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Type: Article
Published: 2023-01-11
Page range: 1-43
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Molecular systematic analysis demonstrates that the threatened southern bell frog, Litoria raniformis (Anura: Pelodryadidae) of eastern Australia, comprises two sub-species

1Department of Zoology, Hungarian Natural History Museum H-1088 Budapest, Baross u. 13, Hungary
2School of Agriculture, Environment and Veterinary Sciences, Charles Sturt University, PO Box 789, Albury, 2640, Australia.
3School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia.
4New South Wales Department of Planning, Industry and Environment, Albury, 2640, Australia.
5South Australian Museum, North Terrace, Adelaide, 5001, Australia.
3School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia.
6School of Environmental and Life Sciences, University of Newcastle, University Drive, Callaghan NSW 2308, Australia
3School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia.
Amphibia mtDNA nuclear DNA Litoria aurea Group Litoria r. raniformis L. r. major divergent evolutionary lineages anthropogenic dispersal

Abstract

In south-eastern Australia, the pelodryadid Litoria aurea Group (sensu Tyler & Davies 1978) comprises three species: Litoria aurea (Lesson, 1829), Litoria raniformis (Keferstein, 1867), and Litoria castanea (Steindachner, 1867). All three species have been subject to declines over recent decades and taxonomic uncertainty persists among populations on the tablelands in New South Wales. We address the systematics of the Group by analysing mitochondrial and nuclear DNA sequences to assess divergence in the Litoria raniformis from across its current range in New South Wales (NSW), Victoria, South Australia (SA) and Tasmania. We also included samples of Litoria castanea from a recently rediscovered population in the southern tablelands of NSW. Our phylogenetic and population genetic analyses show that Litoria raniformis comprises northern and southern lineages, showing deep mitochondrial DNA sequence divergence (7% net average sequence divergence) and can be diagnosed by fixed allelic differences at more than 4,000 SNP loci. Samples of the northern lineage were collected from the Murray-Darling Basin while those of the southern lineage were collected from south-eastern South Australia, southern and south-eastern Victoria and Tasmania. Analysis of the morphology and bioacoustics did not unequivocally delineate the two lineages. The presence of a hybrid backcross individual in western Victoria at the northern margin of the southern lineage, leads us to assign sub-species status to the two lineages, L. r. raniformis for the northern lineage and L. r. major for the southern lineage. Our data do not unequivocally resolve the taxonomic status of L. castanea which will require molecular genetic analyses of museum vouchers from those parts of the range where L. castanea and L. raniformis are no longer extant. Our data also suggest that human mediated movement of frogs may have occurred over the past 50 years. Our genotyping of vouchers collected in the 1970s from the Mount Lofty Ranges in South Australia detected mitochondrial haplotypes of both sub-species and SNP analysis showed that a single Tasmanian specimen was a backcross with L. r. raniformis ancestry. Movement of L. r. raniformis into Tasmania and both sub-species into the Mount Lofty Ranges are not likely due to passive movements of animals through agricultural commerce, but due to the attractiveness of the species as pets and subsequent escapes or releases, potentially of the larval life stage.

 

References

  1. AmphibiaWeb (2021) AmphibiaWeb. University of California, Berkeley, California. Available from: http://amphibiaweb.org (accessed 21 October 2021)
    Anderson, E.C. & Thompson, E.A. (2002) A model-based method for identifying species hybrids using multilocus genetic data. Genetics, 160, 1217−1229. https://doi.org/10.1093/genetics/160.3.1217
    Anstis, M. (2017) Tadpoles and frogs of Australia. New Holland Publishers Pty Limited, Chatswood, 848 pp.
    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
    Armstrong, D.M., Croft S.J. & Foulkes, J.N. (2003) A Biological Survey of the Southern Mount Lofty Ranges, South Australia, 2000–2001. Department for Environment and Heritage, South Australia. [unknown pagination]
    Barker, J. & Grigg, G. (1977) A Field Guide to Australian Frogs. Rigby, Adelaide, 229 pp.
