Abstract
Recent integrative taxonomic studies of the agamid genus Acanthocercus Fitzinger, 1843 have shown that Angola harbors three different taxa, all within the Acanthocercus atricollis (Smith, 1849) species complex—A. cyanocephalus (Falk, 1925) in the northeastern parts of the country, A. margaritae Wagner et al. 2021 in the southern regions, and an unnamed species in the central and northwestern parts of Angola. Using the previously published molecular data as evidence of phylogenetic support and newly collected morphological, meristic and coloration data, we here describe this unnamed lineage as a new species. The new species is morphologically very similar to A. cyanocephalus, but it can easily be differentiated from the latter by the coloration pattern of displaying males, with a blue coloration restricted to the head region, and by its inferior scale counts compared to other species of the A. atricollis complex group. As reported in other studies in this group, male breeding coloration is an effective trait for diagnosing these morphologically conserved species. This description raises the number of Acanthocercus species recognized to 15 and is another contribution revealing the rich but still incompletely described herpetological diversity of Angola.
References
Agarwal, I., Ceríaco, L.M.P., Metallinou, M., Jackman, T. & Bauer, A.M. (2021) How the African house gecko (Hemidactylus mabouia) conquered the world. Royal Society Open Science, 8, 210749. https://doi.org/10.1098/rsos.210749
Angel, F. (1923) Reptiles. In: Rohan-Chabot, J.F. (Ed.), Mission Rohan-Chabot, Angola et Rhodesia (1912–1914). Tome IV. Histoire Naturelle. Fascicule 1 Mammifères (Anatomie Comparée—Embryologie), Oiseaux—Reptiles—Poissons. Imprimerie Nationale, Paris, pp. 157–170, 1 pl.
Baptista, N.L., António, T. & Branch, W.R. (2019) The herpetofauna of Bicuar National Park and surroundings, southwestern Angola: a preliminary checklist. Amphibian and Reptile Conservation, 13 (2), 96–130.
Baptista, N., Pinto, P.V., Keates, C., Edwards, S., Rodel, M., Conradie, W. (2021). A new species of red toad, Schismaderma Smith, 1849 (Anura: Bufonidade), from central Angola. Zootaxa, 5081 (3), 301–332. https://doi.org/10.11646/zootaxa.5081.3.1
Bocage, J.V.B. (1866) Lista de reptis das possessões potuguezas d’Africa ocidental que existem no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes, 1, 37–56.
Bocage, J.V.B. (1879) Subsidios para a fauna das possessões portuguezas d’Africa ocidental – III. Sertão de Angola, do Bihé ao Cassengue. Jornal de Sciencias Mathematicas, Physicas e Naturaes, 7, 90–96.
Bocage, J.V.B. (1895) Herpétologie d’Angola et du Congo. Ministério da Marinha e das Colónias, Lisbonne, 203 pp., 20 pls.
Boulenger, G.A. (1885) Catalogue of the Lizards in the British Museum (Natural History). Volume 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Trustees of the British Museum, London, xii + 436 pp., pls. 1–32.
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. https://doi.org/10.1080/03745480509443656
Branch, W.R. & Conradie, W. (2015) IV. Herpetofauna da região da Lagoa Carumbo. In: Huntley, B. & Francisco, P. (Eds), Avaliação Rápida da Biodiversidade da Lagoa Carumbo, Lunda-Norte—Angola. Ministério do Ambiente de Angola, Luanda, Angola, pp. 194–209.
Branch, W.R., Conradie, W., Vaz Pinto, P. & Tolley, K.A. (2019a) Another Angolan Namib endemic species: a new Nucras Gray, 1838 (Squamata: Lacertidae) from southwestern Angola. Amphibian & Reptile Conservation, 13 (2), 82–95.
Branch, W.R., Vaz Pinto, P., Baptista, N. & Conradie, W. (2019b) The reptiles of Angola: History, diversity, endemism and hotspots. In: Huntley, B.J., Russo, V., Lages, F. & Ferrand, N. (Eds.), Biodiversity of Angola. Science & Conservation: A Modern Synthesis. Springer, Cham, pp. 283–334. https://doi.org/10.1007/978-3-030-03083-4_13
Branch, W.R., Schmitz, A., Lobón-Rovira, J., Baptista, N.L., António, T & Conradie, W. (2021) Rock island melody: A revision of the Afroedura bogerti Loveridge, 1944 group, with descriptions of four new endemic species from Angola. Zoosystematics and Evolution, 87 (1), 55–82. https://doi.org/10.3897/zse.97.57202
Butler, B.O., Ceríaco, L.P.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 (2), 225–240.
