Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Invasional meltdown-under? Toads facilitate cats by removing a naïve top predator

J. Sean Doody https://orcid.org/0000-0002-9023-6206 A * , David Rhind B , Colin M. McHenry C and Simon Clulow D
+ Author Affiliations
- Author Affiliations

A Department of Biological Sciences, University of South Florida – St. Petersburg, St. Petersburg, FL 33701, USA.

B Department of Environment, Parks and Water Security, Northern Territory Government, P.O. Box 496, Palmerston, NT 0831, Australia.

C School of Environmental and Life Sciences, University of Newcastle, Callaghan, NSW 2308, Australia.

D Centre for Conservation Ecology and Genomics, Institute for Applied Ecology, University of Canberra, Bruce, ACT 2617, Australia.

* Correspondence to: jseandoody@gmail.com

Handling Editor: Jonathan Webb

Wildlife Research 51, WR22177 https://doi.org/10.1071/WR22177
Submitted: 25 October 2022  Accepted: 13 March 2023  Published: 14 April 2023

© 2024 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

Australia has been a battleground of invasive versus native species for ~200 years. Two of the most impactful invasive species to Australian animal communities are the toxic cane toad (Rhinella marina) and the predatory feral cat (Felis catus). Australia’s native fauna is evolutionarily naïve to both invaders because neither’s taxonomic group is native to the continent. Both invaders have had severe effects on Australian native animal communities including species extinctions, extirpations, and severe population declines, but until now their effects have generally been thought to be independent of one another.

Aims

We aimed to determine the impacts of invasive cane toads on monitor lizards and feral cats by estimating changes in relative abundance before and after the toad invasion.

Methods

We studied toad impacts at three sites in the Kimberley region of northern Australia. We used two methods for estimating relative abundance: camera traps and track station data. Data included greater than 4000 trap days and included 7 years over an 11-year period.

Key results

As expected, invading cane toads rapidly decimated populations of two species of monitor lizards (97–99% declines), including the top-order predatory Varanus panoptes. Unexpectedly, this loss was associated with a >10-fold mean increase in detection rates of cats by 5 years after the loss of V. panoptes, reflecting relative increases of 3.3–8.7 individual cats per site.

Conclusions

Although some unknown factor may have caused an increase in cats, their similar trophic position and niche to V. panoptes suggests that toads facilitated cats by effectively removing the lizards from the animal community. This interaction likely reflects one type of invasional meltdown, whereby a non-native species (cane toad) facilitated any aspect of another’s (feral cat) invasion (e.g. survival, reproduction, resource acquisition), but the latter has no detected influence on the former (+/0 interaction).

Implications

Because both invaders cause declines in animal populations and are difficult to control, the potentially synergistic tandem of cane toads and feral cats could have chronic, irreversible effects on animal communities.

Keywords: cane toad, exploitation competition, Felis catus, invasive species, lethal toxic ingestion, mesopredator release, predator–prey interactions, Rhinella marina.

References

Abbott I (2008) The spread of the cat, Felis catus, in Australia: re-examination of the current conceptual model with additional information. Conservation Science Western Australia 7, 1-17.
| Google Scholar |

Allen BL, Allen LR, Leung LK-P (2015) Interactions between two naturalised invasive predators in Australia: are feral cats suppressed by dingoes? Biological Invasions 17, 761-776.
| Crossref | Google Scholar |

Australian Bureau of Meteorology (2019) Climate statistics, 2009–2019. Australian Government.

Bellard C, Genovesi P, Jeschke JM (2016) Global patterns in threats to vertebrates by biological invasions. Proceedings of the Royal Society B: Biological Sciences 283, 20152454.
| Crossref | Google Scholar |

Blamires SJ (2004) Habitat preferences of coastal goannas (Varanus panoptes): are they exploiters of sea turtle nests at Fog Bay, Australia? Copeia 2004, 370-377.
| Crossref | Google Scholar |

Braga RR, Gómez-Aparicio L, Heger T, Vitule JRS, Jeschke JM (2018) Structuring evidence for invasional meltdown: broad support but with biases and gaps. Biological Invasions 20, 923-936.
| Crossref | Google Scholar |

