Molecular systematic analysis demonstrates that the threatened southern bell frog, Litoria raniformis (Anura: Pelodryadidae) of eastern Australia comprises two sub-species Author Vörös, Judit Department of Zoology, Hungarian Natural History Museum H- 1088 Budapest, Baross u. 13, Hungary Author Wassens, Skye 0000-0001-8886-8426 School of Agriculture, Environment and Veterinary Sciences, Charles Sturt University, PO Box 789, Albury, 2640, Australia. swassens @ csu. edu. au; https: // orcid. org / 0000 - 0001 - 8886 - 8426 swassens@csu.edu.au Author Price, Luke School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia. Author Hunter, David New South Wales Department of Planning, Industry and Environment, Albury, 2640, Australia. David. hunter @ environment. nsw. gov. au Author Myers, Steven 0000-0001-8885-8770 South Australian Museum, North Terrace, Adelaide, 5001, Australia. & Steven. Myers @ alsglobal. com; https: // orcid. org / 0000 - 0001 - 8885 - 8770 yers@alsglobal.com Author Armstrong, Kyle School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia. & South Australian Museum, North Terrace, Adelaide, 5001, Australia. Author Mahony, Michael J. 0000-0002-1042-0848 School of Environmental and Life Sciences, University of Newcastle, University Drive, Callaghan NSW 2308, Australia michael. mahony @ newcastle. edu. au; https: // orcid. org / 0000 - 0002 - 1042 - 0848 michael.mahony@newcastle.edu.au Author Donnellan, Stephen 0000-0002-5448-3226 South Australian Museum, North Terrace, Adelaide, 5001, Australia. & Steve. donnellan @ samuseum. sa. gov. au; https: // orcid. org / 0000 - 0002 - 5448 - 3226 teve.donnellan@samuseum.sa.gov.au text Zootaxa 2023 2023-01-11 5228 1 1 43 journal article 222253 10.11646/zootaxa.5228.1.1 7b9a994e-a6ec-4efb-a755-e1e8789a2ee0 1175-5326 7524042 DA207B1D-81A2-4176-AA9A-64B1D484C307 Litoria raniformis raniformis ( Keferstein, 1867 ) Figs 12 , 13 Neotype . AMS R45668 , an adult male from Hay , 50 miles on Balranald Road , NSW, Australia . 34.683° S ; 144.05° E . Collected by Harald Ehmann on 9 September 1973 . Diagnosis . Litoria raniformis can be diagnosed from other members of the Litoria aurea Group by the presence of vertebral stripe (vs absent in L. aurea and L. cyclorhyncha ), dorsum with longitudinally aligned raised tubercles and skin ridges (vs absent in L. aurea ); presence of black canthal stripe (vs absent in L. moorei and variably present in L. cyclorhyncha ); pattern-less ventral surface (vs black reticulations ventrally in L. cyclorhyncha ); absence of distinctive large black ringed yellow or cream spots in the groin, and pale yellow spots and/or marbling on the lower surface of the hind limbs (vs present in L. castanea ). Measurements of neotype (mm). SVL: 60.1, HL: 21.4, HW: 21.4, IND: 3.7, NS: 4.1, IOD: 3.3, ED: 5.96, TD: 4.5, FeL: 26.9, TL: 26.7, FoL: 37.8, FLL: 9.8, RLF: 25.2, D1T: 1.3, D4T: 1.5, D1F: 1.5, D3F: 1.3. Description of Neotype . Snout prominent, rounded when viewed from above and in profile. Nostrils more lateral than superior, closer to snout than to eye. Canthus rostralis well defined and straight. Eye relatively large (ED/HL 0.28). Tympanum distinct, circular, length less than eye diameter (TD/ED 0.75). Vomerine teeth short straight plates bridging the gap between the choanae. Tongue approximately rectangular. Fingers long, broad, webbing absent. Subarticular and palmar tubercles prominent.Terminal discs not prominent, barely extending beyond lateral extremities of penultimate phalanx. Fingers in order of length 3>4>1>2. Hindlimb length moderate (TL/SVL 0.48). Toes in order of length 4>5=3>2>1. Webbing on toes I, II, III, V reaches base of terminal toe disc, and to base of the penultimate phalange on toe IV. Subarticular tubercles prominent. Oval inner metatarsal tubercle prominent, approximately one-third length of first toe. Terminal toe discs not prominent, not extending beyond lateral extremities of penultimate phalanx. Distinct pectoral fold present ( Fig. 12B ). FIGURE 12. The neotype designated for Litoria r. raniformis ( Keferstein, 1867 ) AMS R 45668. A ) dorsal, B ) ventral and C ) lateral view. Image by Ron Lovatt © Australian Museum DigiVol. Variation. A summary of variation in 17 mensural traits is presented in Table 6 . Mean SVL: females = 69.3 mm , males = 61.7 mm . Head length equal to head width (HL/HW 0.87–1.1) and