Skip to main content Skip to main navigation menu Skip to site footer
Published: 2020-03-02

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

Department of Natural Sciences, University of Michigan-Dearborn, 4901 Evergreen Road, Dearborn, Michigan 48128, USA
Department of Natural Sciences, University of Michigan-Dearborn, 4901 Evergreen Road, Dearborn, Michigan 48128, USA
Museu de História Natural e da Ciência da Universidade do Porto, Praça Gomes Teixeira, 4099-002 Porto, Portugal Departamento de Zoologia e Antropologia (Museu Bocage), Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Rua da Escola Politécnica 58, 1269–102, Lisboa, Portugal
Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1699, USA.
Scincidae endemism Panaspis mocamedensis sp. nov. southwestern Africa cryptic species taxonomy


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



  1. Angel, F. (1924) Description d’un lezard nouveau d’Afrique Orientale, appartement au genre Ablepharus (Mission Alluaud, 1903–1904). Bulletin du Museum national d’Histoire naturelle, 30, 52–53.

    Angel, F. (1944) Un lézard nouveau du Mont Nimba (Haute-Guinée Française) appartenant au genre Lygosoma (matériaux de la mission Lamotte au Mont Nimba). Bulletin du Muséum d’Histoire naturelle, Paris, 2ème Série, 5, 293–294.

    Baptista, N., António, T. & Branch, W.R. (2018) Amphibians and reptiles of the Tundavala region of the Angolan Escarpment. Biodiversity & Ecology, 6, 397–403.

    Bates, M.F., Branch, W.R., Bauer, A.M., Burguer, M., Marais, J., Alexander, G.J. & De Villiers, M.S. (2014) Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. Suricata 1. South African National Biodiversity Institute, Pretoria, xvii + 459 pp.

    Bauer, A.M., Shea, G. & Günther, R. (2003) An annotated catalogue of the types of scincid lizards (Reptilia, Squamata, Scincidae) in the collection of the Museum für Naturkunde der Humboldt-Universität zu Berlin (ZMB). Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe, 79 (2), 253–321.

    Bauer, A.M., de Silva, A., Greenbaum, E. & Jackman, T.R. (2007) A new species of day gecko from high elevation in Sri Lanka, with a preliminary phylogeny of Sri Lankan Cnemaspis (Reptilia: Squamata: Gekkonidae). Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe, 83, 22–32.

    Bittencourt-Silva, G.B. (2019) A herpetological survey of western Zambia. Amphibian and Reptile Conservation, 13 (2), 1–28.

    Bocage, J.V.B. (1866a) Lista dos reptis das possessões portuguezas d’Africa ocidental que existem no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes, 1, 37–56.

    Bocage, J.V.B. (1866b) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l’Afrique occidentale, qui se trouvent au Muséum de Lisbonne. Jornal de Sciences Mathematicas, Physicas e Naturaes, 1, 57–78.

    Bocage, J.V.B. (1867) Segunda lista dos reptis das possessões portuguezas d’Africa occidental que existem no Museu de Lisboa. Jornal de Sciences Mathematicas, Physicas e Naturaes, 1, 217–228.

    Bocage, J.V.B. (1887) Mélanges erpétologiques. I. Reptiles et batraciens du Congo. Jornal de Sciences Mathematicas, Physicas e Naturaes, 11, 177–192.

    Bocage, J.V.B. (1895) Herpétologie d’Angola et du Congo. Ministério da Marinha e das Colónias, Lisbonne, 203 pp., 20 pls.

    Bocage, J.V.B. (1897) Mammiferos, reptis e batrachios d’Africa de que existem exemplares typicos no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 4, 187–206.

    Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences, 97, 6585–6590.

    Branch, W.R. (1998) Field Guide to Snakes and other Reptiles of Southern Africa. 3rd Edition. Struik Publishers, Cape Town, 399 pp.