    Beeman, K. (1998) Digital signal analysis, editing, and synthesis. In: Animal Acoustic Communication. Springer, Berlin and Heidelberg, pp. 59–103. https://doi.org/10.1007/978-3-642-76220-8_3
    Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L, Meier, R., Winker, K., Ingram, K.K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology and Evolution, 22, 148–155. https://doi.org/10.1016/j.tree.2006.11.004
    Bielby, J., Cooper, N., Cunningham, A.A., Garner, T.W.J. & Purvis, A. (2008) Predicting susceptibility to future declines in the world’s frogs. Conservation Letters, 1, 82–90. https://doi.org/10.1111/j.1755-263X.2008.00015.x
    Blaustein, A.R., Walls, S.C., Bancroft, B.A., Lawler, J.J., Searle, C.L. & Gervasi, S.S. (2010) Direct and indirect effects of climate change on amphibian populations. Diversity, 2, 281. https://doi.org/10.3390/d2020281
    Burns, E.L., Eldridge, M.D.B. & Houlden, B.A. (2004) Microsatellite variation and population structure in a declining Australian hylid Litoria aurea. Molecular Ecology, 13, 1745–1757. https://doi.org/10.1111/j.1365-294X.2004.02190.x
    Burns, E.L., Eldridge, M.D.B., Crayn, D.M. & Houlden, B.A. (2007) Low phylogeographic structure in a widespread endangered Australian frog Litoria aurea (Anura: Hylidae). Conservation Genetics, 8, 17–32. https://doi.org/10.1007/s10592-006-9143-8
    Carey, C. & Alexander, M.A. (2003) Climate change and amphibian declines: is there a link? Diversity and Distributions, 9, 111–121. https://doi.org/10.1046/j.1472-4642.2003.00011.x
    Catullo, R.A., Doughty, P., Roberts, J.D. & Keogh, J.S. (2011) Multi-locus phylogeny and taxonomic revision of Uperoleia toadlets (Anura: Myobatrachidae) from the western arid zone of Australia, with a description of a new species. Zootaxa, 2902 (1), 1–43. https://doi.org/10.11646/zootaxa.2902.1.1
    Catullo, R.A., Doughty, P. & Keogh, J.S. (2014) A new frog species (Myobatrachidae: Uperoleia) from the Northern Deserts region of Australia, with a redescription of U. trachyderma. Zootaxa, 3753 (3), 251–262. https://doi.org/10.11646/zootaxa.3753.3.4
    Catullo, R.A. & Keogh, J.S. (2021) Seen only once: an evolutionarily distinct species of Toadlet (Uperoleia: Myobatrachidae) from the Wessel Islands of northern Australia. Zootaxa, 5057 (1), 52–68 https://doi.org/10.11646/zootaxa.5057.1.3
    Chan, K.O. & Grismer, L.L. (2021) Correcting for body size variation in morphometric analysis. bioRxiv, 2021.05.17.444580. [published online] https://doi.org/10.1101/2021.05.17.444580
    Chapin, F.S., Zavaleta, E.S., Eviner, V.T., Naylor, R.L., Vitousek, P.M., Reynolds, H.L., Hooper, D.U., Lavorel, S., Sala, O.E., Hobbie, S.E., Mack, M.C. & Diaz, S. (2000) Consequences of changing biodiversity. Nature, 405, 234–242. https://doi.org/10.1038/35012241
    Chifman, J. & Kubatko, L. (2014) Quartet inference from SNP data under the coalescent model. Bioinformatics, 30, 3317–3324. https://doi.org/10.1093/bioinformatics/btu530
    Clemann, N. & Gillespie, G.R. (2012) National recovery plan for the southern bell frog Litoria raniformis. Department of Sustainability and Environment, Melbourne.
    Clulow, S. & Swan, M. (2018) A Complete Guide to Frogs of Australia. Australian Geographic, Sydney, 336 pp.
    Copland, S.J. (1957) Australian tree frogs of the genus Hyla. Proceedongs of the Linnean Society of N.S.W., 82, 9–108.
    Cogger, H.G., Cameron E.E. & Cogger, H.M. (1983) Zoological Catalogue of Australia. Australian Government Publishing Service, Canberra. [unknown pagination]
    Courtice, G.P. & Grigg, G.C. (1975) A taxonomic revision of the Litoria aurea complex (Anura: Hylidae) in southeastern Australia. Australian Zoologist, 18, 149–163.
