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Published: 2021-06-29
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A new critically endangered slippery frog (Amphibia, Conrauidae, Conraua) from the Atewa Range, central Ghana

Museum für Naturkunde Berlin – Leibniz Institute for Evolution and Biodiversity Science, Invalidenstraße 43, 10115 Berlin, German
CSIR-Forestry Research Institute of Ghana, P. O. Box 63, Fumesua, Kumasi, Ghana. EDGE of Existence Programme, Zoological Society of London, Regent’s Park, London NW1 4RY, UK.
Université Jean Lorougnon Guédé, UFR Environnement, Laboratoire de Biodiversité et Ecologie Tropicale, Daloa, BP 150, Côte d´Ivoire
Department of Natural History, Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA
Laboratory of Ecology and Ecotoxicology, Faculty of Sciences, University of Lomé, BP 6057 Lomé, Togo
Wild Chimpanzee Foundation (WCF), Liberia Office, FDA Compound, Whein Town, Mount Barclay, Montserrado County, Liberia
Museum für Naturkunde Berlin – Leibniz Institute for Evolution and Biodiversity Science, Invalidenstraße 43, 10115 Berlin, German
Department of Biology & Burke Museum of Natural History and Culture, University of Washington, Seattle, WA, USA
Museum für Naturkunde Berlin – Leibniz Institute for Evolution and Biodiversity Science, Invalidenstraße 43, 10115 Berlin, German
Amphibia Anura biodiversity hotspot conservation integrative taxonomy Upper Guinean Forest West Africa

Abstract

Forty-nine years after the last description of a slippery frog, we describe a seventh species of the genus Conraua. The new Conraua is endemic to the Atewa Range Forest Reserve, central Ghana, and is described based on genetic, bioacoustics, and morphological evidence. Recent molecular phylogenetic and species delimitation analyses support this population as distinct from nominotypical C. derooi in eastern Ghana and adjacent Togo. The new species is sister to C. derooi, from which it differs ~4% in the DNA sequence for mitochondrial ribosomal 16S. Genetic divergences in 16S to other species of Conraua range from 4–12%. The new species is distinguished morphologically from its congeners, including C. derooi, by the combination of the following characters: medium body size, robust limbs, lateral dermal fringing along edges of fingers, cream ventral color with brown mottling, the presence of a lateral line system, indistinct tympanum, the presence of inner, outer, and middle palmar tubercles, and two subarticular tubercles on fingers III and IV. We compare the advertisement calls of the new species with the calls from C. derooi and find that they differ by duration, frequency modulation, and dominant frequency. We discuss two potential drivers of speciation between C. derooi and the new species, including river barriers and fragmentation of previously more widespread forests in West Africa. Finally, we highlight the importance of the Atewa Range Forest Reserve as a critical conservation area within the Upper Guinean biodiversity hotspot.

 

References

