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Issue: Vol. 5374 No. 3: 17 Nov. 2023
Type: Article
Published: 2023-11-17
DOI: 10.11646/zootaxa.5374.3.3
Page range: 361-389
Abstract views: 358
PDF downloaded: 18

The status of Tomopterna pulchra Boulenger, 1896 (Anura: Pyxicephalidae)

Kigelia Solutions/Frank Willems Guiding and Ecology; Chisamba Park; PO Box 12; Fringilla; Zambia
Unit for Environmental Sciences and Management; North-West University; Potchefstroom 2520; South Africa
Amphibia Africa Tomopterna taxonomy Miocene arid corridor

Abstract

Tomopterna pulchra (Boulenger, 1896) was described from “Lake Tanganyika” based on a single specimen. It was synonymised with Tomopterna tuberculosa (Boulenger, 1882) by Loveridge (1957) who noted that it did not fully agree with the description of T. tuberculosa, especially in the difference in dorsal colouration. Genetic differences show that two species are confused, although their advertisement calls and morphology are similar, except for the presence of a pale vertebral stripe mostly in eastern populations. We consider the advertisement calls, morphology, and genetic differences, and remove Tomopterna pulchra (Boulenger, 1896) from the synonymy of Tomopterna tuberculosa. Divergence dating shows that these two species separated in the mid-Miocene. Tomopterna pulchra breeds in rocky habitats, whereas T. tuberculosa breeds in sandy areas, typical of the genus. Based on genetically confirmed specimens, presently T. tuberculosa is known in the west (Angola and Namibia) while T. pulchra is only known from the east (Zambia and Tanzania). The examination of specimens for the presence or absence of a vertebral stripe infers that these species are widespread. Further work is required to determine if the two species are sympatric, and the extent of their ranges.

 

References

  1. Ahl, E. (1927 [“1925”]) Über vernachlässigte Merkmale bei Fröschen. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1925, 40–47.
  2. Anonymous. (1896) The Monthly Record. The Geographical Journal, 7 (4), 425–435. [https://www.jstor.org/stable/i303167]
  3. Balinsky, B.I. (1962) Patterns of animal distribution on the African Continent (summing-up talk). Annals of the Cape Provincial Museums, 2, 299–310.
  4. Bellstedt, D.U., Galley, C., Pirie, M.D. & Linder, H.P. (2012) The migration of the Palaeotropical arid flora: Zygophylloidea as an example. Systematic Botany, 37, 951–959. https://doi.org/10.1600/036364412X656608
  5. Bittencourt-Silva, G.B., Conradie, W., Siu-Ting, K., Tolley, K.A., Channing, A., Cunningham, M., Farooq, H.M., Menegon, M. & Loader, S.P. (2016) The phylogenetic position and diversity of the enigmatic mongrel frog Nothophryne Poynton, 1963 (Amphibia, Anura). Molecular Phylogenetics and Evolution, 99, 89–102. https://doi.org/10.1016/j.ympev.2016.03.021
  6. Bobe, R. (2006) The evolution of arid ecosystems in eastern Africa. Journal of Arid Environments, 66, 564–584. https://doi.org/10.1016/j.jaridenv.2006.01.010
  7. Bogart, J.P., Dawood, A., Becker, F.S. & Channing, A. (2022) Chromosomes in the African frog genus Tomopterna (Pyxicephalidae) and probing the origin of tetraploid Tomopterna tandyi. Genome, 65, 585–604. https://doi.org/10.1139/gen-2022-0053
  8. 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 of the United States of America, 97, 6585–6590. https://doi.org/10.1073/pnas.97.12.6585
  9. Bouckaert, R., Vaughan, T.G., Barido-Sottani, J., Duchêne, S., Fourment, M., Gavryushkina, A., Heled, J., Jones, G., Kühnert, D., De Maio, N. & Matschiner, M. (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS computational biology, 15 (4), e1006650. https://doi.org/10.1371/journal.pcbi.1006650
  10. Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. 2nd Edition. Taylor and Francis, London, 256 pp., 16 pls.