    Department of the Environment, Water, Heritage and the Arts (DEWHA). (2009) Litoria raniformis in Species Profile and Threats Database, Department of the Environment, Water, Heritage and the Arts, Canberra. Available from: http://www.environment.gov.au/sprat (accessed 3 December 2009)
    de Queiroz, K. (2020) An updated concept of subspecies resolves a dispute about the taxonomy of incompletely separated lineages. Herpetological Review, 51, 459–461.
    Donnellan, S.C., McGuigan, K., Knowles, R., Mahony, M. & Moritz, C. (1999) Genetic evidence for species boundaries in frogs of the Litoria citropa species group (Anura: Hylidae). Australian Journal of Zoology, 47, 275–293. https://doi.org/10.1071/ZO99013
    Donnellan, S.C. & Mahony, M.J. (2004) Allozyme, chromosomal and morphological variability in the Litoria lesueuri species group (Anura: Hylidae), including a description of a new species. Australian Journal of Zoology, 52, 1–28. https://doi.org/10.1071/ZO02069
    Donnellan, S.C., Mahony, M.J. & Bertozzi, T. (2012a) A new species of Pseudophryne (Anura: Myobatrachidae) from the central Australian ranges. Zootaxa, 3476, 69–85. https://doi.org/10.11646/zootaxa.3476.1.4
    Donnellan, S.C., Anstis, M., Price, A.H. & Wheaton, L. (2012b) A new species of Crinia (Anura: Myobatrachidae) from the Flinders Ranges, South Australia. Zootaxa, 3499 (1), 1–26. https://doi.org/10.11646/zootaxa.3499.1.1
    Donnellan, S.C., Catalano, S., Pederson, S., Mitchell, K., Suhendran, A., Price, L.C., Doughty, P. & Richards, S.J. (2021) Revision of the Litoria watjulumensis (Anura: Pelodryadidae) group from the Australian monsoonal tropics, including the resurrection of L. spaldingi. Zootaxa, 4933 (2), 211–240. https://doi.org/10.11646/zootaxa.4933.2.3
    Earl, D.A. (2012) Structure harvester: A website and program for visualizing structure output and implementing the Evanno method. Conservation Genetics Resources, 4, 359–361. https://doi.org/10.1007/s12686-011-9548-7
    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput, Nucleic Acids Research, 32, 1792–97. https://doi.org/10.1093/nar/gkh340
    Ehmann, H. (2013) Nullarbor crossing and biodiversity risk: Focus on the spotted-thighed bellfrog. Caring for Country, newsletter of Natural Resources Alinytjara Wiluṟara. Available from: from http://www.naturalresources.sa.gov.au/alinytjara-wilurara/search-results?dlv_AW%20DLV%20search%20results=(keyword=september%20edition%202013%20newsletter) (accessed 12 December 2022)
    Evanno, G., Regnaut, S. & Goudet, J. (2005) Detecting the number of clusters of individuals using the software structure: A simulation study. Molecular Ecology, 14, 2611–2620. https://doi.org/10.1111/j.1365-294X.2005.02553.x
    Ford, M.A. (1989) Vocalisation of the green frogs Litoria aurea and Litoria raniformis, including species-specific male calls. New Zealand Journal of Zoology, 16, 17–23. https://doi.org/10.1080/03014223.1989.10423700
    Fouquet, A., Vences, M., Salducci, M.D., Meyer, A., Marty, C., Blanc, M. & Gilles, A. (2007) Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera species groups. Molecular Phylogenetics and Evolution, 43, 567–582. https://doi.org/10.1016/j.ympev.2006.12.006
    Funk, W.C., Caminer M. & Ron, S.R. (2012) High levels of cryptic species diversity uncovered in Amazonian frogs. Proceedings of the Royal Society B: Biological Sciences, 279, 1806–1814. https://doi.org/10.1098/rspb.2011.1653
    Georges, A., Gruber, B., Pauly, G., White, D., Adams, M., Young, M., Kilian, A., Zhang, X., Shaffer, H. & Unmack, P. (2018) Genome‐wide SNP markers breathe new life into phylogeography and species delimitation for the problematic short‐necked turtles (Chelidae: Emydura) of eastern Australia. Molecular Ecology, 27, 5195–5213. https://doi.org/10.1111/mec.14925.