Burrows ND, Algar D, Robinson AD, Sinagra J, Ward B, Liddelow G (2003) Controlling introduced predators in the Gibson Desert of Western Australia. Journal of Arid Environments 55, 691-713.
| Crossref | Google Scholar |

Carthey AJ, Banks PB (2014) Naiveté in novel ecological interactions: lessons from theory and experimental evidence. Biological Reviews 89, 932-949.
| Crossref | Google Scholar |

Case TJ, Bolger DT, Richman AD (1992) Reptilian extinctions: the last ten thousand years. In ‘Conservation biology: the theory and practice of nature conservation preservation and management’. (Eds PL Fiedler, SK Jain) pp. 91–125. (Chapman and Hall: New York, NY, USA)

Christian KA, Corbett LK, Green B, Weavers BW (1995) Seasonal activity and energetics of two species of varanid lizards in tropical Australia. Oecologia 103, 349-357.
| Crossref | Google Scholar |

Christian K (2004) Varanus panoptes. In ‘Varanoid lizards of the world’. (Eds E Pianka, D King) pp. 423–429. (Indiana University Press, Bloomington, IN, USA)

Clavero M, García-Berthou E (2005) Invasive species are a leading cause of animal extinctions. Trends in Ecology & Evolution 20, 110.
| Crossref | Google Scholar |

Cox JG, Lima SL (2006) Naiveté and an aquatic-terrestrial dichotomy in the effects of introduced predators. Trends in Ecology & Ecology 21, 674-680.
| Crossref | Google Scholar |

Croll DA, Maron JL, Estes JA, Danner EM, Byrd GV (2005) Introduced predators transform subarctic islands from grassland to tundra. Science 307, 1959-1961.
| Crossref | Google Scholar |

Crooks KR, Soulé ME (1999) Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400, 563-566.
| Crossref | Google Scholar |

Davies HF, McCarthy MA, Firth RSC, Woinarski JCZ, Gillespie GR, Andersen AN, Geyle HM, Nicholson E, Murphy BP (2017) Top-Down control of species distributions: Feral cats driving the regional extinction of a threatened rodent in northern Australia. Diversity and Distributions 23, 272-283.
| Crossref | Google Scholar |

Denny EA, Dickman CR (2010) Review of cat ecology and management strategies in Australia. Invasive Animal Cooperative Research Centre, Canberra.

Diamond JM (1989) Quaternary megafaunal extinctions: variations on a theme by paganini. Journal of Archaeological Science 16, 167-175.
| Crossref | Google Scholar |

Diamond J, Case TJ (1986) Overview: introductions, extinctions, exterminations, and invasions. In ‘Community ecology’. (Eds J Diamond, TJ Case) pp. 65–79. (Harper & Row: New York, NY, USA)

Dickman CR (1996) ‘Overview of the impacts of feral cats on Australian native fauna.’ (Australian Nature Conservation Agency: Canberra, ACT, Australia)

Doherty TS, Davis RA, van Etten EJB, Algar DA, Collier N, Dickman CR (2015) A continental-scale analysis of feral cat diet in Australia. Journal of Biogeography 42, 964-975.
| Crossref | Google Scholar |

Doherty TS, Glen AS, Nimmo DG, Ritchie EG, Dickman CR (2016) Invasive predators and global biodiversity loss. Proceedings of the National Academy of Sciences of the United States of America 113, 11261-11265.
| Crossref | Google Scholar |

Doherty TS, Dickman CR, Johnson CN, Legge SM, Ritchie EG, Woinarski JCZ (2017) Impacts and management of feral cats Felis catus in Australia. Mammal Review 47, 83-97.
| Crossref | Google Scholar |

Donadio E, Buskirk SW (2006) Diet, morphology, and interspecific killing in carnivora. The American Naturalist 167, 524-536.
| Crossref | Google Scholar |

Doody JS, Green B, Sims R, Rhind D, West P, Steer D (2006) Indirect impacts of invasive cane toads (Bufo marinus) on nest predation in pig-nosed turtles (Carettochelys insculpta). Wildlife Research 33, 349-354.
| Crossref | Google Scholar |