  11. Boulenger, G.A. (1896) Descriptions of two new frogs from Lake Tanganyika, presented to the British Museum. Annals and Magazine of Natural History, Series 6, 18, 467–468. https://doi.org/10.1080/00222939608680490
  12. 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 (2), 225–240.
  13. Caujape-Castells, J., Janse, R.K., Membrives, N., Pedrola-Monfort, J., Montserrat, J.M. & Ardanuy, A. (2001) Historical biogeography of Androcymbium Eilld. (Colchicaceae) in Africa: Evidence from cpDNA RFLPs. Botanical Journal of the Linnean Society, 136, 379–392. https://doi.org/10.1111/j.1095-8339.2001.tb00581.x
  14. Ceríaco, L.M., de Sá, S.D.A.C., Bandeira, S., Valério, H., Stanley, E.L., Kuhn, A.L., Marques, M.P., Vindum, J.V., Blackburn, D.C. & Bauer, A.M. (2016) 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, 63 (2), 15–61.
  15. Channing, A. & Becker, F. (2019) Correction to the type locality of Tomopterna ahli (Deckert, 1938) (Anura: Pyxicephalidae), with the designation of a neotype. Zootaxa, 4688 (4), 549–560. https://doi.org/10.11646/zootaxa.4688.4.6
  16. Channing, A. & Willems, F. (2018) A new grass frog with rupicolous tadpoles from northern Zambia (Anura: Ptychadenidae). Zootaxa, 4462 (3), 349–366. https://doi.org/10.11646/zootaxa.4462.3.3
  17. Channing, A. & du Preez, L.H. (2020). Taxonomic status of the cryptic sand frog, Tomopterna cryptotis (Anura, Pyxicephalidae). Alytes, 37 (3–4), 1–21.
  18. Channing, A. & Rödel, M.O. (2019) Field guide to the frogs & other amphibians of Africa. Penguin Random House South Africa, Cape Town, 408 pp.
  19. Channing, A., Moyer, D.C. & Dawood, A. (2004) A new sand frog from central Tanzania (Anura: Ranidae: Tomopterna). African Journal of Herpetology, 53 (1), 21–28. https://doi.org/10.1080/21564574.2004.9635495
  20. Coleman, M., Liston, A., Kadereit, J.W. & Abbott, R.J. 2003. Repeat intercontinental dispersal and Pleistocene speciation in disjunct Mediterranean and desert Senecio (Asteraceae) American Journal of Botany, 90, 1446–1454. https://doi.org/10.3732/ajb.90.10.1446
  21. Conradie, W., Keates, C., Verburgt, L., Baptista, N.L. & Harvey, J. (2023) Contributions to the herpetofauna of the Angolan Okavango-Cuando-Zambeze river drainages. Part 3: Amphibians. Amphibian & Reptile Conservation, 17 (1 & 2) [General Section], 19–56 (e325).
  22. Couvreur, T.L.P., Dauby, G., Blach-Overgaard, A., Deblauwe, V., Dessein, S., Droissart, V., Hardy, O.J., Harris, D.J., Janssens, S.B., Ley, A.C., Mackinder, B.A., Sonké, B., Ssosef, M.S.M., Stévart, T., Svenning, J.-C., Wieringa, J.J., Faye, A., Missoup, A.D., Tolley, K.A., Nicolas, V., Ntie, S., Fluteau, F., Robin, C., Guillocheau, F., Barboni, D. & Sepulchre, P. (2021) Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biological Reviews, 96, 16–51. https://doi.org/10.1111/brv.12644
  23. De Sá, R.O., Streicher, J.W., Sekonyela, R. Forlani, M.C., Loader, S.P., Greenbaum, E. Richards, S. & Haddad, C.F.B. (2012) Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology, 12, 241. https://doi.org/10.1186/1471-2148-12-241
  24. Davis, A.L. & Scholtz, C.H. (2010) Speciation and evolution of eco-climatic ranges in the intertropical African dung beetle genus, Diastellopalpus van Lansberge. Biological Journal of the Linnean Society, 99 (2), 407–423. https://doi.org/10.1111/j.1095-8312.2009.01367.x
  25. Dawood, A. & Channing, A. (2002) Description of a new cryptic species of African sand frog, Tomopterna damarensis (Anura: Ranidae), from Namibia. African Journal of Herpetology, 51 (2), 129–134. https://doi.org/10.1080/21564574.2002.9635468
  26. Dawood, A. & Stam, E.M. (2006) The taxonomic status of the monotypic frog genus Anhydrophryne Hewitt from South Africa: a molecular perspective. South African Journal of Science, 102 (5), 249–253.