    Branch, W.R. & Conradie, W. (2015) IV. Herpetofauna da região da Lagoa Carumbo. In: Huntley, B. & Francisco, P. (Eds.), Avaliação Rápida da Biodiversidade da Região da Lagoa Carumbo, Lunda Norte—Angola. Ministério do Ambiente de Angola, Luanda, Angola, pp. 194–209.

    Branch, W.R., Pinto. P.V., Baptista, N. & Conradie, W. (2019a) The Reptiles of Angola: History, Diversity, Endemism and Hotspots. In: Huntley, B, Russo, V., Lages & Ferrand, N. (Eds.) Biodiversity of Angola, Science & Conservation: A Modern Synthesis. Springer, Cham, pp. 283–334.

    Branch, W.R., Conradie, W., Pinto, P.V. & Tolley, K.A. (2019b) Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angol. Amphibian & Reptile Conservation, 13 (2), 82–95. [Special Section]

    Butler, B.O., Ceríaco, L.M., Marques, M.P., Bandeira, S., Júlio, T., Heinicke, M.P. & Bauer, A.M. (2019) Herpetological Survey of Huíla Province, Southwest Angola, Including First Records from Bicuar National Park. Herpetological Review, 50, 225–240.

    Boulenger, G.A. (1887) Catalogue of the Lizards in the British Musuem (Natural History). Vol. 3. Lacertidӕ, Gerrhosauridӕ, Scincidӕ, Anelytropsidӕ, Dibamidӕ, Chamaeleontidӕ. Trustees of the British Museum, London, xii + 575 pp., 40 pls.

    Boulenger, G.A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433–456.

    Boulenger, G.A. (1905) A list of the batrachians and reptiles collected by Dr. W.J. Ansorge in Angola with descriptions of new species. Annals and Magazine of Natural History, Series 7, 16, 105–115, pl. 4.

    Boulenger, G.A. (1909) On the reptiles and batrachians collected by the Tancredi Expedition to Lake Tsana, Abyssinia. Annali del Museo Civico di Storia Naturale di Genova, Series 3, 44, 193.

    Boulenger, G.A. (1913) Descriptions of new lizards in the collection of the British Museum. Annals and Magazine of Natural History, Series 8, 12, 563–566.

    Ceríaco, L.M.P., Bauer, A.M.B., Blackburn, D.C. & Lavres, A.C.F.C. (2014) The herpetofauna of the Capanda Dam region, Malanje, Angola. Herpetological Review, 45 (4), 667–674.

    Ceríaco, L.M.P., Branch, W.R. & Bauer, A.M. (2018a) A new species of African snake-eyed skink (Scincidae: Panaspis) from central and northern Namibia. Zootaxa, 4527 (1), 140–150.

    Ceríaco, L.M.P., de Sá, S.A.C., Bandeira, S., Valério, H., Stanley, E.L., Kuhn, A.L., Marques, M.P., Vindum, J.V., Blackburn, D.C. & Bauer, A.M. (2016a) Herpetological survey of Iona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, southwestern Angola. Proceedings of the California Academy of Sciences, Series 4, 63, 15–61.

    Ceríaco, L.M.P., Marques, M.P. & Bandeira, S.A. (2016b) Anfíbios e Répteis do Parque Nacional da Canganda. Instituto Nacional da Biodiversidade e Áreas de Conservação, Angola & Museu Nacional de História Natural e da Ciência, Lisboa, 97 pp.

    Ceríaco, L.M.P., Marques, M.P., Bandeira, S., Agarwal, I., Stanley, E.L., Bauer, A.M., Heinicke, M.P. & Blackburn, D.C. (2018b) A new earless species of Poyntonophrynus (Anura, Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola. ZooKeys, 780, 109–136.

    Ceríaco, L.M.P., Marques, M.P., Bandeira, S., Blackburn, D.C. & Bauer, A.M. (2018c) Herpetological survey of Cangandala National Park with a synoptic list of the amphibians of Malanje Province, central Angola. Herpetological Review, 49 (3), 408–431.

    Chabanaud, P. (1917) Enumeration des reptiles non encore étudiés de l’Afrique occidentale, appartenant aux collections du Museum, avec la description des espèces nouvelles. Bulletin du Muséum national d’Histoire naturelle, 23, 83–105.

