) based on Angolan material, but without a precise type locality. Southern Angolan and northern Namibian specimens referred to A. cyanocephalus by Wagner et al. (2018), however, represent an unde¬ scribed lineage, treated here as Acanthocercus sp. (see below). Loveridge (1957) erroneously syn- onymized Stellio nigricollis cited by Bocage (1866a) from Angola with Agama cyanogaster (Riip- pell, 1835) and more recently Branch and Conradie (2015) also cited this species from Angola. This certainly represents a misidentification or a lapsus for A. cyanocephalus, since A. cyanogaster is restricted to the Horn of Africa and adjacent parts of East Africa (Uetz and Hosek 2017). Acanthocercus sp. Stellio atricollis: Bocage (1895a:22). Agama atricollis: Monard (193 7b: 5 8). Agama colonorum: Angel (1923:159). Agama cyanogaster. Loveridge (1957:195). Acanthocercus atricollis: Braneh (1998:218), Spawls (2010). Acanthocercus cyanocephalus: Wagner et al. (2018:27). Global distribution: The species is known only from Angola and adjacent north¬ ern Namibia. Occurrences in Angola (Map 246): The species is only known from the plateau areas in the southwestern regions of the country. Huila: “Vila da Ponte” [-14.46667, 16.30000] (Monard 1937b:57, 58; Wagner et al. 2018:46); “Capelongo” [-14.88333, 15.088333] (Wagner et al. 2018:47); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:22; Monard 1937b:57; Loveridge 1957:196). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:57, 58); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:57, 58). Cuando Cubango: “Region du Kwito, affluent du Kubango” [-15.162795, 19.169871] (Angel 1923:158). “Kakindo” [-15.45000, 17.05000] (Monard 1937b:57, 58). IZ”!-: 24“E Map 246. Distribution of Acanthocercus sp. in Angola. Taxonomic and distributional notes: Wagner et al. (2018) included this taxon within their concept of A. cyanocephalus, however, morphological and molecular analyses of freshly collected material in Huila Province suggest that the southwestern Angolan populations of Acanthocercus represent an undescribed lineage, closely related to A. atricollis. A formal description of this new form is being prepared. A specimen reported by Angel (1923) from the Kwito region of Cuando Cubango as Agama colonorum is here presumed to be referrable to this new form, as it is the only large, arboreal agamid occurring in this region of Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 287 Genus Agama Daudin, 1802 Agama aculeata Merrem, 1820 Western Ground Agama Agama aculeata Merrem 1820:53. Leetotype: Speeimen illustrated in Fig. 6, plate 8 of Seba (1735) (eollee- tor unknown), probably lost, designated by Wagner et al. (2012). Type loeality: “Promontorio bonae spei” [= the Cape of Good Hope] South Afriea. Agama aculeata'. Boeage (1866a:43, 1867b:221), Braneh (1998:212), Leaehe et al. (2009:274), Wagner et al. (2012:177), Ceriaeo et al. (2016a:58). Agama armata: Boeage (1870:68, 1879e:88, 1895a:19, 1896a:110, 1896b:127), Boulenger (1905:110). Agama hispida aculeata'. Loveridge (1936a:52), Mertens (1938a:432), Hellmieh (1957a:38, 1957b:51), Mananas (1963:229), Laurent (1964a:40). Agama hispida var. aculeata'. Monard (1937b:59). Agama aculeata aculeata'. Bates et al. (2014:303). Global conservation status (lUCN): Least Concern. Global distribution: Endemic to southern Africa where it has an extensive range from south¬ ern Angola, most of Namibia except the Namib Desert proper, western Botswana and the western half of South Africa. Ocurrences in Angola (Map 247): The species widespreasd in Angola except for the more northerly provinces. Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Boeage 1866a:43, 1895a:19, 1896b:127). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a:40). Moxico: “Lago Cameia” [-11.71667, 20.80000] (Mananas 1963:229), “environs du lac Claundo” [-11.80000, 20.86667] (Laurent 1964a;40), “Calombe, Luso” [-11.83333, 19.93333] (Mana9as 1963:229); “Cassamba” [-13.10000, 20.35000] (Managas 1963:229). Bie: “Silva Porto” [-12.38333, 16.95000] (Mananas 1963:229); “Cubango basin (15)” [-12.78555, 16.75694] (Conradie et al. 2016:8-9, 23). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:59); “Galanga” [-12.06667, 15.15000] (Boeage 1895a:19); “Bela-Vista (Sanguengue)” [-12.36667, 16.20000] (Hellmieh 1957b:51); “Nova Lisboa” [-12.76667, 15.73333] (Mananas 1963:229); “Santo-Amaro” [-12.70000, 15.85000] (Monard 1937b:59); “Cuma” [-12.86667, 15.06667] (Loveridge 1936a:52). Benguela: “Cahata” [-12.35000, 14.81667] (Boeage 1895a: 19) “Quissange” [-12.43333, 14.05000] (Boeage 1895a:19); “Benguella” [-12.58333, 13.41667] (Loveridge 1936a:52); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:59); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:38); “Cubal” [-13.03333, 14.25000] (Mertens 1938:432); “Hanha” [-13.30000, 14.20000] (Boeage 1896:110). Huila: “Rio Cuce” [-13.51667, 15.20000] (Boeage 1895a:19); “Cassoco” [-13.65000, 15.01667] (Boeage 1895a:19); “Caconda” [-13.73333, 15.06667] (Boeage 1879c:88); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:59); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:59); “Boca de Humpata, Sa da Bandeira” [-14.93333, 13.51667] (Laurent 1964a:40); “Chibia, Rio Huila” [-15.20000, 13.68333] (Hellmieh 1957b:51); “Kambisa” [-15.31667, 16.21667] (Monard 1937b:59); “Kului” [-15.41667, 15.73333] (Monard 1937b:59). Map 247. Distribution of Agama aculeata in Angola. 288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Namibe: “Biballa” [-14.76667, 13.36667] (Bocage I895a:I9; Ceriaco et al. 20I6a:58); “Virei- Cahinde” [-15.73333, 12.95000] (Hellmich I957b:5I; Ceriaco et al. 2016a;58); “Molundo” [-15.63333, 15.20000] (Monard 1937b:59); “Chimporo” [-16.03333, 17.15000] (Monard 1937b:59; Ceriaco et al. 2016a:58). Cunene: “Mupanda” [-17.13333, 15.76667] (Monard 1937b:59); “Riv. Mbale” [-15.16667, 16.75000] (Monard 1937b:59); “Kakindo” [-15.45000, 17.05000] (Monard 1937b:59); “Humbe” [-16.68333, 14.90000] (Bocage 1895a:19); “Mupa” [-16.18333, 15.75000] (Monard 1937b:59). Undetermined Locality: “Between Bihe and Quil- lenges” (Ferreira 1905:110); “Between Benguela and Bihe” (Ferreira 1905:110); “Quando” (Bocage 1895a: 19). Taxonomic and distributional notes: The “holotype” of A. aculeata was previously identi¬ fied as ZMB 750 (Denzer et al. 1997). However, Wagner et al. (2012) demonstrated that this spec¬ imen, actually 2in Agama atra Daudin, 1802, was one of four syntypes. They reviewed the complex history of these specimens and selected as a lectotype the only one of the syntypes actually refer¬ able to A. aculeata as currently recognized. The species Agama hispida (Kaup, 1827) mvd Agama armata Peters, 1854 have previously been included in the Angolan fauna by early authors. Agama hispida is, however, endemic to South Africa and adjacent southwestern Namibia (Branch 1998; Bates et al. 2014) and Angolan citations to it are likely referable to A. aculeata or, in some cases, to the superficially simxXdiV Agama anchietae Bocage, 1896, with which it occurs sympatricaly in Southern Angola and Namibia. Agama armata likewise is extralimital, with a distribution in south¬ eastern Africa. Agama anchietae Bocage, 1896 Anchieta’s Agama Agama Anchietae (Bocage 1896b: 129). Syntypes: MBL speeimen numbers unknown (eolleetors J.A. d’Anehieta [Benguella, Catumbella, Dombe] and H. Capello and R. Ivens [Mossamedes]), destroyed by fire 18 Mareh 1978. Type loeality: “Benguella, Catumbella, Dombe et Mossamedes,” Benguella and Namibe provinees, Angola. Agama aeuleata: Boeage (1867b:221). Agama armata: Boeage (1887b:203, 1895a: 19). Agama Anehietae: Boeage (1897a: 194). Agama anehietae: Boulenger and Power (1921:268), Monard (1937b:59), Frade (1963:253), Laurent (1964a:42), Braneh (1998:214), Leaehe et al. (2009:274), Bates et al. (2014:304), Ceriaeo et al. (2016a:21, 58). Agama anehietae anehietae: Sehmidt (1933:9), Parker (1936:131), Laurent (1947:5). Global conservation status (lUCN): Not Evaluated. Global distribution: This species has an extensive geographical range that extends from the northwestern part of South Africa northwards through Namibia and Angola to the southern Demo¬ cratic Republic of the Congo. Ocurrences in Angola (Map 248): The species occurs chiefly in southern lowland regions of Angola, however, there are two isolated records in Bie Province, central Angola, which require confirmation. Bie: “Gauca” [-11.18333, 17.45000] (Schmidt 1933:9); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:9). Benguela: “Lobito” [-12.35000, 13.55000] (Monard 1937b:59); “Catumbella” [-12.43333, 13.55000] (Bocage 1867b:221, 1895a:21, 1896b:129, 1897a:194; Boulenger and Power 1921:269; Laurent 1947:5; Monard 1937b:59); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:221, 1895a:21, 1896b:127, 1897a:194; Laurent 1947:5; Monard 1937b:59); “Huxe” [-12.71667, 13.20000] (Boulenger and Power 1921:269); “Dombe” [-12.95000, 13.10000] (Bocage 1867b:221, 1895a:21, 1896b:127, 1897a:194; Laurent 1947:5; Monard 1937b:59). Huila: “Caconda” [-13.73333, 15.06667] (Boulenger and Power 1921:269; Monard 1937b:59). Namibe: “Maconjo (= Fazenda Mucungo)” [-14.782192, 12.486557] MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 289 (Boulenger and Power 1921:269; Monard I937b:59; Ceriaco et al. 20I6a:58); “Mossamedes” [-15.20000, 12.15000] (Bocage I887b;203, I895a:2I, I896b:I27, 1897a: 194; Monard I937b:59; Ceriaco et al. 20I6a:58); “Iona National Park” [-16.65753, 12.43807] (Ceriaco et al. 2016a:21); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaco et al. 2016a:21); “Namibe Natural Park” [-15.77428, 12.33311] (Ceriaco et al. 2016a:21); “100 km southeast of Mogamedes” [-15.66515, 12.73503] (Laurent 1964a:42; Ceriaco et al. 2016a:58). Undetermined Locality: “Cuangu” (Boulenger and Power 1921:269) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distribntional notes: Map 248. Distribution of in Angola. Boulenger and Power (1921) refered to two specimens from “Caconda” and “Catumbella” in the British Museum collection sent by Bocage, stating that the “Caconda” material belonged to the type series. However, it is clear (Bocage 1896b, 1897a) that this cannot be the case. Specimens of A. anchietae can be confused with the similarly-sized ground dwelling congener ri. aculeata, so all older records should be reexamined. Agama congica Peters, 1877 Congo Agama Global conservation status (lUCN): Not Evaluated. Global distribution: The distribution of this species has not been established, but it is probably limited to the region of northern Angola and adjacent regions of central Africa. Ocurrences in Angola (Map 249): The species is known from northwestern Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:612; Loveridge 1957:191; Wolf¬ gang et al. 1997:313); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:42, 1887a:178, 1895a: 17). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:17, 18). Agama colonorum var. congica (Peters 1877a: 612). Syntypes: ZMB 9169 (3 speeimens) fide Bauer et al. (1995). Type loeality: “Chinehoxo,” Cabinda Provinee, Angola. Agama colonorum'. Boeage (1866a:42, 1895a: 17). Agama planiceps: Boeage (1887a: 178, 1895a:18, 1896a:110. Agama planiceps planieeps: Loveridge (1936a: 56). Agama eolonorum var. eongiea: Denzer et al. (1997:313). Map 249. Distribution of Agama congica in Angola. 290 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: Agama congica was described by Peters (1877) as a variety of Agama colonorum. On the basis of the recommendation of Philipp Wagner, we have rec¬ ognized A. congica as specifically distinct. The distribution and affinities of this taxon are current¬ ly under study. Ceriaco et al. (2014b) considered that specimens of Capanda Dam, Malanje Province, were true congica, but recent reanalysis suggests that these specimens should be consid¬ ered as representatives of an as yet undescribed endemic form (see Agama sp. account). Agama mucosoensis Hellmich, 1957 Mucoso Agama (Endemic) Agama agama mucosoensis (Hellmich 1957a:44, pi. 8). Holotype: ZSM 117/53 (collector W. Hellmich). Type locality: “Mucoso hei Dondo” [= Mucoso, Dondo] Kwanza Norte Province, Angola. Agama agama mucosoensis: Hellmich (1957h:50). Agama mucosoensis: Wagner et al. (2012:184), Ceriaco et al. (2014b:670). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is only known from Angola. Ocurrences in Angola (Map 250): The species is known only from the type locality and near areas in western Angola. Kwanza Norte: “Mucoso” [-9.53333, 14.65000] (Hellmich 1957a;44; Wagner et al. 2012. 18); “Dondo” [-9.66667, 14.41667] (Hellmich 1957b:50; Wagner et al. 2012. 18). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] (Hellmich 1957b:50; Wagner et al. 2012. 18). Taxonomic and distributional notes: Hellmich (1957a) described Agama agama mucosoensis based on a type series of 72 spec¬ imens collected in “Mucoso bei Dondo”, Kwanza Norte Province, recently revalidated by Wagner et al. (2012) as a full species. The only published data for this species are from Kwanza Norte Province, although specimens have recently been deposited in the Museu Map 250. Distribution in Angola. Nacional de Historia Natural e da Ciencia Lisboa, Portugal from “Agucareira’^Bengo) and “Quifangondo” (Luanda), Angola (L. Ceriaco and M. Marques, pers. obs.). Ceriaco et al. (2014b) also suggested that the species might occur in Malanje Province. There is a possibility of confu¬ sion between this species and A. congica, both of which exhibit sexual dimorphism, and all histor¬ ical records should be reexamined with this in mind. Agama planiceps Peters, 1862 Namib Rock Agama Agama planiceps Peters 1862a: 15. Syntypes: ZMB 4200-4201 (eolleetor C. H. Hahn). Type loeality: “Neu- Barmen” [= Gross Barmen, Otjozondjupa Region], Namibia. Agama planiceps: Sehmidt (1933:9), Ceriaeo et al. (2016a:21, 58). Global conservation status (lUCN): Not Evaluated. Global distribution: Agama planiceps occurs in the northwestern quadrant of Namibia and the southwestern quadrant of Angola. Ocurrences in Angola (Map 251): The species is known the true desert areas of Namibe province. Namibe: “Pico Azevedo” [-15.55000, 12.51667] (Schmidt 1933:9; Ceriaco et al. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 291 20I6a:58); “Lion Cave’s at 3.4 km SW of Espinheira camp” [-16.81250, 12.33969] (Ceriaco et al. 20I6a:2I-22); “Omauha Lodge camp” [-16.19872, 12.40008] (Ceriaco et al. 2016a:22); “Iona National Park, north of Tam- bor” [-15.99611, 12.40667] (Ceriaco et al. 2016a:22); “Iona National Park, south side of Curoca River crossing” [-16.30433, 12.41710] (Ceriaco et al. 2016a:22); “Pediva Hot Springs” [-16.12214, 12.56111] (Ceriaco et al. 2016a:22); “Namibe Lubango road, road mark¬ er 59, 1.8 km west by road from Caraculo, north side of the road” [-15.01600, 12.64356] (Ceriaco et al. 2016a;22); “Pico Azevedo” [-15.53400, 12.49197] (Ceriaco et al. 2016a;22). Taxonomic and distributional notes: Bauer et al. (1995) incorrectly gave the current name of the type locality of A. planiceps (Neu Barmen) as “Otjimbingue,” which is actually anoth¬ er Rhenish Mission site to the southwest of Neu Barmen. In Angola the species appears to be confined to the true desert areas in Namibe, being replaced by A. schacki in the northern and east¬ ern parts of the province. However, Namibian ri. planiceps occur broadly throughout arid and semi- arid areas where suitable rocky habitats occur. Monard’s (1937b) record of “ri. planiceps’’' from Cuando Cubango Province is certainly in error and is not included in our map. Map 25 1 . Distribution of Agama planiceps in Angola. Agama schacki Mertens, 1938 Schack’s Rock Agama (Endemic) Agama planiceps schacki (Mertens 1938a:433). Holotype: SMF 25299 (eolleetor W. Sehaek). Type loeality: “Cubal, 900m H. Prov. Benguella, Angola” [= Cubal], Benguela Provinee, Angola. Agama planiceps: Boeage (1887e:210, 1895a: 18, 1896a: 110), Boulenger (1885:358, 1905:110), Mertens (1926:152), Monard (1937b:60). Agama colonorum: Boulenger (1885:357), Ferreira (1905:117, 1906:170). Agama planiceps schacki: Hellmieh (1957a:47, 1957b:52), Mertens (1967:61), Mananas (1963:230), Laurent (1964a:40f Braneh (1998:218). Global conservation status (lUCN): Not Evaluated Global distribution: Agama schacki occurs in the southwestern quadrant of Angola. Ocurrences in Angola (Map 252): The species occurs along the southern areas of the escarp¬ ment (Benguela and Huila provinces), as well as in the lowlands of Benguela Province. Benguela: “Cahata” [-12.35000, 14.81667] (Boeage 1895a: 18); “Morro de Pundo” [-12.38333, 13.88333] (Parker 1936:129); “Quissange” [-12.43333, 14.05000] (Boeage 1887c;210, 1895a;18); “Quin- dumbo” [-12.46667, 14.93333] (Boeage 1895a:18); “Benguella” [-12.58333, 13.41667] (Boulen¬ ger 1885:357; Boeage 1895a;18); “Ebanga” [-12.73333, 14.73333] (Monard 1937b;60); “Lembu, Serra de Selles” [-12.86667, 14.11667] (Ferreira 1905:117, 1906:170); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:47); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:433, 1967:61; Hellmieh 1957b:52); “Marco de Canavezes” [-13.08333, 14.33333] (Laurent 1964a:40); “Hanha” [-13.30000, 14.20000] (Boeage 1896a;110); “Catengue” [-13.03333, 13.73333] (Mertens 1927:152). Huila: “Senhora do Monte, Sa da Bandeira” [-14.93333, 13.43333] (Laurent 1964a:40); “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] (Laurent 1964a;40). Namibe: 292 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Biballa” [-14.76667, 13.36667] (Bocage 1895a: 18; Ceriaco et al. 2016a;58). Taxonomic and distributional notes: Mertens (1938a) diagnosed A. planiceps schacki from the nominotypical form on the basis of a larger, more robust body, a greater number of midbody scale rows (80-90 vs. 63- 76), and a black (rather than red) tail tip. Branch (1998) stated that A. p. schacki replaces the typical form in Angola. Recently collected molecular and morphological data suggests that both forms occur in Angola and that planiceps and schacki, although closely related, are two distinct species. A detailed review of the group is being prepared. Records from Benguela (Boulenger 1885) and Lembu (Ferreira 1906, 1906) originally referred to Agama colonorum have been tentatively assigned to A. schacki on the basis of their distribution. Map 252. Distribution of Agama schacki in Angola. Serpentes Family Typhlopidae Merrem, 1820 Genus Afrotyphlops Broadley and Wallach, 2009 Afrotyphlops angolensis (Bocage, 1866) Angola Blind Snake Onychocephalus angolensis Bocage 1866a:46, 1866b:65. Holotype: MBLT79.1134 (collector F.A.P. Bayao), destroyed by fire on 18 March 1978. Type locality: “le district du Duque de Braganga” [= Calandula], Malanje Province, Angola. Onychocephalus Kraussii: Bocage (1873a:252). Onychocephalus angolensis: Bocage (1879b:95). Typhlops congicus: Bocage (1895a:63). Typhlops punctatus punctatus: Loveridge (1957:242). Typhlops angolensis adolfi: Laurent (1964a: 88). Typhlops angolensis: Laurent (1964c:424), Roux-Esteve (1974a:490; 1974b:46), McDiarmid et al. (1999:90), Spawls et al. (2004:288). Typhlops angolensis angolensis: Managas (1973:189) Rhinotyphlops angolensis: Broadley and Cotterill (2004:44), Chirio and LeBreton (2007:332). Afrotyphlops angolensis: Broadley and Wallach (2009:29), Ceriaco et al. (2014b:672), Wallach et al. (2014:13), Hedges et al. (2014:20). Global conservation status (lUCN): Not Evaluated. Global distribution: The species occurs in both forest and savanna from Cameroon south through the Congo Basin to northern Angola and northeast Zambia and east to Uganda and Kenya. Ocurrences in Angola (Map 253): The species occurs in the northeastern Angola. Kwanza Norte: “Canzele” [-8.30000, 15.18333] (Laurent 1964c:424). Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a:46, 1866b:65, 1873a:252; Loveridge 1957:242; Laurent 1964c:424; Roux-Esteve 1974a:490, 1974b:57; Broadley and Wallach 2009:29; Wallach et al. 2014:13). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964b:88; Roux-Esteve MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 293 I974b:57); “Cassange” [-9.58333, 17.86667] (Bocage I879b:95). Mexico: “Calombe, Luso” [-11.83333, 19.93333] (Managas 1973:189). Huambo: “Nova Lisboa” [-12.76667, 15.73333] (Roux-Esteve I974b:57). Taxonomic and distributional notes: The species was first named by Bocage (1866a), who provided a diagnosis and description, but the status of the species as new was indicated conditionally, as ''Onychocephalus angolensis Nov. sp.?” The name was used without such a caveat in a subsequent publication Bocage (1866b). Both publications appeared in the November 1866 issue of the same journal. See Roux-Esteve (1974b), McDiarmid et al. (1999) and Broadley and Wallach (2009) for compre¬ hensive chresonymies and details of distribu¬ tion elsewhere in Africa. 12*E liyE 20^ E Map 253 Distribution of Afrotyphlops angolensis in Angola. Afrotyphlops anomalus (Bocage, 1873) Angolan Giant Blind Snake (Endemic) Onychocephalus anomalus Bocage 1873a:248, pi. 1, fig. 3. Syntypes: MBL 1177A-D (four specimens), (col¬ lector J.A. d’Anchieta and J.J. da Graga), destroyed by fire 18 March 1978. Type locality: “Huilla, I’inte- rieur de Mossamedes” [= Huila], Huila Province, Angola. Typhlops {Onychocephalus) Anchietae Bocage 1886a: 172. Holotype: MBL 1871, (collector J. Anchieta), destroyed by fire 18 March 1978. Type locality: “Huilla” [= Huila], Huila Province, Angola. Typhlops anchietae-. Boulenger (1893:40, 1915:197), Bocage (1895a:63, 1897a:198), Monard (1937b:103). Typhlops anomalus: Boulenger (1893:47), Bocage (1895a:70), Ferreira (1897b:243) Typhlops anomalus: Monard (1937b: 103, 105). Rhinotyphlops anomalus: Roux-Esteve (1974a:495, 1974b: 193), McDiarmid et al. (1999:78). Megatyphlops anomalus: Broadley and Wallach (2009:48), Wallach et al. (2014:423). Afrotyphlops anomalus: Hedges et al. (2014:20). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 254): The species is restricted to the southwestern areas of the country. Cabinda: “Loango (?)” [-5.15000, 12.16667] (Boulenger 1893:47, 1915:197; Monard 1937b:103). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:103, 105; Broadley and Wallach 2009:48). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:70; Monard 1937b:103; Broadley and Wallach 2009:48); “Cahata” [-12.35000, 14.81667] (Bocage 1895a:70; Monard 1937b:103; Broadley and Wallach 2009:48); “Quindumbo [-12.46667, 14.93333] (Bocage 1895a: 70; Broadley and Wallach 2009:48); “Benguella” [-12.58333, 13.41667] (Bocage 1895a: 70; Boulenger 1915:197; Monard 1937b: 103); “Ebanga” [-12.73333, 14.73333] (Monard 1937b: 103, 105; Broadley and Wallach 2009:48). Huila: “Huilla” [-15.05000, 13.55000] (Bocage 1873b:248, 252, 1886a:172, 1895a:63, 70, 1897a:198; Monard 1937b:103; Roux-Esteve 1974b:194; Broadley and Wallach 2009:48; Wallach et al. 2014:423); “Caconda” [-13.73333, 15.06667] (Bocage 1895a:70; Ferreira 1897:243; Monard 1937b:103; Roux-Esteve 1974b:194; Broadley and Wallach 2009:48); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:103, 105; Broadley and Wallach 2009:48). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a:70; Monard 294 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1937b: 103; Broadley and Wallach 2009:48); “Chiyaka distriet” [vie. -15.08333, 12.73333] (Roux-Esteve 1974b: 194; Broadley and Wal- laeh 2009:48); “Mossamedes” [-15.20000, 12.15000] (Boeage 1873b:248; Boulenger 1893:47). Taxonomic and distributional notes: Hedges et al. (2014) moved all speeies previ¬ ously allocated to Megatyphlops Broadley and Wallach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embed¬ ded within a clade containing the type speci¬ men of the latter. Boulenger (1893, 1915) identified a specimen from “Loango,” a former kingdom within Cabinda Province as M. anomalus. However, this is certainly a misidentification or error in locality, as Roux- Esteve (1974a) demonstrated that this species 1 'l m \ A ^ N Map 254. Distribution of Afrotyphlops anomalus in Angola. was only reliably known from the mountainous regions of Benguela and Huila, south of the Cuan- za River. Wallach et al. (2014) also listed Bie and Cuanza Sul in the range of the species, but this does not appear to be based on previously published records. McDiarmid et al. (1999) included northern Namibia in the stated distribution of this species, but we are not aware of any confirmed specimens outside of Angola. See Broadley and Wallach (2009) for chresonymy and maps of global distribution. Afrotyphlops lineolatus (Jan, 1864) Common Lined Blind Snake Typhlops (Ophthalmidion) lineolatus Jan 1864:24. Holotype, UUMZ 725 (collector A. Afzel), lost fide Roux- Esteve (1974b) and Hahn (1980). Type locality: “Sierra-Leona” [= Sierra Leone]. Typhlops boeagei Ferreira 1904:114. Syntypes: MBL specimen numbers not known (collector F. Newton) (destroyed July 1943 during WWII fide Broadley and Wallach 2009:40). Type locality: “Cabicula, Bom Jesus (margens do Quanza)” [= Bom Jesus], Bengo Province, Angola. Typhlops Boulengeri Boeage (1893:117). Lectotype: BMNH 1946.1.11.18, formerly BMNH 1893.12.27.12 (collector J. Anchieta) designated by Broadley and Wallach (2009:40). Type locality: “Quindumbo, dans Finterieur de Benguella” [= Quindumbo], Benguela Province, Angola. Onychoeephalus liberiensis (part): Boeage (1866a:46, 1873a:252). Onychoeephalus lineolatus'. Boeage (1873a:252, 1866b:65). Typhlops esehriehtii (part): Gunther (1876b:678). Typhlops {Ophthalmidion) Esehriehtii var. lineolata: Peters (1877a:614). Typhlops {Ophthalmidion) Esehriehtii var. intermedia'. Peters (1877a:614). Typhlops {Ophthalmidion) Esehriehtii (part): Peters (1881:147). Typhlops {Ophthalmidion) Kraussii (part): Boeage (1887a: 180). Typhlops lineolatus'. Boulenger (1893:43), Roux-Esteve (1974b:76), McDiarmid et al. (1999:107), Spawls et al. (2004:290). Typhlops punctatus var. intermedia (part): Boeage (1895a:66). Typhlops punetatus var. lineolata'. Boeage (1895a:66). Typhlops punetatus var. lineolatus'. Boeage (1896a: 112), Ferreira (1903:9). Typhlops punetatus (part): Boulenger (1900a:50, 1905:112), Monard (1937b:103, 104), Themido (1941:9). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 295 Typhlops boulengeri: Bocage (1895a:64, 1897a:198), Ferreira (1900a:50, 1906:167), Boulenger (1915:196), Monard (1937b: 103, 104), Laurent (1964a:89). Typhlops punctatus punctatus (part): Parker (1936:120), Mertens (1937a: 11, 1938a:438), Laurent (1950a:7), Hellmieh (1957a:70), Loveridge (1957:242). Typhlops bocagei: Monard (1937b: 103). Typhlops boulengeri boulengeri: Laurent (1964e:414). Typhlops lineolatus lineolatus: Roux-Esteve (1974a:492). Rhinotyphlops lineolatus: Broadley and Cotterill (2004:45). Rhinotyphlops lineolatus lineolatus: Chirio and LeBreton (2007:338). Afrotyphlops lineolatus: Wallaeh and Broadley (2009:41), Wallaeh et al. (2014:14), Hedges et al. (2014:20). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is widely distributed in the savannas of sub-Saharan Africa from Senegal east to Sudan and Ethiopia and south to northern Tanzania, Katanga (Democratic Republic of Congo) and Angola as far south as Benguela. Ocurrences in Angola (Map 255): The species occurs mainly in western Angola, how¬ ever there are records in Lunda Norte Province. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a;614); “Landana” [-5.21667, 12.15000] (Roux-Esteve 1974b:104). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:66, 1887a: 180; Monard 1937b: 103; Roux-Esteve 1974b: 104). Lunda Norte: “riviere Muari (afluente de la Luachi- mo), dans le environs de Dundo” [-7.28333, 20.93333] (Laurent 1950a:7); “Dundo” [-7.36667, 20.83333] (Laurent 1950:7, 1964a:89); “Muita” [-7.80000, 21.45000] (Laurent 1950a:7). Lunda Sul: “Mutianvo” [-11.45000, 19.33333] (Themido 1941:9). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1873a:252; Bocage 1895a:66; Monard 1937b: 103); “Cassange” [-9.58333, 17.86667] (Bocage 1895a:66; Monard 1937b:103); “Malanje” [-9.55000, 16.35000] (Peters 1881:147). Bengo: “Cabicula, Bom Jesus (margem do Quanza)” [-9.16667, 13.56667] (Ferreira 1904:114; Monard 1937b:103; Loveridge 1957:242). Kwanza Norte: “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:112; Monard 19376:103; Roux-Esteve 1974b:104); “Cam- bondo” [-9.15963, 14.65827] (Ferreira 1906:167; Monard 1937b:103, 104); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:167; Monard 1937b: 103, 104; Wallaeh and Broadley 2009:41); “N’dalla Tando, Cazengo” [-9.30000, 14.91667] (Ferreira 1903:9; Monard 19376:103, Roux-Esteve 19746:104; Wallaeh and Broadley 2009:41). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:120); “Congulu” [-10.86667, 14.28333] (Parker 1936:120). Benguela: “Quindumbo” [-12.46667, 14.93333] (Bocage 1893:117, 1895a:64, 1897a:198; Loveridge 1933:216, 1957:241; Monard 19376:103, 104; Laurent 1964a:89, 1964c:414; Roux-Esteve 19746:104; Wallaeh and Broadley 2009:41); “Benguella” [-12.58333, 13.41667] (Boulenger 1915:196; Loveridge 1957:242); “Ebanga” [-12.73333, 14.73333] (Monard 19376:103, 105); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:70); “Cubal” [-13.03333, 14.25000] (Mer- Map 255. Distribution of Afrotyphlops lineolatus in Angola. 296 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II tens I937a:II, I938a:438); “Hanha” [-13.30000, 14.20000] (Bocage I896a:II2; Roux-Esteve 1974b: 104). Undetermined Locality: (Gunther 1876b:678; Ferreira 1900:50);. “Cuango” (Peters 1881:147) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distributional notes: There was long confusion between this taxon and Afrotyphlops punctatus (Leach, 1819) as reflected in the complex synonymies/chresonymies of these two taxa (McDiarmid et al. 1999). Authorship of this species is sometimes attributed to Jan (1863) (e.g., McDiarmid et al. 1999), however, this use is as a nomen nudum. See Broadley and Wallach (2009) for comprehensive chresonymy and distribution in eastern and central Africa. Afrotyphlops mucruso (Peters, 1854) Zambezi Blind Snake Onychocephalus mucruso Peters 1854:621. Leetotype: ZMB 3963 (eolleetor W.C.H. Peters), designated by Loveridge (1933:216). Type loeality: “Macanga” (Peters 1854:621), [= Makanga], Mozambique. Typhlops mucruso: Boulenger (1893:46), Boeage (1895a:67). Typhlops schlegelii mucruso: Loveridge (1933:216, 1957:241). Typhlops schlegeli mucruso: Laurent (1950a:7, 1964a:90), Rhinotyphlops schlegelii dinga (part): Roux-Esteve (1974b: 164). Typhlops sehlegelii mueruso: Broadley (1990:47). Rhinotyphlops sehlegelii (part): MeDiarmid et al. (1999:84). Rhinotyphlops mueruso: Spawls et al. (2004:292), Broadley and Cotterill (2004:45). Megatyphlops mueruso: Broadley and Wallaeh (2009:52), Bates et al. (2014:313), Wallach et al. (2014:423). Afrotyphlops mueruso: Hedges et al. (2014:20). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the savannas and forest-savanna mosaic from East Africa, from coastal Kenya south to central Mozambique, Zimbabwe and Limpopo Province, South Africa, and west through the southern provinces of the Demo¬ cratic Republic of Congo to northeast Angola. Ocurrences in Angola (Map 256): The species occurs in the extreme northeast of the country, in Lunda Norte and Sul provinces. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:7; Roux-Esteve 1974b: 178); “Muita” [-7.80000, 21.45000] (Laurent 1950a:7; “Camisombo” [-8.15000, 20.65000] (Laurent 1964a:89); “Calonda” [-8.41667, 20.53333] (Laurent 1964a:89). Lunda Sul: Map 256. Distribution of Afrotyphlops mucruso in “Alto Chicapa” [-10.88333, 19.23333] (Lau- Angola, rent 1964a: 89). Taxonomic and distributional notes: See Roux-Esteve (1974b), Bauer et al. (1995, 2002) and McDiarmid et al. (1999) for discussions of the original type series of Onychocephalus mucru¬ so. Hedges et al. (2014) moved all species previously allocated to Megatyphlops Broadley and Wal¬ lach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embedded within a clade containing the type specimen of the latter. Previ¬ ously treated as a subspecies, Rhinotyphlops schlegelii mucruso (Peters, 1854), but elevated to full i2‘L 2«''b MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 297 species status and placed in the new genus Megatyphlops by Broadley and Wallach (2009). The status of large typhlopids in Angola previously assigned to Megatyphlops by Broadley and Wallach (2009) as mucruso and schlegelii remains problematic. See Broadley and Wallach (2009) for comprehensive chresonymy and maps of global distribution. Afrotyphlops schlegelii (Bianconi, 1849) Schlegel’s Giant Blind Snake Typhlops schlegelii Bianconi 1849:183, pi. 6, figs. 2, 2a-2d. Holotype: MZUB specimen numbers not known (eolleetor C. Fomasini). Type loeality: “Mozambieo” [=Mozambique] (Inhambane, Mozambique fide Roux-Esteve 1974b). Typhlops (Onyehoeephalus) humbo Boeage 1886a: 171. Syntypes: MBL 1887 [2 speeimens], destroyed by fire 18 Mareh 1978 (eolleetor J.A. d’Anehieta). Type loeality: “Quissange” Benguela Provinee, Angola. Onyehoeephalus Petersii Boeage 1873a:248, pi. 1, fig. 3. Syntypes: MBL 1868 (2 speeimens), destroyed by fire 18 Mareh 1978 (eolleetor J. Anehieta). Type loeality: “Biballa” [= Bibala] Namibe Provinee, Angola. Onyehoeephalus Sehlegelii: Boeage (1873a:250). Typhlops Petersii'. Boeage (1886a: 172, 1895a:68, 1897a:199). Typhlops {Onyehoeephalus) humbo Boeage 1886a: 171. Syntypes: MBL 1887 (2 speeimens) (eolleetor J. Anehieta), destroyed by fire 18 Mareh 1978. Type loeality: “Quissange” Benguela Provinee, Angola. Typhlops humbo'. Boeage (1887e:210, 1895a:66, 1897a: 198), Boulenger (1893:46). Typhlops hottentotus: Boeage 1893:117. Holotype: MBL 1867 (eolleetor J. Anehieta), destroyed by fire 18 Mareh 1978. Type loeality: “Humbe” Cunene Provinee, Angola. Typhlops hottentotus'. Boeage (1895a:69, 1897a: 198), Boulenger (1896:588). Typhlops sehlegeli mueruso: Loveridge (1933:216, 1957:241), Mertens (1938a:438). Typhlops mueruso (= humbo, hottentotus, petersii)'. Monard (1937b: 103). Rhinotyphlops sehlegelii petersii'. Roux-Esteve (1974a:495; 1974b: 166), Braneh (1998:54). Typhlops sehlegelii petersii'. Broadley (1990:48). Rhinotyphlops sehlegelii (part): MeDiarmid et al. (1999:84). Megatyphlops sehlegelii'. Broadley and Wallaeh (2009:48), Bates et al. (2014:314), Wallaeh et al. (2014:423). Afrotyphlops sehlegelii'. Hedges et al. (2014:20) Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southern Mozambique, northern KwaZulu- Natal to Swaziland, through the northeastern provinces of South Africa, central Botswana, northern Namibia and southwestern Angola. Ocurrences in Angola (Map 257): The species occurs in southwestern Angola. Benguela: “Quissange” [-12.43333, 14.05000] (Boeage 1886a:171, 1887c:210, 1895a:66, 1897a; 198; Boulenger 1896:588; Loveridge 1933:216; Monard 1937b: 103; Roux-Esteve 1974:495, 1974b; 180; Broadley and Wallach 2009:51); “Benguella” [-12.58333, 13.41667] (Boulenger 1893:46), “Caimbambo” [-13.01667, 14.01667] (Broadley and Wallach 2009:51); “Cubal” [-13.03333, 14.25000] (Mertens 1938:438). Huila: “Caconda” [-13.73333, 15.06667] (Boeage 1895a;68; Monard 1937b: 103); “10 km SE of Joao de Almeida” [-15.18333, 13.68333] (Broadley Map 257. Distribution of Afrotyphlops schlegelii in and Wallach 2009:51). Namibe: “Biballa” 12*E lU'E irE 24°E 298 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-14.76667, 13.36667] (Bocage I873a:249, 252, I886a:I72, I895a:68, I897a:I99; Loveridge 1933:216; Monard 1937b: 103). Cunene: “Humbe, sur les bords du Cunene (Humbe)” [-16.68333, 14.90000] (Bocage 1893:117, I895a:69; Boulenger 1896:588; Monard I937b:I03; Broadley and Wallach 2009:51); “Erickson’s Drift, Cunene river” [-17.26944, 14.525 E] (Broadley and Wallach 2009:51). Taxonomic and distributional notes: There are three juvenile specimens referable to this species that are extant in the Bianconi collection of Mozambiquan reptiles in Bologna, but none of these seem to be consistent with the type specimen. The date of publication is often confused because the description appeared in the journal issue for August and September 1848, but was not published until April 1849, and because Bianconi (1850:183, pi. 6, fig. 2) published a second description, marked “Nobis” of the same animal. Further, Bianconi (1849) notes that he had described this species, and another, in the session of 1847, but this refers to the oral sessions of the Societa Agraria, e dell’ Accademia delle Scienze dell’ Istituto di Bologna, for which the Nuovi Annali delle Scienze Naturali was the publication outlet. “Hedges et al. (2014) moved all species previously allocated to Megatyphlops Broadley and Wallach, 2009 to Afrotyphlops based on their phylogenetic analysis, which demonstrated that members of the former group are embedded with¬ in a clade containing the type specimen of the latter. Until recently, four subspecies were recog¬ nized under the name Rhinotyphlops schlegelii {brevis, schlegelii, petersii, mucruso) following Roux-Esteve (1974b) and Hahn (1980). Currently Onychocephalus petersii (Bocage, 1873) is placed in the synonymy of Afrotyphlops schlegelii (Bianconi, 1849), whereas the others are recog¬ nized as valid species (Roux-Esteve 1974a; 1974b; Broadley 1990; Branch 1998; Broadley and Wallach 2009). The complex nomenclatural history of this and related taxa, including the interpre¬ tation of type specimens has been reviewed by McDiarmid et al. (1999). See Broadley and Wallach (2009) for comprehensive chresonymy and maps of global distribution. Afrotyphlops schmidti (Laurent, 1956) Schmidt’s Blind-Snake Typhlops schmidti Laurent 1956:71, figs. 9-11, pi. 8, fig. 4. Holotype: MRAC 17996 (collector not men¬ tioned). Type locality: “Nyunzu, Terr. D ’Albertville, Tanganika” [= Nyunza, Tanganyika Province (for¬ merly Katanga Province), Democratic Republic of Congo fide Wallach and Broadley 2009:43]. Typhlops schmidti: Laurent (1964a: 89), McDiarmid et al. (1999:119). Rhinotyphlops schmidti: Broadley et al. (2003:42), Broadley and Cotterill (2004:45) Afrotyphlops schmidti: Broadley and Wallach (2009:43), Wallach et al. (2014: 15), Hedges et al. (2014:20). Global conservation status (lUCN): Not Evaluated. Global distribution: A savanna species distributed in eastern Angola, through northern Zambia and southern Democratic Republic of Congo (Katanga provinces). Ocurrences in Angola (Map 258): The species occurs in eastern Angola. Moxico: “environs du lac Calundo, village du chef Sa- Mussamba (environs du lac Calundo)” 12°E \(CL 2rE 24°E Map 258. Distribution of Afrotyphlops schmidti in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 299 [-11.80000, 20.86667] (Laurent I964a:89; Broadley and Wallach 2009:43); “Cazombo” [-11.88333, 22.91667] (Laurent I964a;89; Broadley and Wallach 2009:43). Taxonomic and distributional notes: The type locality of Typhlops schmidti in the Tangani- ka district (now province) of the Democratic Republic of Congo, has been misinterpreted by some subsequent authors to refer to Tanzania (e.g., McDiarmid et al. 1999). Laurent (1964a) was the first to cite this species from Angola. See Broadley and Wallach (2009) for chresonymy and maps of global distribution. Genus Letheobia Cope, 1868 Letheobia praeocularis (Stejneger, “1893” 1894) Leopoldville Beaked Snake Typhlops praeocularis Stejneger “1893” 1894:709. Holotype: USNM 20799 (eolleetor J.H. Camp). Type loeality: “Leopoldville, or Stanley Pool, Congo Free State” [= Kinshasa or Maleho Pool], Demoeratie Republie of Congo. Typhlops praeocularis lundensis Laurent 1964a:90. Holotype: MD 5929 (eolleetor “Indigenes divers” = various native eolleetors). Type loeality: “Dundo, Lunda,” Lunda Norte, Angola. Rhinotyphlops praeocularis: Roux-Esteve (1974a:498, 1974b:204), MeDiarmid et al. (1999:83). Letheobia praeocularis: Chirio and LeBreton (2007:328), Broadley and Wallaeh (2009:66), LeBreton (2010), Wallaeh et al. (2014:373), Hedges et al. (2014:21). Laurent (1964a) described Typhlops praeocu¬ laris lundensis based on five specimens from “Dundo,” Angola, later considered a synonym of the typical form (Roux-Esteve 1974a, 1974b). Broadley and Wallach (2007a) revived the genus Letheobia from the synonymy of Rhinotyphlops Fitzinger, 1843, but did not include L. praeocularis in their study. Map 259. Distribution of Letheobia praeocularis in Angola. Global conservation status (lUCN): Least Concern. Global distribution: The species is known from western and central Africa, rang¬ ing from Southern Congo to southwestern Democractic Republic of Congo throughout northeastern Angola and probably Nigeria. Ocurrences in Angola (Map 259): The species occurs in northeastern Angola. Lunda Norte: “Dundo, Lunda” [-7.36667, 20.83333] (Laurent 1964a:90; Roux-Esteve 1974a:498, 1974b:204). Taxonomic and distributional notes: 20^ E 2-(°E Family Leptotyphlopidae Stejneger, 1892 Genus Lepto typhlops Fitzinger, 1843 Leptotyphlops kafubi (Boulenger, 1919) Shaba Thread Snake Glaucoma kafubi Boulenger 1919b:186. Syntypes: BMNH 1946.1.11.7-8, formerly BMNH 1919.8.26.13-14, MRAC 2085 (eolleetor L. Stappers). Type loeality: “la riviere Kafuboo, a Elisabethville” [= Kafubu River, Lubumbashi, Haut-Katanga Provinee (formerly Katanga Province)], Democratic Republic of Congo. Leptotyphlops emini emini (part): Laurent (1964a:91). 300 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Leptotyphlops nigricans nigricans (part): Broadley and Watson (1976:490). Leptotyphlops nigricans (part): McDiarmid et al. (1999:39). Leptotyphlops kafubi: Broadley and Broadley (1999:18), Broadley and Wallaeh (2007b: 11), Wallaeh et al. (2014:368). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southeastern Africa, from eastern Angola, the former Katanga Province, Democratic Republic of Congo to Zambia, Malawi and Zimbabwe. Ocurrences in Angola (Map 260): The species has been recorded from eastern Ango¬ la. Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:91; Broadley and Watson 1976:490; Broadley and Broadley 1999:18); “Chutes de la Luisavo, Cabinda, Haul Zambeze” [-11.86667, 23.58333] (Laurent 1964a:91; Broadley and Watson 1976:490; Broadley and Broadley 1999:18). Taxonomic and distributional notes: Broadley and Watson (1976) provided a map with the Angolan records from Laurent (1964a) assigned to Leptotyphlops nigricans (Schlegel, 1839). They described the species range of L. nigricans as discontinuous, with the nominate population occurring in the southern Cape Province (now Western and ^60. Distribution of Leptotyphlops kafubi in Eastern Cape provinces. South Africa), with a small group of locality records in the central portion of the former Transvaal Province separated by a wide gap from the remainder of the distribution, which extends north from central Zambia through eastern Angola to Uganda and South Sudan (Broadley and Watson 1976). Broadley and Broadley (1999) formally diagnosed the L. nigricans group, including the southern African species L.jacobseni Broadley and Broadley, 1999, L. kafubi (Boulenger, 1919) andZ. nigricans. They also restricted L. nigricans to the southern Cape provinces of South Africa and used L. kafubi as a new combination and attributed to it the Angolan records previously assigned to Leptotyphlops emini emini. Leptotyphlops scutifrons (Peters, 1854) Peters’ Thread Snake Stenostoma scutifrons Peters 1854:621. Holotype: ZMB 4826, lost fide Broadley and Watson (1976), Hahn (1980) and Bauer et al. (1995, 2002) (eolleetor W.C.H. Peters). Type loeality: “Sena” (Peters 1854:621), [= Vila da Sena, Zambezi River], Mozambique. Stenostoma seutifrons (part): Peters (1865:261), Boeage (1873b:251, 1895a:71). Stenostoma nigrieans (part): Boeage (1866a:46). Glaueonia seutifrons (part): Ferreira (1904:114), Boulenger (1905:112, 1915:198), Monard (1937b: 106). Leptotyphlops eonjuneta distanti (part): Bogert (1940:13). Leptotyphlops seutifrons seutifrons (part): Broadley and Watson (1976:497), Broadley and Broadley (1999:23). Leptotyphlops seutifrons (part): MeDiarmid et al. (1999:41). Leptotyphlops seutifrons'. Bates et al. (2014:321), Wallaeh et al. (2014:364). Leptotyphlops seutifrons eomplex: Conradie et al. (2016:23). I2*E ki*E 20*E 24*E MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 301 Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from western and eentral Angola, across Zambia and Malawi to southern Tanzania and Mozambique, and south through Namibia, Botswana and Zimbabwe to Swaziland and KwaZulu-Natala and the Eastern Cape in South Africa. Ocurrences in Angola (Map 261): The n-E io‘e za-e species occurs in western Angola. Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a;46; 1873b:251, 1895a:71; Monard 1937b: 106). Kwanza Norte: “Golun- go Alto” [-9.13333, 14.76667] (Ferreira 1903:114; Monard 19376:106); “Zembe” [-9.31667, 14.66667] (Ferreira 1904:114; Monard 1937b: 106); “Cazengo” [-9.33333, 14.76667] (Ferreira 1904:114; Monard 1937b:106). Bie: “Chitau” [-11.25000, 17.16667] (Bogert 1940:13; Broadley and Broadley 1999:26); “Cubango basin (13)” [-13.28061, 16.74722] (Conradie et al. 2016:8- 9, 23). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b: 106). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:71; Monard 1937b: 106); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:106). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:71; Monard 1937b:106); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:106) “Kapelongo” [-14.88333, 15.08333] (Monard 1937b:106); “Huilla (Huila)” [-15.05000, 13.55000] (Bocage 1895a:71; Monard 1937b:106). Namibe: “Biballa (Bibala)” [-14.76667, 13.36667] (Bocage 1873b:251, 1895a:71; Monard 1937b:106); “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:71; Monard 1937b:106). Taxonomic and distributional notes: Bauer et al. (1995) incorrectly regarded ZMB 5706 as a possible syntype. Adalsteinsson et al. (2009) recently identified numerous undescribed cryptic species within the L. scutifrons/incognitus/conjunctus complex that will require detailed analysis to stabilize the taxonomy of the group (Bates et al. 2014). The generic map for Leptotyphlops provided by Adelsteinsson et al. (2009) shows no species in Angola except in far southeastern Cuando Cubango and in eastern Mexico. Some of the specimens referenced in the works cited above may be referable to Namibiana latifrons, so surviving material must be examined. Map 261. Distribution of Leptotyphlops scutifrons in Angola. Genus Namibiana Hedges, Adalsteinsson and Branch, 2009 Namibiana labialis (Sternfeld, 1908) Damara Thread Snake Glaucoma labialis Sternfeld 1908:92. Holotype: ZMB 21691 (eolleetor O. Dempwolff). Type loeality: “Outgo (D. S. W. Afriea)” (Sternfeld 1908:92), [= Outjo, Kunene Region], Namibia. Leptotyphlops labialis Broadley and Broadley (1999:14), McDiarmid et al. (1999:34). Namibiana labialis'. Adalsteinsson et al. (2009:13), Wallaeh et al. (2014:464). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola and northern Nambia. Ocurrences in Angola: The species is known from southern Angola, in Cunene Province near the Namibian border. Cunene: “Miranda” (Broadley and Broadley 1999:14). 302 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: Most authors considered the holotype of this taxon to be lost or unidentified (e.g., McDiarmid et al. 1999; Bauer et al. 2002), however, ZMB 21691, although not marked as a type in the ZMB catalogue, is fully consistent with the description. Adal- steinsson et al. (2009) found deep genetic divergence between Leptotyphlops occidentalis and other species of Leptotyphlops. On this basis and on evidence from shared morphology they designated a new genus, Namibiana for members of the Leptotyphlops rostrata (Bocage, 1866) group {sensu Broadley and Broadley 1999), to which L. occidentalis belongs. Namibiana latifrons (Sternfeld, 1908) Benguela Thread Snake Glaucoma latifrons Sternfeld 1908:93. Leetotype: BMNH 64.6.14.5 (eolleetor J.J. Monteiro) designated by Broadley and Watson (1976). Type loeality: “Benguela”, Benguela Provinee, Angola. Stenostoma nigricans (part): Boeage (1867b:224). Glaucoma scutifrons (part): Boulenger (1893:68). Stenostoma scutifrons (part): Peters (1865:262), Boeage (1895a:71). See notes below regarding Peters’ (1865) applieation of names. St{enostoma\ scutatum Peters (1865:262), unavailable name (see notes below). Glaucoma latifrons'. Monard (1937b: 106). Leptotyphlops seutifrons (part): Bauer et al. (1995:77, 2002:169), MeDiarmid et al. (1999:41). Leptotyphlops latifrons'. Broadley and Broadley (1999:14). Namibiana latifrons'. Adalsteinsson et al. (2009:13), Wallaeh et al. (2014:464). Peters (1854) described Stenostoma scutifrons from the type locality “Sena” in Mozambique. Boulenger (1893) later allocated two sped- mens from “Benguella” to this taxon. Sternfeld Distribution ^fNamihima lalifmns in Angola. (1908) recognized that the east and west African forms were distinctive from one another and pro¬ posed the name latifrons for the Angolan material. Boundy (2014) made the case that Peters him¬ self had validly proposed the name Stenostoma scutatum Peters (1865) for the same Benguela material examined by Boulenger (1893), despite the fact that Peters ultimately concluded that the Mozambiquan and Angolan specimens were conspecific and assignable to S. scutifrons. However, because Peters’ name had not been used subsequently and Glaucoma latifrons Sternfeld, 1908 had Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from western Angola and southwestern Demo¬ cratic Republic of Congo (Bas-Congo). Ocurrences in Angola (Map 262): The species occurs in western Angola. Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1867b:224; 1895a:71; Monard 1937b: 106). Benguela: “Lobito Bay” [-12.35000, 13.55000] (Broadley and Broadley 1999:14); “Catumbella” [-12.43333, 13.55000] (Bocage 1867b:224; 1895a:71; Monard 1937b: 106; Broadley and Broadley 1999:14); “Benguella” [-12.58333, 13.41667] (Peters 1865:621, Boulenger 1893:68; Bauer et al. 1995:77; Broadley and Broadley 1999:14; Wallaeh et al. 2014:464). Taxonomic and distributional notes: MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 303 been in continuous use for over a century, Boundy (2014) argued that the former name should be treated as a nomen oblitum and the later as a nomen protectum, although he did not use those terms. Although Boundy (2014) invoked Article 1 1 of the International Code of Zoological Nomenclature to support his interpretation of the availability of Peters’ name for this taxon, we disagree. Article 11.5 states “To be available, a name must be used as valid for a taxon when proposed, unless it was first published as a junior synonym and subsequently made available under the provisions of Arti¬ cle 11.6.1.” As S. scutatum was first proposed as a synonym and was not treated as an available name prior to 1961, it is unavailable under the Code. Bauer et al. (1995) incorrectly included “Benguella [Angola] (Westafrika)” as part of the type locality for S. scutifrons (Bauer et al. 2002). Adalsteinsson et al. (2009), transferred latifrons to the new genus Namibiana. It is possible that some specimens from western Angola currently assigned to Leptotyphlops scutifrons may represent this species, and such material should thus be carefully rexamined. Namibiana rostrata (Bocage, 1886) Angolan Beaked Thread Snake (Endemic) Stenostoma rostratum: Bocage 1886a: 173. Holotype: MBL specimen number unknown (collector J. d’ Anchi¬ eta), destroyed by fire 18 March 1978. Type locality: “envoye du Humbe, sur les bords du Cunene” [= Humbe] Cunene Province, Angola. Stenostoma rostratum: Bocage (1895a:71, 1897a: 199). Glaucoma rostrata: Boulenger (1915:198). Leptotyphlops rostratus: Broadley and Broadley (1999:18), McDiarmid et al. (1999:40). Namibiana rostrata: Bates (2010), Wallach et al. (2014:464). Global conservation status (lUCN): Data Deficient. Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 263): The species is known from western portions of the country, from the type locality, “Humbe” north to “Luanda.” Malanje: “Vana e Lumbe” (Broadley and Broadley 1999:18). Luanda: “Luanda” [-8.83333, 13.26667] (Broadley and Broadley 1999:18). Huila: “W of Huila” [-15.05000, 13.55000] (Broadley and Broadley 1999:18). Cunene: “envoye du Humbe, sur les bords du Cunene (Humbe)” [-16.68333, 14.90000] (Bocage 1886a:173, 1895a:71, 1897a: 199; Broadley and Broadley 1999:18; Wallach et al. 2014:464). Undetermined Locality: “without precise location” (Bocage 1895a:71; Boulenger 1915:198; Broadley and Broadley 1999:18). Taxonomic and distributional notes: Broadley and Broadley (1999) provided some Map 263. Distribution of Namibiana rostrata in Angola. new data for the species in Angola, increasing the distribution range further north from the type locality. The generic map for Namibiana provided by Adelsteinsson et al. (2009) shows no species occurring north of Kwanza Sul. 304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family Pythonidae Fitzinger, 1826 Genus Python Daudin, 1803 Python anchietae Bocage, 1887 Anchieta’s Dwarf Python Python Anchietae BocagQ 1887d:87. Syntypes: MBLT89-1206, T89-1207 (collector J. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Catumbella” [= Catumbela] Benguela Province, Angola. Python Anchietae: Bocage (1895a:73, 1897a:199). Python anehietae: Boulenger (1893:88,1915:199), Monard ( 1 93 7b : 1 08), Bogert ( 1 940 : 1 8), Frade (1963:253), Laurent (1964a:92), Branch (1998:59), Wallach et al. (2014:607). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Angola and central and northern Namibia. Ocurrences in Angola (Map 264): The species is known from the southwestern Ango¬ la regions of Angola. Benguela: “Hanha road, 18 km from Lobito” [-12.26667, 13.70000] (Laurent 1964a:92); “Catumbella” [-12.43333, 13.55000] (Bocage 1887d:87, 1895a:73, 1897a:199; Boulenger 1893:88, 1915:199; Monard 1937b: 108; Wallach et al. 2014:607); “Hanha” [-12.25000, 13.75000] (Bogert 1940:18). Taxonomic and distributional notes: This species was described by Bocage (1887d) based on two specimens from “Catumbella” collected by Anchieta. Anchieta’s Dwarf Python is poorly recorded in the country and the current gaps in its distribution should be considered artifactual. The lUCN map of its distribution {http://maps.iucnredlist.org/map. ^ html?id= 177539) is entirely incorrect for all but the southernmost part of the species range. Python natalensis Smith, 1840 Southern African Rock Python Python sebae natalensis Smith 1840: pi. 9, first of three aeeompanying unnumbered text pages. Syntypes: BMNH 1946.1.8.3 (formerly 1940.3.27.1) and 1946.1.17.13 (formerly 1940.3.27.2) (fide Broadley 1984). Type loeality: none explieitly stated, although it is “not to be found within hundreds of miles of the bound¬ aries of the [Cape] Colony, and few speeimens have been obtained nearer than Port Natal. “ Given the speeifie epithet, the terra typieal may be interpreted as Port Natal. Python natalensis: Boeage (1895a:72, 1896a: 112), Braneh (1998:59), Broadley (1999:31), Broadley and Cotterill (2004:45), Bates et al. (2014:328), Ceriaeo et al. (2014b:672, 2016a:40), Wallaeh et al. (2014:609), Conradie et al. (2016:23). Python sebae: Monard (1937b: 108), Mertens (1938a:439), Bogert (1940:17, 18), Themido (1941:9), Maeha- do (1979:10, 46), Spawls and Braneh (1995:19), Bellosa et al. (2007:30). Python sebae natalensis: Broadley (1984:362). Global conservation status (lUCN): Not Evaluated. Global distribution: The species ranges across south-central Africa from Angola southeast to South Africa and north to the southeastern Democratic Republic of Congo, Burundi, Tanzania and the Kenya highlands. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 305 Map 265. Distribution of Python natalensis in Angola. Namibe: “Maconjo” [-15.01667, Ocurrences in Angola (Map 265): Pub¬ lished Angolan records are mostly from south¬ ern and western parts of the country. Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1895a:72; Broadley 1984:364). Mexico: “Moxico” [-11.85000, 20.06667] (Machado 1979:10). Benguela: “Benguella” [-12.58333, 13.41667] (Bocage 1895a:72; Broadley 1984:364); “sighting in Ebanga” [-12.73333, 14.73333] (Monard 1937b:108); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:439); “Hanha” [13.30000, 14.20000] (Bocage 1896a:112; Themido 1941:9), “Hanha (North)” [-12.25000, 13.75000] (Bogert 1940:17). HmTa: “Galange” [-13.80000, 16.11667] (Monard 1937b:108); “Vila-da- Ponte” [-14.46667, 16.30000] (Monard 1937b: 108); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:17; Broadley 1984:364). 13.20000](Bocage 1895a:72; Broadley 1984:364); “Rio Giraul” [-15.06833, 12.14222] (Bocage 1896a: 112; Broadley 1984:364); “Beginning of the forested areas, at the start of the climb to Leba Pass (by road), near Bruco village” [-15.12106, 13.18654] (Ceriaco et al. 2016:40). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937b:108; Broadley 1984:364); “sighting in Forte Rogadas” [-16.73333, 14.98333] (Monard 1937b: 108). Cuando Cubango: “Cubago basin (1)” [-16.89413, 17.95766] (Conradie et al. 2016:8-10, 23); “Cuito basin (30a)” [17.50875, 20.06594] (Conradie et al. 2016:9, 10, 23). Taxonomic and distributional notes: Smith (1833) first used the name Python Natalensis as a nomen nudum, giving the locality “from the interior, eastwards of Latakoo [and] in the country about Port Natal” [= east of Kuruman, Northern Cape, South Africa and around Durban, KwaZu¬ lu-Natal], which is given as the type locality by Wallach et al. (2014). Interestingly, Smith (1840) cited the first use of the name as being on page 64 of the South African Quarterly Journal for Octo¬ ber 1833. This is indeed a page in an Andrew Smith paper, but the paper deals exclusively with mammals and page 64 treats elephant shrews specifically (Broadley 1984). The same citation to the original description was provided by McDiarmid et al. (1999). Stimson (1969) considered the terra typica unknown and the types untraced. Wallach et al. (2014) likewise considered that no type had been designated, but considered the specimens to have been associated with the “Cape Town Zool. Soc. Mus.” Although Smith was associated with the nascent South African Museum, the bulk of his herpetological material that has survived was deposited either in London or Edinburgh (FitzSimons 1937; Branch and Bauer 2005). Broadley (1984) indicated that both syntypes were rolled skins preserved in alcohol. The BMNH register notes both from Port Natal, with BMNH 1946.1.8.3 donated by Andrew Smith and BMNH 1946.1.17.13 without a stated collector and noted as “found as stuffed specimen in dry coll.” Python natalensis was for a long time considered as a synonym or subspecies of Python sebae (Gmelin, 1789) (Broadley 1984), but was elevated to specific status by Broadley (1999) based on morphological differences as well the evidence of the overlap in distribution (Broadley and Cotterill 2004). Broadley (1984), Spawls and Branch (1995) and Bellosa et al. (2007) provided maps with the distribution ranges for both species, in which Python natalensis occurs in central and 306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II south Angola, bordered in the north by the Kwanza River and overlaping in Luanda Province with Python sebae, which occurs in northern regions of Angola, including the enclave of Cabinda. Python sebae (Gmelin, 1789) African Rock Python Coluber Sebae Gmelin 1789:1118. Syntypes: specimen described and illustrated by Seba (1735:105, pi. 99, fig. 2) and specimen described by Gronovius (1756:56) Type locality: “Brasiliensis, e regione Guaira” [= Guira, Brazil], Brazil. Python Sebae: Bocage (1866a:47, 1867b:224). Python sebae: Peters (1877:614), Laurent (1954a:38, 1964a:91), Hellmich (1957a:70), van den Audenearde (1966:32), Machado (1979:10), Haacke (1982b:8), Spawls and Branch (1995:19), Bellosa et al. (2007:30). Python sebae sebae: Broadley (1984:362), Broadley (1999:31), Wallach et al (2014:611). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is very widespread in sub-Saharan Africa, from southern Mauritania to southern Chad, South Sudan and Ethiopia, south to Tanzania and Rwanda with is southern limit in northern Angola and the southern Democratic Republic of Congo. Ocurrences in Angola (Map 266): The species distribution is limited to the northern regions of the country. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614). Bengo: “Nambuangongo” [-7.97438, 14.18924] (Machado 1979:10); “Ambriz” [-7.844312, 13.106493] (Broadley 1984:362). Kwanza Norte: “Rio Cuanza, nahe Mucoso” (Hellmich 1957a: 70). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a;38, 1964a:91; Thys van den Audenaerde 1966:32; Broadley 1984:362). Lunda Sul: “Saurimo” [-9.65000, 20.40000] (Haacke 1982b:8). Undetermined Locality: “Without precise location” (Bocage 1866a:47, 1867b:224; Lau¬ rent 1964a;91). Taxonomic and distributional notes: Gmelin (1789) gave two indications to older Map 266. Distribution of Python sebae in Angola. works in his description of Coluber Sebae: “Gron. Mus. 2. N. 11” and “Seb. mus. 2. t. 199. f. 2,” although he provides only one set of scale counts; 272 ventrals and 70 caudals, corresponding to Gronovius ’s specimen. This implies that there were two original syntypes, not a single holotype as incorrectly stated by McDiarmid et al. (1999). Loveridge (1957) refered to the specimen figured by Seba as the type and Wallach et al. (2014) cited this specimen as the lectotype. However, as no pre-2000 work appears to have explicitly recognized this as a lectotype designation, both speci¬ mens must still be considered syntypes. The Seba specimen has been stated to be in Uppsala (Stim- son 1969; McDiarmid et al. 1999), but its actual whereabouts are unknown. The history of Seba’s collections have been discussed in detail (Engel 1937, 1961; Boeseman 1970; Juriev 1981; Adler 1989; Bauer 2002; Bauer and Gunther 2013). Seba’s second collection (the first had been sold to Peter the Great of Russia in 1716; Driessen-van het Reve 2006) was sold after his death at auction (Anonymous 1752). Seba specimens are known or believed to be present in collections in St. Petersburg, London, Leiden (including specimens until recently in Amsterdam), Paris, Copen¬ hagen, Stockholm, Bremen and Berlin (Boeseman 1970; Juriev 1981; Thireau et al. 1998; Bauer MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 307 and Gunther 2013) but few can be traced to particular plate figures, and the lectotype of Python sebae is not among these. Likewise, the specimen noted by Gronovius (1756) is also untraceable. His collection was dispersed as a result of an auction held in Leiden in 1778 (Wheeler 1958). Gronovius refered to Seba’s plate in his own work and repeated the locality “Brasiliensis, e regione Guaira,” however, his measurements and scale counts were based on a specimen in his own collection. FitzSimons (1962) and McDiarmid et al. (1999) incorrectly gave the type locality as “America” and the former author corrected this to “America.” The extraordinary confusion over the type specimens and localities of both P. natalensis and P. sebae suggest that most modem authors have relied on secondary sources for their information. Spawls and Branch (1995:19) and Bellosa et al. (2007) provided a map with the distribution ranges for both large species of Angolan Python in which P. sebae is limited to the northern regions of Angola from northern Mexico to Luanda, where it overlaps with Python natalensis, which occurs in central and south Angola. Family Boidae Gray, 1825 Genus Calabaria Gray, 1858 Calabaria reinhardtii (Schlegel, 1851) Calabar Ground Boa Eryx reinhardtii Schlegel 1851:1, pi. Holotype: ZMB 1471 (eolleetor H. Halleur, don. J.T. Reinhardt). Type loeality: “possessions danoises a la Cote d’or” (Sehlegel 1851:2), [= Ghana], restrieted to “Aqua Pirn” [= Akwapim], Ghana by Hughes and Barry (1969:1010) on the basis of data assoeiated with speeimen. Calabaria Reinhardtii: Boeage (1895a:74). Calabaria reinhardtii: Frade (1963:252). Calabaria reinhardtii: Chirio and LeBreton (2007:364), Wallaeh et al. (2014:130). Charina reinhardtii: MeDiarmid et al. (1999:203). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from West and Central Africa, from Guinea to Cameroon and Central African Republic, south to the Democratic Republic of Congo. Ocurrences in Angola (Map 267): The species occurs in the Cabinda enclave. Cabin¬ da: “Cabinda” [-5.55000, 12.18333] (Frade 1963:252). Taxonomic and distributional notes: Stimson (1969) and McDiarmid et al. (1999) considered the type untraced. Bauer et al. (2002) demonstrated that the specimen was in Berlin, not in Leiden as many had supposed. Although Stimson (1969) provided the correct year of description (1851) he provided the wrong volume of the Bijdragen tot de Dierkunde (vol. 1). Subsequent authors have cited both the wrong year (1848) and the wrong volume number (vol. 1 instead of vol. 3) (McDiarmid et al. 1999; Wallaeh et al. 2014). Most recent literature does not include “Cabinda” in the distribution of Calabaria Distribution of Calabaria reinhardUi in rem/iarr/tn (McDiarmid et al. 1999; Chirio and LeBreton 2007; Wallaeh et al. 2014), however, the proximity and biotic affinities of Cabinda to West and Central Africa account for its presence there. 308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family Viperidae Oppel, 1811 Genus Athens Cope, 1862 Athens squamigera (Hallowell, “1854” 1855) Variable Bush Viper Echis squamigera Hallowell “1854” 1855:193. Holotype: ANSP 6949 (collector M.P. Bellonni-Duchaillu). Type locality: “Near the river Gaboon, Guinea” (Hallowell “1854” 1855:193), Gabon River, Gabon. Athens squamigera: Peters (1881:150), Bocage (1887a:189, 1895a:152), Laurent (1954a:62), Loveridge (1957:304), Frade (1963:252), Spawls and Branch (1995:99), Broadley (1998a: 128), Ernst and Rodel (2002:55), Chirio and LeBreton (2007:596), Dobiey and Vogel (2007:100), Wallach et al. (2014:62) Athens Lueani: Rochebmne (1885:89). Athens squamiger Boulenger (1905:114, 1915:222), Ferreira (1906:169). Athens squamigera squamigera (Hallowell): Laurent (1954a:62), Hellmich (1957b:76), Managas (1981:39). ? Athens squamigera squamigera (Hallowell): Laurent (1964a: 128). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is endemic to West and Central Africa. Ocurrences in Angola (Map 268): The species occurs in northern Angola, including the Cabinda enclave. Cabinda: “Foz do Luan- go” [-5.15000, 12.16667] (Mana9as 1981:39); “Landana” [-6.21667, 12.15000] (Rochebmne 1885:90, Loveridge 1957:304, Mananas 1981:39). Lunda Norte: “Riv. Kundueji, envi¬ rons de Dundo, Lunda” [ -7.36667, 20.83333] (Laurent 1964a: 128); “Dundo, rives de la Lua- chimo” [-7.38333, 20.85000] (Laurent 1954a: 62; Managas 1981:39); “foret de la Lua- chimo” [-7.38333, 20.85000] (Laurent 1964a: 128). Kwanza Norte: “Piri-Dembos” [-8.53333, 14.43333] (Hellmich 1957b:76; Managas 1981:39); “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:114; Ferreira 1906:169; Mananas 1981:39). Undetermined Locality: “Cuango” (Peters 1881:150; Bocage 1887a:189; 1895a:152; Managas 1981:39) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distributional notes: The date of the description of this taxon has variably been given as 1854, 1855 (Wallach et al. 2014), or 1856 (McDiarmid et al. 1999; Broadley 1998a). Although the contents of volume 7 of the Proceedings of the Academy of Natural Sciences of Philadelphia covers the years 1854 and 1855 and the printed cover page indicates 1856, part V of the volume, in which Hallowll’s paper appeared, was printed in 1855. Several taxonomic revisions of the genus Atheris Gray, 1842 were undertaken by Broadley (1996a, 1998a), but the taxonomic status of several entities remain to be established (Ernst and Rodel 2002). A re-examination of museum specimens is necessary to determine if Atheris squamigera (Hallowell, “1854” 1855) is a geographically variable species or a species complex (Broadley 1998a; B. Hughes in Ernst and Rodel 2002). 12*E I^E 24°E Map 268. Distribution of Atheris squamigera in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 309 Genus Bids Gray, 1842 Bids arietans (Merrem, 1820) Puff Adder Vipera {Echidna) arietans Merrem 1820:152. Leetotype: speeimen deseribed and illustrated by Seba (1735:55, pi. 54, fig. 4), now lost (eollector unknown), designated by Wallaeh et al. (2014:61 1). Type loeal- ity: “Vorgeburge der guten Hoffnung/Promontorio bonae spei” (Merrem 1820:152), [= Cape of Good Hope] Western Cape Provinee, Southern South Afriea. Echidna arietans: Boeage (1866a: 53). Clotho arietans: Gunther (1865a:480). Echida arietans: Boeage (1879e:89). Bids arietans: Boeage (1887a:190, 1887e:211), Ferreira (1897b:245), Boulenger (1915:221), Sehmidt (1933:15), Loveridge (1936a:45), Monard (1937b: 141-142), Themido (1941:11), Hellmieh (1957a:75), Spawls and Braneh (1995:114), Braneh (1998:114), Lenk et al. (1999:31), Dobiey and Vogel (2007:104), Barlow et al. (2013:1134), Wallaeh et al. (2014:89), Braneh and Conradie (2015:200), Ceriaeo et al. (2016a:87), Conradie et al. (2016:23). Vipera arietans: Boeage (1895a: 149, 1896a: 113). Cobra lachesis: Mertens (1937a: 16). Bids lachesis: Mertens (1938a:442), Bogert (1940:99), Laurent (1950a: 11, 1954a:62). Bids arietans arietans: Hellmieh (1957b:74), Laurent (1964a: 127), Thys van den Audenaerde (1966:36), Mananas (1981-82:35), Branch and McCartney (1992:2), Bates et al. (2014:331). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from most of sub-Saharan Afriea, extending to the Arabian Peninsula. Ocurrences in Angola (Map 269): The speeies oeeurs in the entire eountry with exeeption of the desert regions of the far south¬ west, as well as some mountainous areas. Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Boeage 1887a:190, 1895a;149; Managas 1981:35). Bengo: “Ambriz” [-7.844312, 13.106493] (Managas 1981:35). Kwanza Norte: “Mucoso, Dondo” [-9.53333, 14.65000] (Hellmieh 1957a:75; Mananas 1981:35). Malanje: “Duque de Bragan^a” [-9.10000, 15.95000] (Boeage 1866a:53, 1895a: 149; Monad 1937b: 141; Managas 1981:35); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016a:87). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:ll, 1954a:62, 1964a: 127; Mananas 1981:35); “Capaia” [-7.33556, 20.21681] (Braneh and Conradie 2015:200); “Muita, Luembe E” [-7.80000, 21.45000] (Laurent 1950a: 11, 1954a:62; Managas 1981:35); “Sombo” [-8.68333, 20.95000] (Laurent 1954:62); “Maludi” [-8.03333, 21.33333] (Thys van den Audenaerde 1966:36). Lunda Sul: “Alto Chieapa” [10.93333, 19.15000] (Laurent 1964a:127; Managas 1981:35). Moxico: “environs du lae Calundo (Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a:127; Managas 1981:35). Bie: “Chitau” [-11.25000, 17.16667] (Sehmidt 1933:15; Bogert 1940:99; Managas 1981:35); “Cuban¬ go basin (9)” [-14.00269, 17.40500] (Conradie et al. 2016:8-9, 23). Huambo: “Bela Vista” [-12.56667, 16.21667] (Managas 1981:35; Hellmieh 1957b:74); “Huambo” [-12.83333, 15.66667] Map 269. Distribution of Bids arietans in Angola. 310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (Bogert 1940:99; Themido 1941:11; Mananas 1981:35); “Cuma” [-12.86667, 15.06667] (Loverid- ge 1936a:45; Managas 1981:35). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1887c:211); “Benguella” [-12.58333, 13.41667] (Boeage 1866a:53); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:75; Managas 1981:35); “Cubal” [-13.03333, 14.25000] (Mertens 1937:16, 1938:442; Managas 1981:35); “Equimina” [-13.20000, 14.78333] (Bocage 1895a:149; Managas 1981:35); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113; Mananas 1981:35). Hmla: “Capelongo” [-14.91667, 15.08333] (Bogert 1940:99; Mana9as 1981:35); “San- geve” [-13.88333, 15.83333] (Monard 19376:141-142; Managas 1981:35); “Rio Calae” [-13.96667, 16.03333] (Bocage 1879c:89, 1895a:149; Monad 1937b:141; Managas 1981:35); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:141, 142; Managas 1981:35); “Kampulu” [-15.21667, 16.11667] (Monard 19376:141-142; Mananas 1981:35); “Rio Cabindon- go” (Bocage 1895a:149; Monad 19376:141; Managas 1981:35). Namibe: “Mossamedes” [-15.20000, 12.15000] (Gunther 1865a:480; Managas 1981:35). Cunene: “ruisseau Mbale (Bale)” [-15.16667, 16.75000] (Monard 1937b:141-142; Managas 1981:35). Cuando Cubango: “approx¬ imately 33 km ESE of Cuito Cuanavale” [-15.28333, 19.50000] (Branch and McCartney 1992:2); “Cuito basin (61)” [-16.98919, 19.40614] “Cuando basin (38)” (not collected) [-17.58830, 22.65694] (Conradie et al. 2016:8, 10, 23). Undetermined Locality: “from almost all localities that he [J. Anchieta] visited” (Bocage 1895a: 149). Taxonomic and distributional notes: Merrem (1820) had indications to numerous earlier works, including Seba (1735) and Gronovius (1756). In addition, his scale counts (ventrals + cau- dals), though similar to those given by Gronovius are not exactly the same, suggesting that he made scale counts on at least one specimen himself Merrem had access to some of Seba’s original spec¬ imens through the collections of Willem Xaver Janssen (1760-1793) and Friedrich Heinrich Graf von Borcke (1776-1825), successive owners of Seba specimens originally sold at auction (Anony¬ mous 1752) (Bauer and Gunther 2013). Surviving B. arietans from this collection (ZMB 2879) does not correspond to the Seba image indicated by Merrem, although it may have been examined by him and could have been part of the basis for his description. Wallach et al. (2014) appear to be the first to have explicitly designated a lectotype for the puffadder. The species has significant phylogeographic structure (Lenk et al. 1999; Barlow et al. 2013; Bates et al. 2014) and the implications for its systematics status are being investigated. Currently, two subspecies are recognized: Bitis arietans arietans (Merrem, 1820) widespread in sub-Saharan Africa and Bitis arietans somalica Parker, 1949 restricted to Somalia (Branch 1998; Bates et al. 2014). Some authors (e.g., Spawls and Branch 1995; Vogel and Dobiey 2007) considered B. ari¬ etans widespread in Angola although with no occurences in northwest regions of the country, whereas Wallach et al. (2014) only cited localities in the eastern regions of Lunda Norte and Mex¬ ico Provinces. See McDiarmid et al. (1999) for a chresonomy of the species. Bitis caudalis (Smith, 1839) Horned Adder Vipera ocellata Smith 1838:92. Syntypes: BMNH 1865.5.4. 153a-e (collector A. Smith). Type locality: “sandy districts north of the Cape Colony,” South Africa. Junior primary homonym of Vipera ocellata Latreille, 1801 [= Vipera aspis hixmdieus, 1758]. Vipera {Cerastes) caudalis Smith 1839: pi. 7, first of two accompanying unnumbered text pages. Syntypes: BMNH 1865.5.4. 153a-e (collector A. Smith). Type locality: “sandy districts north of the Cape Colony,” South Africa (verbatim from Smith 1838). Nomen substitutum for Vipera ocellata Smith, 1838. Cerastes caudalis: Bocage (1867c:227, 1870:68). Vipera caudalis: Gunther (1865a:480), Bocage (1895a: 150). Bitis caudalis: Boulenger (1915:221), Frade (1963:253), Haacke (1984:174), Mananas (1981:37), Visser MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 3II (1981:7), Spawls and Branch (1995:105), Branch (1998:116), Dobiey and Vogel (2007:110), Bates et al. (2014:334), Wallaeh et al. (2014:90). Bitis caudalis caudalis: Laurent (1964a: 128). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from the arid western region of southern Africa, throughout Karoo and Kalahari, from southern Zimbabwe and the western half of South Africa to southern Angola - the northern limit for the species. Ocurrences in Angola (Map 270): The species occurs in the southwestern Angola, with preference of hot and dry open areas in sandy regions. Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1895a: 150; Managas 1981:37). Benguela: “Benguela” [-12.58333, 13.41667] (Mananas 1981:37). Huila: “Huila” [-15.08333, 13.55000] (Mananas 1981:37). Namibe: “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:150; Monard 1937b:141; Mana9as 1981:37); “Mossamedes” [-15.20000, 12.15000] (Gunther 1865a:480; Bocage (1867c:227; Managas 1981:37); “Desert de Mogamedes, a 35 km au sud de la ville” [-15.20000, 12.15000] (Laurent 1964a:128); “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:150; Monard 1937b:141; Managas 1981:37). Undetermined Map 270. Distribution of Bitis caudalis in Angola. Locality: “with no precise location” (Bocage 1870:68). Taxonomic and distributional notes; Although Smith’s (1838) Vipera ocellata was preoccu¬ pied, Smith (1839) noted only “As it does not appear probable that the genis Vipera will ultimate¬ ly be divisible into subgenera, I have thought it advisable to substitute for ocellata. — a term which under such circumstances would cause some confusion, — that of caudalis.'" Preliminary phylo- geographic analysis indicates appreciable regional divergence (A. Barlow et al. unpubished data in Bates et al. 2014). Visser (1981) provided a map with the Transvaal Museum accessions since 1962 for Bitis caudalis. Haacke (1984) subsequently provided a map with some records for the species in Namibe Province, without explicit locality information, however the two maps are very similar and probably represent the same data, showing the southwestern distribution of the species in the country. The Angolan record from “Loanda/Luanda” (Bocage 1895a; Mana9as 1981) is doubtful and certainly represents a misidentification. Bitis gabonica Dumeril, Dumeril and Bibron, 1854 Gabon Adder Echidna Gabonica Dumeril, Dumeril and Bibron 1854:1428, pi. 80 bis, figs. 1-3. Leetotype: MNHH 4012 (eolleetor C.-E. Aubry-Leeomte), designated by Hughes and Barry (1969:1030). Type loeality: “eote du Gabon” (Dumeril, Dumeril and Bibron 1854:1428), Gabon. Echidna rhinoceros'. Boeage (1866a: 53). Vipera (Bitis) rhinoeeros: Peters (1877a:618). Bitis rhinoeeros'. Boeage (1887a: 191). Vipera rhinoeeros'. Boeage (1895a: 149, 1896a:113, 1897b:211). Bitis gaboniea: Boulenger (1915:222), Laurent (1950a:ll), Spawls and Braneh (1995:116-117), Lenk et al. 312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (1999:31), Lenk et al. (2001:94), Chippaux (2006:260), Chirio and LeBreton (2007:602), Dobiey and Vogel (2007:111), Wallach et al. (2014:90), Oliveira et al. (2016:42). Bitis gabonica gabonica: Laurent (1954a:62, 1964a: 128), Loveridge (1957:302), Thys van den Audenaerde (1966:36), Managas (1981:36). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from forest-savanna mosaie of West and Central Africa from Benin to Angola and is very widespread in the Democratic Republic of Congo, extend¬ ing marginally into adjacent areas such as southern Nigeria and northern Zambia, with isolated populations in South Sudan, Kenya, and eastern southern Africa, the southern most occurring in northeastern KwaZulu-Natal. Ocurrences in Angola (Map 271): The species occurs in northern regions of Angola including Cabinda enclave and the northeast¬ ern areas of Moxico Province. Cabinda: “Chinchoxo” [-5.48333, 12.13333] (Peters 1877a;618; Bocage 1895a; 149; Managas 1981:36); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:53, 1887a:191, 1895a:149; Mananas 1981:36). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a:191, 1895a:149; Managas 1981:36). Malanje: “Cacuso” [-9.4994, 15.1613] (Oli¬ veira et al. 2016:42); “Kalandula” [-91956, 15.9026] [-9.2097, 15.8906] [-91907, 15.9159] [-9.3872, 15.9441] (Oliveira et al. 2016:42). Kwanza Norte: “N’dala Tando” [-9.30000, 14.91667] (Mananas 1981:36). Benguela: “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113, 1897b:211; Mananas 1981:36); “Caimbambo” [-13.1677, 13.7663] (Oliveira et al. 2016:42). Moxico: “environs du lac Calundo (Lago Calundo)” [-11.71667, 20.80000] (Laurent 1964a: 128; Managas 1981:36). Lunda Norte: “Dundo” [-7.3667, 20.8267] (Oliveira et al. 2016:42). Taxonomic and distributional notes: McDiarmid et al. (1999) incorrectly cited MNHN 4020 as a holotype specimen. The description was based on a skin sent by Aubry-Lecomte and a speci¬ men from the Menagerie du Museum. Angolan specimens of Bitis gabonica have occasionally been cited as Bitis rhinoceros (Schlegel, 1855) — a closely related species endemic to West Africa (Lenk et al. 1999; Lenk et al. 2001; Chipaux 2006; Bates et al. 2014; Wallach et al. 2014). Map 271. Distribution of Bitis gabonica in Angola. Bitis heraldica (Bocage, 1889) Angolan Adder (Endemic) Vipera heraldica Bocage 1889:127. Holotype: MBL 2127 (eolleetors H.C. Capello and R. Ivens), destroyed by fire 18 Mareh 1978. Type loeality: “sur les bords de la riviere Calae, Pun des affluents du Cunene” [= Calai River banks] Huila Province, Angola. Vipera heraldica: Boeage (1895a: 151). Bitis peringueyi: Boulenger (1896:495, 1905:114, 1915:221), Monard (1937b:143), Bogert (1940:101). Bitis heraldica: Ferreira (1897b:245), Hellmieh (1957b:75), Mertens (1958:146), Mananas (1981:38), Spawls and Braneh (1995:104), Dobiey and Vogel (2007:113), Wallach et al. (2014:91). Global conservation status (lUCN): Not Evaluated. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 313 Global distribution: The species is known from western Angola, chiefly in association with the high plateau. Ocurrences in Angola (Map 272): The species is known from the central highlands in Angola. Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Managas 1981:38). Bengo: “Bengu” [-8.71667, 13.40000] (Monard 1937b:141; Mertens 1958:147; Managas 1981:38; Spawls and Branch 1995:105; Wallach et al. 2014:91). Kwanza Sul: “Mombolo” [-12.16667, 14.83333] (Bogert 1940:101; Mertens 1958:146; Mana- gas 1981:37-38). Huambo: “Bela Vista (San- guengue)” [-12.56667, 16.21667] (Hellmich 1957b:75; Mertens 1958:146; Mana9as 1981:38; Spawls and Branch 1995:105). Bie: “Bie” [-12.38333, 16.95000] (Mertens 1958:147; Managas 1981:38; Spawls and Branch 1995:105) Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:151; Map 272. Distribution of Bitis heraldica in Angola. Ferreira 1897b:245; Monard 1937b:141; Mertens 1958:147; Managas 1981:38; Spawls and Branch 1995:105); “Kalukembe (Caluquembe)” [-13.78333, 14.68333] (Monard 1937b:141, 143; Mertens 1958:147; Mananas 1981:38; Spawls and Branch 1995:105); “sur les bords de la riviere Calae, I’un des affluents du Cunene (Rio Calae)” [-13.96667, 16.03333] (Bocage 1889:127, 1895a:151; Monard 1937b:141; Mertens 1958:147; Managas 1981:38; Spawls and Branch 1995:105). Unde¬ termined Locality: “Between Benguella and Bihe” (Boulenger 1905:114; Monard 1937b: 141). Taxonomic and distributional notes: This species was described by Bocage (1889) based on one specimen from “sur les bords de la riviere Calae, Tun des affluents du Cunene”. This species was later considered by Boulenger (1896) a synonym of Bitis peringueyi (Boulenger, 1888) although Ferreira (1897) doubted this conclusion. Later, Mertens (1958) restablished B. heraldica based on distinctive morphological characters between the two species and refered Angolan mate¬ rial from “Kalukembe” (Monard, 1937b) and from “Namba, Mombolo” (Bogert, 1940) to B. heraldica. Bitis nasicornis (Shaw, 1792) Rhinoceros Viper Coluber Nasicomis: Shaw in Shaw and Nodder 1792: pi. 94, first of three aeeompanying unnumbered pages of Latin deseription). Holotype: BMNH, lost fide Golay et al. (1993) (donated by E. Jenkins). Type loeal- ity: “interiori Afriea” [“likely to be from Ghana” fide Hughes and Barry 1969:1030]. Bitis nasieornis: Boulenger (1915:222), Parker (1936:126), Hellmieh (1957b:76), Loveridge (1957:302), Mananas (1981:36), Spawls and Braneh (1995:118), Lenk et al. (1999:31), Lenk et al. (2001:96), Chirio and LeBreton (2007:604), Dobiey and Vogel (2007:117), Wallaeh et al. (2014:91). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is restricted to the rainforest of West Africa, known from Guinea to Gabon, extending east to Uganda, western Kenya, and northern Tanzania, having its southern limit in the northern regions of Angola and Zambia. Wallach et al. (2014) considered records from Spawls et al. (2004) from northeastern Tanzania to be unconfirmed. Ocurrences in Angola (Map 273): The species is limited to the northwestern of the country. 314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II including the Cabinda enclave, although there are no published records from the latter area. Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b;76). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:126, Managas 1981:36). Taxonomic and distributional notes: This species is often attributed to Shaw (1802), where the species is description is largely repeated, or to Shaw and Nodder (1792) (e.g., Wallach et al. 2014). As noted by McDiarmid et al. (1999), evidence suggests that Shaw alone is responsible for the description. Shaw (1792) specifically stated that the specimen had been presented to the British Museum from Edward Jenkins, who had obtained it from “the master of a Guinea vessel. Hoser (2013) erect- ed a new taxon, B. hoserae, lor B. nasicorms ^ east of the Dahomey Gap, but did not mention Angola or Angolan populations explicitly. Bitis peringueyi (Boulenger, 1888) Peringuey’s Adder Vipera peringueyi Boulenger 1888:141. Holotype: SAM 1852 (eolleetor L.A. Peringuey), lost fide Stander in Golay et al. (1993). Type loeality: “Damaraland, 10 miles east of Walfiseh Bay” [= 10 miles east of Walvis Bay], Erongo Region, Namibia. Bitis peringueyi: Managas (1981:37), Spawls and Braneh (1995:108), Braneh (1998:119), Dobiey and Vogel (2007:118), Wallaeh et al. (2014:92). Global conservation status (lUCN): Least Concern. Global distribution: The species occurs limited to sandy regions of the Namib Desert, in two disjunct populations corresponding to the northern and southern sand seas of the Namib. In Namib¬ ia from Liideritz north to Roikop, and then from northern Namibia northward to Namibe Province, southern Angola. Ocurrences in Angola (Map 274): The species is limited to the Namib Desert in Namibe Province. Namibe: “Porto Alexandre” [-15.80000, 11.83333] (Spawls and Branch 1995). Taxonomic and distributional notes: Boulenger (1896) considered Bitis heraldica (Bocage, 1889) to be a synonym of B. peringueyi which resulted in an incorrect distribution range for the species in Angola (see Bitis heraldica account). The species was cited for southern Angola by several authors (Manias 1981, Branch 1998, Dobiey and Vogel 2007, Wallach et al. 2014), although the local¬ ity “Porto Alexandre”, currently known as Map 274. Distribution of ftto/iermgwej/ in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 315 “Tomboa,” is the first explicit published record for the species in the country (Spawls and Branch 1995). Wallach et al. (2014) erroneously considered B. peringueyi present in Kwanza Sul Province, presumably based on Bogert’s (1940) record of 5. heraldica (as “5. peringueyi”). Genus Causus Wagler, 1830 Causus bilineatus Boulenger, 1905 Two-Striped Night Adder Causus rhombeatus var. bilineatus Boulenger 1905:114. Syntypes: BMNH 1905.5.29.36-40 (eolleetor W.J. Ansorge). Type loeality: “Duque de Braganea, Quissange, Caeonda et Huilla” (Boeage 1895a: 146) [= Calandula, Quissange, Caeonda and Huila] Angola, and “Between Benguela and Bihe” (Boulenger 1905:114), Angola. Causus rhombeatus-. Boeage (1895a:147); Boulenger (1896:468); Sehmidt (1933:15), Hellmieh (1957b:74), Loveridge (1957:299). Causus bilineatus bilineatus'. Laurent (1964a: 125). Causus bilineatus'. Spawls et al. (2004:469), Broadley (1968b:407), Spawls and Braneh (1995:140), Broadley and Cotterill (2004:46), Rasmussen (2005:1), Mananas (1981:33), Dobiey and Vogel (2007:125), Wallaeh et al. (2014:148). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the plateau of South-Central Africa, in Ango¬ la, southern Democratic Republic of Congo and north-west Zambia, reaching Tanzania. Ocurrences in Angola (Map 275): The species occurs from west to eastern Angola. Kwanza Norte: “Piri-Dembos” [-8.53333, 14.43333] (Hellmieh 1957b;74; Manaqas 1981:33; Rasmussen 2005:14); “Canhoca” [-9.25000, 14.68333] (Boulenger 1905:114; Manaqas 1981:33). Malanje: “Duque de Bra- ganqa” [-9.10000, 15.95000] (Boeage 1866a:51, 1895a;145; Monard 1937b;141, Manaqas 1981:33; Rasmussen 2005:14). Moxico: “Rives du lac Calundo (Lago Calun- do)” [-11.80000, 20.86667] (Laurent 1964a;125; Manaqas 1981:33; Rasmussen 2005:14). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:15; Rasmussen 2005:14). Huila: “Caeonda” [-13.73333, 15.06667] (Boeage 1895a:145; Monard 1937b:141; Mana?as 1981:33; Rasmussen 2005:14); Map 275 Distribution of 6, in Angola. “Capelongo” [-14.88333, 15.08333] (Manapas 1981:33); “Huila” [-15.08333, 13.55000] (Boeage 1895a:145; Boulenger 1896:468; Monard 1937b:141; Manaqas 1981:33; Rasmussen 2005:14). Taxonomic and distributional notes: Boeage (1895a) in his remarks on Causus rhombeatus (Lichtenstein, 1823) recognized a striped form based on several individuals from Angola deposit¬ ed in the Museu Boeage collection from “Duque de Braganqa, Quissange, Caeonda et Huilla,” although he did not recognize them as taxonomically distinct. Boulenger (1905) subsequently noted the same striped form described by Boeage based on specimens from “between Benguella and Bihe” and proposed the name C. rhombeatus var. bilineatus. McDiarmid et al. (1999) includ¬ ed Boeage ’s (1895a) specimens among the type series, but Boulenger (1905) clearly refered only to the striped specimens collected by Ansorge and in the BMNH collection. For some time, the 316 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II nomen bilineatus was neglected (e.g., Schmidt 1933; Monard, 1937b; Bogert, 1940; Hellmich 1957a,b) and the corresponding Angolan records were associated with C rhombeatus. Laurent (1964a) revived the name bilineatus, and included his recently described Causus lineatus (Laurent, 1956), which he considered a dwarf subspecies of Causus bilineatus. However, his specimens differ little from those of the type series from “Benguela to Bihe” (Broadley and Cotterill 2004). Loveridge (1957) included both bilineatus and lineatus in the synoymy of C rhombeatus, but Broadley (1968b) placed C b. lineatus in the synonymy of C bilineatus without comment, where it has since remained (Rasmussen 2005). Laurent (1964a) erroneously assigned some Angolan records of C. rhombeatus to C bilineatus (e.g., “Pungo-Andongo” [Boulenger 1905]; “Mombolo” [Bogert 1940]; “Bela Vista”, “Entre Rios” [Hellmich 1957b]) as did Managas (1981). Most of the specimens cited by Laurent (1964a) were examined and revised by Rasmussen (2005). Causus lichtensteinii (Jan, 1859) Forest Night Adder A[spidelaps] (Sepedon) Lichtensteinii Jan 1859:511. Holotype: NMBA, lost (fide Kramer 1978; Golay et al. 1993), destroyed in 1943 during the World War II fide Wallaeh et al. (2014). Type loeality: “Cote-d’Or” [= Ghana]. Causus liehtensteinii: Laurent (1950a:ll, 1964a:127), Mananas (1981:34), Spawls and Braneh (1995:143), Spawls et al. (2004:471), Rasmussen (2005:11), Chirio and LeBreton (2007:606), Dobiey and Vogel (2007:126), Akani et al. (2012:192), Wallaeh et al. (2014:149). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from the central African forests from Guinea and Liberia eastward through Ghana to Nigeria, Cameroon, the Central African Republic, south to Democratic Republic of Congo, northeastern Angola, and northwestern Zambia, and east to Uganda and western Kenya. Ocurrences in Angola (Map 276): The species occurs in the northtem Angola, in the adjacent areas near the border with Democratic Republic of Congo and in the Cabinda encalve, although there are no published records from Cabinda. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a: 127, Mananas 1981:34); “riviere Lukashi, afluente de la rive gauche de la Tshihumbwe (Rio Lukashi)” [-7.53333, 21.08333] (Laurent 1950a:ll; Mananas 1981:34). Taxonomic and distributional notes; Wallaeh et al. (2014) recorded this species as present in Benguela Province, although this almost certainly is based on a misidentifica- tion. Map Angola. 276. Distribution of Causus lichtensteinii in Causus maculatus (Hallowell, 1842) Spotted Night Adder Distiehurus Maeulatus Hallowell 1842:337, pi. 19. Holotype: ANSP 6897 (eolleetor S.M.E. Goheen). Type loeality: “Liberia, Westen Afriea”. Causus rhombeatus-. Boeage (1895a:145), Parker (1936:126), Laurent (1950a:ll, 1954a:61), Hellmieh (1957b:74). Causus maeulatus-. Laurent (1964a:124), Thys van den Audenaerde (1966:36), Managas (1981:32), Spawls MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 317 and Branch (1995:145), Rasmussen (2005:14), Chippaux (2006:241), Trape and Mane (2006:206), Dobiey and Vogel (2007:127), Chirio and LeBreton (2007:608), Wallach et al. (2014:148). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from West-Central Africa from Mauritania to Ethiopia, and south to Angola. Ocurrences in Angola (Map 277): The species occurs in the northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1950a: 11, 1954a;61, 1964a: 124, Thys van den Audenaerde 1966:36, Managas 1891:32; Rasmussen 2005:14); “Barrage de la Luachi- mo” [-7.38333, 20.85000] (Thys van den Audenaerde 1966:36); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:ll, 1954a;61); “Sombo” [-8.68333, 20.95000] (Laurent 1954a:61). Kwanza Norte: “Piri- Dembos” [-8.53333, 14.43333] (Hellmich 1957b:74; Rasmussen 2005:14). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:126). Taxonomic and distributional notes; Bogert (1940) pointed out a possible taxonom¬ ic difference between the savanna C. rhombea- tus (typical form) and the forest dwellers, which was later confirmed by Laurent (1956, 1964a). Laurent (1960) identified C. maculatus as a forest form, distinguishing it from C. rhombeatus, which is sympatric in Angola (Mana9as, 1981). Laurent (1964a) assigned the Angolan records pre¬ viously refered to C. rhombeatus (e.g., “Dundo,” “Muita,” “Sombo” [Laurent 1950a, 1954a], “Quirimbo” [Parker 1936]) to C. maculatus, and refered the specimens cited in Bocage (1895a) from Congo and a part of northern Angolan to C. maculatus. Without further comment. According to Rasmussen (2005) two specimens from the Zoological Museum of the University of Hamburg previously identified as Causus rhombeatus (Lichtenstein, 1 823) from “Piri-Dembos” (Hellmich, 1957b) correspond, in fact, to Causus maculatus. Map 277. Distribution of Causus maculatus in Angola. Causus resimus (Peters, 1862) Green Night Adder Heterophis resimus Peters 1862b:277, pL, figs. 4, 4a, 4b. Holotype: ZMB 4370 (eolleetor A. von Bamin and R. Hartman). Type loeality: “Sennar, vom Gebel-Ghule” [= Jebel Ghule, Sennar Provinee], Sudan. Heterophis resimus = Causus rostratus: Bocage (1870:68). Causus resimus: Bocage (1887c:211), Ferreira (1904:116), Boulenger (1896:469, 1915:220), Loveridge (1936a:44, 1957:299), Parker (1936:127), Monard (1937b:141), Bogert (1940:97), Managas (1981:33), Spawls and Branch (1995:142), Spawls et al. (2004:474). Rasmussen (2005:14), Chirio and LeBreton (2007:610), Dobiey and Vogel (2007:128), Akani et al. (2012:189), Wallach et al. (2014:149). Causus resimus var. angolensis Bocage (1895a: 148). Syntypes: MBL 2025, 2080 (Biballa), MBL 2078, 2084a-b (Quissange), MBL 2079 (Novo Redondo), MBL 2082a-b (Rio Chimba), 2083a-b (Dondo) (fide McDiarmid et al. 1999) and presumably others, (collectors Banyures [Rio Dande, Rio Bengo], F.A.P. Bayao [Dondo], A. da Fonseca [Cazengo], F. Newton [Novo Redondo], J.A. d’Anchieta [Quissange, Rio Chimba, Biballa, Maconjo]), destroyed by fire 18 March 1978. Type locality: “Rio Dande et Rio Bengo ... Dondo ... Cazengo ... Novo Redondo ... Quissange, Rio Chimba, Biballa et Maconjo.” Causus rhombeatus: Hellmich (1957b:74). 318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II regions. Bengo: “Rio Bengo” [-8.71667, 13.40000] (Bocage 1895a; 146; Monard 19376:141; Managas 1981:33); “Cabicula” [-9.16667, 13.56667] (Ferreira 1904:116; Monard 19376:141; Mananas 1981:33); “Cunga” [-9.23333, 13.76667] (Mananas 1981:33). Luanda: “Rio Dande” [-8.46667, 13.38333] (Boeage 1895a:146; Monard 19376:141; Loveridge 1957:299; Mananas 1981:33); Kwanza Norte: “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1896:469; Managas 1891:33); “Cazengo” [-9.33333, 14.76667] (Bocage 1895a: 146; Ferreira 1904:116; Monard 19376:141; Managas 1981:33); “Caculo” [-9.38333, 14.91667] (Per- Map 278. Distribution of in Angola. reira 1904:116); “Dondo” [-9.68333, 14.43333] (Bocage 1895a:146; Monard 19376:141; Mananas 1981:33). Kwanza Sul: “Li6olo-Luati” [-9.98333, 14.90000] (Hellmich 19576:74; Managas 1981:33); “Novo Redondo” [-11.20000, 13.85000] (Bocage 1895a;146; Monard 19376:141; Mana¬ nas 1981:33); “Quirimbo” [-10.68333, 14.26667] (Parker 1936:127; Managas 1981:33); “Congu- lu” [-10.86667, 14.28333] (Parker 1936:127; Mananas 1981:33). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1887c:211, Bocage 1895a:146; Monard 19376:141; Managas 1981:33); “Catengue” [-13.03333, 13.73333] (Rasmussen 2005:14); “Hanha” [-12.25000, 13.75000] (Bogert 1940:97; Mananas 1981:33; Rasmussen 2005:14). Namibe: “Rio Chimba” [-14.30000, 12.40000] (Bocage 1895a:146; Monard 19376:141; Managas 1981:33); “Biballa” [-14.76667, 13.36667 -84] (Bocage 1895a:146; Monard 19376:141: Managas 1981:33); “Maconjo (Macujo)” [-15.01667, 13.20000] (Bocage 1895a:146; Monard 19376:141; Managas 1981:33). Undetermined Locality: “Without precise location” (Bocage 1870:68). Taxonomic and distributional notes: No subspecies are currently recognized (Wallach et al. 2014). Global conservation status (lUCN): Not Evaluated. Global distribution: The species has a strange and seemingly disjunct distribution in Africa, with isolated populations in northern Zaire, eastern Cameroon and the Cameroon-Chad border, Angola, presumably Nigeria, coastal Kenya, Somalia, central and south-western Sudan, and south¬ eastern Ethiopia to Mozambique (see Hughes 1987; Spawls and Branch 1995). Ocurrences in Angola (Map 278): The species occurs in western Angola, in dry and moist savannas and woodlands of coastal Causus rhombeatus (Lichtenstein, 1823) Rhombic Night Adder Sepedon rhombeata Lichtenstein 1823:106. Syntypes: ZMB 2768-69 (eolleetor G.L.E. Krebs) and possibly ZMB 2770 (donor [?] M.H.C. Liehtenstein, possibly G.L.E. Krebs). Type loeality: “Prom. B. sp.” [= Promontorium Bonae Spei = Cape of Good Hope], South Afriea. Restrieted to “area eontained within the magisterial districts of Uitenhage, Kirkwood and Port Elizabeth, Eastern Cape Provinee, South Afriea” by Bauer (2000:55, 57). Causus rhombeatus-. Gunther (1865a:480), Boeage (1866a:51, 1879b:95, 1887a:189, 18876:207, 1895a:145, 1896a:113, 18976:211), Peters (1877a:618, 1881:150), Ferreira (18976:245, 1900a:53, 1906:169), Boulenger (1896:468, 1905:114, 1915:220), Sehmidt (1933:15), Parker (1936:126), Loveridge (1936:44), MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 319 Monard (1937b:141-142), Mertens (1938a:442), Bogert (1940:96), Themido (1941:11), Laurent (1964a:123), Hellmich (1957a:75, 1957b:74), Thys van den Audenaerde (1966:36), Managas (1973:197, 1981:32), Spawls and Braneh (1995:147), Rasmussen (2005:14), Dobiey and Vogel (2007:129), Hampton (2010:235), Broadley (2014:350), Wallaeh et al. 2014:150), Braneh and Conradie (2015:200). Causus ef. rasmusseni: Conradie et al. (2016:23). Global conservation status (lUCN): Not Evaluated. Global distribution: The species has an extensive range in moist savannas of south, central and east Africa, from eastern Nigeria, southern Sudan and Ethiopia southwards to the Swellendam area in the Western Cape, South Africa, but is absent from much of the arid western half and much of the central part, of southern Africa. Ocurrences in Angola (Map 279): The species occurs in the entire country with excep¬ tion of the southern regions and probably Cabinda (see below). Cabinda: ? “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:618; Boca- ge 1895a:145); ? “Landana” [-5.21667, 12.15000] (Bocage 1895a:145). Zaire: “S. Sal¬ vador do Congo” [-6.26667, 14.23333] (Boca¬ ge 1887a: 189, 1895a: 145). Luanda: “Loanda” [-8.83333, 13.26667] (Ferreira 1900a;53). Bengo: “Cacolo ate ao Rio Bengo” (Ferreira 1900:53). Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1881:150; Bocage 1895a:145; Monard 1937b:141); “Pungo- Andongo” [-9.66667, 15.58333] (Gunther 1865a:480; Boulenger 1905:114; Managas 1981:33; Rasmussen 2005:14). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent Map 279. Distribution of Causus rhombeatus in Angola. 1964a:123; Mana9as 1981:32; Thys van den Audenaerde 1966:36; Broadley 2014:350); “Carum- bo” [-7.74422, 19.95467] (Branch and Conradie 2015:200); “Cossa” [-7.90000, 21.36667] (Lau¬ rent 1964a:123; Mana9as 1981:32; Broadley 2014:350); “Riviere Chicapa, pres du pont, 50 km a I’Ouest de Dundo” [-8.10000, 20.51667] (Thys van den Audenaerde 1966:36); “R. Chimenji, affl. de la Chiumbe” [-7.66667, 20.83333] (Thys van den Audenaerde 1966:36); “Cassange” [-9.58333, 17.86667] (Bocage 1895a:145; Monard 1937b:141). Lunda Sul: “Tyihumbwe” [-10.96667, 20.06667] (Monard 1937b:142). Mexico: “Lago Dilolo” [-11.50000, 22.01667] (Mana9as 1973:197); “environs du lac Calundo” [-11.71667, 20.80000] (Laurent 1964a:123; Mana9as 1981:32; Broadley 2014:350); “Calombe, Luso” [-11.83333, 19.93333] (Mana9as 1973:197); “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] (Mana9as 1973:197). Kwanza Norte: “Cazen- go” [-9.33333, 14.76667] (Ferreira 1900a:53); “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:74; Rasmussen 2005:14); “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1896:468; Fer¬ reira 1906:169; Monard 1937b:141; Rasmussen 2005:14); “Canhoca” [-9.25000, 14.68333] (Bou¬ lenger 1905:114; Rasmussen 2005:14); “Territorio de Ambaca” [-9.26667, 15.18333] (Ferreira 1900a:53). Kwanza Sul: “Mombolo” [-12.16667, 14.83333] (Bogert 1940:96; Mana9as 1981:33; Rasmussen 2005:14). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:15; Rasmussen 2005:14). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:142); “Bela-Vista” [-12.36667, 16.20000] (Hellmich 1957b:74; Mana9as 1981:33; Rasmussen 2005:14); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:126); “Santo-Amaro” [-12.70000, 15.85000] (Monard 320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II I937b:I4I-I42). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage I895a:I45; Monard 1937b: I4I); “Benguela” [-12.58333, 13.41667] (Loveridge 1936a:44); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:145; Monard 1937b:141); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:141-142); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:75; Mananas 1981:33); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:442; Rasmussen 2005:14); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113, 1897b:211); “Cabal” (Rasmussen 2005:14). Hulla: “Caconda” [-13.73333, 15.0666] (Bocage 1895a:145; Monard 1937b: 141; Rasmussen 2005: 14); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:141-142); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:141-142); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:96; Ras¬ mussen 2005:14); “Huilla” [-15.05000, 13.55000] (Hampton 2010:235). Namibe: “Rio Cuce” [-13.51667, 15.20000] (Ferreira 1897b:245; Monard 1937b:141); “Mossamedes” [-15.20000, 12.15000] (Bocage 1887a:189, 1895a:145). Cuando Cubango: “Kutatu” [-14.36667, 16.48333] (Monard 1937b:141-142); “Kakindo” [-15.45000, 17.05000] (Monard 1937b:141-142); “Kayun- du” [-15.70000, 17.45000] (Monard 1937b:141-142); “riverine habitat Rio Longa (57)” [-15.45969, 18.76833] (Conradie et al. 2016:8, 12, 23). Undetermined Locality: “Between Benguella and Bihe” (Boulenger 1905:114; Rasmussen 2005:14). Taxonomic and distributional notes: Hughes and Barry (1969) and McDiarmid et al. (1999) incorrectly stated that the types were lost, following information from G. Peters. Bogert (1940) pointed out a possible taxonomically relevant difference between the savanna C. rhombeatus (typ¬ ical form) and its forest dwelling congeners, which was later confirmed by Laurent (1956, 1964a). Laurent (1964a) assigned the Angolan records previously refered to C. rhombeatus (e.g., “Dundo,” “Muita,” “Sombo” [Laurent 1950a, 1954a], “Quirimbo” [Parker 1936]) to C. maculatus and also considered specimens cited in Bocage (1895a) from Congo and a part of northern Angolan to be maculatus, without further comment. The species is widely distributed in Angola, however the Cabinda records, “Chinchoxo” and “Landana,” probably represent misidentifications (Spawls and Branch 1995; Rasmussen 2005; Dobiey and Vogel 2007). Conradie et al. (2016) cited a single spec¬ imen from “Rio Longa,” Cuando Cubango as Causus cf rasmusseni Broadley, 2014, although its taxonomic status remains equivocal, since the topotypic material from C. rasmusseni is not avail¬ able, and the “Rio Longa” specimen is only weakly differentiated from South African material. Family Lamprophiidae Fitzinger, 1843 Genus Amblyodipsas Peters, 1857 Amblyodipsas polylepis (Bocage, 1873) Common Purple-Glossed Snake Calamelaps polylepis Bocage 1873b:216. Holotype: MBL 1878 (collector J.A. d’ Anchieta), destroyed by fire 18 March 1978. Type locality: “Dondo, interieur d’Angola” [= Dondo] Kwanza Norte Province, Angola. Atractaspis Hildebrandtii: Peters (1877a:616). Calamelaps polylepis: Bocage (1895a: 126, 1897a:201), Ferreira (1904:116), Boulenger (1915:214). Calamelaps unieolor Loveridge (1933:260). Calamelaps unieolor polylepis: Loveridge (1944c:164, 1957:281), de Witte and Laurent (1947:32). Amblyodipsas polylepis polylepis: Broadley (1971c:649), Broadley (1990:202), Branch (1998:66). Amblyodipsas polylepis: Broadley and Cotterill (2004:46), Bates et al. (2014:345), Wallach et al. (2014:25). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from coastal Kenya to South Africa, west to Ango¬ la and northern Namibia. Ocurrences in Angola (Map 280): The species is known from western Angola, however, the species may occur country wide with exception of the northern regions. Bengo: “Cabicula” MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 321 [-9.16667, 13.56667] (Ferreira 1904:116; de Witte and Laurent 1947:32; Broadley I97Ia:652). Kwanza Norte: “Cazengo” [-9.33333, 14.76667] (Boeage I895a:I26, I897a:20I; Loveridge 1944c: 164; de Witte and Laurent 1947:32; Broadley 1 97 1 a: 652); “Dondo” [-9.68333, 14.43333] (Boeage I873b:2I6, I895a:I26, I897a:20I; Loveridge 1933:260, I944c:I64, 1957:281; de Witte and Laurent 1947:32; Broadley 197 la: 652, 1990:202). Benguela: “Quissange” [-12.43333, 14.05000] (Boeage 1895a:126, 1897a: 201; Loveridge 1944b: 164; de Witte and Laurent 1947:32; Broadley 1971a:652). Cunene: “Humbe” [-16.68333, 14.90000] (Boeage 1895a: 126, 1897a:201; Loveridge 1944bc:164; de Witte and Laurent 1947:32; Broadley 1971a:652). Undetermined Locali¬ ty: “without precise locality” (Peters 1877a:616; Broadley 1971a:652). Taxonomic and distributional notes: Wallach et al. (2014) confused “Dondo” with “Dundo” from Lunda Norte Province, eastern Angola. There are currently no outstanding taxonomic issues for Amblyodipsas polylepis polylepis (Boeage 1873), although the taxonomic status of the East African subspecies Amblyodipsas polylepis hildebrandtii (Peters, 1877) should be re-evaluated (Bates et al. 2014). Map 280. Distribution of Amblyodipsas polylepis in Angola. Genus Aparallactus Smith, 1849 Aparallactus capensis Smith, 1849 Cape Centipede Eater Aparallactus capensis Smith 1849b: 16. Holotype: BMNH 1946.1.8.62 (formerly 64.12.12.3) (collector A. Smith). Type locality: “the country (Kafifirland) to the eastward of the Cape Colony” [= KwaZulu- Natal], South Africa fide Broadley (1990). Aparalaetus Guentheri (part) Boulenger (1895c: 172). Types: not identified. Type locality: “E. and C. Africa, Angola” corresponding to “Quindumbo”7/Je Boeage (1895a: 127), Benguela Province, Angola. Aparallaetus Boeagii Boulenger (1895c: 173). Types: not identified. Type locality: “Angola,” corresponding to “Novo Redondo” and “Gambos”7/£/e Boeage (1895a: 128), [= Sumbe and Chiange], Kwanza Sul and Huila provinces (respectively), Angola. Aparallaetus punetulolineatus Boulenger (1895c: 173). Type(s): not identified. Type locality: “Angola,” cor¬ responding to “Biballa”7/<7e Boeage (1897a:201) [= Bibala], Angola. Urieehis eapensis: Boeage (1895a: 128). Aparallaetus punetulolineatus: Boulenger (1895d: 173, 1915:217). Aparalaetus guentheri: Boulenger (1895c: 172, 1915:216), Monard (1937b: 129), de Witte and Laurent (1947:115). Aparallaetus punetulolineatus: Boulenger (1895c: 173, 1915:217). Aparallaetus boeagii: Boulenger (1896:259, 1915:216), Monard (1937b: 129). Aparallaetus punetolineatus: Boulenger (1896:259). Urieehis Boeagii: Boeage (1897a:201). Urieehis punetatolineatus: Boeage (1897a:201). Urieehis Guentherii: Boeage (1897a:201). 322 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Aparallactus Bocagii: Boulenger (1905:114). Aparallactus capensis bocagii: Loveridge (1944b:202), de Witte and Laurent (1947:126), FitzSimons (1962:270). Aparallactus capensis capensis: Loveridge (1944e:205, 1957:287). Aparallaetus eapensis punetatolineatus: de Witte and Laurent (1947:128), Laurent (1954a:45). Aparallaetus eapensis: Broadley (1990:154), Spawls et al. (2004:420), Broadley and Cotterill (2004:47), Bates et al. (2014:347), Wallaeh et al. (2014:45). Global conservation status (lUCN): Least Concern. Global distribution: The distribution of this species extends from Tanzania in the north to the eastern Cape Province in the south, and westwards through south-eastern Democratic Republic of the Congo and Zambia to Angola. Ocurrences in Angola (Map 281): The species is apparently distributed from the cen¬ tral-west to southeastern regions, although also extends to the northeastern Lunda Norte Province. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a;45); “Sombo” [-8.68333, 20.95000] (Laurent 1954a:45). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1895a:127-128, 1897a;201; Monard 19376:129; Loveridge 1944b:205; de Witte and Laurent 1947:122; Broadley 1990:156). Benguela: “Bigondo” [-12.06667, 17.41667] (Monard 1937b:129; Loveridge 1944c;202); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:127, 1897a;201; Monard 19376:129; Loveridge 1944c:205, 210; de Witte and Laurent 1947:115). Huila: “Gambos” [-15.76667, Map 281. Distribution of Aparallactus capensis Angola. 14.10000] (Bocage 1895a:127-128, 1897a: 201; Monard 19376:129; Loveridge 1944b; 205; de Witte and Laurent 1947:126; Broadley 1990:156). Cuando Cubango: “approximately 50 km E of Cuito Cunavale” [-15.23333, 19.61667] (Branch and McCartney 1992:2). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a:127, 1897a:201; Monard 19376:128; Loveridge 1 944c :205 ; de Witte and Laurent 1947:128; Broadley 1990:156). Undetermined Locality: “Between Benguella and Bihe” (Boulenger 1915:217; Loveridge 1944c:205; de Witte and Laurent 19476:122). Taxonomic and distributional notes; In the BNMH register, 1946.1.8.62 is listed as “one of the types,” suggesting the possibility that Smith’s description was based on multiple specimens, although details are given for only a single specimen. Bocage (1895a), in his ''Herpetologia d ’An¬ gola et du Congo,'' remarked on some differences of material of Uriechis capensis from the Museu Bocage from typical material, but did not distinguish these individuals as new forms. In the same year, however, Boulenger (1895d), described three new species based on Bocage specimens, Aparallactus bocagii, from “Novo Redondo” [= Sumbe] and “Gambos,” Angola, Aparallactus punetatolineatus from “Biballa,” Angola, and Aparallactus guentheri from “Quindumbo,” Angola and East and Central Africa (Bocage 1897a). Boulenger (1895c) himself did not provide specific localities within Angola for these new taxa, but these may be inferred from Bocage (1895a, 1897a). Loveridge (1944c) and FitzSimons (1962) treated ^4. bocagii as a western subspecies of A. capen- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 323 sis, and considered A. punctatolineatus a synonym, de Witte and Laurent (1947) and Laurent (1954a) maintained the subspeeifie distinetiveness of A. c. punctatolineatus based on what they believed were consistent differences from the nominate form. The latter author eited two new local¬ ities for punctatolineatus in Lunda Norte Provinee. More reeent authors consider A. punctatolin¬ eatus and A. bocagii as synonyms of the typical form (Broadley and Cotterill 2004; Spawls et al. 2004; Bates et al. 2014; Wallaeh et al. 2014). Broadley (1990) examined the syntypes of A. c. bocagii in the Museum Bocage before they were destroyed and noted that they could not be distinguished from the typical form. The speeies A. guentheri is eonsidered valid although it is limited to the eastern Africa and the Angola reeords should be assigned to A. capensis (Broadley 1990; Wallaeh et al. 2014). Genus Atractaspis Smith, 1849 Atractaspis bibronii Smith, 1849 Bibron’s Stiletto Snake Atractaspis bibronii Smith 1849a: pi. 71 (see Notes below). Syntypes: BMNH 1946.1.18.19 (formerly 83.7.30.5) (eolleetor A. Smith) and one unloeated speeimen (fide Wallaeh et al. 2014). Type loeality: “east¬ ern distriets of the Cape Colony” in error (see Notes below). Atractaspis Bibroni'. Boeage (1867e:227, 1870:68, 1895a:141). Atractaspis bibronii'. Boulenger (1896:515, 1915:223), Mertens (1937a: 16, 1938a:442), Tilbury and Braneh (1989:327), Broadley (1991a:498), Spawls and Braneh (1995:31), Spawls et al. (2004:440), Nagy et al. G005:226f Dobiey and Vogel (2007:21), Bates et al. (2014:349f Wallaeh et al. (2014:63), Braneh and Conradie (2015:200). Atractaspis bibroni rostrata: Laurent (1950a:ll, 1950b:33, 1954a:62, 1964a:122), Thys van den Audenaerde (1966:36), Managas (1981:42). Atractaspis bibroni x rostrata'. Broadley (1959:72). Atractaspis bibronii bibronii'. FitzSimons (1962:319), Broadley (1990:219). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is widely distributed in sub-Saharan Afriea, extending from Kenya and southern Somalia, southwards to Tanzania, Demoeratie Republie of Congo and Ango¬ la, western and northern Zambia, Mozambique to Zimbabwe, Botswana, Namibia and eastern por¬ tions of South Afriea, as well as Swaziland. Ocurrences in Angola (Map 282): The species has been reeorded in eentral western Angola and in the northeastern part of the eountry. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:ll, 1954a:62, 1964a; 122; Managas 1981:42); “Dundo, Barra¬ ge de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaerde 1966:36); “Carum- bo” [-7.74422, 19.95467] (Braneh and Conra¬ die 2015:200). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1867e:227, 1895a:141; Monard 1937b:141; Laurent 1950b:32; Managas 1981:41); “Benguella” [-12.58333, 13.41667] (Boeage 1895a;141; Monard 1937b:141; Laurent 1950b:32; Mana¬ nas 1981:41); “Dombe” [-12.95000, 13.10000] (Bocage 1895a:141; Monard 1937b:141; Lau¬ rent 1950b:32; Managas 1981:41); “Cubal” I2''E 20°E ,24“ E Map 282. Distribution of Atractaspis bibronii in Angola. 324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-13.03333, 14.25000] (Monard I937b:I4I; Mertens I937a:I6, I938a:442; Laurent I950b:32; Mananas 1981:41). Taxonomic and distributional notes: Smith’s (1849) plate gives his new species name as Atractaspis bibronii, however, the accompanying three unnumbered pages of text give the name as A. inornatus, except for the last paragraph in which it is implied that the species is named for Bibron. Atractaspis bibronii does not occur in the area given by Smith as the type locality. FitzSi- mons (1962) considered the actual source of the types to be “Natal” and Broadley (1990) likewise considered the type locality “almost certainly Natal,” but Broadley (1991a) suggested that “the northern Cape Province or the western Transvaal” was the likely source of the surviving BMNH syntype, as scale counts of rhte surviving syntype were inconsistent with specimens from Kwa¬ Zulu-Natal. Laurent (1950b) in his Atractaspis revision, treated A. rostrata Gunther, 1868 as a subspecies of A. bibronii and later (Laurent 1964a) cited five specimens from Lunda Norte Province, Angola. Broadley (1959) recognized an intermediate form, A. b. bibronii x rostrata, with presumed extensive intergradation covering most of Zimbabwe (Broadley 1991a). Currently, A. b. rostrata is considered a synonym of A. bibronii, although Nagy et al. (2005) noted that there is considerable intraspecific variation within Atractaspis bibronii, suggesting that several cryptic taxa are currently subsumed under this name. Spawls and Branch (1995) and Dobiey and Vogel (2007) considered Atractaspis bibronii limited to the western regions of Angola, from Luanda to Benguela Province, however, in addition to Laurent (1964a), the species was recently collected by Branch and Conradie (2015) in “Carumbo,” Lunda Norte Province. Atractaspis boulengeri Mocquard, 1897 Boulenger’s Stiletto Snake Atractaspis Boulengeri Mocquard 1897a:54. Holotype: MNHN 1896.562 (collector E. Haug). Type loeality: “les environs de Lambarene, sur le has Ogooue, Gabon” [= Lambarebe, Moyen-Ogooue], Gabon. Atraetaspis boulengeri mixta: Managas (1981:40). Atraetaspis boulengeri: Spawls and Braneh (1995:32), Chippaux (2006:193), Dobiey and Vogel (2007:22), Wallaeh et al. (2014:63). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from across Central Africa from southern Cameroon to the Democratic Republic of Congo. Ocurrences in Angola: The species presumably occurs in northwestern Angola, including the Cabinda enclave, Angola. There are no published records for this species for Angola, however Manaqas (1981), Spawls and Branch (1995) and Dobiey and Vogel (2007) suggested its occurrence in northern Angola. Taxonomic and distributional notes: Mocquard’s original description in the Bulletin du Museum d’Histoire Naturelle (1897a) was reprinted in Bulletin de la Societe Philomathique de Paris in the same year (1897b) (Wallaeh et al. 2014). Atractaspis congica Peters, 1877 Congo Stiletto Snake Atraetaspis eongiea Peters 1877a:616, pi. Figs. 2, 2a-2e. Holotype: ZMB 8644 (eolleetor F.W.A. von Mechow). Type locality: “Chinchoxo (Westafrika)” (Peters 1877a:616) [= Chinchoxo, Cabinda enealve], Angola. Atraetaspis eongiea: Peters (1881:150), Boeage (1887a:187, 1895a:142, 1897b:210), Ferreira (1904:116), Boulenger (1905:114), Schmidt (1933:15), Monard (1937b: 141, 144), Themido (1941:11), Spawls and Branch (1995:33), Branch (1998:63), Broadley and Cotterill (2004:46), Broadley et al. (2003:83), Wallaeh et al. (2014:63). Atraetaspis aterrima: Boeage (1873a:223), Boulenger (1915:223), Chirio and LeBreton (2007:624). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 325 Atractaspis congica congica: Hellmich (1957a:76, 1957b:76), Laurent (1950b:26), Mananas (1981:40), Chi- rio and LeBreton (2007:628). Atractaspis congica orientalis: Laurent (1964a: 122-123), Managas (1981:41), Broadley (1990:221). Map 283. Distribution oi Atractaspis congica in Angola. Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies oeeurs from Cameroon through Demoeratie Republie of Congo to Angola and northern Zambia. Ocurrences in Angola (Map 283): The species is widely distributed across much of Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:616; Bocage 1895a; 142; Laurent 1950b:26; Managas 1981:40); “Cabinda” [-5.55000, 12.18333] (Managas 1981:40). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:123); “Mutianvo” [-11.45000, 19.33333] (Themido 1941:11). Moxico: “environs du lac Calundo (arredores do Lago Calundo)” [-11.80000, 20.86667] (Laurent 1964a: 122; Mana9as 1981:41). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Boulenger 1905:114; Laurent 1950b:27; Managas 1981:40); “Pungo- Adongo” [-9.66667, 15.58333] (Laurent 1950b:27; Managas 1981:40). Kwanza Norte: “Piri-Dembos” [-8.53333, 14.43333] (Hell¬ mich 1957b:76; Managas 1981:40); “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:114; Ferreira 1906:169; Monard 1937b:141; Laurent 1950b:27; Mananas 1981:40); “Cazengo” [-9.33333, 14.76667] (Ferreira 1904:116; Monard 1937b:141). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:15; Laurent 1950b:26; Mananas 1981:40). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b:141, 144; Laurent 1950b:26; Managas 1981:40); “Galanga” [-12.06667, 15.15000] (Bocage 1895a:142; Monard 1937b:141; Laurent 1950b:26; Mana9as 1981:40); “Bela Vista” [-12.56667, 16.21667] (Hellmich 1957b:76; Mana9as 1981:40). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:142; Monard 1937b:141; Laurent 1950b:26; Mana9as 1981:40); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:142; Monard 1937b:141; Laurent 1950b:26; Mana9as 1981:40); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:76); Mana9as 1981:40); “Alto Cubal” [-13.03333, 14.25000] (Hellmich 1957b:76; Mana9as 1981:40); “Hanha” [-13.30000, 14.20000] (Bocage 1897a:210; Laurent 1950b:26; Mana9as 1981:40). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:142; Monard 1937b:141; Laurent 1950b:26; Mana9as 1981:40); “Huila” [-15.08333, 13.55000] (Bocage 1873a:223, 1895a:142; Monard 1937b:141; Mana9as 1981:40). Undetermined Locality: “Cuango” (Peters 1881:150; Laurent 1950b:27; Mana9as 1981:40) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distributional notes: Laurent (1950b) noted that a subspecific distinction probably exists between populations separated by the Congo River, as there are considerable dif- fereces in terms of dorsal and ventral scales between the Angolan populations and those from the north. This species is sympatric with the Reticulate Stiletto Snake, Atractaspis reticulata (Sjostedt, 1896), in northern Angola and around the mouth of the Zaire River (Spawls and Branch 1995). 326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Atractaspis irregularis (Reinhardt, 1843) Variable Stiletto Asp Flaps irregularis Reinhardt 1843:264, pi. 3, figs. 1-3. Holotype: ZMUC 6885 (collector J.R. Chenon). Type locality: “Guinea,” in error, “coast of Ghana” Hughes and Barry (1969:1032), resticted to the “vicin¬ ity of Accra, Ghana” by Rasmussen and Hughes (1997). Atractaspis irregularis: Gunther (1865a:480), Peters (1877a:616), Bocage (1895a: 143), Boulenger (1915:223), Monard (1937b: 141), Frade (1963:253), Spawls and Branch (1995:36), Chippaux (2006:186), Wallach et al. (2014:65). Atractaspis corpulentus: Bocage (1866a:49). Atractaspis irregularis parkeri: Laurent (1950b:17), Managas (1981:41), Chippaux (2006:186). Atractaspis irregularis irregularis: Chirio and LeBreton (2007:634). Atractaspis irregularis: Wallach et al. (2014:65). Global conservation status (lUCN): Least Concern. Global distribution: The species is widely distributed across sub-Saharan Africa from Sierra Leone to the Central African Republic to South Sudan, Eritrea and Ethiopia and south to Tanzania, the Democratic Republic of Congo and north¬ ern Angola. Ocurrences in Angola (Map 284): The species has been reported from the northwest¬ ern regions of the country, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:616; Lau¬ rent 1950b:17; Mananas 1981:41); “Molembo” [-5.33333, 12.20000] (Bocage 1895a:143; Monard 1937b:141; Managas 1981:41). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:143; Monard 1937b:141; Lau¬ rent 1950b:17; Managas 1981:40). Malanje: “Pungo Andongo” [-9.66667, 15.58333] (Giin- ther 1865a;480). Taxonomic and distributional notes: In addition to the nominate form, which occurs in Angola, four other subspecies are currently recognized. Map 284. Distribution of Atractaspis irregularis in Angola. Atractaspis reticulata heterochilus Boulenger, 1901 Reticulate Stiletto Snake Atractaspis heterochilus Boulenger 1901:13, pi. V, fig. 1. Type: MRAC 9321 (eolleetor Capt. Heeq). Type loeality: “des environs d ’Albertville, sur le Tanganika” [= Kalemie, Tanganyika Provinee], Demoeratie Republie of Congo. Atractaspis reticulata heterochilus: Hellmieh (1957b:77), Spawls and Braneh (1995:39). Atractaspis reticulata heterochilus {Atractaspis heterochilus): Managas (1981:42). Atractaspis reticulata heterochilus: Chippaux (2006: 192), Chirio and LeBreton (2007:636), Dobiey and Vogel (2007:30). Atractaspis reticulata: Wallaeh et al. (2014:66). Global conservation status (lUCN): Data Deficient. Global distribution; The subspecies is known from southern Cameroon, through Equatorial Guinea, Congo, large portions of the Democratic Republic of Congo to northwestern Angola, possibly including the Cabinda enclave. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 327 Ocurrences in Angola (Map 285): The species occurs in the northwestern regions of the country. Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:77; Mananas 1981:42). Taxonomic and distributional notes: Chippaux (2006, 2009) incorrectly placed the type locality of Atractaspis heterochilus in Tanzania. Dobiey and Vogel (2007) indicated a presumed range across northwestern Angola including Cabinda, but did not show its distri¬ bution as including the type locality of Albertville, or any ppart of the eastern Demo¬ cratic Republic of Congo. Angola was not included in the distribution as stated by Wal- lach et al. (2014). I2*E 16*E 20“E 24°E Map 285. Distribution of Atractaspis reticulata heterochilus in Angola. Genus Boaedon Dumeril, Bibron and Dumeril, 1854 Boaedon angolensis Bocage, 1895 Angolan House Snake (Endemic) Global conservation status (lUCN): Not Evaluated. Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 286): Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:80; Monard 1937b:113). Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Bocage 1895a:80; Ferreira 1900:48; Monard 1937b: 113); “Cazengo” [-9.33333, 14.76667] (Ferreira 1900a:48); Boodon lineatus var. angolensis Bocage 1895a:80. Syntypes: MBL specimen numbers unknown (collectors F.A.P. Bayao [Duque de Braganga], Padre A. Barroso [St. Salvador du Congo], J.A. d’Anchieta [Ambaca, Quissange, Cahata, Galanga, Caconda, Biballa, Huilla, Gambos, Humbe], destroyed by fire 18 March 1978 [some specimens may have been exchanged and survived in other collections]. Type locality: “Duque de Braganga ... St. Salvador du Congo . . . Ambaca, Quissange, Cahata, Galanga, Caconda, Huilla, Gam¬ bos, Biballa et Humbe”, [= Calandula (Malanje Province) . . . M’Banza Congo (Zaire Province) . . . Amba¬ ca (Kwanza Norte Province) . . . Galanga (Huambo Province) . . . Quissange, Caota (Benguela Province) . . . Caconda, Huila, Chiange (Huila Province) ... Bibala (Namibe Province) and Humbe (Cunene Province)], Angola. I2®E. lO'E 24°E Boodon lineatus var. angolensis: Bocage (1896a: 112, 1897b:211), Ferreira (1900a:51). Boodon lineatus: Ferreira (1897b:244, 1904:114), Monard (1937b: 113). Boaedon lineatus: Wallach et al. (2014:96). Boaedon cf angolensis: Conradie et al. (2016:22). Map 286. Distribution of Boaedon angolensis in Angola. 328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Cacullo” [-9.38333, 14.91667] (Ferreira 1904:114; Monard I937b:II3); Benguela: “Cahata” [-12.35000, 14.81667] (Bocage I895a:78); “Quissange” [-12.43333,14.05000] (Bocage I895a:80); “Hanha” [-13.30000, 14.20000] (Bocage I896a:II2, I897b:2II). Hufla: “Caconda” [-13.73333, 15.06667] (Bocage I895a;80; Ferreira I897b:244; Monard I937b:II3). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage I895a:80). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage I895a:80). Mexico: “upper Cuito River” (photographed) [-I3.I5I39, 18.48056] (Conradie et al. 2016:22). Cuando Cubango: “Cuito basin (50)” [-14.59517, I8.07III] (Conradie et al. 2016:9, 12, 22). Undetermined Locality: “Cacolo ate as margens do Bengo” (Ferreira 1900a:48). Taxonomic and distributional notes: Boaedon angolensis Bocage, 1895, was described by Bocage (1895a) as a variety of Boaedon lineatus Dumeril, Bibron and Dumeril, 1854, but was long forgotten and/or considered as a synonym of Boaedon fuliginosus or B. lineatus. Laurent (1956) mentioned that Angolan populations could be distinguished morphologically from other house snakes, but he did not discuss the availability of B. angolensis. The status of B. angolensis and related forms in Angola is under active review and both its taxonomic status and its distribution will be revised in the near future. Additional literature references to members of the B. fuliginosus complex may belong in the chresonymy of B. angolensis, but the most cannot be resolved without consulting the referred specimens Boaedon variegatus (Bocage, 1867) Variegated House Snake (Endemic) Alopecion variegatum Bocage 1867b:227, 1867d:230. Syntypes: MBL specimen numbers not known. Type locality: “Benguella” and “Novo Redondo” (Bocage 1867d:230) [= Benguela (Benguela Province) and Sumbe (Kwanza Sul Province)] Angola. Boodon lineatus: Bocage (1895a:80), Boulenger (1893:332), Monard (1937b: 113). Boaedon fuliginosus fuliginosus: Loveridge (1957:251). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is endemic to Angola. Ocurrences in Angola (Map 287): Luan¬ da: “Loanda” [-8.83333, 13.26667] (Bocage 1895a:80, Ferreira 1900a:48; Monard 1937b: 113). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1867b:227, 1867d:230, 1895a:80; Monard 1937b:113; Loveridge 1957:251). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a:78). Benguela: “Dombe” [-12.95000, 13.10000] (Bocage 1867b:227, 1895a:80; Monard 1937b: 113); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:227, 1895:78; Bou¬ lenger 1893:333; Loveridge 1957:251); Nami¬ be: “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:80; Monard 1937b:113). Taxonomic and distributional notes: Alopecion variegatum has variously been con¬ sidered as a synonym of other taxa in the Map 287. Distribution of Boaedon variegatus in Angola. B. fuliginosus complex, but recent collections and ongoing reviews support its specific distinctness. The status of B. variegatus and related forms in Angola is under active review and both its taxo¬ nomic status and its distribution will be revised in the near future. Additional literature references MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 329 to members of the B. fuliginosus complex may belong in the chresonymy of B. variegatus, but the most cannot be resolved without consulting the referred specimens. Boaedon fuliginosus complex (Boie, 1827) Brown House Snake Lycodon fuliginosus: Boie (1827: column 551). Type(s): unknown, RMNH?. Type locality: “Erp. de Java ist das Vaterland bekannt” (Boie 1827:column 551). Boie (1827) listed this species under Asian species of Lycodon and cited the unpublished Erpetology de Java as the source for the name. It is unclear why the manuscript title was mn together with “ist der Vaterland bekannt.” This may imply “Java” as the type locality, or it may have been an error for “Der Vaterland is unbekannt.” Boaedon lineatum: Bocage (1866a:49, 1867b:227), Parker (1936:122). Boedon quadrilineatum: Bocage (1879c:89). Boodon quadrilinaetus: Peters (1881:149). Boodon lineatus: Gunther (1865a:480), Bocage (1895a:78), Boulenger (1893:332, 1896:616, 1905:112), Ferreira (1897b:244, 1900a:51, 1903:10, 1904:114, 1906:167), Monard (1937b:113, 117), Themido (1941:9). Boaedon lineatus: Schmidt (1933:13), Loveridge (1936a:22), Mertens (1937a: 12, 1938a:439). Boaedon lineatus lineatus: Bogert (1940:21), Laurent (1950a:7, 1954a:43, 1964a:93), Hellmich (1957a:71, 1957b:60), Thys van den Audenaerde (1966:32). Boaedon fuliginosus fuliginosus (Baedon lineatus): Managas (1973:190). Lamprophis fuliginosus: Trape and Mane (2006:114), Chirio and LeBreton (2007:456), Ceriaco et al. (2016b:85). Boaedon fuliginosus: Kelly et al. (2011:424), Wallach et al. (2014:95), Greenbaum et al. (2015:18), Wallach et al. (2014:95). Global conservation status (lUCN): Not Evaluated. Global distribution: The species complex is known from southern Morocco to South Africa. Ocurrences in Angola (Map 288): Mem¬ bers of this complex are present throughout Angola with the exception of the desert regions of the far southwestern Angola. Cabinda: “Landana” [-5.20000, 12.15000] (Bocage 1895a:78; Monard 1937b:113); “Molembo” [-5.33333, 12.20000] (Bocage 1895a;78; Monard 1937b: 113). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:7, 1954a:43, 1964a:93; Thys van den Audenaerde 1966:32); “Dundo, Barrage de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaer¬ de 1966:32); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1954a:43); “Rio Luinha” [-9.26667, 14.53333] (Ferreira 1906:167; Monard 1937b: 113). Lunda Sul: “Alto Chica- pa” [-10.88333, 19.23333] (Laurent 1964a;93); “Mutianvo” [-11.45000, 19.33333] (Themido . 288- of complex 1941:9). Bengo: “Cabiri” [-8.91667, 13.66667] (Loveridge 1936a:22). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:60); “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:10; Monard 1937b:113); “Dondo” [-9.68333, 14.43333] (Bocage 1895a:78; Monard 1937b:113). Malanje: “Duque de 330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Bragan9a” [-9.10000, 15.95000] (Bocage 1866a:49, 1895a:78; Monard 1937b:113); “Pungo- Andongo” [-9.66667, 15.58333] (Gunther 1865a;480; Boulenger 1893:332-334, 1905:112; Monard 1937b:113); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:85). Mexico: “environs du lae Calundo” [-11.80000, 20.86667] (Laurent 1964a:93); “Rio Calombe, Luso” [-11.83333, 19.93333] (Managas 1973:190). Bie: “Chitau” [-11.43333, 17.15000] (Schmitd 1933:13); “Bihe” [-12.38333, 16.95000] (Bocage 1879c:89). Huambo: “Bela-Vista (Sanguen- gue)” [-12.36667, 16.20000] (Hellmich 1957b:60); “Huambo” [-12.76667, 15.73333] (Themido 1941:9). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:122); “Lembu (Serra de Selles)” [-12.86667, 14.11667] (Ferreira 1904:114). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:78; Monard 1937b:113); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:227, 1895a:78; Boulenger 1893:333); “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:71); “Cubal” [-13.03333, 14.25000] (Mertens 1937a:12, 1938a:439; Hellmich 1957b:60); “Hanha” [-12.25000, 13.75000] (Bogert 1940:21). Huila: “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:113, 117); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:113, 117); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:78); “Gambos” [-15.76667, 14.10000] (Bocage 1895a:80). Cunene: “fl. Mbale” [-15.16667, 16.75000] (Monard 1937b:113, 117). Cuan- do Cubango: “Kutatu” [-14.36667, 16.48333] (Monard 1937b:113, 117); “Kakindo” [-15.45000, 17.05000] (Monard 1937b:113, 117). Undetermined Locality: “without precise location” (Bocage 1867b:227); “Cuango” (Peters 1881:149) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distributional notes: The Boaedon fuliginosus complex, includes Boaedon lineatus Dumeril, Bibron and Dumeril, 1854, Boaedon fuliginosus (Boie, 1827) and Boaedon capensis (Bibron and Dumeril, 1854) as well as the taxa B. angolensis and B. variegatus, listed above. Recent molecular analyses conducted by Kelly et al. (2011) demonstrated extensive genet¬ ic variation in the widespread B. fuliginosus, suggesting several cryptic species are present. Sever¬ al names are available for putative taxa within the B. fuliginosus complex, and additional genetic sampling and extensive morphological analyses are needed to update the taxonomy of this group (Greenbaum et al. 2015). Broadley et al. (in prep) are trying to evaluate species boundaries in the group and the concomitant application of names. Wallach et al. (2014) assigned the Angolan records to B. fuliginosus, however, we have separately listed those records which appear to be unambiguously assignable to B. angolensis and B. variegatus. Citation records in this account may also refer to one of these other taxa, but this cannot be determined without a moe thorough revision. Boaedon olivaceus (Dumeril, 1856) Olive House Snake Holuropholis olivaceus Dumeril 1856:466. Holotype: MNHN 3408 (collector C.E. Auhry-Lecomte). Type locality: “Gabon.” Holuropholis olivaceus: Peters (1877a:615). Boodon olivaceus: Bocage (1895a:81). Boaedon olivaceus: Boulenger (1915:202), Laurent (1954a:43), Loveridge (1957:251), Frade (1963:252), Thys van den Audenaerde (1966:32), Wallach et al. (2014:97). Lamprophis olivaceus: Chiro and LeBreton (2007:460). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from west and central Africa, from Guinea to Central African Republic, south to the Democratic Republic of the Congo, and northern Angola and east to Rwanda and Uganda. Ocurrences in Angola (Map 289): The species occurs in the Cabinda enclave and Lunda MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 331 Norte Pronvice. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters I877a:6I5; Boca- ge I895a:8I); “Cabinda” [-5.55000, 12.18333] (Frade 1963:252). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954:43; Thys van den Audenaerde 1966:32). Taxonomic and distributional notes: Kelly et al. (2011) provided molecular phylo¬ genetic data to support the inclusion of this taxon along with the B. fuliginosus complex within a monophyletic Boaedon, separate from Lamprophis sensu stricto. Il-E 20°E 24'>E Map 289. Distribution of Boaedon olivaceus in Angola. Genus Bothrophthalmus Peters, 1863 Bothrophthalmus lineatus Peters, 1863 Red-Black Striped Snake Bothrophthalmus lineatus {nomem nudum) Schlegel in Lichtenstein and von Martens 1856:27. Type loeality: “Goldkuste.” Elaphis {Bothrophthalmus) lineatus Peters 1863:287. Holotype: ZMB 1820 (“aus dem Museum zu Leiden”). Type loeality: “Guinea” [= Ghana]7/t/e Hughes and Barry (1969:1013). Bothrophthalmus lineatus lineatus'. Laurent (1950b:8, 1954:44, 1964a:93), Loveridge (1957:249), Thys van den Audemaede (1966:32). Bothrophthalmus lineatus'. Spawls et al. (2004:316), Chirio and LeBreton (2007:386), Wallaeh et al. (2014:112). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Uganda, southwest to northern Angola, west to Guinea, with sporadic records from Uganda and west Rwanda. Ocurrences in Angola (Map 290): The species is known from extreme northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950b:8, 1954a:44; Thys van den Audemaede 1966:32); “Riv. Luachi- mo, Dundo” [-7.38333, 20.85000] (Laurent 1964a:93); “Riv. Mussungue, environs de Dundo” [-7.41667, 20.83333] (Laurent 1964a:93). Taxonomic and distributional notes: The original use of the name was as a nomen nudum in the Nomenclator of Lichtenstein and von Martens (1856), who attributed the name ll-E 1hS°E 20'>E 24°E Map 290. Distribution of Bothrophthalmus lineatus in Angola. 332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II to Schlegel, but provided no details other than a locality. Peters’ (1863) description used this name, again attributed to Schlegel, but as the description itself was that of Peters, he alone is the author¬ ity for the nomen. Authorship of the Nomenclator has been cited in various ways. Lichtenstein alone is often given credit for names in this work, but the preface states that the majority of the preparation of the Nomenclator, including the descriptions, was due to von Martens (see Harris and Kluge 1984; Ulber 2003). David Weinland also contributed to the publication (Adler 2012), although his status as an author on the publication is unresolved. Genus Dromophis Schlegel, 1837 Dromophis lineatus (Dumeril, Bibron and Dumeril, 1854) Lined Olympic Snake Dryophylax lineatus Dumeril, Bibron and Dumeril 1854:1124. Syntypes: MNHN 7643-7644 (eolleetor B. d’Amaud). Type loeality: “Nil blane” (Dumeril, Bibron and Dumeril 1854:1126), [= Upper Nile, Jon- glei or Central Equatoria Provinee, South Sudan fide Wallaeh et al. 2014]. Dromophis lineatus: Bogert (1940:79), Hughes (2004:74), Broadley and Cotterill (2004:49), Wallaeh et al. (2014:240). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Senegal east to Ethiopia, south to Angola, Zambia and northern Malawi. Ocurrences in Angola: Recorded in the literature from “Angola” with no precise information (Bogert (1940:79); Loveridge (1940:9); Hughes (2004:74). Taxonomic and distributional notes: Bogert (1940) cited the species for “Angola” without information about the collection site. The genus Dromophis has been treated as a synonym of Psammophis by Kelly et al. (2008), but has been recognized as valid by Hughes 2004, 2013), Wallaeh et al. (2014) and most other modem aurthors. Likewise the history of the nomen D. lin¬ eatus has been intertwined with that of Psammophis sibilans (Linnaeus, 1758) (see Hughes 2004 for a summary and comment). Genus Gonionotophis Boulenger, 1893 Gonionotophis brussauxi (Mocquard, 1889) Mocquard’s File Snake Godionotus brussauxi Mocquard 1889:146. Holotype: MNHN 1890.54 (collector E. Bmssaux). Type loeali¬ ty: “Loudinia-Niara, sur le fleuve Niai, entre Loango sur le litoral, et Brazzaville sur le fleuve Congo” [= between Ludinia on the Niara River], Congo. Gonionotophis brussauxi: Laurent (1954a:44), Chippaux (2006:91), Chirio and LeBreton (2007:428), Vidal et al. (2008:57), Kelly et al. (2011:425), Wallaeh et al. (2014:308), Lanza and Broadley (2014:89), Broad¬ ly et al. (2018:7). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Cameroon to the Democratic Republic of Congo, and extreme northeastern Angola. Ocurrences in Angola (Map 291): The species is known from extreme northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:44). Taxonomic and distributional notes: The Dundo record of Laurent (1954a) probably repre¬ sents the southernmost limit of the species. Loveridge (1939) remarked on the close relationship between the two file snake genera Mehelya and Gonionotophis Boulenger, 1893, and later, Vidal et al. (2008) and Kelly et al. (2011) determined that Gonionotophis brussauxi (Mocquard, 1889), type species of Gonionotophis, is nested within Mehelya. In order to maintain monophyletic gen¬ era, Kelly et al. (2011) transferred all species placed in Mehelya Csiki, 1903 to Gonionotophis, MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 333 which has priority over all other available names (Lanza and Broadley 2014). More recently, Broadley et al. (2018) split Goniono- tophis into four genera, with truss auxi being the only Angolan species remaining in the genus. Il-E 14“ E Map 291. Distribution of Gonionotophis brussauxi in Angola. Genus Hemirhagerrhis Boettger, 1896 Hemirhagerrhis viperina (Bocage, 1873) Western Bark Snake Psammophylax viperinus Bocage 1873a:222. Holotype: MBL 1715 (collector J.A. d’ Anchieta), destroyed by fire 18 March 1978. Type locality: “Dombe” [= Dombe Grande], Benguela Province, Angola. Psammophylax nototaenia: Bocage (1895a: 109). Amplorhinus nototaenia'. Boulenger (1915:211). Amplorhinus nototaenia {Psammophylax nototaenia)'. Monard (1937b: 128). Hemirhagerrhis nototaenia viperinus'. Bogert (1940:76), FitzSimons (1962:208), Laurent (1964a: 112), Broadley (1990:120). Hemirhagerrhis viperina'. Broadley (1997c: 164), Branch (1998:86), Broadley and Hughes (2000:7), Wallach et al. (2014:321). Hemirhagerrhis nototaenia'. Branch (1998:86), Wal¬ lach et al. (2014:320). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola to northern Namibia. Ocurrences in Angola (Map 292): The species occurs in the southwestern Angola. Huambo: “Huambo” [-12.83333, 15.66667] (Bogert 1940:75; Broadley 1997c: 164; Broadley and Hughes 2000:9). Benguela: “Dombe” [-12.95000, 13.10000] (Bocage 1873a:222, 1895a:109; Monard 1937b:128; FitzSimons 1962:208; Broadley 1990:120, 1997c: 161, 164; Broadley and Hughes 2000‘9)' “Hanha” [-12 25000 13 75000] Distribution of Hemirhagerrhis viperina in ’ ’ Angola. 334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (Bogert 1940:76; Broadley 1997c: 1 64; Broadley and Hughes 2000:9). Hufla: “Humpata, Fazenda Bumbo” [-15.20000, 13.00000] (Laurent 1964a: 1 12; Broadley 1997c: 164; Broadley and Hughes 2000:9); “8 km from Tundavala to Sa da Bandeira” [-15.83333, 13.40000] (Broadley 1997c: 164; Broadley and Hughes 2000:9); “5 km S of Chibemba” [-15.75000, 14.08333] (Broadley 1997c:164; Broadley and Hughes 2000:9). Namibe: “Huxe” [-12.71667, 13.20000] (Broadley 1997c: 164; Broadley and Hughes 2000:9); “Lungo” [-14.31667, 13.20000] (Broadley 1997c: 164; Broadley and Hughes 2000:9); “Munhino” [-14.91667, 13.00000] (Bogert 1940:76; Broadley 1997c:164; Broadley and Hughes 2000:9); “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:109; Broadley 1997c:161, 164; Broadley and Hughes 2000:9); “Caraculo” [-15.01667, 12.66667] (Broadley 1997c: 164; Broadley and Hughes 2000:9); “Capangombe” [-15.10000, 13.15000] (Bocage 1895a: 109; Broadley 1997c: 161, 164; Broadley and Hughes 2000:9). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a:109; Broadley 1997c:161). Taxonomic and distributional notes: Bocage (1873a) described Psammophylax viperinus based on one individual from “Dombe” collected by Anchieta in Benguela Province. Subsequent¬ ly he (Bocage 1895a) synonymized P. viperinus with P. nototaenia (Gunther, 1864) and recorded six specimens from southwestern Angola: two from “Maconjo,” one from “Capangombe,” two from “Humbe,” and the “Dombe” viperinus speciemen. Broadley (1997c) examined the material in Museu Bocage in 1968, and found only five specimens, the one from “Dombe” and two each from “Maconjo” and “Capangombe” (the reason for the discrepancy in the number of specimens from Capangombe is unknown). He suggested that the missing specimens from “Humbe” may have been typical H. nototaenia. However, H. nototaenia extends from Kenya to South Africa and Mozambique, whereas H. viperina is widely distributed from southwestern Angola to northern Namibia (Broadley 1990, 1997c; Broadley and Hughes 2000), suggesting that all records from Angola are correctly interpreted as H. viperina. Genus Hypoptophis Boulenger 1908 Hypoptophis wilsonii Boulenger, 1908 Wedge-Snouted Burrowing Snake Hypoptophis wilsonii Boulenger 1908:93. Holotype: BMNH 1946.1.2.99 (formerly BMNH 1908.6.16.5) (eol- leetor H. Wilson). Type loeality: “Inkongo, on the Sankum River, in the Kassai Province of the Congo” (Boulenger 1908:93), [= Inkongo, Kasai Oriental Province], Democratic Republic of Congo. Hypoptophis wilsoni katangae: de Witte and Lau¬ rent (1947:93), Laurent (1950a: 10). Hypoptophis wilsonii-. Broadley et al. (2003:91), Broadley and Cotterill (2004:47), Wallach et al. (2014:337). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from from the Congo Basin south to Katanga in Democratic Republic of Congo, western Zambia and northeastern Angola. Ocurrences in Angola (Map 293): The species is only reported from “Dundo” in the extreme northeast of the country near the MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 335 border with the Demoeratie Republic of Congo. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 10). Taxonomic and distributional notes: This species was originally described from the central Congo basin and is considered rare being known only from few specimens (Broadley et al. 2003). Muller (1911) described Michellia katangae, from “Kituri, Katanga district,” Congo, and de Witte and Laurent (1947) considered it a subspecies of Hypoptophis wilsonii Boulenger, 1908, which is currently viewed as a monotypic (Wallach et al. 2014). Wallach et al. (2014) did not include Ango¬ la in the species distribution. Genus Limaformosa Broadley, Tolley, Conradie, Wishart, Trape, Burger, Kusamba, Zassi-Boulou and Greenbaum, 2018 Limaformosa capensis (Smith, 1847) Southern File Snake Heterolepis capensis Smith 1847a: pi. 55, first of two accompanying unnumbered text pages. Holotype: lost fide FitzSimons (1937). Neotype: BMNH 1891.9.15.9 (donated by W.L. Selater, SAM), designated by Broadley (2005) fiide Lanza and Broadley (2014:90). Holotype loeality: “eastern distriets of the Cape Colony” [= South Afriea], locality rejected by Broadley (2005); Neotype locality: “Delagoa Bay” [= Maputo Bay, Maputo Province], Mozambique. Simoeephalus eapensis: Monard (1937b: 111, 119). Mehelya eapensis eapensis: Loveridge (1939:143, 1957:253), Laurent (1964a:94), Braneh and McCartney (1992:2), Broadley and Cotterill (2004:48). Mehelya eapensis: Broadley (2005:232), Chirio and LeBreton (2007:476), Wallaeh et al. (2014:424). Gonionotophis eapensis: Kelly et al. (2011:424), Lanza and Broadley (2014:90). Limaformosa eapensis: Broadley et al. (2018:5). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Southeastern Africa, north to eastern Ango¬ la and Zambia, southeastern Democratic Republic of Congo (former Katanga Province), Malawi and southeastern Tanzania as far north as the eastern Usambara Mountains. Wallach et al. (2014) credited Lanza and Broadley (2014) with the neotype designation, but this was originally proposed by Broadley (2005). Kelly et al. (2011) argued that Gonionotophis is the appropriate name for species previously assigned to Mehelya Csiki, 1903, but more recently Broadley et al. (2018) erected the new genus Limaformosa, with capensis as its type species. Records from Ocurrences in Angola (Map 294): The species is known from the southeastern part of the country. Mexico: “Rives du lac Calundo, 105 km a Lest de Luso (Lake Calundo)” [-11.80000, 20.86667] (Laurent 1964a:94). Huila: “Vila-da-Ponte” [-14.46667, 16.30000] (Monard 1937b: 113, 119; Loveridge 1939:144). Cuando Cubango: “vinicity of Cuito Cuanavale - approximately 14 km NE of Mapupa” [-17.38333, 20.08333] (Branch and McCartney 1992:2). Taxonomic and distributional notes: ll^E 20°E 24”E I0”S Map 294. Distribution of Limaformosa capensis in Angola. 336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II further north in Central Africa (Chippaux 2006; Chirio and LeBreton 2007) require confirmation (Wallach et al. 2014). Limaformosa vernayi (Bogert, 1940) Angola File Snake Mehelya vernayi Bogert 1940:28, figs, la-e, 2. Holotype: AMNH 51795 (collector A. S. Vemay, H. Lang and R. Boulton). Type locality: “Hanha” [Hanha do Norte], Benguela Province, southwestern Angola. Mehelya vernayi: Broadley (1990:99), Haacke 24“e (1981:221), Branch (1998:79), Wallach et al. ' (2014:427). Gonionotophis vernayi: Kelly et al. (2011:425). Limaformosa vernayi: Broadley et al. (2018:5). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from western Angola and northern Namibia. Ocurrences in Angola (Map 295): The species has only been recorded from the type locality “Hanha” in Benguela Province, although is probably more widespread species in the western Angola. Benguela: “Hanha” [-12.25000, 13.75000] (Bogert 1940:29; Haacke 1981:221; Wallach et al. 2014:427). Taxonomic and distributional notes: See Limaformosa capensis. Map Angola. 295. Distribution of Limaformosa vernayi in Genus Lycophidion Fitzinger, 1843 Lycophidion hellmichi Laurent, 1964 Lyeophidion hellmiehi Laurent 1964a:95, fig. 29. Holotype: MD 3824 (collector H. Baumann). Type locality: “Kapolopopo, desert de Mo^amedes ... ± 15.55 S 12.42 E, + 450 m” (coordinates provided by Laurent 1964a: 19) [= Capolopopo, Namib Desert], Namibe Province, Angola. Lyeophidion hellmiehi: Haacke (1970:281), Laurent (1968:479), Broadley (1990:95, 1991b:212, 1992b:35, 1996b: 17), Branch (1998:77), Bauer et al. (2001:78), Wallach et al. (2014:401). Lyeophidion eapense eapense: Hellmich (1957a:71). Global conservation status (lUCN): Data Deficient. Global distribution: The species is known from western Angola and northern Namibia. Ocurrences in Angola (Map 296): The species occurs in southwestern Angola. Hellmich ’s Wole Snake ll^E 1&°E 20°E 24”E Map 396. Distribution of Lycophidion hellmichi in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 337 Benguela: “Entre-Rios” [-13.01667, 14.63333] (Hellmich I957a:7I; Laurent I964a:95; Broadley I996b:I8); “Quissange” [-12.43333, 14.05000] (Broadley 1996b:18). Namibe: “Kapolopopo desert de Mo9amedes (Kapolopopo)” [-15.91667, 12.70000] (Laurent 1964a:95; Broadley 1990:95, 1991b;212, 1996b:18). Taxonomic and distributional notes: This species was described by Laurent (1964a) based on the holotype from “Kapolopopo” and a ZSM paratype from Entre Rios. The northern distribu¬ tional limits for this species remain unknown due to limited collectiong in Angola (Bauer et al. 2001). Lycophidion laterale Hallowell, 1857 Flat Wolf Snake Lycophidion laterale Hallowell 1857:58. Holotype: ANSP 10268 (eolleetor H.A. Ford). Type loeality: “Gaboon eountry” (Hallowell 1857:59), Gabon. Lyeophidion laterale'. Boeage (1866a:49, 1895a:82), Boulenger (1896:616), Ferreira (1903:10), Monard (1937b:lll), Broadley (1992b:35, 1996b:ll), Chippaux (2006:77), Wallaeh et al. (2014:401). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from central-west Africa, from Cote d’Ivoire and possibly Senegal to western Ugan¬ da and south to the Democratic Republic of Congo and northern Angola. Ocurrences in Angola (Map 297): The species is known only from two records in northwestern Angola, including the Cabinda enclave. Cabinda: “Molembo” [-5.33333, 12.20000] (Boeage 1866a:49, 1895a:82; Broa¬ dley 1996b: 11). Kwanza Norte: “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:10; Monard 1937b: 113; Broadley 1996b:ll). Taxonomic and distributional notes: None. I2"E 1&°E 20°E 24”E Map 397. Distribution of Lycophidion laterale in Angola. Lycophidion meleagre Boulenger, 1893 Speckeled Wole Snake Lyeophidium meleagris Boulenger 1893:337, pi. 21, figs. 2, 2a-2b. Leeototype: BMNH 1946.1.14.40 (formerly BMNH 43.14.8.? fide BMNH register) (eolleetor unknown) designated by Broadley (1996b:9). Type loeality: “Ambrizette” and “Ambriz” [= N’Zeto and Ambriz], northwestern Angola, restrieted by neotype designation to “Ambrizette” [= N’Zeto], Zaire Provinee, Angola. Lyeophidium meleagris'. Boeage (1895a: 82), Ferreira (1904:115), Boulenger (1915:202), Monard (1937b: 111), Hellmieh (1957b:62), Loveridge (1957:252), Frade (1963:252). Lyeophidion meleagre'. Broadley (1992b:33, 1996b: 11), Broadley (1996b: 11), Spawls et al. (2004:325), Wal¬ laeh et al. (2014:402). Global conservation status (lUCN): Not Evaluated. Global distribution: The species has an apparently disjunct distribution, with published records from northwestern Angola, southeastern Democratic Republic of Congo, southeastern Kenya and northeastern Tanzania. Ocurrences in Angola (Map 298): The species is rectricted to northwest Angola, including 338 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II the Cabinda enclave. Cabinda: “Chinxasio” (Broadley 1996b: 11) (probably Chinchoxo?). Zaire: “Ambrizette” [-7.23333, 12.86667] (Boulenger 1893:337, 1915:202; Bocage 1895a:82; Monard 1937b: 113; Loveridge 1957:252; Broadley 1996b: 11). Luanda: “Luanda” [-8.83333, 13.26667] (Broadley 1996b: 11). Bengo: “Ambriz” [-7.844312, 13.106493] (Boulenger 1893:337, 1915:202; Bocage 1895a:82; Monard 1937b:113; Loveridge 1957:252; Broadley 1996b: 11); “Cabiri” [-8.91667, 13.66667] (Ferreira 1904:115; Broadley 1996b:ll). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] (Hellmich 1957b:62; Broadley 1996b: 11). Taxonomic and distributional notes: The wide and disjunct distribution of this species calls for further taxonomic investigation. Lycophidion multimaculatum Boettger, 1888 Lycophidium Capense mut. multimaculata Boettger 1888:67. Leetotype: SMF 17973 (formerly SMF 7340, a) (eolleetor P. Hesse) designated by Mertens (1922:179). Type loeality: “bei Povo Nemlao ... bei Povo Netonna, naehst Banana,” restrieted by leetotype designation to “Banana, Unterer Kongo” [= Banana, lower Congo], Bas-Congo Provinee, Demoeratie Republie of Congo. Lycophidion Horstockii: Boeage (1866a:49, 1870:68). Lycophidion capense: Peters (1877a:615, 1881:149), Boeage (1895a:81, 1896a:112), Ferreira (1904:115, 1906:167), Boulenger (1893:616, 1905:112), Monard (1937b:117). Lyeophidion semiannulis: Ferreira (1897b:243), Monard (1937b: 113). Lyeophidion eapense eapense: Sehmidt (1933:13), Bogert (1940:30), Hellmieh (1957a:61, 1957b:71). Lyeophidion eapense multimaeulatum: Broadley (1990:94, 1991b:214), Laurent (1964a:94, 1968:474). Lyeophidion multimaeulatum: Braneh and MeCartney (1992:1), Broadley (1992b:36, 1996b: 19), Braneh (1998:77), Broadley and Cotterill (2004:48), Chippaux (2006:79), Chirio and LeBreton (2007:470), Wal- laeh et al. (2014:402). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Cameroon and Central African Republic to Tanzania, southwest to Democratic Republic of Congo, southern Angola and adjacent Zambia, and the Caprivi Strip in Namibia. Ocurrences in Angola (Map 288): The records are mostly from eastern Angola. However, the species distribution may comprise the entire country including Cabinda Province, with exception of the desert regions of the far southwestern Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Broadley 1996b:19); “Landana” [-5.21667, 12.15000] (Broadley 1996b:19); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:49, 1895a:81; Broadley 1996b:19). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:81). Luanda: “Luanda” [-8.83333, 13.26667] (Broadley 1996b: 19). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:49, 1895a:81; Boulenger 1893:616; Broadley 1996b:19); “Malanje” [-9.55000, 16.35000] (Bocage 1866a:149, 1895a:81); “Pungo-Adongo” [-9.66667, 15.58333] (Broadley 1996b: 19). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a:95, Map 298. Distribution of Lycophidion meleagre in gola. Spotted Wolf Snake MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 339 Map 299. Distribution of Lycophidion multimaculatum in Angola. 1968:474); “Calonda” [-8.41667, 20.53333] (Laurent I964a:94; 1968:474); “Lunda Dis¬ trict” (Broadley 1996b: 19). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a:95; 1968:474). Mexico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:95; 1968:474); “Macondo, Haut Zambeze” [-12.55000, 23.76667] (Laurent 1964a:95; 1968:474). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Ferreira 1906:167; Broadley 1996b: 19); “Golungo Alto” [-9.13333, 14.76667] (Ferreira 1906:167). Bie: “32 km E of Dande, Gauca” [-11.18333, 17.45000] (Schmidt 1933:13; Broadley 1996b:19); “Chitau” [-11.43333, 17.15000] (Schmidt 1933:13; Laurent 1968:474; Broadley 1996b: 19). Huambo: “Bela-Vista” [-12.36667, 16.20000] (Hellmich 1957b:61; Broadley 1996b:19); “Galanga” [-13.73333, 15.06667] (Bocage 1895a:81; Laurent 1968:474). Benguela: “Entre-Rios” [-12.26667, 17.41667] (Hellmich 1957a:61, 71; Broadley 1996b: 19); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:112). Hufla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:81; Boulenger 1893:616; Laurent 1968:474; Broadley 1996b:19); “Rio Cuce” [-13.51667, 15.20000] (Ferreira 1897b:243; Monard 1937b:113); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:117); “Gambos” [-15.76667, 14.10000] (Broadley 1996b:19); “28 km N of Sa da Bandeira” [-14.916667, 13.499914] (Broadley 1996b: 19). Namibe: “Mossamedes” [-15.20000, 12.15000] (Bocage 1895a:81; Laurent 1968:474; Broadley 1996b:19). Cuando Cubango: “approximately 50 km E of Cuito Cuanavale” [-15.23333, 19.61667] (Branch and McCartney 1992:1; Broadley 1996b: 19). Undetermined Locality: “Between Benguela and Bihe” (Boulenger 1905:112; Broadley 1996b: 19). Taxonomic and distributional notes: The majority of the specimens originally assigned to Lycophidion capense ^Smith, 1831, in Angola (Peters 1877a, 1881, Bocage 1895a, 1896a; Ferreira 1904, 1906; Boulenger 1893, 1905, Schmidt 1933; Monard 1937b:117; Bogert 1940; Hellmich 1957a, 1957b) refer to Lycophidion multimaculatum Boettger, 1888. Modem authors (e.g., Broadley 1996b; Branch 1998) have typically followed Broadley (1992b) in recognizing L. multi¬ maculatum as a full species. Given that the currently understood species boundary between this species and L. capense is suspiciously consistent with political boundaries, further investigation of populations of wolf snakes in and adjacent to the Caprivi Strip in northeastern Namibia is required. Lycophidion ornatum Parker, 1936 Ornate Wolf Snake Lycophidion ornatum Parker 1936:122. Holotype: BMNH 1946.1.14.28 (formerly BMNH 1936.8.1.690) (eol- leetor K. Jordan). Type loeality: “Congulu” [= Congulo], Kwanza Sul Provinee, Angola. Lycophidion capense ornatum'. Loveridge (1957:252). Lyeophidion ornatum'. Laurent (1968:470), Broadley (1992b:35, 1996b: 16), Spawls et al. (2004:325), Broadley and Hughes (1993:12), Chippaux (2006:80), Chirio and LeBreton (2007:474), Wallaeh et al. (2014:411). Global conservation status (lUCN): Least Concern. 340 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global distribution: The species is known from Nigeria to South Sudan and Kenya, south to Tanzania and west to the Dem¬ ocratic Republic of Congo and Angola. Ocurrences in Angola (Map 300): The species is only known from the type locality “Congulu” in Kwanza Sul Province, however, its distribution may comprise all northern regions of the country. Kwanza Sul: “Congu¬ lu” [-10.86667, 14.28333] (Parker 1936:122; Loveridge 1957:252; Laurent 1968:470; Broadley and Hughes 1993:12; Broadley 1996b: 16; Chippaux 2006:80; Wallach et al. 2014:411). Taxonomic and distributional notes: Loveridge (1957) treated L. ornatum as a sub¬ species of Lycophidion capense (Smith, 1831), but it was later elevated to specific status again by Laurent (1968). ll^E 1&°E 20“E I4«E Map 300. Distribution of Lycophidion ornatum in Angola. Genus Mehelya Csiki, 1903 Mehelya poensis (Smith, 1847) Western Forest File Snake Heterolepis poensis Smith 1847a: page 2 of 2 unnumbered pages aeeompanying plate 55 [Heterolepis capen- sisl Holotype, BMNH 1946.1.14.20 (formerly BMNH 47.4.4.? fide BMNH register) (colleetors T. Vogel and J. Ansel [Niger Expedition]). Type loeality: “Fernando Po” [= Bioko], Gulf of Guinea, Equatorial Guinea. Heterolepis biearinatus: Boeage (1866a:49). Heterolepis poensis: Ferreira (1906:168). Simoeephalus poensis: Monard (1937b: 111). Mehelya poensis: Hellmieh (1957b:61). Mehelya poensis: Loveridge (1939:145, 1957:253), Laurent (1950a: 8), Thys van den Audenaerde (1966:32), Spawls et al. (2004:332), Broadley and Cotterill (2004:48), Chippaux (2006:84), Chirio and LeBreton (2007:482), Broadley et al. (2018:6). Gonionotophis poensis: Kelly et al. (2011:425), Braneh and Conradie (2015:200), Wallaeh et al. (2014:426). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Guinea-Bissau east to South Sudan and south to the southern Democratic Republic of Congo and northern Angola. Ocurrences in Angola (Map 301): The species occurs in northern regions of Angola, including the Cabinda enclave. Cabinda: I2”E 20°E I4'E Map 301. Distribution of Mehelya poensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 341 “Cabinda” [-5.55000, 12.18333] (Loveridge 1939:146). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Thys van den Audenaerde 1966:32); “Cammbo” [-7.75294, 19.95672] (Branch and Conradie 2015:200); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent I950a:8). Bengo: “Cabi- cula (Cazengo)” [-9.16667, 13.56667] (Ferreira 1906:168; Monard I937b:II3, Loveridge 1939:146). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich I957b:6I). Taxonomic and distributional notes: Broadley et al. (2018) recently used molecular and morphological data to revise the African file snakes, splitting the genus Gonionotophis and resur¬ recting Mehelya, with poensis as its type species. Genus Polemon Jan, 1858 Polemon collaris (Peters, 1881) Collared Snake-Eater Microsoma collare Peters 1881:148. Holotype: ZMB 10045 (eolleetor F.W.A. von Meehow). Type loeality: “Cuango,” “Maeange” [= Malanje] Malanje Provinee, Angola. Microsoma collare: Boeage (1887a:182, 1895a:124), Boulenger (1896:251). Miodon collaris: Boulenger (1905:114, 1915:215), Ferreira (1906:169), de Witte and Laurent (1947:70). Miodon gabonensis: Hellmieh (1957a:72, 1957b:63). Polemon eollaris: Chippaux (2006:207), Wallaeh et al. (2014:561). Polemon eollaris eollaris: Chirio and LeBreton (2007:644). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Nigeria to Uganda and south to Angola. Ocurrences in Angola (Map 302): The zve species occurs in western Angola. Malanje: “Maeange” [-9.55000, 16.35000] (Peters 1881:148; de Witte and Laurent 1947:70; Hellmich 1957a:72; Chippaux 2006:207; Chirio and LeBreton 2007:644; Wallaeh et al. 2014:561). Kwanza Norte: “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:114; Ferreira 1906:169; de Witte and Laurent 1947:70; Hellmich 1957a:72); “Cazengo” [-9.33333, 14.76667] (Boeage 1887a:182, 1895a: 124; de Witte and Laurent 1947:70; Hellmich 1957a: 72). Huambo: “Bela- Vista” [-12.36667, 16.20000] (Hellmich 1957b:53). Benguela: “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a:124; de Witte and Laurent 1947:70; Hellmich 1957a:72); “Entre Rios” [-13.01667, 14.63333] (Hellmich Map 302. Distribution of Pctemcn cc/fam in Angola. 1957a:72). Undetermined Locality: “Cuango” (Peters 1881:148; de Witte and Laurent 1947:70, Chippaux 2006:207; Chirio and LeBreton 2007:644; Wallaeh et al. 2014:561) — Malanje Province, near the Francisco Jose Waterfalls and the border with the Democratic Republic of Congo [-7.61789, 17.27817]. Taxonomic and distributional notes: Peters (1881) mentioned both “Cuango” and “Maeange” [= Malanje] in the original description. Von Meehow collected along the Cuango River in Malanje Province, specifically near the Francisco Jose Waterfalls. Because the Cuango extends into other provinces and because there are also villages with this name, the origin of Peters’ type of Microsoma collare has sometimes been confused. 342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Prosymna Gray, 1849 Prosymna ambigua Bocage, 1873 East African Shovel-Snout Prosymna ambiguus Bocage 1873b:218. Holotype: MBL 610 (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Duque de Braganga” [= Calandula], Malanje Province, Angola. Prosymna ambigua brevis Laurent 1954a:50, figs. 12-14. Holotype: MD 2177 (collector A. Barros Macha¬ do), lost (see Notes below). Type locality: “Dundo,” Lunda Norte Province, Angola. Prosymna meleagris: Bocage (1866a:47). Prosymna ambigua: Boulenger (1893:248, 1915:208), Bocage (1895a:99), Loveridge (1933:244), Monard (1937b: 114), Wallach et al. (2014:568), Ceriaco et al. (2016b:83). Prosymna bocagii: Boulenger (1915:208). Prosymna ambigua brevis: Laurent (1964a:108), Thys van den Audenaerde (1933:34), Loveridge (1958:157). Prosymna ambigua ambigua: Laurent (1950a:9, 1954a:52), Loveridge (1958:151), Broadley (1980:534), Chippaux (2006:142). Global conservation status (lUCN): Least Concern. Global distribution; The species is known from central Cameroon, Gabon and southern Congo, east through the southern half of Democratic Republic of Congo, northern Angola and Zambia to South Sudan, southwestern Kenya and Uganda and south to Malawi and Zimbabwe. Ocurrences in Angola (Map 303): The species occurs in the northern regions of the country. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950:9, 1954a:50, 1964a: 108; Loveridge 1958:158; Thys van den Audenaerde 1966:34; Broadley 1980:538); “Sombo” [-8.68333, 20.95000] (Laurent 1954:50; Loveridge 1958:158; Broadley 1980:538). Malanje: “Duque de Braganqa” [-9.10000, 15.95000] (Bocage 1866a:47, 1873a:218, 1895a:99; Loveridge 1933:244, 1958:151-154; Monard 1937b:114; Broadley 1980:538; Chippaux 2006:142; Wallach et al. 2014:568); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:83). Taxonomic and distributional notes: Broadley (1980) synonymized the subspecies P. a. brevis Laurent, 1954 with the typical Map 303. Distribution of Prosymna amNgwa in Angola. form. In January 2017 the holotype of P. a. brevis could not be located in the Museu Dundo. Prosymna angolensis Boulenger, 1915 Angola Shovel-Snout Prosymna angolensis Boulenger 1915:208. Syntypes: MBL specimen numbers unknown, lost fide Broadley (1980). Neotype: NHMW 19275:2 designated by Broadley (1980:497). Type locality: Angola by implica¬ tion (Boulenger 1915:208), restricted to “Huila” by Loveridge (1958:149), changed to “Caconda” by subsequent designation of a neotype Broadley (1980:497, 513). Prosymna frontalis: Bocage (1873b:218, 1895a:98, pi. 11, fig. 2), Boulenger (1893:248, 1896:641). Prosymna ambigua: Monard (1931:104, 1937b: 123). Prosymna angolensis: Monard (1937b: 114, 122), Loveridge (1958:149), Bogert (1940:59), Hellmich (1957b:66), FitzSimons (1962:161), Broadley (1980:512, 1990:227), Branch (1998:84), Broadley et al. (2003:187), Wallach et al. (2014:568). Prosymna ambigua ambigua: Mertens (1938a:439), Loveridge (1958:151). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 343 Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Angola, northern Namib¬ ia including the Caprivi Strip, Zambia west of the Zambezi, northern Botswana, and western Zimbabwe. Ocurrences in Angola (Map 304): The species occurs in the southwestern Angola, extending northwards along the coast to Luan¬ da. Luanda: “Luanda” [-8.83333, 13.26667] (Broadley 1980:515). Huambo: “Bela- Vista” (Missao di Dondi) [-12.36667, 16.20000] (Hellmich 1957b:66; Loveridge 1958:154; Broadley 1980:515, 1990:229). Benguela: “Quibula” [-12.28333, 14.68333] (Bocage 1895a:98); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:98; Loveridge 1958:151); Map 304. Distribution of “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:98; Loveridge 1958:151); “Benguella” [-12.58333, 13.41667] (Bocage 1895a:98); “Eban- ga” [-12.73333, 14.73333] (Monard 1937b:122; Loveridge 1958:151; Broadley 1980:515); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:439; Loveridge 1958:151; Broadley 1980:515). Huila: “Capelongo” [-14.91667, 15.08333] (Bogert 1940:59; Loveridge 1958:153-154; Broadley 1980:515); “Caconda” [-13.73333, 15.06667] (Bocage 1895a:98; Loveridge 1958:151; Broadley 1980:515, 1990:227, 229); “Vila-da-Ponte” [-14.46667, 16.30000] (Monard 1937b:123; Loveridge 1958:154; Broadley 1980:515); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:98; Loveridge 1958:149, FitzSimons 1962:161). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1873a:218, 1895a:98; Loveridge 1958:151); “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:98; Loveridge 1958:151; Broadley 1980:515); “Mossamedes” [-15.20000, 12.15000] (Bocage 1973b:218, 1895a:99; Boulenger 1893:248; Loveridge 1958:151; Broadley 1980:515). Undetermined Local¬ ity: “no locality” (Broadley 1980:515). Taxonomic and distributional notes: Boulenger (1915) applied the new name Prosymna angolensis to a series of Angolan snakes listed under the name Prosymna frontalis (Peters, 1867) by Bocage (1873b, 1895a). Boulenger did not nominate any type locality other than Angola but Loveridge (1958) subsequently restricted the type locality to “Huila.” Broadley (1980) reviewed the species. He had visited the Museu Bocage collection in Lisbon in 1968 and did not find any specimens from Bocage’s series from “Huila”, but did locate four individuals from “Caconda.” Although these specimens had been destroyed before Broadley ’s (1980) review, he restricted the type locality to “Caconda,” by designating a neotype from this locality in the Naturhistorisches Museum Wien. Angolan records from “Vila-da-Ponte” (Monard 1931, 1937b), “Cubal” (Mertens 1938a), and “Capelongo” (Loveridge 1958) previously identified as belonging to Prosymna ambigua (Bocage, 1873) are probably misidentifications and should be assigned to P angolensis. Il'E 16°E Prosymna frontalis (Peters, 1867) South-Western Aerican Shovel-Snout Temnorhynchus frontalis Peters 1867a:236, pi. Leeototype: ZMB 5763a (eolleetor C.H. Hahn), designated by Mertens (1955:94). Type loeality: “Otjimbingue in Sudwestafriea” [= Otjimbingwe, Erongo Region], Namibia. 344 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Prosymna frontalis: Broadley (1980:517, 1990:229), Branch (1998:85), Bates et al. (2014:389), Wal- lach et al. (2014:569). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southwestern Angola to Namibia, southwards to the Northern Cape, Souh Africa. Ocurrences in Angola (Map 305): The species occurs in southwestern Angola. Benguela: “Benguela” [-12.58333, 13.41667] (Broadley 1980:519). Taxonomic and distributional notes: This species was recorded for Angola by Bocage (1873a, 1895a) and Boulenger (1893, 1896), but these records likely represent Prosymna angolensis Boulenger, 1893 (see P. angolensis account). Broadley (1980) cited one specimen from “Benguela” deposited in the um of Natural History, Pretoria, South Africa. Il'-E 16“E 20“E 24'‘E Map 305. Distribution of Prosymna frontalis in Angola. Transvaal Musuem, now Ditsong National Muse- Prosymna visseri FitzSimons, 1959 Visser’s Shovel-Snout Ocurrences in Angola (Map 306): The species’ distribution range comprises Benguela and Namibie Provinces as well the adjacent areas of Huila and Cunene Provinces. Benguela: “20 km S of Balabaia” [-12.01667, 13.96667] (Broadley 1980:544). Huila: “5 km S of Chibemba” [-15.785418, 14.106551] (Broadley 1980:544). Namibe: “near Caracul, S. Angola” [-15.01667, 12.66667] (FitzSimons 1959:408; Broadley 1980:544; Bauer et al. 2001:76; Wallach et al. 2014:571). Taxonomic and distributional notes: This species was previously considered an Angolan endemic (e.g., Broadley 1980) but it has since been found at numerous localities in arid to semi-arid northwestern Namibia. Prosymna visseri FitzSimons 1959:408. Holotype: TM 24531 (collector C. Koch). Type locality: “near Cara¬ cul, S. Angola” (FitzSimons, 1959:408) [= near Caraculo], Namibe Province, Angola. Prosymna visseri: Broadley (1980:543), Branch (1998:85), Bauer et al. (2001:76), Branch (1998:85), Wallach et al. (2014:571). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is ii»e i&°e 20“e 24"e known from southwestern Angola and northern Namibia. Map 306. Distribution of Prosymna visseri in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 345 Genus Psammophis Bole, 1825 Psammophis angolensis (Bocage, 1872) Dwarf Sand Snake Amphiophis angolensis Bocage 1872:82. Holotype: MBL 1822 (collector F.A.P. Bayao), destroyed by fire 18 March 1978. Type locality: “Dondo dans I’interieur d’Angola” [= Dondo], Kwanza Norte, Angola. Ablabes Homeyeri Peters (1877a:620). Holotype: ZMB 9209 (collector A. von Homeyer). Type locality: “Pungo Andongo,” Malanje Province, Angola. Amphiophis angolensis: Bocage (1895a: 113, 1897a:201). Psammophis angolensis: Boulenger (1896:170), Boettger (1898:104); Boulenger (1915:213); Schmidt (1933:14), Loveridge (1933:252, 1940:68, 1957:280), Laurent (1950a:9, 1954a:59, 1964a: 114), FitzSi- mons (1962:235), Frade (1963:253), Auerbach (1987:171), Brandstatter (1996:40), Broadley (1962a:834, 1977b:26, 1990:148, 2002a:98), Broadley and Cotterill (2004:50), Spwals et al. (2004:403), Kelly et al. (2008:1053), Bates et al. (2014:374), Wallach et al. (2014:575). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is widespread in sub-Saharan Afriea, from Ethiopia and Tanzania, through the Democratic Republic of Congo, through Zambia, Malawi, Mozambique, westwards to Angola and southwards through northern Namibia, Botswana, and Zimbabwe to northeastern South Africa (Limpopo and Mpumalanga provinces). Ocurrences in Angola (Map 307): The species occurs in the entire country with excep¬ tion of the Namibe Province and the Cabinda enclave. Zaire: “Ambrizette” [-7.23333, 12.86667] (Bocage 1895a;113, 1897a;201; Boettger 1898:104; Loveridge 1940:68). Kwanza Norte: “Dondo” [-9.68333, 14.43333] (Bocage 1872:82, 1895a:113, 1897a:201; Loveridge 1933:252, 1940:68, 1957:280; FitzSimons 1962:235; Auerbach 1987:171, Broadley 1962a:834, 1977b:26, 1990: 148, 2002a: 1 1 8; Wallach et al. 2014:575). Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1881:149; Bocage 1895a: 113, 1897a:201; Loveridge 1940:68); “Pungo- Andongo” [-9.66667, 15.58333] (Peters 1877a:620; Bocage 1895a:113, 1897a:201; Loveridge 1940:68, 1957:280; Broadley 307 Disuibution of 1977b:26, 2002a: 11 8). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Bocage 1895a:113; Laurent 1950a:9, 1954a:59, 1964a:114; Loveridge 1957:280). Mexico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:114); “Reserve de chasse de Cameia” [-11.83333, 21.00000] (Laurent 1964a:114). Kwanza Sul: “Novo Redondo” [-11.20000, 13.85000] (Bocage 1895a: 113; Loveridge 1940:68). Benguela: “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:113, 1897a:201; Loveridge 1940:68). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:113, 1897a:201; Loveridge 1940:68). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a:113, 1897a:201; Schmidt 1933:14; Loveridge 1940:68). Taxonomic and distributional notes: The type locality, Dondo, has been variously misiden- tified as “Danda, Loanda Distr.” (Loveridge 1940, Broadley 1962a) “Donda, Loanda Distr.” (Auer¬ bach 1987), “Donda i.e. Dundo, Loanda distr.” (FitzSimons 1962a; Broadley 1990) or even 346 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Dondo, Luanda” (Crawford-Cabral and Mesquitela 1989). Psammophis angolensis is a wide¬ spread species with several peculiar autapomorphies. It was originally described in the genus Amphiophis Bocage, 1872, and Brandstatter (1995) suggested that this should be revived as a monotypic subgenus. However, Kelly et al. (2008) did not follow this suggestion in their phyloge¬ netic anaylsis of Psammophis. Psammophis ansorgii Boulenger, 1905 Link-Marked Sand Racer (Endemic) Psammophis Ansorgii Boulenger 1905:113, pi. 4, fig. 4. Holotype: BMNH 1946.1.8.49 (formerly BMNH 1905.5.29.31) (eolleetor W.J. Ansorge). Type loeality: “between Benguella and Bihe,” Angola. Psammophis jallae: Loveridge (1940:62). Psammophis ansorgii'. Hellmieh (1957b:69), Broadley (1977b:26, 2002a:98), Brandstatter (1996:42), Wallaeh et al. (2014:576). ii^e 2o°e 24"e Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola. Ocurrences in Angola (Map 308): The species is endemic to the central highlands of Angola. Huambo: “Bela-Vista” [-12.36667, 16.20000] (Hellmieh 1957b:69). Undeter¬ mined locality: “Between Benguella to Bihe” (Boulenger 1905:113; Loveridge 1940:62; Broadley 1977b:26, 2002a:98; Wallaeh et al. 2014:576). Taxonomic and distributional notes: Loveridge (1940) placed Psammophis ansorgii in the synonymy of Psammophis jallae Perac- ca, 1896, but Broadley (1977b) considered it a full species, as have subsequent authors. Map 308. Distribution of Psammophis ansorgii in Angola. Psammophis jallae Peracca, 1896 Jalla’s Sand Snake Psammophis jallae Peracca 1896:2, 2 figs. Syntypes: MZUT 1633 and 1871 (formely MZUT 1823a-b) (collector L. Jalla). Type locality: “Strada da Kazungula a Buluwaio” (Peracca 1896:2), [= road from Kazungula, Zambia to Bulawayo], Zimbabwe. Psammophis Rohani Angel (1922:116). Type: MNHP 20-198 (collector J. F. de Rohan-Chabot). Type locali¬ ty: “Region de la riviere Lumuna, affluent de Loengoue” [= Lumuna River, tributary of the Luina and Quando Rivers], Angola. Psammophis rohani'. Angel (1923:166), Monard (1937b: 128). Psammophis jallae'. Loveridge (1940:62), FitzSimons (1962:237), Brandstatter (1996:65), Broadley (1977b: 12, 1990:138, 2002a:93), Branch (1998:90), Bates et al. (2014:376), Wallaeh et al. (2014:576). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from central and southeastern Angola, western Zambia, northeastern Namibia, Botswana, western and central Zimbabwe and Limpopo Province of South Africa. Also from the southern portion of the former Katanga Province, Democratic Republic of Congo {fide Wallaeh et al. 2014). Ocurrences in Angola (Map 309): The species occurs in central and southeastern Angola. Bie: “Bigondo” [-12.06667, 17.41667] (Loveridge 1940:64); “Benguella to Bihe” (Loveridge MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 347 1940:64). Cuando Cubango: “Lumuna regions, Loengoue affluent” (Angel 1921:118; 1923:168; Monard 1937b:128; Loveridge 1940:62; Broadley 1977b: 12, 2002a:93). Taxonomic and distributional notes: Angel (1923) provided three views of the head (figs. 10-12) of the speeies he deseribed the year previously. The date of deseription has been given as 1921, but Angel (1923) explieit- ly stated that the assoeiated issue of the Bul¬ letin de la Soeiete Zoologique de Franee had appeared only in Mareh 1922. Psammophis rohani was synonymized with P. jallae Perraea, 1896 by Loveridge (1940). Psammophis leopardinus (Bocage, 1887) Psammophis sibilans van leopardinus Bocage 1887a:206. Lecototype: MBL 1798 (collector J.A. d’ Anchieta), designated hy Broadley (1977b: 18), destroyed by fire 18 March 1978. Type locality: “Catumbella” (Bocage 1887b:206) [= Catumbela] Benguela Province, Angola. Psammophis sibilans van C leopardina: Bocage (1895a: 117). Psammophis sibilans van E brevirostris: Bocage (1895a: 11 8). Psammophis brevirostris: Boulenger (1915:213); Monard (1937b: 133), Themido (1941:10). Psammophis brevirostris brevirostris: Brandstatter (1996:45). Psammophis sibilans sibilans: Loveridge (1957:279). Psammophis sibilans leopardinus: Broadley (1977b: 18). Psammophis brevirostris leopardinus: Brandstatter (1996:48), Branch (1998:91). Psammophis leopardinus: Broadley (1990:143, 2002a:95), Kelly et al. (2008:1048), Hughes and Wade (2002:75), Bates et al. (2014:374), Wallach et al. (2014:577). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from dry savanna and semi-desert regions in southwestern Angola and central and northern Namibia. Ocurrences in Angola (Map 310): This species occurs chiefly in southwestern Angola. Luanda: “Luanda” [-8.83333, 13.26667] (Broadley 2002:111; Hughes and Wade 2002:77). Huambo: “Bella Vista (Missao di Dondi)” [-12.36667, 16.20000] (Broadley 2002:111; Hughes and Wade 2002:77); “Huambo” [-12.76667, 15.73333] (Themido 1941:10). Benguela: “Lobito bay” [-12.35000, 13.5500] (Broadley 1977b:19, 2002a:lll; Hughes and Wade 2002:77); “Catum¬ bella” [-12.43333, 13.55000] (Bocage 1895a:117; Loveridge 1957:279; Broadley 1990:143, 2002a:lll; Hughes and Wade 2002:77); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:117); “Catengue” [-13.03333, 13.73333] (Broadley 2002:111; Hughes and Wade 2002:77). Huila: “Caconda” [-13.73333, 15.06667] (Broadley 2002:111; Hughes and Wade 2002:77); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:133; Broadley 2002:111); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:114); “Vila da Ponte” [-14.46667, 16.30000] (Monard 1937b:133; Broadley 2002:111); “Capelongo” [-14.88333, 15.08333] (Hughes and Wade 2002:77). Namibe: “Capangombe” [-15.10000, 13.15000] (Bocage 1887b:206; Broadley 2002:111); “Mossamedes” I2"E 16.°E 2«°E Map 309. Distribution of Psammophis jallae in Angola. Leopard Sand Snake 348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II [-15.20000, 12.15000] (Bocage I887b:206, I895a:II4; Loveridge 1957:279); “Iona” [-16.28106, 12.22291] (Broadley 2002:111; Hughes and Wade 2002:77). Cunene: “Oncocua, 37 km NE on way to Otchinzau” [-16.65606, 13.42370] (Broadley 2002:111; Hughes and Wade 2002:77). Cuando Cuban¬ go: “Kayundu” [-15.70000, 17.45000] (Monard 19376:133). Undetermined Locali¬ ty: “Cuanza” (Broadley 2002:111). Taxonomic and distributional notes: In addition to his Catumbella specimen, Bocage (1887b) based his description on a second specimen from, “Tinterieur de Mossamedes” collected by Capello and Ivens. Brandstatter (1996) recognized P. sibilans as occurring no further south than the northern part of Tanzania and proposed the reinstatement of P. brevi- rostris Peters, 1881 as a full species, rather than a subspecies of P. sibilans, with leopardinus treated as a subspecies of it. Currently P. leopardinus is recognized as a valid species (Broadley 1977b, 2002a; Kelly et al. 2008; Branch 1998; Wallach et al. 2014). The record from “Calombe” has been assigned to P. leopardinus although the habitat does not correspond to the typical dry savanna and semi-desert regions where the species is com¬ monly found. Psammophis mossambicus Peters, 1882 Olive Whip Snake Psammophis sibilans var. mossambica Peters 1882e:122. Leetotype: ZMB 2468A, designated by Broadley (2002:96). Type loeality: “auf der Insel Mossambique und auf dem gegenuberliegenden Festlande, auf der Halbinsel Caba^eira und Mesuril [...] auf den Querimba-Inseln [...] und in Boror” (eolleetor W.C.H. Peters), restrieted to “der Insel Mossambique” [= Mozambique Island] by leetotype designation. Psammophis sibilans: Boeage (1866a:48, 1895a: 114, 1896a: 113), Peters (1877a:615), Ferreira (1904:116), Boulenger (1905:113, 1915:213), Sehmidt (1933:14), Monard (1937b: 131), Loveridge (1936a:38). Psammophis elegans: Boeage (1867b:226). Psammophis sibilans sibilans: Mertens (1938a:441), Bogert (1940:70), Loveridge (1940:30), Laurent (1964a: 113), Hellmieh (1957b:70), Managas (1973:196). Psammophis phillipsii: Broadley (1977b:24), Braneh and MeCartney (1992:2), Brandstatter (1996:55), Hugh¬ es (1999:64), Kelly et al. (2008). Psammophis mossambicus: Braneh (1998:92), Broadley (2002:96), Broadley et al. (2003:167), Broadley and Cotterill (2004:50), Kelly et al. (2008:1048), Wallaeh et al. (2014:577), Ceriaeo et al. (2016b:87), Conradie et al. (2016:22). Psammophis ef mossambicus: Braneh and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from South Sudan to South Africa, west through Swaziland in the east, in the west to northern Namibia (see Notes). Ocurrences in Angola (Map 311): The species occurs throughout the country, including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Loveridge 1940:30); “Cabinda” [-5.55000, 12.18333] (Ferreira 1904:116; Loveridge 1940:30). Zaire: “San Antonio” [-6.13333, 12.36667] (Loveridge 1940:30). Bengo: “Ambriz” [-7.844312, 13.106493] ll«E 2rE 24" E Map 310. Distribution of Psammophis leopardinus in Angola. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 349 (Loveridge 1940:30; Broadley 2002:113). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage I866a:48, 1 895a: 1 14; Loveridge 1940:30); “Loanda island” [-8.83333, 13.26667] (Loveridge 1940:30; Broadley 2002:113); “Quieamba Park” [-9.18333, 13.38333] (Broadley 2002:113). Kwanza Norte: “Cazengo” [-9.33333, 14.76667] (Fer- reria 1904:116; Loveridge 1940:30). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Boulenger 1905:113; Loveridge 1940:30; Broadley 2002:113); “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1905:113; Loveridge 1940:30; Broadley 2002:113); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016b:87); “Mulon- do, Luanda Res., Malanje” (Broadley 2002:113). Lunda Sul: “Alto Chicapa” 311. Distribution of [-10.88333, 19.23333] (Laurent 1964a:113). Mexico: “Lago Dilolo” [-11.50000, 22.01667] (Managas 1973:196); “evirons du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:113); “Calombe, Luso” [-11.83333, 19.93333] (Mananas 1973:196); “reserve de chasse de Cameia” [-11.83333, 21.00000] (Laurent 1964a:113); “Cazom- bo” [-11.88333, 22.91667] (Laurent 1964a:113). Kwanza Sul: “Libolo/Luati” [-9.98333, 14.90000] (Hellmich 1957b:70); “Chingo” [-11.20000, 13.85000] (Ferreira 1904:116). Bie: “Chi- tau” [-11.43333, 17.15000] (Schmidt 1933:14; Loveridge 1940:30; Broadley 2002:113); “Cuban¬ go basin (16b)” [-12.54222, 16.67694] (Conradie et al. 2016:8,9, 22). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a: 114; Loveridge 1940:30); “Bela- Vista (Sanguengue)” [-12.36667, 16.20000] (Hellmich 1957b:70); “Chipipa 13 kmN, Hamubo” [-12.55000, 15.73333] (Broadley 2002:113); “Cuma” [-12.85889, 15.06722] (Loveridge 1936a:38, 1940:30; Broadley 2002:113). Benguela: “Lobito bay” [-12.33333, 13.58333] (Bogert 1940:79); “Catumbella” [-12.43333, 13.55000] (Loveridge 1940:30); “Quissange” [-12.43333, 14.05000] (Loveridge 1940:30); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:226); “Ebanga” [-12.73333, 14.73333] (Monard 1937b:131, Loveridge 1940:30); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:441; Loveridge 1940:30; Hellmich 1957b:70; Broadley 2002:113); “Katange” [-13.03333, 13.73333] (Loveridge 1940:30); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113; Loveridge 1940:30; Broadley 2002:113); “Capaia to Cubal” (Broadley 2002:113). Huilla: “Rio Cuce” [-13.51667, 15.20000] (Bocage 1895a:114; Loveridge 1940:30); “Caconda” [-13.73333, 15.06667] (Bocage 1895a:114; Loveridge 1940:30); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:131; Loveridge 1940:30); “Quillengues” [-14.06667, 15.08333] (Loveridge 1940:30); “Kuvangu” = “Vila-da-Ponte” [-14.46667, 16.30000] (Monard 1937b:131; Loveridge 1940:30); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:70; Broadley 2002:113); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:114; Loveridge 1940:30); “Tchiuinguira, Huila” [-15.16667, 13.30000] (Broadley 2002:113); “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] (Laurent 1964a:113), “Negola 6 km S, Huila” (Broadley 2002:113). Namibe: “Mossamedes” [-15.20000, 12.15000] (Loveridge 1940:30; Broadley 2002:113); “Rio Coroca” [-15.78333, 12.06667] (Bocage 1895a:114); “Port Alexander” [-15.80000, 11.83333] (Loveridge 1940:30). Cunene: “fl. Mbale” [-15.16667, 16.75000] (Monard 1937b:131); “Mupa” [-16.18333, 15.75000] (Monard 1937b:131; I2"E 16“E 20“E 24<-E 350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Loveridge 1940:30); “Mupanda” [-17.13333, 15.76667] (Monard I937b:I3I; Loveridge 1940:30); “Tala Kilau, Mossamedes” (Broadley 2002:113). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b:131; Loveridge 1940:30); “Kayundo” [-15.70000, 17.45000] (Loveridge 1940:30); “vineity of Cuito Cuanavale - approximatley 75 km W of Mavinga” [-15.78333, 19.70000] (Branch and McCartney 1992:2); “Cubango basin (48)” [-14.68136, 17.44530] (Conradie et al. 2016:9, 12, 22). Undetermined Locality: “without precise location” (Laurent 1954:59, 1964a: 113); “Cascalada 6 km NW” (Broadley 2002:113); “Marimo 10 km W of Machado” (Broadley 2002:113). Taxonomic and distributional notes: The taxonomy of the Psammophis sibilans complex, to which this species belongs, remains confused (Kelly et al. 2008; Wallach et al. 2014) and we have here treated Angolan specimens previously refered to P. sibilans (Linnaeus, 1758) (except those attributed to P. zambiensis) and P. phillipsii (Hallowell, “1844” 1845) as P mossambicus (e.g., Loveridge 1953; Broadley 1977b; Broadley et al. 2003). Brandstatter (1996) and Hughes (1999) considered P phillipsii to be restricted to West Africa, and Branch (1998) was the first to use the name P. mossambicus Peters, 1882 to refer to the former southern and East African populations of P. phillipsii. Broadley (2002a) likewise recognized that the Angolan material initially assigned to P. sibilans and P. phillipsii should be attributed to P. mossambicus. Psammophis namibensis Broadley, 1975 Namib Sand Snake Psammophis leightoni namibensis Broadley 1975b:9, pi. 1. Holotype: TM 37093 (collector W.D. Haacke). Type locality: “Hams in the Uri-Hauchab Mountains, Diamond Area No. 2, South West Africa” [= Karas Region], Namibia. Psammophis leightoni namibensis: Broadley (1977b: 11, 1990:136), Brandstatter (1996:68), Branch (1998:90). Psammophis namibensis: Broadley (2002:91), Bates et al. (2014:378), Wallach et al. (2014:577). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known scrubland, occurring along the southwest Africa, Namib Desert into western Namibia, to Namaqualand, in the Northern Cape Province, South Africa. Ocurrences in Angola (Map 312): The species occurs in the southwestern Angola in the Namib Desert. Namibe: “Mossamedes” [-15.20000, 12.15000] (Broadley 1975b:ll; 2002a: 107); “Pico Acezevedo” [-15.55000, 12.51667] (Broadley 2002:107); “Rio Coroca” [-15.78333, 12.06667] (Broadley 1975b:ll; 2002a:107); “Foz do Cunene” [-17.28333, 11.80000] (Broadley 1975b: 11; 2002a: 107); “Cunene Forde, 15 km NE, Iona Res.” (Broadley 2002:107). Taxonomic and distributional notes: Broadley (1975b) initially described this form as a new subspecies of Psammophis leightoni Boulenger, 1902, but later considered it a full species (Broadley 2002). Kelly et al. (2008), from arid regions, including desert and succulent from southwestern Angola extending through the II^E Ili'E 20^E 24'*E Map 312. Distribution of Psammophis namibensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 351 however, noted that Psammophis namibensis and Psammophis leightoni might represent a single species (Bates et al. 2014; Wallach et al. 2014). Psammophis notostictus Peters, 1867 Karoo Sand Snake Psammophis moniliger var. notostictus Peters 1867a:237. Syntypes: ZMB 5756a-b (eolleetor C.H. Hahn). Type loeality: “Otjimbingue in Sudwestafriea” [= Otjimbingwe, SE Karibib Distriet, Erongo Region], Namibia. Psammophis sibilans var. stenocephalus Boeage (1887a:205). Holotype: MBL speeimen, number unknown (eolleetors H.C. Capello and R. Ivens). Type loeality: “interieur de Mossamedes” (Boeage 1887b:205), eorreeted to “Rio Coroea” by Boeage (1895a: 116), Namibe Provinee, Angola. Psammophis sibilans var. B (var. stenocephala): Boeage (1895a: 116). Psammophis notostictus: Boulenger (1895a:538, 1896:156, 1915:213), Monard (1937b: 128), FitzSimons (1962:223), Frade (1963:253), Broadley (1975b:13, 1977b:10, 2002a:87), Brandstatter (1996:77), Braneh (1998:90), Bates et al. (2014:377); Wallaeh et al. (2014:577); Ceriaeo et al. (2014b:41). Psammophis sibilans notostictus: Loveridge (1940:44). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Karroid bushveld, Namib Desert and fynbos areas from southwestern Angola, south through Namibia to the Cape Peninsula and eastwards through the Karoo to the Albany District, Eastern Cape Province. Ocurrences in Angola (Map 313): The species occurs in the southwestern Angola in the Namib Desert. Namibe: “Espinheira” [-16.78717, 12.35764] (Ceriaeo 2016b: 41); “Pico Azevedo” [-15.53400, 12.49197] (Ceria- co 2016b: 41); “Sao Nicolao River” [-14.26667, 12.36667] (Boulenger 1896:156; Loveridge 1940:44; Broadley 1975a: 14, 2002a; 103); “interieur de Mossamedes (=Rio Coroea)” [-15.78333, 12.06667] (Boeage 1887a;205, 1895a:116; Monard 1937b:128; Loveridge 1940:44; Broadley 1975b: 14, 1977b:10, 2002a;87,103). Taxonomic and distributional notes: Psammophis sibilans var. stenocephalus Boeage 1887, was was synonymized with Psammophis notostictus Peters, 1867 by Boulenger (1895a). Psammophis notostictus was described as a variety of Psammophis moniliger (Daudin, 1803) by Peters (1867a) but was also treated as a variety of P. sibilans (Linnaeus, 1758) by several authors (see Broadley 1975b). Boeage (1895a) referred to P. sibilans var. B, but in the same account equated this with his P. s. var. stenocephalus Boeage, 1887. Boulenger (1896) treated P. notostictus as a full species, and was followed by several authors until Loveridge (1940) again considered it as a subspecies of P sibilans. Mertens (1955) restored notostictus to its current specific rank. Map 313. Distribution of Psammophis notostictus in Angola. Psammophis subtaeniatus Peters, 1882 Stripe-Bellied Sand Snake Psammophis sibilans var. subtaeniata Peters 1882e:121. Leetoype: ZMB 9992A designated by Broadley (1977b: 13). Type loeality: “sowohl im Innem des Landes bei Tette, als naher der Kiiste, wie bei Boror,” 352 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II restricted to “Tete, Mogambique” by Broadley (1966b). Psammophis bocagii Boulenger (1895a:538). Syntype: BMNH 67.7.23.22 (collector Donaldson Smith). Type locality: “Angola” later given as “Benguella,” Angola (Boulenger 1896:161). Psammophis sibilans var. A (var. subtaeniata): Bocage (1895a: 116). Psammophis bocagii: Boulenger (1896:161, 1915:213), Monard (1937b: 131), Bogert (1940:82). Psammophis Bocagii: Bocage (1897a:201). Psammophis subtaeniatus subtaeniatus: Loveridge (1940:55), Broadley (1966b:6, 1977b: 13), Brandstatter (1996:99). Psammophis subtaeniatus: Branch (1998:91), Broadley (1990:342, 2002b:93), Bates et al. (2014:379), Wallach et al. (2014:580), Conradie et al. (2016:22). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from southern Angola and northern Namibia, east through Botswana to southern Zambia, Zimbabwe, parts of western Mozambique, the northeastern provinces of South Africa and eastern Swaziland. Ocurrences in Angola (Map 314): The species occurs in southern Angola. Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1895a:115, 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Benguella” [-12.58333, 13.41667] (Boulenger 1896:161; Loveridge 1940:55; Broadley 1966b:8, 1977b: 13, 2002a:108). Huila: “Molundo” [-15.01667, 13.20000] (Monard 1937b:131; Loveridge 1940:55). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a:115, 1897a:201; Loveridge 1940:55); “Chao de Chella” [-14.88944, 13.27417] (Broadley 2002:108); “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:115, 1897a:201; Loveridge 1940:55; Broadley 2002:108). Cunene: “Mupa” [-16.18333, 15.75000] (Bocage 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Cahama 5 km SE” [-16.28333, 14.30000] (Broadley 2002:108); “Humbe” [-16.68333, 14.90000] (Bocage 1895a:115, 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Forte Roqadas” [-16.71667, 15.01667] (Monard 1937b:131; Loveridge 1940:55; Broadley 2002:108); “Dongwenna, Mossamedes” [-17.01667, 14.71667] (Broadley 2002:108). “Mupanda” [-17.13333, 15.76667] (Monard 1937b:131; Loveridge 1940:55; Broadley 2002:108); “Cunene” (Bocage 1895a: 114; Loveridge 1940:55; Broadley 2002:108). Cuando Cubango: “Cuito drainage” [-17.57333, 23.26000] (Conradie et al. 2016:9-10, 22). Undetermined Locality: “Cuanza” (Bocage 1895a: 11 5); “Rio Bengo” (Bocage 1895a: 11 5, 1897a:201; Loveridge 1940:55; Broadley 2002:108); “Pen Pen” (Broadley 2002:108). Taxonomic and distributional notes: Psammophis subtaeniatus has been known by a variety of names, Boulenger (1895a) restricted the name subtaeniatus to the eastern form and erected a new name, Psammophis bocagii (Boulenger 1895), for the western race based on specimens from Angola (Broadley 1977b). The description appears as part of a key and no precise locality or num¬ ber of specimens is given, however, the fact that a range of scale counts is provided demonstrates that there was originally a syntype series. Boulenger (1896) indicated that the single BMNH spec¬ imen had come from Bocage in Lisbon, so it is likely that at least one syntype remained in the MBL Map 314. Distribution of Psammophis subtaeniatus in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 353 collection, perhaps not marked as such. Bocage (1895a) referred to P. sibilans var. A, but in the same aceount equated this with P. s. var. subtaeniatus Peters, 1882. Loveridge (1940) used P subtaeniatus subtaeniatus for the western form and placed P. bocagii in its synonymy. Psammophis trigrammus Gunther, 1865 Western Sand Snake Psammophis trigrammus Gunther 1865b:95, pi. 2, fig. e. Holotype: BMNH 1946.1.8.12 (formerly BMNH 64.7.11.4) (eolleetor J. Monteiro). Type loeality: “on the banks of river S. Nieolao (Little Fish Bay, West Afriea)” [= Rio Sao Nieolau], Namibe Provinee, Angola. Psammophis trigrammus: Boeage (1887b:206), Loveridge (1940:23), FitzSimons (1962:221), Broadley (1977b:9, 1990:133, 2002a:87), Brandstatter (1996:104), Braneh (1998:89), Bates et al. (2014:379), Wallaeh et al. (2014:581). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from southern Angola and western Namibia, reaching its southern limit in the extreme north of the Richtersveld, South Africa. Ocurrences in Angola (Map 315): The species is restrieted to Namibe Province. Namibe: “Catara River” [-13.55000, 12.55000] (Broadley 2002:103); “on the banks of river S. Nieolao (Little Fish bay. West Afri- ka)” [-14.26667, 12.36667] (Gunther 1865b;95); “Rio de Sao Nieolau, Mossamedes (Rio San Nieolau)” [-14.26667, 12.36667] (Bocage 1887b:206; Loveridge 1940:23; FitzSimons 1962:221; Broadley 1977b:9, 1990:133, 2002ba:86, 103; Wallaeh et al. 2014:581); “Rio Coroca” [-15.78333, 12.06667] (Broadley 2002:103); “Iona Reserve, 7 km to Oncocua” [-16.85831, 12.61275] (Broadley 2002:103). Taxonomic and distributional notes: Bocage (1887b) corrected the type locality to “Mossamedes” [= Rio Sao Nieolau, Namibe]. 20°E 24'E Map 315. Distribution of Psammophis trigrammus in Angola. Psammophis zambiensis Hughes and Wade, 2002 Zambian Whip Snake Psammophis zambiensis Hughes and Wade 2002:75. Holotype: BMNH 1959.1.1.81 (eolleetor H.J. Bredo). Type loeality: “supposedly from “Abereom” (= Mbala) area of Zambia . . . but likely to be from Mwem- Wantipa”. Psammophis sibilans sibilans: Laurent (1950a:9, 1954a:59), Thys van den Audenaerde (1966:34). Philothamnus irregularis irregularis: Managas (1973:191). Psammophis zambiensis: Broadley and Cotterill (2004:49), Wallaeh et al. (2014:582). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known with certainty from the southern Demoeratic Republic of Congo and northern Zambia. Literature records of Psammophis sibilans that may be referable to P. zambiensis exist from the Caprivi Strip of Namibia and adjacent northern Botswana, northern Zimbabwe, Malawi, Zambezia Province of Mozambique and northestem Angola. Ocurrences in Angola (Map 316): None eonfirmed but Psammophis zambiensis is expected to occur in northeastern regions of the country. The following records are P. ^sibilans" records that 354 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II may be referable to P. zambiensis. Mexico: “Calombe” [-11.83333, 19.93333] (Managas 1973:191). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:9, 1954a:59, 1964a: 113; Thys van den Audenaerde 1966:34); “Barrage de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaer¬ de 1966:34); “Dundo, R. Mussungue, aff. Luachimo” [-7.41667, 20.83333] (Thys van den Audenaerde 1966:34); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1954a:59). Taxonomic and distributional notes: Hughes and Wade (2002) found significant dif¬ ferences between specimens of '‘Heopardinus^^ from Angola and from Zambia and described P. zambiensis to accommodate the latter. Some eastern records of P. leopardinus could corre¬ spond to this new species. Hughes and Wade (2002) presented information from Desmond Vesey-FitzGerald via Donald G. Broadley that substantiated the likely origin of the holotype from Mweru- Wantipa in the Mporokoso District [Zambia]. Hughes and Wade (2002) plotted localities of Psammophis 'sibilans' from the literature, which could include material assignable to P. zambi¬ ensis. Two such Angolan records correspond to literature references from Laurent (1950, 1954a, 1964a) and Thys van den Audenaerde (1966). Wallach et al. (2014) included only the former province of Katanga (Democratic Republic of Congo) and northern Zambia in the distribution of the species, corresponding to the verified records given in the description. Genus Psammophylax Fitzinger, 1843 Psammophylax acutus (Gunther, 1888) Striped Beaked Snake Psammophis acutus Gunther 1888:327, pi. 19, fig. d. Holotype: BMNH 1946.1.2.81 (formerly BMNH 1864. 7.13.36) (collector F.M.J. Welwitsch). Type locality: “Pungo Andongo,” Malanje Pronvince, Angola. Psammophis oxyrhynchus: Gunther (1865a:480, 1895:89). Rhagerhis acuta: Bocage (1895a: 111). Rhamphiophis acutus: Boulenger (1896:148, 1905:113, 1915:212), Loveridge (1933:252), Hellmich (1957b:71), Managas (1973:195), Broadley et al. (2003), Spawls et al. (2004:398), Chippaux (2006:170). Rhamphiophis acutus {Rhagerhis acuta): Monard (1937h:128). Rhamphiophis acutus wittei: Laurent (1964a: 111). Rhamphiophis acutus acutus: Loveridge (1957:277), Laurent (1964a: 111), Broadley (1971h:3), Chirio and Ineich (1991:220). Rhamphiophis acutus jappi: Broadley (1971d:4). Psammophylax tritaeniatus tritaeniatus: Mananas (1973:194). Psammophylax acutus acutus: Kelly et al. (2008:1052). Psammophylax acutus jappi: Kelly et al. (2008:1052). Psammophis acutus: Wallach et al. (2014:575). Psammophylax acutus: Conradie et al. (2016:22). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Gabon to the former province of Katanga (Democratic Republic of Congo) to Angola and east to northern Malawi, western Tanzania and ll-E 20°E ,24'’ E Map 316 Distribution of Psammophis zambiensis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 355 Burundi. Records from northern Namibia (M. Griffin in Wallach et al. 2014) have not been verified or vouchered. Ocurrences in Angola (Map 317): The species occurs from southwestern regions of Huila Province to central and northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a:lll; Broadley 1971d:4); “Cassange” [-9.58333, 17.86667] (Bocage 1895a:lll; Broadley 1971d;3). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a; 111; Broadley 1971d:3); “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:lll; Broadley 1971b:3). Moxico: “Calombe, Luso” [-11.83333, 19.93333] (Managas 1973:194, 195). Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Gunther 1865a:480; 1888:327, 3*’' Distribution of Psammophylax acums in 1895:89; Bocage 1895a: 111; Boulenger 1896:148; Loveridge 1933:252; Monard 1937b; 128; Broadley 1971d:3; Chirio and Ineich 1991:220; Chippaux 2006:170; Wallach et al. 2014:575). Huambo: “Bela-Vista” [-12.36667, 16.20000] (Hellmich 1957b:71; Broadley 1971d:3). Benguela: “Bigondo” [-12.06667, 17.41667] (Monard 1937b:128). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:lll; Monard 1937b:128; Broadley 1971d:3); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:lll; Monard 1937b:128; Broadley 1971d:3). Cuando Cubango: “Longa River (23)” [-14.58972, 18.17083] (Conradie et al. 2016:8-9, 22); “Longa River (51)” [-14.58970, 18.1711] (Conradie et al. 2016:8, 12, 22). Undetermined locality: “Between Benguella and Bihe” (Boulenger 1905:113; Broadley 1971d:3). Taxonomic and distributional notes: Broadley (1971b) recognized three subspecies: Rhamphiophis a. acutus, R. a.jappi (Broadley, 1971) andR «. togoensis (Matschie, 1893) and pro¬ visionally synonymized Rhamphiophis acutus wittei (Laurent, 1956) with the nominate form. Recently Kelly et al. (2008) showed that Rhamphiophis is diphyletic and transfered R. acutus and its subspecies to the genus Psammophylax. The specimen from “Dundo” (MD 5193-A) identified by Laurent (1964a) as R. a. acutus and later as R. a.jappi by Broadley (197 Id) is here interpreted as the typical form. We recently examined the specimen identified by Mananas (1973) as Psam¬ mophylax tritaeniatus tritaeniatus (Gunther, 1868) from “Calombe, Luso” deposited in the Institu¬ te de Investigagao Cientifica Tropical, Lisboa and it is likewise referable to Psammophylax a. acu¬ tus. Wallach et al. (2014) erroneously allocated R. acutus to Psammophis. Psammophylax rhombeatus ocellatus (Bocage, 1873) Spotted Skaapsteker (Endemic) Psammophylax ocellatus Bocage 1873a:22L Holotype: MBL specimen number unknown (collector J.A. d’Anchieta), specimen lost fide Broadley (1977c). Type locality: “I’interieur de Mossamedes (Gambos)” [= Gambos], Huila Province, Angola. Psammophis rhombeatusl: Bocage (1867b:224). Psammophylax rhombeatus: Bocage (1895a: 108), Boulenger (1896:138, 1915:211), Bates et al. (2014), Wallach et al. (2014:582). Trimerorhinus rhombeatus: Monard (1937b: 126, 128). Psammophylax rhombeatus oeellatus: Broadley (1977c:20, 1990:121), Branch (1998:88). 356 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Global conservation status (lUCN): Not Evaluated. Global distribution: The subspecies is endemic to southern Angola. Ocurrences in Angola (Map 318): The subspecies occurs in southwestern Angola near Namibia border. Benguela: “Dombe” [-12.95000, 13.10000] (Bocage 1867b:224). Huila: “Gambos” [-15.76667, 14.10000] (Bocage 1873a:221, 1895a: 108; Monard 1937b: 128; Broadley 1977d:20-21). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a:108; Boulenger 1896:139, Monard 1937b:128; Broadley 1977c:21). Undeter¬ mined Locality: “No specific locality (Coll. Anchieta)” (Broadley 1977d:21). Taxonomic and distributional notes: Broadley (1977c) and subsequent authors (Broadley 1990; Branch 1998; Bates et al. 2014) have considered P. r ocellatus (Linnaeus, 1758) as a valid subspecies. The type of Psam- mophylax ocellatus Bocage, 1873 was not located by Broadley during his visit to the Museu Bocage in 1968 (Broadley 1977b). Wallach et al. (2014) noted that Broadley (1977d) had restrict¬ ed the type locality oi P r. rhombeatus to “SW Western Cape Prov., South Africa” but Broadley took no such formal action, merely noting that the type specimens “agree well” with with specimens of P. rhombeatus rhombeatus from the southwestern Cape Province, from wherer the types doubtless originated. Map 318. Distribution of Psammophylax rhombeatus ocellatus in Angola. Psammophylax tritaeniatus (Gunther, 1868) Striped Skaapsteker Rhagerrhis tritaeniata: Gunther 1868:423, pi. 19, fig. H. Holotype: BMNH 1946.1.2.78 (collector Warwick). Type locality: “probably from South-eastern Africa”, (“presumably Rhodesia [Zimbabwe] or the Trans¬ vaal” [= Limpopo or Mpumalanga], South Africa fide Broadley 1977c:32). Rhagerrhis tritaeniata-. Bocage (1873b:220, 1879b:95, 1887c:210, 1895a: 110, 1896a: 112, 1897b:211), Ferreira (1897b:244). Cerastes tritaeniatus tritaeniatus'. Mertens (1937a: 14, 1938a:441), Bogert (1940:77). Trimerorhinus rhombeatus tritaeniatus'. Monard (1937b: 130). Psammophylax tritaeniatus tritaeniatus'. Hellmich (1857b:71), Loveridge (1957:276), Laurent (1964a:110). Psammophylax tritaeniatus'. Broadley (1977c:32, 1990:124), Branch (1998:88), Wallach et al. (2014:583). Global conservation status (lUCN): Least Concern. Global distribution: The species endemic to sub-Saharan Africa, from the former province of Katanga, Democratic Republic of Congo, Angola and Zambia to southern Tanzania and south through Zimbabwe, western Mozambique, Botswana, northern Namibia and northeastern South Africa. Ocurrences in Angola (Map 319): The species distribution probably comprises the most southern regions of Angola, with exception of the Namibe Province, as well as Mexico. Moxico: “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:110; Broadley 1977c:36). Benguela: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:110; Broadley 1977c:36); “Quissange” [-12.43333, 14.05000] (Bocage 1887c:210, 1895a:110; Broadley 1977c:36); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:110; Broadley 1977c:36); “Ebanga” [-12.73333, 14.73333] (Monard MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 357 1937b: 130; Broadley I977c:36); “Cubal (Alto Cubal)” [-13.03333, 14.25000] (Mertens 1937a:14, 1938a:441; Hellmich 1957b:71; Broadley 1977c:36); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:112, 1897b:211). Hulla: “Caconda” [-13.73333, 15.06667] (Bocage 1895a: 110; Ferreira 1897b:244; Broa¬ dley 1977c:36); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:130; Broadley 1977c:36); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:77; Broadley 1977c:36); “Huilla” [-15.05000, 13.55000] (Bocage 1873b:220, 1895a: 110); “Jau, envi¬ rons de Sa da Bandeira” [-15.20000, 13.51667] (Laurent 1964a: 110; Broadley 1977c:36); “Gambos” [-15.76667, 14.10000] (Bocage 1873b:220, 1895a:110). Cunene: “Forte Rogadas” [-16.71667, 15.01667] (Laurent 1964a: 110; Broadley 1977a:36); “Humbe” [-16.68333, 14.90000] (Bocage 1895a:110; Broadley 1977c:36); “Dongwenna” [-17.01667, 14.71667] (Broadley 1977c:36); “Mupanda” [-17.13333, 15.76667] (Monard 1937b:130; Broadley 1977c:36); “36 km NW of Humbe” (Broadley 1977b:36). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b:130). Undetermined Locality: “without precise location” (Bocage 1879b:95). Taxonomic and distributional notes: Managas’s (1973) reference to Psammophylax tritae- niatus tritaeniatus (Gunther, 1868) from “Calombe, Luso” in the Institute de Investigagao Cienti- fica Tropical, Lisboa is referable to Psammophylax a. acutus. Genus Pseudaspis Fitzinger, 1826 Pseudaspis cana (Linnaeus, 1758) Mole Snake Coluber canus Linnaeus 1758:221. Holotype: NHR Lin-22, formely MAFR (ex Mus. Drottingholm). Type loeality: “Indiis” [= India] (Linnaeus 1758:221), in error, eorreeted to “Cape of Good fide Wallaeh et al. (2014:585), South Afriea. Ophirhina Anehietae: Boeage 1882a:300. Syntypes: MBL speeimen numbers unknown (eolleetor J.A. d’Anehieta), destroyed by fire 18 Mareh 1978. Type loeality: “Caeonda, dans finterieur de Benguella” [= Caeonda] Huila Provinee, Angola. Pseudaspis cana: Boeage (1895a: 100), Braneh (1998:80), Boulenger (1915:204), Monard (1937b: 118), Bogert (1940:42), Loveridge (1957:266), FitzSimons (1962:162), Frade (1963:253), Broadley (1990:108), Broadley et al. (2003:147), Broadley and Cotterill (2004:49), Kelly et al. (2009:46), Bates et al. (2014:392), Wallaeh et al. (2014:585). Pseudaspis cana anehietae: Laurent (1956:141). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from throughout southern Africa, extending north to Angola, Zambia, the former Katanga Province of the Democratic Republic of Congo, Burundi, Rwanda and the Kenya highlands. Ocurrences in Angola (Map 320): The species occurs in the central-southern areas. Kwanza Sul and Huila Province. It is probable that the distribution of this species reachs the southeastern I2"E 16.°E 2«°E Map 319. Distribution of Psammophylax tritaeniatus in Angola. 358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II regions of the country. Kwanza Sul: “Mombo- lo” [-12.16667, 14.83333] (Bogert 1940:42; Laurent 1956:141). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a;100; Monard 1937b:113; Laurent 1956:141). Hulla: “Rio Cuce” [-13.51667, 15.20000] (Bocage 1895a:100; Monard 1937b:113; Laurent 1956:141); “Caconda” [-13.73333, 15.06667] (Bocage 1882a:300, 1895a: 100; Monard 1937b: 113; Laurent 1956:141; Loveridge 1957:266); “Sangeve” [-13.88333, 15.83333] (Monard 1937b:113, 118; Laurent 1956:141); “Kuvangu (Vila-da-Ponte)” [-14.46667, 16.30000] (Monard 1937b:113, 118; Laurent 1956:141). Map 320. Distribution of Pseudaspis cana in Angola. Taxonomic and distributional notes; Kelly et al. (2009) revived the family Pseudas- pididae Dowling and Duellman, 1978 for two monotypic genera, Pseudaspis {P. cana) and Pythonodipsas (P carinata Gunther, 1868). This clade has, however, subsequently been treated as a subfamily, Pseudaspidinae, by Pyron et al. (2010, 2013). Laurent (1956) recognized P. c. anchietae (Bocage, 1882) as a valid subspecies, but Broadley (1990) considered evidence for the recognition of a taxonomically distinct northern form to be unconvincing. Il'E l&'E 20°E 24”E Genus Pythonodipsas Gunther, 1868 Pythonodipsas carinata Gunther, 1868 Pythonodipsas carinata Gunther 1868:427. Holotype: man). Type loeality: “Zambezi” (Gunther 1868:427), in error, corrected to “Damaraland, South West Africa” by Broadley (1971c:679). Pythonodipsas carinata: FitzSimons (1962:181), Broadley (1971c:679, 1990:110), Branch et al. (1997:89), Branch (1998:82), Kelly et al. (2009:46), Bates et al. (2014:388), Wallach et al. (2014:612). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is restricted to gravel plains and arid savannah from the Bethanie District in southern Namib¬ ia, northwards into the coastal desert of south¬ ern Angola. Ocurrences in Angola (Map 321): The species is occurs at the edges of the Angolan Namib Desert in the southern portion of Namibe Province. Western Keeled Snake BMNH 1946.1.4.70, formerly SAM (collector J. Chap- ll-'E 16°E 20°E irE Map 321. Distribution of Pythonodipsas carinata in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 359 Namibe: “Quarter Degree Square I5I2Aa” [=Saco; -15.125, 12.125] (Branch et al. 1997:94); “Quarter Degree Square I6I2Cd” [-16.875, 12.375] (Branch et al. 1997:94); “Quarter Degree Square 1613Ca” [-16.625, 12.125] (Branch et al. 1997:94). Taxonomic and distributional notes: Kelly et al. (2009) revived the family Pseudaspididae Dowling and Duellman, 1978 for Pseudaspis and Pythonodipsas but this clade has subsequently been treated as a subfamily, Pseudaspidinae, by Pyron et al. (2010, 2013). Despite the few litera¬ ture refereces for this species in Angola, it is expected to be widespread in the Namib Desert of the southwest (e.g., FitzSimons 1962; Broadley 1990; Branch et al. 1997; Branch 1998; Branch 1998; Bates et al. 2014; Wallach et al. 2014). Branch et al. (1997) provided the first literature records from southwestern Angola, all as quarter degree points plotted on a map. Genus Xenocalamus Gunther, 1868 Xenocalamus bicolor machadoi Laurent, 1954 Machado’s Quill-Snouted Snake Xenocalamus bicolor machadoi Laurent 1954a:45, figs. 9-11. Holotype: MD 2082 (collector A. Barros Machado). Type locality: “Dundo,” Kwanza Norte Province, northeastern Angola. Xenocalamus Mechovii: Boulenger (1905:113). Xenocalamus bicolor pernasutus: de Witte and Laurent (1947:46). Xenocalamus bicolor machadoi: Broadley (1971a:680), Branch (1998:68). Xenocalamus bicolor. Wallach et al. (2014:780). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and the western Katanga region. Democratic Republic of Congo. Ocurrences in Angola (Map 322): The species distribution may comprise the entire country with exception of the northwestern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:45; Broad¬ ley 197 la: 681). Benguela: “Benguella” [-12.58333, 13.41667] (Laurent 1954a:45). Undetermined locality: “Between Benguella and Bihe” (Boulenger 1905:113; de Witte and Laurent 1947:46; Broadley 1971a:681). Taxonomic and distributional notes: Laurent (1954a) dQScnhQd Xenocalamus bicol¬ or machadoi as a new subspecies based on a female holotype specimen from “Dundo,” Lunda Norte Province, from the Barros Machado collection in the Museu do Dundo, as well as a paratype from “Sandoa, dist. Lualaba, Congo Beige” and a specimen from Boulenger (1905) identified as Xenocalamus mechovii from “Between Benguella and Bihe.” It is expected that the nominate form, Xenocalamus bicolor bicolor, also occurs in Angola, presumably in the southeastern regions of the country, although it has not yet been documented. Figueroa et al. (2016) synonymized Xenocalamus with Amblyo- dipsas, however, as their data were limited, we retain the highly distinctive Xenocalamus as valid pending further data. Map 322. Distribution oi Xenocalamus bicolor machadoi in Angola. 360 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Xenocalamus mechowii mechowii Peters, 1881 Elongate Quill-Snouted Snake Xenocalamus Mechowii Peters 1881:147. Holotype: ZMB 10044 (eollector F.W. von Mechow). Type loeali- ty: “Malanje,” Malanje Provinee, Angola. Xenocalamus Mechovii: Boulenger (1915:214). Xenocalamus mechowi: Monard (19376:127), Branch (1998:69), Broadley (2003:88), Chippaux (2006:213), Wallach et al. (2014:781). Xenocalamus mechowi mechowi'. de Witte and Laurent (1947:49), Broadley (1971a:681) Laurent (1954a:45). Xenocalamus mechowii inornatus: Branch and McCartney (1992:2). Global conservation status (lUCN): Not Evaluated. Global distribution: The species as a whole Republic of Congo. Through eastern Angola, to northeastern Namibia, northern Botswana and northwestern Zimbabwe. Ocurrences in Angola (Map 323): The species probably occurs throughout the entire country with the exception of the western regions along the coast. Lunda Norte: “Sombo (Kassekue river, right affluent of Chiumbe)” [-8.68333, 20.95000] (Laurent 1954a:45). Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1881:147; de Witte and Laurent 1947:49; Chippaux 2006:213). Taxonomic and distributional notes: Within Angola Broadley (1971a), considered the typi¬ cal form to be restricted to northern Angola. Figueroa et al. (2016) synonymizQd Xenocala¬ mus with Amblyodipsas, however, as their data were limited, we retain the highly distinctive Xenocalamus as valid pending further data. is known from southern Congo and Democratic I2”E 20“E 24“E Map 323. Distribution of Xenocalamus mechowii mechowii in Angola. Xenocalamus mechowii inornatus de Witte and Laurent, 1947 Inornate Elongate Quill- Snouted Snake Xenocalamus mechowii inornatus de Witte and Laurent 1947:51. Holotype: SAM 19719 (collector unknown). Type locality: “Ovamboland (Sud-Ouest africain)” [= Ovamboland, Namibia]. Xenocalamus mechowii inornatus'. Branch and McCartney (1992:2). Global conservation status (lUCN): Not Evaluated. Global distribution: This subspecies is known from northern and northeastern Namibia, and adjacent regions of Zambia, Botswana, Zimbabwe and Angola. Ocurrences in Angola (Map 324): This subspecies is confirmed form a single record in the southeast of the country, but it may occur more broadly. Cuando Cubango: “vinicity of Cuito Cua- navale- approximatley 45 km S of Lupire” [-15.08333, 19.58333] (Branch and McCartney 1992:2). Taxonomic and distributional notes: Branch and McCartney (1993) recorded X. mechowii inornatus from Cuando Cubango Province and considered theirs to be first record of the southern race in Angola, although there are several records from immediately adjacent to Cuando Cubango in the Caprivi Strip and western Zambia. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 361 ll«E 2(»”E 24'>E Map 324. Distribution of Xenocalamus mechowii inor- natus in Angola. Family Elapidae Boie, 1827 Genus Aspidelaps Fitzinger, 1843 Aspidelaps lubricus cowlesi Bogert, 1940 Angolan Coral Snake Matrix lubrica Laurenti 1768:80. Lectotype: specimen described and illustrated by Seba (1735:44, pi. 43, fig. 3) fide Wallach et al. (2014:55, see Notes below), lost fiide Broadley in Golay et al. (1993:113) although status considered unknown by Broadley and Baldwin (2006). Type locality: restricted to “Africa” based on the lectotype designation (Laurenti 1768:80) and more specifically to “ex Promontorio Bonae Spei/Cap de Bonne-Esperance” [= Cape of Good Hope, Western Cape Province], South Africa (Seba 1735:44). Aspidelaps lubrieus cowlesi Bogert 1940:94, fig. 17A-D. Holotype: AMNH 32801 (collectors A.S. Vemay, H. Lang and R. Boulton). Type locality: “Munhino (101 km east of Mossamedes, via railroad)” (Bogert 1940:95) [= Muninho, 101 km east of Namibe], Namibe Province, Angola. Aspidelaps lubricus cowlesi: FitzSimons (1962:278), Managas (1981:21), Broadley (1990:281), Spawls and Branch (1995:54), Branch (1998:104), Broadley and Baldwin (2006:167), Dobiey and Vogel (2007:32), Wallach et al. (2014:55). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Angola and Namibia as far south as Liideritz. Ocurrences in Angola (Map 325): The species occurs in southwestern Angola, near the bor¬ der between Cunene Province and Namibia. Namibe: “Munhino (101 km east of Mossamedes, via railroad)” [-14.91667, 13.00000] (Bogert 1940:94; Managas 1981:21; Broadley 1990:281; Broadley and Baldwin 2006: 167). Taxonomic and distributional notes: Laurenti (1768) cited only a specimen figured by Seba (1735) for both Matrix lubrica as a whole and for his “Var. p.” although another form from Suri¬ name was noted, so there must have originally been syntypes. The history of Seba’s collections have been discussed in detail (Engel 1937, 1961; Boeseman 1970; Juriev 1981, Adler 1989; Bauer 2002; Bauer and Gunther 2013). Seba’s second collection (the first had been sold to Peter the Great of Russia in 1716; Driessen-van het Reve 2006) was sold after his death at auction (Anonymous 362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1752). Seba specimens are known or believed to be present in collections in St. Petersburg, London, Leiden (including specimens until recently in Amsterdam), Paris, Copenhagen, Stockholm, Bremen and Berlin (Boeseman 1970; Juriev 1981; Thireau et al. 1998; Bauer and Gunther 2013) but few can be traced to particular plate figures, and the lectotype of Matrix lubrica is not among these. Wallach et al. (2014) cited no source for the designation of a lectotype, and indeed, no lectotype seems to have been formally designated previously, although most authors have only cited the African syntype (or Seba’s figure) as type material, apparently unaware that the original description was based on more than one speci¬ men. We interpret Wallach et al.’s use of the term lectotype as an intended lectotype desig¬ nation, but as it lacks an express statement of deliberate designation (Art. 74.7.3) this designation is not valid, although it is appropriate, as the other specimen(s) implied are apparently referable to some other taxon, presumably from South America. Broadley and Baldwin (2006) referred Aspidelaps lubricus infuscatus Mertens, 1954, which had generally been recognized as the taxon occurring in northern Namibia, to the synonomy of^. /. cowlesi. Map 325. Distribution of Aspidelaps lubricus cowlesi in Angola. Genus Dendroaspis Schlegel, 1848 Dendroaspis jamesoni (Traill, 1843) Jameson’s Mamba Flaps jamesoni Traill 1843:54. Holotype: NMSZ 1869.3.147, formely TST and RSM (collector T.S. Traill family). Type locality: “Demerara, Equinoxial America” (Traill 1843:53) [= Guyana, South America], in error, restricted to “West Africa” by Mertens (1938b:49). Dendraspis Welwitschii Gunther 1865b:97, pi. 2, fig. A. Holotype: BMNH:64.7. 13.39 (collector F. Wel- witsch). Type locality: “Golungo Alto”, Kwanza Norte Province, Angola. Dendraspis negleetus Bocage 1888:141, fig. 4. Syntypes: MBL specimen numbers unknown (collector not stated), destroyed by fire 18 March 1978. Type locality: “L’Afrique occidental de la Senegambie a Ango¬ la (au nord du Quanza)”, [= West Africa from Senegal and Gambia to Angola (north of Quanza)]. Dendraspis Welwitsehii: Bocage (1866a:51, 1888:143). Dendraspis negleetus: Bocage (1895a: 138), Ferreira (1900a:53). Dendroaspis jamesonii: Boulenger (1896:436, 1915:220), Parker (1936:126), Monard (1937b:137), Frade (1963:252), Spawls and Branch (1995:47), Dendroaspis jamesoni jamesoni: Laurent (1950a: 10, 1954a:61, 1964a: 121), Hellmich (1957b:73), Thys van den Audenaerde (1966:36), Chirio and LeBreton (2007:570). Dendroaspis jamesonii jamesonii {Flaps jamesonii): Managas (1981:30). Dendroaspis jamesoni: Dobiey and Vogel (2007:42), Wallach et al. (2014:219), Vaz Pinto and Branch (2015:45). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Ghana eastwards to southern Sudan, South Sudan and southwestern Kenya (Kakamega) and south to northern and central Angola, the south¬ ern Democratic Republic of Congo, and possibly northern Zambia. Wallach et al. (2014) consid- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 363 ered that records from Benin, Cote d’Ivoire and northern Zambia required confirmation. Ocurrences in Angola (Map 326): The species is known from north and central Ango¬ la. Cabinda: “Landana” [-5.21667, 12.15000] (Bocage 1895a:138); “Cabinda” [-5.55000, 12.18333] (Bocage 1895a: 138; Mananas 1981:30). Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:138; Managas 1981:30). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 10, 1954a:61, 1964a: 121; Thys van den Audenaer- de 1966:36); “Dundo, barrage de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaer- de 1966:36). Luanda: “Loanda” [-8.83333, 13.26667] (Ferreira 1900a:53). Bengo: “de Cacolo ao rio Bengo” (Ferreira 1900:53). Malanje: “Pungo-Andongo” [-9.66667, Distribution oi Dendroaspis jamesoni in 15.58333] (Bocage 1895a:138). Kwanza Norte: “Piri-Dembos” [-8.53333, 14.43333] (Hellmich 1957b:73; Mananas 1981:30); “Golungo Alto (Golungo-alto, dans llnterieur d’Angola, au nord du Quanza)” [-9.13333, 14.76667] (Gunther 1865b:97; Bocage 1888:143; 1895a:138; Boulenger 1896:436; Monard 1937b:137, Managas 1981:30); “Ambaca” [-9.26667, 15.18333] (Ferreira 1900a:53); “N’dalla Tando” [-9.30000, 14.91667] (Ferreria 1903:13; Mananas 1981:30); “Dondo” [-9.68333, 14.43333] (Managas 1981:30). Kwanza Sul: “on a track between Kwanza bridge “Filomena da Camara” and Farm Cabuta” [-9.78528, 14.81389] {pers. comm. Kelse Alexandre in Vaz Pinto and Branch 2015:45); “main road from Luanda to Huambo at Munenga, approximately 12 km south of the turning to Calulo” [-10.12083, 14.66500] (Vaz Pinto and Branch 2015:45); “Quirimbo” [-10.68333, 14.26667] (Parker 1936:126; Mana9as 1981:30); “Congulu” [-10.86667, 14.28333] (Parker 1936:126; Managas 1981:30). Huambo: “Bela Vista” [-12.36667, 16.20000] (Hellmich 1957b:73; Mananas 1981:30). Undetermined Locality: “Angola (au nord du Quanza)” (Bocage 1888:142). Taxonomic and distributional notes: Hughes and Barry (1969) listed two syntypes (RSM 1869.3.147 and BMNH 1946.1.20.43) of D. jamesoni, but the original description specified a sin¬ gle type (see Wallach et al. 2014). Bocage (1895a) recognized two species of mambas in Angola, Dendroaspis angusticeps (Smith, 1849) md Dendroaspis neglectus (Bocage, 1888), the latter cur¬ rently considered a junior synonym of D. jamesoni. The species Dendroaspis angusticeps has been restricted to East and Southern Africa, and Bocage’s records assigned to this species in fact repre¬ sent the Black Mamba, Dendroaspis polylepis Gunther, 1864. In Angola, Spawls and Branch (1995) and Dobiey and Vogel (2007) considered D. jamesoni present only to the north of Luanda. Il'E 20°E 24^E Dendroaspis polylepis (Gunther, 1864) Black Mamba Dendraspis polylepis Gunther 1864:310. Holotype: BMNH 1946.1.20.53 (formerly 64.6.28.15) (eolleetor J. Kirk). Type loeality: “regions bordering the Zambezi, ineluding those of the Nyassa Lake” [= Zambesi River], Mozambique fide Loveridge (1953a:290). Dendraspis angustieepsl: Boeage (1866a: 52). Dendraspis angustieeps: Peters (1877a:617, 1888:149), Boeage (1888:143, 1895a: 140), Boulenger 364 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (1915:220), Schmidt (1933:15), Monard (1937b:137), Bogert (1940:92), Frade (1963:253). Dendroaspis polylepis polylepis: Loveridge (1957:294), FitzSimons (1962:310), Baynham (2010:25), Conradie et al. (2016:22). Dendroaspis polylepis polylepis (Dendroaspis polylepis): Mananas (1981:30). Dendroaspis polylepis: Hakansson and Madsen (1983:186), Broadley (1990:297), Spawls and Branch (1995:49), Broadley and Cotterill (2004:48), Chippaux (2006:235), Chirio and LeBreton (2007:572), Dobiey and Vogel (2007:44), Bates et al. (2014:398), Wallach et al. (2014:220), Branch and Conradie (2015:200), Ceriaco et al. (20 16b: 87). Global conservation status (lUCN): Least Concern. Global distribution: The species has an enormous range throughout the savannas of sub- Saharan Africa, occuring from Senegal eastwards to Sudan, Eritrea, Ethiopia and Somalia, then south to the Eastern Cape in South Africa, and west to Namibia and Angola, but absent from the equatorial forests of west and central Africa and from desert areas. Range in West Africa highly fragmented. Ocurrences in Angola (Map 327): The species mainly occurs in the west and central areas of Angola, however, it is also widely dis¬ tributed in the eastern regions of the country. There are no published records from most coastal areas or from the Namib Desert. Zaire: “Luanda Pil, approximately 6.5 km west of Soyo airport” [-6.155556, 12.26611] (Bayn¬ ham 2010:25). Lunda Norte: “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200). Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1881:149); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:87). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:15). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1888:143; Mana9as 1981:30); “Quindumbo” [-12.46667, 14.93333] (Bocage 1888:143, 1895a: 140; Monard 1937b: 137; Managas 1981:30); “Cahata” [-12.35000, 14.81667] (Bocage 1895a:140; Monard 1937b:137; Managas 1981:30); “Hanha” [-12.25000, 13.75000] (Bogert 1940:92). HmTa: “Molundo” [-15.63333, 15.20000] (Schmidt 1933:15). Cuando Cubango: “Cuando basin (37)” (observation record) [-17.67833, 22.61475] (Conradie et al. 2016:9, 10, 22). Taxonomic and distributional notes: For some time Dendroaspis polylepis (Gunther, 1864) was wrongly identified as Dendroaspis angusticeps (Smith, 1849) in Angola (Bocage 1866a, 1888, 1895a; Peters 1877a, 1888; Boulenger 1915; Schmidt 1933; Bogert 1940; Frade 1963). This is clearly a misidentification, as D. angusticeps has a restricted distribution in Eastern southern Africa (Managas 1981; Wallach et al. 2014). Branch and Conradie (2015) recently collected D. polylepis from Lake Carumbo, in Lunda Norte Province, the most eastern record of the species in Angola and Branch et al. (in Conradie et al. 2016) suggested that the species is likely more widely distrib¬ uted in southern Angola than previously believed. Map 327. Distribution of Dendroaspis polylepis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 365 Genus Elapsoidea Bocage, 1866 Elapsoidea guentherii Bocage, 1866 Gunther’s Garter Snake Elapsoidea Guntherii Bocage 1866a:50, 1866b:70, pi. 1, figs. 3, 3a-3b. Lectotype: MBL T-130, fonnerly MBL 1935 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978, designated by Parker (1949:97). Type locality: restricted to “Cabinda” (Bocage 1866a:50, 1866b:70), Angola through the lectotype desig¬ nation. Elapsoidea Guntherii: Bocage (1873b:224). Elapsoidea Giintheri: Bocage (1895a: 129, 1897b:202). Elapeehis guentheri: Boulenger (1896:359). Elapsoidea giintheri: Loveridge (1936a:41), Bogert (1940:86). Elapsoidea sundevallii guntherii: Loveridge (1944c:222, 1957:290). Elapsoidea sundevallii giintheri [part]: Parker (1949:93). Elapsoidea giintheri giintheri: Laurent (1964a: 117). Elapsoidea sundevalli semiannulata: Haacke and Finkeldey (1967:3). Elapsoidea guentheri: Broadley (1971e:600), Spawls and Branch (1995:60), Branch (1998:105) Elapsoidea guentheri {Elapsoidea Guntherii): Managas (1891:22). Elapsoidea guentherii: Chippaux (2006:216), Dobiey and Vogel (2007:49), Wallach et al. (2014:262). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies ranges from the lower Congo, south to Angola and east to the southern Democratic Republic of Congo, western Zambia and Zimbabwe. Ocurrences in Angola (Map 328): The species occurs from the northwest to the central plateaus of Angola. Cabinda: “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:59, 1866b;70, 1873b:224, 1895a: 129, 1897b:202; Loveridge 1936a:41, 1944c:222, 1957:290; Parker 1949:95; Broadley 1971e;601; Managas 1981:22; Chippaux 2006:216; Wallach et al. 2014:262). Kwanza Norte: “N’dala Tando” [-9.30000, 14.91667] (Broadley 1971e:601; Managas 1981:22). Moxico: “Rives du lac Calundo (Lago Calundo)” [-11.71667, 20.80000] (Laurent 1964a: 11 7; Broadley 1971e:601; Mana9as 1981:22). Benguela: “with no precise identification but probably from Hanha” [-12.25000, 13.75000] (Bogert 1940:86; Broadley 1971e:601). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a: 129; Loveridge 1936a:41; Broadley 1971e:601; Mananas 1981:22). Taxonomic and distributional notes: The species was described as new by Bocage in two different papers (1866a,b), both of which appeared in the November 1866 issue of the same jour¬ nal. However, the generic name, although used in both papers, was actually defined and introduced as new only in the second paper. Haacke and Finkeldey (1967) regarded E. semiannulata Bocage, 1882 as the only member of the genus occurring in Angola and southern Congo. However, both E. semiannulata and E. guentherii are both present (e.g., Broadley 1971e; Mana9as 1981; Spawls and Branch 1995; Chippaux 2006; Dobiey and Vogel 2007; Wallach et al. 2014). Map 328. Distribution of Elapsoidea guentherii in Angola. 366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Elapsoidea semiannulata semiannulata Bocage, 1882 Angolan Garter Snake Elapsoidea semi-annulata Bocage 1882a:303. Holotype: MBL 1938 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978. Type locality: “Caconda” Huila Province, Angola. Elapsoidea decosteri huilensis Laurent (1964a: 11 8). Type: Holotype, MD 5552 (collector W. Kisker, coll. Prof. H. Baumann leg.). Type locality: “Fazenda Bumbo, Humpata (Huila),” Huila Povince, Angola. Elapsoidea Guntherii: Bocage (1873b:224). Elapsoidea Giintheri: Bocage (1895a: 129; 1897b:202). Elapsoidea giintheri var. semiannulata'. Ferreira (1900a: 52). Elapsoidea giintheri'. Schmidt (1933:14), Mertens (1938a:442). Elapeehis guentheri: Monard (1937b: 137). Elapsoidea sundevallii semiannulata'. Loveridge (1936:41, 1944c:220). Elapsoidea sundevallii giintheri [part]: Parker (1949:93). Elapsoidea semiannulata'. Broadley (1998b: 15), Chippaux (2006:217), Dobiey and Vogel (2007:53), Wallach et al. (2014:236), Elapsoidea ? sundevallii semiannulata'. Hellmich (1957a:73). Elapsoidea semiannulata semiannulata'. Broadley (1971e:610, 1998b: 15), Spawls and Branch (1995:64). Elapsoidea semiannulata semiannulata {Elapsoidea semi-annulata)'. Mananas (1981:23). Global conservation status (lUCN): Not Evaluated. Global distribution: The nominotypical subspecies is known from Angola, Zambia west of the Zambezi and northern Namibia. Ocurrences in Angola (Map 329): Wide¬ spread, with most records in the west of the country, but replaced in Cabinda and possibly elsewhere in the far north by E. s. moebiusi. Luanda: “Loanda” [-8.83333, 13.26667] (Fer¬ reira 1900a;52). Bengo: “de Cacolo ao rio Bengo” (Ferreira 1900a:52). Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Ferreira 1900a:52); “N’dalla Tando” [-9.30000, 14.91667] (Broadley 1971e;611; Managas 1981:23). Moxico: “Cazengo” [-9.33333, 14.76667] (Ferreira 1900a:52; Loveridge 1944c:221); “Vila Luso” [-11.78333, 19.91667] (Managas 1981:23). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:14; Loveridge 1944c: 221; Broadley 1971e:611; Managas 1981:23). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a:129, 1897b:202; Broadley 1971e:611; Managas 1981:23); “Dondi” [-12.53333, 16.25000] (Loveridge 1944c:221; Broadley 1971b:611; Managas 1981:23). Benguela: “Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:73; Broadley 1971e:611; Managas 1981:23); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:442; Loveridge 1944c:221; Broadley 1971e:611; Mana9as 1981:23). Huila: “Kampulu (environs de Kasinga)” [-15.21667, 16.11667] (Monard 1937b:137; Loveridge 1944c:221; Broadley 1971e:611; Managas 1981:23); “Caconda” [-13.73333, 15.06667] (Bocage 1882b:303;, 1895a:129; Loveridge 1936a:41; 1944c:221; Parker 1949:95; Broadley 1971e:611, 1998b:16; Managas 1981:23; Chip¬ paux 2006:217; Chirio and LeBreton 2007:576; Wallach et al. 2014:263); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:137; Loveridge 1944c:221; Broadley 1971e:611; Managas Map 329. Distribution of Elapsoidea semiannulata semi¬ annulata in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 367 1981:23); “Huilla” [-15.05000, 13.55000] (Bocage I873b:224, I895a:I29, I897b:202; Broadley I97Ie:6II; Managas 1981:23); “Fazenda Bumbo, Humpata” [-15.20000, 13.00000] (Laurent I964a:II8; Broadley 1971e:661; Managas 1981:23); “Gambos” [-15.76667, 14.10000] (Bocage 1873a:224, 1895a:12, 1897a:202; Loveridge 1944c:221; Broadley 1971e:611, 1998b:16; Managas 1981:23). Namibe: “Maconjo” [-15.01667, 13.20000] (Bocage 1895a:129, 1897c:202; Loveridge 1944c:221; Broadley 1971e:611, 1998b:16; Mananas 1981:23). Taxonomic and distributional notes: The species was described by Bocage (1882a) as Elap- soidea semi-annulata based on two specimens from “Caconda” collected by J. Anchieta. Bocage (1895a) later changed the orthography to Elapsoidea semiannulata. Laurent (1964a) described a new subspecies from Angola, Elapsoidea decosteri huilensis, which currently is recognized as synonym of this taxon. Elapsoidea semiannulata moebiusi (Werner, 1897) Moebius’ Garter Snake Elapechis moebiusi Werner 1897:400. Holotype: ZMB 13802 (eolleetor J.N.F.J. Graf von Zeeh). Type loeal- ity: “Kete” [Kete, or Kete Kraehi, was in the German eolony of Togo at the time of the deseription, but after 1914 beeame part of the Gold Coast, and following independenee in 1957, Ghana]. Elapsoidea semiannulata moebiusi'. Broadley (1971e:609), Spawls and Braneh (1995:64), Broadley (1998b: 16). Elapsoidea semiannulata moebiusi (Elapeehis moebiusi)'. Mananas (1981:23). Elapsoidea semiannulata'. Trape and Mane (2006:190), Dobiey and Vogel (2007:53), Wallaeh et al. (2014:236). Elapsoidea semiannulata moebiusi'. Chirio and LeBreton (2007:576). Global conservation status (lUCN): Not Evaluated. Global distribution: The subspecies is known from from Senegal and the extreme south of Mauritania, west to southern Chad, and the Central African Republic, and from disjunct popula¬ tions south to western Congo, western and southwestern Democratic Republic of Congo, and far northern Angola. Ocurrences in Angola (Map 330): Elap¬ soidea. s. moebiusi is known from northern regions including the Cabinda enclave. Cabin¬ da: “Cabinda” [-5.55000, 12.18333] (Broadley 1998b: 16, 18 [Fig.l]; Dobiey and Vogel 2007:53). Taxonomic and distributional notes: Broadley (1971e, 1998b) Spawls and Branch (1995) and Dobiey and Vogel (2007) recong- nized two subspecies for the country, the typical form E. s. semiannulata in more south¬ ern regions and E. s. moebiusi (Werner, 1 897) in Cabinda or Cabinda and northern Zaire and Uige provinces. Chirio and LeBreton (2007) mistakenly noted “Caconda” as the type locali¬ ty of E. s. moebiusi. Map 330. Distribution moebiusi in Angola. of Elapsoidea semiannulata 368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Naja Laurenti, 1768 Naja anchietae Bocage, 1879 Anchieta’s Cobra Naja Anchietae Bocage 1879a:98, 1879c:89. Lectotype: MBL 1987 (collectors H.C. Capello and R. Ivens), designated by Broadley (1995:31), destroyed by fire 18 March 1978. Type locality: “Caconda,” Huila Province, Angola. Naja Anchietae-. Bocage (1879c:98, 1895a: 133, 1897a:202). Naja anehietae: Ferreira (1900ab:134), Monard (1937b: 138), Bogert (1940:90), Frade (1963:252), Broadley (1968c:7), Broadley and Cotterill (2004:47), Broadley and Wuster (2004:101), Dobiey and Vogel (2007:63), Ceriaco et al. (2014b:671), Ceriaco et al. (2016b:87). Naja haje anehietae'. Mertens (1937:23), Bogert (1940:90), Laurent (1964a:118), Managas (1981:26), Broadley (1966c:23), Spawls and Branch (1995:70), Trape et al. (2009:20). Naja anulifera aneheitae: Broadley (1995:31). Naja (Uraeus) anehietae'. Wallach et al. (2009:31). Uraeus anehietae: Wallach et al. (2014:762). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from western parts of southern Afriea, from Ango¬ la and northern Namibia through western Zambia and northern Botswana to northwestern Zimbabwe and Lake Bangweulu, northeastern Zambia. Ocurrences in Angola (Map 331): The speeies oeeurs in mostly in the southern Ango¬ la however, there are some records from more northern provinces. Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Laurent 1964a;118, Mananas 1981:26); “Alto Chicapa” [-10.93333, 19.15000] (Laurent 1964a;118, Managas 1981:26). Malanje: “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:671); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:87). Benguela: “Hanha” [-13.30000, 14.20000] (Ferreira 1900b: 134; Managas 1981:26). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1879a:98, 1879c:89, 1895a:133, 1897a:202; Ferreira 1900b: 134; Mananas 1981:26; Broadley 1995:31); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b:138; Map 331. Distribution of Naja anchietae in Angola. Managas 1981:26); “Vila da Ponte” [-14.46667, 16.30000] (Managas 1981:26); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:138; Managas 1981:26); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:90; Managas 1981:26); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:133, 1897a:202; Ferreira 1900b:134; Mananas 1981:26); “vu I’espece dans la region du Kului superieur” [-15.41667, 15.73333] (Monard 1937b:138). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 1937b:138; Mananas 1981:26); “Humbe” [-16.68333, 14.90000] (Bocage 1895a:133, 1897a:202; Ferreira 1900b:134; Managas 1981:26). Undetermined Locality: “Rio Cuando” (Bocage 1895a:133, 1897a:202; Mananas 1981:26). Taxonomic and distributional notes: Wallach et al. (2014) attributed the lectotypoe designa¬ tion for Naja anchietae to Broadley and Wallach (2004), however, Broadley (1995) had already refered to MBL 1987 as the lectotype a decade earlier. Mertens (1937a), Bogert (1940) and subse- MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 369 quent authors treated Naja anchietae as a subspeeies of Naja haje (Linnaeus, 1758), until Broadley (1995) ineluded it as a subspecies of Naja annulifera Peters, 1854. Recent studies suggest that these two taxa are morphologically and genetically distinct evolutionary lineages that should be considered as two separate species (Broadley and Wiister 2004). Wallach et al. (2009) assigned the non-spitting African cobras to the subgenus Uraeus Wagler, 1830 and subsequently treated Uraeus as a full genus (Wallach et al. 2014). Most literature has considered N. anchietae to be restricted in Angola to the south (e.g., Broadley 1966c, 1968c, 1995; Spawls and Branch 1995; Broadley and Wiister 2004; Broadley and Cotterill 2004; Dobiey and Vogel 2007; Trape et al. 2009:20; Wallach et al. 2014) but there are now numerous records from more northerly areas (Laurent 1964a; Managas 1981; Ceriaco et al. 2014b, 2016b). Naja annulata Buchholz and Peters, 1876 Ringed Water Cobra Naja annulata Buchholz and Peters 1876 in Peters 1876:119. Holotype: ZMB 8772 (eolleetor R. Buehholz). Type loeality: “Dorfe Mbusu (Eliva Sonangeam am Ogowe)” [= Eliva Sonange, near Ogooue-Maritime], Gabon. Boulengerina annulata: Spawls and Braneh (1995:55), Broadley et al. (2003:106), Nagy et al. (2005:224), Wiister et al. (2007:445), Wallaeh et al. (2014:119). Boulengerina annulata annulata: Dobiey and Vogel (2007:37). Naja {Boulengerina) annulata: Wallaeh et al. (2009:31), Braneh and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. Global distribution: The the nominate subspecies is known from southern Cameroon to the southern Central African Republic and throughout the northern and central portions of the Demo¬ cratic Republic of Congo. It extends southwards through People’s Republic of Congo, Gabon and Equatorial Guinea, and reaches Angola along the lower Zaire River. The subspecies N. a. stormsi (Dollo, 1886) occurs extralimitally in the areas around Lakje Tanganyika and Lake Kivu. Ocurrences in Angola (Map 332): The species presumably occurs in the Cabinda enclave and in the northernmost portions of the country. Lunda Norte: “Along the edge of the Luele River, near the base camp, 5 km from the Carumbo Lagoon” [-7.75294, 19.95672] (Branch and Conradie 2015:200). Taxonomic and distributional notes: Water cobras {Boulengerina) have long been known to make Naja paraphyletic (Slowinski and Keogh 2000, Nagy et al. 2005, Wiister et al. 2007, Wallach et al. 2009). Wiister et al. (2007) recovered the species Paranaja multi- fasciata (Werner, 1902) as the sister taxon of Boulengerina annulata (Buchholz and Peters, 1876) and Naja melanoleuca Hallowell, 1857 and noted that the three constitute a clade of primarily forest inhabiting cobras. Wiister et al. Map 332. Distribution of Naja annulata in Angola. (2007), therefore, considered both Boulengerina and Paranaja as synonyms oiNaja. Wallach et al. (2009) recognized this clade as the subgenus Boulengerina Dollo, 1886, whereas Wallach et al. (2014) treated this group as generically distinct. Spawls and Branch (1995) and Dobiey and Vogel (2007) both suggested that the species probably occurs in Cabinda and in the Zaire region of north- 370 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II west Angola. A subadult Naja annulata from Lunda Norte Province (Branch and Conradie 2015), is the first voucheredd record for the country. Naja melanoleuca Hallowell, 1857 Forest Cobra Naja haje (var. melanoleuca) Hallowell 1857:61. Syntypes: ANSP 6875-78 (collector H. A. Ford). Type loca¬ lity: “Gaboon country, West Africa” [= Gabon]. Naja haje\ Bocage (1866a:51, 1895a:132), Peters (1877a:618). Naja haje haje: Loveridge (1957:291). Naja melanoleuca: Boettger (1898:120), Boulenger (1905:114, 1915:219), Ferreira (1900b:133, 1903:12), Parker (1936:126), Bogert (1940:87), Laurent (1950a: 10, 1954a:60), Hellmich (1957b:72), FitzSimons (1962:300), Broadley (1968c:5), Mananas (1981:27), Broadley (1990:291), Spawls and Branch (1995:71), Branch (1998:107), Broadley and Cotterill (2004:47), Chippaux (2006:225), Trape and Mane (2006:198), Dobiey and Vogel (2007:69), Bates et al. (2014:404), Wallach et al. (2014:120), Ceriaco et al. (2017:141). Naja melanoleuca melanoleuca: Laurent (1964a: 120), Thys van den Audenaerde (1966:35), Chirio and LeBreton (2007:582). Naja (Boulengerina) melanoleuca: Wallach et al. (2009:31). Boulengerina melanoleuca: Wallach et al. (2014:120). Global conservation status (lUCN): Not Evaluated. Global distribution: The N. melanoleuca species complex is known from forest and savanna through west, central and east Africa, from Senegal east to western Ethiopia, southern Somalia, south to Angola on the west coast and KwaZulu-Natal, South Africa on the east coast. Populations in the east of the range are partly disjunct. With the recent elevation of N. subfulva to specific status (Ceriaco et al. 2017), the global distribution of V. melanoleuca will need to be reevaluated. Ocurrences in Angola (Map 333): The species occurs from the northeast of the coun¬ try, including areas adjacent with Zambia in Mexico Province (pers. obs.) to the northwest, including Cabinda, and throughout western Angola, exclusive of arid regions. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Bocage 1866a:51; Ferreira 1900b:133; Mananas 1981:27); “Cabinda” [-5.55000, 12.18333] (Peters 1877a:618; Bocage 1895a: 132, Boett¬ ger 1898:120). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 10, 1954a;60, 1964a; 120; Thys van den Audenaer¬ de 1966:35; Mananas 1981:27); “Dundo, R. Capemba” (Thys van den Audenaerde 1966:35); “Dundo, Barrage de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaer¬ de 1966:35); “Dundo, Cacanda” [-7.40000, Map 333. Distribution ofiVayame/ano to in Angola. 21.80000] (Thys van den Audenaerde 1966:35); “Cossa” [-7.93333, 21.38333] (Thys van den Audenaerde 1966:35); “R. Camaiala” [-8.05000, 18.61667] (Thys van den Audenaerde 1966:35). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a;51, 1895a:132; Ferreira 1900b:133; Mananas 1981:27); “Pungo-Andongo” [-9.66667, 15.58333] (Boulenger 1905:114; Mananas 1981:27). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:72; Mananas 1981:27; Ceriaco et al. 2017:141); “Ambaca” [-9.26667, 15.18333] (Mananas 1981:27); MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 371 “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:12; Mananas 1981:27); “Cazengo” [-9.33333, 14.76667] (Mana9as 1981:27). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] (Hellmieh I957b:72; Managas 1981:27; Ceriaeo et al. 2017); “Quirimbo” [-10.68333, 14.26667] (Parker 1936:126; Managas 1981:27); “Congulu” [-10.86667, 14.28333] (Parker 1936:126; Mana- gas 1981:27). Huambo: “Galanga” [-12.06667, 15.15000] (Boeage I895a:I32). Benguela: “Caha- ta” [-12.35000, 14.81667] (Boeage I895a:I32; Ferreira I900b:I33; Managas 1981:27); “Hanha” [-12.25000, 13.75000] (Bogert 1940:87; Mananas 1981:27). Hufla: “Sanguengue” [-12.36667, 16.20000] (Hellmieh 1957b:72); “Huilla” [-15.05000, 13.55000] (Ferreira 1900b:133; Mananas 1981:27). Namibe: “Capangombe” [-15.10000, 13.15000] (Ferreira 1900b:133; Mananas 1981:27). Taxonomic and distributional notes: Broadley and Cotterill (2004) referred to Naja melanoleuca as a “species complex” that might contain several cryptic species. It was being reviewed by Broadley et al. at the time of the senior author’s death. Wallach et al. (2009) assigned Naja melanoleuca to the subgenus Boulengerina and later (Wallach et al. 2014) treated Boulenge- rina as generically distinct. Although Spawls and Branch (1995) and Dobiey and Vogel (2007) considered N. melanoleuca to be restricted to northern Angola, including the Cabinda enclave, it is much more widely spread throughout the country. Ceriaeo et al. (2017) presented molecular evi¬ dence to support Naja subfulva Laurent, 1955, as a valid species and not a mere “savannah form” or subspecies of N. melanoleuca as considered by many previous authors. The two species are eas¬ ily told apart by a combination of morphological characters and coloration, but to separate the Angolan records, many of them from before the original description of subfulva, will require a detailed review of all available specimens. Despite N. melanoleuca being considered more of a forest dweller and subfulva a savannah species, they are sympatric in parts of Angola (LMPC pers. obs.), frustrating attempts to infer the identity of records based solely on distribution. Naja mossambica Peters, 1854 Mozambique Spitting Cobra N[aja]. mossambica Peters 1854:625. Leetotype: ZMB 2811 (eolleetor W.C.H. Peters), designated by Golay et al. (1993: 188). Type loeality: “Tette and Sena” [= Tete, Vila de Sena], Mozambique, restrieted to “Tette” as a result of leetotype designation. Naja mossambica mossambica: Managas (1981:28). Naja mossambica: Broadley (1966e:25, 1974:156), Broadley (1990:295), Spawls and Braneh (1995:74), Broadley and Cotterill (2004:47), Dobiey and Vogel (2007:71), Wurster et al. (2007:438) Naja mossambica mossambica: Managas (1981:28). Naja (Afronaja) mossambica: Wallaeh et al. (2009:32), Conradie et al. (2016:22). Afronaja mossambica: Wallach et al. (2014:10). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is widespread in eastern and southern Africa, occuring from southern Tanzania southwestwards to extreme southeastern Angola, northern Namibia and Botswana, and southwards to eastern South Africa and Swaziland. Ocurrences in Angola (Map 334): The species occurs in the extreme southeastern regions of Cuando Cubango Province and there is an isolated record in Namibe Province. Namibe: “Macon- jo” [-15.01667, 13.20000] (Broadley 1974:156; Managas 1981:28). Cuando Cubango: “Cuito basin (30a)” [-17.50875, 20.06594] (Conradie et al. 2016:9, 10, 22); “Cuando basin (44b)” [-17.56916, 23.27305] (Conradie et al. 2016:9, 10, 22). Taxonomic and distributional notes: Dobiey and Vogel’s (2007) distribution map shows that the Mozambique Spitting Cobra in Angola is restricted to the extreme southeastern regions of Cuando Cubango Province, near the Zambian and Namibian borders. However, Broadley (1974, 372 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1990) reported a reeord from “Maconjo” that represents an isolated western Angolan locality, where it is sympatric with Naja (Afronaja) nigricincta Bogert, 1940. Wallach et al. (2009) proposed nomenclatural changes regarding the genus Naja: the subgeneric name Naja is appli¬ cable to the Asiatic cobras, whereas African spitting cobras, including N. mossambica, were placed in a new subgenus, Afronaja Wallach, Wiister and Broadley, 2009. Wallach et al. (2014) usQd Afronaja at the generic rank. Map 334. Distribution of Naja mossambica in Angola. Naja multifasciata Werner, 1902 Many-Banded Cobra Naia multifasciata Werner 1902:347. Holotype: IRSNB 2015 (eolleetor L. Rom). Type loeality: “der oberen Maringa” [= Upper Maringa River, Tshuapa], Demoeratie Republie of Congo. Paranaja multifasciata: Spawls and Braneh (1995:82), Nagy et al. (2005:224), Dobiey and Vogel (2007:80), Wiister et al. (2007:445). Naja (Boulengerina) multifasciata: Wallaeh et al. (2009:31). Boulengerina multifasciata: Wallaeh et al. (2014:121). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Cameroon to Congo. Ocurrences in Angola: The species is expected to occur in the Cabinda enclave and poten¬ tially Zaire Province in northwestern Angola (Spawls and Branch 1995; Dobiey and Vogel 2007) but there are no vouchered records. Taxonomic and distributional notes: Wiister et al. (2007) recovered the species Paranaja multifasciata (Werner, 1902) as the sister taxon to Boulengerina annulata (Buchholz and Peters, 1876) plus Naja melanoleuca Hallowell, 1857 and they considered both Paranaja and Boulenge¬ rina as synonyms of Naja. Althoughh long placed in the monotypic genus Paranaja, Wallach et al. (2009), based on phylogenetic information, assigned Naja multifasciata to the subgenus Boulen¬ gerina and later (Wallach et al. 2014) treated Boulengerina as generically distinct. Naja nigricincta Bogert, 1940 Western Barred Spitting Cobra Naja nigricollis var. fasciata Bocage 1895a: 136. Lectotype: MBL 1968 (collector A.P. De Carvalho), destroyed by fire 18 March 1978, designatedby Broadley (1974:158). Type locality: “Benguella ... Dondo . . . Capangombe,” Angola, restricted to “Benguella” through lectotype designation. Preoccupied by Naja fasciata Laurenti, 1768, syn. Naja naja (Linneaus, 1758). Naja nigricollis nigricinctus Bogert 1940:89, pi. 1, fig. 1. Holotype: AMNH 51823 (collector A. S. Vemay, H. Lang and R. Boulton). Type locality: “Muninho (101 km east of Mossamedes via railroad),” Namibe Province, Angola. Naja nigricollis: Bocage (1866a:51, 1866b:71, 1867b:228), Ferreria (1900b: 134), Monard (1937b: 136-137). Naja nigricollis nigricinctus: FitzSimons (1962:303). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 373 Naja mossambica nigricincta: FitzSimons (1962:306), Managas (1981:28). Naja nigricincta'. Broadley (1966c:25), Wurster et al. (2007:445). Naja nigricollis nigricincta'. Broadley (1968c:7), Spawls and Branch (1995:78). Naja nigricincta nigricincta'. Broadley (1974:158, 1990:294), Dobiey and Vogel (2007:72). Naja (Afronaja) nigricincta'. Wallach et al. (2009:32). Afronaja nigricincta'. Wallach et al. (2014:10). Global conservation status (lUCN): Not Evaluated. Global distribution: The nominate form of the speeies is known from southwestern Angola and eentral and northern Namibia. Naja nigricincta woodi Pringle, 1955 is extralimital, oeeurring in southern Namibia and western South Africa. Ocurrences in Angola (Map 335): The species is known from southwestern Angola. Benguela: “Benguela” [-12.58333, 13.41667] (Bocage 1866a:51, 1866b:71, 1867b:228, 1895a:136; Monard 1937b;136; Broadley 1974:158; Managas 1981:28); “Rio Equimina” [-13.20000, 14.78333] (Broadley 1974:158; Mananas 1981:28). Namibe: “Munhino” [-14.91667, 13.00000] (Bogert 1940:89; Fitz¬ Simons 1962:306; Broadley 1968c:7, 1974:158, 1990:294; Managas 1981:28; Wal¬ lach et al. 2014:10); “Maconjo” [-15.01667, 13.20000] (Broadley 1974:158; Managas 1981:28); “Capangombe” [-15.1 0000, 13.15000] (Bocage 1895a:136); “Foz do Cune- ne” [-17.28333, 11.80000] (Broadley 1974:158; Managas 1981:28). ™ . j j* X M X* 1 X Map 335. Distribution of Vo/a «/er/cmcto in Angola. Taxonomic and distributional notes: Bocage (1895a) recognized several varieties of Naja nigricollis Reinhardt, 1843 based on coloration (var. occidentalis; var. melanoleuca', Yar.fasciata). Naja nigricollis nigricinctus Bogert, 1940 is a synonym of Bocage’s (1895a) Yav.fasciata, but since the older name was preoccupied by Naja fasciata Laurenti (1768), a synonym ofNaja naja (Linnaeus, 1758), Broadley (1966c) accept¬ ed Bogert’s (1940) as the valid name of this species, treating it as a subspecies of N. nigricollis. Years later he treated this form as a subspecies of A. mossambica Peters, 1854 (Broadley 1968c, 1974). Bocage had recorded A. n. y2lv. fasciata from “Dondo,” however Broadley (1974) examined the Museu Bocage specimens and only found one specimen from that locality, which he nominat¬ ed as the lectotype of Naja nigricollis var. occidentalis (Bocage 1895a) (= Naja nigricollis Rein¬ hardt, 1843) whereas the A. nigricollis var. fasciata speciemens were from “Benguela” and “Maconjo,” Spawls and Branch (1995) still recognized nigricincta as a subspecies of nigricollis, whereas Wiister et al. (2007) treated it as a separate evolutionary species, with A. n. woodi as a subspecies. Naja nigricollis Reinhardt, 1843 Black-Necked Spitting Cobra Naja nigricollis Reinhardt 1843:269, pi. 3, figs. 5-7. Lectotype: ZMUC 65204 (collector J.R. Chenon) desig¬ nated by F. W. Braestmp in Hughes and Barry (1969:1027). Type locality: “Guineakysten” (Reinhardt 1843:271) [= coast of Ghana fide Hughes and Barry (1969:1027)], probably near “Christiansborg Castle, now a suburb of Accra,” [Ghana], where the collector was stationed (Rasmussen and Hughes 1997:15). 374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Naja nigricollis var. occidentalis Bocage 1895a: 135. Lectotype: MBL 1963 (collector F.A.P. Bayao), destroyed by fire 18 March 1978, designated by Broadley (1974:156). Type locality: “Dondo ... Quis- sange, Quillengues, Huilla et Humbe [Angola] ... Bissau [Guinea-Bissau],” restricted to “Dondo” [= Sumbe] Kwanza Norte Province, Angola through the designation of the lectotype. Naja nigricollis var. melanoleuca Bocage 1895a: 136 (non Hallowell, 1857). Lectotype: MBL 1972 (collector J.A. d’Anchieta), destroyed by fire 18 March 1978, designated by Broadley (1974:156). Type locality: “Catumbella et . . . Caconda” restricted to “Caconda,” Angola through the designation of the lectotype. Naja nigricollis-. Bocage (1867b:228, 1896a:113), Peters (1881:149), Boettger (1898:120), Ferreria (1900b: 134), Boulenger (1905:114), Schmidt (1933:14), Monard (1937b: 136-137), Broadley (1966c:25), Mananas (1981:28), Spawls and Branch (1995:78), Dobiey and Vogel (2007:74), Wiirst et al. (2007:438). Naia nigricollis-. Boulenger (1896:378). Naja nigricollis nigricollis-. Mertens (1938a:442), Laurent (1950a: 10, 1954a:60), Hellmich (1957a:74, 1957b:73), Loveridge (1957:292), FitzSimons (1962:302), Broadley (1974:156, 1990:292). Naja nigricollis occidentalis-. Laurent (1964a: 119), Thys van den Audenaerde (1966:35), Broadley (1968c:7). Naja (Afronaja) nigricollis-. Wallach et al. (2009:32). Afronaja nigricollis-. Wallach et al (2014:10). Global conservation status (lUCN): Not Evaluated. Global distribution; The speeies is widely distributed from southern Mauritania and Senegal east to Ethiopia and Somalia, and south to Zambia, Malawi, the southern Democratie Republic of Congo, and Angola. Absent from the forests of Central and West Africa, with disjunct populations along the lower Congo River, in southwestern Angola and elsewhere. Ocurrences in Angola (Map 336): The species very widespread, from coastal areas to the interior highlands of Angola. Zaire: “Ambrizette” [-7.23333, 12.86667] (Bocage 1895a: 135; Boettger 1898:120; Mananas 1981:28). Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1881:149; Bocage 1895a: 135; Managas 1981:28). Kwanza Norte: “Golungo Alto” [-9.13333, 14.76667] (Boulenger 1905:114; Monard 1937b: 136); “Dondo” [-9.68333, 14.43333] (Bocage 1895a: 135; Ferreira 1900b: 134; Monard 1937b: 136; Loveridge 1957:292; Hellmich 1957b:73; Broadley 1968c:7; Mana9as 1981:28). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] (Hellmich 1957b:73; Managas 1981:28); “Dondo a Humbe” (Ferrei¬ ra 1900b:134). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:14). Benguela: 1867b:228, 1895a:135; Ferreira 1900b:134; Map 336. Distribution of Naja nigricollis in Angola. Catumbella” [-12.43333, 13.55000] (Bocage Mananas 1981:28); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:135; Ferreira 1900b:134; Monard 1937b:136; Loveridge 1957:292; Broadley 1968c:7; Managas 1981:28); “Chivitidi/Entre Rios” [-13.01667, 14.63333] (Hellmich 1957a:74; Managas 1981:28); “Cubal” [-13.03333, 14.25000] Mertens 1938:442; Managas 1981:28); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113; Managas 1981:28). Huila: “Cacon¬ da” [-13.73333, 15.06667] (Bocage 1895a:135; Ferreira 1900b:134; Monard 1937b:136; Managas 1981:28); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:135; Monard 1937b:136; Loverid¬ ge 1957:292; Broadley 1968c:7; Managas 1981:28); “Kuvangu/Kubangu/Vila da Ponte” MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 375 [-14.46667, 16.30000] (Monard I937b:I36-I37; Managas 1981:28); “Capelongo” [-14.88333, 15.08333] (Managas 1981:28); “Huilla” [-15.05000, 13.55000] (Bocage I895a:I35; Monard I937b:I36; Loveridge 1957:292; Broadley I968c:7; Mana9as 1981:28); “Osi” [-15.08333, 15.40000] (Monard I937b:I36-I37; Managas 1981:28); “Gambos” [-15.76667, 14.10000] (Mana- 9as 1981:28). Namibe: “Capangombe” [-15.10000, 13.15000] (Ferreira I900b:I34; Monard I937b:I36; Managas 1981:28). Cunene: “Humbe” [-16.68333, 14.90000] (Boeage I895a:I35; Ferreira 1900b: 134; Monard 1937b: 136; Loveridge 1957:292; Broadley I968c:7; Mana9as 1981:28); “Mupanda” [-17.13333, 15.76667] (Monard I937b:I36-I37; Managas 1981:28). Cuan- do Cubango: “Kayundu” [-15.70000, 17.45000] (Monard I937b:I36-I37; Mananas 1981:28). Taxonomic and distributional notes: Bocage’s (1895a) locality of “Ambrizette” in Zaire Province has been considered questionable and has been excluded from some distribution maps for the species (Spawls and Branch 1995; Dobiey and Vogel 2007). However, Dobiey and Vogel (2007) do include Cabinda within the range of the species, although we are not aware of any published records. These authors also indicate occurrence in northcentral Namibia, although only a single specimen, from Katima Mulilo in the eastern Caprivi Strip has been documented from that country (Broadley 1974). The taxonomy of N. nigricollis still requires further study (Wiister et al. 2007). Wallach et al. (2009) placed the African spitting cobras in a new subgenus, Afronaja Wallach, Wiister and Broadley, 2009, with Naja nigricollis as the type species and Wallach et al. (2014) have treated this as a full genus. Naja subfulva Laurent, 1955 Savanna Cobra Naja melanoleuca subfulva Laurent 1955:61. Holotype: MRAC 17514 (collector Van der Borght). Type loca¬ lity: “Lwiro, 1850 m, Terr, de Kabare, Kivu,” Democratic Republic of the Congo. Naja subfulva: Ceriaco et al. (2017:141). Global conservation status (lUCN): Not Evaluated. Global distribution: The N. melanoleuca species complex is known from forest and savanna through west, central, and east Africa, from Senegal east to western Ethiopia, southern Somalia, south to Angola on the west coast and KwaZulu-Natal, South Africa on the east coast. Populations in the east of the range are partly disjunct. With the recent elevation of N. subfulva to specific status (Ceriaco et al. 2017), the global distribu¬ tion of both this taxon and N. melanoleuca sensu stricto will need to be reevaluated. Ocurrences in Angola (Map 337): The species should occur across the country, except in the more heavily forested areas in the north, but currently the only confirmed records are for Kwanza Norte, Kwanza Sul and Malanje provinces. Malanje: “Cacuso” [-9.421023, 15.746679] (Ceriaco et al. 2017:141). Kwan¬ za: Norte: “Piri-Dembos” [-8.56667, 14.50000] (Ceriaco et al. 2017:141). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] (Ceriaco et al. 2017:141). Taxonomic and distributional notes: See Naja melanoleuca account. Il-E t6°E 20°E 24“ E Map 337. Distribution of Naja subfulva in Angola. 376 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Pseudohaje Gunther, 1858 Pseudohaje goldii (Boulenger, 1895) African Tree Cobra Naia Goldii Boulenger 1895d:34. Holotype: BMNH 1946.1.21.43 (formerly BMNH 1895.5.3.53) (eolleetor W.H. Crosse). Type loeality: “Territories of the Royal Niger Company, near Asaba, about 150 miles up the Niger” (Boulenger 1895d:32-33) [= vineinity of Asaba, Niger River], southern Nigeria. Naia goldii: Boulenger (1915:219). Naja goldii: Parker (1936:126). Pseudohaje goldii: Bogert (1942:7), Laurent (1950a: 10, 1954a:61), Hellmieh (1957b:73), Spawls and Braneh (1995:84), Spawls et al. (2004:460), Chippaux (2006:230), Chirio and LeBreton (2007:590), Dobiey and Vogel (2007:81), Wallaeh et al. (2014:591). Pseudohaje goldii {Naia goldii): Managas (1981:28). Bogert (1942) reviewed the Genus Pseudohaje and considered the species Pseudohaje goldii Boulenger, 1895 to be confined to West African forests. Additional distributional data on P. goldii are presented in the maps in Spawls and Branch (1995), Spawls et al. (2004), Chip¬ paux (2006) and Dobiey and Vogel (2007). Broadley (1990) presented evidence refuting Map 338. Distribution of ftewJo/jaye go/ J// in Angola. the occurrence of R goldii in Namibia as proposed by Mertens (1955). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from west and central African forests, from Nige¬ ria in the northwest, eastward through Cameroon and the Democratic Republic of Congo to Ugan¬ da, western Kenya, and possibly Rwanda (Spawls et al. 2004) and southward to northern Angola. Isolated populations in the Cote d’Ivoire and Ghana and possibly Togo (Segniagbeto et al. 2011). Ocurrences in Angola (Map 338): The species occurs in the northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Lau¬ rent 1950a; 10, 1954a:61; Managas 1981:28). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:73; Managas 1981:28). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:126; Man- agas 1981:28). Taxonomic and distributional notes; Family Colubridae Oppel, 1811 Genus Chamaelycus Boulenger, 1919 Chamaelycus parkeri (Angel, 1934) Parker’s Banded Snake Oophilositum parkeri Angel 1934:417. Holotype: MNHN 1934-11 (eolleetor G. Babault). Type loeality: “Kabulire (dans les forets situees a I’Ouest du lae Kivu)” [= Kabulire, Sud-Kivu] Demoeratie Republie of Congo. Oophilositum parkeri: Parker (1936:123). Chamaelyeus parkeri: Chippaux (2006:72), Chirio and LeBreton (2007:392), Wallaeh et al. (2014:155). Global conservation status (lUCN): Not Evaluated. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 377 Global distribution: The species is distributed in Central Africa, from northern Congo to the Democratic Republic of Congo and Angola. Ocurrences in Angola (Map 339): The species has only been recorded from “Congu- lu,” Kwanza Sul Province in western Angola but it is probable that the species occurs further north in the country. Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:123). Taxonomic and distributional notes: Further surveys in the country may yield additional records of this species in northern Angola. I1«E 16°E 20“E 24<’E Map 339. Distribution of Chamaelycus parkeri in Angola. Genus Crotaphopeltis Fitzinger, 1843 Crotaphopeltis hotamboeia (Laurenti, 1768) Red-Lipped Snake Coronella hotamboeia Laurenti 1768:85. Holotype: specimen illustrated by Seba (1734:54, pi. 33, fig. 6), specimen not located. Type locality: “in India orientali” [= East Indies] 7/t/e Laurenti (1768:85), [Ceylon = Sri Lanka]7/Je Seba (1734), in error, corrected to “Africa” by Loveridge (1957:271). Crotaphopeltis rufeseens: Bocage (1866a:49, 1895a: 122), Peters (1881:149), Bocage (1896a: 113, 1897b:211). Leptodira rufeseens: Gunther (1876b:679), Peters (1877a:615), Bocage (1879b:95). Crotaphopeltis semiannulatus: Bocage (1895a: 122). Leptodira hotamboeia: Ferreira (1903:12, 1904:116, 1906:169), Boulenger (1905:112), Monard (1937b:129). Crotaphopeltis hotamboeia: Parker (1936:125), Branch and McCartney (1992:2), Broadley et al. (2003:203), Broadley and Cotterill (2004:51), Chippaux (2006:143), Spawls et al. (2004:379), Trape and Mane (2006:88), Chirio and LeBreton (2007:398), Wallach et al. (2014:197), Branch and Conradie (2015:200), Ceriaco et al. (2016b:77), Conradie et al. (2016:19). Crotaphopeltis hotamboeia hotamboeia: Mertens (1938:440), Hellmich (1957a:72, 1957b:68), Laurent (1950:9; 1964a:110). Global conservation status (lUCN): Not Evaluated. Global distribution: This species is one of the most widespread African snakes, typically occupying moist savannas of sub-Saharan Africa. Ocurrences in Angola (Map 340): The species occurs in the entire country (including the Cabinda enclave), with exception of the desert regions of the far southwestern Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:49, 1895a:122). Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:122). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Peters 1881:149); “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200); “Cassange” [-9.58333, 17.86667] (Bocage 1879b:95). Lunda Sul: “Alto Cuilo, galerie forestiere des sources du ruisseau Tcha-Muchito” [-10.01667, 19.45000] (Laurent 1964a:110). Bengo: “Dande” [-8.46667, 13.38333] (Bocage 1895a:122). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:49); “Cafuxi” [-9.24000, 17.17167] (Boulenger 1906:169); “Cangandala National Park” [-9.84606, 16.72233] 378 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Map 340. Distribution of Crotaphopeltis hotamboeia in Angola. (Ceriaco et al. 201 6b: 77). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hell- mich 1957b:68); “N’golla Bumba” [-9.03333, 14.60000] (Ferreira 1906:169); “Golungo Alto” [-9.13333, 14.76667] (Ferreira 1906:169); “Cambondo” [-9.15963, 14.65771] (Ferreira 1906:169); “Canhoca” [-9.25000, 14.68333] (Boulenger 1905:112); “Rio Lui- nha” [-9.26667, 14.53333] (Ferreira 1906:169); “N’dalla Tando” [-9.30000, 14.91667] (Bocage 1895a:122); “Dondo” [-9.68333, 14.43333] (Ferreira 1903:12). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:125); “Chingo” [-11.20000, 13.85000] (Ferreira 1904:116). Mexico: “Maeondo” [-12.55000, 23.76667] (Laurent 1964a: 110). Huambo: “Bela- Vista” [-12.36667, 16.20000] (Hellmieh 1957b:68). Bie: “Cubango basin (12b)” [-13.59638, 16.87722] (Conradie et al. 2016:8-9, 19). Benguela: “Quissange” [-12.43333, 14.05000] (Boeage 1895a:122); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:122); “Benguella” [-12.58333, 13.41667] (Bocage 1895a:122); “Katala” [-12.88333, 14.01667] (Ferreira 1906:169); “Entre Rios” [-13.01667, 14.63333] (Hellmieh 1957a:72); “Alto Cubal” [-13.03333, 14.25000] (Mertens 1938:440; Hellmieh 1957b:68); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:113, 1897b:211). Hufla: “Sangeve” [-13.88333, 15.83333] (Monard 1937b: 129); “Caconda” [-13.73333, 15.06667] (Bocage 1895a:122); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a:122); “Kuvangu/Vila da Ponte” [-14.46667, 16.30000] (Monard 1937b:129); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:122; Hellmieh 1957b:68); “Kampulu (region de Kasinga)” [-15.21667, 16.11667] (Monard 1937b:129); “Gambos” [-15.76667, 14.10000] (Bocage 1895a:122). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a: 122). Cunene: “Humbe” [-16.68333, 14.90000] (Bocage 1895a: 122). Cuando Cubango: “approximately 23 km SE of Cuito Cuanavale” [-15.33333, 19.33333] (Branch and McCartney 1992:2). Undetermined Locality: “Cuango” (Peters 1881: 149; Bocage 1895a: 122) (Malanje Province impossible to georeference: See History Section for more detailed information). Taxonomic and distributional notes: Wallach et al. (2014) indicated that no holotype had been indicated for this species. However, Laurenti (1768) provided an indication to an image in Seba (1734) and the individual pictured is the holotype. The history of Seba’s collections have been discussed in detail (Engel 1937, 1961; Boeseman 1970; Juriev 1981; Adler 1989; Bauer 2002; Bauer and Gunther 2013). Seba’s second collection (the first had been sold to Peter the Great of Russia in 1716; Driessen-van het Reve 2006) was sold after his death at auction (Anonymous 1752). Seba specimens are known or believed to be present in collections in St. Petersburg, London, Leiden (including specimens until recently in Amsterdam), Paris, Copenhagen, Stock¬ holm, Bremen and Berlin (Boeseman 1970; Juriev 1981; Thireau et al. 1998; Bauer and Gunther 2013) but few can be traced to particular plate figures, and the lectotype of Coronella hotamboeia is not among these. This species has a wide distribution in sub-Saharan Africa (Broadley and Cotterill 2004) and a phylogeographic analysis is therefore desirable to investigate the possibility of cryptic species (Bates et al. 2014). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 379 Genus Dasypeltis Wagler, 1830 Dasypeltis palmarum (Leach, 1818) Palm Egg Eater Coluber palmarum Leach 1818:408. Holotype: 1946. 1.2.35 (collector J. Cranch). Type locality: “in palm trees at Embomma” (Leach 1818:409) [= Boma], Democratic Republic of Congo. Dasypeltis palmarum: Gunther (1865a:480), Peters (1877a:615). Dasypeltis seabra var. inornatum: Bocage (1867b:227). Dasypeltis seabra var. palmarum: Bocage (1895a: 106), Boulenger (1905:112), Dasypeltis seabra var. palmarum (var. inornatus): Ferreira (1903:10). Dasypeltis seabra: Parker (1936:125). Dasypeltis seabra seabra: Bogert (1940:85). Dasypeltis palmarum: Gans (1959:219), Chippaux (2006:115), Wallach et al. (2014:208). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from Angola, People’s Republie of Congo, and Demoeratie Republie of Congo. Ocurrences in Angola (Map 341): The speeies oeeurs in the western Angola. The speeies is distributed from the Cabinda enelave to Benguela Provinee, mostly near the eoast. Cabinda: “Chinehoxo” [-5.10000, 12.10000] (Peters 1877a:615; Boeage 1895a; 106; Gans 1959:220); “Chiloango” [-5.18333, 12.18333] (Gans 1959:220); “Landana” [-5.21667, 12.15000] (Gans 1959:220). Luanda: “Caeua- eo, Luanda Disk” [-8.78333, 13.36667] (Gans 1959:220). Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Gunther 1865a:480; Boulenger 1905:112; Monard 1937b:113; Managas 1973:193; Gans 1959:220). Kwanza Norte: “Ambaea” [-9.26667, 15.18333] (Boca¬ ge 1895a:106; Monard 1937b:113; Managas 1973:193); “Rio Muembege, proximo de N’dalla Tando (Cazengo)” (Ferreira 1903:10); Distribution of Dasypeltis palmanm in “N’dala Tando” [-9.30000, 14.91667] (Monard 1937b:113; Managas 1973:193); “Dondo” [-9.68333, 14.43333] (Gans 1959:220; Managas 1973:193). Kwanza Sul: “Congulu” [-9.66667, 15.58333] (Parker 1936:125; Gans 1959:220). Benguela: “Lobito Bay (baia do Lobito)” [-12.33333, 13.58333] (Bogert 1940:85; Managas 1973:193; Gans 1959:220); “Catumbella” [-12.43333, 13.55000] (Bocage 1867b:227, 1895a:106; Monard 1937b:113; Gans 1959:220); “Quissange” [-12.43333, 14.05000] (Bocage 1895a:106; Monard 1937b:113); “Dombe” [-12.95000, 13.10000] (Bocage 1867b:227, 1895a:106; 1937b:113; Gans 1959:219). Huila: “Caconda” [-13.73333, 15.06667] (Gans 1959:219). Namibe: “Mossa- medes” [-15.20000, 12.15000] (Gans 1959:219). Taxonomic and distributional notes: None. Dasypeltis seabra (Linnaeus, 1758) Common Egg Eater Coluber seaber Cirmatus 1758:223. Holotype: NHR, formerly MAFR, (Mus. Drottingholm), lost fide AndQr- sson (1899) and Gans (1959:87). Type locality: “in Indiis” [India], in error, corrected to “South Africa” by Flower (1933:818). 380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Dasypeltis fasciolata: Peters (1877a:615). Dasypeltis scabra: Boeage (1895a: 106, 18976:210), Boulenger (1893:356, 1896:648), Ferreira (1904:115), Monard (19376:113, 123), Mertens (1938a:440), T6emido (1941:10), Broadley and Cotterill (2004:52), Spawls et al. (2004:418), Bates et al. (2014:411), Wallaeh et al. (2014:208), Ceriaeo et al. (2016a:81). Dasypeltis scabra var. medici: Boeage (1895a: 106). Dasypeltis scabra var. 3 (var. fasciolata?): Boeage (1895a: 106). Dasypeltis scabra var. inornata: Ferreira (1906:168). Dasypeltis scabra scabra: Laurent (1954a:60, 1964a: 116), Hellmie6 (19576:72), T6ys van den Audenaerde (1966:35), Managas (1973:192). Global conservation status (lUCN): Least Coneem. Global distribution: The species is widely distributed, extending from the south-western Cape northwards to Zambia and the southern half of the Democratic Republic of the Congo, further along the eastern half of the continent to Ethiopia and adjacent parts of eastern Sudan, with isolated records in Egypt and the Arabian Peninsula. Ocurrences in Angola (Map 342): The species occurs in the entire country (including Cabinda enclave), with exception of the desert regions of the far southwestern Angola. Cabin¬ da: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Managas 1973:193); “Molembo” [-5.33333, 12.20000] (Boeage 1895a:106; Mananas 1973:193); “Cabinda” [-5.55000, 12.18333] (Boeage 1895a: 106; Mana9as 1973:193). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Boeage 1895a:106; Managas 1973:193). Kwanza Norte: “Piri- Dembos” [-8.56667, 14.50000] (Hellmich 1957b:72); “Cambondo” [-9.15963, 14.65771] (Boulenger 1893:356; Mana9as 1973:193). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:60, 1964a: 116; Thys van den Audenaerde 1966:35; Mana9as 1973:193); Map 342. Distribution of Dasypeltis scabra in Angola. “entre Capaia et Carumbo” (Laurent 1964a: 117). Lunda Sul: “Mutianvo” [-11.45000, 19.33333] (Themido 1941:10). Luanda: “Cacuaca” [-8.78333, 13.36667] (Boulenger 1893:356). Malanje: “Cangandala National Park” [-9.84606, 16.72233] (Ceriaeo et al. 2016a:81). Moxico: “Calombe, Luso” [-11.83333, 19.93333] (Mana9as 19732:192). Huambo: “Bela-Vista” [-12.36667, 16.20000] (Hellmich 1957b:72). Benguela: “Catumbella” [-12.43333, 13.55000] (Boeage 1895a:106; Mana9as 1973:193); “Quissange” [-12.43333, 14.05000] (Boeage 1895a:106; Mana9as 1973:193); “Quindumbo” [-12.46667, 14.93333] (Boeage 1895a:106; Monard 1937b:113; Mana- 9as 1973:193); “Cubal” [-13.03333, 14.25000] (Mertens 1938a:440); “Hanha” [-13.30000, 14.20000] (Boeage 1897b:210). Huila: “Caconda” [-13.73333, 15.06667] (Boeage 1895a:106; Monard 1937b:113); “Vila-da-Ponte/Kuvangu” [-14.46667, 16.30000] (Monard 1937b:113, 123); “Gambos” [-15.76667, 14.10000] (Boeage 1895a:106; Monard 1937b:113; Mana9as 1973:193). Cuando Cubango: “Kakindo” [-15.45000, 17.05000] (Monard 1937b: 113, 125; Mana9as 1973:193). Taxonomic and distributional notes: Gans (1959) suggested that the most likely source of the type specimen, based on its described morphology, would be “South West Africa [Namibia] or MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 381 the northern Transvaal [Limpopo Province, South Africa].” However, these areas are extremely unlikely to have yielded specimens destined for Europe in the 1730s. The taxonomy of this species in southern Africa is being investigated, and the presence of cryptic taxa has been suggested (M. Bates et al. in prep.). Genus Dipsadoboa Gunther, 1858 Dipsadoboa shrevei (Loveridge, 1932) Shreve’s Tree Snake Crotaphopeltis shrevei Loveridge 1932b:83. Holotype: MCZ R-32471 (eolleetor K.H. Prior). Type loeality: “Missao de Dondi, Bella Vista, via. Lobito, Angola” [= Caehiungo], Huambo Provinee, Angola. Crotaphopeltis shrevei: Sehmidt (1933:13), Barbour and Loveridge (1946:97), Laurent (1951:210, 1964a: 110), Broadley (1968d:138). Dipsadoboa shrevei shrevei: Rasmusssen (1986:59). Dispadoboa shrevei: Broadley et al. (2003:210), Broadley and Cotterill (2004:50), Spawls et al. (2004:383), Wallaeh et al. (2014:228). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from lower Congo and Angola, east through the Democratic Republic of Congo and north¬ ern Zambia to southeast Tanzania. Ocurrences in Angola (Map 343): Pub¬ lished species records are in the central interior of Angola. Moxico: “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:110; Rasmusssen 1986:64). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:13; Ras¬ musssen 1986:64). Huambo: “Bella Vista, Missao de Dondi” [-12.56667, 16.21667] (Loveridge 1932b; 83; Barbour and Loveridge 1946:97; Rasmusssen 1986:64; Wallaeh et al. 2014:228). Taxonomic and distributional notes: Rasmussen’s (1993) distribution records in adjacent regions suggest, that despite the lack of literature records, it is possible that this species might extend to the Cabinda encalve. Map 343. Distribution of Dipsadoboa shrevei in Angola. Genus Dispholidus Duvernoy, 1832 Dispholidus typus typus (Smith, 1828) Boomslang Bueephalus typus Smith 1828:2. Syntype: RSM, identified by FitzSimons (1937:263), however, not reeorded as a type by Herman et al. (1990). Type loeality: “eastern distriets of South Afriea” (Smith 1828:2), restrieted to “Old Latakoo [= Lattakoo or Lithako, approx. 27°S, 24°E, i.e. btwn. Kumman and Taungs, N. Cape Provinee],” [= Northern Cape or Northwest Provinee], South Afriea. Bueephalus typus: Boeage (1866a:48, 1870:68, 1879b:95), Peters (1881:149), Ferreira (1900a:52). Bueephalus eapensis: Boeage (1895a: 121). Dispholidus {Bueephalus) typus: Ferreira (1897b:244). Dispholidus typus: Sehmidt (1933:14), Boulenger (1905:113, 1915:213), Loveridge (1936a:40), Monard (1937b: 128, 134), Mertens (1938:441), Bogert (1940:68), Themido (1941:10), FitzSimons (1962:196), Mananas (1973:193), Broadley (1990:252), Spawls and Braneh (1995:20), Braneh (1998:99), Broadley 382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II and Wallach (2002:72), Broadley and Cotterill (2004:51), Broadley et al. (2003:218), Chippaux (2006:159), Trape and Mane (2006:96), Eimermacher (2012:1), Ceriaco et al. (2016b:79), Wallaeh et al. (2014:236). Dispholidus typus punctatus: Braneh and MeCartney (1992:2). Dispholidus typus occidentalis: Chirio and LeBreton (2007:422). Global conservation status (lUCN): Not Evaluated. Global distribution: The nominotypical subspeies is known from Senegal east across the Sahel to the Horn of Africa, south to the southwestern Cape, excluding areas of rain forest, grass¬ land and desert. Ocurrences in Angola (Map 344): The nominotypical subspecies distribution com¬ prises almost the entire country including Cabinda Province, with the exception of the desert regions of far southwestern Angola, and the area occupied by D. t. punctatus. Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a:121; Monard 19376:128). Luanda: “Loanda” [-8.83333, 13.26667] (Fer¬ reira 1900a: 52). Malanje: “Duque de Bragan- 5a” [-9.10000, 15.95000] (Bocage 1866a;48, 1895a:121; Monard 1937b:128); “Malanje” [-9.55000, 16.35000] (Peters 1881:149; Boca¬ ge 1895a: 121); “Cangandala National Park” [-9.84606, 16.72233] (Ceriaco et al. 2016b:79). Lunda Sul: “Tyihumbwe” [-10.96667, 20.06667] (Monard 19376:128, 134); “Mutian- vo” [-11.45000, 19.33333] (Themido 1941:10). ^44. Distribution of Dispholidus typus typus in Kwanza Norte: “Ambaca” [-9.26667, 15.18333] (Bocage 1895a:121; Ferreira 1900a:52; Monard 19376:128); “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:12); “Cazengo” [-9.33333, 14.76667] (Ferreira 1900a:52). Kwanza Sul: “Mombolo” [-12.16667, 14.83333] (Bogert 1940:68). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:14). Huambo: “Galanga” [-12.06667, 15.15000] (Bocage 1895a:121; Monard 19376:128); “Santo-Amaro” [-12.70000, 15.85000] (Monard 19376:134). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1895a: 121; Monard 19376:128); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:121; Monard 19376:128); “Ebanga” [-12.73333, 14.73333] (Monard 19376:134); “Cubal” [-13.03333, 14.25000] (Mertens 1938:441). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1895a:121; Ferreira 1897:244; Loveridge 1936:40; Monard 19376:128); “Kalukembe” [-13.78333, 14.68333] (Monard 19376:128, 134); “Kuvangu/Vila da Ponte” [-14.46667, 16.30000] (Monard 19376:128, 134); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:68); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:121; Monard 19376:128). Cunene: “Mupa” [-16.18333, 15.75000] (Monard 19376:134); “Humbe” [-16.68333, 14.90000] (Bocage 1895a:121; Monard 19376:128). Cuando Cubango: “approximately 24 km E of Cuito Cuanavale” [-15.18333, 19.38333] (Branch and McCartney 1992:2). Undetermined Locality: “Rio Cuando” (Bocage 1895a:121, Monard 19376:128); “Between Benguella and Bihe” (Boulenger 1905:113; Monard 19376:128). Taxonomic and distributional notes: Dispholidus typus is often cited as having been described in 1829 (e.g., FitzSimons 1962) in The Zoological Journal, however, the description first ll-E 20°E 24” E MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 383 appeared one year earlier in the South African Commercial Advertiser in a short paper that has been reprinted by Ulber (2003) and Braneh and Bauer (2005). FitzSimons’ restrieted type loeality is derived from the stated type loeality of Bucephalus viridis Smith, 1841, a subjeetive junior syn¬ onym of B. typus Smith, 1828. Four subspeeies: D. t. typus, D. t. viridis Smith, 1828, D. t. kivuen- sis Laurent, 1955 andZ). t. punctatus Laurent, 1955 were reeognised in the past. Preliminary genet- ie analyses earried out by Eimermaeher (2012) indieate that multiple distinet lineages may be pres¬ ent, some of which should be resurrected and elevated to species status, including D. t. punctatus Laurent, 1955 from “Dund,” Angola. Eimermaeher and Broadley (in prep.) are currently undertak¬ ing a detailed revision of the genus. All Angolan material should be reevaluated in order to deter¬ mine where species boundaries lie. Eimermaeher ’s (2012) sampling was sparse in southwestern Africa, but his data suggested that central Namibian material was referable to to D. t. viridis (Smith, 1838). If this form also reaches southern Angola and is likely valid, as suggested by Eimermaeher (2012), then it, rather than D. t. typus, would be the widespread Boomslang in Angola. However, as no published records from Angola have previously been referred to D. t. viridis, we continue to use the nominotypical epithet for Angolan specimens pending relevant genetic data. Dispholidus typus punctatus Laurent, 1955 Spotted Boomslang Dispholidus typus punctatus Laurent 1955:129. Holotype: RGMC 17395 (currently MRAC) (collector A. Bar- ros Machado). Type locality: “Dundo,” Lunda Norte Province, Angola. Dispholidus typus: Laurent (1950a: 10, 1954a:57), Mananas (1973:193). Dispholidus typus punctatus: Hellmich (1957b:68), Laurent (1964a: 114), Thys van den Audenaerde (1966:35), Broadley (1990:255), Spawls and Branch (1995:21), Branch (1998:99), Broadley and Wallach (2002:72), Broadley et al. (2003:218), Broadley and Cotterill (2004:51), Eimermaeher (2012:2), Branch and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. Global distribution: The subspecies is known from forest regions of southern Democratic Republic of Congo, Angola and Zambia. Ocurrences in Angola (Map 345): Dispholidus typus punctatus occurs mainly in the eastern regions of the country. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a: 10, 1954a:57, 1955:129, 1964a: 114; Thys van den Audenaerde 1966:34; Broadley and Wallach 2002:72; Eimermaeher 2012:23); “Dundo, R. Mussungue” [-7.41667, 20.83333] (Thys van den Audenaerde 1966:35); “Mwaoka (± 45 km S. Dundo)” [-7.65000, 20.85000] (Thys van den Audenaerde 1966:34); “Carumbo” [-7.74422, 19.95467] (Branch and Conradie 2015:200); “R. Camaiala” [-8.05000, 18.61667] (Thys van den Audenaerde 1966:34); “R. Capemba” [-9.25167, 19,55808] (Thys van den Audenaerde 1966:35). Lunda Sul: “Alto Cuilo” [-10.01667, 19.55000] (Lau¬ rent 1964a: 114); “Chutes du Cuango-Muque, Alto Chicapa” [-10.76667, 19.20000] (Laurent 1964a: 114). Mexico: “Fazenda Santa Cruz, Luso” [-11.78333, 19.91667] (Managas 12*E l(i*E 2(}°£ 24*E Map 345. Distribution of Dispholidus typus punctatus in Angola. 