Abstract
A new species of Tomopterna is described from Banhine National Park, southern Mozambique. It differs from all other sand frogs by an uncorrected p-distance of 4.9–8.7% for 16S rRNA. The new species can be distinguished from all other described species of sand frogs by: males 37 mm SVL or less, undivided subarticular tubercles, more than three phalanges free of webbing on the fourth toe, a continuous glandular ridge below the tympanum, a distinct tympanic membrane, the nostrils closer to the snout tip than the eye, the rounded canthus rostralis, a round tympanum, the absence of an outer metatarsal tubercle, an interrupted inter-orbital bar, small dorsal warts, the absence (or weakness) of vomerine teeth, the solid pigmented throat in mature males, the distinct palmar tubercles, the smooth ventral surface, and barring on the limbs. This description adds a further amphibian to the list of Mozambican frogs, which has increased significantly in recent years.
References
- Bittencourt-Silva, G.B. (2019) A herpetological survey of western Zambia. Amphibian & Reptile Conservation, 13 (e181), 1–28.
- Bittencourt-Silva, G.B., Bayliss, J. & Conradie, W. (2020) First herpetological surveys of Mount Lico and Mount Socone, Mozambique. Amphibian & Reptile Conservation, 14 (2) [General Section], e247, 198–217.
- Blackburn, D.C., Nielsen, S.V., Barej, M.F., Doumbia, J., Hirschfeld, M., Kouamé, N.G., Lawson, D., Loader, S., Ofori-Boateng, C., Stanley, E.L. & Rödel, M.-O. (2020) Evolution of the African slippery frogs (Anura: Conraua), including the world’s largest living frog. Zoologica Scripta, 49, 684–696. https://doi.org/10.1111/zsc.12447 DOI: https://doi.org/10.1111/zsc.12447
- 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 DOI: https://doi.org/10.1139/gen-2022-0053
- Bossuyt, F., Brown, R.M., Hillis, D.M., Cannatella, D.C. & Milinkovitch, M.C. (2006) Phylogeny and Biogeography of a cosmopolitan frog radiation: Late Cretaceous diversification resulted in continent-scale endemism in the family Ranidae. Systematic Biology, 55, 579–594. https://doi.org/10.1080/10635150600812551 DOI: https://doi.org/10.1080/10635150600812551
- Braae, A. (2007) Genetic structuring in the clicking stream frog (Strongylopus grayii); evidence of a cryptic species? B.Sc Honours Thesis, University of Cape Town, Cape Town. [unknown pagination]
- Branch, W.R. (2004) Herpetological Survey of The Niassa Game Reserve: Report prepared for Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa. Bayworld, Port Elizabeth. [unknown pagination]
- Buruwate, T.C. & Lloyd-Jones, D.J. (2024) Amphibian and Reptile Diversity of Niassa Special Reserve, Northern Mozambique. Journal of East African Natural History, 113 (1), 1–18. https://doi.org/10.2982/028.113.0101 DOI: https://doi.org/10.2982/028.113.0101
- Ceríaco, L.M.P., 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, Series 4, 63, 15–61.
- 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 DOI: https://doi.org/10.11646/zootaxa.4688.4.6
- Channing, A., Dehling, J.M., Lötters, S. & Ernst, R. (2016) Species boundaries and taxonomy of the African river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa, 4155 (1), 1–76. https://doi.org/10.11646/zootaxa.4155.1.1 DOI: https://doi.org/10.11646/zootaxa.4155.1.1
- Channing, A. & Du Preez, L.H. (2020) Taxonomic status of the cryptic sand frog, Tomopterna cryptotis (Anura, Pyxicephalidae). Alytes, 37, 1–21.
- Channing, A. Moyer, D.C. & Dawood, A. (2004) A new sand frog from Tanzania (Anura: Ranidae: Tomopterna). African Journal of Herpetology, 53, 21–28. https://doi.org/10.1080/21564574.2004.9635495 DOI: https://doi.org/10.1080/21564574.2004.9635495
- Channing, A., Schmitz, A., Zancolli, G., Conradie, W. & Rödel, M.-O. (2022) Phylogeny and taxonomy of the African frog genus Strongylopus (Anura: Pyxicephaliidae). Revue suisse de Zoologie, 129, 243–281. https://doi.org/10.35929/RSZ.0074 DOI: https://doi.org/10.35929/RSZ.0074
- Channing, A. & Rödel, M.-O. (2019) Field Guide to the Frogs and other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.
