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Pests are often viewed as having consistently strong and negative effects on biodiversity values and agricultural productivity, especially if they have been introduced from elsewhere. However, pests do not necessarily have just negative impacts, and there is evidence that in some situations their effects can be beneficial. The positive effects of pests arise when they become deeply embedded in ecological communities and are involved in webs of direct and indirect interactions with other species. In this paper, I first outline the concepts of direct and indirect interactions, and then describe two case studies that illustrate how these interactions develop between pests and native animals. In the first case study, house mice (Mus domesticus) introduced to Boullanger Island in Western Australia have direct effects on a small dasyurid marsupial (Sminthopsis griseoventer boullangerensis), but also exert indirect negative and indirect positive effects on four species of insular skinks. In the second case study, high levels of activity of domestic house cats (Felis catus) in suburban bushland in Sydney are associated with reduced richness of bird species. However, high cat activity also appears to depress the rate of egg predation in above-ground bird nests, apparently by suppressing the activities of small egg predators such as rats. In these and other examples, control of the putative pest would have unintended negative consequences on some native species, making decisions about management more difficult. I propose a preliminary framework to predict the likelihood of indirect interactions occurring at different times after a pest has been introduced, at different places, and at different pest densities, and use this to suggest options for management of the pest species.

Arthur, W. and Mitchell, P. 1989. A revised scheme for the classification of population interactions. Oikos 56: 141-143.
Baldock, F. C., Alexander, L. and More, S. J. 2003. Estimated and predicted changes in the cat population of Australian households from 1979 to 2005. Australian Veterinary Journal 81: 289-292.
Banks, P. B., Dickman, C. R. and Newsome, A. E. 1998. Ecological costs of feral predator control: foxes and rabbits. Journal of Wildlife Management 62: 766-772.
Barratt, D. G. 1998. Predation by house cats, Felis catus (L.), in Canberra, Australia. II. Factors affecting the amount of prey caught and estimates of the impact on wildlife. Wildlife Research 25: 475-487.
Bencini, R., McCulloch, C., Mills, H. R. and Start, A. N. 2001. Habitat and diet of the dibbler ( Parantechinus apicalis) on two islands in Jurien Bay, Western Australia. Wildlife Research 28: 465-468.
Braysher, M. 1993 Managing Vertebrate Pests: Principles and Strategies. Australian Government Publishing Service: Canberra.
Burrows, N. D., Algar, D., Robinson, A. D., Sinagra, J., Ward, B. and Liddelow, G. 2003. Controlling introduced predators in the Gibson Desert of Western Australia. Journal of Arid Environments 55: 691-713.
Caughley, G. and Gunn, A. 1996 Conservation Biology in Theory and Practice. Blackwell Science: Oxford.
Caughley, J., Monamy, V. and Heiden, K. 1994 Impact of the 1993 Mouse Plague. Occasional Paper Series No. 7, Grains Research and Development Corporation: Canberra.
Chapple, R. 2005. The politics of feral horse management in Guy Fawkes River National Park, NSW. Australian Zoologist 33: 233-246.
Courchamp, F., Langlais, M. and Sugihara, G. 1999. Cats protecting birds: modelling the mesopredator release effect. Journal of Animal Ecology 68: 282-292.
Crooks, K. R. and Soulé, M. E. 1999. Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400: 563-566.
Crowther, M. S., Dickman, C. R. and Lynam, A. J. 1999 Sminthopsis griseoventer boullangerensis (Marsupialia: Dasyuridae), a new subspecies in the S. murina complex from Boullanger Island, Western Australia. Australian Journal of Zoology 47: 215-243.
CSIRO. 1991 The Insects of Australia, 2nd edition. Melbourne University Press: Melbourne.
Dickman, C. R. 1988. Body size, prey size, and community structure in insectivorous mammals. Ecology 69: 569-580.
Dickman, C. R. 1992. Conservation of mammals in the Australasian region: the importance of islands. Pp. 175-214 in Australia and the Global Environmental Crisis, edited by J. N. Coles and J. M. Drew. Academy Press: Canberra.