  27. Dawood, A. & Uqubay, S.M. (2004) A molecular phylogeny of the sand frog genus Tomopterna (Amphibia: Anura: Ranidae) based on mitochondrial 12S and 16S rRNA sequences. African Zoology, 39, 145–151.
  28. de Sá, R.O., Streicher, J.W., Sekonyela, R., Forlani, M.C., Loader, S.P., Greenbaum, E., Richards, S. & Haddad, C.F. (2012) Molecular phylogeny of microhylid frogs (Anura: Microhylidae) with emphasis on relationships among New World genera. BMC Evolutionary Biology, 12, 1–21. https://doi.org/10.1186/1471-2148-12-241
  29. De Witte, G.F. (1930) Liste des batraciens du Congo Belge (Collecton du Musée du Congo Belge à Tervuren). Première partie. Revue de Zoologie et de Botanique Africaines. Tervuren, 19, 232–273.
  30. Deckert K. (1938) Beiträge zur Osteologie und Systematik ranider Froschlurche. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1938, 127–184.
  31. Dubois, A. (1987 “1986”) Miscellanea taxinomica batrachologica (I). Alytes, 5, 7–95.
  32. Freitag, S. & Robinson, T.J. (1993) Phylogeographic patterns in mitochondrial DNA of the ostrich (Struthio camelus). The Auk, 110, 614–622. https://doi.org/10.2307/4088425
  33. Freitas, E.S., Bauer, A.M., Siler, C.D., Broadley, D.G. & Jackman, T.R. (2018) Phylogenetic and morphological investigation of the Mochlus afer-sundevallii species complex (Squamata: Scincidae) across the arid corridor of sun-Saharan Africa. Molecular Phylogenetics and Evolution, 127, 280–287. https://doi.org/10.1016/j.ympev.2018.04.020
  34. Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F., De Sa, R.O., Channing, A., Wilkinson, M., Donnellan, S.C. & Raxworthy, C.J. (2006) The amphibian tree of life. Bulletin of the American Museum of natural History, 297, 1–291. https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
  35. Frost, D.R. (2023) Amphibian Species of the World: An Online Reference. Version 6.2. 2 October 2023. Electronic Database. American Museum of Natural History, New York, New York. Available from: https://amphibiansoftheworld.amnh.org/index.php. (accessed 2 October 2023)
  36. Günther, A.C.L.G. (1865 “1864”) Descriptions of new species of batrachians from West Africa. Proceedings of the Zoological Society of London, 1864, 479–482.
  37. Haas, A, Mitgutsch, C., Hertwig, S., Dawood, A. & Channing, A. (2006) The tadpole of Tomopterna luganga Channing, Moyer & Dawood, 2004 (Anura: Ranidae). Salamandra, 42 (1), 1–12.
  38. Hasegawa, M., Kishino, H. & Yano, T. (1985) Dating the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution, 22, 160–174. https://doi.org/10.1007/BF02101694
  39. Heinicke, M.P., Ceríaco, L.M., Moore, I.M., Bauer, A.M. & Blackburn, D.C. (2017) Tomopterna damarensis (Anura: Pyxicephalidae) is broadly distributed in Namibia and Angola. Salamandra, 53 (3), 461–465.
