Abstract
We show that the Wotjulum frog, Litoria watjulumensis (Copland, 1957), comprises two deeply divergent mitochondrial DNA lineages that are also reciprocally monophyletic for a nuclear gene locus and have discrete distributions. The taxa are differentiated in multivariate analysis of shape but show no appreciable differences in colour and pattern. The two taxa differ substantially in the degree of female biased sexual size dimorphism, with the western taxon showing considerably more pronounced dimorphism. We subsequently resurrect Litoria (Hyla) spaldingi (Hosmer, 1964) for populations from east of the Daly River system in the Northern Territory through to western Queensland and restrict L. watjulumensis to populations from the Kimberley region of north-western Australia and the Victoria River system of the western Northern Territory. The complex advertisement call of L. spaldingi is described for the first time.
References
Anstis, M., Price, L.C., Roberts, J.D., Catalano, S., Doughty,P., Hines, H.B. & Donnellan, S.C. (2016) Revision of the Australian water holding frog (Cyclorana platycephala, Anura: Hylidae), with a description of a new species and subspecies. Zootaxa, 4126 (4), 451–479.
https://doi.org/10.11646/zootaxa.4126.4.1
Copland, S.J. (1957) Australian tree frogs of the genus Hyla. Proceedings of the Linnean Society of New South Wales, 82, 9–108.
de Queiroz, K. (1998) The general lineage concept of species, species criteria and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms, Species and Speciation. Oxford University Press, Oxford, pp. 57–75.
Doughty, P. (2011) An emerging frog diversity hotspot in the northwest Kimberley of Western Australia: another new frog species from the high rainfall zone. Records of the Western Australian Museum, 26, 209–216.
https://doi.org/10.18195/issn.0312-3162.26(2).2011.209-216
Doughty, P. & Anstis, M. (2007) A new species of rock dwelling hylid frog (Anura: Hylidae) from the eastern Kimberley region of Western Australia. Records of the Western Australian Museum, 23, 241–257.
https://doi.org/10.18195/issn.0312-3162.23(3).2007.241-257
Doughty, P., Palmer, R., Cowan, M. & Pearson D.J. (2012) Biogeographic patterns of frogs of the Kimberley islands, Western Australia. Records of the Western Australian Museum, Supplement 81, 109–124.
https://doi.org/10.18195/issn.0313-122x.81.2012.109-124
Doughty, P., Bourke, G., Tedeschi, L.G., Pratt, R.C., Oliver, P.M., Palmer, R.A. & Moritz, C. (2018) Species delimitation in the Gehyra nana (Squamata: Gekkonidae) complex: cryptic and divergent morphological evolution in the Australian Monsoonal Tropics, with the description of four new species. Zootaxa, 4403 (2), 201–244,
https://doi.org/10.11646/zootaxa.4403.2.1
Edgar, RC. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.
https://doi.org/10.1093/nar/gkh340
Francis, J. (2013) Ecological studies of anuran larvae in temporary freshwaters of the Australian wet-dry tropics. PhD Thesis, Scholl of Animal Biology, University of Western Australia, Perth, xiii + 236 pp.
Fujita, M.K., McGuire, J.A., Donnellan, S.C. & Moritz C. (2010) Diversification and persistence in arid environments: Australian biogeography of the Bynoe’s gecko (Heteronotia binoei; Gekkonidae). Evolution, 64, 2293–2314.
https://doi.org/10.1111/j.1558-5646.2010.00993.x
Horner, P. (2007) Systematics of the snake-eyed skinks, Cryptoblepharus Wiegmann (Reptilia: Squamata: Scincidae)—an Australian-based review. Records of the Museum of the Northern Territory, Supplement 3, 2–98.
Hoskin, C., Grigg, G., Stewart, D. & McDonald, S. (2015) Frogs of Australia—a complete electronic field guide to Australian frogs. Ug Media, Australia. Available from: ugmedia.com.au/field-guides/frogs-of-australia/ (accessed 19 January 2021)
Hosmer, W. (1964) A new frog of the genus Hyla from Northern Territory, Australia. American Museum Novitates, 2182, 1–7.