Dickman, C. R. 1996 Overview of the Impacts of Feral Cats on Australian Native Fauna. Australian Nature Conservation Agency: Canberra.
Dickman, C. R. 1999. Rodent-ecosystem relationships: a review. Pp. 113-133 in Ecologically-based Management of Rodent Pests, edited by G. R. Singleton, L. A. Hinds, H. Leirs and Z. Zhang. Australian Centre for International Agricultural Research: Canberra.
Dickman, C. R. 2006a. Species interactions: direct effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Dickman, C. R. 2006b. Species interactions: indirect effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Dickman, C. R. 2006c. House cats in the Australian environment: behavior, impacts, and management. Berryman Bulletin.
Dickman, C. R. and Lunney, D. (eds) 2001 A Symposium on the Dingo. Royal Zoological Society of New South Wales: Mosman.
Dickman, C. R. and Murray, B. R. 2006. Species interactions: complex effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Disney, H. J. de S. and Smithers, C. N. 1972. The distribution of terrestrial and freshwater birds on Lord Howe Island, in comparison with Norfolk Island. Australian Zoologist 17: 1-11.
Ewel, J. J. and Putz, F. E. 2005. A place for alien species in ecosystem restoration. Frontiers in Ecology and the Environment 2: 354-360.
Fitzgerald, B. M. 1988. Diet of domestic cats and their impact on prey populations. Pp. 123-147 in The Domestic Cat: the Biology of its Behaviour, edited by D. C. Turner and P. Bateson. Cambridge University Press: Cambridge.
Fitzgerald, B. M., Karl, B. J. and Veitch, C. R. 1991. The diet of feral cats ( Felis catus) on Raoul Island, Kermadec group. New Zealand Journal of Ecology 15: 123-129.
Flecker, A. S. and Townsend, C. R. 1994. Community-wide consequences of trout introduction in New Zealand streams. Ecological Applications 4: 798-807.
Fuller, P. J. and Burbidge, A. A. 1987. Discovery of the dibbler, Parantechinus apicalis, on islands at Jurien Bay. Western Australian Naturalist 16: 177-181.
Gentle, M. N. 2005. Factors affecting the efficiency of fox ( Vulpes vulpes) baiting practices on the Central Tablelands of New South Wales. PhD thesis, University of Sydney: Sydney.
Gillies, C. and Clout, M. 2003. The prey of domestic cats ( Felis catus) in two suburbs of Auckland City, New Zealand. Journal of Zoology 259: 309-315.
Glen, A. S. and Dickman, C. R. 2005. Complex interactions among mammalian carnivores in Australia, and their implications for wildlife management. Biological Reviews 80: 387-401.
Grayson, J. and Calver, M. 2004. Regulation of domestic cat ownership to protect urban wildlife: a justification based on the precautionary principle. Pp. 169-178 in Urban Wildlife: More than meets the Eye, edited by D. Lunney and S. Burgin. Royal Zoological Society of New South Wales: Mosman.
Grayson, J., Calver, M. C. and Styles, I. 2002. Attitudes of suburban Western Australians to proposed cat control legislation. Australian Veterinary Journal 80: 536-543.
Grigg, G. 2002. Conservation benefit from harvesting kangaroos: status report at the start of a new millennium. A paper to stimulate discussion and research. Pp. 53-76 in A Zoological Revolution: Using Native Fauna to Assist in its own Survival, edited by D. Lunney and C. R. Dickman. Royal Zoological Society of New South Wales: Mosman.
Kuchling, G. 1997. Managing the last survivors: integration of in situ and ex situ conservation of Pseudemydura umbrina. Pp. 339-344 in Proceedings: Conservation, Management and Restoration of Tortoises and Turtles, 11-16 July 1993, edited by J. van Abbema. New York Turtle and Tortoise Society: New York.
Kuchling, G. 2000. Conservation strategies for remnant turtle populations: the Western Australian swamp turtle Pseudemydura umbrina recovery programme. Stapfia 69, Zugleich Kataloge des O. Ö. Landesmuseums N.F. 149: 179-188.