    Geyle, H.M., Hoskin, C.J., Bower, D.S., Catullo, R., Clulow, S., Driessen, M., Daniels, K., Garnett, S.T., Gilbert, D., Heard, G.W., Hero, J-M., Hines, H.B., Hoffmann, E.P., Hollis, G., Hunter, D.A., Lemckert, F., Mahony, M., Marantelli, G., McDonald, K.R., Mitchell, N.J., Newell, D., Roberts, J.D., Scheele, B.C., Scroggie, M.P., Vanderduys, E., Wassens, S., West, M., Woinarski, J.C.Z & Gillespie, G.R. (2021) Red hot frogs: identifying the Australian frogs most at risk of extinction. Pacific Conservation Biology, 28, 211–223. https://doi.org/10.1071/PC21019
    Gillespie, G. R., Roberts, J. D., Hunter, D., Hoskin, C.J., Alford, R.A., Heard, G.W., Hines, H., Lemckert, F., Newell, D. & Scheele, B.C. (2020) Status and priority conservation actions for Australian frog species. Biological Conservation, 247, 108543. https://doi.org/10.1016/J.BIOCON.2020.108543
    Gruber, B., Unmack, P.J., Berry, O.F. & Georges, A. (2018) dartr: An R package to facilitate analysis of SNP data generated from reduced representation genome sequencing. Molecular Ecology Resources, 18, 691–699. https://doi.org/10.1111/1755-0998.12745
    Hamer, A. & Organ, A. (2008) Aspects of the ecology and conservation of the Growling Grass Frog Litoria raniformis in an urban-fringe environment, southern Victoria. Australian Zoologist, 34, 393–407. https://doi.org/10.7882/AZ.2008.017
    Häupl, M. & Tiedemann, F. (1978) Vertebrata 1. Typenkatalog der Herpetologischen Sammlung. Kataloge der Wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien, 2, 7–34.
    Häupl, M., Tiedemann, F. & Grillitsch, H. (1994) Vertebrata 1. Katalog der Typen der Herpetologischen Sammlung nach dem Stand vom 1. Janner 1994. Kataloge der Wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien, 9, 1–42.
    Heard, G.W., Robertson, P. & Scroggie, M. (2004) The ecology and conservation status of the Growling Grass Frog (Litoria raniformis) within the Merri Creek Corridor. Second Report: Additional field survey and site monitoring. Wildlife Profiles Report to DSE and Yarra Valley Water. Wildlife Profiles Pty Ltd., Heidelberg, Victoria. [unknown pagination]
    Heard, G., Robertson, P. & Scroggie, M. (2008) Microhabitat preferences of the endangered growling grass frog Litoria raniformis in southern Victoria. Australian Zoologist, 34, 414–425. https://doi.org/10.7882/az.2008.019
    Heard, G., Scroggie, M. & Clemann, N. (2010) Guidelines for managing the endangered Growling Grass Frog in urbanising landscapes. Arthur Rylah Institute for Environmental Research, Heidelberg. [unknow pagination]
    Heard, G.W., Scroggie, M.P. & Malone, B.S. (2012) The life history and decline of the threatened Australian frog, Litoria raniformis. Austral Ecology, 37, 276–284. https://doi.org/10.1111/j.1442-9993.2011.02275.x
    Hebert, P.D.N., Cywinska, A., Ball, S.L. & DeWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences 270, 313–321. https://doi.org/10.1098/rspb.2002.2218.
    Hero, J.-M., Murray Littlejohn, M. & Marantelli, G. (1991) Frogwatch field guide to Victorian frogs. Dept. of Conservation and Environment, East Melbourne, Vic. 108 pp.
    Hero, J.M. & Morrison, C. (2004) Frog declines in Australia: global implications. Herpetological Journal, 14, 175–186.
    Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology Evolution, 35, 518–522. https://doi.org/10.1093/molbev/msx281
    Hoskin, C.J. (2004) Australian microhylid frogs (Cophixalus and Austrochaperina): phylogeny, taxonomy, calls, distributions and breeding biology. Australian Journal of Zoology, 52, 237–269. https://doi.org/10.1071/ZO03056
    Hoskin, C.J. (2007) Description, biology and conservation of a new species of Australian tree frog (Amphibia: Anura: Hylidae: Litoria) and an assessment of the remaining populations of Litoria genimaculata Horst, 1883: systematic and conservation implications of an unusual speciation event. Biological Journal of the Linnean Society, 91, 549–563. https://doi.org/10.1111/j.1095-8312.2007.00805.x
    Hoskin, C.J. (2012) Two new frog species (Microhylidae: Cophixalus) from the Australian Wet Tropics region, and redescription of Cophixalus ornatus. Zootaxa, 3271 (1), 1–16. https://doi.org/10.11646/zootaxa.3271.1.1
    Hoskin, C.J., Hines, H.B., Meyer, E., Clarke, J. & Cunningham, M. (2013) A new treefrog (Hylidae: Litoria) from Kroombit Tops, east Australia, and an assessment of conservation status. Zootaxa, 3646 (4), 426–446. https://doi.org/10.11646/zootaxa.3646.4.6
    Hunter, D., Clemann, N., Coote, D., Gillespie, G., Hollis, G., Scheele, B., Phillips, A. & West, M. (2018) Frog declines and associated management response in south-eastern mainland Australia and Tasmania. In: Heatwole, H. & Rowley, J. (Eds.), Amphibian Biology. Vol. 11. Amphibian Conservation in Australia, New Zealand and Pacific Islands. Surrey Beatty and Sons, Baulkham Hills, New South Wales, pp 39–58.
    Ineich, I., Bochaton, C. & Policard, J.P. (2015) «Les Grenouilles du Résident»: on the date of introduction of Litoria aurea (Lesson,1826) (Hylidae) into New Caledonia and Wallis Island (Wallis and Futuna) by Europeans. Herpetology Notes, 8, 495–499.
    Jombart, T. (2008) Adegenet: a R package for the multivariate analysis of genetic markers. Bioinformatics, 24, 1403–1405. https://doi.org/10.1093/bioinformatics/btn129
    Kalyaanamoorthy, S., Minh, B.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
    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., Mentjies, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649. https://doi.org/10.1093/bioinformatics/bts199
    Keely, C.C., Hale, J.M., Heard, G.W., Parris, K.M., Sumner, J., Hamer, A.J. & Melville, J. (2015) Genetic structure and diversity of the endangered growling grass frog in a rapidly urbanizing region Royal Society open science, 2, 140255. https://doi.org/10.1098/rsos.140255
    Keferstein, W. (1867) Ueber einige neue oder seltene Batrachier aus Australien und dem tropischen Amerika. Nachrichten von der Gesellschaft der Wissenschaften zu Göttingen, 1867, 341–361.
    Kilian, A., Wenzl, P., Huttner, E., Carling, J., Xia, L., Blois, H., Caig, V., Heller-Uszynska, K., Jaccoud, D., Hopper, C., Aschenbrenner-Kilian, M., Evers, M., Peng, K., Cayla, C., Hok, P. & Uszynski, G. (2012) Diversity Arrays Technology: a generic genome profiling technology on open platforms. Methods in Molecular Biology, 888, 67–89. http://doi.org/10.1007/978-1-61779-870-2_5.