384 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1973:193-194); “environs du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a:114); “Calom- be, Luso” [-11.83333, 19.93333] (Managas 1973:193-194); “Cazombo” [-11.88333, 22.91667] (Laurent 1964a:114). Huambo: “Bela-Vista” [-12.36667, 16.20000] (Hellmich 1957b:68); “Nova Lisboa” [-12.76667, 15.73333] (Laurent 1955:129). Benguela: “Alto Cubal” [-13.03333, 14.25000] (Hellmich 1957b:68). Taxonomic and distributional notes: Although Broadley and Wallach (2002) considered punctatus a synonym of D. typus, Eimermacher (2012) indicated that multiple distinct lineages of boomslang should be probably be resurrected and elevated to species status, including D. t. punc¬ tatus. On this basis we list this form as a distinct taxon, but retain its subspecific status pending fur¬ ther data. All Angolan material should be reevaluated in order to determine where species bound¬ aries lie. It seems likely on biogeographic grounds that some of the more southern and western records previously attributed to D. t. punctatus may, in fact, belong to the nominotypical form or to D. t. viridis (see Notes und Dispholidus typus typus). Genus Grayia Gunther, 1858 Grayia caesar (Gunther, 1863) Caesar’s African Water Snake Xenurophis caesar Gunther 1863:357, pi. 6, fig. c. Holotype: BMNH 1946.1.6.38 (formerly BMNH 1843.1.10.7) (eolleetor W. Raddon). Type loeality: “Fernando P6” [= Bioko], Equatorial Guinea. Grayia caesar: Chippaux (2006:94), Chirio and LeBreton (2007:432), Wallaeh et al. (2014:310). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Cameroon, Central African to Democratic Republic of the Congo. Ocurrences in Angola (Map 346): The species has been recorded from the Cabinda enclave. Cabinda: “Mouth of the Loango” [-5.15000, 12.16667] (Boulenger 1894:288). Taxonomic and distributional notes: This species was only recorded from Angola by Boulenger (1894), this record could correspond to Grayia smithii (Leach, 1818), however, the extralimital records of G. caesar (Chippaux 2006; Chirio and LeBreton 2007; Wallach et al. 2014) are consistent with its occurrence in the Cabinda enclave. Il'E 1&°E 20°E 24'E Map 346. Distribution of Grayia caesar in Angola. Grayia ornata (Bocage, 1866) Ornate African Water Snake Maerophis ornatus Bocage (1866a:47, 1866b:67). Syntypes: ZMB 7772 (formerly MBL), two additional MBL speeimens, numbers unknown (eolleetor F.A.P. Bayao), destroyed by fire 18 Mareh 1978. Type loeality: Type loeality: “Duque de Braganga dans I’interior d’Angola” (Boeage 1866a:47) [= Calandula], Malanje Provinee, Angola. Glaniolestes ornatus: Peters (1877a:614). Graya ornata: Boeage (1897a:200, 1895a: 104), Boulenger (1915:207), Loveridge (1936a:34), Laurent (1950a:9, 1954a:44, 1964a:102), Thys van den Audenaerde (1966:34), Broadley (1983:21), Broadley et al. (2003:179), Broadley and Cotterill (2004:49), Chippaux (2006:95), Chirio and LeBreton (2007:434), Wal¬ laeh et al. (2014:310). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 385 Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies extends from Cameroon south to Angola, west to the former Katanga Province of the Democratic Republic of Congo and northwestern Zambia. Ocurrences in Angola (Map 347): The species is limited to northern regions of the country including the Cabinda encalve. Cabin¬ da: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:614; Bocage 1895a: 104). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Bocage 1895a:104; Laurent 1950a:9, 1954a:44, 1964a: 10; Thys van den Audenaerde (1966:34); “in a tributary of the right side of the Tchihumbwe about 50 km west of Dundo” [-7.38333, 20.85000] (Laurent 1950:9); “Dundo (Mussungue river)” [-7.41667, 20.83333] (Thys van den Audenaerde 1966:349); “Luachimo” [-7.53333, 21.08333] (Laurent 1950a:9). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:47, 1866b:67, 1895a: 104, 1897a:200, Loveridge 1936a:34; Broadley 1983:21; Chippaux 2006:95; Chirio and LeBreton 2007:434; Wallach et al. 2014:310). Taxonomic and distributional notes: The species was described as new by Bocage in two different papers (1866a, b), both of which appeared in the November 1866 issue of the same jour¬ nal. Grayia smythii (Leach, 1818) Smyth’s African Water Snake Coluber Smythii Leach 1818:409. Holotype: BMNH 1946.1.1.67 (formerly BMNH 48.9.15.10) or BMNH 1946.1.5.16 (collector J. Cranch, don. W.E. Leach), see Boulenger (1894:287). Type locality: “near Embomma” (Leach 1818:409), [= vicinity of Boma, Bas-Congo Province] Democratic Republic of Congo. Grayia triangularis: Bocage (1866a:47, 1895a: 102). Grayia smithii: Boulenger (1894:287, 1915:207). Grayia smithi: Ferreira (1906:168), Laurent (1950a:8, 1964a: 102), Thys van den Audenaerde (1966:34), Trape and Mane (2006:104), Chirio and LeBreton (2007:436) Grayia smythii: Spawls et al. (2004:337), Chippaux (2006:93), Wallach et al. (2014:311). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Lake Victoria west to Senegal, north into the southern Sudan and southwest to northern Angola. Ocurrences in Angola (Map 348): The species occurs in the northern Angola. Cabinda: “Mouth of the Loango” [-5.15000, 12.16667] (Boulenger 1894:287). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a:102); “Barrage de la Luachimo” [-7.38333, 20.85000] (Thys van den Audenaerde 1966:34); “Dundo, riviere Luachimo” [-7.38333, 20.85000] (Laurent 1950a:8). Bengo: “Rio Dande” [-8.46667, 13.38333] (Bocage 1895a: 102). Kwanza Norte: “Cam- bondo” [-9.15963, 14.65771] (Ferreira 1906:168). Taxonomic and distributional notes: Two possible specimens are noted as being the type (see above). In the BMNH registers both (under their 1946 reregistration numbers) are listed as being a type of C. smythii or G. silurophaga (or Grayia smithii or G. silurophaga). Grayia I2"E 16.°E 2«°E Map 347. Distribution of Grayia ornata in Angola. 386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II silurophaga Gunther, 1858 is a junior subjec¬ tive synonym of G. smythii (Leach, 1818). According to Boulenger (1894), Gunther prob¬ ably inadvertently included the type of Leach’s species in the type series of his own new species. Although Wallach et al. (2014) consid¬ ered Boulenger’s (1894) comments as consti¬ tuting a lectotype designation, it seems more probable that only a single specimen was present from the collection made by Cranch. However, it is impossible to determine this from the description itself The correct version of the specific epithet of this species has been questioned. Although many authors have used G. smithii, the description employs Smythii. Although there is compelling evidence from the original publication (several other new species of animals are named with the patronym Smithii) that the species was probably named after Professor Christen Smith, who accompanied the expedition (see Laurent 1956), Wallach et al. (2014) are correct that Article 32.5 of the International Code of Zoological Nomenclature prohibits an emendation under the present circumstances. Map 348. Distribution of Grayia smythii in Angola. Grayia tholloni Mocquard, 1897 Thollon’s African Water Snake Grayia Tholloni Mocquard 1897b: 11. Syntypes: MNHN [2 specimens] specimen numbers unknown, and Musee de Toulouse [2 speeimens], speeimen numbers unknown (eolleetor F.-R. Thollon). Type loeality: “Congo frangais ... Brazaville” (Moequard 1897:11), [= Brazaville], Congo. Grayia tholloni: Laurent (1954a:44), Loveridge (1957:267), Broadley (1983:22), Broadley et al. (2003:182), Spawls et al. (2004:338), Broadley et al. h-e i&°e 2o°e 24"e (2003:182), Chippaux (2006:95), Trape and Mane (2006:105), Chirio and LeBreton (2007:438), Wallaeh et al. (2014:311). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Senegal through Ethiopia and southern Sudan south through the Great Lakes rigion to western Tanzania, northern Zambia and northeastern Angola. Ocurrences in Angola (Map 349): The species occurs in the extreme northeast of the country. Lunda Norte: “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1954a:44). Taxonomic and distributional notes: The presumed MNHN syntypes are not listed in the Paris Museum online collections database. Map 349. Distribution of Grayia tholloni in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 387 Genus Hapsidophrys Fischer, 1856 Hapsidophrys smaragdinus (Schlegel, 1837) Emerald Snake Dendrophis smaragdina Schlegel 1837:237. Leetotype: RMNH 909 (eolleetor D.F. Esehrieht), designated by Hoogmoed in Hughes and Barry (1969:1017). Type loeality: “Cote d’or” [= Ghana], given as “Cote de Guinee i.e., eoastal Ghana’' fide Hughes and Barry (1969:1017) by leetotype designation. Hapsidophrys smaragdina: Peters (1877a:615), Boeage (1887a: 186, 1895a:96), Spawls et al. (2004:364), Chippaux (2006:125), Chirio and LeBreton (2007:446). Gastropyxis smaragdina: Boulenger (1915:206), Thys van den Audenaerde (1966:33), Laurent (1954a:49, 1964a: 108), Hellmieh (1957b:65). Hapsidophrys smaragdinus: Wallaeh et al. (2014:314). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from western and central Africa, from Gambia to Angola and Uganda and Tanzania, with an isolated population on Sao Tome Island. Ocurrences in Angola (Map 350): The species occurs in the northern Angola, includ¬ ing the Cabinda enclave. Cabinda: “Chincho- xo” [-5.10000, 12.10000] (Peters 1877a:615; Boeage 1887a; 186, 1895a;96); “Cabinda” [-5.55000, 12.18333] (Boeage 1895a:96). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a;49, 1964a; 108; Thys van den Audenaerde 1966:33); “Sombo riv. Tchium- bue” [-8.68333, 20.95000] (Laurent 1954a:49). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:65); “N’dalla Tando” [-9.30000, 14.91667] (Ferrreira 1903:10); “Cazengo” [-9.33333, 14.76667] (Boeage 1895a:96). Taxonomic and distributional notes: The description notes numerous specimens vailable to Schlegel (1837), however, only two specimens collected by Esehrieht, including Map 350. Distribution of Hapsidophrys smaragdinus in Angola. the leetotype, were associated with an explicit locality. Genus Hormonotus Hallowell, 1857 Hormonotus modestus (Dumeril, Bibron and Dumeril, 1854) Uganda House Snake Lamprophis modestus Dumeril, Bibron and Dumeril 1854:429. Holotype: RMNH 304 (eolleetor H.S. Pel). Type loeality: “eote de Guinee,” restrieted to “Daboerom, Ghana” fide Hughes and Barry (1969:1015). Hormonotus modestus: Boulenger (1896:617, 1915:204), Parker (1936:125), Loveridge (1957:253), Spwals et al. (2004:329), Chirio and LeBreton (2007:450), Wallaeh et al. (2014:331). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is widespread in tropical West and Equatorial Africa from Guinea to Uganda and southwestern Kenya and into Angola. Ocurrences in Angola (Map 351): The species is only known from northwestern Angola. Cabinda: “Mouth of the Loango” [-5.15000, 12.16667] (Boulenger 1896:617). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:125). 388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Taxonomic and distributional notes: None. Il'E 16°E 20°E Map 351. Distribution of Hormonotus modestus in Angola. Genus Lycodonomorphus Lichtenstein, 1823 Lycodonomorphus subtaeniatus Laurent, 1954 Eastern Congo White-Bellied Water Snake Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Democratic Republic of Congo and northern Angola. Its presence in the Peo¬ ple’s Republic of Congo is likely but uncon¬ firmed. Ocurrences in Angola (Map 352): The species is known from northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a: 3 8). Taxonomic and distributional notes: Lycodonomorphus subtaeniatus upembae Lau¬ rent, 1954 has been recognized as specifically distinct and has been transferred to Boaedon based on phylogenetic information by Green- baum et al. (2015). Lycodonomorphus subtaeniatus subtaeniatus Laurent 1954a:38, figs. 1-4. Holotype: MRAC 14864 (former¬ ly RGMC 14864) (eolleetor N’Kele). Type loeality: “Keseki, pres de Kwamouth” [= Keseki, near Kwamouth], Demoeratie Republie of Congo. Lyeodonomorphus subtaeniatus: Broadley and Cot- i2'>e ^o^e >4«e term (2004:48), Wallaeh et al. (2014:399). Map 352. Distribution of Lycodonomorphus subtaenia¬ tus in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 389 Genus Mopaneveldophis Figueroa, McKelvy, Grismer, Bell and Lailvaux, 2016 Mopaneveldophis zebrinus (Broadley and Schatti, “1997” 1999) Mopaneveld Snake Coluber zebrinus Broadley and Schatti “1997” 1999:172, figs. 2-3, 5. Holotype: SMWN 8046 (collectors M. Lindeque and M. Griffin). Type locality: “near the Cunene River, at Ruacana, western Owamboland, Namibia.” Coluber zebrinus: Bauer et al. (2001:79). Hemerophis zebrinus: Wallach et al. (2014:319). Mopaneveldophis zebrinus: Figueroa et al. (2016:23). Global conservation status (lUCN): Not Evaluated. Global distribution; The species is known from northwestern Namibia. Ocurrences in Angola: The species is expected to occur in southern areas of Namibe and Cunene provinces, near the Cunene River. Taxonomic and distributional notes; Although the date on the paper in which this species was described is 1997, there was a significant delay in publication and the actual date is 1999 (fide B. Schatti, pers. comm., Oct. 2016). Bauer et al. (2001) suggested that this species would likely eventually be added to the herpetofauna of Angola, based on the similarities of its typical habitat with the adjacent southwestern regions of Angola. Wallach et al. (2014) were the first to allocate this species to Hemerophis, but provided no justification for this. Subsequently Figueroa et al. (2016), based on molecular phylogenetic data, erected a new genus, Mopaneveldophis, to accom¬ modate the species. Genus Philothamnus Smith, 1840 Philothamnus angolensis Bocage, 1882 Angolan Green Snake Philothamnus angolensis Bocage 1882b:7. Syntypes: MBL T92.1456 and others (eolleetors J.A. d’Anehieta and J. Horta), destroyed fire 18 Mareh 1978. Type loeality: “Capangombe” and “Angola,” restrieted to “Capangombe,” Namibe Province, by Loveridge (1951:9). Ahaetulla irregularis: Gunther (1865a:480). Philothamnus irregularis: Peters (1881:149), Bocage (1882b:6, 1887b:205, 1895a:85, 1896a:112), Ferreira (1906:167). Chlorophis angolensis: Boulenger (1893:95, 1915:205), Frade (1963:252). Chlorophis irregularis: Boulenger (1893:96), Parker (1936:125), Monard (1937b:114, 121), Mertens (1938a:439), Bogert (1940:53). Philothamnus angolensis: Bocage (1897a:200), Hughes (1985:518), Broadley (1990:238), Branch (1998:94), Broadley and Cotterill (2004:51), Spawls et al. (2004:552), Chippaux (2006:135), Chirio and LeBreton (2007:500), Bates et al. (2014:416), Wallaeh et al. (2014:542). Chlorophis (Philothamnus) irregularis: Ferreira (1903:10). Philothamnus Guntheri: Ferreira (1906:168). Philothamnus irregularis var. angolensis: Ferreira (1900a:51, 1904:115). Chi. angolensis (Philothamnus angolensis): Monard (1937b: 114). Philothamnus irregularis irregularis: Loveridge (1951:9, 1957:261), Hellmich (1957b:64), FitzSimons (1962:144), Managas (1973:191). Global conservation status (lUCN): Not Evaluated. Global distribution; Cameroon east to Sudan and South Sudan, south to northern Namibia in the west and northeastern KwaZulu-Natal, South Africa in the east. Ocurrences in Angola (Map 353): The species is known chiefly from western Angola, how¬ ever, its distribution may comprise most of country, exclusive of the desert regions of far south¬ western Angola. Zaire: “St. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1895a;85). Uige: 390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II “Fazenda Otilia, Encoge” [-7.55000, 15.03333] (Managas 1973:191). Lunda Norte; “Dundo” [-7.36667, 20.83333] (Laurent I950a:8, I954a:47, 1964a: 106; Thys van den Audenaer- de 1966:32); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent I954a:47). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage I895a:85; Ferreira I900a;5I; Monard 1937b: 1 14). Bengo: “Cacolo ao Rio Bengo” (Ferreira I900a;5I). Malanje: “Duque de Bra- gan9a” [-9.10000, 15.95000] (Bocage I866a;49, I882b:5, I895a;85; Monard 1937b: 1 14); “Pungo-Andongo” [-9.66667, 15.58333] (Gunther I865a:480; Bocage I882b:5, I895a:85; Ferreira 1906:167; Monard 1937b: 1 14). Mexico: “Lago Dilolo” [-11.50000, 22.01667] (Managas 1973:191). Huambo: “Bela- Vista” [-12.36667, 16.20000] Map 353. Distribution of Philothamnus angolensis in Angola. (Hellmich 1957b:64); “Mt. Moco” [-12.41667, 15.18333] (Parker 1936:125). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:64); “Ambaca” [-9.26667, 15.18333] (Ferreira 1900a:51); “Cazengo” [-9.33333, 14.76667] (Ferreira 1900a;51); “Cambondo” [-9.15963, 14.65771] (Boulenger 1893:356; Ferreira 1906:167, Monard 1937b:114, Mana9as 1973:193); “N’dalla Tando” [-9.30000, 14.91667] (Ferreira 1903:10). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:125); “Chingo” [-11.20000, 13.85000] (Ferreira 1904:115; Monard 1937b:114). Bie: “Silva Porto” [-12.38333, 16.95000] (Mana9as 1973:191). Benguela: “Quissange” [-12.43333, 14.05000] (Bocage 1895a:85; Monard 1937b:114); “Quindumbo” [-12.46667, 14.93333] (Bocage 1895a:85; Monard 1937b:114); “Alto Cubal” [-13.03333, 14.25000] (Mertens 1938a:439; Hellmich 1957b:64); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:112). Huila: “Caconda” [-13.73333, 15.06667] (Boulenger 1893:96; Bocage 1895a:85; Monard 1937b:114); “Capelongo” [-14.91667, 15.08333] (Bogert 1940:53); “Kuvangu/Vila-da-Ponte” [-14.46667, 16.30000] (Monard 1937b: 121); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:85, 1887b:205, 1895a:85; Monard 1937b:114). Namibe: “Capangombe” [-15.10000, 13.15000] (Bocage 1882b:6, 7, 1897a:200; Loveridge 1951:9, 1957:261; Broadley 1990:238; Chippaux 2006:135; Chirio and LeBreton 2007:500; Wallach et al. 2014:52); “Mossamedes” [-15.20000, 12.15000] (Bocage 1887b:205, 1895a:85). Cunene: “Ruis- seauMbale” [-15.16667, 16.75000] (Monard 1937b:121); “Mupa” [-16.18333, 15.75000] (Monard 1937b:121). Undetermined Locality: “Cuango” (Peters 1881:149) (Malanje Province impossible to georeference; See History Section for more detailed information). Taxonomic and distributional notes: The majority of Angolan literature records for Philothamnus irregularis (Leach, 1819) are probably referable to P. angolensis or P. hoplogaster, two species with which it has commonly been confused (Hughes 1985). Philothamnus irregularis is essentially a West Africa form ranging from Senegal to northern Cameroon and east to south¬ western Chad. Its distribution south of Cameroon and in East Africa is a matter of contention (Loveridge 1951; Rodel and Schmitz 2010). Current literature, including Hughes (1985), Chippaux (2006), Trape and Mane (2006) and Wallach et al. (2014), have not included Angola in the distri¬ bution of P. irregularis, although Wallach et al. (2014) included southern and western provinces of the Democratic Republic of Congo in its range. Based largely on the maps of Hughes (1985), we MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 391 have tentatively alloeated literature reeords of P. irregularis to either P. angolensis or P hoplo- gaster, although all of these should be verified and some may refer to other eongeners. Hughes (1985) plotted numerous reeords for P angolensis, ehiefly in the west of the eountry, based on museum specimens and one literature record from Cuando Cubango near the Namibian border, unfortunately without providing precise localities. Philothamnus carinatus (Andersson, 1901) Thirteen-Scaled Green Snake Chlorophis Andersson 1901:9. Syntypes: NHR 1972 (collector Knoppel), NRH 1973-76 (collectors K.W. Knutson and G. Valdau) [5 specimens]. Type locality: “Mapanja, Cameroon” and “Cameroon” restricted to “Mapanja” [Sud-Ouest Province, Cameroon] yitie Mertens (1964:227). Chlorophis heterodermus carinatus: Thys van den Audenaerde (1966:32). Philothamnus carinatus: Broadley and Cotterill (2004:51), Spawls et al. (2004:354), Chippaux (2006:135), Chirio and LeBreton (2007:504), Wallach et al. (2014:543). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from from Guinea to eastern Kenya and Tanzania, south to the former Katanga and northeastern Angola. Ocurrences in Angola (Map 354): The species has only been recorded from Dundo in northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Thys van den Aude¬ naerde 1966:32). Taxonomic and distributional notes: The Dundo record from Thys van den Audenaerde (1966) appears to be the only explicitly named locality cited for the species in Angola, how ever Wallach et al. (2014) state that the species is also distributed in the northwestern regions of Angola in Cabinda Province, perhaps based on the map of Hughes (1985). Philothamnus dorsalis (Bocage, 1866) Leptophis dorsalis Bocage 1866a:48, 1866b:69. Lecotype: MBL specimen number unknown (collector J.A. d’Anchieta), designated by Bocage (1882b: 10), destroyed by fire 18 March 1978. Type locality: “Cabin¬ da” and “Loanda” (Bocage 1866a:48; 1897a:200), but given as “Duque de Bragan^a” and “Molembo” by Bocage (1866b:70), restricted to “Molembo (cote de Loango),” Cabinda Province, through designation of lectotype. Leptophis dorsalis: Bocage (1867b:226). Ahaetulla dorsalis: Gunther (1876b:679). Philothamnus dorsalis: Bocage (1882b: 10, 1887a: 185, 1895a:92, 1897a:200), Boulenger (1893:101, 1915:206), Ferreira (1897b:244), Hughes (1985:520), Chippaux (2006:133), Chirio and LeBreton (2007:506), Wallach et al. (2014:544). Philothamnus semivariegatus dorsalis: Bogert (1940:56), Loveridge (1933:238, 1951:11, 1957:262), Hellmich (1957b:65). Global conservation status (lUCN): Not Evaluated. Il-E 16.°E 20°E I4®E Map 354. Distribution of Philothamnus carinatus in Angola. Striped Green Snake 392 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Map 355. Distribution of Philothamnus dorsalis in Angola. Global distribution: The species is known from Angola, Cameroon, Congo, Democratic Republic of Congo and Gabon. Ocurrences in Angola (Map 355): The species occurs in the central-north western Angola. Cabinda: “Molembo” [-5.33333, 12.20000] (Bocage 1866b;69, 1882b:10, 1895a:92; Loveridge 1933:238, 1951:11, 1957:262; Chirio and LeBreton 2007:506; Wal- lach et al. 2014:544); “Cabinda” [-5.55000, 12.18333] (Bocage 1866a:48, 1895a:92, 1897a:200). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1881a:185, 1895a:92). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1866a:48, 1867b:226, 1882b:10, 1895a:92, 1897a:200; 1957:262). Bengo: “Rio Dande” [-8.46667, 13.38333] (Bocage 1882b: 10, 1895a:92). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866b:69; Loveridge 1933:238); “Pungo-Andongo” [-9.66667, 15.58333] (Bocage 1882b:10; 1895a:92). Kwanza Norte: “Dondo” [-9.66667, 14.41667] (Hellmich 1957b:65). Kwanza Sul: “Libolo” [-9.98333, 14.90000] (Hellmich 1957b:65). Benguela: “Catumbella” [-12.43333, 13.55000] (Bocage 1867c:226, 1882b:10, 1895a:92); “Benguella” [-12.58333, 13.41667] (Bocage 1867b:226, 1882b:10, 1895a:92; Boulenger 1893:101); “Dombe” [-12.95000, 13.10000] (Bocage 1867c:226; Hughes 1985:520). Huila: “Caconda” [-13.73333, 15.06667] (Ferreira 1897b:244; Hughes 1985:520). Undetermined Locality: “no locality” (Ferreira 1897b:244); “Carangigo” [-13.03333, 13.73333] (Boulenger 1893:101); “Benguella to Ogouoe” (Boulenger 1893:101). Taxonomic and distributional notes: The species was described as new by Bocage in two different papers (1866a, b), both of which appeared in the November 1866 issue of the same jour¬ nal. Unlike many of the descriptions in 1866a, however, that of Leptophis dorsalis is a nomen nudum as there is no description, only a statement that it is similar to Leptophis Chenoni [= Philothamnus irregularis]. In the first paper, two specimens from “Cabinda”, (collected by Anchieta) and “Loanda” (collected by Bayao) were noted. In the description that made the name available, Bocage (1866b) cited the “Cabinda” specimen as being from “Molembo” and that col¬ lected by Bayao from “Duque de Bragan^a.” Bocage (1882b) refered to the Molembo specimen as the type, and mentioned numerous other specimens, but none from Duque de Braganga. In Bocage’s (1897a) type catalogue he again referred only to “Cabinda” and “Loanda.” The locality “Duque de Bragan9a” may have been an error, as Bocage never cited it in his subsequent works. However, ZMB 6467 is marked in the Berlin catalogue as a type specimen from “Angola” obtained from Bocage, and it is also possible that the “Duque de Bragan^a” specimen had been traded away before Bocage’s (1882b) paper on Philothamnus. Bogert (1940), Loveridge (1933, 1951, 1957), and Hellmich (1957b) considered P. dorsalis as a subspecies of P. semivariegatus, however, according to Hughes (1985) this is untenable on geographic grounds, since the two co-occur in the lower Congo. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 393 Philothamnus heterodermus (Hallowell, 1857 Emerald Green Snake Chlorophis heterodermus Hallowell 1857:54. Holotype: ANSP 5219. Type loeality: “Gaboon eountry” [= Gabon]. Philothamnus heterodermus: Bocage (1895a:89), Hughes (1985:514), Chippaux (2006:130), Chirio and LeBreton (2007:508), Cerlaco et al. (2014b:671), Wallach et al. (2014:544). Chlorophis heterodermus: Parker (1936:125). Philothamnus heterodermus heterodermus: Loveridge (1951:6), Hellmieh (1957b:64). Hughes (1985) regarded a reeord from Karibib in eentral Namibia as valid, however, we rejeet this, as only P. angolensis Boeage, 1882 R ornatus Boeage, 1872, andR semivariegatus (Smith, 1840) are now believed to oeeur in Namibia. Map 356. Distribution of Philothamnus heterodermus in Angola. Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from Guinea-Bissau to Uganda, Burundi, Rwana- da and Tanzania, and south through Gabon, Congo, the Demoeratie Republie of Congo, and Angola. Ocurrences in Angola (Map 356): The species occurs in northwestern Angola, includ¬ ing the Cabinda encalve. Malanje: “Capanda” [-9.72841, 15.34585] (Ceriaco et al. 2014b:671). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmieh 1957b:64; Hughes 1985:514). Kwanza Sul: “Congulu” [-10.86667, 14.28333] (Parker 1936:125; Hughes 1985:514). Taxonomic and distributional notes: Philothamnus heterolepidotus (Gunther, 1863) Slender Green Snake Ahaetulla heterolepidota Gunther 1863:286. Holotype: BMNH 1946.1.10.21 (eolleetor unknown). Type loea¬ lity: “Affiea,” restrieted to “West Afriea” by Boulenger (1894:96). Lepthophis heterolepidota: Boeage (1866a:48, 1866b:69). Philothamnus heterolepidotus: Boeage (1879b:95, 1882b:8-9, 1887a:185, 1895a:88), Loveridge (1951:10, 1957:261), Laurent (1954a:48), Hughes (1985:525), Broadley et al. (2003:195), Spawls et al. (2004:356), Chippaux (2006:131), Chirio and LeBreton (2007:510), Wallaeh et al. (2014:544). Chlorophis heterolepidotus: Boulenger (1893:95, 1905:112, 1915:205), Monard (1937b:120), Laurent (1950a:8, 1964a: 105), Thys van den Audenaerde (1966:33). Philothamnus irregularis irregularis: Managas (1873:191). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Sierra Leone to Sudan, Kenya and western Ethiopia south to the Democratic Republic of Congo, Angola, and Zambia. Ocurrences in Angola (Map 357): The species occurs in the entire country with exception of the arid desert areas. Cabinda: “Cabinda” [-5.55000, 12.18333] (Bocage 1895a: 88). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a: 185, 1895a:88). Luanda: “Loanda” [-8.83333, 13.26667] (Bocage 1887a:185). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a:8, 1954a:48); “Dundo, R. Mussungue” [-7.41667, 20.83333] (Thys van den Audenaerde 394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II 1966:33); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1 950a; 8); “Cassange” [-9.58333, 17.86667] (Bocage I895a:88; Monard 1937b; 1 14). Mexico: “environs du lac Calundo, near Cameia” [-11.80000, 20.86667] (Laurent 1964a; 105); “Naricumbi, Reserve de chasse de Cameia” [-11.83333, 21.00000] (Laurent 1964a; 105); “Margens do rio Lungue, Bingo-Moxico” [-12.45000, 20.05000] (Mana- gas 1973:191). Malanje: “Duque de Bragan- 9a” [-9.10000, 15.95000] (Bocage 1866a:48, 1866b:69, 1882b:8, 1895a:88; Boulenger 1905:112; Monard 1937b:114). Kwanza Norte: “Dondo” [-9.68333, 14.43333] (Bocage 1882a:8, 1895a:88; Monard 1937b:114). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b: 114, 120). Benguela: “Quibu- la” [-12.28333, 14.68333] (Bocage 1895a:88; Monard 1937b: 114). Huila: “Caconda” Map 357. Distribution of Philothamnus heterolepidotus in Angola. [-13.73333, 15.06667] (Bocage 1882b;8, 1895a;88; Monard 1937b:114). Cuando Cubango: “Kutatu” [-14.36667, 16.48333] (Monard 1937b: 114, 120). Undetermined Locality: “Cuango” (Bocage 1887a: 185) (Malanje Province impossible to georeference; See History Section for more detailed information). Taxonomic and distributional notes; Wallach et al. (2014) noted that the BMNH catalogue lists BMNH 1946.1.10.25 as a type, but the specimen under this number is actually listed as Ahaetulla gracillima Gunther, 1888, a junior synonym of P. heterolepidotus (Gunther, 1863). According to Hughes (1985) Philothamnus heterolepidotus is difficult to distinguish from P. irreg¬ ularis (Leach, 1819). Loveridge (1951) also noted this and remarked that this species was, for some time, misidentified as P. natalensis (Smith, 1848) or P. ornatus (Bocage, 1862) [= Philothamnus]. Monard’s (1937b) specimens from “Bimbi,” Huambo Province and “Kutatu” Cuando Cubango Province, could possibly belong to ornatus (Loveridge 1951). Philothamnus hoplogaster (Gunther, 1863) Southeastern Green Snake Ahaetulla hoplogaster Gunther 1863:286. Syntypes: BMNH 1946.1.6.1 (formerly BMNH 62.3.14.6), 1946.1.6.6 and 1946.1.6.14 (formerly BMNH 62.3.14.12) (eolleetor T. Ayres) [BMNH 1946.1.6.6 is list¬ ed in the BMNH register as having the loeality “Off Nataf’ and no eolleetor is listed]. Type loeality: “Port Natal” (Gunther 1863:286), [= Durban, eastern KwaZulu-Natal], South Affiea. Ahaetulla irregularis [part]: Gunther (1865a:480). Leptophis Chenoni: Boeage (1866a:49). Philothamnus irregularis: Peters (1877a:615), Boeage (1882b:6, 1895a:85), Ferreira (1906:167). Philothamnus hoplogaster: Boeage (1887a: 186; 1895a:86), Branch (1998:94), Hughes (1985:517), Spawls et al. (2004), Bates et al. (2014:417), Wallach et al. (2014:545), Conradie et al. (2016:19). Chlorophis irregularis: Parker (1936:125), Monard (19376:114), Themido (1941:10), Laurent (1950a:8). Chlorophis sp. (hoplogasterl): Monard (19376:114, 120). Philothamnus irregularis irregularis: Hellmich (1957b:64), Laurent (1954a:47). Chlorophis hoplogaster: Frade (1963:253), Laurent (1964a: 103), Thys van den Audenaerde (1966:32). Chlorophis irregularis shiranus: Thys van den Audenaerde (1966:32), Laurent (1964a: 103). Global conservation status (lUCN): Not Evaluated. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 395 Global distribution: The species is widespread in sub-Saharan Africa from Cameroon to South Sudan and south to Angola in the west and western South Africa in the east. Ocurrences in Angola (Map 358): The species occurs appears to be widespread, par¬ ticularly in the central and northern regions of Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Bocage 1882b:6, 1895a:85); “Cabinda” [-5.55000, 12.18333] (Bocage 1895a:85). Zaire: “S. Salvador do Congo” [-6.26667, 14.23333] (Bocage 1887a: 186; 1895a:86). Lunda Norte: “Camis- sombo” [-8.15000, 20.65000] (Laurent 1964a: 106). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Laurent 1964a:106); “sources du Cuilo, Alto Chicapa” [-10.91667, 19.33333] (Laurent 1964a: 103); “Mutianvo” [-11.45000, 19.33333] (Themido 1941:10). Malanje: “Malanje” [-9.55000, 16.35000] (Bocage 1882b:6, 1895a:85; Monard 1937b:114). Huambo: “Bimbi” [-11.81667, 15.83333] (Monard 1937b: 120). Cuando Cubango: “Cuito River system” (photographic 2016:19). ll^E 1&°E 20°E 24'>E Map 358. Distribution of Philothamnus hoplogaster in Angola. record) [-14.14889, 18.81000] (Conradie et al. Taxonomic and distributional notes: The majority of Angolan literature records for Philothamnus irregularis (Leach, 1819) are probably referable to P. angolensis ov P. hoplogaster, two species with which it has commonly been confused (Hughes 1985). Philothamnus irregularis is essentially a West Africa form ranging from Senegal to northern Cameroon and east to south¬ western Chad. Current literature, including Hughes (1985), Chippaux (2006), Trape and Mane (2006) and Wallach et al. (2014), have not included Angola in the distribution of P. irregularis. Based largely on the maps of Hughes (1985), we have tentatively allocated literature records of P. irregularis to either P. angolensis or P. hoplogaster, although all of these should be verified and some may refer to other congeners. Hughes (1985) provided a distribution map for Philothamnus hoplogaster with literature records, museum records and previouisly unpiblished D. G. Broadley records. Hughes (1985) provisionally accepted a record from Damaraland, northern Namibia as valid, however, we follow FitzSimons (1962) and Broadley (1983, 1990) in rejecting this. Only P. angolensis Bocage, 1882, P ornatus Bocage, 1872, andi? semivariegatus (Smith, 1840) are con¬ firmed to occur in Namibia. Philothamnus nitidus loveridgei (Laurent, 1960) Loveridge’s Green Bush Snake Ahaetulla nitida Gunther 1863:286. Syntypes: BMNH 1946.1.10.24 (collector Col. E. Sabine) and BMNH 1946.1.6.2 (formerly BMNH 71.4.21.5) (collector unknown). Type locality: “Demerara” [Guyana], in error, corrected to “Lagos” [Nigeria] by Boulenger (1894:101] Wallach et al. (2014:546) and “not known”. Philothamnus nitidus loveridgei Laurent (1960:40). Holotype: MRAC (formerly RGMC) 18423 (collector R. F. Laurent). Type locality: “Itula, 650 m., Territoire de Shabunda, Kivu” [South Kivu Province, Demo¬ cratic Republic of Congo]. Chlorophis nitidus loveridgei: Laurent (1964a: 106), Thys van den Audenaerde (1966:33). 396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Philothamnus nitidus: Spwals et al. (2004:359), Chippaux (2006:132), Chirio and LeBreton (2007:516), Wallach et al. (2014:546). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from West and Central Africa, form Sierra Leone to the Democratic Republic of Congo and adjacent eastern Angola, east to southwestern Kenya and southwestern Tanzania. The subspecies P. n. loveridgei occupies the eastern portion of the range, from the eastern Congo eastwards, including all of the Democratic Republic of Congo and north¬ eastern Angola. Ocurrences in Angola (Map 359): The species has been recorded only from “Dundo,” Lunda Norte Province. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1964a: 106; Thys van den Audenaerde 1966:33). Taxonomic and distributional notes: Hughes (1985) noted that BMNH 1946.1.10.24 is one of the types of P. nitidus, the individual supposedly from Demerara, but that it is actu¬ ally referable to the species from Sao Tome, P. thomensis Bocage, 1882. Both BMNH 1946.1.10.28 and 1946.1.6.2, regarded as the syntypes of P. nitidus by Wallach et al. (2014) are indicated as types of Ahaetulla lagoensis in the BMNH register, with the explicit locality “Lagos.” This conflicts with the statement by Wallach et al. (2014) that BMNH 1946.1.10.28 is listed as a type of P nitidus in the BMNH register. It is clear that some confusion still surrounds the types of P. nitidus and similar forms in the BMNH collection. The majority of recent authors (e.g., Spwals et al. 2004; Chippaux 2006; Chirio and LeBreton 2007) have not included Angola in the distribution of Philothamnus nitidus. However, Laurent (1964a), and later Thys van den Audenaerde (1966) reported specimens identi¬ fied as Philothamnus nitidus loveridgei Laurent, 1960 from Dundo, and this locality was accepted and plotted by Hughes (1985). The nominotypical subspecies has not been reported from Angola, but according to Hughes (1985) it occurs very near to Cabinda. It is likely that P. n. loveridgei also occurs in Zaire Province, based on the proximity of records from the southern bank of the lower Congo River in the Democratic Republic of Congo. Map 359. Distribution loveridgei in Angola. of Philothamnus nitidus Philothamnus ornatus Bocage, 1872 Ornate Green Snake Philothamnus ornatus Bocage 1872:80. Syntypes: MBL [2 speeimens from Huila] (eolleetor J.A. d’Anehieta) and MBL [from Caeheu] (eolleetor C. Hopffer), all destroyed by fire 18 Mareh 1978. Type loeality: “Huil- la,” Angola and “Caeheu, sur la eote de Guinee, Afrique oeeidentale” [= Caeheu], Guinea-Bissau, restriet- ed to “Huilla, Angola,” Huila Provinee by Bogert (1940:51). Philothamnus ornatus: Boeage (1882b: 16, 1895a:93, 1897a:200), Hellmieh (1957b:65), Broadley (1990:236), Braneh (1998:94), Spawls et al. (2004:359), Chippaux (2006:134), Chirio and LeBreton (2007:518), Wallaeh et al. (2014:546), Conradie et al. (2016:19). Chlorophis ornatus: Boulenger (1893:93, 1905:112), Monard (1937b:119), Bogert (1940:51), Frade (1963:252). Philothamnus irregularis ornatus: Loveridge (1958:84, 1951:8). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 397 Global conservation status (lUCN): Not Evaluated. Global distribution; The species is distributed from eastern Zimbabwe, Malawi, and eastern Zambia, through northern Botswana and the Caprivi Strip of Namibia to western Angola. Disjunct records exist from Cameroon and the far northeastern Democratic Republic of Congo. Ocurrences in Angola (Map 360): The species occurs from northwestern to southern Angola. Huambo: “Bela- Vista” [-12.36667, 16.20000] (Hellmich 1957b;65); “Huambo” [-12.83333, 15.66667] (Bogert 1940:51). Huila: “Caconda” [-13.73333, 15.06667] (Bocage 1882b: 16, 1895a:93; Monard 19376:114; Loveridge 1958:84); “Kalukembe” [-13.78333, 14.68333] (Monard 19376:114, 119); “Huilla” [-15.05000, 13.55000] (Bocage 1872:80, 1882b:16, 1895a:93, 1897a:200; Monard 19376:114; Loveridge 1951:8, 1958:84; Broadley 1990:236; Chippaux 2006: 134; Chirio and LeBreton 2007:518; Wal- lach et al. 2014:546). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 19376:114, 119; Loveridge 1958:84); “Cunene” (Bocage 18826:17, 1895a;93; Monard 19376:114; Loveridge 1958:84). Cuando Cubango: “Kutatu” [-14.36667, 16.48333] (Monard 19376:114, 119; Loveridge 1958:84); “Cuito basin (24)” [-14.06622, 18.46722] (Conradie et al. 2016:8-9, 19). Undetermined Locality: “Between Benguela and Bihe” (Boulenger 1905:112; Monard 19376:114; Loveridge 1958:84). Taxonomic and distributional notes: Bocage (1872) gaive “Cacheu” as the type locality of P. ornatus although both Bogert (1940) and Loveridge (1958) suspected that this locality corre¬ sponded to an error in labeling and considered “Huilla” as the type locality. Loveridge (1951) treat¬ ed P. ornatus as a race of Philothamnus irregularis (Leach, 1819), whereas Hughes (1985) considered it a full species. Loveridge (1951) noted that Monard’s (19376) specimens of Chlorophis heterolepidotus (Gunther, 1863) from “Bimbi” and “Kutatu,” could possibly belong to this species (Loveridge 1958). Map 360. Distribution of Philothamnus ornatus Angola. Philothamnus semivariegatus (Smith, 1840) Spotted Bush Snake Dendrophis {Philothamnus) semivariegata Smith 1840: pis. 59-60, first of two unnumbered pages of aeeom- panying text. Leetotype: NMSZ 1859.13.1257a, formerly RSM (eolleetor A. Smith), designated by Bogert (1940:56). Type locality: “Bushman Flat” and “the country beyond Kurrichaine,” South Africa,” restrict¬ ed to “Bushman Flat” by designation of leetotype. Philothamnus Smithir. Bocage 1882b: 12, fig. 5. Syntypes: MBL specimen numbers unknown (collectors Fer¬ reira Borges [Bissau], d’Anchita [Catumbella, Huilla, Humbe, Capangombe, Maconjo]). Type locality: “Bissau (Guinee du Cap- Vert) et . . . plusieurs localites d’ Angola” (“Catumbella [Benguela Province] . . . Huilla [Huilla Province] . . . Humbe [Cunene Province] . . . Capangombe [Namibe Province] . . . Maconjo [Namibe Province]”). Ahaetulla Bocagii Gunther 1888:326. Holotype: BMNH specimen number unknown (collector Lieutenant V.L. Cameron). Type locality: “Angola.” Philothamnus semivariegatus: Boulenger (1893:99), Bocage (1895a:90, 1896a:112), Ferreira (18976:244), Monard (19376:122), Schmidt (1933:13), Thys van den Audenaerde (1966:33), Hughes (1985:522), 398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Branch (1998:93), Chippaux (2006:129), Trape and Mane (2006:140), Chirio and LeBreton (2007:520), Wallach et al. (2014:547). Philothamnus semivariegatus semivariegatus: Mertens (1938:439), Laurent (1950a:8, 1954a:48); Loveridge (1951:11, 1957:262), Haaeke (1985:7). Philothamnus sp. (alf. semivariegatus): Thys van den Audenaerde (1966:33). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is extremely widely distributed in sub-Saharan Africa from Senegal and Niger to Sudan and Ethiopia, and south to South Africa. Ocurrences in Angola (Map 361): The species has been reported from across the entire country except parts of the northwest and the desert regions of far southwestern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a;8, 1954a;48 1964a: 107; Thys van den Audenaerde 1966:33). Moxico: “envi¬ rons du lac Calundo” [-11.80000, 20.86667] (Laurent 1964a: 107). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:13). Benguella: “Catumbella” [-12.43333, 13.55000] (Bocage 1882b:12, 1895a:90); “Cubal” [-13.03333, 14.25000] (Mertens 1938:439); “Hanha” [-13.30000, 14.20000] (Bocage 1895a:90). Huilla: “Rio Cuce” [-13.51667, 15.20000] (Ferreira 1897b;244); “Quillengues” [-14.06667, 15.08333] (Bocage 1 895a:90); “Kuvangu/Vila-da-Ponte” [-14.46667, 16.30000] (Monard 1937b:122); “Huilla” [-15.05000, 13.55000] (Bocage 1882b:12, 1895a;90). Namibe: “Maconjo” [-15.01667, 13.20000] (Bocage 1882b:12); “Capangombe” [-15.10000, 13.15000] (Bocage 1895a:90; Haaeke 1985:8). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937b;122); “Humbe” [-16.68333, 14.90000] (Bocage 1882b:12, 1895a:90; Boulenger 1893:99). Cuando Cubango: “Kutatu” [-14.36667, 16.48333] (Monard 1937b:122); “Kakindo” [-15.45000, 17.05000] (Monard 1937b: 122). Taxonomic and distributional notes; Bocage (1882b) mentioned specimens of Philothamnus Smithii from “Huilla”, “Humbe”, “Catumbella” and “Maconjo” and noted that the individuals from the last two localities resemble Philothamnus variegata (Smith, 1840) [= Philothamnus semivariegatus (Smith, 1840)]. However, due the impossibility of the comparison with variegata specimens at the time there was no confirmation of their specific identity. Later, Gunther (1888) described a new species as Ahaetulla Bocagii based on a specimen from Angola without a precise location, collected by Lieutenant Cameron. Map 361. Distribution of Philothamnus semivariegatus in Angola. Genus Rhamnophis Gunther, 1862 Rhamnophis aethiopissa Gunther, 1862 Large-Eyed Green Tree Snake Rhamnophis aethiopissa Gunther 1862:129, pi. 10. Holotype: BMNH 1946.1.4.99 (formerly BMNH 61.12.30.61) (don. Capt. R.H. Beddome). Type loeality: “West Afriea.” Rhamnophis ituriensis Sehmidt (1923:81, fig. 4). Holotype: AMNH R-12505 (eolleetor H. Lang and J.P. Chapin). Type loeality: “Niapu” (Sehmidt 1923:81-82), Demoeratie Republie of Congo. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 399 Rhamnophis aethiopissa: Boulenger (1915:206), Roux-Esteve (1965:65), Broadley and Howell (1991:66), Broadley and Wallach (2002:64), Spawls et al. (2004:366), Wallach et al. (2014:633). Rhamnophis aethiopissa aethiopissa: Hellmich (1957b:66). Rhamnophis aethiopissa ituriensis: Laurent (1950a:8, 1954a:50, 1964a: 108), Thys van den Audenaerde (1966:33). Tharsops aethiopissa: Chippaux (2006:109). Global conservation status (lUCN): Not Evaluated. Global distribution: The speeies is known from Guinea east to the Demoeratie Republic of Congo, Rwanda, Uganda, and western ,6-e 20“e 24“e the Kenya, and south to northern Angola and northwestern Zambia. Ocurrences in Angola (Map 362): The species occurs in the northern Angola including the Cabinda enclave. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:50, 1964a: 108); “sur un arbuste dans la foret des sources de la Mussungue (affluent de la Lua- chimo) pres de Dundo” [-7.41667, 20.83333] (Laurent 1950a: 8, Thys van den Audenaerde 1966:33); “Calonda, Camissombo” [-8.41667, 20.53333] (Laurent 1964a: 108). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:66). Taxonomic and distributional notes: Roux-Esteve (1965) placed the subspecies Rhamnophis aethiopissa ituriensis Schmidt, 1923 in the synonymy of the typical form. Map 362. Distribution of Rhamnophis aethoipissa in Angola. Genus Scaphiophis Peters, 1870 Scaphiophis albopunctatus Peters, 1870 African Shovel-Nosed Snake Seaphiophis albopunetatus Peters 1870:645, pi. 1, figs. 4, 4a-4e. Holotype: ZMB 6945 (eolleetor Jahn). Type loeality: “Keta (Guinea)” [= Keta, Ghana] 7/t/e Wallaeh et al. 2014:650]. Seaphiophis albopunetatus: Laurent (1950a:9), Broadley (1994:4), Chirio and LeBreton (2007:546), Wallaeh et al. (2014:650), Braneh and Conradie (2015:200). Global conservation status (lUCN): Not Evaluated. Global distribution; The species is from northern Zambia and southern Democratic Republic of Congo to Kenya and South Sudan in the east and Ghana in the west. Ocurrences in Angola (Map 363): The species occurs in the extreme northeast of the coun¬ try in Lunda Norte Province. Lunda Norte: “Capaia” [-7.33556, 20.21681] (Branch and Conradie 2015:200); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:9). Taxonomic and distributional notes: Peters’ (1870) type locality of Keta, Guinea has cau¬ sed a great deal of confusion. Loveridge (1936a) accepted the locality as being “Kita, Guinea” [= Kita, Mali]. Hughes and Barry (1969), Broadley (1994) and Wallach et al. (2014) all interpreted the locality as being Keta [= Kete], Ghana. However, there is indeed a Keta, Guinea as well, although it is small and unlikely to have been the source of Peters’ type. Further, as noted by Hughes and Barry (1969), the collector, Jahn, was based at the Bremen Mission in Keta, Ghana. Wallach et al. (2014) noted several occurrences of the species west of Ghana, the limit of the 400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II species according to Broadley (1994). These may well be the result of continuing confusion over the placement of Keta. Given the localities known from Lunda Norte, it seems likely that the species occurs in other northern regions of Angola. ii*n i6°t: 2i"'K Map 363. Distribution of Scaphiophis albopunctatus in Angola. Genus Telescopus Wagler, 1830 Telescopus finkeldeyi Haacke, 2013 Telescopus finkeldeyi Haacke 2013:281. Holotype: TM 53542 (collector J.A. van Rooyen). Type locality: “Rossing Uranium mine area, Swako- mund [sic] district (2214Db) Namibia.” Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from Angola and Namibia. Ocurrences in Angola (Map 364): The species occurs in southwestern Angola. Namibe: “5 km north Namibe” [-15.20000, 12.15000] (Haacke 2013:285). Taxonomic and distributional notes: Some earlier records of T. semiannulatus poly- stictus in Namibia actually refer to this recent¬ ly described species. Damara Tiger Snake 1I°E l(i®E 20°£ I4'*E Map 364. Distribution of Telescopus finkeldeyi in Angola. Telescopus semiannulatus semiannulatus Smith, 1849 Common Tiger Snake Telescopus semiannulatus Smith 1849: pi. 72, first of two unnumbered pages of accompanying text. Holotype: BMNH 1946.1.2.55 (formerly BMNH 1871.4.21.1) (collector A. Smith) fide Wallach et al. (2014). Type locality: Not stated, “South Africa” by inference (Loveridge 1957:270). Telescopus semiannulatus semiannulatus: Loveridge (1957:270), Hellmich (1957b:71), Thys van den Aude- naerde (1966:34). Telescopus semiannulatus: Wallach et al. (2014:698). MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 401 Global conservation status (lUCN): Not Evaluated. Global distribution: The species is wide¬ ly distributed from Angola and the southern Democratic Republic of Congo to Kenya and Tanzania and south through Namibia northern and eastern South Africa. Ocurrences in Angola (Map 365): The species occurs in northern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Thys van den Audenaerde 1966:34); “Mucoloje” [-7.50000, 20.91667] (Thys van den Aude¬ naerde 1966:34). Kwanza Sul: “Libolo-Luati” [-9.98333, 14.90000] (Hellmich 1957b:71). Taxonomic and distributional notes: FitzSimons (1937) stated that the type was not located in either the BMNH or RSM. Wallach et al. (2014) incorrectly stated that Loveridge (1957) had specified “Loanda, South Africa [= La Locanda, E Western Cape Prov., SW South Africa, 33°51’S, 22°3rE, elevation 350 m]” as the restricted type locality of T. semiannulatus. Loanda, is however, the locality listed in the BMNH register for BMNH 1871.4.21.1. The subspecies T. s. polystictus Mertens, 1954 occurs in western Namibia and the northwestern Northern Cape Province of South Africa. Although it approaches the Angolan border (Broadley 1990), there are no published records from Angola. Likewise, the nominotypical form occurs in the western Caprivi Strip, immediately adjacent to southeastern Angola and should therefore be expected in Cuando Cubango. Genus Thelotornis Smith, 1849 Thelotornis capensis oatesi (Gunther, 1881) Oates’ Twig Snake Thelotornis Capensis Smith 1849:19. Holotype: not located, presumed lost fide FitzSimons (1937) and Broadley (2001) (eolleetor A. Smith). Type loeality: “Kafihrland and the eountry towards Port Natal” [= Durban], South Afriea. Dryiophis oatesii Gunther 1881:330, eolor pi. D. Holotype: BMNH 1946.1.9.76. (formerly BMNH 78.7.31.1) (eolleetor C.G. Oates). Type loeality: “Matabeleland” [= western Zimbabwe] Broadley (2001:61). Dryiophis Kirtlandii: Boeage (1895a: 119). Thelotornis kirtlandii'. Loveridge (1936a:39). Thelotornis Kirtlandi'. Monard (1937b: 135). Thelotornis eapensis'. Bogert (1940:70), Laurent (1954a:58), Wallaeh et al. (2014:714). Thelotornis kirtlandii eapensis'. Loveridge (1944b:154, 1953a:277, 1957:275). Thelotornis kirtlandii oatesii'. Hellmieh (1957b:69). Thelotornis eapensis oatesii'. Laurent (1964a: 116), Spawls and Braneh (1995:23), Broadley (1979:130, 2001:61), Broadley and Wallaeh (2002:70), Broadley and Cotterill (2004:51), Eimermaeher (2012:63), Bates et al. (2014:423), Conradie et al. (2016:19). Global conservation status (lUCN): Least Concern. Global distribution: The species as a whole is known from Southern Angola and northern Namibia, west through northern Botswana, Zambia and southeast Democratic Republic of Congo to Zimbabwe, western Mozambique and Malawi, and south into the northeastern provinces of li-'E 1&°E 20“E 14* E Map 365. Distribution of Telescopus semiannulatus semiannulatus in Angola. 402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II South Africa. Thelotornis capensis oatesi occupies much of this range exclusive of South Africa, eastern Botswana and adjacent areas. Ocurrences in Angola (Map 366): The subspecies occurs in southern and central-east Angola. Lunda Norte: “Sombo” [-8.68333, 20.95000] (Laurent 1954:58). Lunda Sul: “Alto Chicapa” [-10.88333, 19.23333] (Lau¬ rent 1964a:116; Broadley 2001:68). Benguela: “Quisange” [-12.43333, 14.05000] (Bocage 1895a:119; Loveridge 1944b:159); “Hanha” [-12.25000, 13.75000] (Bogert 1940:70; Loveridge 1944b: 159; Broadley 2001:68). HmTa: “Caconda” [-13.73333, 15.06667] (Loveridge 1936a:40, 1944b: 159; Broadley 2001:68); “Quillengues” [-14.06667, 15.08333] (Bocage 1895a: 119; Loveridge 1944b:159); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:135; Loveridge 1944b:159); “Huilla” [-15.05000, 13.55000] (Bocage 1895a: 119; Broadley 2001:68), “Joa Almeida” [-15.18333, 13.68333] (Broadley 2001:68). Namibe: “Biballa” [-14.76667, 13.36667] (Bocage 1895a:119; Loveridge 1944b:159). Cunene: “Chimporo” [-16.03333, 17.15000] (Monard 1937b:135; Loveridge 1944b:159); “Dorbush am Cunene, Chitado (thorny bush in Chitado)” [-17.30000, 13.90000] (Hellmich 1957b:69; Broadley 2001:68). Cuando Cubango: “Cuito basin (52)” [-14.59333, q] (Conradie et al. 2016:9, 12, 19). Undetermined Locality: “Rio Quando” (Bocage 1895a: 11 9; Loveridge 1944b: 159). Taxonomic and distributional notes: Bocage (1895a) noted that some Angolan specimens in the Museu Bocage collections previously identified as Dryiophis Kirtlandii from “Quissange,” “Biballa,” “Rio Quando,” “Quillangues,” and “Huilla” resembled the individuals described by Gunther (1881) as Dryiophis Oatesi from “Matabeleland,” Zimbabwe. Loveridge (1944b) consid¬ ered two races of Thelotornis kirtlandii (Hallowell, 1 844), one from the north as the typical form and one from the south as Thelotornis kirtlandii capensis, which might represent the southern records of Bocage (1895a) and Monard (1937b). This association should correspond to Thelotor¬ nis capensis oatesi as suggested by Loveridge (1953a) and represented in Broadley (1979). Broadley (1979) and Broadley and Wallach (2002) presented distribution maps but did not list the localities plotted. The status of T c. oatesi as a valid taxon remains problematic and has not been resolved by morphological and genetic analyses (Eimermacher 2012). I2"E 16.°E 20°E I4'E. Map 366. Distribution of Thelotornis capensis oatesi in Angola. Thelotornis kirtlandii (Hallowell, 1844) Forest Twig Snake Leptophis Kirtlandii Hallowell 1844:62. Holotype: ANSP 5271 (collector E. Blanding, don. W. Blanding). Type locality: “Liberia, West Africa.” Dryophis Kirtlandi: Bocage (1866a:48). Thelotornis Kirtlandi'. Ferreira (1900a:52), Boulenger (1915:213), Parker (1936:125), Themido (1941:10), Laurent (1954a:59, 1964a: 116), Thys van den Audenaerde (1966:34). Thelotornis kirtlandii eapensis: Loveridge (1944c: 154). Thelotornis kirtlandii kirtlandii'. Loveridge (1944c: 149, 1957:274), Hellmich (1957b:69). Thelotornis kirtlandii'. Spawls and Branch (1995:24), Broadley (2001:57), Broadley and Wallach (2002:69), MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 403 Eimennacher (2012:64), Broadley and Cotterill (2004:51), Spawls et al. (2004:390), Broadley and Cotter- ill (2004:51), Chirio and LeBreton (2007:552). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from from Guinea-Bissau east through forested areas of West Africa and the Congo basin to Uganda and South Sudan, south to northern Angola, northwestern Zambia and south-central Tanzania. Ocurrences in Angola (Map 367): The species occurs in the northern Angola. Luan¬ da: “Luanda” [-8.83333, 13.26667] (Ferreira 1900a: 52; Themido 1941:10). Lunda Norte: “He Bena-Mai, riv. Luachimo, pres de Dundo (Bena-Mai island)” [-7.35000, 20.83333] (Lau¬ rent 1954a;59; Broadley 2001:66); “Dundo” [-7.36667, 20.83333] (Broadley 2001:66). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Bocage 1866a:48, 1895a:119; Loveridge 1944b: 154). Kwanza Norte: “Piri- Dembos” [-8.56667, 14.50000] (Hellmich 1957b: 69; Broadley 2001:66); “Ambaca” [-9.26667, 15.18333] (Ferreira 1900a:52); “Canhoca” [-9.25000, 14.68333] (Broadley 2001:66); “Cazengo” [-9.33333, 14.76667] (Ferreira 1900a:52; Loveridge 1944c: 154). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:125; Loveridge 1944c:154, 159; Broadley 2001:66). Taxonomic and distributional notes: Broadley (1979) and Broadley and Wallach (2002) pre¬ sented distribution maps but did not list the localities plotted. Map Angola. 367. Distribution of Thelotornis kirtlandii in Genus Thrasops Hallo well, 1858 Thrasops flavigularis (Hallowell, 1852) Yellow-Throated Bold-Eyed Tree Snake Dendrophis flavigularis Hallowell 1852b:205. Holotype: ANSP 5243 (collector H.A. Ford), lost fide Malnate (1971). Type locality: “Liberia, Western Africa,” in error, corrected to “Gabon” by Loveridge (1944b: 132). Thrasops flavigularis'. Peters (1877a:615), Bocage (1895a:97), Loveridge (1944c: 132), Hellmich (1957b:65), Frade (1963:252), Broadley and Wallach (2002:63), Chippaux (2006:108), Chirio and LeBreton (2007:558), Wallach et al. (2014:716). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from southwestern Nigeria through extreme east¬ ern Democratic Republic of Congo and northwestern Angola. Also on Bioko Island, Equatorial Guinea. Ocurrences in Angola (Map 368): The species occurs in the northwestern Angola including the Cabinda enclave. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Bocage 1895a:97; Loveridge 1944c:134); “Cabinda” [-5.55000, 12.18333] (Frade 1963:252). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hellmich 1957b:65). Taxonomic and distributional notes: None. 404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Il'E 16°E 20°E Map 368. Distribution of Thrasops flavigularis in Angola. Thrasops jacksonii Gunther, 1895 Black Tree Snake Thrasops Jacksonii Gunther 1895:528. Holotype: BMNH 1946.1.4.8 (formerly BMNH 94.2.1.7) (eolleetor F.J. Jaekson). Type loeality: “Kavirondo” [= southern Western and Nyanza provinees], Kenya. Thrasops jacksoni jacksoni: Thys van den Audenaerde (1966:33). Thrasops jacksonii-. Broadley and Wallaeh (2002:64), Broadley et al. (2003:214), Spawls et al. (2004:365), Chirio and LeBreton (2007:560), Wallaeh et al. (2014:716). Global conservation status (lUCN): Not Global distribution: The species is known from southern Cameroon, east through the Congo basin and southern Central African Republic to Uganda and western Kenya and south to northwestern Zambia, northeastern Angola, and the former Katanga Province in the Democratic Republic of Congo. Ocurrences in Angola (Map 369): The species occurs in northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Thys van den Audenaerde 1966:33), “Riv. Cama- iala” [-8.05000, 18.61667] (Thys van den Audenaerde 1966:33). Taxonomic and distributional notes: The Angolan records of Thys van den Audenaerde (1966) appear to be the only confirmation of the occurrence of Thrasops jacksonii in Angola. Evaluated. 1I°E l(j«E 2i>°E 24"E Map 369. Distribution of Thrasops jacksonii in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 405 Genus Toxicodryas Hallowell, 1857 Toxicodryas blandingii (Hallowell, “1844” 1845) Blanding’s Tree Snake Dipsas Blandingii Hallowell “1844” 1845: 170. Holotype: ANSP 10083 (collector E. Blanding, don. W. Bland- ing), lost fide Hughes and Barry (1969:1020). Type locality: “Liberia, West Africa.” Boiga blandingii: Laurent (1950a:9, 1954a:57, 1964a: 109), Thys van den Audenaerde (1966:34), Spawls and Branch (1995:26), Broadley and Cotterill (2004:50), Spawls et al. (2004:373). Toxicodryas blandingii: Trape and Mane (2006:168), Chirio and LeBreton (2007:562), Wallach et al. (2014:719). Global conservation status (lUCN): Not Evaluated. Global distribution; The species is known from Senegal and Guinea, east to west¬ ern Kenya, south to northern Angola, the south¬ ern Democratic Republic of Congo, and north¬ ern Zambia. Ocurrences in Angola (Map 370): The species is known from northeastern Angola. Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1950a;9, 1954a:57, 1964a: 109; Thys van den Audenaerde 1966:34); “Andrada” [-7.70000, 21.38333] (Laurent 1964a;109). Taxonomic and distributional notes: Some authors including Peters (1877a), Bocage (1895a), Parker (1936) and Hellmich (1957b) erroneously reported this species from northwestern Angola. Records from “Chin- choxo,” “Piri-Dembos,” and “Quirimbo” formerly identified as T. blandingii should be assigned to Toxicodryas pulverulenta (Fischer, 1856). Toxicodryas pulverulenta (Fischer, 1856) Dipsas pulverulenta Fischer 1856:81, pi. 3, figs, la-lc. Lectotype: ZMH 4376, formerly ZMH 339 (collector Dr. Davis), designated by Wallach et al. (2014:720). Type locality: “Edina, Grand Bassa County in Liberia (West-Afrrka),” corrected to “St. Thome,” Sao Tome and Principe, Gulf of Guinea” by Ladiges in Hugh¬ es and Barry (1969:1020), which is itself in error; the original type locality is probably correct as stated (see notes below). Dipsas pulverulenta: Peters (1877a:615), Bocage (1887a: 186, 1895a: 123). Dipsas Blandingii: Peters (1877a:615), Bocage (1895a: 124). Boiga blandingi: Parker (1936:125). Boiga blandingii: Hellmich (1957b:67), Loveridge (1957:268). Boiga pulverulenta: Parker (1936:109), Hellmich (1957b:68), Loveridge (1957:269), Frade (1963:252), Spawls et al. (2004:374). Toxicodryas pulverulenta: Chippaux (2006:155), Chirio and LeBreton (2007:564). Toxicodryas pulverulentus: Wallach et al. (2014:719). Global conservation status (lUCN): Not Evaluated. Global distribution: The species is known from West and Central Africa, from Guinea to Cameroon, east to Uganda and Kenya, and south to the southern Democratic Republic of Congo and Angola. Map 370. Distribution of Toxicodryas blandingii in Angola. Fischer’s Cat Snake 406 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Ocurrences in Angola (Map 371): The species is known from northwestern Angola. Cabinda: “Chinchoxo” [-5.10000, 12.10000] (Peters 1877a:615; Bocage 1887a;186, 1895a:123-124); “Cabinda” [-5.55000, 12.18333] (Frade 1963:252). Kwanza Norte: “Piri-Dembos” [-8.56667, 14.50000] (Hell- mich 1957b;67-68). Kwanza Sul: “Quirimbo” [-10.68333, 14.26667] (Parker 1936:125); “Congulu” [-10.86667, 14.28333] (Parker 1936:125). Taxonomic and distributional notes: Authors including Peters (1877a), Bocage (1895a), Parker (1936) and Hellmich (1957b) confused this species with Toxicodryas blandingii (Hallowell, “1844” 1845), resulting in erroneous estimations of species distribu¬ tions. Toxicodryas blandingii is limited to extreme northeastern Angola (Spawls and Branch 1995; Wallach et al. 2014) whereas the distribution of T. pulverulenta (Fischer, 1856) com¬ prises a large portion of the northwestern parts of the country. In the original description Fischer (1856) referred to two specimens (one subsequently lost fide Hallermann 1998) from “Edina, Grand Bassa County in Liberia (West-Afrika),” collected by Dr. Davis and donated to the Hamburg Museum. Hughes and Barry (1969), following a personal communication from the curator at that time, Werner Ladiges, corrected the type locality of Toxicodryas pulverulenta to “St. Thome,” pre¬ sumably Sao Tome and Principe, Gulf of Guinea. Schatti and Loumont (1992) and Wallach et al. (2014) followed Hughes and Barry (1969) and Chippaux (2006) incorrectly referred to “Saint Thome, Liberia” as the type locality of the species (there is no locality with this or a similar name in Liberia). The suggestion by Ladiges was certainly the result of a labeling mistake. The species has never subsequently been found on Sao Tome, despite recent extensive collecting by the Cali¬ fornia Academy of Sciences and Museu Nacional de Historia Natural e da Ciencia, and it has never been cited in any of the works on the herpetofauna of the island (e.g., Bocage 1879, 1886b, 1886c, 1886d, 1890, 1905; Greef 1884; Bedriaga 1892; Mananas 1958; Capocaccia 1961; Schatti and Loumont 1992; Nill 1993; Ceriaco et al. 2018). Therefore, we recommend that original type local¬ ity, “Edina, Liberia,” should stand as is. Family Natricidae Bonaparte, 1838 Genus Limnophis Gunther, 1865 Limnophis bicolor Gunther, 1865 Bicolored Swamp Snake Limnophis bicolor Gunther 1865b:96, pi. 2, fig. C. Syntypes: BMNH 1946.1.14.53-54 (formerly BMNH 64.10.28.16) (eolleetor F.A.P. Bayao, don. J.V. Barboza du Boeage) [2 speeimens]. Type loeality: “pro- vinee Duque de Braganee (Angola)” [= Calandula], Malanje Provinee, Angola. Limnophis bicolor. Boeage (1866a:47, 1866b:68, 1879b:95), Bogert (1940:36), Hellmieh (1957b:63), Mertens (1963:438), Broadley (1974:8), Braneh (1998:82), Broadley et al. (2003:175), Wallaeh et al. (2014:376). Helicops bicolor. Boeage (1895a:75, 1896a:112, 1897a:200), Boulenger (1905:112, 1915:201), Sehmidt (1933:12), Monard (1937b: 116). Limnophis bicolor bicolor. Laurent (1964a: 100). Il'E L&“E 20°E 24''E Map 371. Distribution of Toxicodryas pulverulenta in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 407 Global conservation status (lUCN): Not Evaluated. Global distribution; The speeies is known from the southern Demoeratie Republie of Congo, the Angolan highlands, western Zambia and westernmost Zimbabwe. Ocurrences in Angola (Map 372): The species is known from mainly from central Angola. Cabinda: “Luango” [-5.15000, 12.16667] (Bocage 1895a:75). Lunda Sul: “Poste de Cacolo, Alto Cuilo, dans une petit grotte d’ou sort une cascade sur le cours du Na- Ipanha, afluente du Cuilo” [-10.01667, 19.55000] (Laurent 1964a: 100); “Marais du Khoka, afluente du Kutele, sous-affluent du Cuango, Alto Cuilo” [-10.93333, 19.08333] (Laurent 1964a; 100); “Tyihumbwe” [-10.96667, 20.06667] (Monard 1937b:116). Malanje: “Duque de Braganga” [-9.10000, 15.95000] (Gunther 1865b:96; Bocage 1866a;47, 1866b;68, 1895a:75, 1897a:200; Mertens 1963:437); “Rio Loando” [-11.55000, 18.15000] (Bocage 1879b;95). Bie: “Chitau” [-11.43333, 17.15000] (Schmidt 1933:12). Map 372. Distribution of Limnophis bicolor in Angola. Huambo; “Bela- Vista” [-12.36667, 16.20000] (Hellmich 1957b;63; Mertens 1963:437). Bengue- la: “Cahata” [-12.35000, 14.81667] (Bocage 1895a:75); “Quindumbo” [-12.46667, 14.9333] (Bocage 1895a:75); “Hanha” [-13.30000, 14.20000] (Bocage 1896a:112). Huila; “Caconda” [-13.73333, 15.06667] (Bocage 1895a:75); “Kalukembe” [-13.78333, 14.68333] (Monard 1937b;116); “Kuvangu” [-14.46667, 16.30000] (Monard 1937b:116); “Huilla” [-15.05000, 13.55000] (Bocage 1895a:75); “Kakulakaze (afluente du Kului)” (Monard 1937b:116). Taxonomic and distributional notes; Thys van den Audenaerde (1965) considered the occur¬ rence of the species in the province of Bas-Congo, Democratic Republic of Congo as doubtful. Limnophis bangweolicus (Mertens, 1936) Bangweulu Water Snake Helicops bangweolicus Mertens 1936:284. Holotype: SMF 22172 (eolleetor F. Haas). Type loeality: “Nsom- bo, Nordende des Bangweolo-Sees, Nord Rhodesien” [= Nsombo, Lake Bangweulu], Zambia. Limnophis bicolor bangweolicus: Laurent (1964a: 100). Limnophis bangweolicus: Braneh (1998:82), Broadley et al. (2003:175), Broadley and Cotterill (2004:50), Wallaeh et al. (2014:376), Conradie et al. (2016:19). Global conservation status (lUCN): Not Evaluated. Global distribution; The species is known from Zambia, west to the former Katanga Province of the Democratic Republic of Congo and eastern Angola as well as the Caprivi Strip, Namibia, northern Botswana and western Zimbabwe. Ocurrences in Angola (Map 373): The species is known from just two published records, although it probably occurs in other regions of eastern Angola. Moxico; “environs du lac Calun- do” [-11.80000, 20.86667] (Laurent 1964a:100). Cuando Cubango: “floodplain of the Cuando River (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9, 10, 22). Taxonomic and distributional notes: None. 408 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II ll-E 1&°E 20°E 24" E Map 373. Distribution of Limnophis bangweolicus in Angola. Genus Natriciteres Loveridge, 1953 Natriciteres bipostocularis Broadley, 1962 Southwestern Forest Marsh Snake Natriciteres olivaceus bipostocularis Broadley 1962b:785. Holotype: NMSR/M 3524 (collector L.D.E.F. Vesey-FitzGerald). Type locality: “Chisansa, Abercon, Northern Rhodesia” [= Mbala], Zambia. Coronella olivacea: Peters (1882c: 115). Mizodon olivaceus: Bocage (1895a:74). Tropinodotus fuliginoides: Monard (1937b: 113, 115). Neusterophis olivaceus uluguruensis: Bogert (1940:35). Natrieiteres olivaceus uluguruensis: Hellmich (1957b:62), Loveridge (1957:256). Natrieiteres olivaeea bipostoeularis: Broadley (1963:5, 1966d:7). Natrieiteres variegata bipostocularis: Branch (1998:81). Natriciteres bipostocularis: Broadley et al. (2003:172), Broadley and Cotterill (2004:50), Wallach et al (2014:465). Global conservation status (lUCN): Not Evaluated. Global distribution; The species is known from northeastern Zambia, extending west through the southern Democratic Repub¬ lic of Congo to the highlands of central Ango¬ la. Ocurrences in Angola (Map 374): The species occurs in the central-western in Angolan highlands. Malanje: “Malanje” [-9.55000, 16.35000] (Peters 1882c:115; Bocage 1895a:74; Broadley 1966d:7). Huam- bo: “Bela Vista” [-12.36667, 16.20000] (Hellmich 1957b:62; Broadley 1966d:7). Map 374. Distribution of Natriciteres bipostocularis in Angola. MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 409 Hulla: “Kalukembe” [-13.78333, 14.68333] (Monard I937b:II3, 115 Broadley I966d:7). Taxonomic and distributional notes: Described by Broadley (1962d) as a subspecies of Natriciteres olivacea (Peters, 1845) from northeastern Zambia, the species was first considered as a montane race close to Natriciteres olivacea ulugurensis Loveridge, 1935, itself now regarded as a synonym of N. variegata (Peters, 1861) or as a subspecies of N. variegata (Peters, 1861) (Broadley 1966d; Broadley et al. 2003; Broadley and Cotterill 2004). Recently, N. bipostocularis, N. pembana (Loveridge, 1935) and A. sylvatica, Broadley, 1966, the last two also once considered subspecies of N. variegata, were recognized as full species (Broadley et al. 2003; Broadley and Cotterill 2004). Natriciteres olivacea (Peters, 1854) Olive Marsh Snake Coronella olivacea Peters 1854:622. Holotype: ZMB 4803 (eolleetor W.C.H. Peters). Type loeality: “Tette” [= Tete], Mozambique. Neusterophus atratus (Peters 1877a:614). Holotype, ZMB 9177 (eolleetor Afrieanische Gesellsehaft). Type loeality: “Chinehoxo (Westafrika)” [= Chinehoxo], Cabinda enelave, Angola. Coronella olivaeea: Boeage (1866a:47, 1866b:66). Coronella (Mizodon) olivaeea'. Peters (1877a:614). Tropidonotus olivaeeus: Boulenger (1915:112). Neusterophis olivaeeus olivaeeus'. Laurent (1950a:7). Natrieiteres olivaeeus olivaeeus'. Laurent (1954a:44). Natrieiteres olivaeeus olivaeea'. Hellmieh (1957b:62), Loveridge (1957:256). Natrieiteres olivaeea'. Broadley (1966d:9), Broadley and Cotterill (2004:50), Chirio and LeBreton (2007:496), Wallaeh et al. (2014:465), Conradie et al. (2016:22). Global conservation status (lUCN): Least Concern. Global distribution: The species is known from Senegal east to the Sudan and Ethiopia, south through southern Somalia to southern Mozambique and west through Zimbabwe, northern Botswana and the Caprivi Strip to Gabon and Angola. Ocurrences in Angola (Map 375): The species occurs throughout more mesic parts of Ango¬ la, although most records are from the north of the ocuntry. Cabinda: “Chinehoxo” [-5.10000, 12.10000] (Peters 1877a:614; Loveridge 1957:256). Lunda Norte: “Dundo” [-7.36667, 20.83333] (Laurent 1954a:44; Broadley 1966d:9); “Muita (Luembe E)” [-7.80000, 21.45000] (Laurent 1950a:7, 1954a:44; Broadley 1966d:9). Kwanza Norte: “Dondo” [-9.66667, 14.41667] (Hellmieh 1957b:62; Broadley 1966d:9). Malanje: “Pungo-Andongo” [-9.66667, 15.58333] (Boeage 1895a:74; Boulenger 1905:112; Broadley 1966b:9). Cuando Cubango: “flood- plains of the Cuando River (44a)” [-17.57333, 23.26000] (Conradie et al. 2016:9, 10, 22). Undetermined Locality: “North of Cuanza River” (Broadley 1966db:9). Taxonomic and distributional notes: A recent record from Cuando Cubango (Conradie et al. 2016) represents the first published occur¬ rence of the species from the south of Angola. ll^E 20°E 24”E Map 375. Distribution of Natriciteres olivacea in Angola. 410 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Acknowledgments The present work would not have been possible without the help and support of many people and institutions. We apologize in advance if we have inadvertently omitted anyone from the list that follows. From North American institutions, we particularly thank to the curators, collection managers and researchers of all institutions visited or consulted, namely Ned Gilmore (ANSP), Darrel Frost, Frank Burbrink, David Kirziran and Arianna Kuhn (AMNH), Jens Vindum, Lauren Scheinberg, Alan E. Leviton and Robert C. Drewes (CAS), James Hanken, Jonathan Losos , Jose Rosado, Joe Martinez, and Linda Ford (MCZ), Carol Spencer, Michelle Koo and Jimmy McGuire (MVZ), Alan Resetar (FMNH), Steve Rogers (CM), and Bob Reynolds, Addison Wynn, Rayna Bell and Roy McDiarmid (USNM). Important contributions were made by friends and colleagues in Portuguese institutions. As the present Atlas was originally presented as a M.Sc. thesis in the Conservation Biology M.Sc. program from Universidade de Evora, Evora, by the first author, we want to thank the professors in that program, and particularly Joao Rabaga and Paulo Sa-Sousa for their support and comments during the initial stages of this Atlas. From the Museu Nacional de Historia Natural e da Ciencia, we thank the staff and all of our colleagues, in particular Cristiane Bastos-Silveira, Isabela Queiroz, Alexandra Cartaxana, Alexandra Margal, Maria Judite Alves and Luis Filipe Lopes for their constant support for this project. From the Museu de Historia Natural da Universidade do Porto, we thank the director of the institution, Nuno Ferrand de Almeida, for providing access to the col¬ lection under his responsibility, and we particularly thank our friend Luzia Sousa, former curator of the zoological section, for her outstanding knowledge and dedication to the preservation of such an admirable collection. From the former Institute de Investiga^ao Cientifica Tropical, we thank Luis Mendes and Bivar de Sousa for their support and valuable discussions and suggestions during our data collection and research on both the herpetological collection and archive of the institution. From the Museu da Ciencia of Universidade de Coimbra, we thank the director, Carlota Simoes, and the manager of the zoological collections. Ana Rufmo, for their support during our visit to their museum. Other private individuals, especially Joao Crawford-Cabral and David Luna de Carvalho provided important data and information on topics related to the bio¬ geography and history of science of Angola. Several colleagues and friends from German institutions provided both specimen access and data regarding their collections and research in Angola and Africa, these include Gunther Kohler and Linda Acker (SMF), Uwe Fritz, Raffael Ernst and Markus Auer (SMF), Jakob Hallerman (ZMH), Michael Franzen and Frank Glaw (ZSM), Frank Tillack, Rainer Gunther and Mark- Oliver Rodel (ZMB), Wolfgang Bohme (ZFMK), Fritz Geller-Grimm (MWNH) and Philipp Wag¬ ner (now of the Allwetterzoo, Munster). From France, we thank Alain Dubois, Annemarie Ohler, Roger Bour and Marc Cugnet (MNHN) for their continued support, shared ideas and access to their rich collections. From Switzerland, we thank Andreas Schmitz (MHNG) as well as Amaud Maed- er (MHNC) for providing access to the collections under their care, and Urs Wiiest (NMBA) for collection data. From the United Kingdom, Patrick Campbell and Jeff Streicher (BMNH) respond¬ ed promptly and courteously to numerous requests for information and hosted our visits to London. From Belgium, we thank Danny Meirte (MRAC) and Georges Lenglet (IRSNB). From Austria we thank Heinz Grillitsch and Silke Schweiger (NHMW), from The Netherlands, Pirn Amtzen and Esther Dondorp (RMNH). In Sweden, we thank Goren Nilson (GNM), Sven Kullander (NRM) and Ericas Mejlon (UUMZ), and in Italy, Stefano Scab (MSNM), Franco Andreone (MZUT) and the staff of MZUB for data on specimens in their care. MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 4II From South Africa, we thank Michael Franeis Bates (NMB), Bill Branch and Werner Conradie (PEM), Lauretta Mahlangu, Lemmy Mashinini and Wulf Haacke (TM), Jofred Opperman (SAM), and Beryl Wilson (MMK) for information on their colleetions and other relevant data regarding Angolan and Southern Afriea herpetofauna. From Zimbabwe, Shiela Broadley and the late Don Broadley (NMZB), and from Namibia, Mathilda Awases and Eugene Marais (SMNW). From Angola, we would like to thank the Baptista family espeeially to Alvaro, “Varito” and young Pedro, for their outstanding and ineredibly organized logistical help during our field surveys in Namibe Provinee and tourism operator Candido Cameiro for kindly providing Figure 8. From the Faeuldade de Cieneias of Universidade Agostinho Neto, we thank Joao Serodio de Almeida, Michel Morais, Soki Kuedikuenda and Filomena Mateus, with whom we have activelky eollabo- rated and exehanged ideas regarding Angolan biodiversity. Soki Kuedikuenda was also the former direetor of INBAC, with whom the current eollaboration was initiated. From the Universidade Metodista de Angola, we thank Luis Sebastian, Diogo Figueiredo and Teresa da Silva Neto, dean of the university, for their institutional support at the beginning of this project. From the Museu Naeional de Historia Natural, we want to thank both the former and eurrent directors of the insti¬ tution, Francisca da Costa and Belmira Seeo de Oliveira Gumbe, respeetively, and all of their staff for their enthusiastie and professional support during our visits to their eollections and also during various publie lectures held in the institution. From the Museu do Dundo, we thank the director, Fonseca Sousa and his staff for opening the doors of their extraordinary eolleetion and allowing us to study Raymond Laurent’s type material. Our aeknowledgments extend to the Ministry of Culture, speeifieally to her Exeeleney the Minister of Culture, Carolina Cerqueira, and to the National Director of the Museums of Angola, Ziva Domingos, for allowing our investigations in both aforementioned institutions. From the Gabinete de Aproveitamento do Medio Kwanza (GAMEK), we thank Rafael Neto for his support during field surveys in the Lauea Dam area, which resulted in the eolleetion of important specimens. Alice Ponciano (Odebrecht), Cristina Rebelo (EnvGreen) and the environment team of Lauea Dam (Vicente Jose, Helena Pinto, et al.) provided invaluable support and friendship during the aforementioned surveys. The authors also wish to thank Daniel “Gastrula” Machado, Almeida Jose, Iracelma Machado, Fidel Viegas, and Sara and David Elizalde for their friendship and camaraderie. The Embassy of the United States of America in Luanda, and particularly Gustavo Guerrero and Todd Katsche, have provided institu¬ tional help whenever needed. From the Ministry of Environment, we wish to thank to both the former and current Ministers of the Environment, Their Excellencies Maria de Fatima Jardim and Paula Francisco Camuhoto, respectively, for their institutional support and permission to conduct research in the country. In addition, the former National Director for Biodiversity, now Secretary of State for Biodiversity, Joaquim Manuel offered his support since the beginning of the Atlas project. Provincial and com¬ munal governors have facilitated our field-work aeross various provinces of the country. The current National Director for Biodiversity, Nascimento Antonio, must also to be thanked. We espe¬ eially want to aeknowledge the friendship and support of our eolleagues from the Institute Naeional da Biodiversidade e Areas de Conservagao (INBAC), namely its eurrent direetor, Aristo- fanes Pontes, and viee-direetor, Maria Loa, and their eolleagues Sango de Sa, Suzana Bandeira, Hilaria Maehado, Celsia Afrieano, Ivania Castro, Miguel Xavier, Elizeth Gonsalves. The former direetor of the Institute, Abias Huongo, during his time leading the institute also provided outstanding support for our projeet. The authors also thank to all of the Angolan wildlife rangers and park offieers (Miguel Savituma, Antonio “Muloge” Lopes, Roland Goetz, among others) for their help and support during our field surveys and for their outstanding work on the preservation of the magnifieent biodiversity of Angola. Without INBAC and the dedieation of its staff to the study and proteetion of Angolan biodiversity none of our work would have been possible. 412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Finally, the authors thank to their respective families and friends for their support during the preparation of this Atlas. Funding for this work came from the California Academy of Sciences, Villanova University through the Gerald M. Lemole Endowed Chair Fund, the JRS Biodiversity Foundation, and the United States National Science Foundation (NSF DEB-1019443, 1202609, 1560667, 1556255, 1556559, and 1556585) through grants to Aaron M. Bauer, David C. Blackburn and Matthew P. Heinicke. Mariana P. Marques is currently supported by FCT contract SFRH/BD/129924/2017. Literature Cited Adalsteinsson, S., W.R. Branch, S. Trape, L.J. Vitt, and S.B. Hedges. 2009. 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Notes on the taxonomy and distribution of the African cobras of the genus Naja. Jour¬ nal of the Herpetologieal Assoeiation of Afriea, 2(l):21-30. Broadley, D.G. 1966d. A review of the genus Natrieiteres Loveridge (Serpentes: Colubirdae). Arnoldia {Rhodesia), 2(35): 1-11. Broadley, D.G. 1968a. A revision of the African genus Typhlosaurus Wiegmann (Sauria: Scincidae). Arnoldia {Rhodesia), 3(36): 1-20. Broadley, D.G. 1968b. The venomous snakes of Central and South Africa. Pages 403-435, in W. Bucherl, E.E. Buckley, and V. Deulofeu eds., Venomous Animals and their Venoms. Volume L Venomous Verte¬ brates. Academic Press, New York, New York, USA. xxiv + 707 pp. Broadley, D.G. 1968c. A review of the African cobras of the genus Naja (Serpentes: Elapinae). Arnoldia {Rhodesia), 3(29): 1-14. Broadley, D.G. 1968d. Anew species of Crotaphopeltis (Serpentes: Colubridae) from Barotseland, Zambia. Fieldiana, Zoology, 5 1(10): 135-139. Broadley, D.G. 1969. 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However, in the time between the original submission and final acceptance of this manuscript several additional papers relating to the composition and distribution of the Angolan herpetofauna have been published or are in press and may appear in advance of the present atlas. Although many more publications will follow in short order and this work will soon be out of date, we thought it useful to note at least those of which we are aware. Baptista, N., T. Antonio, and W.R. Branch. 2018. Amphibians and reptiles of the Tundavala region of the Angolan Esearpment. Biodiversity & Ecology 6:397-403. Ceriaco, L.M.P., S. Bandeira, M.P. Marques, I. Agarwal, E.L. Stanley, A.M. Bauer, M.P. Heinicke, and D.C. Blackburn. 2018. Anew earless speeies of Poyntonophrynus (Anura: Bufonidae) from the Serra da Neve Inselberg, Namibe Provinee, Angola. ZooKeys 780:109-136. Ceriaco, L.M.P., S.D.A.C de Sa, and A.M. Bauer. 2018. 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MARQUES ET AL.: AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 457 Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola: Atlas of historical and bibliographic records (1840-2017) APPENDIX 458 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Table Al. Checklist of Amphibians and Reptiles recorded for Angola. (lUCN RL) Glob¬ al status according to the lUCN Red List of Threatened Species (2017): “EN” Endangered, “VU” vulnerable, “LC” least concern, “DD” data deficient, “NE” not evaluated. Taxa listed as “ — ” are those for which uncertainties preclude inclusion in the current lUCN assessments. SpL'c ics/S u bspcc ics Commem name Fmdemicity lUCN RL A IN UR A Dumeril, 1805 Family PIPIUAE Gray, 1825 Genus Xenopus Waglcr, 1827 Xenopus andrei Loumonl, 1983 Andre’s Clawed Frog — LC Xempm cf. epilrnpicalis Fischberg, Colombclli and Picard, 1982 Congolese Clawed Frog — LC Xenoptts nutelleh (Peters, 1844) Muller’s Clawed Frog — LC Xeimpits peteraii Bocagc, 1 895 Peters' Clawed Frog — LC Xempus poweri Hewitt, 1927 Power’s Clawed Frog — NF. Xempits sp. — — - Family BUFOINIIME Cray, 1825 Genus Mcrtensophrync Tilien, I960 Merfensophyne mdampkmra (Schmidt and Inger, 1959) Dark-Sided Toad — LC Xferiensophrytie afV. mocqitardi (Angd, 1924) Mocq Liard 's Toad — DD Genus PoytUimophrynus Frost et al.. 2006 Poymomphrymts tiomhemss (Bocage. 1895) Donibe Toad — LC Poyntonophrytma ^rnmdiso} uie (Poynton and Haackc, 1993 ) Grandison’sToad Endemic DD Poyntonophymts kavan^^emis (Poynton anti Broadlcy, 1988) Kavango Toad — LC Genus Schismatierma Smith, 1849 Schismudenna carem (Smith, 1848) African Red Toad — LC Genus Scierophrys Tschudi, 1838 Sckrophrys huchnen (Peters, 1882) Buchner's Toad — DD Sciemphrys ftmerea (Bocagc, 1866) Angolan Toad — LC Sderophiys ^armani ( Meek, 1897) Garman’s Toad — LC Sekfophrys giiduratis (Power, 1927) Guttural Toad — LC Sekrophrys kmairii (Boulenger, 1901) Lema ire’s Toad — LC Sekrophrys power! (Hewitt, 1935) Power's Toad — LC Sckrophiys lY/a { M ertens, 1937) Mertens’ Striped Toad — LC Sekrophrys regtdaris (Reuss, 1833) African Common Toad — LC Family MICROHYIJr>AE Gunther, 1858 (1843) Genus Phrytiomaiuis Peters, 1867 Phrvnomamis a films Bou tenge r, i9n'i Spotted Rubber Frog — LC Phiynomands anneehms Werner, 1910 Marbled Rubber Frog — LC Phiynomoniis bifasciaHiS (Smith, 1 847) Banded Rubber Frog — LC MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 459 Family BREVICIPITIDAE Bonaparte, 1850 Genus Breviceps Merrem, 1820 Breviceps poweri Parker, 1934 Power's Rain Frog — LC Breviceps sp. - - — * Family HEMISOTIUAE tope, 1867 Genus fiemism Wagler, 1S27 Hem ism gitineensis tope, 1865 Guinea Snout-Burrower — LC Hem ism mannonifus (Peters, 1854) Marbled Snout-Burrower — LC Family HYPEROl.IIDAE Laurent, 1943 Genus Afrixuius Laurent, 1944 Afrixalm dorsaiis (Peters, 1 R75) Striped Spiny Reed Frog — LC Afrixulm fuivovittatus (Cope, “1860” 1861) Four-Lined Spiny Reed Frog — LC Afrixahts osorioi (Ferreira, 1906) Osorio’s Spiny Reed Frog — LC Afrixahts quadrivitfatm (Werner, 1908 “1907”) Four-Lined Spiny Reed Frog LC Afrixahts \vhiei ( Laurent, 1941) dc Witte's Spiny Reed Frog — LC Genus Cryptothyiax Laurent and Conrbaz, 1950 Crvpiothyiax greshofii (Schitthuis, 1889) Greshoff s Wax Frog — LC Genus HyperoHus Rapp, 1842 HyperoHus adspersus Peters, 1 877 Sprinkled Long Reed Frog — LC Hyperoiius angoiensis Steindachner, 1^7 Angolan Reed Frog — LC HyperoHus bengueUensis (Bocage, 1893) Benguela Long Reed Frog — LC HyperoHus bicolor Ahl, 193 1 Tw'o -Colored Reed Frog Endemic DD HyperoHus bocagei Steindachner, 1867 Bocage' s Reed Frog — LC HyperoHus cheiaemis Conrad ie. Branch. Measy and Tolley, 201 2 Chela Mountain Reed Frog Endemic DD HyperoHus ciriereus Monard, 1937 Ashy Reed Frog Endemic LC HyperoHus cintmmnmeo ventris Bocage, 1866 Cinnamon-Bellied Reed Frog — LC HyperoHus concohr (Hallo well, 1844) Variable Reed Frog — LC HyperoHus darteveilei Laurent, 1943 Dartevelle’s Reed Frog — LC HyperoHus fuscigula Bocage, 1 866 Brown -Throated Reed Frog Endemic Dl> HyperoHus glamlicohr Peters, 1878 Peters’ Reed Frog - LC HyperoHus gu laris Ahl, 1931 Luanda Reed Frog Endemic DD Hyperoiius kivuemis Ahl, 1 93 1 Kivu Reed Frog — LC HyperoHus langl Noble, 1 924 Lang’s Reed Frog — LC HyperoHus maesius Rochebrune, 1885 Cabinda Reed Frog Endemic DD HyperoHus misufus Gunther, 1 865 Large-Nosed Long Reed Frog — LC HyperoHus nil idulus Peters, 1875 Plain Reed Frog — LC HyperoHus oceilatm (Giinther, 1858) Golden -Eyed Reed Frog — LC HyperoHus plat\’ceps (Boulenger, 1900) Benito River Reed Frog — LC HyperoHus poUi Laurent, 1 943 Tshimbulu Reed Frog — DD 460 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II ilypemlhis profchei Rochcbrunc, 18S5 Rochcbrunc’s Reed Frog Endemic DD HyperoUus pusiilus (Cope, 1862) Water Lilly Reed Frog — LC Hypefolkfs qumpievitiafus Bocage, 1866 Five-Striped Reed Frog — LC Hypei oihis raynmmii Conradic, Branch and Tolley, 201 3 Raymond’s Reed Frog Endemic DD HyperoUus rhkophihss Rochcbrunc, 1885 African Reed Frog Endemic DD HyperoUus sieimiachneri Bocagc, 1866 Steindaehner’s Rccd Frog ■ — ■ LC HyperoHus xnUietuis Laurent, !964a Vilhena's Reed Frog Endemic DD Genus Kassim Girard, 1853 Kassirta kuvangeosis (Monard, 1937) Knvangu Kassina — LC Kassina maculosa (Slemleld, 1917) Marbled Running Frog — LC Kassina senegalensis (Dumeril and Bibron, 1841) Senegal Kassina — LC Family ARTHROLEPTIDAE Mivart, 1869 Arthroiepth Smith, 1849 Anhrolepfis curquejai Ferreira. 1906 Carqueja's Squeaker Endemic DD Arthrolepfis kimeerei de Witte, 1921 Lameer’s Squeaker — LC Arfhrokpits spinalis Boulcnger, 1919 Tanganyika Scrcccliing Frog — DD Arihrolepds stenodactvhss Pfeffer, 1893 Common Squeaker — LC Arihrolepds xenochirus Boulenger, 1905 Plain Squeaker — LC Genus LepUfpelis Glint her, 1859 LepiopeUs anchietae (Bocage, 1873) Anchieta’s Tree Frog Endemic LC LepiopeUs uubryi (DunierH, 1856) Gabon Forest Tree frog — LC LepiopeUs bocagii (GOiilhcr, 1865) Bocage’s Tree Frog — LC LepiopeUs cynnamomeus (Bocagc, 1893) Angola Forest Tree frog — LC LepiopeUs jonkmi Parker, 1936 C ongulu Forest Tree frog Endemic DD LepiopeUs marginahts (Bocagc, 1895) Quissanguc Fo rest Tree frog Endemic DD LepiopeUs notaius (Peters, 1875) C ommon Forest Tree frog — LC LepiopeUs virkUs (Giinther, “1868” 1869) Rusty Forest Tree fr og — LC Genus Trkbobmrachus Boulenger, 1900 Tricobairaebus rohusius Boulenger, 1900 Hairy Frog — LC Family PTYCHADENIDAE Dnbois, 1987 Genus HUdehrutidda Nieden, 1907 Hitdebramiiia ornalissinui (Bocage, 1879) Angola Ornate Frog Endemic DD Hihiebrartdiia ornata (Peters, 1878) Ornate Frog — LC Genus Piydmdena Boulenger, 1917 Piychadem anchietae (Bocage, 1867) Anchieta’s Grass Frog — LC Pivchaderia ansorgii (Boulenger, 1905) Ansorge’s Grass Frog — LC Pivchadem bunodernia (Boulenger, 1907) Rough Grass Frog — LC MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 461 Ptychadena ^l amlisonae Laurent, 1954 Grandison’s Grass Frog — LC Ptychadena guihei Laurent, 1954 Guibe’s Grass Frog — LC Ptychadena keiiifigi (Monard, 1937) Kei ling’s Grass Frog — LC Ptyc'hafiena cf. mascareniensis (Dumeril and Bibron, 1841) Mascarene Grass Frog ' — ‘ LC Ptychadem cf. mos/sambica (Peters, 1854) Mozambique Grass Frog ' — LC Ptychadena oj^Thynchns (Smith, 1 849) Sharp-Nosed Grass Frog — LC Ptychadena petplicaki Laurent, 1964 Many-Ridged Grass Frog — LC Ptychadena parosissi/na (Steindachner, 1867) Striped Grass Frog — LC Ptychadena sitbpunctata (Bocage, 1 866) Spotted Grass Frog — LC Ptychadena taeniosceHs Laurent, 1954 Small Grass Frog — LC Pt}>chadena upemhae (Schmidt and liiRcr, 1959) Upemba Grass Frog — LC Ptychadena tiiung\yemis (Loveridge, 1932) Udzungwa Grass Frog — LC Family PHRYNOBATRACHIDAE Laurent, 1941 Genus Phtymfbatraehm Giiiither, 1862 Phtynabatracims brevipalniatus (Ahl, 1925 “1923”) Ahl’s Screeching Frog Endemic DD Phn nobatmehm ciypiotis Schmidt and Inger, 1 959 Cryptic River Frog — DD Phtyno batrachm mababiensb FitzSimons, 1932 Mababc Puddle Frog — LC Phrynobatrachm minutus (Boulenger, 1895) Ethiopian Dwarf Puddle Frog — LC Phry’nohatracbm naialensis (Smith, 1 849) Natal Dwarf Puddle Frog — LC Phtynobatrachtfs parvidus (Boulenger, 1905) Dwarf River Frog — LC Phry^nohatraebifs plicatm (Gunther, 1 85S) Coast River Frog — LC Family PYXICEPHALIDAE Bonaparte, 1850 Genus Dubois, 1987 Amietia angokmis (Bocage, 1866) Angola River Frog — LC Genus Boulenger, 1917 Anbrki sp. - — NE Genus Pyxieephaim Tschudi, 18,38 Pyxicephalus edidis Peters, 1854 Edible Bullfrog — LC Genus Tomoptenui Dumeril and Bibron, 1841 Tomoptema ctyptoiis (Boulenger, 1907) Tremelo Sand Frog — LC Toinopterna damarennis Dawood and Charming, 2002 Damaraland Sand Frog — DD Tomoptei'm kmgerensis Passmore and Caituthers, 1975 Knocking Sand Frog — LC Tomopterna tandyi Channing and Bogart, 1996 Tandy’s Sand Frog — LC 462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Tomoplerna tuberculosa (Boulenger, 1882) Rough Sand Frog — LC Family DICROGLOSSIDAE Anderson, 1871 Genus Hopiohatrachas Peters, 1863 ! hpkibiitnichus occipitaUs (Gtlnthcr, 1858) African Crowned Bullfrog — LC Family RANI DAE Ratsch, 17% Genus Amuirma Dubois, 1992 Anmirana alhohbrss (Halloweil, 1856) White-Lipped Frog — LC Amuinma dariingi (Boulcngcr, 1902) Darling’s W hite- Lipped Frog — LC Amnimm lemairet (de Witte, 1921) Lemaire’s W'hite-Lipped Frog — LC Atiunram lepus (Andersson, 1903) Andersson’s W'hite-Lipped Frog — LC Amninma parketiam (Mertens, 1938) Parker’s W'hite-Lipped Frog Endemic DD Family Rll ACOPIIORIDAR Hoffman, 1932 (1858) Genus Chtramutuh Peters, 1854 Chiromantis xerampeiina Peters, 1854 Grey Foam- Nest Tree frog — LC REPIILIA ORDER CHELONII Brongniart, 1800 Fa mily C 1 1 1: LON 1 1 D A E Op pel, 1 8 1 1 Genus Curetta Rafinesque, 1814 Caretta caretta (Linnaeus, 1758) 1 Loggerhead Sea Turtle | — 1 Genus Chelania Brongniart, 1800 Chehma mydas (Linnaeus, 1758) 1 Green Sea T urlle | — 1 EN Genus Eretmocbelys Fitzinger, 1843 Eretmochetvs (Linnaeus, 1766) llawksbill Sea Turtle — CR Genus Lcpidocbelys Fit linger, 1843 Lepkiocheiys oHvacea (Eschschollz, 1829) Olive Ridley Sea Turtle — VT) Fa mi ly l>E R M OC 1 1 E L Y 1 DA E Fitz i nger, 1 843 Genus Dermaeheiys Blainville, 1816 Dcrmochelys coriacea ( Vandelli, 1761) Leatherback Sea Turtle — VLl Family PELOMEDUSIDAE Cope, 1868 Genus Pehmedttsa Waglcr, 1830 Pehmedma suhnifu (Bonnaterre, 1789) II el meted Terrapin — NE Genus Pelashs Wagler, 1830 Pelitssos bechitanicus FitzSimons, 1932 Okavango Mud Turtle — NE Peiusios cusianeits (Schweigger, 1812) W'est African Mud Turtle — NE Peiusios chapini Laurent, 1965 Central African Mud Turtle — NE Peiusios gabonemis (Dumdril, 1 856) African Forest Turtle — NE Peiusios nanus Laurent, 1956 A fr i can D warf M ud Turt 1 e — NE Peiusios rhodesianus Hewitt, 1927 Variable Mud Turtle — LC MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 463 Family TESTlJDIlNinAE UaJsch. 1788 G emits Cych/dermu Peters, 1854 Cychdenm mthry>i (Dumeril, 1856) Aubry ’s Softshell Turtle — NE Genus KinLvys Bell, 1827 KiiiLxy’s beiliaiia Gray, 1831 Bdfs Hinge-Back Tortoise — NE Kinixys erosa ( Schweigger, 1812) Forest Hinge-Back Tortoise — DO KinLxys spekii Gxsiy, 1863 Speke's Hinge-Back Tortoise — NE (ienns Sfig/twcitefys Gray, 1873 Stiymochelys paida!ix (Bell, 1828) Leopard Tortoise — LC Family TRIONYCHIDAE Fitzinger, 1826 Genus Cyclodcrma Peters, 1854 Cyciadenna cmbryi (Dumeril, 1856) Aubry^’s Softshdl Turtle — VU Genus Trhnw Geoffroy Saint-Hilaire, 1809 Tnomx friHrtguii (Forskiii, 1775) African Soflshdl Turtle — VU Order CROCODVLIA Gmelin, 1789 Family CROCODYIJDAE Cuvier, 1808 Genus Cmcodylus Laiirenti, 1768 Crocody/us mhtictis Laurcnti, 1768 Nile Crocodile — LC (^enus Mechtops Gray, 1844 Mecixtops cf, catophractu-s (Cuvier, 1 825) West African Slender- Snouted Crocodile — CE Genus Osft’ofuemii'i Cope, 1861 Osfeolaenms ieimspis Cope, 1 861 A frican I3warf Crocod i 1 c — VU ORDER SQUAMATA Oppel, 1811 Family CJEKKONIDAE Gray, 1825 Gunm A/roedura Loveridgc, 1944 Afroedura bogeni complex Loveridge, 1944 Bogert's Rock Gecko — NE Genus Bauer, Good and Brandi, 1997 Afrogecko amorgii (Roulenger, 1907) Ansorge's Gecko Endemic NE Genus ChondradactylHx Peters, 1870 Chondtodactyius Jdzsimofm (Loveridge, 1947) Button-Scaled Thick-Toed Gecko — NE ChmdrodacfyitfJi hevigalus (Fischer 1888) Button-Scaled Gecko — NE Chondmdcictyhts pidirzerae (Schmidt, 1933) Pulitzer's Thick-Toed Ciecko — NE (Jenus //emidacfylu.'i Okeis, 1817 Heniidacfyhis mgidaiits complex II alio well, 1852 African Tropical Gecko — NE Heniidaay/us hayomi 1 Socage, 1 893 Bayao’s Gecko tiidemic NE HemidadvhfS benguedemh Bocage, 1 893 Benguda Gecko Endemic NE ffemidactyhis longkepbuhts Bocage, 1 873 Long-Headed Tropical Gecko — NE Heniidtictyliis: nwbottia (Moreau De Jonnes, 1818) Tropical House Gecko — NE Heniidactvfus ef. miiriceus F^eters, 1870 Guinean Spiny Tropical Gecko — NE 464 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus Kotekatuis Heittieke, Duza, Greenbaum, Jackman and Bauer, 2014 Kolekanos plumicaitdifs (Haacke, 2008) Feather-Tailed Gecko Endemic NE Genus Lyf^oductylus Gray, 1864 Lvgodacivhis angoiensis Bocage, r896 Angola Dwarf Gecko — NE Lygodaciylus brudfiehis Hewitt. 1932 Bradfield’s Dwarf Gecko — NE Lygodaclyhi.\ capetisis (Smith, 1 849) Cape Dwarf Gecko — NE Lvgodactvhfs chobiensts FitzSimons, 1932 Okavango Dwarf Gecko — LC Lygodaciylu& kmrencei Hewitt, 1926 Lawrence’s Dwarf Gecko — NE Genus Pachyduttyi us W i eg ni a n n , 1834 PachYdactvhis augoiemis Loveridge, 1944 Angolan Thick-Toed Gecko Endemic NE Pacbydacrylm camcid icus Fil/.Simons, 1959 Arigola Banded Thick -Toed Gecko — NE Pachydactyius oreophdus complex Me L ach Ian and S pence, 1967 Kaokoland Rock Gecko — NE Pachydacfyhts pimciaius complex Peters, 1854 Speckled Thick-Toed Gecko — NE Pachydactyius ningei (Anderssoni, 1908) Namib Web-Footed Gecko — LC Pachydactyius scherzi Mertens, 1954 Scherz’s Thick-Toed Gecko — NE Pachydactyius scuiaius Hewitt, 1927 Scaly Thick-Toed Gecko — NE Pachydactyius vamyii (Sleyn and Haacke, 1966) Namib Desert Gecko — NE Pachydactyius wahlbergii (Peters, 1869) Wahiberg’s Kalahari Gecko — NE Genus Rhuptropus Peters, 1869 Rhoptropus afer Peters, 1 869 Namib Day Gecko — NE Rhoptropus barmirdi Hewitt, 1926 Barnard’s Namib Day Gecko — NE Rhoptropus benguellemis Mertens, 1938 Benguela Namib Day Gecko Endemic NE Rhoptropus bipot asm FitzSimons, 1957 Fitzsimons’ Namib Day Gecko — NE Rhoptropus boidtom Schmidt. 1933 Boulton's Namib Day Gecko — NE Rhoptropus montamts Laurent, 1964 Mountain Namib Day Gecko Endemic NE Rhoptropus taeniostictus Laurent, 1964 Angolan Namib Day Gecko Endemic NE Rhoptropus sp. — Endemic - Family AMPHISBAEINIDAE Gray, 1825 Genus Dalapbia Gray, 1865 Dalophia angoteusis Gans, 1976 Angolan Worm Lizard — NE Dalophia cllenhcrgeri (Angel, 1920) Ellenberger’s Worm Lizard — NE Dalophia pisfUluiti (Boeltger, 1895) Blunt-Tailed Worm Lizard — NE Genus Monopcfds Smith, 1848 Monopeltis anchietae (Bocage, 1873) Anchieta’s Worm Lizard — LC Monopellis htfuscata Broad ley, 1997 Infuscate Wedge-Snouted Worm Lizard — NE Monopelt Is luandae Gans, 1976 Luanda Worm Lizard Endemic NE MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 465 Mormpeltis perplexiis Gans, 1976 W'edge-Snouted Worm Lizard Endemic NE MotropelHs vamierysii de Witte, 1 922 Vandery'st’s Wonn Lizard — NE Monopehis welwslschsi (Gray, 1865) W'elwitsch’s Worm Lizard Endemic NE Geitiis Cope, 1885 Zvi^aspis rtigra Broad Icy and Cans, 1969 Black Round-Snouicd Worm Lizard — LC Zygaspis (piudrifrom (Peters, 1862) Kalahari Round- Snouted Worm Lizard — NE Family LACERTIDAE Bonaparte, 1831 Genus Helioholas Fit finger, 1843 Heiioboltis Ittgitbhs (Smith, 1838) Bush veld Lizard — NE Genus HitlmpisGtny, 1863 Hoiaspis guenthers Gray, 1 863 Blue-Tailed Tree Lizard — NE Genus L-htuftropis Peters, 1854 Ichnotropis hiviUakt hivsitaia Bocage, 1866 Angolan Rough- Scaled Lizard — NE ichnotropi^ bivsUata pallida Laurent, 1964 Pale Rough-Scaled Lizard Endemic NE Icimotropis capensis capetisis (Smith, 1838) The Cape Rough-Scaled Lizard — NE Ichnotropis capensis overlaeli de Witte and Laurent, 1 942 Lunda Rough- Sea led Lizard Endemic NE Ichnoiropis microlepidoia Marx, 1956 Marx’s Rough-Sealed Lizard Endemic NE Genus Meroks Ciray, tS3S Mecoles anchietae ( Bocage, 1 867) Anchieta’s Dune Lizard — NE Mcroks resknlattis (Bocage, 1867) Reticulate Sand Lizard — NE Meroles squanntlosus (Peters, 1854) Common Rough-Scaled Lizard — NE Genus Nucras Gray, 1838 Nncras scalaris Laurent, 1964 Scaled Sand veld Lizard Endemic DD Nucras aff. tessellaia (Smith. 1838) W ester n Sand veld Lizard — - Genus Pedioplanis Fitzinger, 1843 Pediopahm henguetlemis (Bocage, 1867) Bocage’s Sand Lizard — NE Pediopalms haackei Con rad ie. Measey, Branch and Tolley, 2012 llaacke's Sand Lizard Endemic NE Pcdiopnhm htmdeyi Conrad ie, Measey, Branch and Tolley, 2012 Huntley’s Sand Lizard Endemic NE Family CORI>Y LI DAE Mertens, 1937 Genus Chmmesuara Sclineider, 1801 Chamaesattra angtdna oHgopholis Laurent, 1964 Angolan Snake Lizard — NE Chamaesaiira miopropus Boulctiger, 1894 Zambian Snake Lizard — NE Genus Cardylm Laurenti, 1768 Cordylm angolemis (Bocage, 1895) Angolan Girdled Lizard Endemic NE Cordylus machadoi Laurent, 1 964 Machado’s Girdled Lizard — NE Cordylm namakidytis Stanley, Ceriaco, Bandeira, Valerio, Bates and Branch, 2016 Kaokoveld Girdled Lizard Endemic NE 466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family GERRHOSAlJRIDAE Fiuinger, 1843 Genus Cordyiosaurtis Gray, 1865 |1S66) Conivhsam us sublessellatus (Smith 1844) Dwarf Plated Lizard — LC Genus Gerrlwsaurus Wiegmanti, 1828 Getrhosaut'iis aurilus Boeltger, 1887 Kalahari Plated Lizard — NE Cenhoxaurifs hulsi Laurent, 1 954 Laurent’s Plated Lizard — NF Gerrhosaurus mulfiliNearits ttocage, 1866 Kwanza Keeled Plated Lizard Endemic NE Gerrhasaurm cf, nigrolineatus HaUowelL 1857 Black'Lined Plated Lizard — NE Getrhosaiinfs skoogi Andersson, 1916 Desert Plated Lizard — LC Genus !\fafobos(iurus Bates and Tolley, 2013 Matobosaw'ifs fmldahni (de Grys, 1938) Western Giant Plated Lizard — NE Genus Te/mdactylus Merrem, 1820 Tetrudacnltis ellenbergeri (Angel, 1922) lillenberger's Plated Snake- Lizard — NE Family SCINCIDAE Gray, 1825 Genus Cuvier, 1816 |1S17| A cofitkix jappi (Broadley, 1968) Baroiseland Blind Legless Skink — NE AcoiUkiS kgaiagadi Lamb, Biswas and Bauer, 2010 Kalahari Legless Skink — NE Acofidas occidentafis FiuSimons, 1941 Savanna Legless Skink — LC Genus Euniecia Boeage, 1870 Eiiinecia anchietae andueiae Bocage, 1870 W'eslem Serpenti form Skink ■ — NE Etmecia at^ddeiae major Laurent, 1964 Major Western Serpentiforni Skink Endemic NE Genus t'eylinia Cray, 1845 Feylima currori Gray, 1 845 Western Forest Feyiinia — NE Feyiinia degans (1 tail 0 well, 1852) Elegant Feyiinia — NE Feyiima gramissqmmis Muller, 1910 Large-Sealed Feyiinia — NE Genus Lepidothyrh Cope, 1 892 Lepidothyris hinkeii joei Wagner, Boh me, Fauwels and Schmiu, 2009 Joe’s Red-Flanked Skink — NE Genas Leptosiaphm Schmidt, 1943 Lepmsiaphos de^ ittei (Loveridge, 1934) de Witte’s Five-Toed Skink — NR Genas Lithttya Horton, 1972 Luhuya ivemii {Bocage, 1 879) Ivens’ Skink — NE Genus Meiamtseps Boulenger,. 1897 Mehnosep.s ocddenialis (Peters, 1 877) Western Limbless Skink — NE Genus Moddits Gunther, 1864 Moddus sundevadii (Smith, 1849) Sundevall’s Writhing Skink — LC Genus Panaspis Cope, 1868 Panaspis hreviceps (Peters, 1873) Peters’ Snake-Eyed Skink — NE MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 467 Panaspis cabimiae (Bocage, Cabinda Snake-Eyed Skink — DD Panasph macuHcoHb Jacobsen and Broadley, 2000 Speckle-Lipped Snake-Eyed Skink NE Panmph a IT, wuhibergii (Smith, IS49) Wahlberg's Snake-Eyed Skink — NE (Mentis Sepsina Bocage, 1866 Sepsim ai^goietisis Bocage, 1 866 Angolan Reduced- Li mb Skink — NE Sepsina biiyuni (Bocage, 1866) BaySo's Reduccd-Limb Skink — NE Sepsina copei 1873 Cope's Reduced-Limb Skink Endemic NE Genus Trachylepis Fitzinger, 1843 Trachylepis ucutsiahris (Peters, 1 862) Wedge-Snouted Skink — NE Trachylepis aljinis (Gray, 1838) Senegal Skink — NE Trachylepis cf. alhoptmcfata (Bocage, 1867) Angolan Variable Skink — NE Trachylepis hayami (Bocage, 1872) BaySo's Skink — DD Trachylepis binokita (Boeagc, 1867) Ovambo Tree Skink — NE Trachylepis bocagii (Boulongcr, 1887)’ Bocage 's Skink — LC Trachylepis cbimbana (Bonlcnger, 1 887) Chimban Skink NE Trachylepis damarana (Peters, 1870) Kalahari Variable Skink — NE Trachylepis boeschi (Mertens, 1954) Hoesch's Skink — NE Trachylepis cT kKertiformis (J*eters, 1854) Bronze Rock Skink — LC Trachylepis laevis (Boulenger, 1907) Angolan Blue-Tailed Skink — NE Trachylepis macuUlahris (Gray, 1845) Speckle-Lipped Skink — NE Trachylepis cf. megahtra (Peters, 1878)^ Grass-Top Skink — NE Trachylepis motianii nom. tiov. Marques, Ceriaeo. Blackburn and Bauer Monard's Skink Endemic NE Trachylepis occidenkilis (Peters, 1867) Western Three-Striped Skink — NE Trachylepis pimendata (Bocage, 1872) Speckled Sand Skink — NE Trachylepis spslogaster (Peters, 1882)' Kalahari Tree Skink — NE Trachylepis sulcata ( Peters, 1 867) Western Rock Skink — NE Trachylepis wahlbetgli (Peters, “1 869” 1 870) Wahlberg's Striped Skink — NE Genus Bocage, 1873 Typhlacotuinsjohmonit Andersson, 1916 Johnson's Burrowing Skink — NE Typhlacmfias puncfatlssimm hogertll Laurent, 1 964 Rogert’s Dotted Blind Dart Skink Endemic NE Typhlacoatias punctafisshn us ptmemtissinms Bocage, 1873 Doited Blind Dart Skink — NE Typhlacoatias rohani Angel, 1923 Rohan’s Blind Dart Skink — NE Typhlacoutias rudebeckl Haacke, 1997 Rudebeck’s Blind Dart Skink Endemic NE 468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Family VARANIHAE Hardwifkf and Ciray, 1S24 Genus Varnnus Mcrrcm, 1820 Vanimtfi tilhigakins (ifhigulark (Diiudin, 1802) White-Throated Monitor — LC Varatiux alhigiihf k angoiamis Schmidt, 1933 Angolan Wiiitc-Throated Monitor — LC' Varamtx tuhtk’iix (Linnaeus, 1758) Nile Monitor — LC Family CHAMAELEONIDAE Gray, 1825 (>enus Chamaek’a Laurenti, 1768 Charttaeleo cimhiefae Bocage, 1872 Anchieta's Chameleon — LC Chanttieieo dUepis tfuUeiisix Bocage, 1886 Quilo Flap-Neck Chameleon — LC Chtiniaeieo gfuciUs etiemei Schmidt, 1919 Etienne’s Ciracet'ul Chameleon — LC Chamaeieo ftainacfuemix Smith, 1831 Namaqua Chameleon — LC Genus Trhcervs Swainsun, 1839 Triocems oweni (Gray, 1831) Owen's Chameleon — LC Family AGAMIDAE Gray, 1827 Genus Acatilfutcercus Fitzlnger, 1843 AccmthocerctfS cymocephalux (Falk, 1925) Angolan Tree Agama — LC Acoinhocerais sp. — — NE Genus Oaiidln, 1802 Agtintu octihafci Merrem, 1 820 W'estern Ground Agama — LC Agania anchietae Bocage, 1 896 Anchieta’s Agama — NE Agowjo cofjgfco Peters, 1877 Congo Agama — NE Agama miicosoensk llellmich. 1957 Mucoso Agama Endemic NE Agiimci plonicepx Peters, 1862 Namib Rock Agama — NE Agarua schacki Mertens, 1938 Schack’s Rock Agama Endemic NE Aga/iia sp. — Endemic NE SERPE^TES Family TYPHLOPIDAE Merrem, 1820 Genus Afrotyphti/px Broad ley and W'allach, 2009 Afmp'phhps angolensis { Bocage, 1866) Angola Blind Snake — NE Afrohphlops anomaliis (Bocage, 1873) Angolan Giant Blind Snake Endemic NE Afrotyphhps lineohiiiis (Jan, 1864) Common Lined Blind Snake — NE Afrofyphhpx miicmso (Peters, 1854) Zambezi Blind Snake — NE A frotypfiiops xchleget ii ( B ianco ni , 1849) SchlegePs Giant Blind Snake — NE Afronphhpx schmhii! (Laurent, 1956) Schmidt's Blind Snake — NE Genus Letheabia Cope, 1868 Lelheohia praeociiiaris (Stejneger, “1893" 1894) Leopoldville Beaked Snake — LC MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 469 Family LEPTOTYPHLOPIDAE Stejnt-ger, 1892 Genus Leptotyphhps Fit/.inger, 1843 Leptotypbhps kafubi (Boulenger, 1919) Shaba Thread Snake — NE Lepiotvphhps {Peters, 1854)’ Peters’s Thread Snake — • NE Genus Namibhm Hedges, Adiilsteiiissoii and Branch, Z0U9 Namibkma labktUs (Stem fold, 1908) Daniara Thread Snake — NE Ncmiibkma tafifrons {Stcmfeld, 1908) Bengueia Thread Snake — NE Namibima rostra fa (Bocage, 1 886) Angolan Beaked Thread Snake Endemic DD Family PYTHONIDAE Fitxinger, 1826 Genus Pytboir Dandin, 1803 Python anchietae Bocage, (887 Anchieta’s Dwarf Python — LC Python natalensis Smith, 1 840 Southern African Rock Pylhon — NE Python sebae (Gmelin, 1 789) African Rock Pylhon — NE Family BOIDAE Cray, 1825 Genus Caiaharia Gray, 1858 Cahboria reinhardtii {Schlegel, 1851) Calabar Ground Boa — NE Family VIPERIDAE OppcI, 1811 Genus Athens Cope, 1 862 Athens septantigera (flallowell, “1844" 1845) Variable Bush Viper — NE Genus Bids Gray, 1842 Bit is arietans (Merrem, 1830) PufT Adder — NE Biiis catidafis (Smith, 1839) Homed Adder — NE Bills gahonico Dumeril, Bibron, and Dumeril 1854 Gabon Adder — ■ NE Bids heraldica (Bocage, 1889) Angolan Adder Endemic NE Bids nask'ornis (Shaw, 1792) Rhinoceros Viper — NE Bids peri ngtie}h (Bouleoger, 1888) Peringuey’s Adder — LC Genus Caiisus Wagicr, 1830 Caimis hilineafiis Boulcngcr, 1905 Two-Striped Night Adder — NE Causiis Uchtensteind (Jan, 1 859) Forest Night Adder “ NE Caiisits imicidafiis (Hallowell, 1842) Spotted Night Adder — NE Caitsns resifiius (Peters, 1862) Green Night Adder — NE Causiis rhombeatus (Lichtenstein. 1823) Rhombic Night Adder — NE Family LAMPROPHHDAE Fitzinger, 1843 Genus/4wfA/|.'(Jt((p,¥ffs Peters, 1857 Atnblvotfipsas polylepis (Bocage, 1873) Common Purple-Glossed Snake — LC Genus ApparaBucins Sinilh, 1849 Apparalactits capensis Smith. 1 849 Cape Centipede Eater — LC Genus Atractaspis Smith, 1849 Atractaspis bibron ii Smith, 1849 Bibron’s Stiletto Snake — NE 470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Airi!cfaspi& boulengeri Mocquard, 1897 Boulenger’ s Stiletto Snake — NE Alractaspi.H cotigica Peters, 1877 Congo Stiletto Snake — NE Atractaspss irregularis (Reinhardt, 1843) Variable Stiletto Snake — LC Airacfaspis reticuiafa heiertychilus Boylcnser, 1901 Reticulate Stiletto Snake — DD Genus Htmedon Uumcrtl, Hihrctn and l>umenl, 1854 Boaedon angolemis Bocage, 1895 Angolan House Snake Endemic NE Boaedon lutiginosus complex (Bote, 1827) Brown House Snake — NE Boaedon olivaceiis (Dumeril, 1856) Olive House Snake — NE Boaedon vanegatus (Bocage, 1867) Variegated House Snake Endemic NE Genus Boihrophthahnus Peters, 1863 Bothrophihalmus Uneatus Peters, 1863 Red-Black Striped Snake — NE Genus Oromophis Schlegcl, 1S37 Dromophis lineaiits (Dumeril, Bibron and Dumeril, 1854) Lined Olympic Snake — NE Genus Gonioiwtuphis Buulenger, 1893 Cnntonotopbis hrimauxi (Mwquard, 1889) Moequard’s File Snake — NF Genus Hemirbagerrhis Boettger, 1896 liemifhageirhis viper ina (Bocage, 1873) Western Bark Snake — NE Genus Hypoptophis Buulenger 1908 Hypoptophh u Usonii Boulenger, 1908 Wedge-Snouted Burrowing Snake — NE Genus Umaformosa liroadley, Tolley, Conradie, Wishart, I’rapc, Burger, Rusamba, Zassi- Boulou, and Creenbaum, 2018 Umaformosa capensls (Smith, 1847) Southern Fite Snake — LC Umaformoset vernayi (Bogert, 1940) Angola File Snake — NE Genus Kit/inger, 1843 Lycophidion heiimichi Laurent, 1 964 Hcllmich’s Wolf Snake — DD Lycophidion laieraie I lallowell, 1857 Flat Wolf Snake — NE Lycophidion meleagre Boulenger, 1893 Speckeled Wolf Snake — NE Lycophidion midtimacidafittn Boeitger, 1888 Spot ted W'o 1 f S nake — NE Lycophidion ornatwn Parker, 1936 Ornate Wolf Snake — LC Genus Csiki, 1903 Meheiya poensis (Smith, 1847) Western Forest File Snake — NE Genus Pole/non .Ian, 1858 Pofemon collaris (Peters, 1 88 1 ) Collared Snake-Eater — NE Genus Prffsymiia Gray, 1849 Prosymna auibigm Bocage, 1 873 East African Shovel -Snout — LC Prosymna ango/ensis Boulenger, 1915 Angola Shovel-Snout — LC Prosymna frotitalis (Peters, 1 867) South-Western African Shovel- Snout — LC Prosymna vi.sseri FitzSimons, 1959 Visser's Shovel -Snout — NF MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 471 Genus Psammoplm Bole, IS25 Psammaphif; angolensix (Hocage, 1872) Dwarf Sand Snake NE Psammophis amoi-gii Boulenger, 1905 Link-Marked Sand Racer Endemic NE Psammophis jailae Pcracca, 1 896 Jalla's Sand Snake — NE Psammophis leopardimts (BtKage, 1 887) Leopard Sand Snake — ■ NE Psammophis mossambictis Peters, 1882 Olive Whip Snake — NE Psammophis namibattsis Broad ley, 1975 Namib Sand Snake — NE Psammophis notosiktus Peters, 1 867 Karoo Sand Snake — NE Psammophis suhtaenialus Peters, 1882 Stripe-Bellied Sand Snake — LC Psammophis Ihgrammus Gunther, 1865 W'estern Sand Snake — ■ NE Psammophis zambiensis Hughes and Wade, 2002 Zambian Whip Snake — NE Genus Psammaphylax Fitzinger, 1843 Psammophyiax acutiis (GOnther, 1888) Striped Beaked Snake — NE Psammophvlax rhambeatus ocetiatus (Bocage, 1873) Spotted Skaapsteker Endemic NE Psammophvlax tritaeniatus (Gunther, 1 868) Striped Skaapsteker ‘ — LC Genus Pseudasph Fitzinger, 1826 Pseudaspis cana (Linnaeus, 1758) Mole Snake — NE Genus Pythonadipsas Gunther, 1868 Pvthonodipsas curirtatu GUnther, 1868 W'estern Keeled Snake — NE Genus Xem/citiamus Ciinther, 1868 Xemcaiamtis bicaior machadoi Laurent, 1954 Machado’s Quill -Snouted Snake — NE Xenocalamns mechowii inornaius de Witte and Laurent, 1947 Inornate Elongate Quill- Snouted Snake — NE Xenacalamns mechowii mechowii Peters, 1881 Elongate Quill -Snouted Snake — ■ NE Family ELAPIDAE Boie, 1827 Genus /Is/JH/e/fl/fs Fitzingcr, 1843 Aspidelaps iuhricus cowiesi Bogert, 1940 Angolan Coral Snake — NE Genus Dendroaspis Schkgel, 1848 Dendroaspisjamesoni (Traill, 1843) Jameson’s Mamba -- NE Dendroaspis polyiepis (Gunther, 1 864) Black Mamba — LC (knus Elapsoideti Bneage, 1866 Elapsoidea gaemherii Boeage, 1866 Gunther’s Garter Snake — NE Eiapsoidea semiamulafa moebhisi (Werner, 1897) Moebius' Garter Snake — NE Eiapsoidea semianmdata semiannuiata Bocage, 1 882 Angolan Garter Snake — NE 472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II (lOnus Najii Laurenti, I76}i Ndjd {inchieUie l^ocagc, 1 879 Anchieta's Cobra NE iWiija ammia/a Buchholz and PcIcts, 1876 Ringed Water Cobra — NE Na/(i mckmoleHca Hallowcll. 1857 Forest Cobra - NE Nnja mossiirnhictf Peters, 1 854 Mozambique Spitting Cobra — NE S’ajii imilfifaxckikt Werner, 1902 Many-Banded Cobra — NE Ncija iiiffricinctii liogerl, 1940 Western Barred Spitting Cobra — NE nigricfiUis Reinhardt, 1 843 Rlaek-Necked Spitting Cobra — NE xuhjhh’ii Laurent, 1955 Savanna Cobra — NE Genus Pseuilohiije Giinther, 1858 Pxeudothije gohtii (Boulenger, 1895) African Tree Cobra — NE Family CQLUBRiDAE OppcI, 181 1 Genus Clmniuefyfus Bnuleiif’cr, 1919 Chamadrais parkeii (Angel. 1934) Parker’s Banded Snake — NE Genus Cmraphiipeftix Fitxinger, 184,3 Crokfphopellis hoki/nboeki (Laurenti, 1768) Red-Lipped Snake — NE Genus Dasypehis Wagler, 1830 Dctxype/lix palnumnH ( Leach, 1818) Palm Egg Eater = NE DdsypeUix scsihid (Linnaeus, 1758) Common F.gg Rater — LC Genus Dipsmk/btta Giinther, 1858 Dtpsadobifa xltrevei (Loveridge. 1932) Shrcve's Tree Snake — NE Genus Dixpfwfidiix Ouvernoy, 1832 Dixphalidns fyptix lypitx (Smith, 1828) Booms lang — NE Dispholidtfx tvpifx piinctaltis Laurent, 1955 Spotted Boom slang — NE Genus CiniyUi Giinther, 1858 Crayki caesur (Giinther, 1863) Caesar's African Water Snake — NE Gmyki y/virtto ( Bocage, 1866) Grnate African Water Snake NE Gtayki xmythk (Leach. 1818) Smyth’s African Water Snake — NE Gmyia tholloiii Moequard, 1897 Thol Ion's African Water Snake — NE Genus Hapxidi)phr}>!i Fischer, 1856 Hapxidop/in-x xmanigdiiiiis (Schleccl, 1837) Emerald Snake — NE Genus flortiioiiolHS llallowell, 1857 ffonnonotiix madexltix (DLiineril, Bibron and Dumcril, 1854) Uganda House Snake — NE Genus kyciuldtnuuftrpftm Liehtenstein, 1823 Lyivdtmoiiiorphiix stihlaemktfx Laurent, 1954 Eastern Congo White-Bellied Water Snake — LC Genus Mffpawveidaphis Figueroa, McKeh'y, Grismer, Bell and Lailvaus, 2016 Mopamvddophix zehrimis (Broadley and Schalti, "1997” 1999) Mopaneveld Snake — NE MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 473 Genus Phihthamnus Sniiili, 184tt Philaihamnus an^olemis Bocage, 1882 Angolan Green Snake — NE Philolhammts cannams (Andersson, I%1) Thirleen*SeaIed Green Snake — NE Philrtihamnitx dorsalk (Bocage, 1 866) Striped Green Snake — NE Philoihamnus heterodermtis (HallowdI, ! 857) Emerald Green Snake — NE Phshthamims heiero!epklotm (Gtinrhcr, 1863) Slender Green Snake ■ — NE Phitoihamnm hophgaater (Ciuntlier, 1863) Southeastern Green Snake — NE Phihlhammis mUdus ioveridgei Laureni, 1960 Loveridge’s Green Bush Snake — NE Phihihamnm ormtus Bocage, 1 872 Ornate Green Snake — NE Philothamnus semivariegatvs {Smilh, 1 840) Spotted Bush Snake — NE Genus Rhainnttphis GUnthir, 1862 Rhamnophis aethioplssa GOnthcr, 1862 Large-Eyed Green Tree Snake — NE Genus Scdp/thphis Peters, 1870 Scaphhphis a/bopimckitti.'? Peters, 1870 African Shovel-Nosed Snake — NE Genus Tetescopus Wagler, 1830 TelexcopiLs fiukeliieyi 1 laacke, 2013 Daniara Tiger Snake — NE Telescopus semiamttifatttJs semianmilafm Smith, 1 849 Common Tiger Snake — NE Genus Thrtisups Ha Howe II, 1858 Thrasops fJavigiilans (Hallow'd), 1852) Y el lo w-T hroated Bo Id-Eyed Tree Snake NE Thrasops Jackson ii Gunther, 1 895 Black Tree Snake — NE Genus Tftefolornis Smith, 1849 TVie/oj'oriJi'jf capensis oalesi (Gunther, 1881) Oates’ Twig Snake • — LC Thelafornis kii /landii (HalknvcIL 1 844) Forest Twig Snake — NE Genus Taxicadryas HallowdI, 1857 Toxicodnm hlandmgd (HallowdI, “1844” l’845) Blanding's Tree Snake — NE Toxicodnas piilventlema (Fischer, 1 856) Fischer's Cat Snake — NE Family NATRICIDAE Bonaparte, 1838 Genus Limiwp/tis Gunther, 1865 Limnopins hangwealicus (Mertens, 1936) Bangweulu Water Snake — NE Limnophis bicolor GUnthcr, 1 865 Bicolored Swamp Snake — NE Genus Nafriciferes Loveridge, 1953 Nalriciteres hiposloculans Broad ley, 1962 Southwestern Forest Marsh Snake — NE ^'atriciferes oiivacea (Peters, 1854) Olive Marsh Snake — LC 474 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Table A2. Amphibian and reptile taxa that have been removed from the speeies list for Angola. This ineludes taxa that have undergone revision, resulting in name ehanges applieable to Angolan forms. Species References citing nanie Sec accoiint(s) ANURA Dumeril, lltOS Faniilj PIPIDAF, Gray, 1S25 Geaiis XeHflyjm' Wagler, 1827 Xertopus caicaruitts Peters, 1875 Peters (I877a:6 18). .V. cf. epiiropicalis Xenopiisfrasen Bouknger, IW5 Laurent (1950a; 13, 1954a:70), Cd (1977; 16), Kobe! (1981:120), Ruas( 1996:20), Charming (200 1:240), Fa-ley et al. (201 1:22), Cbanninget al. (2012:294), Wagner ctal. (2013:206), Ernst et a!. (2015: 147). Xenopits sp. Xenopiif iXswpus) laevis (Daudin. 1802) Gunther ( IS65a:480), Boulenger( 1905:107), Monard (I937a:25. 