Doody JS, Green B, Rhind D, Castellano CM, Sims R, Robinson T (2009) Population-level declines in Australian predators caused by an invasive species. Animal Conservation 12, 46-53.
| Crossref | Google Scholar |

Doody JS, Hall M, Rhind D, Green B, Dryden G (2012a) Varanus panoptes (Yellow-spotted monitor) Diet. Herpetological Review 43, 491-492.
| Google Scholar |

Doody JS, Lloyd R, Rhind D (2012b) Varanus panoptes (Yellow-spotted monitor) Diet and prey capture. Herpetological Review 43, 339-340.
| Google Scholar |

Doody JS, Castellano CM, Rhind D, Green B (2013) Indirect facilitation of a native mesopredator by an invasive species: are cane toads re-shaping tropical riparian communities? Biological Invasions 15, 559-568.
| Crossref | Google Scholar |

Doody JS, Mayes P, Clulow S, Rhind D, Green B, Castellano CM, D’Amore D, McHenry C (2014) Impacts of the invasive cane toad on aquatic reptiles in a highly modified ecosystem: the importance of replicating impact studies. Biological Invasions 16, 2303-2309.
| Crossref | Google Scholar |

Doody JS, Soanes R, Castellano CM, Rhind D, Green B, McHenry CR, Clulow S (2015a) Invasive toads shift predator–prey densities in animal communities by removing top predators. Ecology 96, 2544-2554.
| Crossref | Google Scholar |

Doody JS, James H, Colyvas K, Mchenry CR, Clulow S (2015b) Deep nesting in a lizard déjà vu devils’ corkscrews: first helical reptile burrow and deepest vertebrate nest. Biological Journal of the Linnean Society 116, 13-26.
| Crossref | Google Scholar |

Doody JS, McHenry C, Mahony S, Taylor S, Clulow S (2015c) Varanus panoptes (Yellow-spotted Monitor). Diet. Herpetological Review 46, 263-264.
| Google Scholar |

Doody JS, Sawyer G, Hands K (2015d) Varanus panoptes (Yellow-spotted Monitor). Diet and foraging behavior. Herpetological Review 46, 95-96.
| Google Scholar |

Doody JS, Rhind D, Green B, Castellano C, McHenry C, Clulow S (2017) Chronic effects of an invasive species on an animal community. Ecology 98, 2093-2101.
| Crossref | Google Scholar |

Doody JS, McHenry C, Letnic M, Everitt C, Sawyer G, Clulow S (2018) Forecasting the spatiotemporal pattern of the cane toad invasion into north-western Australia. Wildlife Research 45, 718-725.
| Crossref | Google Scholar |

Doody JS, Soennichsen KF, James H, McHenry C, Clulow S (2021a) Ecosystem engineering by deep-nesting monitor lizards. Ecology 102, e03271.
| Crossref | Google Scholar |

Doody JS, McHenry C, Rhind D, Gray C, Clulow S (2021b) Impacts of invasive cane toads on an Endangered marsupial predator and its prey. Endangered Species Research 2021(46), 269-277.
| Crossref | Google Scholar |

Elton CS (1958) ‘The ecology of invasion by plants and animals.’ (Springer: Methuen, London)

Feit B, Gordon CE, Webb JK, Jessop TS, Laffan SW, Dempster T, Letnic M (2018) Invasive cane toads might initiate cascades of direct and indirect effects in a terrestrial ecosystem. Biological Invasions 20, 1833-1847.
| Crossref | Google Scholar |

Fisher DO, Johnson CN, Lawes MJ, Fritz SA, McCallum H, Blomberg SP, VanDerWal J, Abbott B, Frank A, Legge S, Letnic M, Thomas CR, Fisher A, Gordon IJ, Kutt A (2014) The current decline of tropical marsupials in Australia: is history repeating? Global Ecology and Biogeography 23, 181-190.
| Crossref | Google Scholar |