Lilith, M., Calver, M., Styles, I. and Garkaklis M. 2006. Protecting wildlife from predation by owned domestic cats: application of a precautionary approach to the acceptability of proposed cat regulations. Austral Ecology 31: 176-189.
Low, T. 1999 Feral Future. Penguin Books: Melbourne.
Low, T. 2004 The New Nature. Penguin Books: Melbourne.
MacDougall, A. S. and Turkington, R. 2005. Are invasive species the drivers or passengers of change in degraded ecosystems? Ecology 86: 42-55.
Matthews, A., Dickman, C. R. and Major, R. E. 1999. The influence of fragment size and edge on nest predation in urban bushland. Ecography 22: 349-356.
McLeod, R. 2004 Counting the Cost: Impact of Invasive Animals in Australia, 2004. Cooperative Research Centre for Pest Animal Control: Canberra.
Meek, P. D. 1998. Food items brought home by domestic cats Felis catus (L.) living in Booderee National Park, Jervis Bay. Proceedings of the Linnean Society of New South Wales 120: 43-47.
Miller, S., Bencini, R., Mills, H. and Moro, D. 2003. Food availability for the dibbler ( Parantechinus apicalis) on Boullanger and Whitlock Islands, Western Australia. Wildlife Research 30: 649-654.
Molsher, R. 1999. The ecology of feral cats ( Felis catus) in open forest in New South Wales: interactions with food resources and foxes. PhD thesis, University of Sydney: Sydney
Morin, P. J. 1999 Community Ecology. Blackwell Science: Oxford.
Newsome, A. 1990. The control of vertebrate pests by vertebrate predators. Trends in Ecology and Evolution 5: 187-191.
Newsome, A., Pech, R., Smyth, R., Banks, P. and Dickman, C. 1997 Potential Impacts on Australian Native Fauna of Rabbit Calicivirus Disease. Environment Australia: Canberra.
NSW National Parks and Wildlife Service. 2002 Draft Wild Horse Management Plan for the Alpine Area of Kosciuszko National Park. NSW National Parks and Wildlife Service: Sydney.
Olsen, P. 1998 Australia's Pest Animals: New Solutions to Old Problems. Kangaroo Press: Sydney.
Pearson, D. E. and Callaway, R. M. 2003. Indirect effects of host-specific biological control agents. Trends in Ecology and Evolution 18: 456-461.
Proulx, S. R., Promislow, D. E. L. and Phillips, P. C. 2005. Network thinking in ecology and evolution. Trends in Ecology and Evolution 20: 345-353.
Reark 1994 The Metropolitan Domestic Cat: a Survey of the Population Characteristics and Hunting Behaviour of the Domestic Cat in Australia. Petcare Information and Advisory Service: Melbourne.
Risbey, D. A., Calver, M. C., Short, J., Bradley, J. S. and Wright, I. W. 2000. The impact of cats and foxes on the small vertebrate fauna of Heirisson Prong, Western Australia. II. A field experiment. Wildlife Research 27: 223-235.
Sinclair, A. R. E., Krebs, C. J., Fryxell, J. M., Turkington, R., Boutin, S., Boonstra, R., Seccombe-Hett, P., Lundberg, P. and Oksanen, L. 2000. Testing hypotheses of trophic level interactions: a boreal forest ecosystem. Oikos 89: 313-328.
Soulé, M. E., Bolger, D. T., Alberts, A. C., Wright, J., Sorice, M. and Hill, S. 1988. Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. Conservation Biology 2: 75-92.
Start, A. N., Moro, D., Adams, M. and Bencini, R. 2006. Dunnarts from Boullanger Island: new evidence and reassessment of a taxonomic issue with resource implications. Australian Mammalogy 28: 51-58.
Tidemann, C. R., Yorkston, H. D. and Russack, A. J. 1994. The diet of cats, Felis catus, on Christmas Island, Indian Ocean. Wildlife Research 21: 279-286.
