Review of the past distribution and decline of the Southern Bell Frog* Litoria raniformis in New South Wales Skye Wassens Institute for Land, Water and Society, School of Environmental Sciences, Charles Sturt University, Locked bag 588, Wagga Wagga, NSW, 2678, Australia. [email protected] The Southern Bell Frog Litoria raniformis was formally widespread throughout south eastern Australia. Poor knowledge of the past and present distribution of populations and the pattern of decline currently limits our ability to plan conservation measures. The past and present distribution of L. raniformis in New T South Wales (NSW) is described through the use of museum records, wildlife databases, reports and surveys. Survey identified a further 10 populations, all in lower Murrumbidgee floodplain (Lowbidgee), and C confirmed the absence of L. raniformis in past habitats along the middle reaches of the Murrumbidgee River. A Based on past records, L. raniformis declines have been substantial. The eastern edge of the geographic range has shifted approximately 500 km west from the South Eastern Highlands, and the northern edge R of its range has shifted 200 km south from the most northerly population in Willandra National Park. T Litoria raniformis is currently restricted to the semi-arid zone of NSW with populations occurring in the S Lower Murrumbidgee and Murray River, the Wakool Irrigation Area and the Coleambally Irrigation Area. B The wide geographic range of L. raniformis prior to the decline means that a single causal factor is unlikely. A Former habitats throughout the original range have been highly modified as a result of a range of factors, including flooding regulation, livestock grazing and exotic introductions. Key words: Litoria raniformis, historic distribution, range decline Introduction The identification of the extent and timing of a species’ Prior to the early 1980’s the Southern Bell Frog (Litoria decline can yield important clues about the key agents raniformis) was considered to be widespread and that have acted upon it over time (Caughley and Gunn locally abundant throughout south-eastern Australia 1996). To determine these agents, declines need to be and Tasmania (Barker and Grigg 1977). It occupied measured at spatial and temporal scales large enough to a range of habitat types including small permanent separate natural fluctuations in population numbers from wetland systems, streams, dams, irrigation areas and anthropogenic declines (Shaffer et al. 1998). In the absence large ephemeral wetlands (Mahony 1999). of long-term monitoring programs, determination of the long-term changes in a species’ geographic distribution When local extinctions were first noted in the early can be achieved through comparisons of past and present 1990’s in the South Eastern Highlands of New South records of a species’ distribution (Mingozzi and Esteve Wales (NSW), they were described as sudden, and the 1997; Shaffer et al. 1998). Records that are suitable for actual causes for the declines were poorly understood this purpose include natural history collections (Shaffer (Osborne et al. 1996). In contrast, declines and local et al. 1998), catch rates (Zwanenburg 2000), bag numbers extinctions in Tasmania appeared to occur as a result of (Thirgood et al. 2000) pelt sales (Jimenez 1996; Riley et wetland degradation, with some declines exacerbated by al. 2004), natural history accounts (Mingozzi and Esteve the 1980’s drought (Ashworth 1998). 1997) or a combination of the above. Declines in the abundance and distribution of L. A long-term approach to measuring changes in distribution raniformis in South Australia, Victoria and inland areas is especially important for amphibian species which often of NSW have not been formerly documented through exhibit chaotic population dynamics that are strongly dedicated surveys but are thought to be significant (Tyler linked to the availability of temporary resources (Blaustein 1997; Mahony 1999; Pyke 2002). The aim of this review et al. 1994; Biek et al. 2002). For these species separation is to describe, in detail the past and present distribution of “natural” and anthropogenically driven declines is of L. raniformis and its habitats in NSW, with special important, especially when correlations between the emphasis on records from the Murrumbidgee floodplain decline and the onset of potential causal agents are sought and its tributaries in the South West Slopes, Riverina (Kiesecker et al. 2001). Natural history collections and literature have been used to determine the status and and Murray-Darling Depression biogeographic regions. patterns of declines of riverine frogs (Gillespie and Hines This review builds on the general national distribution 1999) and the Green and Golden Bell Frog (Litoria aurea) and decline maps of Mahony (1997) and Pyke (2002) (Gillespie 1996; Osborne et al. 1996) in Australia. using natural history and National Parks and Wildlife *Referred to as the Growling Grass Frog in Victoria Australian 446 Zoologist volume 34 (3) October 2008 Past distribution and decline of the Southern Bell Frog Service Atlas database records reports, informal records there were discrepancies between map coordinates and and field surveys of important wetland systems in location information, the latter was used. Where no map the mid