  40. Heinz, M.D., Brennan, I.G., Jackman, T.R. & Bauer, A.M. (2021) Phylogeny of the genus Chondrodactylus (Squamata: Gekkonidae) with the establishment of a stable taxonomy. Bulletin of the Museum of Comparative Zoology, 163, 151–210. https://doi.org/10.3099/0027-4100-163.5.151
  41. Heled, J. & Drummond, A.J. (2012) Calibrated tree priors for relaxed phylogenetics and divergence time estimation. Systematic Biology, 61, 138–149. https://doi.org/10.1093/sysbio/syr087
  42. Hillis, D.M., Mable, B.K., Larson, A., Davis, S.K. & Zimmer, E.A. (1996) Nucleic acids IV: Sequencing and cloning. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics. 2nd Edition. Sinauer Associates, Sunderland, Massachusetts, pp 321–381
  43. Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2017) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35, 518–522. https://doi.org/10.1093/molbev/msx281
  44. Jürgens, N. (1997) Floristic biodiversity and history of African arid regions. Biodiversity and Conservation, 6, 495–514. https://doi.org/10.1023/A:1018325026863
  45. K. Lisa Yang Center for Conservation Bioacoustics (2023) Raven Pro: Interactive Sound Analysis Software. Version 1.6.5. The Cornell Lab of Ornithology, Ithaca, New York, New York. Available from: https://ravensoundsoftware.com/ (accessed 1 November 2023)
  46. Kalyaanamoorthy, S., Minh, Q., Wong, T.K.F., Von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589. https://doi.org/10.1038/nmeth.4285
  47. Kirk-Spriggs, A.H. & McGregor, G. (2009) Disjunctions in the Diptera (Insecta) fauna of the Mediterranean Province and southern Africa and a discussion of biogeographical considerations. Transactions of the Royal Society of South Africa, 64, 32–52. https://doi.org/10.1080/00359190909519236
  48. Kissling, W.D., Black-Overgaard, A., Zwaan, R.E. & Wagner, P. (2016) Historical colonization and dispersal limitation supplement climate and topography in shaping species richness of African lizards (Reptilia: Agaminae). Scientific Reports, 6, 34014. https://doi.org/10.1038/srep34014
  49. Kosuch, J., Vences, M., Dubois, A., Ohler, A. & Böhme, W. (2001) Out of Asia: mitochondrial DNA evidence for an oriental origin of tiger frogs, genus Hoplobatrachus. Molecular Phylogenetics and Evolution, 21 (3), 398–407. https://doi.org/10.1006/mpev.2001.1034
  50. Lancaster, N. (1981) Paleoenvironmental implications of fixed dune systems in southern Africa. Palaeogeography Palaeoclimatology Palaeoecology, 33, 327–346. https://doi.org/10.1016/0031-0182(81)90025-0
  51. Laurent, R.F. (1954) Reptiles et batraciens de la région de Dundo (Angola) (Deuxième note). Das publicações culturais da companhia de diamantes de Angola, 23, 35–84.
  52. Laurent, R.F. (1964) Reptiles et Amphibiens de l’Angola. Trosième contribution. Museu do Dondo, 67, 1–160.
  53. Lehtinen, R.M., Richards, C.M. & Nussbaum, R.A. (2004) Origin of a complex reproductive trait: phytotelm breeding in mantelline frogs. Ecology and Evolution of Phytotelm-Breeding Anurans. Museum of Zoology, University of Michigan, Ann Arbor, Michigan, pp. 45–54.
  54. Lobón-Rovira, J., Vaz Pinto, P., Becker, F.S., Tolley, K.A., Measey, J., Bennet, B., Boon, B., de Sá, S. & Conradie, W. (2022) An updated herpetofaunal species inventory of Iona National Park in southwestern Angola. Check List, 18, 289–321. https://doi.org/10.15560/18.2.289
  55. Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya; Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, 117, 153–362.
  56. Meester, J. (1965) The origins of southern African mammal fauna. Zoologica Africana, 1, 87–93. https://doi.org/10.1080/00445096.1965.11447302
  57. Mertens, R. (1971) Die Herpetofauna Südwest-Afrikas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 529, 1–110
  58. Monard, A. (1937 “1936”) Contribution à la batrachologie d’Angola. Bulletin de la Société Neuchâteloise des Sciences Naturelles, 62, 5–59.