Williams, K., Parer, I., Coman, B., Burley, J. and Braysher, M. 1995 Managing Vertebrate Pests: Rabbits. Australian Government Publishing Service: Canberra.
Wolfe, K. M., Mills, H. R., Garkaklis, M. J. and Bencini, R. 2004. Post-mating survival in a small marsupial is associated with nutrient inputs from seabirds. Ecology 85: 1740-1746.
Zavaleta, E. S., Hobbs, R. J. and Mooney, H. A. 2001. Viewing invasive species removal in a whole-ecosystem context. Trends in Ecology and Evolution 16: 454-459.
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References

Arthur, W. and Mitchell, P. 1989. A revised scheme for the classification of population interactions. Oikos 56: 141-143.
Baldock, F. C., Alexander, L. and More, S. J. 2003. Estimated and predicted changes in the cat population of Australian households from 1979 to 2005. Australian Veterinary Journal 81: 289-292.
Banks, P. B., Dickman, C. R. and Newsome, A. E. 1998. Ecological costs of feral predator control: foxes and rabbits. Journal of Wildlife Management 62: 766-772.
Barratt, D. G. 1998. Predation by house cats, Felis catus (L.), in Canberra, Australia. II. Factors affecting the amount of prey caught and estimates of the impact on wildlife. Wildlife Research 25: 475-487.
Bencini, R., McCulloch, C., Mills, H. R. and Start, A. N. 2001. Habitat and diet of the dibbler ( Parantechinus apicalis) on two islands in Jurien Bay, Western Australia. Wildlife Research 28: 465-468.
Braysher, M. 1993 Managing Vertebrate Pests: Principles and Strategies. Australian Government Publishing Service: Canberra.
Burrows, N. D., Algar, D., Robinson, A. D., Sinagra, J., Ward, B. and Liddelow, G. 2003. Controlling introduced predators in the Gibson Desert of Western Australia. Journal of Arid Environments 55: 691-713.
Caughley, G. and Gunn, A. 1996 Conservation Biology in Theory and Practice. Blackwell Science: Oxford.
Caughley, J., Monamy, V. and Heiden, K. 1994 Impact of the 1993 Mouse Plague. Occasional Paper Series No. 7, Grains Research and Development Corporation: Canberra.
Chapple, R. 2005. The politics of feral horse management in Guy Fawkes River National Park, NSW. Australian Zoologist 33: 233-246.
Courchamp, F., Langlais, M. and Sugihara, G. 1999. Cats protecting birds: modelling the mesopredator release effect. Journal of Animal Ecology 68: 282-292.
Crooks, K. R. and Soulé, M. E. 1999. Mesopredator release and avifaunal extinctions in a fragmented system. Nature 400: 563-566.
Crowther, M. S., Dickman, C. R. and Lynam, A. J. 1999 Sminthopsis griseoventer boullangerensis (Marsupialia: Dasyuridae), a new subspecies in the S. murina complex from Boullanger Island, Western Australia. Australian Journal of Zoology 47: 215-243.
CSIRO. 1991 The Insects of Australia, 2nd edition. Melbourne University Press: Melbourne.
Dickman, C. R. 1988. Body size, prey size, and community structure in insectivorous mammals. Ecology 69: 569-580.
Dickman, C. R. 1992. Conservation of mammals in the Australasian region: the importance of islands. Pp. 175-214 in Australia and the Global Environmental Crisis, edited by J. N. Coles and J. M. Drew. Academy Press: Canberra.
Dickman, C. R. 1996 Overview of the Impacts of Feral Cats on Australian Native Fauna. Australian Nature Conservation Agency: Canberra.
Dickman, C. R. 1999. Rodent-ecosystem relationships: a review. Pp. 113-133 in Ecologically-based Management of Rodent Pests, edited by G. R. Singleton, L. A. Hinds, H. Leirs and Z. Zhang. Australian Centre for International Agricultural Research: Canberra.
Dickman, C. R. 2006a. Species interactions: direct effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Dickman, C. R. 2006b. Species interactions: indirect effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Dickman, C. R. 2006c. House cats in the Australian environment: behavior, impacts, and management. Berryman Bulletin.
Dickman, C. R. and Lunney, D. (eds) 2001 A Symposium on the Dingo. Royal Zoological Society of New South Wales: Mosman.
Dickman, C. R. and Murray, B. R. 2006. Species interactions: complex effects. In press in Ecology: an Australian Perspective, 2nd edition, edited by P. Attiwill and B. Wilson. Oxford University Press: Melbourne.
Disney, H. J. de S. and Smithers, C. N. 1972. The distribution of terrestrial and freshwater birds on Lord Howe Island, in comparison with Norfolk Island. Australian Zoologist 17: 1-11.
Ewel, J. J. and Putz, F. E. 2005. A place for alien species in ecosystem restoration. Frontiers in Ecology and the Environment 2: 354-360.
Fitzgerald, B. M. 1988. Diet of domestic cats and their impact on prey populations. Pp. 123-147 in The Domestic Cat: the Biology of its Behaviour, edited by D. C. Turner and P. Bateson. Cambridge University Press: Cambridge.
Fitzgerald, B. M., Karl, B. J. and Veitch, C. R. 1991. The diet of feral cats ( Felis catus) on Raoul Island, Kermadec group. New Zealand Journal of Ecology 15: 123-129.
Flecker, A. S. and Townsend, C. R. 1994. Community-wide consequences of trout introduction in New Zealand streams. Ecological Applications 4: 798-807.
Fuller, P. J. and Burbidge, A. A. 1987. Discovery of the dibbler, Parantechinus apicalis, on islands at Jurien Bay. Western Australian Naturalist 16: 177-181.
Gentle, M. N. 2005. Factors affecting the efficiency of fox ( Vulpes vulpes) baiting practices on the Central Tablelands of New South Wales. PhD thesis, University of Sydney: Sydney.
Gillies, C. and Clout, M. 2003. The prey of domestic cats ( Felis catus) in two suburbs of Auckland City, New Zealand. Journal of Zoology 259: 309-315.
Glen, A. S. and Dickman, C. R. 2005. Complex interactions among mammalian carnivores in Australia, and their implications for wildlife management. Biological Reviews 80: 387-401.
Grayson, J. and Calver, M. 2004. Regulation of domestic cat ownership to protect urban wildlife: a justification based on the precautionary principle. Pp. 169-178 in Urban Wildlife: More than meets the Eye, edited by D. Lunney and S. Burgin. Royal Zoological Society of New South Wales: Mosman.
Grayson, J., Calver, M. C. and Styles, I. 2002. Attitudes of suburban Western Australians to proposed cat control legislation. Australian Veterinary Journal 80: 536-543.
Grigg, G. 2002. Conservation benefit from harvesting kangaroos: status report at the start of a new millennium. A paper to stimulate discussion and research. Pp. 53-76 in A Zoological Revolution: Using Native Fauna to Assist in its own Survival, edited by D. Lunney and C. R. Dickman. Royal Zoological Society of New South Wales: Mosman.
Kuchling, G. 1997. Managing the last survivors: integration of in situ and ex situ conservation of Pseudemydura umbrina. Pp. 339-344 in Proceedings: Conservation, Management and Restoration of Tortoises and Turtles, 11-16 July 1993, edited by J. van Abbema. New York Turtle and Tortoise Society: New York.
Kuchling, G. 2000. Conservation strategies for remnant turtle populations: the Western Australian swamp turtle Pseudemydura umbrina recovery programme. Stapfia 69, Zugleich Kataloge des O. Ö. Landesmuseums N.F. 149: 179-188.
Lilith, M., Calver, M., Styles, I. and Garkaklis M. 2006. Protecting wildlife from predation by owned domestic cats: application of a precautionary approach to the acceptability of proposed cat regulations. Austral Ecology 31: 176-189.
Low, T. 1999 Feral Future. Penguin Books: Melbourne.
Low, T. 2004 The New Nature. Penguin Books: Melbourne.
MacDougall, A. S. and Turkington, R. 2005. Are invasive species the drivers or passengers of change in degraded ecosystems? Ecology 86: 42-55.
