COMMENSAL SMALL MAMMAL SPECIES AND BAIT PREFERENCES IN URBAN AREAS OF PENANG ISLAND
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Published:
Aug 28, 2019
Keywords:
pests rodents urban area
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Abstract
Physical trapping is an important first step in controlling pest populations and species identification; and bait preferences are crucial information for effective management. This study was conducted to explore species diversity and bait preferences of commensal species in urban areas on Penang Island. Live trapping was carried out in eight sampling sites on Penang Island in commercial and residential areas. Eight different type of baits were used; apple and peanut butter (APB), white bread and peanut butter (BPB), salted fish (SF), salted fish and peanut butter (SFPB), white bread and fish extract (BFE), fried chicken leftover (FCL), vegetables and peanut butter (VPB) and sausages (S). The most successful bait was the high protein bait, fried chicken leftover (FCL), with a capture success rate of 35.56%. We also analysed the bait preferences for the different species of small mammals captured in our study. Five species of small mammals were captured in our study sites: Rattus norvegicus, Rattus rattus, Mus musculus, Bandicota bengalensis and Suncus murinus. Norway rats, R. norvegicus was the most trapped species in commercial areas whereas lesser bandicoot rats, B. bengalensis was the most captured species in residential areas.
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Amni, W. N., Ravindran, S., Saufi, S., Hamid, N. H., Zainal Abidin, C. M. R., Ahmad, A. H., & Salim, H. (2019). COMMENSAL SMALL MAMMAL SPECIES AND BAIT PREFERENCES IN URBAN AREAS OF PENANG ISLAND. