Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2022-04-05
Page range: 401-430
Abstract views: 724
PDF downloaded: 38

Systematic revision of Afrogecko ansorgii (Boulenger, 1907) (Sauria: Gekkonidae) from western Angola

CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Portugal. 2Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4099-002 Porto, Portugal. 3BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485-661 Vairão, Portugal.
Port Elizabeth Museum (Bayworld), Beach Road, Humewood, Port Elizabeth, 6013, South Africa. Department of Nature Conservation Management, Natural Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, George, South Africa.
CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Portugal. BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485-661 Vairão, Portugal. Fundação Kissama, Rua Joquim Kapango, 49 1º Luanda, Angola.
Zoology and Entomology Molecular Lab (ZEML), Department of Zoology and Entomology, Rhodes University, Makhanda, 6140, South Africa.
Zoology and Entomology Molecular Lab (ZEML), Department of Zoology and Entomology, Rhodes University, Makhanda, 6140, South Africa.
Physics Department, Stellenbosch University, Stellenbosch, 7602, South Africa. CT Scanner Facility, Stellenbosch University, Stellenbosch, 7602, South Africa.
Port Elizabeth Museum (Bayworld), Beach Road, Humewood, Port Elizabeth, 6013, South Africa.
Reptilia Leaf-toed gecko taxonomy CT-scan skull osteology phylogeny


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



  1. Baptista, N.L., Tolley, K.A., Bluhm, M., Finckh, M. & Branch, W.R. (2020) Rediscovery, range extension, habitat and phylogenetic relation of the endemic Scaled Sandveld Lizard Nucras scalaris Laurent, 1964 (Sauria: Lacertidae) in the central Angolan plateau. African Journal of Herpetology, 69 (1), 12–28.
    Bauer, A.M., Good, D.A. & Branch, W.R. (1997) The taxonomy of the Southern African leaf-toed geckos (Squamata: Gekkonidae), with a review of Old World “Phyllodactylus” and the description of five new genera. Proceedings of the California Academy of Sciences, 49 (14), 447–497.
    Bauer, A.M. & Menegon, M. (2006) A new species of prehensile-tailed gecko, Urocotyledon (Squamata: Gekkonidae), from the Udzungwa Mountains, Tanzania. African Journal of Herpetology, 55 (1), 13–22.
    Bauer, A.M., Beach-Mehrotra, M., Bermudez, Y., Clark, G.E., Daza, J.D., Glynne, E., Hagyari, J.M.H., Holovacs, N., Kanasiro, A., Lofthus, A.J., Pierce, Z.W., Aaliyah, R., Syed, S., Vallejo-Pareja, M.C., Walker, B.A. & Willett, J. (2018) The tiny skull of the Peruvian gecko Pseudogonatodes barbouri (Gekkota: Sphaerodactylidae) obtained via a divide-and-conquer approach to morphological data acquisition. South American Journal of Herpetology, 13 (2), 102–116. 00113.1
    Bauer, A.M., Lamb, T. & Branch, W.R. (2006) A revision of the Pachydactylus serval and P. weberi groups (Reptilia: Gekkota: Gekkonidae) of Southern Africa, and with the description of eight new species. Proceedings of the California Academy of Sciences, 57 (23), 595–709.
    Branch, W.R. & Bauer, A.M. (1992) The status of Phyllodactylus peringueyi (Reptilia: Gekkonidae). The Journal of the Herpetological Association of Africa, 40 (1), 21–21.
    Branch, W.R. & Bauer, A.M. (1994) Phyllodactylus peringueyi (Reptilia: Gekkonidae): Its taxonomic history, rediscovery, and phylogenetic affinities. Annals of the South African Museum, 104 (2), 13–30.
    Branch, W.R., Guyton, J.A., Schmitz, A., Barej, M.F., Naskrecki, P., Farooq, H., Verburgt, L. & Roedel, M.O. (2017) Description of a new flat gecko (Squamata: Gekkonidae: Afroedura) from Mount Gorongosa, Mozambique. Zootaxa, 4324 (1), 142–160.
    Branch, W.R., Conradie, W., Vaz Pinto, P. & Tolley, K. (2019b) Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from south-western Angola. Amphibian and Reptile Conservation, 12 (2), 85–95.
    Branch, W.R., Schmitz, A., Baptista, N., Lobón-Rovira, J., Antonio, T. & Conradie, W. (2021) Rock island melody: a revision of the Afroedura bogerti Loveridge, 1944 group, with the description of four new endemic species for Angola. Zoosystematics and Evolution, 97 (1), 55–82.
    Branch, W.R., Vaz Pinto, P., Baptista, N. & Conradie, W. (2019a) The Reptiles of Angola: History, Diversity, Endemism and Hotspots. In: Huntley, B., Russo, V., Lages, F. & Ferrand, N. (Eds.), Biodiversity of Angola, Science & Conservation: A Modern Synthesis. Springer, Cham, Switzerland, pp. 283–334.
    Boulenger, G.A. (1907) Descriptions of three new lizards and a new frog, discovered by Dr. W. J. Ansorge in Angola. The Annals and Magazine of Natural History, 19 (7), 212–214.
    Bruford, M.W., Hanotte, M., Brookfield, J.F.Y. & Burke, T. (1992) ‘Single locus and multilocus DNA fingerprint’. In: Hoelzel, A.R. (Ed.), Molecular Genetic Analysis of Populations: A Practical Approach. IRL Press, Oxford, pp. 225–270.
    Burgess, N., D’Amico Hales, J., Underwood, E., Dinerstein, E., Olson, D., Itoua, I., Schipper, J., Ricketts, T. & Newman, K. (2004) Terrestrial ecoregions of Africa and Madagascar: a conservation assessment. Island Press, Washington, D.C. 524 pp. [xxiii + 501]
    Ceríaco, L.M., Agarwal, I., Marques, M.P. & Bauer, A.M. (2020a) A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species. Zootaxa, 4746 (1), 1–71.
    Ceríaco, L.M., Heinicke, M.P., Parker, K., Marques, M.P. & Bauer, A.M. (2020b) A review of the African snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species. Zootaxa, 4747, 77–112.
    Ceríaco, L.M., Tolley, K.A., Marques, M.P., Heinicke, M.P. & Bauer, A.M. (2020c) A dwarf among giants: phylogenetic position of the elusive Angolan Adder (Bitis heraldica) and biogeographic affinities of Angolan Afromontane regions. African Journal of Herpetology, 69 (1), 108–118.
    Chernomor, O., Von Haeseler, A. & Minh, B.Q. (2016) Terrace Aware Data Structure for Phylogenomic Inference from Supermatrices. Systematic Biology, 65 (6), 997–1008.
    Collyer, M.L. & Adams, D.C. (2018) RRPP: An r package for fitting linear models to high-dimensional data using residual randomization. Methods in Ecology and Evolution, 9 (7), 1772–1779.
    Conroy, C.J., Papenfuss, T., Parker, J. & Hahn, N.E. (2009) Use of Tricaine Methanesulfonate (MS222) for Euthanasia of Reptiles. Journal of the American Association for Laboratory Animal Science, 48, 28–32.
    Daza, J.D., Abdala, V., Thomas, R. & Bauer, A.M. (2008) Skull anatomy of the miniaturized gecko Sphaerodactylus roosevelti (Squamata: Gekkota). Journal of Morphology, 269 (11), 1340–1364.
    Daza, J.D. & Bauer, A.M. (2012) A new amber-embedded sphaerodactyl gecko from Hispaniola, with comments on morphological synapomorphies of the Sphaerodactylidae. Breviora, 529 (1), 1–28.
    Dixon, J.R. & Kroll, J.C. (1974) Resurrection of the generic name Paroedura for the phyllodactyline geckos of Madagascar, and description of a new species. Copeia, 24–30.
    Evans, S.E. (2008) The skull of lizards and tuatara. In: Gans C., Gaunt, S. & Adler, K. (Eds.), Biology of the Reptilia, Volume 20, Morphology. Society for the Study of Amphibians and Reptiles, Ithaca, pp. 1–347.
    Feng, J., Han, D., Bauer, A.M. & Zhou, K. (2007) Interrelationships among Gekkonid geckos inferred from mitochondrial and nuclear gene sequences. Zoological Science, 24, 656–665.
    FitzSimons, V.F. (1959) Some new reptiles from southern Africa and southern Angola. Annals of the Transvaal Museum, 23, 405–409.
    Funnell, S., Shrum, M., Ellis, E. & Andreone, F. (2012) A new record of the phantom gecko Matoatoa spannringi from Sainte Luce, SE Madagascar augments the species’ known range. Herpetology Notes, 5, 151–153.
    Gamble, T., Greenbaum, E., Jackman, T.R., Russell, A.P. & Bauer, A.M. (2012) Repeated origin and loss of adhesive toepads in geckos. PloS one, 7 (6), e39429.
    Gonçalves, F.M., 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.
    Haacke, W.D. (2008) A new leaf-toed gecko (Reptilia: Gekkonidae) from south-western Angola. African Journal of Herpetology, 57 (2), 85–92.
    Hall, T.A. (1999) ‘BioEdit, a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT’, Nucleic Acids Symposium Series (Oxford), 41, 95–98.
    Hallerman, J., Ceríaco, L.M., Schmitz, A., Ernst, R., Conradie, W., Verburgt, L., Marques, M.P. & Bauer A.M. (2020) A review of the Angolan House snakes, genus Boaedon Duméril, Bibron and Duméril (1854) (Serpentes: Lamprophiidae), with description of three new species in the Boaedon fuliginosus (Boie, 1827) species complex. African Journal of Herpetology, 69 (1), 29–78.
    Hawlitschek, O., Scherz, M.D., Ruthensteiner, B., Crottini, A. & Glaw, F. (2018) Computational molecular species delimitation and taxonomic revision of the gecko genus Ebenavia Boettger, 1878. The Science of Nature, 105 (7–8), 49.
    Heinicke, M.P., Daza, J., Greenbaum E., Jackman, T.F. & Bauer, A.M. (2014) Phylogeny, taxonomy and biogeography of a circum-Indian Ocean clade of leaf-toed geckos (Reptilia: Gekkota), with a description of two new genera. Systematics and Biodiversity, 12 (1), 23–42.
    Heinicke, M.P., Turk, D. & Bauer, A.M. (2017) Molecular phylogeny reveals strong biogeographic signal and two new species in a Cape Biodiversity Hotspot endemic mini-radiation, the pygmy geckos (Gekkonidae: Goggia). Zootaxa, 4312 (3), 449–470.
    Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) ‘UFBoot2: Improving the ultrafast bootstrap approximation’. Molecular Biology and Evolution, 35 (2), 518–522.
    Huntley, B.J., Russo, V., Lages, F. & Ferrand, N. (2019) Biodiversity of Angola: Science & Conservation: A Modern Synthesis. Springer Nature, Cham, 549 pp.
    IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. Second edition. Gland, Switzerland and Cambridge, IUCN, iv + 32 pp.
    Jackman, T.R., Bauer, A.M., Greenbaum, E., Glaw, F. & Vences, M. (2008) Molecular phylogenetic relationships among species of the Malagasy-Comoran gecko genus Paroedura (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 46 (1), 74–81.
    Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., Von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14 (6), 587–589.
    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 (6), 1547–1549.
    Lobón-Rovira. J., Vaz Pinto, P., Becker, F.S., Tolley, K.A., Measey, J., Bennet, B., Boon, B., Sá, S., & Conradie, W. (2022) An updated herpetofaunal species inventory of Iona National Park in southwestern Angola. Check List, 18. (in press)
    Lobón-Rovira, J. & Bauer, A.M. (2021) Bone-by-bone: A detailed skull description of White-headed dwarf gecko Lygodactylus picturatus (Peters, 1870). African Journal of Herpetology, 70 (2) 1–20.
    Lobón-Rovira, J., Conradie, W., Buckley Iglesias, D., Ernst, R., Baptista, N. & Vaz Pinto, P. (2021) Between sand, rocks and branches: an integrative taxonomic revision of Angolan Hemidactylus Goldfuss, 1820, with description of four new species. Vertebrate Zoology, 71, 465–501.
    Loveridge, A. (1944) New geckos of the genera Afroedura, new genus, and Pachydactylus from Angola. American Museum Novitates, 1254, 1–4.
    Loveridge, A. (1947) Revision of the African lizards of the Family Gekkonidae. Bulletin of the Museum of Comparative Zoology, 98, 1–469., pls. 1–7.
    Marques, M.P., Ceríaco, L.M.P., Stanley, E.L., Bandeira, S., Agarwal, I. & Bauer, A.M. (2019) A new species of girdled lizard (Squamata: Cordylidae) from Serra da Neve inselberg, Namibe province, southwestern Angola. Zootaxa, 4668 (4), 503–524.
    Marques, M.P., Ceríaco, L.M.P., Buehler, M.D., Bandeira, S., Janota, J.M. & Bauer, A.M. (2020) A revision of the Dwarf Geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with the description of three new species. Zootaxa, 4583 (3), 301–352.
    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Trees. Gateway Computing Environments Workshop, 1–8.
    Minh, B.Q., Schmidt, H.A. Chernomor, O., Schrempf, D., Woodhams, M.D, von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37, 1530–1534.
    Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32 (1), 268–274.
    Nicolau, G.K., Petford, M., Edwards, S., Busschau, T., Lynch, K., Kemp, L., Balmer, J., Hundermark, C.R., Weeber, J. & Conradie, W. (2021) New insights into the geographical distribution, ecology and conservation status of South Africa’s endemic Coastal Leaf-toed Gecko, Cryptactites peringueyi (Boulenger, 1910). Herpetology Notes, 14, 439–450.
    Palumbi, S.R. (1996) The polymerase chain reaction. In: Hillis, D., Moritz, C. & Mable, B. (Eds.), Molecular systematics. 2nd edition. Sunderland, Sinauer Associates, Massachusetts, pp. 205–247.
    Portik, D.M., Travers, S.L., Bauer, A.M. & Branch, W.R. (2013) A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mount Namuli, an isolated ‘sky island’ of northern Mozambique. Zootaxa, 3710 (5), 415–435.
    Pyron, R.A., Burbrink, F.T. & Wiens, J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93.
    R Core Team. (2021). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: (accessed 12 November 2021)
    Rocha, S., Harris, D.J. & Posada, D. (2011) Cryptic diversity within the endemic prehensile-tailed gecko Urocotyledon inexpectata across the Seychelles Islands: patterns of phylogeographical structure and isolation at the multilocus level. Biological Journal of the Linnean Society, 104 (1), 177–191.
    Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenback, J.P. (2012) MrBayes 3.2 : Efficient Bayesian Phylogenetic Inference and Model Choice Across a Large Model Space. Software for Systematic and Evolution, 61 (3), 539–542.
    Stanley, E.L., Ceríaco, L.M.P., Bandeira, S., Valerio, 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.
    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6 : Molecular Evolutionary Genetics Analysis V. 6.0. Molecular Biology and Evolution, 30 (12), 2725–2729.
    Travers, S.L., Jackman, T.R. & Bauer, A.M. (2014) A molecular phylogeny of Afromontane dwarf geckos (Lygodactylus) reveals a single radiation and increased species diversity in a South African montane center of endemism. Molecular Phylogenetics and Evolution, 80, 31–42.
    Vaz Pinto, P., Luis Veríssimo, L. & Branch, W.R. (2019) Hiding in the bushes for 110 years: rediscovery of an iconic Angolan gecko (Afrogecko ansorgii Boulenger, 1907, Sauria: Gekkonidae). Amphibian & Reptile Conservation, 13 (2), 29–41.
    Vaz Pinto, P., Conradie, W., Becker, F. & Lobón-Rovira, J. (2021) Updated distribution of Kolekanus plumicaudus (Sauria: Gekkonidae), with some annotation on its natural history. Herpetology Notes, 14, 1207–1212.
    Vidal, N. & Hedges, S.B. (2005) The phylogeny of squamate reptiles (lizards, snakes, and amphisbaenians) inferred from nine nuclear protein-coding genes. Comptes Rendus - Biologies, 328 (10–11), 1000–1008.
    Vienne, D.M., De, Giraud, T. & Martin, O.C. (2007) A congruence index for testing topological similarity between trees. Bioinformatics, 23 (23), 3119–3124.
    Whiting, A.S., Bauer, A.M. & Sites, J.W. (2003) Phylogenetic relationships and limb loss in sub-Saharan African scincine lizards (Squamata: Scincidae). Molecular Phylogenetics and Evolution, 29 (3), 582–598.
    Xia, X. (2013) DAMBE5 : A Comprehensive Software Package for Data Analysis in Molecular Biology and Evolution. Molecular Biology and Evolution, 30 (7), 1720–1728.