Flannery T (1994) ‘The future eaters: an ecological history of the Australasian lands and people.’ (New Holland: Sydney)

Frank ASK, Johnson CN, Potts JM, Fisher A, Lawes MJ, Woinarski JCZ, Tuft K, Radford IJ, Gordon IJ, Collis M-A, Legge S (2014) Experimental evidence that feral cats cause local extirpation of small mammals in Australia’s tropical savannas. Journal of Applied Ecology 51, 1486-1493.
| Crossref | Google Scholar |

Gurevitch J, Padilla DK (2004) Are invasive species a major cause of extinctions? Trends in Ecology & Evolution 19((9)), 470-474.
| Crossref | Google Scholar |

Hardman B, Moro D, Calver M (2016) Direct evidence implicates feral cat predation as the primary cause of failure of a mammal reintroduction programme. Ecological Management & Restoration 17, 152-158.
| Crossref | Google Scholar |

Hebblewhite M, White CA, Nietvelt CG, McKenzie JA, Hurd TE, Fryxell JM, Bayley SE, Paquet PC (2005) Human activity mediates a trophic cascade caused by wolves. Ecology 86, 2135-2144.
| Crossref | Google Scholar |

Hoffmann BD, Broadhurst LM (2016) The economic cost of managing invasive species in Australia. NeoBiota 31, 1-18.
| Crossref | Google Scholar |

James AI, Eldridge DJ, Hill BM (2009) Foraging animals create fertile patches in an Australian desert shrubland. Ecography 32, 723-732.
| Crossref | Google Scholar |

Johnson CN (2006) ‘Australia’s Mammal Extinctions: A 50,000 Year History.’ (Cambridge University Press)

Kutt AS (2012) Feral cat (Felis catus) prey size and selectivity in north-eastern Australia: implications for mammal conservation. Journal of Zoology (London) 287, 292-300.
| Crossref | Google Scholar |

Legge S, Woinarski JCZ, Dickman CR, Doherty TS, McGregor H, Murphy BP (2020) Cat ecology, impacts and management in Australia. Wildlife Research 47, i-vi.
| Crossref | Google Scholar |

Letnic M, Dickman CR (2006) Boom means bust: interactions between the El Niño/Southern Oscillation (ENSO), rainfall and the processes threatening mammal species in arid Australia. Biodiversity & Conservation 15, 3847-3880.
| Crossref | Google Scholar |

Letnic M, Webb JK, Shine R (2008) Invasive cane toads (Bufo marinus) cause mass mortality of freshwater crocodiles (Crocodylus johnstoni) in tropical Australia. Biological Conservation 141, 1773-1782.
| Crossref | Google Scholar |

Lever C (2001) ‘The cane toad: the history and ecology of a successful colonist’. (Westbury Academic and Scientific Publishing: Yorkshire)

Losos JB, Greene HW (1988) Ecological and evolutionary implications of diet in monitor lizards. Biological Journal of the Linnean Society 35, 379-407.
| Crossref | Google Scholar |

Losos JB, Ricklefs RE (2010) ‘The theory of island biogeography revisited.’ (Princeton University Press: Princeton, NJ)

Loss SR, Marra PP (2017) Population impacts of free-ranging domestic cats on mainland vertebrates. Frontiers in Ecology and the Environment 15, 502-509.
| Crossref | Google Scholar |

Loss SR, Will T, Marra PP (2013) The impact of free-ranging domestic cats on wildlife of the United States. Nature Communications 4, 1396.
| Crossref | Google Scholar |

Low T (1999) ‘Feral future: the untold story of Australia’s exotic invaders.’ (Viking: Australia)

Madsen T, Ujvari B, Shine R, Olsson M (2006) Rain, rats and pythons: climate-driven population dynamics of predators and prey in tropical Australia. Austral Ecology 31, 30-37.
| Crossref | Google Scholar |

McLeod R (2004) Counting the cost: impact of invasive animals in Australia 2004. Cooperative Research Centre for Pest Animal Control, Canberra. p. 82.