1938:55, 76), Nellraich ( I957a:22), Inger (1959:540), Kobel (1981 :120), Gavetti and Andreone (1993:41), Frdey et al. (2011 :22), X. petersii Family BlIFOMDAE Gray, 1825 Genus TseKudi, 18,^8 Bii/o [= Sdtropho’s] guineemis OUnther,"l858” 1859 Peters ( 1 877a:6 1 8), Bcfcage ( 1 879b:89), Scleroplin, Frdey et al. (201 1:23). Frost (2016), Cerlaco el al. f2016a: 19). Sclerophr\>s pusilla Bufo [= Scleropljr}'s] paniherinus Dum^ril and Bibron, 1841 Bocage ( 1 866a:56). ScIeroplir}’s regularis Bu/o [= Scleropbrys] spinosvs (Daudin. 1803) Bocage ( 1 S67b,:227), Loveridge ( 1 936a:82). Scierapliij’s gutfuralis Family BREVlCiriTlUAE IJomii>arle, 1851) Genus Breviceps Merrem, 1S20 Breviceps athpersns Peters, 1882 Poynton (1982:67, 1992: 68). Frost (1985:356, 2016), Channing (2001:213), Poynlon and Broadicy (1985a;523), Ruas (2002:142). du Pneezand Carrulhers (2009: 108), Fneley et al. (201 1 :27), Conrad ie et al, (20 1 6: 1 1 ), Breviceps Sp. Breviceps gibbosHS (Linnaeus, 1758) Bocage (1870:68, 1873b:227), Breviceps sp. Breviceps mossambicus Peters, 1854 Bocage ( 1 895a; 1 82). Parker ( 1 934; 194), Monard (1 937a:29, 1 938:56. 8 1 ), Hellmich ( 1 957a:30), Loveridge ( 1 957:357), Inger (1959:532), Laurent (1964a; 156), Cei (1977:17, 18). Poynton (1982:67, 1992:68), Frost (1985:356, 2016), Gavetti and Andreone (1993:1 14), Ruas (1996:22), Channing (2001:213), duu Preezand Carrulhers (2009:108), Freley et al. (2011:27). Ruas (2002:142). Breviceps sp. Family HE.MISOTIDAE Cope, 1867 Genus Hemisas Wagler, 1827 Hemisus giiflaluni 1842) Bocage ( 1 S95a: 1 84), Monard (1 938:56), Fradc ( 1 963:254). H. gitineensis MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 475 Ht’inisus sudanense Steindachner, 1864 Boulenger { 1 882: 1 78). H. marmoratux Faniily FEROLIIDAE Lsiiirtnt, l■^43 tic nils Eniirtnl, l')44 Xfegalixiis [= dfrixalux] feptosomua Peters, 1 877 Boulenger {1882:129). AfrixaSvx quadrivittiitiix tic nils tfypendim Kapp, 1S42 NyfyeroHus cincnvenirif! Cope, 1862 [-HyperaliHS argiix Peters. 1854] Bocage { 1 895a: 1 68), Boulenger ( 1 905: 1 1 0). Noble { 1 92.1:252). Monard (1938:9.1). linger (19.59:541 ). Cei (1977:17). H. hocagei Hyperoiius Bouletiger, l‘H)l Laurent (1964a; 155). Cei (1977:17), H. adspersus HyperoliUi gmueri Alil, 1951 [ = Hyperontts marginadis Peters, 1854] Merteiis (I937a:2!)). H. iingokmii- Hyperoliui marmtratus Kapp. 1842 Ganllier()864a:480), Bocage (1 866a: 55. i879c:89. l8S6b;74; 1 895a: 1 64, 1 S96a: 113,1 897b:2 1 1 ). Peters (1881:1 50), Boulenger ( 1 882: 1 2 1 , 1 90S: 1 09), Ferreira ( 1 904; 1 12, 1 906: 1 60). Noble ( 1 921:253 ). Loveridge ( 1 936a: 106, 1 953a:350). Schmidt ( 1 936: 1 3 1 ). Monard ( 1 9.17a:35. 1 938:89), Merlens (l93Sa:427), Barbour and Loveridge (1946:127), luger (1959:541), Laurenl (1961:88), Pcrrci (I976a:27). Cei (1977:17), Gevetti and A ndreone (1993:103), Freteyet al (201 1:32). Ceriaco et al. (20 14b;669). N. cingolefisis Hyperolitis microps Giinlhcr, 1 S64 Bocage (I866a:55, 1866b;75), Frade (1963:254), Noble (1923:253). H. ptisilkis l/yperolitis pctrallelus Gilnther, 1858 Peters (I877a:61 8). Laurent ( 1 943a: 14; 1961:89). Cei ( 1 977: 1 7). Fretey ci al (201 1 :32), Ceriaco et al. (20 1 4b:669), Frost (2016). H. ungolensis Uyperolius piiticfidatia Bocagc, 1895 Bocage (1897a;204), Ferreira ( 1904: 1 12). Noble (1923:253). Loveridge (193 6a: 107, I936b;405), Mcrtens ( 1937:20), Monard ( 19.38:86). Frade (1963:254), Perrei (1976a:27), Frosi ( 1985:216), Aniiet (2005:275). a. nuxuliif: Family AR 1 HKOLEin il>AE Mivart, 1869 tic n us Leptffpelis G uni her, 1859 Lcpiopeli.i parhfKagii Poynton and Broadley, 1987 Poynicin and Rrnadley (1987:171), Sehiote and van Daele (2QO,3;146). L hfKagii Family r'[ yCllAI)EMr>AE Dtibnis, 1987 tie mis Ply ctniile fill Btinlenger, 1917 Pi\Tluidena amorgei (Bouicnger, 1905) Laurenl ( 1 954a: 74). P. perpUcam Ratiii [= PpThademi] bibroiiii Hallowell, 1845 Monard (I937a;5l, 1938:109), Laurenl (1950a: 14 ). P. grandisonae, P. guihei 476 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Plyvluittena chiysogasrer Laurt:nL 1954 Fretey et al. (2011 :4 1 ). P. guihei Ram [= Pfyvhadena] mascareniemis (Dumeril and Uibron. I841J Boulengertl866a:53, 1882:52, 1905:107), Bocage ( 1895a: 160), Monard (19373:50, 1938:108), GuM and Lammote (1957:978). Loveridge (1957:342), Ruas (1996: 25. 2002:144), Cei (1977:16, 17), Frost (1985:473), Ghanning (2001 :329). Pickersgill (2007a; 128), Frdiey cl al. (201 1:41 ), Frost (2016), Conrad ie et al. (2016: 18). Ptychadem anchietae, P. cf, mmcareniemis, P. poroshsima Pl^’chatk'm! puniitio ( Bou Icnge r, 1920) Larger (2001 :342), Frctcy cl al. (20 1 1 :4 1 ). P. luensosceth Family f'YXICFPUALlDAF, Bonaparte, ISSfl Genus Amietia Dubois, ]9{t7 Ram [= Amietki\ fusdg!ita (Dumeril and Bibron, 1841) Ingcr (1959:540), Schmidt and Ingcr (1959:48). Amietia aiigolensis Ccniis Boiilciigcr, IM17 Aubtiii musuko Ohicrand Kaz,^di, 1990 Channing (2011 :2S4), Frcley et al. (20 1 1 :42), Clianining ot al. (2012:329), Frost (2016), Aubria sp. Aubria suhsigiitaia (Duimeril, 1856) Motiard(l937a:47, 1938:104), Cd (1977:17), Ferret (1996:96). Aubria Sp. Genus Pyxk^pbalas Tscitutli, 18-^8 Pyxicephatus adspentts Tschudi. 1838 Bocage (1895a: 157), Monard (I937a:46, 1938:103), Inger ( 1 959:54 1 ). 1 radc ( 1 963:254). Cei ( 1977: 1 7). l••rosl ( 1 985:477). Channing (2001:346), du Preez and Carulhers (2009:414), Channing et al. (2012:123). P, eduUs REPTILIA ORDER CHEI OMl Bron^niarl, ««0fil Family PELOMEDL'SIDAE Cojie, 1868 Genus Pekimeduxa Waller, 1830 Pmton^x PehmeihiSit] gehafie (Ruppell, 1835) Bocage (1870:68), P.submfa Genus Peiusios \Va(;ler, 1830 Sterttothcierys (- Pthisios] sinuiitus (Smilh. 1838) Bocage ( 1 895a:4). Monard ( 1 937b: 148). Schmidt (1933:3). P. rhcHtesiamis Pelusios subnigcr (Bonnaterre, 1789) Bocage { 1 895a:3), Peters ( 1 877a;6 1 1 ), Monard (1931:109, 1937b: 148). Schmidl ( 1933:3), Mcnens ( 1 938a:430), l overidge (1941 b:49 1 . 1 957; 1 75), Frade (l%3:252.253). P. bechuanicus, P. casumeus, P. numts^ P. rbodesiuniis tJrdcrSyUAMA rA OppcI, 1811 Family GEKKOMDAE Gray, 1825 Genus CUondradactylus Peters, 1870 1 ioittoductyius 1= PuchythayhiS ~ Ctmndrixiact\'hts\ bibronii (Smith, 1846) Bocage (I867b:220, I867c:227, !895a:l5. IS87b;202. 1887c:209). Boulengcr (1885:201 ), Merlens (1926:152), Monard (I937b;53). C. piditzerae, C. kie\'igutuii MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 477 Chondrodactyius tumeri (Gray, 1864) Branch (1998:254). Heinz (2011:30), Bates el al. (2014:104), Cerlaco et al. (20146:670). C. laevigatus Genus Hemidactytus OKaa, 1817 Hemidacty‘!ux hrookii Gray. 1 845 Spawls et al. (2004:86). H. a»gulatus comple-x Hcmidactylux p!at%vephahis Peters, 1854 Bocage(lS66a:42, 1866b:60, 1870:68, 1873b;209). H. longicephcdus Genus Paehydut'tylus VViegmann, 1834 Pachydacty'hi'i oceUafm (Cuvier, 1817) [= P. ^es(js (Sparnnan, 1778)] Bocage{!867b:220. IS95a:16). Boulenger( 1885:205, 1905:110), Frade (1963:253). P. punctatus complex PiK'hydactyhis &ervitl Wemer, 1910 Monard(l93l 90, I937b;54). P. pmetatiis complex Family AMPHISBAEMDAE Gray, I82S Genus (84S Monopeltis capemis Smith, 1 84S Bocage (18736:216, IS95a:28), Loveridge (I94k:425). Gans (1967:85). M infuscata Family LACEKI1UAE liunaparte, 1831 Genus Nucraa Gray, 1838 Nticras ornata (Gray, 1 864) Broadley(l%5b:23). N. afT, texseihta Genus Fediopiuttis Fitzinger, 1843 Pediopkmis namaquensii (Duineril and Bibron, 1839) Boulenger { 1 887:91 ), Bocage ( 1 895a:3 1 ), Branch ( 1 998; 1 72), Makofchaci al. (2007:623). P. benguiiiettsis Pedioplwis wnt/ato (Smith, 1838) Laurent {1964a: 60). P. htiackei, P. hundeyi Family CORDYLIRAE Mertens, 1937 Genus CftHWiaeiatf™ Schneider, 18(J1 Chamoesuunt miscrok^pis (Cope, 1 862) Bocage ( 1 895a:25>, Monard ( 1 937b:6 1 ), Hellmich ( 1 957b:52). C. niiopropui Genus Cordylm l,atirea(i, 1768 Cordylus tropidostermm (Cope, 1869) Broadley (1971:22). C. angolemis Family SCINCIDAE Gray, 1825 Genus ,'ic'oiff/a.v Cuvier, 1816 “1817” plumhem Bianconi, 1849 Monard (I937b:96). A, oeddetitaih Genus Lepidolbyris Cope, 1892 Ly^gosoma [= MocMus = Lepid(/fhyns\ fermndi (Burton, 1836) Laurent { I964a:78), Chirio and LeBreton (2007:266). Lepidothyfis hinkeli joei (lenus Moehlui Giiniher, 1864 Xtochfus qfer (Peters, 1 854) Bocage { 1 867b:222, 1 867c:227). Ei/meces [= Mochlux] re/joH/ijfw.'i (Smith, 1849) Bocage ( 1 S79c;88). Mochfu.i xunde],'ailii 478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genuii Bn cage, lS6fi Scehtes f= Stysj;™] biftes (Linnai^us, 1766) Gtlnllier (1864a:480). Sepsina httyoni Genus Tracfiylepis Fititinger, I84J Eiipyepes [= Trachylepia] gravenluvstii (Duineril and Bibroii, 1839) Bccage (lS66a;44). T. hayomi Xhhuya [ ^ Eitprepix = Trachylepis] (ftiitt(}ueiaeiiiai(t (Lkliteiislem, 1823) Bo ulenger {1887:198, 1905:1 1 1), Bncage (1866a:44), liellniich (l957b:54),Brygofl (1985a 90), T. {}inot(ir(i, T hneagii Ettprepes [= Tnich_viepis] peirotefii (Dumcriland Bibron, 183'?) Peters ( 1 877a;614). Bocage ( 1 895a;39). Trachylepis macjilHabris Familj CHAMAELEONIDAE Gray, 1825 Genus Chumttetea Lauroiiti, 1768 Chtwiaeico svrtegaiensis Daiidin, 1802 GOnlher (I864a:480). Bocage { 1 870:68), Peters {1881 : 47). Lovendge{ 1957:97), C. grticilis eliemwi Family AGAMIDAE Gray, 1827 Genus Againa Daudin, 18112 Agama agama (Linnaeus. 1 758). Agamn colonvriim Daudin, 1802. Agamci ocdpifalis Gray, 1 827. Agama pkiicauda Peters. 1 877 GOniher { 1 S64a:480), BcN::age ( 1 866a :4 2, 1 895 a: 1 7). Bouleuger (1885:357), Ferreira (]900a:50. I‘>04;1!17, 1905:1 17, 1906:170), Angel (1923:159), Hcllmich {1957a:41. 1957b:50). Loveridge {1957:191). .4. congicd.A. slutcki, A. sp.. Accmihocercus cyanocepliaiits Genus Acattlliocercn:!! Fluinger, 18d,3 SfeHio 1“ Agtmti - Acmthocercun] «Crfco/to (Smith, 1849) Bocage (lS66a:43, I879b;95. I895ai22). Peters (1881 ; 147), Boulenger {1885:356, 358, 1905:1 10), Ferreira {1900a ;49. 1 903: 1 5), Angel { 1 923: 1 58). Schmidt ( 1 933:9), Parker ( 1 936: 1 32), Monard ( 1 937b:58, (vO), Themido (1941 :7), Laurent (I950a:l2, I964a:.38), Klausewit?. ( 1957:161), Fradc (1963:253), Mananas ( 1963:228), Branch (1998:218), Spawts (2010). Accitifhacemis ntinocepholits Agama [= /ft’Hfif/!f»cercifs] eyemogaster (Ruppel, 1835) Loveridge { 1 957: 1 95), Branch and Conradie (20 1 5:200). Accinr/iocemis cytinocephiilus Family TVPHLQPIDAE Merreni, 1S20 Genus A/roryphfops Broadley and Wallach, 2009 Typhhpx [= Afronphhps] ( 1 each, 1 S 1 9) Bocage { 1 866b:46, 187.3a:253, 1 895 a:66), Gunther ( I876b:678), Boulenger (1900a:50. 1905: 1 12). Parker (1936:120), Mertens ( 1937a. 1 1, 1938a:438), Monard ( 1 937b; 1 03. 1 04), Themido (1941 :9), Laurent { 1 950a:7), HeIJmich {1957a:70), Loveridge (1957:242). A. angoieiws. ,4. lineohtus Family I.EPTOTVPm.OPlIIAF StcjncRer, 1892 Genus Leptatyphhpx Fii/.inger, \ iNfimihiiiua Hedges, Adalsiein.sson, and Branch, 2009 Leplohphiops conjitnclci disfimti (Boulcnger, 1892) Bogert(1940;l3). L .iciili/rons MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 479 Leptonphhps emini (Bouicnger, 1890} Ijiurenlt 1964a: 91). kafuhi LeptonpMops mgi-icam (Schlege!, 1839) Bocage ( I866a;46, 1867b:224), Broadley and Watson (1976:490), McDiamiid et al, (I999;39). L. kafuhi^ L scuiifrom^ Namibiana lanfrons Family ViPEKIDAE Oppel. 181 1 Bhis CIray, 1842 Echidna [= Vipera = Bin's] r/?iiKicerc?j(Schlegel, 1855) Bocage (ia66a:53, 1887a: 191. 1895a: 149, 1896a: 113. 18976:21 1). Peters (I877a:618}, B. gabonica Family LAMPKOPHIIDAE Fitiinger, 1843 Pttcrs, IS57 Cahmeiaps [= Amhiyodipsas] wf/cotof (Reinhardl. 1843) Loveridge (1933:260), A. polyfcpis Smith , 1849 AparallactHS tineaUis (Peters. 1870) Botilenger(l895d:t73, 1896:259. 1915:217). Bocage (1897a: 201). de Witte and Laurent ( 1947:128), Laurent (I954a:45). A. capensis Cenus.4rm‘rM.^^rj! Smith, 1849 Atraciaspis aterrima Gunther, 1863 Bocage (1873a:223), Boulenger (1915:223), Chirio and LeBreton (2007:624). A. congica Airaclaspis corpaknta Peters, 1877 Bocage (1866a:49), A. irregularis Genus Boaedtm Diinicril, Bilimn and Dum^ril, 1854 Boaedon [= Lamprophis = Boodan] iinealus Dumeril. Rihrcn and Dum^ril, 1854 Gunther (1864a:480), Bocage ( I895a:78, 80), Boulenger (1893:332, 1896:616, 1905:1 12), Ferreira (18976:244, 1 900a:5 1 . 1 903: 10, 1 904: 11 4, 1906: 1 67), Monard ( 1 937b: 113. 1 17), Schmidt (1933:13), Loveridge ( I936a:22), Mertens ( 1 937a: 12, 1938a:439), Bogcrt ( 1 940:2 1 ). Themido (1941 :9), Laurent {1950a: 7. 1954a:43, I964a:93), Hellmich {I957a:71, 19576:60), Thys van den Aiidenaerde (1966:32), Wailach el al. (2014:96). Boaedon aagolensis, B. variegalus, B. fidiginasus Genus Lycophidioii Fitzinger, 1843 Lycaphidhn capense (Smith, 1 83 1 ) Hellmich ( 1 957a:7 1 ). Petera ( 1 877a:6 1 5, 1 88 1 : 1 49). Bocage (IS95a;8L 1 896a: 1 12), Ferreira (1904: 1 15, 1906: 167). Boulenger ( 1 893:616. 1 905; 1 1 2), Monard ( 1 9376: 1 1 7). Schmidt (1933:13), Bogert (1940:30), llellmicli (I957a:61. 19576:71). L heifmichi^ L multimacuhtum L)Tophidiofi semiannule Peters, IS54 Ferreira (I897b:243), Monard (1937b: 1 13). L. muttimucukaum (■enus Pn/emnH ,]an, 1858 i\fitxion [- Palemon] gabonensis Dumeril, 1856 Hellmich (1957a:72, 1957b;63). P. cotiaris Genus Prasyitnia Gray, 1849 Prosynifia meleagris (Reinhardt, 1843) Bocage ( 1 866a;47). P. ambigua 480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II Genus psaimmtphis Boic, 1S25 Psommophis brexirostris Peters. 1S3I Boulenger (1915:213), Monard (I9j7b:l33). Theinido { 1 94 1 ; 1 0), B randstatter ( 1 996:45 ). P. ieopurdinm Pxammophis efegam (Shaw, 1802) Bocage(1867b:226). P. mosxumhkiss Pxammopliis philtipsii (1 lallowell, 1844) Broadely (1977a:24), Branch and McCartney (1992:2). Brandstatter( 1996:55). Hughes (1999:64), Kelly etal. (2008), P. ssiosxmsbkiss Pxatwnopim xibUaitx (Linnaeus, 1758) Bocage ( 1 866a;48, 1895a: 11 4, I896a;l 13). Peters ( IS77a;6 15), Ferreira (1904:1 16). Boulenger( 1905:1 13. 1915:213), Schmidl (1933:14), Monard (I937b:l31 ). Loveridge ( I936a:38. 1957:279), Merlens ( 1 938a:44 1 ), Bogerl ( 1 940:70), Loveridge (1940:30), LiiuTeiit(l950a;9, 1954a;59, 1964a: 11 3). fldlmieh (1957b: 70), Thys van den Audenaerde (1966:34), Mananas (1973:196). P. kiiffai'dimis, P. zambkmsss, P. mosicsmbkiis Genus Psummitphytux Ht/.iiiRcr, 1845 Rhampimphis oxyrbyndms (Reinhardt. 1843) GDnther(1864a:480, 1895:89). Pxam»ti?pbylax aciiliss Family ELAPIDAE Boit, 1827 Genus tyetnif9sspis Schlegel, 1848 Dendruspis [= Dendroaspix] anjjw.f/ice/J.s (Smith, 1849) Bocagc(1866a:52, 1888:143, 1 895a; 1 40), Peters ( 1 877a;6 1 7, 1 888: 1 49), Boulenger ( 1 9 1 5:220), Schmidt ( 1 933: 1 5), Monard { 1 937b: 137), Bcigert ( 1 940:92). Frade ( 1 963:253). Deisdmaxpis polylepis Genus Nujn Lnurenti, 1768 ,V«ya/!«ye (Linnaeus, 1758) Bocage ( 1 866a;5 1 , 1 895a: 132), Peters ( 1 S77a:6 18), Loveridge (1957:291). jV. mehnolenco Family COLLBRIDAE Opfiel, 1811 Genus lymypePh \5 aRler, 1830 Dasypi'liis fascioksm Peicrs, 1 868 (= Dasyijetds medics Bianconi, 1859] Peters ( 1 877a:61 5). Bocage ( 1 895a; 1 06). D. scahra Genus .Smith, 1840 AlwefSitta [- Pbilotlninmus\ irse^ukiris (Lcacb. 1819) Giinlhcr ( 1 864a;480), Peters ( 1 877a;6 1 5, 1 88 1 : 149), Bocagc {i882b:6. I887b:205. I895a:85. 1896a:l 12). Ferreira (1903:10, 1906:167). Boulenger (1893:96), Parker (1936:125). Monard ( 1 937b: 1 1 4, 1 2 1 ), Mertens ( 1 938a:439). Bogert ( 1 940:53), Thcmldo (1941:10), Laurent (195 0a:8. I954a:47) Loveridge (1951:9, 1957:261), liellmich (19576:64), I'iizSimons (1962:144). Manafas (1973:191). P. irregss/aris. P. helerolepidoliss, P. bophgaxter MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 481 Taxonomic Index A A[lgyra] capensis 216 Ablabes Homeyeri 345 angolensis 345 Ablephams 26, 247, 248, 249 aeneus 247 cabindae 26, 247, 248 Cabindae 247 wahlbergii 249 Wahlbergii 249 Acanthocercus 7, 284, 285, 286 atricollis 284, 285, 286 cyanocephalus 284, 285, 286 cyanogaster 284, 286 sp 286 Acontias 7, 35, 46, 237, 238, 239, 241 elegans 241 jappi 237 kgalagadi 35, 46, 237, 238 kgalagadi 35 lineatus 238 occidentalis 238, 239 percivali 238, 239 ocidentalis 238 plumbeus 238 occidentalis 238 Afrana angolensis 149 Afrixalus 5, 85, 86, 87, 88 dorsalis 85 leptosomus 85, 87 regularis 85 equatorialis 87 fluvovittatus 87 leptosomus 87 fulvovittatus 85, 86, 87 leptosomus 86 leueostietus 87 osorioi 86, 87, 88 quadrivittatus 85, 86, 87 wittei 88 Afroablephams 247, 248, 249 Afroedura 6, 30, 177, 178 bogerti 30, 177, 178 ef. bogerti 177 karroiea 30, 177 bogerti 30, 177 Afrogeeko 6, 29, 41, 178, 179, 188 ansorgii 41, 178, 179 plumieaudus 188 Afronaja 371, 372, 373, 374, 375 mossambiea 371 nigrieineta 373 nigrieollis 374 Afrotyphlops 7, 41, 45, 292, 293, 294, 295, 296, 297, 298 angolensis 292 angolensis 292, 293 anomalus 41, 293, 294 lineolatus 294, 295 mueruso 296 punetatus 296 sehlegelii 297, 298 sehmidti 45, 298 Agama 7, 28, 31, 32, 41, 284, 286, 287, 288, 289, 290, 291, 292 aeuleata 287, 288, 289 aeuleata 287 agama 32, 290 mueosoensis 32 mueosoensis 290 anehietae 288, 289 anehietae 288 Anehietae 288 armata 287, 288 atrieollis 284, 286 atrieollis 284 eolonorum 28, 286, 289, 290, 291, 292 eongiea 289 eongiea 28, 289, 290 eyanoeephala 284 eyanogaster 284, 286 hispida 287, 288 aeuleata 287 mueosoensis 41, 290 planieeps 31, 289, 290, 291, 292 planieeps 289 sehaeki31,291,292 sehaeki 291, 292 sp 290 Agamidae 7, 37, 38, 41, 284 Ahaetulla 28, 389, 391, 393, 394, 396, 397 boeagii 28 Boeagii 397 dorsalis 391 graeillima 394 heterolepidota 393 hoplogaster 394 irregularis 389, 394 482 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II nitida 395 Alopecion variegatum 328 Amblyodipsas 7, 320, 321, 359, 360 polylepis 320, 321 hildebrandtii 321 polylepis 320, 321 AmietiaS, 138, 149, 150, 151 angolensis 138, 149, 151 cf. angolensis 149 Amietophrynus 69, 70, 71, 72, 74, 75, 77 buchneri 69 funereus 70 garmani 7 1 gutturalis 72 lemairii 74 maculatus 75 power! 74 regularis 77 Amnirana 6, 41, 45, 158, 159, 160, 161, 162 albolabris 158 darling! 158, 159 lemairei 158, 160 lepus 160, 161, 162 parkeriana 41, 45, 161 Amphiophis 345, 346 Amphisbaena 31, 212, 213 ambuellensis 31, 21 2, 213 quadrifrons 212 Amphisbaenidae 6, 37, 38, 41, 205 Amplorhinus 333 noto taenia 333 Amy da 174 triunguis 174 triunguis 174 Anelytrops elegans 241 Angolosaums 33, 234, 235 skoogi 234, 235 Aparallactus 7, 321, 322 bocagii 321, 322, 323 Bocagii 321, 322 capensis 321, 322, 323 bocagii 322, 323 capensis 322 punctatolineatus 322 guentheri 321, 322, 323 Guenther! 321 punctatolineatus 322, 323 punctolineatus 321 punctulolineatus 32 1 Aporosaura 218, 219 anchietae 218 Arthroleptidae 5, 37, 38, 41, 117 Arthroleptis 5, 29, 41, 45, 117, 118, 119, 120, 121, 146, 148 boulengeri 119 carquejai 41, 45, 117, 118 lameerei 45, 118, 119 minutus 146 parvulus 29, 146 plicatus 148 spinalis 119 stenodactylus 120 variabilis 118 xenochims 29, 120, 121 Aspidelaps 8, 30, 316, 361, 362 (Sepedon) Lichtensteinii 316 lubricus 30, 361, 362 cowlesi 30, 361, 362 infuscatus 362 Atheris 7, 308 Lucani 308 squamiger 308 squamigera 308 squamigera 308 Atractaspis 7, 28, 45, 48, 320, 323, 324, 325, 326, 327 angolensis 327 aterrima 324 bibroni 323 Bibroni 323 bibronii 323, 324 bibronii 323, 324 rostrata 323, 324 boulengeri 324 mixta 324 Boulengeri 324 congica 28, 324, 325 congica 325 orientalis 325 corpulentus 326 heterochilus 326, 327 Hildebrandtii 320 irregularis 45, 326 irregularis 326 parkeri 326 reticulata 48, 325, 326, 327 heterochilus 48, 326, 327 rostrata 324 Aubria 6, 46, 151, 152 masako 151 sp46, 151, 152 subsigillata 151 B Bids 7, 26, 41, 46, 47, 53, 309, 310, 311, 312, 313, MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 483 314 arietans 47, 53, 309, 310 arietans 309, 310 somalica 310 caudalis 310, 311 caudalis 311 gabonica 311, 312 gabonica 312 Gabonica 311 heraldica 41, 46, 312, 313, 314 hoserae 314 lachesis 309 nasicomis 313, 314 peringueyi 46, 312, 313, 314, 315 rhinoceros 311, 312 Boaedon 7, 41, 327, 328, 329, 330, 331, 388 angolensis 41, 327, 328, 330 capensis 330 cf. angolensis 327 fuliginosus 328, 329, 330, 331 fuliginosus 328 lineatum 329 lineatus 327, 328, 329, 330 olivaceus 330, 331 quadrilinaetus 329 quadrilineatum 329 variegatum 41 variegatus 328, 329, 330 Boidae 7, 37, 38, 307 Boiga 405 blandingi 405 blandingii 405 pulverulenta 405 Boodon 327, 328, 329, 330 lineatus 327 angolensis 327 olivaceus 330 Bothrophthalmus 7, 331 lineatus 331 lineatus 331 Boulengerina 35, 369, 370, 371, 372 annulata 369 annulata 369 melanoleuca 370 multifasciata 372 Brachymerus bifasciatus 80 Breviceps 5, 46, 81, 82 adspersus 81, 82 gibbosus 8 1 gibosus 81 mossambicus 81, 82 mossambicus-adspersus 8 1 mossambicus/adspersus 8 1 power! 82 sp46, 81 Brevicipitidae 5, 37, 38, 81 Bucephalus 381, 383 capensis 381 typus 381, 383 viridis 383 Bufo 28, 31, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78 benguelensis 70 benguellensis 7 1 buchneri 69 Buchneri 69 carens 69 cinereus 75 decorsei 70 dombensis 67 funereus 69, 70, 71, 75, 76 funereus 70 garmani 7 1 , 74 grandisonae 67 guineensis 77 gutturalis 71, 72 guturallis 78 kavangensis 68 lemairei 74 lemairii 73, 74 maculatus 75, 78 melanopleura 66 mocquardi 67 Mocquardi 66 pantherinus 77 power! 74 pseudogarmani 75 pusillus 76 pusilus 75 regularis 31, 71, 72, 73, 74, 75, 77, 78 gutturalis 72, 73 humbensis 31, 71, 72 power! 74 pusillus 75 regularis 72, 77 spinosus 72, 73 taitanus 67 Bufonidae 5, 37, 38, 41, 66 c Calabaria 7, 45, 307 reinhardtii 45, 307 Reinhardtii 307 Calamelaps 320 484 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II polylepis 320 unicolor 320 polylepis 320 Cassiniopsis 115 kuvangensi 115 kuvangensis 115 Cassionopsis (see as Kassina) 3 1 kuvangensis 3 1 Catia afrieana 237 Causus 7, 29, 35, 315, 316, 317, 318, 319, 320 bilineatus 29, 315, 316 bilineatus 315 lineatus 316 ef. rasmusseni 319, 320 liehtensteinii 316 lineatus 316 maeulatus 316, 317, 320 rasmusseni 35, 320 resimus 317, 318 angolensis 317 rhombeatus 315, 316, 317, 318, 319, 320 bilineatus 315 rostratus 317 Cerastes 310, 356 eaudalis 310 tritaeniatus 356 tritaeniatus 356 Chamaeleo 7, 279, 280, 281, 282, 283, 284 anehietae 279 Capellii 279 dilepis 279, 280, 281,283 dilepis 280 quilensis 279, 280, 281 roperi 28 1 graeilis 280, 282, 283 etiennei 282, 283 graeilis 282 namaquensis 283 oweni 284 Oweni 284 quilensis 280 senegalensis 279, 282, 283 senegalensis 282 tubereulifems 283 Chamaeleon 279, 282, 283 anehietae 279 Anehietae 279 dilepis 279 Dilepis 279 etiennei 282 graeilis 282 namaquensis 283 parvilobus 279 quilensis 279 senegalensis 282 Chamaeleonidae 7, 38, 279 Chamaelyeus 8, 376, 377 parkeri 376, 377 Chamaesaura 6, 225, 226, 227 anguina 225, 226 anguina 226 oligopholis 225, 226 maerolepis 226, 227 miopropus 226 miopropis 227 miopropus 226, 227 Charina reinhardtii 307 Chelonil 6, 162 Chiromantis 6, 160, 162 lepus 160 xerampelina 162 Chlorophis 389, 391, 393, 394, 395, 396, 397 (Philothamnus) irregularis 389 angolensis 389 earinatus 391 heterodermus 391, 393 earinatus 391 heterolepidotus 393, 397 hoplogaster 394 irregularis 389, 394 shiranus 394 nitidus 395 loveridgei 395 omatus 396 Chondrodaetylus 6, 46, 179, 180, 181, 182 angulifer 180, 181, 182 bibronii 181, 182 ef. pulitzerae 180, 181 fitzsimonsi 179, 180, 182 laevigatus 181, 182 laevigatus 182 turner! 182 pulitzerae 46, 180, 181 turner! 181, 182 Cinixys 170, 171 belliana 170 erosa 171 Clotho arietans 309 Cobra 309 laehesis 309 Colopus 198, 199 wahlbergii 198, 199 wahlbergii 198 Wahlbergii 198 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 485 Coluber 24, 306, 313, 357, 379, 385, 389 canus 357 Nasicomis 313 palmarum 379 scaber 379 Sebae 306 zebrinus 389 Colubridae 8, 37, 38, 376 Corachodichus 120 stenodactylus 120 stenodactylus 120 Cordylidae 6, 37, 38, 41, 225 Cordylosaums 6, 214, 229, 230 subtessellatus 214, 229, 230 trivittatus 229 Cordylus 6, 30, 36, 41, 46, 47, 227, 228, 229 angolensis 41, 46, 227, 228 cordylus 227, 229 angolensis 227 machadoi 47, 228 namakuiyus 30, 36, 41, 47, 229 tropidostemum 227 tropidostemum 227 vittifer 228 machadoi 228 Coronella 377, 378, 408, 409 (Mizodon) olivacea 409 hotamboeia 377, 378 olivacea 408, 409 Crocodilus 174, 176 cataphractus 176 cataphractus 176 niloticus 174 vulgaris 174 Crocodylidae 6, 37, 38, 174 Crocodylus 6, 47, 48, 53, 174, 175, 176, 177 cataphractus 176 frontatus 177 niloticus 47, 48, 174, 175 chamses 174, 175 suchus 48, 175 tetraspis 177 Crotaphopeltis 8, 377, 378, 381 hotamboeia 377, 378 hotamboeia 377 rufescens 377 semiannulatus 377 shrevei 381 Crumenifera pusilla 111 Cryptoblepharus walbergii [sic] 249 Cryptopodus Aubryi 173 Cryptothylax 5, 45, 88 greshoffii 45, 88 Ctenotus australis 266 Cyclo-derma aubryi 45 Cycloderma 6, 173 aubryi 173 Aubryi 173 Cystignathus 26, 117, 122, 123 Bocagii 26, 122, 123 Senegalensis 117 D Dactylethra 61, 62 laevis 62 Muelleri 61 mulleri 62 Multeri 62 Dalophia 6, 25, 31, 34, 46, 205, 206, 207, 211 angolensis 34, 205, 206, 207 colobura 207 ellenbergeri 205, 206, 207 grand 207 kaynuamarum 207 kuanyamarum 207 mossambica 207 mossambicus 207 pistillum31,46, 205, 206, 207 transvaalensis 207 welwitschii 25, 211 Welwitschii 211 Dasia olivacea 269 Dasypeltis 8, 24, 379, 380 fasciolata 380 inomata 380 medici 380 palmarum 379 scabra 379, 380 inomatum 379 inomatus 379 palmarum 379 scabra 379 Dendraspis 25, 362, 363 angusticeps 363 neglectus 362 polylepis 363 welwitschii 25 Welwitschii 362 Dendroaspis 8, 25, 362, 363, 364 angusticeps 363, 364 jamesoni 25, 362, 363 jamesoni 362 jamesonii 362 jamesonii 362 486 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II neglectus 363 polylepis 363, 364 polylepis 364 Dendrophis 387, 397, 403 (Philothamnus) semivariegata 397 flavigularis 403 Dendrophis smaragdina 387 Dicroglossidae 6, 37, 38, 157 Dicroglossus occipitalis 157 Dipsadoboa 8, 381 shrevei 381 Dipsas 405 Blandingii 405 pulvemlenta 405 Dispholidus 8, 381, 382, 383, 384 (Bucephalus) typus 381 punctatus 384 typus 381, 382, 383, 384 kivuensis 383 occidentalis 382 punctatus 382, 383, 384 typus 383, 384 viridis 383, 384 Distichums Maculatus 316 Dromophis 7, 332 lineatus 332 Dryiophis 401, 402 Kirtlandii 401, 402 Oatesi 402 oatesii 40 1 Dryophylax lineatus 332 Dumerilia Bayonii 25 1 E Echidna 309, 311 arietans 309 rhinoceros 311 Echis squamigera 308 Elapechis guentheri 365, 366 Elaphis (Bothrophthalmus) lineatus 331 Elapidae 8, 37, 38, 361 Elaps 326, 362 irregularis 326 jamesoni 362 jamesonii 362 Elapsoidea 8, 365, 366, 367 decosteri 366, 367 huilensis 367 guentheri 365 guentherii 365 giintheri 365, 366 guntheri 365 semiannulata 366 Guntheri 365, 366 Guntherii 365, 366 Guntherii 365 semi-annulata 366, 367 semiannulata 365, 366, 367 moebiusi 367 semiannulata 366, 367 sundevalli 365 semiannulata 365 sundevallii 365, 366 guntheri 365, 366 guntherii 365 semiannulata 366 Empagusa 275 Emys castanea 165 Engystoma marmoratum 83 Eremias 213, 222, 223, 224 benguelensis 222, 223 benguellensis 222, 223 lugubris 213 namaquensis 222, 223 sp 222 undata 222, 223, 224 undata 222, 223, 224 Eryx reinhardtii 307 Eumeces 239, 246 reticulatus 246 Eumecia 7, 41, 239, 240, 245 anchieta 239 anchietae 41, 239, 240 anchietae 239 major 239, 240 Anchietae 239 Eumices [sic] (Riopa) sunderallii 245 Euphlyctis occipitalis 157 Euprepes 26, 28, 244, 245, 247, 252, 253, 254, 256, 257, 258, 259, 260, 262, 263, 264, 265, 266, 267, 268, 269, 270 (Eupr.) Perrotetii 263 (Euprepis) striatus 268 spilogaster 268 (Mabuia) breviceps 247 (Mabuia) megalums 264 acutilabris 252 affinis 259, 260 anchietae 26, 264 Anchietae 263 angolensis 254 australis 266 Bayonii 256 binotata 257 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 487 binotatus 257, 258 Blandingii 253 damaranus 260 Ivensi 244 ivensii 26 lacertiformis 262 maculialbris 263 maculilabris 263 notabilis 263, 264 occidentalis 266 olivaceus 269 Oliveiri 254 albo-punctatus 254 Olivieri 256 albo-punctatus 256 perottetii 264 petersi 258, 259 Petersi 258 petersii 259 punctatissimus 270 punctulatus 267 quinquetaeniata 259 quinquetaeniatus 258 sulcatus 269 varia 265 vittatus 266 Wahlbergii 270 Euprepis 254, 256, 258, 265 affinis 254 angolensis 256 blandingii 254 quinquetaeniata 258 raddoni 254 Eutropis dissimilis 258, 259 F Feylinia 7, 45, 240, 241, 242 currori 240, 241, 242 elegans 241 grandisquamis 242 Currori 240, 241, 242 elegans 241, 242 grandisquamis 242 grandisquamis 45, 242 polylepis 240 G Gastropyxis smaragdina 387 Gecko 186 Mabouia 186 Gekkonidae 6, 37, 38, 41, 177 Geochelone 172, 173 pardalis 172, 173 babcocki 172, 173 pardalis 173 Gerrhosauridae 6, 37, 38, 41, 229, 235 Gerrhosaurus 7, 29, 32, 33, 41, 46, 229, 230, 231, 232, 233, 234, 235, 236 auritus 230, 231, 232 bulsi 230, 231,232 cf. nigrolineatus 233 flavigularis 233 nigrolineatus 233 intermedius 232, 234 multilineatus 41, 230, 231, 232, 234 nigro-lineatus 233 nigrolineatus 32, 230, 231, 232, 233, 234 ahlefeldti 32, 233, 234 nigrolineatus 231, 233 robustus 235, 236 skoogi 29, 33, 46, 234, 235 validus 235, 236 maltzahni 235 Glaniolestes omatus 384 Glauconia 299, 300, 301, 302, 303 kafubi 299 labialis 301 latifrons 302 rostrata 303 scutifrons 300, 302 Godionotus brussauxi 332 Gonionotophis 7, 332, 333, 335, 336, 340, 341 brussauxi 332, 333 capensis 335 poensis 340 vemayi 336 Grayia 8, 45, 384, 385, 386 caesar 45, 384 ornata 384 silurophaga 385, 386 smithi 385 smithii 384, 385, 386 Smithii 386 smythii 385, 386 Smythii 385, 386 tholloni 386 Tholloni 386 triangularis 385 Gymnopus aegyptiacus 173 H Hapsidophrys 8, 387 smaragdina 387 488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II smaragdinus 387 Helicops 406, 407 bangweolicus 407 bicolor 406 Heliobolus 6, 213 lugubris 213, 214 Hemerophis 389 zebrinus 389 Hemidactylus 6, 41, 47, 182, 183, 184, 185, 186, 187, 190 angulatus 182, 183, 187 ansorgii 187, 188 bayonii41, 183, 184 Bayonii 183 benguellensis 41, 184 bocagei 185 bocagii 185 brooki 182 brookii 182, 183, 184 angulatus 182, 183, 184 capensis 190 cf. muriceus 187 intestinalis 187, 188 longicephalus 185, 186, 187, 188 mabouia47, 184, 185, 186, 187 mabouia 186 muriceus 187, 188 platycephalus 185, 186 Hemirhagerrhis 7, 333 nototaenia 333, 334 viperina 333, 334 Hemisotidae 5, 37, 38, 83 Hemisus 5, 83, 84 guineensis 83, 84 microps 83, 84 guttatum 83 marmoratum 83, 84 marmoratus 83, 84 (?) angolensis 84 guineensis 83 sudanense 84 Heterolepis 335, 340 bicarinatus 340 capensis 335 poensis 340 Heterophis resimus 317 Hildebrandtia 5, 26, 41, 46, 128, 129 angolensis 128, 129 myotympanum 128 omata46, 128, 129 omata 129 omatissima 128 ornatissima 41, 128, 129 Holaspis 6, 214, 230 guentheri 214, 230 guentheri 214 Holuropholis olivaceus 330 Hoplobatrachus 6, 26, 157 occipitalis 26, 157 Hormonotus 8, 387, 388 modestus 387, 388 HylaAubryi 122 Hylambates 88, 121, 122, 123, 124, 125, 126 anchietae 121 angolensis 122, 123 Aubryi 122 bocagei 123 leucopunctata 123, 124 bocagii 123 cynnamomeus 124 greshofFii 88 Greshoffii 88 marginatus 125 notatus 126 viridis 126 Hylarana 158, 159, 160, 161 albolabris 158, 160 cf. darlingii 159 darling! 159 lemairei 160 parkeriana 161 Hylarthroleptis 143 brevipalmatus 143 graueri 143 Hyperoliidae 5, 37, 38, 41, 85 Hyperolius 5, 26, 28, 31, 35, 41, 45, 46, 55, 85, 86, 87, 88, 89, 90, 91, 92, 94, 95, 96, 97, 98, 99, 100, 101, 102, 103, 104, 105, 106, 107, 108, 109, 110, 111, 112, 113, 114, 115, 148 (Hyperolius) [subgenus] 92, 94, 97, 98, 100, 102, 104, 105, 106, 108, 109, 110, 111, 112, 113, 114, 115 acuticeps 107 adspersus 28, 89, 90, 96 angolanus 109, 110 angolensis 26, 28, 31, 90, 91, 92, 94, 96, 103, 111 argus 98 benguelensis-nasutus complex 95 benguellensis 89, 90, 94, 95, 96, 105, 107, 108 bicolor 41, 96 bocagei 26, 96, 97, 98, 105 cf adspersus 89 cf steindachneri 115 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 489 chelaensis 35, 41, 46, 98 cinctiventris 97, 98 cinereus 31, 41, 98, 99, 100, 113 cinnamomeo-ventris 100, 101 cinnamomeoventris 100 cinnamomeoventris 100, 101, 102, 110 citrinus 91 concolor 101, 102 dartevellei 102 decoratus 91 dorsalis 85 erythromelanus 31, 91, 92 fasciatus 109 ferreirai 109 fulvovittata 88 fulvovittatus 85, 88 fuscigula 41, 102 glandicolor 103 granulatus 89, 90 graueri 91 gularis 41, 103 huillensis 90, 91 insignis 90, 91, 94 kivuensis 104, 105, 112 kivuensis 104 multifasciatus 104 langi 105 leptosomus 28, 87 quadrivittatus 87 lucani 108, 109 Lucani 108 machadoi 114, 115 maestus 41, 105, 106 Maestus 105 marmoratus 26, 90, 91, 94, 96, 103, 105, 106, no. 111 albofasciatus 91 albomfus 9 1 angolensis 26, 90, 91, 110 huillensis 91 insignis 91, 96 marungaensis 91 parallelus 91 vermiculatus 91 marungaensis 91 microps 111, 112 microstictus 91 microtictus 91 modestus 101 multifasciatus 104, 113 nasutus 26, 89, 90, 94, 95, 96, 102, 106, 107, 108 adspersus 89, 102 nasutus 106 nasutus complex 95, 96, 102 nitidulus 108 ocellatus 45, 108, 109 osorioi 86 oxyrhynchus 94, 96 parallelus 90, 91, 92, 94 albomfus 91 huillensis 92 insignis 91 plicifems 91 toulsoni 91 parallelus-marginatus 92, 94 parallelus-marginatus subgroup 92, 96 platycephalus 105 langi 105 platyceps 109, 110 angolanus 109 angolensis 110 plicatus 148 poiii no protchei 41, 111 punctulata 107 punctulatus 106 pusillus 111, 112 quinquevittatus 104, 105, 112, 113 quinquevittatus 112 raymondi 35, 41, 100, 113 rhizophilus 41, 113, 114 Rhizophilus 113 seabrai 97 steindachneri 114, 115 steindachneri 114 Toulsinii 9 1 toulsonii 26 Toulsonii 90 tristis 100 vermiculatus 28, 90, 91 vilhenai 41, 115 viridiflavus 98, 103 viridis 107 Hypoptophis 7, 334, 335 wilsoni 334 katangae 334 wilsonii 334, 335 I Ichnotropis 6, 32, 41, 45, 214, 215, 216, 217, 218, 220 bivittata 214, 215, 216, 217, 218 bivittata 214, 215 pallida 216 bivittatus 215, 216 490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II capensis 41, 215, 216, 217 bivittata 215 capensis 216, 217 overlaeti 41, 217 dumerilii 216 Dumerilii 214 longipes 215 microlepidota 41, 45, 218 overlaeti 45, 217, 218 spp 216 squamulosa 220 Ixalus eoneolor 101 K Kaokogeeko 198 vanzyli 198 Kassina 5, 31, 34, 46, 115, 116, 117 angeli 117 ef. maeulosa 116 kuvangensis 46, 115, 116 maeulosa 34, 116 senegalensis 117 angeli 117 mierops 117 Kinixys 6, 51, 52, 169, 170, 171 belliana 52, 169, 170, 171 spekii 171 Belliana 170 belliana 170 ef. belliana 170 erosa 171 spekii 169, 170, 171 Kolekanos 6, 41, 46, 188 plumieaudus 41, 188 L Laeerta213,277, 278 lugubris 213 monitor 277, 278 nilotiea 277, 278 tesselata 221 Laeertidae 6, 37, 38, 41, 213 Lampreremias lugubris 213 Lamprophiidae 7, 37, 38, 41, 320 Lamprophis 329, 330, 331, 387 fuliginosus 329 modestus 387 olivaeeus 330 Lepidostemon (Phraetogonus) Anehietae 207 Lepidothyris 7, 45, 242, 243 femandi 243 hinkeli 45, 242, 243 joei 45, 242, 243 Lepthophis heterolepidota 393 Leptodira 377 hotamboeia 377 mfeseens 377 Leptopelis 5, 26, 41, 45, 121, 122, 123, 124, 125, 126, 127 anehietae 45, 121, 122, 125 angolensis 41, 123, 124 aubryi 122, 125 boeagei 123, 124 boeagii 26, 122, 124, 125 ef anehietae 121 ef boeagii 123 eynnamomeus 124 jordani 41, 45, 125 marginatus 41, 125 nordequatorialis 122 notatus 126 oryi 122 parboeagii 123, 124 tessmanni 126 viridis 124, 126, 127 einnamomeus 124 Leptophis 26, 391, 392, 394, 402 Chenoni 392, 394 dorsalis 26, 391, 392 Leptosiaphos 7, 243, 247 (Laeertaspis) brevieeps 247 dewittei 243 Leptotyphlopidae 7, 37, 38, 41, 299 Leptotyphlops 7, 299, 300, 301, 302, 303 eonjuneta 300 distant! 300 emini 299, 300 emini 299, 300 kafubi 299, 300 labialis 301 latifrons 302 nigrieans 300 nigrieans 300 oeeidentalis 302 rostrata 302 rostratus 303 seutifrons 300, 301, 302, 303 seutifrons 300 seutifrons/ineognitus/eonjunetus eomplex 301 Letheobia 7, 299 praeoeularis 299 Limaformosa 7, 335, 336 eapensis 335, 336 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 491 vemayi 336 Limnodytes albolabris 158 Limnophis 8, 26, 406, 407, 408 bangweolicus 407, 408 bicolor 26, 406, 407 bangweolicus 407 bicolor 406 Lubuya 7, 26, 239, 244, 245 ivensii 244 Lycodon 329 fuliginosus 329 Lycodonomorphus 8, 388 subtaeniatus 388 subtaeniatus 388 upembae 388 Lycophidion 7, 28, 48, 336, 337, 338, 339, 340 capense 336, 338, 339, 340 capense 336, 338 multimaculatum 338 omatum 339, 340 hellmichi 48, 336 Horstockii 338 laterale 337 meleagre 28, 337, 338 multimaculatum 338, 339 omatum 339, 340 semiannulis 338 Lycophidium 337, 338 Capense 338 meleagris 337 multimaculata 338 Lygodactylus 6, 30, 188, 189, 190, 191, 192 angolensis 188, 189, 191 bradfieldi 189, 190 capensis 189, 190, 191 capensis 190 chobiensis 191 grotei 191 laurae 30, 188, 189 lawrencei 191, 192 picturatus 191 chobiensis 191 Lygosoma 239, 242, 243, 244, 245, 246, 247 (Eumecia) Anchietae 239 (Hinulia) compressicauda 244 (Panaspis) breviceps 247 (Siaphos) compressicauda 243 anchietae 239 dewittei 243 femandi 242 Ivensii 244 modesta 246 modesta 246 modestum 246 sundevalii 246 Lygosominae 248 M Mabouia 28 Mabuia 29, 244, 253, 254, 256, 258, 259, 263, 265, 266, 267, 269, 270 ansorgii 29 Ansorgii 269 bayonii 256 Bayonii 256 bocagi 269 ansorgei 269 bocagii 258, 259 chimbana 259 ivensii 244 laevis 29, 263 maculilabris 263 occidentalis 266 Petersi 258 punctulata 267 quinquetaniata 258 raddonii 253 Raddonii 253 striata 256, 265, 270 angolensis 256, 265 sulcata 269 ansorgii 269 varia 254 Mabuya 31, 244, 245, 252, 253, 254, 256, 257, 258, 259, 260, 261, 262, 263, 264, 265, 266, 267, 268, 269, 270 acutilabis 252 afFinis 254 angolensis 265 bayonii 256, 257 bayonii 256 huilensis 256, 257 binotata 257 bocagei 258 bocagii 258 bocagii 258 chimbana 259, 260 hoeschi 261 ivensi 244, 245 septemlineata 244, 245 ivensii 244, 245 lacertiformis 262 laevis 263 longiloba 267 492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II longiloba 267 maculilabris 263 maculilabris 263 megalura 264, 265 occidentalis 266 punctulata 267 quinquetaeniata 257, 258 binotata 257 quinquetaeniata 258 radoni 254 spilogaster 268 striata 31, 259, 260, 265, 268, 270 angolensis 31, 259, 265 ehimbana 259 spilogaster 268 striata 270 wahlbergi 270 suleata 269 ansorgii 269 varia 254 varia 254 variegata 267 punetulata 267 Maerophis omatus 384 Maltzahni 235 Matobosaums 7, 235, 236 maltzahni 235, 236 Meeistops 6, 48, 176 eataphraetus 48, 176 ef. eataphraetus 176 Megalixalus 87, 116 fomasinii 87 eongieus 87 maeulosus 116 Megalixus 87, 88 leptosomus 87 quadrivittatus 87 wittei 88 Megatyphlops 293, 294, 296, 297, 298 anomalus 293, 294 muemso 296, 297 sehlegelii 297 sehlegelli 297 Mehelya 7, 30, 332, 335, 336, 340, 341 eapensis 335 eapensis 335 poensis 340, 341 poensis 340 vemayi 30, 336 Melanoseps 7, 245 oeeidentalis 245 zairensis 245 Meroles 6, 46,218,219, 220 anehietae 46, 218, 219, 220 retieulatus 219 squamulosus 220 Mertensophryne 5, 46, 66, 67 aff. moequardi 66 melanopleura 66 moequardi 46, 66, 67 Mesalina benguelensis 222 Miehellia katangae 335 Mierohylidae 5, 37, 38, 79 Mierosoma 28, 341 eollare 28, 341 Miodon 341 eollaris 341 gabonensis 341 Mizodon 408, 409 olivaeeus 408 Moehlus 7, 242, 243, 245, 246 afer 245 femandi 242 sundevalii 246 sundevalii 246 sundevallii 245 Monitor 277 nilotieus 277 saums 277 Monopeltis 6, 25, 31, 34, 41, 46, 205, 206, 207, 208, 209,210,211 anehietae 31, 46, 207, 208, 209 Anehietae 207 eapensis 208, 209 eapensis 208 devisi 31, 207, 208 ellenbergeri 205 Ellenbergeri 206 giganteus 211 grant! 31, 205, 206 kuanyamamm 31, 206 transvaalensis 206 tmasvaalensis 205 infuseata 46, 208, 209 luandae 34, 41, 209 okavangensis 31, 207, 208 perplexus 34, 41, 209 pistillum 206 sphenorhynehus 208 vanderysti 210 vilhenai 210 Vanderysti 210 welwitsehii 41, 205, 211 Welwitsehii 211 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 493 Mopaneveldophis 8, 389 zebrinus 389 N Naia 372, 374, 376 goldii 376 Goldii 376 multifasciata 372 Naja 8, 30, 35, 53, 368, 369, 370, 371, 372, 373, 374, 375, 376 (Afronaja) mossambica 371 (Afronaja) nigricincta 372, 373 (Afronaja) nigricollis 374 (Boulengerina) annulata 369 (Boulengerina) melanoleuca 370 (Boulengerina) multifaseiata 372 (Uraeus) anehietae 368 anehietae 368, 369 Anehietae 368 annulata 35, 369, 370 annulifera 369 anulifera 368 faseiata 372 goldii 376 haje 368, 369, 370 haje 370 melanoleuea 370 melanoleuea 369, 370, 371, 375 melanoleuea 370 subfulva 375 mossambiea 371, 372, 373 mossambiea 371 nigrieineta 373 multifaseiata 372 nigrieineta 372, 373 nigrieineta 373 nigrieollis 30, 372, 373, 374, 375 faseiata 372, 373 melanoleuea 373, 374 nigrieineta 30, 373 nigrieinetus 372, 373 nigrieollis 374 oeeidentalis 373, 374 woodi 373 subfulva 370, 371, 375 Namibiana 7, 41, 46, 48, 301, 302, 303 labialis 301 latifrons 301, 302 rostrata 41, 46, 48, 303 Natrieidae 8, 37, 38, 406 Natrieiteres 8, 28, 408, 409 bipostoeularis 408, 409 olivaeea 28, 408, 409 bipostoeularis 408 ulugurensis 409 olivaeeus 408, 409 olivaeea 409 olivaeeus 409 uluguruensis 408 pembana 409 sylvatiea 409 variegata 408, 409 bipostoeularis 408 Matrix 361, 362 lubriea 361, 362 Neusterophis 28, 408, 409 atratus 28 olivaeeus 409 olivaeeus 409 Neusterophus atratus 409 Nueras 6,41,48, 221,222 aff. tessellata 221, 222 holubi 222 intertexta 221 holubi 221 omata 221 sealaris 41, 48, 221 taeniolata 222 tessellata 221, 222 livida 222 taeniolata 221 tessellata 222 o Onyehoeephalus 292, 293, 294, 296, 297, 298 angolensis 292, 293 anomalus 293 Kraussii 292 liberiensis 294 lineolatus 294 mueruso 296 petersii 298 Petersii 297 Sehlegelii 297 Oophilositum parkeri 376 Ophirhina Anehietae 357 Osteoblepharon 177 Osteolaemus 6, 45, 48, 176, 177 osbomi 177 tetraspis 45, 48, 176, 177 osbomi 177 494 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II P Pachydactylus 6, 30, 41, 46, 179, 180, 181, 182, 192, 193, 194, 195, 196, 197, 198, 199 amoenoides 195, 196 angolensis 41, 192, 193, 198 bibronii 30, 180, 181, 182 pulitzerae 30, 180, 181 tumeri 180, 181 capensis 180 caraculicus 193 cf. oreophilus 194 fitzsimonsi 179 geitje 195 laevigatus 179, 180, 181 fitzsimonsi 179, 180 laevigatus 180, 181 pulitzerae 180 tessellatus 180 oeellatus 194, 195 oreophilus 194 paraseutatus 192 pulitzerae 182 punetatus 194, 195, 196, 197 bmnnthaleri 194 punetatus 194 rangei 46, 196, 198 seherzi 196, 197 seutatus 30, 192, 193, 197, 198 angolensis 30, 192, 193, 198 seutatus 193 serval 194, 195 stellatus 181, 182 tessellatus 179, 180 vanzyli 196, 198 wahlbergii 198 Paehyrhynehus Anehietae 218 Palmatogeeko 196, 198 rangei 196 vanzyli 198 Panaspis 7, 26, 35, 48, 247, 248, 249, 250 aff wahlbergii 249 anaeus 248 brevieeps 247 eabindae 48, 247, 248 maeulieollis 35, 248, 249, 250 wahlbergii 249, 250 Paranaja 369, 372 multifaseiata 369, 372 Paratetradaetylus Ellenbergeri 236 Pedioplanis 6, 35, 41, 46, 222, 223, 224, 225 benguellensis 222, 223 haaekei 35, 41, 223, 224, 225 huntelyi 224, 225 huntleyi 35, 41, 224, 225 namaquensis 222, 223 undata 222, 224, 225 Pelomedusa6, 51, 162, 163, 164 galeata 162 submfa 162, 163, 164 submfa 162 Pelomedusidae 6, 37, 38, 162 Pelusios 6, 45, 51, 52, 164, 165, 166, 167, 168 adansoni 167 beehuanieus 164 beehuanieus 164 eastaneus 52, 165, 166, 168 eastaneus 165 ehapini 165 rhodesianus 168 seyehellensis 165 ehapini 165, 166 derbianus 165, 166, 167 gabonensis 45, 166, 167 nanus 166, 167 nigrieans 167, 168 rhodesianus 167 rhodesianus 165, 166, 167, 168 sinuatus 167 sinuatus 167 subniger 164, 165, 167, 168 Pentonyx 162, 166 gabonensis 166 Gehafie 162 Philothamnus 8, 26, 28, 353, 389, 390, 391, 392, 393, 394, 395, 396, 397, 398 angolensis 389, 390, 391, 393, 395 earinatus 391 dorsalis 391, 392 Guntheri 389 heterodermus 393 heterodermus 393 heterolepidotus 393, 394 hoplogaster 390, 391, 394, 395 irregularis 389, 390, 391, 392, 393, 394, 395, 396, 397 angolensis 389 irregularis 389, 393, 394 omatus 396 natalensis 394 nitidus 395, 396 loveridgei 395, 396 omatus 393, 394, 395, 396, 397 semivariegatus 391, 392, 393, 395, 397, 398 dorsalis 391 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 495 semivariegatus 398 Smithii 397 sp398 thomensis 396 variegatus 28 Phrynobatrachidae 5, 37, 38, 41, 143 Phrynobatrachus 5, 29, 41, 143, 144, 145, 146, 147, 148, 149 auritus 149 brevipalmatus 41, 143 cf. natalensis 147 cf. parvulus 146 cryptotis 144, 146 mababiensis 144, 145, 146 minutus 144, 145, 146 natalensis 147, 148 parvulus 144, 145, 146 plicatus 148, 149 Pbrynomantis 5, 46, 79, 80, 81 affinis 46, 79 annectens 79 bifasciata 80 bifasciatus 79, 80, 81 bifasciatus 80 microps 81 Pbrynomems 79, 80 affinis 79 annectens 79 Pbyllodactylus (see also Afrogecko) 29, 178 ansorgii 29 Ansorgii 178 Pipidae 5, 37, 38, 59 Polemon 8, 341 collaris 341 collaris 341 Polydaedalus 277 Poyntonopbrynus 5, 41, 46, 67, 68 dombensis 67, 68 grandisonae 41, 46, 67, 68 kavangensis 46, 68 vertebralis 68 Prosymna 8, 29, 342, 343, 344 ambigua 342, 343 ambigua 342 brevis 342 ambiguus 342 angolensis 29, 342, 343, 344 bocagii 342 frontalis 342, 343, 344 meleagris 342 visseri 344 Psammobates 173 Psammopbis 8, 25, 26, 29, 41, 46, 332, 345, 346, 347, 348, 349, 350, 351, 352, 353, 354, 355 acutus 25, 354 angolensis 345, 346 ansorgii 29, 41, 346 Ansorgii 346 bocagii 352, 353 Bocagii 352 brevirostris 347 brevirostris 347 leopardinus 347 cf mossambicus 348 elegans 348 irregularis 353 irregularis 353 jallae 46, 346, 347 leigbtoni 350, 351 namibensis 350 leopardinus 347, 348 moniliger 351 notostictus 351 mossambicus 348, 349, 350 namibensis 350, 351 notostictus 351 oxyrbyncbus 354 pbillipsii 348, 350 rbombeatus 355 robani 346, 347 Robani 346 sibilans 26, 332, 347, 348, 350, 351, 352, 353, 354 brevirostris 347 leopardina 347 leopardinus 26, 347 mossambica 348 notostictus 351 sibilans 347, 348, 353 stenocepbala 351 stenocepbalus 351 subtaeniata 351, 352 stenocepbalus 26 subtaeniatus 46, 351, 352, 353 subtaeniatus 352 trigrammus 353 zambiensis 350, 353, 354 Psammopbylax 8, 25, 41, 333, 334, 354, 355, 356, 357 acutus 354, 355, 357 acutus 354, 357 jappi 354 nototaenia 333, 334 ocellatus 355, 356 496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II rhombeatus 41, 355, 356 ocellatus 41, 355, 356 rhombeatus 356 tritaeniatus 354, 355, 356, 357 tritaeniatus 354, 355, 356, 357 viperinus 333, 334 Pseudaspidinae 358, 359 Pseudaspis 8, 357, 358, 359 cana 357, 358 anchietae 357, 358 Pseudohaje 8, 376 goldii 376 Ptychadena 5, 29, 31, 47, 129, 130, 131, 132, 133, 134, 135, 136, 137, 138, 139, 140, 141, 142, 143 (Ptychadena) anchietae 130 (Ptychadena) ansorgii 130 (Ptychadena) bunoderma 131 (Ptychadena) chrysogaster 133, 134 (Ptychadena) guibei 133 (Ptychadena) keilingi 134 (Ptychadena) mascareniensis 135 (Ptychadena) oxyrhynchu 137 (Ptychadena) perplicata 138 (Ptychadena) porosissima 139 (Ptychadena) pumilio 141 (Ptychadena) subpunctata 140 (Ptychadena) upembae 142 (Ptychadena) uzungwensis 142 anchietae 129 ansorgei 130, 138 ansorgi 130 ansorgii 130, 131 bibroni 133, 136 bibronii 132 bunoderma 29, 31, 131, 132 cf. grandisonae 132 cf. mascareniensis 134 cf. mossambica 136 chrysogaster 133 guibei 133 grandisonae 132, 133 guibei 133, 134 hylaea 135, 136 keilingi 134 loveridgei 138, 139 macareniensis 136 mascareniensis 47, 134, 135, 139 bibroni 135 hylaea 134, 135 mossambica 136 nilotica 135 oxyrhynchus 136, 137 perplicata 138 porosissima 138, 139 pumilio 141 smith! 141 subpunctata 139, 140, 142 taenioscelis 141 upembae 142 machadoi 142 uzungwensis 142, 143 Ptychadenidae 5, 37, 38, 41, 128 Python 7, 51, 52, 53, 304, 305, 306, 307 anchietae 304 Anchietae 304 natalensis 51, 304, 305, 307 Natalensis 305 sebae 53, 304, 305, 306, 307 natalensis 304 sebae 306 Sebae 306 sp 52 Pythonidae 7, 37, 38, 304 Pythonodipsas 8, 358, 359 carinata 358 Pyxicephalidae 5, 37, 38, 149 Pyxicephalus 6, 46, 129, 152, 153, 155, 156 adspersus 152, 153 edulis 152 edulis 46, 152, 153 mgosus 155, 156 tuberculosus 155 R Rana 26, 29, 31, 81, 91, 128, 129, 130, 131, 132, 133, 134, 135, 136, 138, 139, 140, 142, 143, 149, 151, 152, 153, 155, 156, 157, 158, 159, 160, 161, 162 (Amnirana) albolabris 158 (Amnirana) darling! 159 (Amnirana) lemairei 160 (Amnirana) parkeriana 161 (Aubria) subsigillata 151 (Ptychadena) bibroni 132 (Ptychadena) buneli 131 (Ptychadena) keilingi 134 (Ptychadena) mascareniensis 135, 139 (Ptychadena) oxyrhynchus 136 oxyrhynchus 136 (Pyxicephalus) adspersus 152, 153 (Rana) angolensis 149 (Rana) darling! 159 (Tomoptema) cacondana 155 (Tomoptema) signata 155 (Tomoptema) tuberculosa 155 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 497 adspersa 152 albolabris 158, 159, 160, 161, 162 acutirostris 161, 162 adiscifera 159, 160 lemairei 160 parkeriana 161, 162 anchietae 129 angolensis 129, 149 ansorgei 130 ansorgii 130 Ansorgii 130 bragantina 26, 157 buneli 31, 132 bunodemia 131 chapini 149 chrysogaster 133 guibei 133 cryptotis 29, 153 darling! 158, 159, 160 Delalandi 149 fusca 161 acutirostris 161 fuscigula 149 angolensis 149 grandisonae 132 keilingi 3 1 Lemairei 160 loveridgei 139 marmorata 91 huillensis 91 maseareniensis 129, 132, 134, 135, 139, 142, 143 maseareniensis 135 porosissima 139 subpunetata 139, 142 uzungwensis 142, 143 mio tympanum 129 mossambiea 136 mossambieus 81 oeeipitalis 157 omata 128, 129 omatissima 128, 129 omatissima 26, 128, 129 oxyrhyneha 136 oxyrhynehus 129, 136 oxyrhrynehus 129 oxyrhynehus 136 parkeriana 161 porosissima 138, 139 Rana (Hildebrandtia) 128 myotympanum 128 omatissima 128 mgosa 155 subpunetata 139, 140, 143 subsigillata 151 tubereulosa 155, 156 upembae 142 Ranidae 6, 37, 38, 41, 158 Rappia 85, 86, 87, 88, 89, 90, 91, 94, 95, 96, 97, 100, 101, 102, 106, 108, 109, no. 111, 112, 114 benguellensis 94 bivitatta 110 bivittata 100, 101, 109 boeagei 96 maeulata 96 boeagii 96 einetiventris 96 einnamomei-ventris 100 einnamomeiventris 100 faseiata 109, 110 fulvo-vittata 85 fulvovittata 85, 88 fulvovittatus 88 fuseigula 102 insignis 91 marmorata 90, 91 marginata 90 taeniolata 90 variegata 90 mierops 111 nasuta 95, 106 nobrei 89, 90 oeellata 108 osorioi 86, 87 platyeeps 109, 110 angolensis 109, 110 plieifera 90, 91 punetulata 106 quinquevittata 112 seabrai 96, 97 steindaehneri 114 tristis 100, 101 Rhaeophoridae 6, 37, 38, 162 Rhagerhis aeuta 354 Rhagerrhis tritaeniata 356 Rhamnophis 8, 398, 399 aethiopissa 398, 399 aethiopissa 399 ituriensis 399 ituriensis 398 Rhamphiophis 354, 355 aeutus 354, 355 aeutus 354, 355 jappi 354, 355 498 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II togoensis 355 wittei 354, 355 Rhinotyphlops 292, 293, 295, 296, 297, 298, 299 angolensis 292 anomalus 293 brevis 298 lineolatus 295 lineolatus 295 mucmso 296, 298 petersii 298 praeocularis 299 schlegelii 296, 297, 298 dinga 296 mucmso 296 petersii 297 schmidti 298 Rhoptropus 6, 30, 31, 41, 45, 46, 47, 199, 200, 201, 202, 203, 204, 205 afer 199, 201 benguellensis 201 bamardi 200, 202, 204 benguellensis 41, 45, 47, 200, 201, 204 biporosus 201, 202, 204 boultoni 30, 31, 200, 201, 202, 203, 204 benguellensis 31, 200, 201 boultoni 202 montanus 203 braconnieri 199 bradfieldi 200 montanus 41, 203, 204 sp 204 taeniostictus 41, 200, 204 Riopa 239, 243, 246, 247, 248 (Eumecia) anchietae 239 (Panaspis) cabindae 247 anchietae 239 sundevalii 246 sundevalii 246 sundevalii 246 s Scapateira reticulata 219 Scaphiophis 8, 399, 400 albopunctatus 399, 400 Scapteira 219, 220 reticulata 220 serripes 219 Scelotes 250, 251,252 bayonii 252 bipes 251, 252 ungolensis 250 Schismaderma 5, 69 carens 69 Schoutedemella spinalis 119 Schoutedenella 118, 120 lameerei 118 xenochira 120 xenochims 120 Scincidae 7, 37, 38, 41, 237 Scincodipus congicus 25 1 Sclerophrys 5, 28, 31, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78 buchneri 28, 69, 70, 71 flinera 76 funerea 70 garmani 71, 72, 75 humbensis 72 gutturalis 72, 73, 78 humbensis 72 lemairii 73, 74 maculata 75, 76 poweri 72, 74, 75 pseudogarmani 75 pusilla 75, 76, 78 regularis 31, 72, 77, 78 Sepedon316, 318 rhombeata 318 Sepsina 7,41,46, 250, 251,252 (Dumerilia) Bayonii 25 1 angolensis 46, 250 bayonii 25 1 copei 41, 46, 252 Copei 252 copii 252 Siaphos dewittei 243 Simocephalus 335, 340 poensis 340 Simocephalus capensis 335 Sphenorhina elegans 241 Stellio 284, 285, 286 atricollis 284, 286 nigricolis 285 nigricollis 284, 286 Stenhorhynchus natalensis 147 Stenostoma 300, 302, 303 nigricans 300, 302 rostratum 303 scutatum 302, 303 scutifrons 300, 302, 303 Stemothaems 164, 165, 166, 167, 168 Adansoni 167 derbianus 165, 167 Derbianus 165 gabonensis 166, 167 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 499 nigricans 164, 166, 167, 168 sinuatus 167 Stigmochelys 6, 172, 173 pardalis 172, 173 T Telescopus 8, 400, 401 finkeldeyi 400 semiannulatus 400, 401 polystictus 400, 401 semiannulatus 400, 401 Temnorhynchus frontalis 343 Testudinidae 6, 37, 38, 169 Testudo 162, 163, 171, 172, 173 erosa 171 galeata 163 pardalis 172 pardalis 172 subrufa 163 Subrufa 162 triunguis 173 Tetradactylus 7, 31, 236, 237 africanus 237 boulengeri 236 lundensis 236 ellenbergeri 31, 236, 237 boulengeri 236, 237 ellenbergi 236 lundensis 31, 236, 237 Tharsops aethiopissa 399 Thelotomis 8, 401, 402, 403 eapensis 401, 402 oatesi 401, 402 oatesii 401 Capensis 401 Kirtlandi 401, 402 kirtlandii 401, 402, 403 eapensis 401, 402 kirtlandii 402 oatesii 401 Thrasops 8, 403, 404 flavigularis 403, 404 jaeksoni 404 jaeksonii 404 jaeksoni 404 Jaeksonii 404 Tiliqua affinis 253 Tomoptema 6, 29, 36, 46, 153, 154, (Tomoptema) eryptotis 153 (Tomoptema) krugerensis 155 (Tomoptema) tubereulosa 156 eryptotis 153, 154, 155 damarensis 36, 46, 154 delalandii 155 kmgerensis 154, 155 mgosa 155 signata 155 tandyi 154, 155 tubereulosa 155, 156 Tomompeltis eolobura luluae 205 Tomuropeltis 211 Toxieodryas 8, 405, 406 blandingii 405, 406 pulvemlenta 405, 406 pulvemlentus 405 Traehylepis 7, 9, 26, 28, 29, 31, 41, 45, 46, 48, 55, 244, 245, 252, 253, 254, 255, 256, 257, 258, 259, 260, 261, 262, 263, 264, 265, 266, 267, 268, 269, 270, 271 aeutilabris 46, 252, 253 aff. spilogaster 268, 269 affinis 253, 254 albopunetata 256 angolensis 266 bayonii 48, 256, 257 bayonii 256 huilensis 256 keniensis 256 Bayonii 256 binotata 257, 258 boeagii 28, 258, 259 ef. albopunetata 255 ef. albopunetatus 254 ef. laeertiformis 262 ef megalura 264, 265 ef spilogaster 268 ehimbana 259, 260 damarana 260, 261 Gravenhorstii 256 hoesehi 46, 261, 262 ivensii 244, 245 laeertiformis 46, 260, 262 laevis 263 maeulilabris 26, 28, 45, 263, 264 megalura 265 monardi 9, 31, 41, 46, 55, 256, 265, 266 oeeidentalis 266 puleherrima 259 punetulata 46, 267, 268 155,156 quinquetaeniata 259 spilogaster 268, 269 striata 270, 271, 272 suleata 29, 269, 270 ansorgii 269, 270 500 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 65, Supplement II nigra 270 sulcata 270 varia 254, 255, 256, 261, 262, 266 variegata 268 wahlbergi 272 wahlbergii 270, 271, 272 Tricbobatracbus 5, 35, 127 cf. robustus 127 robustus 127 Trimerorbinus 355, 356 rbombeatus 355, 356 tritasniatus 356 Trioceros 7, 45, 284 oweni 45, 284 Trionycbidae 6, 37, 38, 173 Trionyx 6, 24, 173, 174 triunguis 173, 174 Tropidonotus olivaceus 409 Tropidosaura Dumerilii 216 Tropinodotus fuliginoides 408 Tupinambis albigularis 275 Typblacontias 7, 29, 41, 46, 272, 273, 274, 275 bogerti 273 jobnsonii 272 punctatissimus 41, 272, 273, 274 bogerti 41, 273, 274 punctatissimus 272, 273 rboani 46 robani 29, 46, 274 mdebecki 41, 274, 275 Typblopidae 7, 37, 38, 41, 292 Typblops 292, 293, 294, 295, 296, 297, 298, 299 (Onycbocepbalus) Ancbietae 293 (Onycbocepbalus) bumbo 297 (Opbtbalmidion) Escbricbtii 294 intermedia 294 lineolata 294 (Opbtbalmidion) Kraussii 294 (Opbtbalmidion) lineolatus 294 ancbietae 293 angolensis 292 adolfi 292 angolensis 292 anomalus 293 bocagei 294, 295 boulengeri 295 boulengeri 295 Boulengeri 294 congicus 292 escbricbtii 294 bottentotus 297 bumbo 297 lineolatus 294, 295 lineolatus 295 mucmso 296, 297 Petersii 297 praeocularis 299 lundensis 299 punctatus 292, 294, 295 intermedia 294 lineolata 294 lineolatus 294 punctatus 292, 295 scblegeli 296 mucmso 296 scblegelii 296, 297 mucmso 296, 297 petersii 297 scbmidti 298, 299 Typblosaums 237, 238 jappi 237 lineatus 237, 238 jappi 237 u Uraeus 368, 369 ancbietae 368 Uriecbis 321, 322 capensis 321, 322 Guentberii 321 punctatolineatus 321 V Varanidae 7, 37, 38, 275 Varanus 7, 30, 31, 46, 47, 51, 53, 275, 276, 277, 278 (Empagusa) exantbematicus 275 angolensis 275 (Polydaedalus) niloticus 277 albigularis 30, 46, 51, 53, 275, 276, 277 albigularis 275 angolensis 30, 46, 53, 275, 277 exantbematicus 275, 277 albigularis 275 angolensis 275 niloticus 47, 277, 278 niloticus 277 ocellatus 275 omatus 278 Vipera 26, 309, 310, 311, 312, 314 (Bids) rhinoceros 311 (Cerastes) caudalis 310 (Echidna) arietans 309 arietans 309 MARQUES ET AL.; AMPHIBIANS AND TERRESTRIAL REPTILES OF ANGOLA 501 aspis 310 caudalis 310 heraldica 26, 312 ocellata 310, 311 peringueyi 314 rhinoceros 311 Viperidae 7, 37, 38, 41, 308 X Xenocalamus 8, 28, 45, 359, 360, 361 bicolor 45, 359 bicolor 359 machadoi 45, 359 pemasutus 359 mechovii 359 Mechovii 359, 360 mechowi 360 mechowi 360 mechowii 28, 45, 360, 361 inomatus 360, 361 mechowii 360 Mechowii 360 Xenopus 5, 59, 60, 61, 62, 63, 64, 65 (Silurana) epitropicalis 60 allofraseri 59, 61 amieti 59, 61 andrei 59, 61 boumbaensis 59 calcaratus 60 cf. epitropicalis 60 cf. petersii 62 epitropicalis 60 eysoole 59 fraseri 59, 60, 61 itombwensis 59 laevis 62, 64, 65 petersi 62 petersii 64, 65 poweri 62, 64, 65 lenduensis 59 longipes 59 mellotropicalis 60 muelleri 61, 62 parafraseri 61 petersii 62, 65 Petersii 62 poweri 64, 65 pygmaeus 59, 61 mwenzoriensis 59 sp 60, 61 tropicalis 60 vestitus 59 victorianus 65 wittei 59 Xenurophis caesar 384 z Zonurus 227 angolensis 227 cordylus 227 Zygaspis 6, 31, 211, 212 nigra 211, 212 quadri-frons 212 quadrifrons31,211,212, 213 capensis 211 quadrifrons 212 CALIFORNIA ACADEMY OF SCIENCES INSTRUCTIONS TO AUTHORS Authors planning to submit papers for consideration for publication in the Academy’s Proceedings, Occasional Papers, or Memoir series must follow the directions given below in preparing their submissions. 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COMMENTS Address editorial correspondence or requests for pricing information to the Editor, Scientific Publications Office, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118 U.S.A. or via email to the Editor, Scientific Publications, at aleviton@calacademy.org or gwilliams@calacademy.org Table of Contents Mariana P. Marques, Luis M.P. Ceriaco, David C. Blackburn, and Aaron M. Bauer: Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola . . .1-480 Taxonomic Index . 481-501