    Chirio, L. & LeBreton, M. (2007) Atlas des Reptiles du Cameroun. Publications Scientifiques du Muséum national d’Histoire naturelle/IRD Éditions, Paris, 686 pp.

    Conradie, W., Bills, R. & Branch, W.R. (2016) The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian & Reptile Conservation, 10 (2), 6–36.

    Conradie, W., Branch, W.R., Measey, G.J. & Tolley, K.A. (2012a) A new species of Hyperolius Rapp, 1842 (Anura: Hyperoliidae) from the Serra da Chela Mountains, south-western Angola. Zootaxa, 3269 (1), 1–17.

    Conradie, W., Branch, W.R. & Tolley, K.A. (2013) Fifty shades of grey: giving colour to the poorly known Angolan Ashy reed frog (Hyperoliidae: Hyperolius cinereus), with the description of a new species. Zootaxa, 3635 (3), 201–223.

    Conradie, W., Measey, G.J., Branch, W.R. & Tolley, K.A. (2012b) Revised phylogeny of African sand lizards (Pedioplanis), with the description of two new species from south-western Angola. African Journal of Herpetology, 61 (2), 91–112.

    Cope, E.D. (1868) Observations on reptiles of the old world. Proceedings of the Academy of Natural Sciences of Philadelphia, 20 (1868), 316–323.

    de Queiroz, K. (1999) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, pp. 57–75.

    De Witte, G.F. (1933) Batraciens et reptiles recueillis par M.L. Burgeon au Ruwenzori, au Kivu et au Tanganika. Revue de Zoologie et de Botanique Africaines, 24, 111–123.

    Ernst, R., Nienguesso, A.B.T., Lautenschlager, T., Barej, M.F., Schmitz, A. & Hölting, M. (2014) Relicts of a forested past: Southernmost distribution of the hairy frog genus Trichobatrachus Boulenger, 1900 (Anura: Arthroleptidae) in the Serra do Pingano region of Angola with comments on tis taxonomic status. Zootaxa, 3779 (2), 297–300.

    Ernst, R., Schmitz, A., Wagner, P., Branquima, M.F. & Hölting, M. (2015) A window to Central African forest history: distribution of the Xenopus fraseri subgroup of the Congo Basin, including a first record of Xenopus andrei from Angola. Salamandra, 51 (2), 147–155.

    Ferreira, J.B. (1904) Reptis e amphibios de Angola da região ao norte do Quanza (Collecção Newton—1903). Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 7, 111–117.

    FitzSimons, V.F.M. (1937) Notes on the reptiles and amphibians collected and described from South Africa by Andrew Smith. Annals of the Transvaal Museum, 17, 259–274.

    Fuhn, I.E. (1969) The “polyphyletic” origin of the genus Ablepharus (Reptilia, Scincidae): a case of parallel evolution. Journal of Zoological Systematics and Evolutionary Research, 7 (1), 67–76.

    Fuhn, I.E. (1972) Révision du phylum forestier du genre Panaspis Cope (Reptilia, Scincidae, Lygosominae). Revue Roumaine de Biologie, série de Zoologie, 17 (4), 257–272.

    Gonçalves, F.M.P., Braine, D., Bauer, A.M., Valério, H., Marques, M.P. & Ceríaco, L.M. (2019) Rediscovery of the poorly known Angolan Adder, Bitis heraldica (Bocage, 1889) (Serpentes: Viperidae): New records, live photographs and first case history of envenomation. Herpetological Review, 50 (2), 241–246.

    Grandvaux-Barbosa, L.A. (1970) Carta Fitogeográfica de Angola. Instituto de Investigação Científica de Angola, Luanda, 323 pp.

    Greer, A.E. (1974) The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, 31, 1–67.

    Groth, J. G. & Barrowclough, G. F. (1999) Basal divergences in birds and the phylogenetic utility of the nuclear RAG-1 gene. Molecular Phylogenetics and Evolution, 12, 115–123.

    Heinicke, M.P., Ceríaco, L.M.P., Moore, I.M., Bauer, A.M. & Blackburn, D.C. (2016) Tomopterna damarensis (Anura: Pyxicephalidae) is broadly distributed in Namibia and Angola. Salamandra, 53 (3), 461–465.

    Hoogmoed, M.S. (1980) Herpetologische waarnemingen in Ghana. VII. Lacerta, 38, 88–95.

    Huntley, B., Russo, V., Lages, F. & Ferrand, N. (2019) Biodiversity of Angola, Science & Conservation: A Modern Synthesis. Springer, Cham, 549 pp.

    Jesus, J., Harris, D.J. & Brehm, A. (2007) Relationships of Afroablepharus Greer, 1974, skinks from the Gulf of Guinea islands based on mitochondrial and nuclear DNA: patterns of colonization and comments on taxonomy. Molecular Phylogenetics and Evolution, 45 (3), 904–914.

    Jacobsen, N.H.G. & Broadley, D.G. (2000) A new species of Panaspis Cope (Reptilia: Scincidae) from southern Africa. African Journal of Herpetology, 49 (1), 61–71.

    Kilunda, F.K., Conradie, W., Wasong, D.V., Jin, J.-Q, Peng, M.-S., Murphy, R.W., Malonza, P.K. & Che, J. (2019) Revalidation and resurrection of Panaspis massaiensis (Angel, 1924) and the description of a new species of Panaspis Cope (Squamata: Scincidae) from south-eastern Kenya. Zootaxa, 4706 (2), 255–274.

    Kuhn, A. (2016) Systematics of the Namib Day Geckos (Squamata: Gekkonidae: Rhoptropus). Unpublished Master’s Thesis, Villanova University, Villanova, Pennsylvania, 226 pp.

    Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35, 1547–1549.

    Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2016) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773.

    Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez, R. & Thompson, J.D. (2007) Clustal W and Clustal X version 2.0. Bioinformatics, 23, 2947–2948.

    Loveridge, A. (1932) New races of a skink (Siaphos) and frog (Xenopus) from the Uganda Protectorate. Proceedings of the Biological Society of Washington, 45, 113–115.

    Loveridge, A., (1952) A startlingly turquoise-blue gecko from Tanganyika. Journal of the East Africa Natural History Society and National Museum, 20, 446.

    Marques, M.P., Ceríaco, L.M.P., Blackburn, D.C. & Bauer, A.M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola—Atlas of historical and bibliographic records (1840–2017). Proceedings of the California Academy of Sciences, Series 4, 65 (supplement 2), 1–501.

    Marques, M.P. Ceríaco, L.M.P., Bandeira, S., Pauwels, O.S.G. & Bauer, A.M. (2019a) Description of a new long-tailed skink (Scincidae: Trachylepis) from Angola and the Democratic Republic of Congo. Zootaxa, 4568 (1), 51–68.

    Marques, M.P. Ceríaco, L.M.P., Stanley, E.L., Bandeira, S., Agarwal, I. & Bauer, A.M. (2019b) A new species of girdled lizard (Squamata: Cordylidae) from Serra da Neve inselberg, Namibe province, southwestern Angola. Zootaxa, 4668 (4), 503–524.

    Medina, M.F., Bauer, A.M., Branch, W.R., Schmitz, A., Conradie, W., Nagy, Z.T., Hibbitts, T.J., Ernst, R., Portik, D.M., Nielsen, S.V., Colston, T.J., Kusamba, C., Behangana, M. & Rödel, M.-O. (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100, 409–423.

    Monard, A. (1949) Vertébrés nouveaux du Cameroun. Revue Suisse de Zoologie, 56 (38), 731–745.

    Müller, L. (1910) Beiträge zur Herpetologie Kameruns. Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie der Wissenschaften, 24, 543–626, 1 pl.

    Nguyen, L.T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2014) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.

    Nieden, F. (1923) Neues Verzeichnis der Kriechtiere (außer den Schlangen) von Deutsch-Ostafrika, I. Teil: Reptilia. Mitteilungen aus dem Zoologischen Museum in Berlin, 7 (1), 51–100.

    Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics, 2nd Edition. Sinauer Associates, Sunderland, Massachusetts, pp. 205–247.

    Parker, H.W. (1936) Dr. Karl Jordan’s expedition to South-West Africa and Angola : herpetological collection. Novitates Zoologicae, 40, 115–146.

    Perret, J.L. (1973) Contribution à l’étude des Panaspis (Reptilia, Scincidae) d’Afrique occidentale avec la description de deux espèces nouvelles. Revue Suisse de Zoologie, 80, 595–630.

    Perret, J.L. (1975) La differentiation dans le genre Panaspis Cope (Reptilia: Scincidae). Bulletin de la Société Neuchâteloise des Sciences Naturelles, 98, 5–16.

    Peters, W.C.H. (1873) Über eine neue Schildkrötenart, Cinosternon effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1873, 603–618.

    Peters, W.C.H. (1877) Übersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum übergeben sind. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1877, 311–621, 1 pl.

    Roll, U., Feldman, A., Novosolov, M., Allison, A., Bauer, A.M., Bernard, R., Böhm, M, Castro-Herrera, F., Chirio, L., Collen, B., Colli, G.R., Dabool, L., Das, I., Doan, T.M., Grismer, L.L., Hoogmoed, M., Itescu, M., Kraus, F., LeBreton, M., Lewin, A., Martins, M., Maza, E., Meirte, D., Nagy, Z.T., Nogueira, C.C., Pauwels, O.S.G., Pinhceira-Donoso, D., Powney, G.D., Sindaco, R., Tallowin, O.J.S., Torres-Carvejal, O., Trape, J-F., Vidan, E., Uetz, P., Wagner, P., Wang, P., Wang, Y., Orlme, D.L., Grenyer, R. & Meiri, S. (2017) The global distribution of tetrapods reveals a need for targeted reptile conservation. Nature Ecology Evolution, 1, 1677–1682.

    Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863 (1), 1–28.

    Schmidt, K.P. (1919) Contributions of the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909–1915. Part 1, Turtles, crocodiles, lizards and chamaeleons. Bulletin of the American Museum of Natural History, 39, 385–624, pls. 7–32.

    Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701.

    Smith, A. (1849) Appendix. In: Smith, A. (Ed.), Illustrations of the zoology of South Africa, Reptilia, 1849, pp. 1–28 + [i].

    Smith, M.A. (1937) A review of the genus Lygosoma (Scincidae: Reptilia) and its allies. Records of the Indian Museum, 39 (3), 213–234.

    Soares, L.B., Ceríaco, L.M.P., Marques, M.P., Bastos-Silveira, C., Scheinberg, L.A., Harris, D.J., Brehm, A. & Jesus, J. (2018) Review of the leaf-litter skinks (Scincidae: Panaspis) from the Gulf of Guinea oceanic islands, with the description of a new species. African Journal of Herpetology, 67 (2), 132–159.

    Stanley, E.L., Ceríaco, L.M.P., Bandeira, S., Valério, H., Bates, M.F. & Branch, W.R. (2016) A review of Cordylus machadoi (Squamata: Cordylidae) in southwestern Angola, with the description of a new species from the pro-Namib desert. Zootaxa, 4061 (3), 201–226.

    Trape, J.F., Trape, S. & Chirio, L. (2012) Lézards, crocodiles et tortures d’Afrique occidentale et du Sahara. IRD, Marseille, 503 pp.

    Werner, F. (1902) Über westafrikanische Reptilien. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 52, 332–348.

    Werner, F. (1909) Beschreibung neuer Reptilien aus dem Kgl. Naturalienkabinett in Stuttgart. Jahresheft des Vereins für vaterländische Naturkunde im Württemburg, 65, 55–63.

    Werner, F. (1914) Über Eidechsen, darunter zwei neue, aus dem angloägyptischen Sudan. Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse, 57 (18), 393–94.