- Conradie, W., Bittencourt-Silva, G.B., Engelbrecht, H.M., Loader, S.P., Menegon, M., Nanvonamuquitxo, C., Scott, M. & Tolley K.A. (2016) Exploration into the hidden world of Mozambique’s sky island forests: new discoveries of reptiles and amphibians. Zoosystematics and Evolution, 92 (2), 163–180. https://doi.org/10.3897/zse.92.9948 DOI: https://doi.org/10.3897/zse.92.9948
- Conradie, W., Bittencourt-Silva, G.B., Farooq, H.M., Loader, S.P., Menegon, M. & Tolley, K.A. (2018a) New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs. African Journal of Herpetology, 67, 61–85. https://doi.org/10.1080/21564574.2017.1376714 DOI: https://doi.org/10.1080/21564574.2017.1376714
- Conradie, W., Verburgt, L., Portik, D.M., Ohler, A., Bwong, B.A. & Lawson. L.P. (2018b) A new reed frog (Hyperoliidae: Hyperolius) from coastal northeastern Mozambique. Zootaxa, 4379 (2), 177–198. https://doi.org/10.11646/zootaxa.4379.2.2 DOI: https://doi.org/10.11646/zootaxa.4379.2.2
- 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, 129–134. https://doi.org/10.1080/21564574.2002.9635468 DOI: https://doi.org/10.1080/21564574.2002.9635468
- Dawood, A., Channing, A. & Bogart, J.P. (2002) A molecular phylogeny of the frog genus Tomopterna in southern Africa: examining species boundaries with mitochondrial 12S rRNA sequence data. Molecular Phylogenetics and Evolution, 22, 407–413. https://doi.org/10.1006/mpev.2001.1060 DOI: https://doi.org/10.1006/mpev.2001.1060
- 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.
- 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.
- 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 DOI: https://doi.org/10.1186/1471-2148-12-241
- Du Preez, L.H., Netherlands, E.C., Rödel, M.-O. & Channing, A. (2024) A new bullfrog from southern Africa (Pyxicephalidae, Pyxicephalus Tschudi, 1838). African Journal of Herpetology, 73, 61–89. https://doi.org/10.1080/21564574.2023.2296654 DOI: https://doi.org/10.1080/21564574.2023.2296654
- Farooq, H.O. & Conradie, W. (2015a) A second record of Scolecomorphus kirkii Boulenger, 1883 (Gymnophiona: Scolecomorphidae) for Mozambique. Herpetology Notes, 8, 59–62.
- Farooq, H., Liedtke, H.C., Bittencourt-Silva, G., Conradie, W & Loader, S.P. (2015b) The distribution of Mertensophryne anotis with a new record in northern Mozambique. Herpetology Notes, 8, 305–307.
- Farooq, H., Nanvonamuquitxo, C., Nassongole, B., Conradie, W., Bills, R., Soares, A. & Antonelli, A. (2022) Shedding light on a biodiversity dark spot: survey of amphibians and reptiles of Pemba Region in northern Mozambique. Herpetological Conservation and Biology, 17, 423–432.
- Frost, D.R. (2024) Amphibian Species of the World: an Online Reference. Version 6.2. American Museum of Natural History, New York, New York. Electronic Database. Available from: https://amphibiansoftheworld.amnh.org/index.php (accessed 18 December 2024) https://doi.org/10.5531/db.vz.0001 DOI: https://doi.org/10.5531/db.vz.0001
- Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., De Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.O., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P. Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370. https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2 DOI: https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
- Guindon, S., Dufayard, J.-F., Lefort, V., Anismova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321. https://doi.org/10.1093/sysbio/syq010 DOI: https://doi.org/10.1093/sysbio/syq010
- Haas, A., Mitgutsch, C., Hertwig, S., Dawood, A. & Channing, A. (2006) The tadpole of Tomopterna luganga Channing, Moyer & Dawood, 2004 (Anura: Ranidae). Salamandra, 42, 1–12.