approximately one-third of SVL (HL/SVL 0.32–0.45). Eye relatively large, diameter equal to or greater than eye to naris distance (ED/EN 0.99– 1.44). Pupil horizontal when constricted ( Fig. 13 ). Tympanum length greater than half eye diameter (TD/ED 0.68- 0.92). Hindlimb length moderate and variable (TL/SVL 0.41–0.64). Oval inner metatarsal tubercle prominent. Light brown nuptial pad wraps around inner surface of digit I from base of digit to base of penultimate phalange (SAMA R59848 ). Small vocal slits present in buccal cavity at base of tongue parallel to lateral margin of tongue. FIGURE 13. Images, in life of Litoria r. raniformis . A) Gayini, Lowbidgee floodplain, NSW, Damian Michael; B ) Balranald, NSW Stephen Mahony; C ) 9km N of Lake Cullulleraine, NSW, Peter Robertson; D) Gayini, Lowbidgee floodplain, NSW, Damian Michael; E) Lake Victoria, NSW, Peter Robertson; F ) Wallpolla Island, west of Mildura, Vic. Peter Robertson. Dorsum mildly granular with varying coverage of low tubercles. Upper surface of limbs smooth or with varying coverage of low tubercles. Flanks with dense coverage of low tubercles. Chin, abdomen, undersurface of limbs, and lateral aspect of body coarsely granular or granularity confined to lateral margins of abdomen. Colour pattern. Variation in colour is described from images taken in life ( Fig. 13 ). Head smooth with either uniform colour or heavily marked with dark patches. Iris gold. Canthus rostralis typically uniformly colored with the light tones of the dorsum. Darker tympanum contrasts with lighter loreal region. Prominent green to bronze mid-dorsal stripe extends from the snout or from between the eyes to the cloaca. Back covered in prominent dark tubercles arranged either in lines or irregularly. Dorsal colour varies from green to bronze sometimes with darker patches. A clearly demarked line with a black lower margin runs from the nostril through the eye, above the tympanum and then along the dorso-lateral margin to the groin. In the dorso-lateral region its upper margin is enhanced by prominent cream to white tubercles. Flanks with prominent cream or white coloured coarse granules in upper half. Colour pattern of limbs is a continuation of dorsal colour and pattern, leading edge of lower leg with prominent black margin. Back of thighs awash with emerald green to blue, moderately granular proximally with few to moderate numbers of prominent light coloured tubercles. Belly and chin cream or white, undersurfaces of limbs either cream or with light green flush, with dark patches. Groin regions with similar pattern and colour to back of thigh or less frequently with yellow spots with black outline, particularly tableland populations in the east of the species range. Throat dusky in mature males. Distribution. Southern New South Wales , northern Victoria and the Murray Valley in South Australia . It occurs in the IBRA regions: Murray Darling Depression, NSW South Western Slopes, Riverina, South Eastern Highlands, and Victorian Midlands. There is absence of records of L. raniformis from the alpine regions in north-eastern Victoria and south-eastern New South Wales and in the mallee country of western Victoria and eastern South Australia . Identity of vouchers from the southern tablelands of NSW assigned to L. raniformis by Thomson et al. (1996) needs confirmation with ancient DNA analyses. Our genetic data identified L. r. raniformis at seven locations in the central Mount Lofty Ranges south-east of Adelaide, all collected between 1972 and 1979. Ecology and reproductive biology. Populations occur in swamps dominated by eucalypts ( Eucalyptus camaldulensis , E. largiflorens ), Lignum, Typha, and Nitre Goosefoot ( Chenopodium nitrariaceum ), and also in irrigated rice crops ( Wassens 2008 ). Common features of occupied wetlands are that they are large, continuous areas containing both permanent and ephemeral waterbodies that undergo regular flooding and are surrounded by areas containing suitable refugia in the form of ground debris, vegetation cover and cracking soils ( Wassens 2008 ). Frogs concentrate in refugia prior to flooding, then disperse across the landscape during flooding events ( Wassens 2008 ). Breeding is triggered by flooding during spring or summer ( Wassens 2008 ; Wassens et al. 2010 , Turner 2022 ). The larval phase may be as short as two months ( Wassens 2008 ).