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics Applications Note, 28, 1647–164.
https://doi.org/10.1093/bioinformatics/bts199
Köhler, J., Jansen, M., Rodríguez, A., Kok, P.J.R., 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), 1–124.
https://doi.org/10.11646/zootaxa.4251.1.1
Kutt, A. & Woinarski, J.C.Z. (2007) The effects of grazing and fire on vegetation and the vertebrate assemblage in a tropical savanna woodland in north-eastern Australia. Journal of Tropical Ecology, 23, 95–106.
https://doi.org/10.1017/S0266467406003579
Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.
https://doi.org/10.1093/molbev/mss020
Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93.
https://doi.org/10.1006/jtbi.2000.2043
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 November 2010, New Orleans, Louisiana, pp. 1–8.
https://doi.org/10.1109/GCE.2010.5676129
Moritz, C., Fujita, M.K., Rosauer, D., Agudo, R., Bourke, G., Doughty, P., Palmer, R., Pepper, M., Potter, S., Pratt, R., Scott, M., Tonione, M. & Donnellan, S. (2016) Multilocus phylogeography reveals nested endemism in a gecko across the monsoonal tropics of Australia. Molecular Ecology, 25, 1354–1366.
https://doi.org/10.1111/mec.13511
Oliver, P.M., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386.
https://doi.org/10.1186/1471-2148-10-386
Oliver, P.M., Prasetya, A.M., Tedeschi, L.G., Fenker, J., Ellis, R.J., Doughty, P. & Moritz, C. (2020) Crypsis and convergence: integrative taxonomic revision of the Gehyra australis group (Squamata: Gekkonidae) from northern Australia. PeerJ, 8, e7971
https://doi.org/10.7717/peerj.7971
Rabosky, D.L., Hutchinson, M.N., Donnellan, S.C., Talaba, A.L. & Lovette, I.J. (2014) Phylogenetic disassembly of species boundaries in a widespread group of Australian skinks (Scincidae: Ctenotus). Molecular Phylogenetics and Evolution, 77, 71–82.
https://doi.org/10.1016/j.ympev.2014.03.026
Rambaut, A. & Drummond, A.J. (2007) Tracer. Version 1.4. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 19 January 2021)
Renner, S.S. (2016) return to Linnaeus’s focus on diagnosis, not description: the use of DNA characters in the formal naming of species. Systematic Biology, 65, 1085–1095.
https://doi.org/10.1093/sysbio/syw032
Reynolds, S.J., Dostine, P.L. & Griffiths A.D. (2010) Trial of a survey methodology to monitor frog assemblages in the Darwin region. Report number 26/2010D. Northern Territory Government, Northern Territory of Australia. [unknown pagination, ISBN 978-1-921519-86-4]
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
Rowley, J.J.L., Callaghan, C.T., Cutajar, T., Portway, C., Potter K., Mahony, S, Trembath, D.F., Flemons, P. & Woods, A. (2019) FrogID: Citizen scientists provide validated biodiversity data on frogs of Australia. Herpetological Conservation and Biology, 14, 155-170.
Stamatakis, A. (2014) RAxML Version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731–2739.
https://doi.org/10.1093/molbev/msr121
Thorpe, R.S. (1976) Biometric analysis of geographic variation and racial affinities. Biological Reviews, 51, 407–452.
https://doi.org/10.1111/j.1469-185X.1976.tb01063.x
Tyler, M.J. (1968) A taxonomic study of hylid frogs of the Hyla lesueuri complex occurring in northwestern Australia. Records of the South Australian Museum, 15, 711–727.
Tyler, M.J., Crook, G.A. & Davies, M. (1983) Reproductive biology of the frogs of the Magela Creek System, Northern Territory. Records of the South Australian Museum, 18, 415–440.
Tyler, M.J. & Davies, M. (1978) Species-groups within the Australopapuan hylid frog genus Litoria Tschudi. Australian Journal of Zoology, Supplemental Series, Number 63, 1–47.
https://doi.org/10.1071/AJZS063
Tyler, M.J., Davies, M. & Martin, A.A. (1978) A new species of hylid frog from the Northern Territory. Transactions of the Royal Society of South Australia, 102, 151–157.
Tyler, M.J. & Doughty, P. (2009) Frogs of Western Australia. 4th Edition. Western Australian Museum Press, Perth, 160 pp.
Vences, M., Thomas, M., Van der Meijden, A.Y., Chiari, Y. & Vieites, D.R. (2005) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology, 2, 5.
https://doi.org/10.1186/1742-9994-2-5
Woinarski, J.C.Z. & Gambold, N. (1992) Gradient analysis of a tropical herpetofauna: distribution patterns of terrestrial reptiles and amphibians in Stage III of Kakadu National Park, Australia. Wildlife Research, 19, 105.
https://doi.org/10.1071/WR9920105
Woinarski, J.C.Z., Armstrong, M., Price, O., McCartney, J., Griffiths, A.D. & Fisher, A. (2004) The terrestrial vertebrate fauna of Litchfield National Park, Northern Territory: monitoring over a 6-year period and response to fire history. Wildlife Research, 31, 587–596.
https://doi.org/10.1071/WR03077