- Heinicke, M.P., Ceríaco, L.M.P., Moore, I.M., Bauer, A.M. & Blackburn, D.C. (2017) Tomopterna damarensis (Anura: Pyxicephalidae) is broadly distributed in Namibia and Angola. Salamandra, 53, 461–465.
- 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 DOI: https://doi.org/10.1093/molbev/msx281
- Jacobsen, N.H.G., Pietersen, E.W. & Pietersen, D.W. (2010) A preliminary herpetological survey of the Vilanculos Coastal Wildlife Sanctuary on the San Sebastian Peninsula, Vilankulo, Mozambique. Herpetology Notes, 3, 181–193.
- Jones, S.E., Clause, J.K., Geeraert, L., Jamie, G.A., Sumbane, E., van Berkel, T. & Jocque, W. (2017) The Njesi Plateau expedition: a biological assessment of Mt Chitagal, Mt Sanga and the Njesi Plateau in Niassa Province, Mozambique: BES Report 6.3. Biodiversity Inventory for Conservation, Glabbeek. [unknwon pagination]
- K. Lisa Yang Center for Conservation Bioacoustics (2022) Raven Pro: Interactive Sound Analysis Software. Version 1.6.5. The Cornell Lab of Ornithology, Ithaca, New York. Available from: https://ravensoundsoftware.com/ (accessed 23 December 2024)
- 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, 398–407. https://doi.org/10.1006/mpev.2001.1034 DOI: https://doi.org/10.1006/mpev.2001.1034
- Lehtinen, R.M., Richards, C.M. & Nussbaum, R.A. (2004) Origin of a complex reproductive trait: phytotelm breeding in mantelline frogs. In: Lehtinen, R.M. (Ed.), Ecology and Evolution of Phytotelm-Breeding Anurans. Museum of Zoology, University of Michigan, Ann Arbor, Michigan, pp. 45–54.
- Lobón-Rovira, J., Vaz Pinto, P.S., Becker, F., 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 (2), 289–321. https://doi.org/10.15560/18.2.289 DOI: https://doi.org/10.15560/18.2.289
- Minh, B.Q., Nguyen, M.A.T. & Von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024 DOI: https://doi.org/10.1093/molbev/mst024
- 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 DOI: https://doi.org/10.1093/molbev/msu300
- Ohler, A. & Frétey, T. (2014) Going back to Rovuma: The frog fauna of a coastal dry forest, and a checklist of the amphibians of Mozambique. Journal of East African Natural History, 103, 73–124. https://doi.org/10.2982/028.103.0203 DOI: https://doi.org/10.2982/028.103.0203
- Pietersen, D.W., Pietersen, E.W. & Haacke, W.D. (2013) First herpetological appraisal of the Parque Nacional de Banhine, Gaza Province, southern Mozambique. Annals of the Ditsong National Museum of Natural History, 3, 153–163.
- Pietersen, D.W. (2014) New distribution records of herpetofauna in Mozambique south of the Zambezi River, including additional records from Parque Nacional de Banhine. Annals of the Ditsong National Museum of Natural History, 4, 174–180.
- Portik, D.M., Mulungu, E.A., Sequeira, D. & McEntee, J.P. (2013) Herpetological surveys of the Serra Jeci and Namuli massifs, Mozambique, and an annotated checklist of the southern Afromontane archipelago. Herpetological Review, 44, 394–406.
- Poynton, J.C. (1964) The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum, 17, 1–334.
- Poynton, J.C. (1966) Amphibia of northern Mozambique. Memórias do Instituto de Investigação Cientifica de Moçambique. Série A Ciências Biológicas, 8, 13–34.
- Poynton, J.C. & Broadley, D.G. (1991/0. Amphibia Zambesiaca 5. Zoogeography. Annals of the Natal Museum, 32, 221−277.