    Knowles, R., Mahony, M., Armstrong, J. & Donnellan, S.C. (2004) Systematics of sphagnum frogs of the genus Philoria (Anura: Myobatrachidae) in eastern Australia. Records of the Australian Museum, 56, 57–74. https://doi.org/10.3853/j.0067-1975.56.2004.1391
    Köhler, J., Vieites, D.R., Ronald, R.M., García, F.H., Glaw, F., Steinke, D. & Vences, M. (2005) New amphibians and global conservation: A boost in species discoveries in a highly endangered vertebrate group. BioScience, 55, 693–696. https://doi.org/10.1641/0006-3568(2005)055[0693:NAAGCA]2.0.CO;2
    Köhler, J., Jansen, M., Rodríguez, A., Kok, P., Toledo, L., Emmrich, M., Glaw, F., Haddad, C. & Rödel, M. & 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
    Kopelman, N.M., Mayzel, J., Jakobsson, M., Rosenberg, N.A. & Mayrose, I. (2015) CLUMPAK: a program for identifying clustering modes and packaging population structure inferences across K. Molecular Ecology Resources, 15, 1179–1191. https://doi.org/10.1111/1755-0998.12387
    Leaché, A.D., Banbury, B.L., Felsenstein, J., de Oca, A.N. & Stamatakis, A. (2015) Short tree, long tree, right tree, wrong tree: new acquisition bias corrections for inferring SNP phylogenies. Systematic Biology, 64, 1032–1047. https://doi.org/10.1093/sysbio/syv053
    Lehtinen, R.M., Nussbaum, R.A., Richards, C.M., Cannatella, D.C. & Vences, M. (2007) Mitochondrial genes reveal cryptic diversity in plant-breeding frogs from Madagascar (Anura, Mantellidae, Guibemantis). Molecular Phylogenetics and Evolution, 44, 1121–1129. https://doi.org/10.1016/j.ympev.2007.05.020
    Littlejohn, M. & Martin, A.A. (1969) Mating call as an aid to the identification of the frogs of the Melbourne area. Victorian Naturalist, 86, 126–127.
    Littlejohn, M. & Martin, A.A. (1974) The amphibia of Tasmania. In: Williams, W.D. (Ed.), ‘Biogeography and ecology of Tasmania’. Junk, The Hague, pp. 251–289.
    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
    Loftus-Hills, J.J. (1971) Comparative aspects of auditory function in Australian anurans. Australian Journal of Zoology, 21, 353–367. https://doi.org/10.1071/ZO9730353
    Mace, G.M. (2004) The role of taxonomy in species conservation. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 359, 711–719. https://doi.org/10.1098/rstb.2003.1454
    Mahony, M. (1999) Review of the declines and disappearances within the bell frog species group (Litoria aurea species group) in Australia. In: Campbell, A. (Ed.), ‘Declines and Disappearances of Australian Frogs’. Environment Australia: Canberra, pp. 81–93.
    Mahony, M.J., Knowles, R., Foster, R. & Donnellan, S.C. (2001) Systematics of the Litoria citropa (Anura: Hylidae) complex in northern New South Wales and southern Queensland, Australia, with the description of a new species. Records of the Australian Museum, 53, 37–48.
    10.3853/j.0067-1975.53.2001.1322
    Mahony, M.J., Hamer, A.J., Pickett, E.J., McKenzie, D.J., Stockwell, M.P., Garnham, J.I., Keely, C.C., Deboo, M.L., O’Meara, J., Pollard, C.J., Clulow, S., Lemckert, F.L, Bower, D.S. & Clulow, J. (2013) Identifying conservation and research priorities in the face of uncertainty: a review of the threatened bell frog complex in eastern Australia. Herpetological Conservation Biology, 8, 519–538.
    Mahony, M.J., Moses, B., Mahony, S., Lemckert, F. & Donnellan, S.C. (2020) A new species of frog in the Litoria ewingii species group (Anura: Pelodryadidae) from south-eastern Australia. Zootaxa, 4858 (2), 201–230. https://doi.org/10.11646/zootaxa.4858.2.3
    Mahony, M.J., Penman, T., Bertozz,i T., Lemckert, F., Bilney, R. & Donnellan, S.C. (2021a) Taxonomic revision of south-eastern Australian giant burrowing frogs (Anura: Limnodynastidae: Heleioporus Gray). Zootaxa, 5016 (4), 451–489. https://doi.org/10.11646/zootaxa.5016.4.1
    Mahony, M.J., Hines, H.B., Mahony, S., Moses, B., Catalano, S.R., Myers, S. & Donnellan, S.C. (2021b) A new hip-pocket frog from mid-eastern Australia (Anura: Myobatrachidae: Assa). Zootaxa, 5057 (4), 451–486. https://doi.org/10.11646/zootaxa.5057.4.1