and lower Murrumbidgee Floodplain. These coordinates were supplied, the coordinates of the nearest included some wetlands where L. raniformis had been town were used. recorded previously. The quality of data ranged from good to very poor with some museum records lacking location data or collection Methods dates these were excluded from further analysis.. There is a single record for L. raniformis from Pambula in the South Frog Surveys Eastern Corner biogeographic region (Australian Museum Surveys were conducted in wetland systems where Collection) which is outside the accepted range for L. L. raniformis could potentially occur, or where there raniformis (Mahony 1999). This record has been excluded were past records. This included the wetlands along from consideration because there is a high probability that the Murrumbidgee River between Wagga Wagga and the specimen has been incorrectly labelled. Balranald, as well as the Coleambally and Murrumbidgee Once error checking was complete, the coordinates for Irrigation Areas. Australian Bureau of Meteorology all locations where L. raniformis had been recorded were (BOM) weather records for Coleambally, Narrandera, imported into a GIS database (ArcMap version 8.3 ESRI). Hay and Balranald were obtained for the study periods. The database also included; NSW rivers, NSW and ACT All surveys were conducted at night during warm biogeographic regions (Environment Australia 2000), conditions, when daytime temperatures exceeded 25oC. NSW wetland systems (Environment Australia 2001), A total of 44 sites were surveyed between October 2001 NSW state borders, and NSW towns and localities. and November 2002 and then again between October 2003 and January 2004. Information on the distribution of L. raniformis from all data sources was mapped using the revised Interim Due to the size of the study region and the large number Biogeographic Regionalisation for Australia (IBRA) of wetlands to be surveyed, a rapid survey approach was (Environment Australia 2000) (hereafter referred to adopted. Surveys were conducted after dark, employing a as biogeographic regions). This approach was used for minimum of two 5-minute audio surveys, one 2-minute two reasons. Firstly, biogeographic regions more closely call playback and two active visual surveys along 100 m reflect the biological and geological character of parts transects per 10 hectares of waterbody. Audio surveys of L. raniformis’ range than government or topographic involved standing quietly by the waters’ edge and listening units. Secondly, the use of biogeographic regions was for calls for 5-minute periods. If no calls were heard, call considered useful because of the sparse nature of frog playback was employed for two minutes followed by a records in NSW. This meant that the failure to record further 5-minute listening period. Active visual searches L. raniformis within individual wetlands where it had were conducted along 100-m transects, 1 metre from and occurred previously might occur simply because the area parallel to the waters’ edge. A 50-watt spotlight was used was never re-surveyed or that the results of surveys were in a side to side motion to search a 1-metre wide area on not reported. In contrast, the probability of L. raniformis either side of the transect and in fringing vegetation. being recorded at least once within a whole biogeographic Road transect surveys were also conducted in the region over a 10-year period is quite high. Coleambally Irrigation Area (CIA), the Hay Plain and the There are six biogeographic regions in NSW that occur Murrumbidgee Irrigation Area because the comprehensive within the accepted range of L. raniformis. These are the network of roadways allowed transects to be driven South Eastern Highlands, Australian Alps, NSW South between rice bays and along irrigation canals, which are Western Slopes, Riverina, Cobar Peneplain and the often utilised by L. raniformis (AMBS Consulting 2000). Murray-Darling Depression. This technique involved five minute audio-surveys every 5 km in any area that contained water such as irrigation Results canals and dams as well as spotlighting along the road edges while travelling at less than 10km/hour. Survey results Historical records Litoria raniformis was located in two distinctly different wetland types in the Lowbidgee region in the Murray- Records for L. raniformis voucher specimens collected in Darling Depression and western edge of the Riverina NSW between 1900 and 2003 were obtained from the biogeographic region. These were River Red Gum Australian Museum, South Australian Museum, Museum (Eucalyptus camaldulensis) and Black Box (E. largiflorens)/ Victoria, and the Australian National Wildlife Collection Lignum (Muelenbeckia florulenta). Both River Red Gum (CSIRO). The NSW National Parks and Wildlife Services and Black Box/Lignum wetlands are normally subject to Atlas database records for L. raniformis were obtained in flooding in 4 out of every 5 years (Kingsford and Thomas October 2003. Information on the present status of L. 2001). raniformis was obtained from published and unpublished fauna surveys and articles. Litoria raniformis was present on rice growing properties within the CIA, occurring at all 19 sites surveyed. Litoria The longitude and latitude and easting northing coordinates raniformis