  1. Addo-Fordjour, P., Osei, B.A. & Kpontsu, E.A. (2015) Butterfly community assemblages in relation to human disturbance in a tropical upland forest in Ghana, and implications for conservation. Journal of Insect Biodiversity, 3(6), 1–18. https://doi.org/10.12976/jib/2015.3.6 Amiet, J.-L. (1989) Quelques aspects de la biologie des amphibiens anoures du Cameroun. Année Biologique, 28, 73–136. Amiet, J.-L. (1990) Images d’amphibiens camerounais. II. L’enfouissement et la phonation bouche ouverte chez Conraua crassipes (Buchholz & Peters, 1875). Alytes, 8, 99–104. Amiet J.-L. & Goutte S. (2017) Chants d’Amphibiens du Cameroun. J.-L. Amiet & Editions Petit Génie. Saint-Nazaire, France, 280 pp + 4CDs. AmphibiaWeb (2020) Conraua goliath: Goliath frog, University of California, Berkeley, CA, USA. Available from: http://amphibiaweb.org/species/4691 (accessed 13 March 2021). Barbour, T. & Loveridge, A. (1927) Some undescribed frogs and a new gecko from Liberia. Proceedings of the New England Zoölogical Club, 10, 13–18. Barbour, T. & Loveridge, A. (1930) Reptiles and amphibians from Liberia. In: Strong, R.P. (Ed.), The African republic of Liberia and the Belgian Congo, based on the observations made and material collected during the Harvard African Expedition 1926–1927, Vol. 2. Greenwood Press, New York, pp. 769–786. Barej, M.F., Rödel, M.-O., Gonwouo, L.N., Pauwels, O.S., Böhme, W., & Schmitz, A. (2010) Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae). Zootaxa, 2340, 1–49. https://doi.org/10.11646/zootaxa.4072.4.6 Blackburn, D.C. & Wake, D.B. (2011) Class Amphibia Gray, 1825. In: Zhang, Z.-Q. (Ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148, 39–55. https://doi.org/10.11646/zootaxa.3148.1.8 Blackburn, D.C., Nielsen, S.V., Barej, M.F., Doumbia, J., Hirschfeld, M., Kouamé, N.G., Dwight, L., 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 Burger, M., Branch, W.R. & Channing, A. (2004) Amphibians and reptiles of Monts Doudou, Gabon: species turnover along an elevational gradient. Memoirs of the California Academy of Sciences, 28, 145–186. Center for Conservation Bioacoustics. (2019) Raven Pro: Interactive Sound Analysis Software (Version 1.6.1) [Computer software]. Ithaca, NY: The Cornell Lab of Ornithology. Available from http://ravensoundsoftware.com/. Channing, A., Rödel, M.-O. & Channing J. (2012) Tadpoles of Africa. The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, Germany, 402 pp. Channing, A. & Rödel, M.-O. (2019) Field guide to the frogs and other amphibians of Africa. Vol. 1. Struik Nature, Cape Town, South Africa, 408 pp. Couvreur, T.L., Dauby, G., Blach-Overgaard, A., Deblauwe, V., Dessein, S., Droissart, V., Hardy, O., Harris, D., Janssens, S., Ley, A.C., Mackinder, B.A., Sonké, B., Sosef, 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 Duminil, J., Brown, R.P., Ewédjè, E.-E.B.K., Mardulyn, P., Doucet, J.-L. & Hardy, O.J. (2013) Large-scale pattern of genetic differentiation within African rainforest trees: insights on the roles of ecological gradients and past climate changes on the evolution of Erythrophleum spp (Fabaceae). BMC Evolutionary Biology, 13, 195. https://doi.org/10.1186/1471-2148-13-195 Escher, K. (1925) Das Verhalten der Seitenorgane der Wirbeltiere und ihrer Nerven beim Übergang zum Landleben. Acta Zoologica, 6, 307–414. Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N.J. (2007) Underestimation of species richness in neotropical frogs revealed by mtDNA analyses. PLoS One, (2) 10, e1109. https://doi.org/10.1371/journal.pone.0001109 Frétey, T. & Blanc, C.P. (2000) Liste des amphibiens d’Afrique centrale. Cameroun, Congo, Gabon, Guinée-Equatoriale, République Centrafricaine, République Démocratique du Congo, São Tomé et Príncipe. Les dossiers de l’ADIE, Série Biodiversité, 2, 1–39. Fritzsch, B., Drewes, R.C. & Ruibal, R. (1987) The retention of the lateral-line nucleus in adult anurans. Copeia, 1987, 127–135. Frost, D.R. (2021) Amphibian Species of the World: An Online Reference. Version 6.1 Electronic Database accessible at https://amphibiansoftheworld.amnh.org/index.php. American Museum of Natural History, New York, USA (accessed 7 January 2021). 