    Mahony, M.J., Hines, H.B., Bertozzi, T., Mahony, S. & Donnellan, S.C. (2022) A sixth species of northern Philoria (Anura: Myobatrachidae) from the uplands of the World Heritage forests of eastern Australia. Zootaxa, 5104 (2), 209–241. https://doi.org/10.11646/zootaxa.5104.2.3
    Mann, R., Hyne, R., Selvakumaraswamy, P. & Barbosa, S. (2010) Longevity and larval development among southern bell frogs (Litoria raniformis) in the Coleambally irrigation area implications for conservation of an endangered frog. Wildlife Research, 37, 447–455. https://doi.org/10.1071/WR10061
    Mcdonald, K.R., Rowley, J.J.L., Richards, S.J.& Frankham, G.J. (2016) A new species of treefrog (Litoria) from Cape York Peninsula, Australia. Zootaxa, 4171 (1), 153–169. https://doi.org/10.11646/zootaxa.4171.1.6
    Meegaskumbura, M., Bossuyt, F., Pethiyagoda, R., Manamendra-Arachchi, K., Bahir, M., Milinkovitch, M.C. & Schneider, C.J. (2002) Sri Lanka: An amphibian hot spot. Science, 298, 379–379. https://doi.org/10.1126/science.298.5592.379
    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 Evolution, 32, 268–274. https://doi.org/10.1093/molbev/msu300
    Osborne, W.S., Littlejohn, M.J. & Thomson, S.A. (1996) Former distribution and apparent disappearance of the Litoria aurea complex from the Southern Tablelands of New South Wales and the Australian Capital Territory. Australian Zoologist, 30, 190–198. https://doi.org/10.7882/AZ.1996.011
    Pepper, M., Keogh, J.S. & Chapple, D.G. (2016) Molecular biogeography of Australian and New Zealand reptiles and amphibians. In: Ebach, M.C. (Ed.), Handbook of Australasian Biogeography. CRC Press, Boca Raton, Florida, pp. 295–328.
    Pfenninger, M. & Schwenk, K. (2007) Cryptic animal species are homogeneously distributed among taxa and biogeographical regions. BMC Evolutionary Biology, 7, 121. https://doi.org/10.1186/1471-2148-7-121
    Pritchard, J.K., Stephens, M. & Donnelly, P. (2000) Inference of population structure using multilocus genotype data. Genetics, 155, 945–959. https://doi.org/10.1093/genetics/155.2.945
    Pyke, G.H. (2002) A review of the biology of the southern bell frog Litoria raniformis (Anura: Hylidae). Australian Zoologist, 32, 32–48. https://doi.org/10.7882/AZ.2002.003
    Rambaut, A. & Drummond, A.J. (2007) Tracer v1.4. http://beast.bio.ed.ac.uk/Tracer.
    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.
    Robertson, P., Heard, G.W. & Scroggie, M.P. (2002) The ecology and conservation status of the Growling Grass Frog (Litoria raniformis) within the Merri Creek Corridor. Interim Report: Distribution, Abundance and Habitat Requirements. Report produced for the Department of Natural Resources and Environment. Wildlife Profiles Pty Ltd, Heidelberg, Victoria. [unknown pagination]
    Robinson, M. (1993) A field guide to frogs of south eastern Australia. From Port Augusta to Fraser Island including Tasmania. Australian Museum/Reed Books, Chatswood, 112 pp.
    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. https://doi.org/10.3897/zookeys.912.38253
    Rowley, J., Mahony, M.J., Hines, H.B., Myers, S., Aplin, K.P., Price, L. & Donnellan, S.C. (2021) Two new species from the Litoria rubella species group from eastern Australia. Zootaxa, 5071 (1), 1–41. https://doi.org/10.11646/zootaxa.5071.1.1
    Sadlier, R.A. & Pressey, R.L. (1994) Reptiles and amphibians of particular conservation concern in the western division of New South Wales –a preliminary review. Biological Conservation, 69, 41–54. https://doi.org/10.1016/0006-3207(94)90327-1
    Sanders, M.G. (2021) Photographic field guide to Australian frogs. CSIRO Publishing, Melbourne, 376 pp.
    Steindachner, F. (1867) Amphibien. In Reise der Osterreichieschen Freggate Novara um die Erde in den Jaren 1857–1859. Vienna Zoologische Thiel, 1, 1–70.