Medina FM, Bonnaud E, Vidal E, Tershy BR, Zavaleta ES, Josh Donlan C, Keitt BS, Le Corre M, Horwath SV, Nogales M (2011) A global review of the impacts of invasive cats on island endangered vertebrates. Global Change Biology 17, 3503-3510.
| Crossref | Google Scholar |

Medway DG (2004) The land bird fauna of Stephens Island, New Zealand in the early 1890s, and the cause of its demise. Notornis 51, 201-211.
| Google Scholar |

Murphy BP, Woolley L-A, Geyle HM, Legge SM, Palmer R, Dickman CR, Augusteyn J, Brown SC, Comer S, Doherty TS, Eager C, Edwards G, Fordham DA, Harley D, McDonald PJ, McGregor H, Moseby KE, Myers C, Read J, Riley J, Stokeld D, Trewella GJ, Turpin JM, Woinarski JCZ (2019) Introduced cats (Felis catus) eating a continental fauna: the number of mammals killed in Australia. Biological Conservation 237, 28-40.
| Crossref | Google Scholar |

Oakwood M (2004) The effect of cane toads on a marsupial carnivore, the northern quoll, Dasyurus hallucatus. Unpublished Report to Department of Environment and Heritage, Darwin.

Phillips BL, Greenlees MJ, Brown GP, Shine R (2010) Predator behaviour and morphology mediates the impact of an invasive species: cane toads and death adders in Australia. Animal Conservation 13, 53-59.
| Crossref | Google Scholar |

Pimentel D, Lach L, Zuniga R, Morrison D (2000) Environmental and economic costs of nonindigenous species in the United States. BioScience 50, 53-65.
| Crossref | Google Scholar |

Pimentel D, McNair S, Janecka J, et al. (2001) Economic and environmental threats of alien plant, animal, and microbe invasions. Agriculture, Ecosystems & Environment 84, 1-20.
| Crossref | Google Scholar |

Pinch K, Madsen T, Ujvari B (2017) No signs of Na+/K+-ATPase adaptations to an invasive exotic toxic prey in native squamate predators. Austral Ecology 42, 929-933.
| Crossref | Google Scholar |

Price-Rees SJ, Brown GP, Shine R (2010) Predation on toxic cane toads (Bufo marinus) may imperil bluetongue lizards (Tiliqua scincoides intermedia, Scincidae) in tropical Australia. Wildlife Research 37, 166-173.
| Crossref | Google Scholar |

Pyšek P, Hulme PE, Simberloff D, Bacher S, Blackburn TM, Carlton JT, Dawson W, Essl F, Foxcroft LC, Genovesi P, Jeschke JM, Kühn I, Liebhold AM, Mandrak NE, Meyerson LA, Pauchard A, Pergl J, Roy HE, Seebens H, van Kleunen M, Vilà M, Wingfield MJ, Richardson DM (2020) Scientists’ warning on invasive alien species. Biological Reviews 95, 1511-1534.
| Crossref | Google Scholar |

Radford IJ, Woolley L-A, Dickman CR, Corey B, Trembath D, Fairman R (2020) Invasive anuran driven trophic cascade: an alternative hypothesis for recent critical weight range mammal collapses across northern Australia. Biological Invasions 22, 1967-1982.
| Crossref | Google Scholar |

Rhind D, Doody JS, Pritchard R, McHenry C (2013) Varanus panoptes (Yellow-spotted monitor) Cannibalism. Herpetological Review 44, 687-688.
| Google Scholar |

Rhind D, Sawyer G, Trembath D, Parrott M, Doody JS (2014) Varanus panoptes (Yellow-spotted monitor). Diet and behavior. Herpetological Review 45, 335-336.
| Google Scholar |

Sala OE, Stuart Chappin F, III, Armesto JJ, Berlow E, Bloomfield J, Dirzo R, Huber-Sanwald E, Huenneke LF, Jackson RB, Kinzig A, Leemans R, Lodge DM, Mooney HA, Oesterheld M, Poff NL, Sykes MT, Walker BH, Walker M, Wall DH (2000) Global biodiversity scenarios for the year 2100. Science 287, 1770-1774.
| Crossref | Google Scholar |