  59. Nyblade, A.A. & Robinson, S.W. (1994) The African Superswell. Geophysical Research Letters, 21, 765–768. https://doi.org.10.1029/94GL00631
  60. 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, 268–274. https://doi.org/10.1093/molbev/msu300
  61. Parker, H.W. (1936) Dr. Karl Jordan’s expedition to South-West Africa and Angola: Herpetological collections. Novitates Zoologicae, 40, 115–146.
  62. Passmore, N.I. & Carruthers, V.C. (1975) A new species of Tomopterna (Anura: Ranidae) from the Kruger National Park, with notes on related species. Koedoe, 18, 31–50. https://doi.org/10.4102/koedoe.v18i1.913
  63. Perkins, J.S. (2020) Take me to the river along the African drought corridor: Adapting to climate change. Botswana Journal of Agriculture and Applied Sciences, 14, 60–71. https://doi.org/10.37106/bojaas.2020.77
  64. Poynton, J.C. (1964) The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum, 17, 1–334.
  65. Poynton, J.C. (1995) The “arid corridor” distribution in Africa: a search for instances among amphibians. Madoqua, 19, 45–48.
  66. Poynton, J.C. (2000) Evidence for an Afrotemperate amphibian fauna. African Journal of Herpetology, 49, 33–41. https://doi.org/10.1080/21564574.2000.9650014
  67. Poynton J.C. & Broadley D.G. (1985) Amphibia Zambesiaca 2. Ranidae. Annals of the Natal Museum, 27, 115–181.
  68. Prendini, L. (2001) Phylogeny of Parabuthus (Scorpiones, Buthidae). Zoologica Scripta, 30, 13–35. https://doi.org/10.1046/j.1463-6409.2001.00056.x
  69. Rambaut, A. (2018) FigTree, a graphical viewer of phylogenetic trees. Version 1.4.4. Institute of evolutionary biology, University of Edinburgh. [program]
  70. Reeder, J.E. (2019) Amphibians of northern KwaZulu-Natal: A phylogenetic study. M. Sc. Dissertation, North-West University, Potchefstroom, 102 pp.
  71. Sabaj, M.H. (2022) Codes for Natural History Collections in Ichthyology and Herpetology. Online Supplement. Version 9.0. 14 February 2022. American Society of Ichthyologists and Herpetologists, Washington, D.C. Electronically accessible. Available from: https://asih.org (accessed 1 November 2023)
  72. Schmidt, K.P. & Inger, R.F. (1959) Amphibians exclusive of the genera Afrixalus and Hyperolius. Exploration du Parc National de l’Upemba. Mission G.F. de Witte (1946–1949), 56, 1–264.
  73. 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. https://doi.org/10.1093/aesa/87.6.651
  74. Swofford, D.L. (2002) PAUP* Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]
  75. Thiv, M., Van der Niet, T., Rutschmann, F., Thulin, M., Brune, T. & Linder, H.P. (2011) Old–New World and trans-African disjunctions of Thamnosa (Rutaceae): Intercontinental long-distance dispersal and local differentiation in the succulent biome. American Journal of Botany, 98, 76–87. https://doi.org/10.3732/ajb.1000339
  76. Tolley, K.A., Braae, A. & Cunningham, M. (2010) Phylogeography of the Clicking Stream Frog Strongylopus grayii (Anura, Pyxicephalidae) reveals cryptic divergence across climatic zones in an abundant and widespread taxon. African Journal of Herpetology, 59 (1), 17–32. https://doi.org/10.1080/04416651003744943
  77. Tolley, K.A., Telford, N.S., Makhubo, B.G., Power, R.J. & Alexander, G.J. (2023) Filling the gap: Noteworthy herpetological discoveries in North West Province, South Africa. Zoosystematics and Evolution, 99 (1), 101–116. https://doi.org/10.3897/zse.99.90181
  78. Trifinopoulos, J., Nguyen, L.-T., Von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, W1, W232–W235. https://doi.org/10.1093/nar/gkw256
  79. Temminck, C.J. & Schlegel, H. (1838) Les Batrachiens. In: Siebold, P.F. (Ed.), Fauna Japonica, Reptilia. Vol. 144. Müller, J. et co., Amsterdam, pp. 105–137, pls. 3–8.