was not located in any of the wetlands that form for each voucher specimen and database record were part of the Mid Murrumbidgee wetland system. checked against town and locality information. Where Australian October 2008 Zoologist volume 34 (3) 447 Wassen Description of historic and current There is some disagreement as to whether L. raniformis distribution remains in the Monaro region at the southern end of the South Eastern Highlands. Mahony (1999) notes that L. In total, 174 records of L. raniformis were obtained from raniformis was present in the area in the 1990’s, however the dedicated surveys, museum records, the NSW NPWS surveys of this region in the late 1990’s failed to locate L. ATLAS database and four reports (Osborne et al. 1996; raniformis (Osborne, pers. com. 2004). White and Pyke 1999; AMBS Consulting 2000; Wassens 2006). The majority of records were for the Riverina South West Slopes bioregion followed by the South Eastern Highlands. Only Litoria raniformis appears to have been widespread four records were from the Cobar Peneplain, which may throughout the South Western Slopes of NSW, particularly be a result of the limited surveys conducted in this region along the Murrumbidgee and Murray River systems and (Table 1). their tributaries until the early 1980’s. However, due to Based on the combination of known past and present the limited records and small number of frog surveys habitats occupied by L. raniformis in NSW, declines conducted, it is difficult to determine exactly when occurred almost simultaneously in the northern and local extinctions occurred after this. Jansen and Healey eastern parts of the species’ geographic range (Table (2003) surveyed wetlands along the Murrumbidgee River, 1). As a result of these local extinctions, the eastern including historical L. raniformis habitats such as Berry edge of the geographic range of L. raniformis has shifted Jerry Lagoon and the Mid Murrumbidgee Wetlands, approximately 500 km west from the South Eastern from Gundagai in the east of the Southern Western Highlands, and the northern edge of its range has shifted Slopes Biogeographic region to Hay in the Riverina 200 km south from the most northerly population in Biogeographic region, and failed to locate L. raniformis. Willandra National Park. Surveys of wetlands along the Murrumbidgee tributaries in the South West Slopes between 2002 and 2004, South Eastern Highlands and Australian Alps including Buckingbong State Forest and Medium Swamp The most northerly records of L. raniformis are from the where L. raniformis had been recorded previously, also South Eastern Highlands near Bathurst west of the Great failed to locate it (Leigh Thompson, Johnstone Centre Dividing Range (Fig. 1). Litoria raniformis was recorded at Environmental Consulting, pers. com.). a number of wetlands this region during the 1970’s but was The status of populations of L. raniformis along the locally extinct by the 1980’s (Ehmann and White 1997). Murray River in the South West Slopes is uncertain. It Litoria raniformis was also formerly widespread in the appears to have been quite widespread in wetlands around southern part of the South Eastern Highlands (Osborne Woomargama National Park, near Albury (Caughley and et al. 1996). Prior to 1980 it was considered to be relatively Gall 1985), but there have been no confirmed records common in the ACT, occurring in natural wetlands such of this species from this area since 1979. There is also as Lake George near Collector and artificial wetlands in an unconfirmed record of L. raniformis occurring in the Canberra including Lake Ginninderra and Lake Burley Wonga wetlands near Albury in 1999 (NPWS, 2003) but Griffin (Osborne et al. 1996). Further south, L. raniformis the accuracy of this record is questionable (Matt Herring, was recorded on multiple occasions near Cooma and Charles Sturt University, pers. com.). Nimmitabel prior to 1979. In the extreme south of the biogeographic region, L. raniformis was recorded from the Cobar Peneplain Monaro Lakes near Bombala on 11 separate occasions Litoria raniformis was first collected from the Cobar between 1955 and 1980 (Australian Museum Collection). Peneplain biogeographic region in Willandra National Park These wetlands are a series of permanent and ephemeral near Hillston in 1928 (Australian Museum Collection) upland wetlands ranging in size from 0.5 to 215 ha and again in 1985 (NPWS, 2003). Within Willandra (Australian Nature Conservation Agency 1996). On National Park L. raniformis is likely to have occurred in the western side of the Australian Alps L. raniformis was the Willandra Creek and ephemeral wetlands associated recorded around Talbingo until 1980 and near Khancoban with Willandra and Yangellawah Creeks, which are in the early 1970’s. distributaries of the Lachlan River. Litoria raniformis was Table 1. Number of L. raniformis records, including new data collected during the surveys, in each biogeographic region in NSW and the ACT during each decade since 1900. 