Salo P, Korpimäki E, Banks PB, Nordström M, Dickman CR (2007) Alien predators are more dangerous than native predators to prey populations. Proceedings of the Royal Society B: Biological Sciences 274, 1237-1243.
| Crossref | Google Scholar |

Saunders DA (2006) Cats and cockatoo chicks. In ‘The state of Australia’s birds 2006: invasive species’. (Eds P Olsen, A Silcocks, M Weston). p. 13. (Birds Australia: Hawthorn East, Victoria)

Shanmuganathan T, Pallister J, Doody JS, McCallum H, Robinson T, Sheppard A (2010) Biological control of the cane toad in Australia: a review. Animal Conservation 13, 16-23.
| Crossref | Google Scholar |

Shannon R, Mendyk RW (2009) Aquatic foraging behavior and freshwater mussel (Velesunio sp.) predation by Varanus panoptes panoptes in Central-Western Queensland. Biawak 3, 85-87.
| Google Scholar |

Shine R (1986) Food habits, habitats and reproductive biology of four sympatric species of varanid lizards in tropical Australia. Herpetologica 42, 346-360.
| Google Scholar |

Shine R (2010) The ecological impact of invasive cane toads (Bufo marinus) in Australia. The Quarterly Review of Biology 85, 253-291.
| Crossref | Google Scholar |

Shine R, Doody JS (2011) Invasive species control: understanding conflicts between researchers and the general community. Frontiers in Ecology and the Environment 9, 400-406.
| Crossref | Google Scholar |

Short J (2016) Predation by feral cats key to the failure of a long-term reintroduction of the western barred bandicoot (Perameles bougainville). Wildlife Research 43, 38-50.
| Crossref | Google Scholar |

Sih A, Bolnick DI, Luttbeg B, Orrock JL, Peacor SD, Pintor LM, Preisser E, Rehage JS, Vonesh JR (2010) Predator-Prey naivete, antipredator behavior, and the ecology of predator invasions. Oikos 119, 610-621.
| Crossref | Google Scholar |

Simberloff D, Rejmanek M (2011) ‘Encyclopedia of biological invasions.’ (University of California Press)

Simberloff D, Von Holle B (1999) Synergistic interactions of nonindigenous species: invasional meltdown? Biological Invasions 1, 21-32.
| Crossref | Google Scholar |

Spencer EE, Newsome TM, Dickman CR (2017) Prey selection and dietary flexibility of three species of mammalian predator during an irruption of non-cyclic prey. Royal Society Open Science 4, 170317.
| Crossref | Google Scholar |

Stobo-Wilson AM, Murphy BP, Cremona T, Carthew SM (2019) Contrasting patterns of decline in two arboreal marsupials from Northern Australia. Biodiversity and Conservation 28, 2951-2965.
| Crossref | Google Scholar |

Sutherland DR, Glen AS, de Tores PJ (2011) Could controlling mammalian carnivores lead to mesopredator release of carnivorous reptiles? Proceedings of the Royal Society B: Biological Sciences 278, 641-648.
| Crossref | Google Scholar |

Sweet SS, Pianka ER (2007) Monitors, mammals, and Wallace’s line. Mertensiella 16, 79-99.
| Google Scholar |

Tingley R, Ward-Fear G, Schwarzkopf L, Greenlees MJ, Phillips BL, Brown G, Clulow S, Webb J, Capon R, Sheppard A, Strive T, Tizard M, Shine R (2017) New weapons in the toad toolkit: a review of methods to control and mitigate the biodiversity impacts of invasive cane toads (Rhinella marina). The Quarterly Review of Biology 92, 123-149.
| Crossref | Google Scholar |

Ujvari B, Madsen T (2009) Increased mortality of naïve varanid lizards after the invasion of non-native cane toads (Bufo marinus). Herpetological Conservation and Biology 4, 248-251.
| Google Scholar |

Ujvari B, Mun H-C, Conigrave AD, Bray A, Osterkamp J, Halling P, Madsen T (2013) Isolation breeds naivety: island living robs Australian varanid lizards of toad-toxin immunity via four-base-pair mutation. Evolution 67, 289-294.
| Crossref | Google Scholar |