Matthews, A., Dickman, C. R. and Major, R. E. 1999. The influence of fragment size and edge on nest predation in urban bushland. Ecography 22: 349-356.
McLeod, R. 2004 Counting the Cost: Impact of Invasive Animals in Australia, 2004. Cooperative Research Centre for Pest Animal Control: Canberra.
Meek, P. D. 1998. Food items brought home by domestic cats Felis catus (L.) living in Booderee National Park, Jervis Bay. Proceedings of the Linnean Society of New South Wales 120: 43-47.
Miller, S., Bencini, R., Mills, H. and Moro, D. 2003. Food availability for the dibbler ( Parantechinus apicalis) on Boullanger and Whitlock Islands, Western Australia. Wildlife Research 30: 649-654.
Molsher, R. 1999. The ecology of feral cats ( Felis catus) in open forest in New South Wales: interactions with food resources and foxes. PhD thesis, University of Sydney: Sydney
Morin, P. J. 1999 Community Ecology. Blackwell Science: Oxford.
Newsome, A. 1990. The control of vertebrate pests by vertebrate predators. Trends in Ecology and Evolution 5: 187-191.
Newsome, A., Pech, R., Smyth, R., Banks, P. and Dickman, C. 1997 Potential Impacts on Australian Native Fauna of Rabbit Calicivirus Disease. Environment Australia: Canberra.
NSW National Parks and Wildlife Service. 2002 Draft Wild Horse Management Plan for the Alpine Area of Kosciuszko National Park. NSW National Parks and Wildlife Service: Sydney.
Olsen, P. 1998 Australia's Pest Animals: New Solutions to Old Problems. Kangaroo Press: Sydney.
Pearson, D. E. and Callaway, R. M. 2003. Indirect effects of host-specific biological control agents. Trends in Ecology and Evolution 18: 456-461.
Proulx, S. R., Promislow, D. E. L. and Phillips, P. C. 2005. Network thinking in ecology and evolution. Trends in Ecology and Evolution 20: 345-353.
Reark 1994 The Metropolitan Domestic Cat: a Survey of the Population Characteristics and Hunting Behaviour of the Domestic Cat in Australia. Petcare Information and Advisory Service: Melbourne.
Risbey, D. A., Calver, M. C., Short, J., Bradley, J. S. and Wright, I. W. 2000. The impact of cats and foxes on the small vertebrate fauna of Heirisson Prong, Western Australia. II. A field experiment. Wildlife Research 27: 223-235.
Sinclair, A. R. E., Krebs, C. J., Fryxell, J. M., Turkington, R., Boutin, S., Boonstra, R., Seccombe-Hett, P., Lundberg, P. and Oksanen, L. 2000. Testing hypotheses of trophic level interactions: a boreal forest ecosystem. Oikos 89: 313-328.
Soulé, M. E., Bolger, D. T., Alberts, A. C., Wright, J., Sorice, M. and Hill, S. 1988. Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. Conservation Biology 2: 75-92.
Start, A. N., Moro, D., Adams, M. and Bencini, R. 2006. Dunnarts from Boullanger Island: new evidence and reassessment of a taxonomic issue with resource implications. Australian Mammalogy 28: 51-58.
Tidemann, C. R., Yorkston, H. D. and Russack, A. J. 1994. The diet of cats, Felis catus, on Christmas Island, Indian Ocean. Wildlife Research 21: 279-286.
Williams, K., Parer, I., Coman, B., Burley, J. and Braysher, M. 1995 Managing Vertebrate Pests: Rabbits. Australian Government Publishing Service: Canberra.
Wolfe, K. M., Mills, H. R., Garkaklis, M. J. and Bencini, R. 2004. Post-mating survival in a small marsupial is associated with nutrient inputs from seabirds. Ecology 85: 1740-1746.
Zavaleta, E. S., Hobbs, R. J. and Mooney, H. A. 2001. Viewing invasive species removal in a whole-ecosystem context. Trends in Ecology and Evolution 16: 454-459.
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