Ceríaco, L.M.P., Bauer, A.M., 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., Marques, M.P. & Bandeira, S.A. (2016) Anfíbios e répteis do Parque Nacional da Cangandala. Instituto Nacional da Biodiversidade e Áreas de Conservação, Angola & Museu Nacional de História Natural e da Ciência, Lisboa, 97 pp.
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. https://doi.org/10.11646/zootaxa.4527.1.12
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. https://doi.org/10.3897/zookeys.780.25859
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 and reptiles of Malanje Province, Central Angola. Herpetological Review, 49 (3), 408–431.
Ceríaco, L.M.P., 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. https://doi.org/10.11646/zootaxa.4746.1.1
Ceríaco, L.M.P., Heinicke, M.P., Parker, K.L., 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 (1), 77–112. https://doi.org/10.11646/zootaxa.4747.1.3
Ceríaco, L.M.P., Agarwal, I., Marques, M.P. & Bauer, A.M. (2020c) A correction to a recent review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two additional species. Zootaxa, 4861 (1), 92–106. https://doi.org/10.11646/zootaxa.4861.1.6
Ceríaco, L.M.P., Santos, B.S., Marques, M.P., Bauer, A.M. & Tiutenko, A. (2021) Citizen science meets specimens in old formalin filled jars: A new species of Banded Rubber Frog, genus Phrynomantis (Anura: Microhylidae) from Angola. Alytes, 38 (1–4), 18–48.
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, southwestern Angola. Zootaxa, 3269 (1), 1–17. https://doi.org/10.11646/zootaxa.3269.1.1
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 southwestern Angola. African Journal of Herpetology, 61 (2), 91–112. https://doi.org/10.1080/21564574.2012.676079
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. https://doi.org/10.11646/zootaxa.3635.3.1
Conradie, W., Deepak, V., Keates, C. & Gower, D.J. (2020a) Kissing cousins: a review of the African genus Limnophis Günther, 1865 (Colubridae: Natricinae), with the description of a new species from north-eastern Angola. African Journal of Herpetology, 69 (1), 79–107. https://doi.org/10.1080/21564574.2020.1782483
Falk, K. (1925) Herpetologische Berichte aus Angola (Portugiesisch West-Afrika). Blätter für Aquarien- und Terrarienkunde, 36, 81–83.
Ferreira, J.B. (1900) Sobre alguns exemplares pertencentes à fauna do norte de Angola. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 6, 48–54.
Ferreira, J.B. (1903) Reptis de Angola da região norte do Quanza da collecção Pereira do Nascimento (1902). Jornal de Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 7, 9–16.
Frade, F. (1963) Linhas gerais da distribuição dos vertebrados em Angola. Memórias da Junta de Investigação do Ultramar, Segunda Série, 43, 241–257.
Grandvaux-Barbosa, L.A. (1970) Carta Fitogeografica de Angola. Instituto de Investigação Científica de Angola, Luanda, 323 pp.
Greenbaum, E., Allen, K.E., Vaughan, E.R., Pauwels, O.S.G., Wallach, V., Kusamba, C., Muninga, W.M., Aristote, M.M., Mali, F.M.M., Badjedjea, G., Penner, J., Rödel, M.-O., Rivera, J., Sterkhova, V., Johnson, G., Tapondjou N.W.P. & Brown, R.M. (2021) Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata: Colubridae) with descriptions of two new cryptic species from Central Africa. Zootaxa, 4965 (1), 1–44. https://doi.org/10.11646/zootaxa.4965.1.1
Hallerman, J., Ceríaco, L.M.P., 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. https://doi.org/10.1080/21564574.2020.1777470
Klausewitz, W. (1957) Eidonomische Untersuchungen über die Rassenkreise Agama cyanogaster und Agama atricollis. 2. Die Unterarten von Agama atricollis. Senckenbergiana biologica, 38, 157–174.
Laurent, R.F. (1950) Reptiles et batraciens e la région de Dundo (Angola du Nord-Est) (Première note). Publicações Culturais da Companhia de Diamantes de Angola. s.n., Lisboa, 17 pp.
Laurent, R.F. (1964) Reptiles et amphibiens de l’Angola (Troisième contribution). Publicações Culturais da Companhia de Diamantes de Angola, 67, 1–165.
Leaché, A.D., Wagner, P., Linkem, C.W. , Böhme, W., Papenfuss, T.J., Chong, R.A., Lavin, B.R., Bauer, A.M., Nielsen, S.V., Greenbaum, E., Rödel, M.O., Schmitz, A., LeBreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E.A., El Din, S.B., Lemmon, A.R. & Burbrink, F.T. (2014) A hybrid phylogenetic—phylogenomic approach for species tree estimation in African Agama lizards with applications to biogeography, character evolution, and diversification. Molecular Phylogenetics and Evolution, 79, 215–230. https://doi.org/10.1016/j.ympev.2014.06.013
Lobón-Rovira, J., Conradie, W., Iglesias, D.B., Ernst, R., Verisimmo, L., Baptista, N. & Pinto, P.V. (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. https://doi.org/10.3897/vz.71.e64781
Loveridge, A. (1957) Check list of the reptiles and amphibians of east Africa (Uganda, Kenya, Tanganyika, Zanzibar). Bulletin of the Museum of Comparative Zoology Harvard, 117 (2), 153–362.
Manaças, S. (1963) Sáurios de Angola. Memórias da Junta de Investigação do Ultramar, 43 (2), 223–240.
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 (1850–2017). Proceedings of the California Academy of Sciences, 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 the Congo. Zootaxa, 4568 (1), 51–68. https://doi.org/10.11646/zootaxa.4568.1.3
Marques, M.P., Ceríaco, L.M.P., Stanley, E.L., Bandeira, S.A., Agarwal, I. & Bauer, A.M. (2019b) A new species of Girdled Lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola. Zootaxa, 4668 (4), 503–524. https://doi.org/10.11646/zootaxa.4668.4.4
Marques, M.P., Ceríaco, L.M.P., Buehler, M.D., Bandeira, S.A., 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, 4853 (3), 301–352. https://doi.org/10.11646/zootaxa.4853.3.1
Monard, A. (1937) Contribution à l’herpétologie d’Angola. Arquivos do Museu Bocage, 8, 19–154.
Parrinha, D., Marques, M.P., Heinicke, M.P., Khalid, F., Parker, K.L., Tolley, K.A., Childers, J.L., Conradie, W.C., Bauer, A.M. & Ceríaco, L.M.P. (2021) A revision of Angolan species in the genus Pedioplanis Fitzinger (Squamata: Lacertidae), with the description of a new species. Zootaxa, 5032 (1), 001–046. https://doi.org/10.11646/zootaxa.5032.1.1
Parker, H.W. (1936) Dr. Karl Jordan’s expedition to South-West Africa and Angola: herpetological collections. Novitates Zoologicae, 40, 115–146.
Peters, W.C.H. (1881) Zwei neue von Herrn Major von Mechow während seiner letzten Expedition nach west-Afrika entdeckte Schlangen und eine übersicht der von ihm mitgebrachten herpetologischen Sammlung. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1881, 147–150.
Schmidt, K.P. (1933) The reptiles of the Pulitzer-Angola expedition. Annals of the Carnegie Museum, 25, 1–15, pls. 1–2.
Smith, A. (1849) Appendix to Illustrations of the zoology of South Africa; consisting chiefly of figures and descriptions of the objects of natural history collected during an expedition into the interior of South Africa, in the years 1834, 1835, and 1836; fitted out by “The Cape of Good Hope Association for Exploring Central Africa”. Smith, Elder and Co., London, 28 pp. https://doi.org/10.5962/bhl.title.120442
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. https://doi.org/10.11646/zootaxa.4061.3.1
Tamar, K., Scholz, S., Crochet, P.-A., Geniez, P., Meiri, S., Schmitz, A., Wilms T. & Carranza, S. (2016) Evolution around the Red Sea: Systematics and biogeography of the agamid genus Pseudotrapelus (Squamata: Agamidae) from North Africa and Arabia. Molecular Phylogenetics and Evolution, 97, 55–68. https://doi.org/10.1016/j.ympev.2015.12.021
Themido, A.A. (1941) Répteis e batráquios das colónias portuguesas (Catálogo das colecções do Museu Zoológico de Coimbra). Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 119, 1–28.
Uetz, P., Freed, P. & Hošek, J. (2021) The Reptile Database. Available from: http://www.reptile-database.org (accessed 16 September 2021)
Wagner, P., Greenbaum, E. & Bauer, A.M. (2012) A new species of the Acanthocercus atricollis complex (Squamata: Agamidae) from Zambia. Salamandra, 48 (1), 21–30.
Wagner, P., Greenbaum, E., Bauer, A.M., Kusamba, C. & Leaché, A.D. (2018) Lifting the blue-headed veil – integrative taxonomy of the Acanthocercus atricollis species complex (Squamata: Agamidae). Journal of Natural History, 52, 771–817. https://doi.org/10.1080/00222933.2018.1435833
Wagner, P., Butler, B., Ceríaco, L.M.P. & Bauer, A.M. (2021) A new species of the Acanthocercus atricollis (Smith, 1849) complex (Squamata: Agamidae). Salamandra, 57 (4), 449–463.