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 total Australian Alps 1 1 1 4 7 South Eastern Highlands 1 2 10 17 16 46 Cobar peneplain 1 2 1 4 South West Slopes 1 4 16 1 1 23 Riverina 3 4 11 13 1 9 18 59 Murray-Darling Depression 3 1 2 2 15 9 32 171 Total NSW & ACT 2 0 1 4 10 29 59 22 25 27 Australian 448 Zoologist volume 34 (3) October 2008 Past distribution and decline of the Southern Bell Frog Figure 1. The distribution of L. raniformis populations over 20 year time intervals in each biogeographic region: AA Australian Alps, SEH South eastern Highlands, SWS South West Slopes, RIV Riverina, MDD Murray –Darling Depression. last recorded in Willandra National Park in 1985 and has but has not been recorded since. In the south east, L. not been recorded since, despite surveys by students from raniformis was found at Barron Box Swamp near Griffith the University of Canberra (Dr Will Osborne, University in 1967, which at this time was a large ephemeral wetland of Canberra, pers. com.). Willandra Creek, which is system (Australian National Wildlife Collection). This the main waterway in the park, is presently subject to wetland was surveyed in 2003 along with all other major flow regulation and contains water in all but severe wetland systems in the Murrumbidgee Irrigation Area, but drought conditions (New South Wales National Parks no L. raniformis individuals were located (Wassens 2006). and Wildlife Service 1989). The Willandra National Park Barron Box Swamp was converted for use as permanent 1989 management plan does not mention L. raniformis or water storage in the late 1970’s (Briggs et al. 1997). In the potential habitats (New South Wales National Parks and middle reaches of the Murrumbidgee River, L. raniformis Wildlife Service 1989). There are a number of records was collected from oxbow lagoons between Narrandera from the southern edge of this region, mainly from and Darlington Point between 1957 and 1979. These wetland systems west of Murrami (Museum Victoria). sites included the mid Murrumbidgee wetlands, Yanco Agricultural High School wetlands and Darlington Riverina Point wetlands (Australian Museum Collection; South Litoria raniformis was widespread throughout the Riverina Australia Museum). Jansen and Healey (2003) surveyed biogeographic region and individuals still persist in historical L. raniformis sites between Narrandera and Hay, wetlands along the lower Murrumbidgee and Murray including Darlington Point wetlands, and did not locate Rivers (NPWS, 2003). However, declines have been any individuals of this species. extensive from other parts of the Riverina biogeographic Litoria raniformis populations persist in the Coleambally region, in the north L. raniformis occurred in wetland Irrigation Area (CIA) south of the Murrumbidgee (AMBS systems off the Lachlan River. It was recorded from the Booligal wetlands in 1978 (Australian National Wildlife Consulting 2000), where it was first recorded in 1932. Given Collection) and Wooran near Booligal in 1980 (Australian that the CIA was not developed until the late 1960’s, it is Museum Collection). Litoria raniformis probably went difficult to determine whether these are founder populations locally extinct from wetlands around Booligal in the early from the Murrumbidgee floodplain or represent an expansion 1980’s after the ephemeral wetlands that were present of populations that may have existed along Coleambally after the 1974 floods dried up. It was also recorded near Creek (now the CIA outfall drain) and Buckingbong State Oxley in 1972 and Hill End in 1961 (Museum Victoria) Forest (Caughley and Gall 1985). Australian October 2008 Zoologist volume 34 (3) 449 Wassen Litoria raniformis persisted in the Murrumbidgee Irrigation reduced flooding frequency as result of flow regulation Area (MIA) immediately north of Murrumbidgee River (Briggs et al. 1997; Jansen and Robertson 2001). In in the Riverina biogeographic region until the early the former case, an outbreak of Chytrid is likely to 1980’s (National Museum Collection), but has since have played a significant role in local extinctions of disappeared (Wassens 2006). L. raniformis (Osborne et al. 1996). However, there is little evidence that chytrid contributed to declines Litoria raniformis occurred in River Red Gum in the inland regions of NSW because the disease is wetland systems on the Murray River in the Riverina much less likely to be present in areas with higher mean Biogeographic region, and was recorded in the Mulwala temperatures (Woodhams et al. 2003). and Boomanoomana State Forests in 1979 (Caughley and Gall 1985), and the Barmah-Millewa State forest Gambusia has been shown to prey heavily on L. aurea in 1959 and 1963 (Victoria Museum). The status of eggs and small tadpoles in tank experiments ( Morgan L. raniformis populations in these wetland systems is and Buttemer 1996; Pyke and White 2000).Predation uncertain as surveys have been very limited. However, of tadpoles by the introduced fish Gambusia (Gambusia recent frog surveys of the Barmah-Millewa Forest in holbrooki) has been implicated in the declines of a number 2003-2004 did not record L. raniformis (Ward 2004), of frog species in Australia, including L. raniformis (Morgan and the lack of any records since 1979 suggests that local and Buttemer 1996; Pyke and White 2000). However, in extinction may have also occurred in the Mulwala and natural environments viable L. raniformis populations Boomanoomana State Forests. In the extreme west of commonly persist with relatively high Gambusia densities the Riverina biogeographic region, L. raniformis persists (Osborne et al. 1996; Ashworth 