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.C., 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. Fuchs, J. & Bowie, R.C. (2015) Concordant genetic structure in two species of woodpecker distributed across the primary West African biogeographic barriers. Molecular Phylogenetics and Evolution, 88, 64–74. https://doi.org/10.1016/j.ympev.2015.03.011 Gartshore, M. (1984) The status of the montane herpetofauna of the Cameroon highlands. In: Stuart, S.N. (Ed.), Conservation of Cameroon Montane Forests. International Council for Bird Preservation, London, pp. 204–240. Ghartey-Tagoe, F., Ekumah, B., Pappoe, A.N.M. & Akotoye, H.K. (2021) Effects of anthropogenic activities on land-use dynamics in an upland tropical evergreen forest in Ghana. African Geographical Review, 40, 163–179. https://doi.org/10.1080/19376812.2020.1785318 Guibé, J. & Lamotte, M. (1958) La réserve naturelle intégrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius). Mémoires de l’Institut français d’Afrique Noire, 53, 241–273. Hassanin, A., Khouider, S., Gembu, G.C., Goodman, S.M., Kadjo, B., Nesi, N., Pourrut X., Nakouné, E. & Bonillo C. (2015) The comparative phylogeography of fruit bats of the tribe Scotonycterini (Chiroptera, Pteropodidae) reveals cryptic species diversity related to African Pleistocene forest refugia. Comptes Rendus Biologies, 338, 197–211. https://doi.org/10.1016/j.crvi.2014.12.003 Herrmann, H.-W., Böhme, W., Herrmann, P.A., Plath, M., Schmitz, A. & Solbach, M. (2005) African biodiversity hotspots: the amphibians of Mt. Nlonako, Cameroon. Salamandra, 41, 61–81. Herrmann, H.-W. & Edwards, T. (2006) Conraua goliath (Goliath Frog) skittering locomotion. Herpetological Review, 37, 202–203. Hillers, A., Loua, N.-S. & Rödel, M.-O. (2008) A preliminary assessment of the amphibians of the Fouta Djallon, Guinea, West Africa. Salamandra, 44, 113–122. Hillers, A., Boateng, C.O., Segniagbeto, G.H., Agyei, A.C. & Rödel, M.-O. (2009) Assessment of the amphibians in the forests of southern Ghana and western Togo. Zoosystematics and Evolution, 85, 127–141. https://doi.org/10.1002/zoos.200800019 Hodgetts, N.G., Essilfie, M.K., Adu-Gyamfi, A., Akom, E., Kumadoh, J. & Opoku, J. (2016) Bryophytes of Atewa Forest, Eastern Region, Ghana. Journal of Bryology, 38, 211–222. https://doi.org/10.1080/03736687.2016.1145525 Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, 25, 29–38. Hulselmans, J.L.J. (1972) Contribution à l’herpétologie de la République du Togo, 4. Description de Conraua derooi, n. sp. (Amphibia). Revue de Zoologie et de Botanique Africaines, 84, 153–159. Hutchison, V.H. (1998) The Goliath Frog (Conraua goliath): physiological ecology of the largest anuan. Extended abstracts of the 1998 International Symposium on Animal Adaptation, Institute of Zoology, Academia Sinica, Taipei, Taiwan, Republic of China. 5 pp. IUCN. (2012) IUCN Red List Categories and Criteria: Version 3.1. Second edition. Gland, Switzerland and Cambridge, UK: IUCN. iv + 32pp Jacquet, F., Nicolas, V., Colyn, M., Kadjo, B., Hutterer, R., Decher, J., Akpatou, B., Cruaud, C. & Denys, C. (2014) Forest refugia and riverine barriers promote diversification in the West African pygmy shrew (Crocidura obscurior complex, Soricomorpha). Zoologica Scripta, 43, 131–148. Jongsma, G.F., Barej, M.F., Barratt, C.D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M., Leaché, A.D., Penner, J., Portik, D.M., Zassi-Boulou, A.-G., Rödel, M.-O. & Blackburn, D.C. (2018) Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa. Molecular Phylogenetics and Evolution, 120, 274–285. 10.1016/j.ympev.2017.12.006 Knoepffler, L.P. (1985) Le comportement fouisseur de Conraua crassipes (Amphibien anoure) et son mode de chasse. Biologia Gabonica, 1, 239–245. Köhler, J., Jansen, M., Rodríguez, A., Kok, P., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F.B., Rödel, M.-O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251, 1–124. https://doi.org/10.11646/zootaxa.4251.1.1 Kouamé, N.G., Boateng, C.O. & Rödel, M.-O. (2007) A rapid survey of the amphibians from the Atewa Range Forest Reserve, Eastern Region, Ghana. In: McCullough, J., Alonso, L.E., Naskrecki, P., Wright, H.E. & Osei-Owusu, Y. (Eds.), A rapid biological assessment of the Atewa Range Forest Reserve, Eastern Ghana. RAP Bulletin of Biological Assessment 47, Conservation International, Washington D.C, pp 76–83. Kusimi, J.M. (2015) Characterizing land disturbance in Atewa range forest reserve and buffer zone. Land Use Policy, 49, 471–482. https://doi.org/10.1016/j.landusepol.2015.08.020 Lamotte, M. (1971) Le Massif des Monts Loma (Sierra Leone), Fasciule I, XIX. Amphibiens. Mémoires de l’Institut fondamental d’Afrique Noire, 86, 397–407. Lamotte, M. & Perret, J.-L. (1968) Révision du genre Conraua Nieden. Bulletin de l’Institut fondamental d’Afrique noire, Série A, 30, 1603–1644. Lamotte, M., Perret J.-L. & Dzieduszycka, S. (1959) Contribution à l’étude des batraciens de l’Ouest Africain IX. – Les formes larvaires de Petropedetes palmipes, Conraua goliath et Acanthixalus spinosus. Bulletin de l’Institut français d’Afrique noire, Série A, 21: 762–776. Largen, M.J. & Spawls, S. (2010) The amphibians and reptiles of Ethiopia and Eritrea. Edition Chimaira, Frankfurt am Main, 693 pp. Lawson, D.P. (1993) The reptiles and amphibians of the Korup National Park project, Cameroon. Herpetological Natural History, 1, 27–90. Leaché, A.D., Oaks, J.R., Ofori-Boateng, C. & Fujita, M.K. (2020) Comparative phylogeography of West African amphibians and reptiles. Evolution, 74, 716–724. https://doi.org/10.1111/evo.13941 Lindsell, J., Agyei, R., Bosu, R., Decher, J., Hawthorne, W., Marshall, C., Ofori-Boateng, C. & Rödel, M.-O. (2019) The Biodiversity of Atewa Forest Research Report. A Rocha Ghana, Accra, Ghana, 88 pp. Lötters, S., Gossmann, V., Obame, F. & Böhme, W. (2001) Zur Herpetofauna Gabuns, Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche. herpetofauna, 23 (133), 19–34. McCullough, J., Alonso, L.E., Naskrecki, P., Wright, H.E. & Osei-Owusu, Y. (2007) A rapid biological assessment of the Atewa Range Forest Reserve, Eastern Ghana. RAP Bulletin of Biological Assessment, 47, Conservation International, Washington D.C., 193 pp. Mittermeier, R.A., Gil, P.R., Hoffmann, M., Pilgrim, J., Brooks, T., Mittermeier, C.G., Lamoreux, J. & da Fonseca, G.A.B. (Eds.) (2004) Hotspots revisited. Earth’s biologically richest and most endangered terrestrial ecoregions. CEMEX/ Agrupación Sierra Madre, Mexico City, 309 pp. Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858. Naskrecki, P. (2008a) A new ricinuleid of the genus Ricinoides Ewing (Arachnida, Ricinulei) from Ghana. Zootaxa, 1698, 57–64. Naskrecki, P. (2008b) Sylvan katydids (Orthoptera: Tettigoniidae: Pseudophyllinae) of the Guinean Forests of West Africa hotspot: an overview and descriptions of new species. Zootaxa, 1712, 1–41. Naskrecki, P. (2008c) New species of arboreal predatory katydids from West Africa (Orthoptera: Tettigoniidae: Meconematinae). Zootaxa, 1732, 1–28. Neff, N.A. & Marcus, L.F. (1980) A survey of multivariate methods for systematics. New York: American Museum of Natural History. New York, USA, 243 pp. Nicolas, V., Akpatou, B., Wendelen, W., Kerbis Peterhans, J., Olayemi, A., Decher, J., Missou A.-D., Denys, C., Barriere, P., Cruaud, C. & Colyn, M. (2010) Molecular and morphometric variation in two sibling species of the genus Praomys (Rodentia: Muridae): implications for biogeography. Zoological Journal of the Linnean Society, 160, 397-419. Ofori-Boateng, C., Leaché, A.D., Obeng-Kankam, B., Kouamé, N.G., Hillers, A. & Rödel, M.-O. (2018) A new species of Puddle Frog, genus Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from Ghana. Zootaxa, 4374, 565–578. https://doi.org/10.11646/zootaxa.4374.4.6 OpenStreetMap. (2020) Available from: https://www.openstreetmap.org (accessed 18 December 2020). Paluh, D.J., Stanley, E.L. & Blackburn, D.C. (2020) Evolution of hyperossification expands skull diversity in frogs. Proceedings of the National Academy of Sciences of the United States of America, 117, 8554–8562. https://doi.org/10.1073/pnas.2000872117 Penner, J., Wegmann, M., Hillers, A., Schmidt, M. & Rödel, M.-O. (2011) A hotspot revisited – a biogeographical analysis of West African amphibians. Diversity and Distribution, 17, 1077–1088. https://doi.org/10.1111/j.1472-4642.2011.00801.x Perret, J.L. (1957) Observations sur Rana goliath Blgr. Bulletin de la Société Neuchâteloise des Sciences naturelles, 80, 195–202. Perret, J.-L. (1960) Études herpétologiques africaines II. Bulletin de la Société Neucthâteloise des Scienes Naturelles, 83, 93–100. Perret, J.-L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbücher (Abteilung für Systematik, Ökologie und Geographie der Tiere), 8, 289–464. Plath, M., Solbach, M. & Herrmann, H.-W. (2004) Anuran habitat selection and temporal partitioning in a montane and submontane rainforest on southwestern Cameroon – first results. Salamandra, 40, 239–260. Portik, D.M., Bell, R.C., Blackburn, D.C., Bauer, A.M., Barratt, C.D., Branch, W.R., Burger, M., Channing, A., Colston, T.J., Conradie, W., Dehling, J.M., Drewes, R.C., Ernst, R., Greenbaum, E., Gvoždík, V., Harvey, J., Hillers, A., Hirschfeld, M., Jongsma, G.F.M., Kielgast, J., Kouete, M.T., Lawson, L.P., Leaché, A.D., Loader. S.P., Lötters, S., van der Meijden, A., Menegon, M., Müller, S., Nagy, Z.T., Ofori-Boateng, C., Ohler, A., Papenfuss, T.J., Rößler, D., Sinsch, U., Rödel, M.-O., Veith, M., Vindum, J., Zassi-Boulou, A.-G. & McGuire, J.A. (2019) Sexual dichromatism drives diversification within a major radiation of African amphibians. Systematic Biology, 68, 859–875. https://doi.org/10.1093/sysbio/syz023 R Core Team. (2018) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/. Rödel, M.-O. (2003) The amphibians of Mont Sangbé National Park, Ivory Coast. Salamandra, 39, 91–110. Rödel, M.-O. & Bangoura, M.A. (2004) A conservation assessment of amphibians in the Forêt Classée du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with the description of a new Amnirana species (Amphibia Anura Ranidae). Tropical Zoology, 17, 201–232. https://doi.org/10.1080/03946975.2004.10531206 Rödel, M.-O., Bangoura, M.A. & Böhme, W. (2004) The amphibians of south-eastern Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa, 17, 99–118. Rödel, M.-O., Barej, M.F., Hillers, A., Leaché, A.D., Kouamé, N.G., Ofori-Boateng, C., Assemian, N.E., Tohé, B., Penner, J., Hirschfeld, M., Doumbia, J., Gonwouo, L.N., Nopper, J., Brede, C., Diaz, R., Fujita, M.K., Gil, M., Segniagbeto, G.H., Ernst, R. & Sandberger, L. (2012) The genus Astylosternus in the Upper Guinea rainforests, West Africa, with the description of a new species (Amphibia: Anura: Arthroleptidae). Zootaxa, 3245, 1–29. https://doi.org/10.11646/zootaxa.3245.1.1 Rödel, M.-O. & Branch, W.R. (2002) Herpetological survey of the Haute Dodo and Cavally forests, western Ivory Coast, Part I: Amphibians. Salamandra, 38, 245–268. Rödel, M.-O., Kouamé, N.G., Doumbia, J. & Sandberger, L. (2011) A new beautiful squeaker frog (Arthroleptidae: Arthroleptis) from West Africa. Zootaxa, 3011, 16–26. https://doi.org/10.11646/zootaxa.3011.1.2 Rödel, M.-O., Kosuch, J., Veith, M. & Ernst, R. (2003) First record of the genus Acanthixalus Laurent, 1944 from the Upper Guinean rain forest, West Africa, with the description of a new species. Journal of Herpetology, 37, 43–52. Sabater-Pi, J. (1985) Contribution to the biology of the giant frog (Conraua goliath, Boulenger). Amphibia-Reptilia, 6, 143–153. Schäfer, M., Tsekané, S.J., Tchassem, F.A.M., Drakulić, S., Kameni, M., Gonwouo, N.L. & Rödel, M.-O. (2019) Goliath frogs build nests for spawning–the reason for their gigantism? Journal of Natural History, 53, 1263–1276. https://doi.org/10.1080/00222933.2019.1642528 Schiøtz, A. (1964) A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 127, 1–17. Segniagbeto, G.H., Assou, D., Dendi, D., Rödel, M.-O., Ohler, A., Dubois, A. & Luiselli, L. (2017) The distribution and local density of the critically endangered frog, Conraua derooi Hulselmans, 1972 in Togo, West Africa. The Herpetological Bulletin, 141, 23–27. Segniagbeto, G.H., Bowessidjaou, J.E., Dubois, A. & Ohler, A. (2007) Les Amphibiens du Togo: état actuel des connaissances. Alytes, 24, 72–90. Seidu, I., Danquah, E., Ayine Nsor, C., Amaning Kwarteng, D. & Lancaster, L.T. (2017) Odonata community structure and patterns of land use in the Atewa Range Forest Reserve, Eastern Region (Ghana). International Journal of Odonatology, 20, 173–189. https://doi.org/10.1080/13887890.2017.1369179 Shelton, P.M. (1970) The lateral line system at metamorphosis in Xenopus laevis (Daudin). Development, 24, 511–524. Siaw, D.E.K.A. & Dabo, J. (2007) A rapid botanical survey of the Atewa Range Forest Reserve, Ghana. In: McCullough, J., Alonso, L.E., Naskrecki, P., Wright, H.E. & Osei-Owusu, Y. (Eds.), A rapid biological assessment of the Atewa Range Forest Reserve, Eastern Ghana. RAP Bulletin of Biological Assessment 47, Conservation International, Washington D.C, pp. 47–49. U.S. Geological Survey. (2020) Shuttle radar topography mission (SRTM). Available from: https://www.usgs.gov (accessed 18 December 2020). Venables, W.N. & Ripley, B.D. (2002) Modern Applied Statistics with S, Fourth edition. Springer, New York. ISBN 0-387-95457-0. http://www.stats.ox.ac.uk/pub/MASS4/. Vieites, D.R., Wollenberg, K.C., Andreone, F., Köhler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar’s biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106, 8267–8272. https://doi.org/10.1073/pnas.0810821106 Wallace, A.R. (1854) On the monkeys of the Amazon. Annals and Magazine of Natural History, 14(84), 451–454. https://doi.org/10.1080/037454809494374 Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. https://ggplot2.tidyverse.org. Womack, M.C. & Bell, R.C. (2020) Two-hundred million years of anuran body-size evolution in relation to geography, ecology and life history. Journal of Evolutionary Biology, 33, 1417–1432. https://doi.org/10.1111/jeb.13679