  80. Van der Meijden, A., Crottini, A., Tarrant, J., Turner, A. & Vences, M. (2011) Multi-locus phylogeny and evolution of reproductive modes in the Pyxicephalidae, an African endemic clade of frogs. African Journal of Herpetology, 60 (1), 1–12. https://doi.org/10.1080/21564574.2010.523904
  81. Van Zinderen Bakker, E.M. (1975) The origin and palaeoenvironment of the Namib Desert Biome. Journal of Biogeography, 2, 65–73. https://doi.org/10.2307/3038074
  82. Vences, M., Glaw, F., Kosuch, J., Das, I. & Veith, M. (2000) Polyphyly of Tomopterna (Amphibia: Ranidae) based on sequences of the mitochondrial 16S and 12S rRNA genes, and ecological biogeography of Malagasy relict amphibian groups. Diversité et endéémisme aMadagascar/Diversity and endemism in Madagascar. Mémoires de la Société de Biogéographie, Pairs, pp. 229–242.
  83. Vences, M., Köhler, J., Crottini, A. & Glaw, F. (2010) High mitochondrial divergence meets morphological and bioacoustic conservatism: Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar. Bonn zoological Bulletin, 57, 241–255.
  84. Visser, J.H., Bennett, N.C. & Jansen van Vuuren, B. (2019) Phylogeny and biogeography of the Bathyergidae: a review of patterns and processes. PeerJ, 7, e7730. https://doi.org/10.7717/peerj.7730
  85. 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 (Squamatra: Agamidae). Journal of Natural History, 52, 771–817. https://doi.org/10.1080/00222933.2018.1435833
  86. Wasonga, D.V. & Channing, A. (2013) Identification of sand frogs (Anura: Pyxicephalidae: Tomopterna) from Kenya with the description of two new species. Zootaxa, 3734 (2), 221–240. https://doi.org/10.11646/zootaxa.3734.2.7
  87. Wichura, H., Bousquest, R., Oberhänsli, R., Strecker, M.R. & Trauth, M.H. (2010) Evidence for middle Miocene uplift of the East African Plateau. Geology, 38, 543–546. https://doi.org/10.1130/G31022.1
  88. Wilson, L. & Channing, A. (2019) A new sand frog from Namaqualand, South Africa (Pyxicephalidae: Tomopterna). Zootaxa, 4609, 225–246. https://doi.org/10.11646/zootaxa.4609.2.2
  89. Wilfert, L., Kaib, M., Durka, W. & Brandl, R. (2006) Differentiation between populations of a termite in eastern Africa: implications for biogeography. Journal of Biogeography, 33, 1993–2000. https://doi.org/10.1111/j.1365-2699.2006.01556.x
  90. Winterbottom, J.M. (1967) Climatological implications of avifaunal resemblances between south western Africa and Somaliland. Palaeoecology of Africa and the Surrounding Islands, 2, 77–79.
  91. Wüster, W. & Broadley, D.G. (2007) Get an eyeful of this: a new species of giant spitting cobra from eastern and north-eastern Africa (Squamata: Serpentes: Elapidae: Naja). Zootaxa, 1532 (1), 51–68. https://doi.org/10.11646/zootaxa.1532.1.4
  92. Zimkus, B.M. & Larson, J.G. (2011) Examination of the molecular relationships of sand frogs (Anura: Pyxicephalidae: Tomopterna) and resurrection of two species from the Horn of Africa. Zootaxa, 2933 (1), 27–45. https://doi.org/10.11646/zootaxa.2933.1.2
  93. Zhang, P., Liang, D., Mao, R.L., Hillis, D.M., Wake, D.B. & Cannatella, D.C. (2013) Efficient sequencing of anuran mtDNAs and a mitogenomic exploration of the phylogeny and evolution of frogs. Molecular Biology and Evolution, 30 (8), 1899–1915. https://doi.org/10.1093/molbev/mst091