1998; Heard et al. 2004). in wetlands near the confluence of the Murray and Rufus The apparent contradiction between tank experiments Rivers, possibly in Lake Victoria (NPWS). and field studies probably occurs because of the strong interaction between predation rates by Gambusia, aquatic Litoria raniformis populations also persist in the Wakool vegetation cover and wetland permanence. Periodic region, north of the Murray River within both irrigation drying reduces fish densities in wetlands (Baber et al. infrastructure and natural Black Box wetlands (Matthew 2002), while predation rates may increase with reduced Herring, Charles Sturt University, pers. com.). Formal aquatic vegetation cover (Lawler et al. 1999). Because frog surveys of the Murray River and associated wetlands of this, the impact of Gambusia needs to be considered have been limited; however, Matthew Herring conducted in conjunction with other factors, particularly wetland general fauna surveys at 94 wetlands in Northern degradation and hydrological changes. Victoria and the NSW Murray Catchments and failed to detect L. raniformis (Matthew Herring, Charles Sturt Limitations of the data University, pers. com.). In addition, the Heartlands Eastern Billabong Wildlife Survey conducted around The use of historical records to describe patterns of Henty in 2000-2001 also failed to detect L. raniformis declines has a number of limitations. Firstly, records (Matt Herring, Charles Sturt University, pers.com.) of L. raniformis in inland NSW are limited and have usually been conducted at a single point in time. In Murray-Darling depression many cases poor survey repetition makes it difficult Records from the Murray-Darling Depression are limited to accurately place the declines within a particular but this may be a result of poor survey effort; Litoria decade. In addition, some of the wetlands where L. raniformis remains locally abundant in the Lowbidgee raniformis once occurred have not been resurveyed. In region at the confluence of the Murray and Darling Rivers cases where L. raniformis populations were small, they and down stream along the Murray River into South could have been overlooked during surveys, as was the Australia (NPWS database). case for a L. aurea population south of Queanbeyan (Wassens and Mullins 2001). Conclusion A second problem with the use of historic records is Litoria raniformis formerly occurred across a large area, the lack of habitat data. The habitat requirements of L. occupying a diverse range of habitats from alpine raniformis populations in inland NSW outside of irrigation streams to ephemeral wetlands in the semi-arid zone areas have received little attention and it is unclear how (Pyke 2002).Given the widespread nature of declines populations might have responded to changes in wetland and the range of habitats affected, a number of different condition as a result of flow regulation and cattle grazing. threatening processes are likely to have contributed In addition, it is not clear when these habitat changes to declines in different areas. These differences are occurred in particular wetlands, whether they coincided especially evident when considering differences in with L. raniformis declines, and if they did, whether these declines between populations in the South Eastern changes were causal or coincidental to the declines. Highlands and riverine populations in the South Great care must be taken when interpreting the timing of West Slopes. Declines of the populations in the South decline, particularly on the basis of limited data, because Eastern Highlands did not coincide with significant the last recorded populations and extinctions are likely changes in wetland condition (Osborne et al. 1996). to represent the end point of a protracted decline period In contrast, former habitats within the South West marked by local extinctions of smaller populations (Hobbs Slopes have been highly modified, either through and Mooney 1998) rather then a single catastrophic event their conversion permanent water storages or through occurring over a short time frame. Australian 450 Zoologist volume 34 (3) October 2008 Past distribution and decline of the Southern Bell Frog Acknowledgements This study was funded by the CRC for Sustainable Rice in establishing survey techniques. We thank land Production and Murray-Darling Basin Commission. S. managers for giving us access to the study sites. This Wassens was supported by an Australian Postgraduate research was conducted under NSW National Parks Award. Robyn Watts, Amy Jansen and Alistar and Wildlife Service Licence (S3053). Ethics approval Robertson provided comments on early drafts of this was granted by Charles Sturt University Animal Care manuscript. Graham Pyke and Will Osborne assisted and Ethics Committee (01/034) References AMBS Consulting 2000. Biodiversity benchmarking survey of the Gillespie, G. R. 1996. Distribution, habitat and conservation Coleambally Irrigation Area. Unpublished report to Coleambally status of the Green and Golden Bell Frog Litoria aurea (Lesson Irrigation Corporation. 1829) (Anura: Hylidae) in Victoria. Australian Zoologist 30: 199-207 Ashworth, J.M. 1998. An appraisal of the conservation status of Litoria raniformis (Kefferstein) in Tasmania. 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