Ujvari B, Casewell NR, Sunagar K, Arbuckle K, Wuster W, Lo N, O’Meally D, Beckmann C, King GF, Deplazes E, Madsen T (2015) Widespread convergence in toxin resistance by predictable molecular evolution. Proceedings of the National Academy of Sciences of the United States of America 112, 11911-11916.
| Crossref | Google Scholar |

Ward S (2012) Threatened species of the Northern territory. Mitchells’ water monitor. Varanus mitchelli. Northern Territory Government, Darwin, Australia.

Ward S, Woinarski J, Griffiths T, McKay JL (2006) Threatened species of the Northern territory. Mertens’ water monitor. Varanus mertensi. Northern Territory Government, Darwin, Australia.

Ward S, Woinarski J, Griffiths AD, McKay JL (2012) Threatened species of the Northern Territory. Yellow-spotted monitor. Varanus panoptes. Northern Territory Government, Darwin, Australia.

Weavers B (1989) Diet of the lace monitor lizard (Varanus vadus) in south-eastern Australia. Australian Zoologist 25, 83-85.
| Crossref | Google Scholar |

Webb GJW, Manolis SC (2010) Australian Freshwater Crocodile Crocodylus johnstoni. Pp. 66–70 in Crocodiles. Status Survey and Conservation Action Plan. 3rd edn, Ed. by S.C. Manolis and C. Stevenson. Crocodile Specialist Group: Darwin.

Whitford WG (1998) Contribution of pits dug by goannas (Varanus gouldii) to the dynamics of banded mulga landscapes in eastern Australia. Journal of Arid Environments 40, 453-457.
| Crossref | Google Scholar |

Wilson EO (1989) Threats to biodiversity. Scientific American 261, 108-116.
| Crossref | Google Scholar |

Woinarski JCZ, Oakwood M, Winter J, Burnett S, Milne D, Foster P, Myles H, Holmes B (2008) Surviving the toads: patterns of persistence of the northern quoll Dasyurus hallucatus in Queensland. Report to The Australian Government’s Natural Heritage Trust.

Woinarski JCZ, Legge S, Fitzsimons JA, Traill BJ, Burbidge AA, Fisher A, Firth RSC, Gordon IJ, Griffiths AD, Johnson CN, McKenzie NL, Palmer C, Radford I, Rankmore B, Ritchie EG, Ward S, Ziembicki M (2011) The disappearing mammal fauna of northern Australia: context, cause, and response. Conservation Letters 4, 192-201.
| Crossref | Google Scholar |

Woinarski JCZ, Murphy BP, Legge SM, Garnett ST, Lawes MJ, Comer S, Dickman CR, Doherty TS, Edwards G, Nankivell A, Paton D, Palmer R, Woolley LA (2017) How many birds are killed by cats in Australia? Biological Conservation 214, 76-87.
| Crossref | Google Scholar |

Woinarski JCZ, Murphy BP, Palmer R, Legge SM, Dickman CR, Doherty TS, Edwards G, Nankivell A, Read JL, Stokeld D (2018) How many reptiles are killed by cats in Australia? Wildlife Research 45, 247-266.
| Crossref | Google Scholar |

Woinarski JCZ, Legge SM, Dickman CR (2019) ‘Cats in Australia: companion and killer.’ (CSIRO Publishing: Melbourne)

Woinarski JCZ, Legge SM, Woolley LA, Palmer R, Dickman CR, Augusteyn J, Doherty TS, Edwards G, Geyle H, McGregor H, Riley J, Turpin J, Murphy BP (2020) Predation by introduced cats Felis catus on Australian frogs: compilation of species records and estimation of numbers killed. Wildlife Research 47, 580-588.
| Crossref | Google Scholar |

Yip SJS, Rich M-A, Dickman CR (2015) Diet of the feral cat, Felis catus, in central Australian grassland habitats during population cycles of its principal prey. Mammal Research 60, 39-50.
| Crossref | Google Scholar |