    Stuart, B.L., Inger, R.F. & Voris, H.K. (2006) High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs. Biology Letters, 2, 470–474. https://doi.org/10.1098/rsbl.2006.0505
    Swofford, D.L. (2003) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]
    Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution, 38, 3022–3027. https://doi.org/10.1093/molbev/msab120
    Taylor, C.M., Keppel, G., Peters, S., Hopkins, G.R. & Kerr, G.D. (2018) Establishment and potential spread of the introduced spotted-thighed frog, Litoria cyclorhyncha (Ranoidea cyclorhynchus), in South Australia. Transactions of the Royal Society of South Australia, 142, 86–101. https://doi.org/10.1080/03721426.2018.1444911
    Taylor, C.M., Keppel, G., O’Sullivan, S., Peters, S., Kerr, G.D. & Williams, C.R. (2020) Indiscriminate feeding by an alien population of the spotted-thighed frog (Litoria cyclorhyncha) in southern Australia and potential impacts on native biodiversity. Australian Journal of Zoology, 67, 59–72. https://doi.org/10.1071/ZO19042
    Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M., Beaumont, L.J., Collingham, Y.C., Erasmus, B.F.N., de Siqueira, M.F., Grainger, A., Hannah, L., Hughes, L., Huntley, B., van Jaarsveld, A.S., Midgley, G.F., Miles, L., Ortega-Huerta, M.A., Townsend Peterson, A., Phillips, O.L. & Williams, S.E. (2004) Extinction risk from climate change. Nature, 427, 145–148. https://doi.org/10.1038/nature02121
    Thomson, G.M. (1922) The Naturalisation of Animals and Plants in New Zealand. Cambridge University Press, Cambridge. [unknown pagination]
    Thomson, S.A., Littlejohn, M.J., Robinson, W.A. & Osborne, W.S. (1996) Taxonomy of the Litoria aurea complex: a re-evaluation of the Southern Tableland populations of the Australian Capital Territory and New South Wales. Australian Zoologist, 30, 158–169. https://doi.org/10.7882/AZ.1996.008
    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
    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
    Turner, A. (2022) Environmental predictors of disease risk for Litoria raniformis populations in south-eastern Australia. Doctoral thesis, Charles Sturt University, Albury. [unknown pagination]
    Turner, A., Wassens, S. & Heard, G. (2021) Chytrid infection dynamics in frog populations from climatically disparate regions. Biological Conservation, 264, 109391. https://doi.org/10.1016/j.biocon.2021.109391
    Tyler, M.J. (1997) The Action Plan for Australian Frogs. Wildlife Australia, a branch of Environment Australia, Commonwealth Department of Environment, Sport and Territories, Canberra. [unknown pagination]
    Tyler, M. J. & Davies, M. (1978) Species-groups within the Australopapuan hylid frog genus Litoria Tschudi. Australian Journal of Zoology, Supplementary Series, Number 63, 1–47.
    Uetz, P. (2021) The reptile database. Available from: http://www.reptile-database.org (accessed 20 October 2021.
    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 U.S.A., 106, 8267–8272. https://doi.org/10.1073/pnas.081082110
    Vörös, J., Mitchell, A., Waldman, B., Goldstien, S. & Gemmell, N. (2008) Crossing the Tasman Sea: Inferring the introduction history of Litoria aurea and Litoria raniformis (Anura: Hylidae) from Australia into New Zealand. Austral Ecology, 33, 623–629. https://doi.org/10.1111/j.1442-9993.2007.01828.x
    Waldman, B., van de Wolfshaar, K.E., Klena, J.D., Andjic, V., Bishop, P.J. & Norman, R.J.D.B. (2001) Chytridiomycosis in New Zealand frogs. Surveillance, 28, 9–11.
    Wassens, S. (2008) Review of the past distribution and decline of the southern bell frog Litoria raniformis in New South Wales. Australian Zoologist, 34, 446–452. https://doi.org/10.7882/AZ.2008.022
    Wassens, S., Hall, A., Osborne, W. & Watts, R. (2010) Habitat characteristics predict occupancy patterns of the endangered amphibian Litoria raniformis in flow-regulated flood plain wetlands. Austral Ecology, 35, 944–955. https://doi.org/10.1111/j.1442-9993.2010.02106.x
    White, A.W. & Pyke, G.H. (1999) Past distribution of Litoria aurea and Litoria castanea in the Bathurst-Orange Area of New South Wales. Herpetofauna, 29, 2–9.