- Rambaut, A. (2018) FigTree - Tree Figure Drawing Tool. Version 1.4.4. Available from: http://tree.bio.ed.ac.uk/ (accessed 12 January 2019)
- Reeder, J.E. (2019) Amphibians of Northern KwaZulu-Natal: A Phylogenetic Study. M.Sc. Dissertation, North-West University, Potchefstroom. [unknown pagination]
- Rödel, M.-O. & Hallermann, J. (2006) Rediscovery and identity of Phrynobatrachus monticola Fischer, 1884. African Journal of Herpetology, 55, 69–74. https://doi.org/10.1080/21564574.2006.9635540 DOI: https://doi.org/10.1080/21564574.2006.9635540
- Stalmans, M. (2003) Plant communities and landscapes of the Parque Nacional de Banhine Mozambique. African Wildlife Foundation, Johannesburg. [unknown pagination]
- Stalmans, M. & Wishart, M. (2005) Plant communities, wetlands and landscapes of the Parque Nacional de Banhine, Moçambique. Koedoe, 48 (2), 43–58. https://doi.org/10.4102/koedoe.v48i2.99 DOI: https://doi.org/10.4102/koedoe.v48i2.99
- Stecher, G., Tamura, K. & Kumar, S. (2020) Molecular Evolutionary Genetics Analysis (MEGA) for macOS. Molecular Biology & Evolution, 37, 1237–1239. https://doi.org/10.1093/molbev/msz312 DOI: https://doi.org/10.1093/molbev/msz312
- Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA 11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution, 38, 3022–3027. https://doi.org/10.1093/molbev/msab120 DOI: https://doi.org/10.1093/molbev/msab120
- Tolley, K.A., Telford, N.S., Makhubo, B.G., Power, R.J. & Alexander, G.J. (2022) Filling the gap: Noteworthy herpetological discoveries in North West Province, South Africa. Zoosystematics and Evolution, 99, 101–116. https://doi.org/10.3897/zse.99.90181 DOI: https://doi.org/10.3897/zse.99.90181
- Turner, A.A., De Villiers, A.L., Dawood, A. & Channing, A. (2004) A new species of Arthroleptella Hewitt, 1926 (Anura: Ranidae) from the Groot Winterhoek Mountains of the Western Cape Province, South Africa. African Journal of Herpetology, 53, 1–12. https://doi.org/10.1080/21564574.2004.9635493 DOI: https://doi.org/10.1080/21564574.2004.9635493
- Trifinopoulos, J., Nguyen, L.-T., Von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maxi- mum likelihood analysis. Nucleic Acids Research, 44, W1, W232–W235. https://doi.org/10.1093/nar/gkw256 DOI: https://doi.org/10.1093/nar/gkw256
- Van der Meijden, A., Vences, M., Hoegg, S. & Meyer, A. (2005) A previously unrecognized radiation of ranid frogs in Southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 37, 674–685. https://doi.org/10.1016/j.ympev.2005.05.001 DOI: https://doi.org/10.1016/j.ympev.2005.05.001
- 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, 1–12. https://doi.org/10.1080/21564574.2010.523904 DOI: https://doi.org/10.1080/21564574.2010.523904
- Van Dijk, D.E. (1966) Systematic and field keys to the families, genera and described species of southern Africam anural tadpoles. Annals of the Natal Museum, 18, 231–286.
- 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. In: Lourenco, W.R. & Goodman, S.M. (Eds.), Diversity and endemism in Madagascar. Memoires de la Société de Biogéographie, Paris, pp. 229–242.
- 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 DOI: https://doi.org/10.11646/zootaxa.3734.2.7
- Weinell, J.L., Portik, D.M. & Bauer, A.M. (2017) Noteworthy records of amphibians and reptiles from Mozambique. African Herp News, 65, 3–10.
- Willems, F. & Channing, A. (2023) The status of Tomopterna pulchra Boulenger, 1896 (Anura: Pyxicephalidae). Zootaxa, 5374 (3), 361–389. https://doi.org/10.11646/zootaxa.5374.3.3 DOI: https://doi.org/10.11646/zootaxa.5374.3.3
- Wilson, L. & Channing, A. (2019) A new sand frog from Namaqualand, South Africa (Pyxicephalidae: Tomopterna). Zootaxa, 4609 (2), 225–246. https://doi.org/10.11646/zootaxa.4609.2.2 DOI: https://doi.org/10.11646/zootaxa.4609.2.2
- 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 DOI: https://doi.org/10.1093/molbev/mst091
- 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 DOI: https://doi.org/10.11646/zootaxa.2933.1.2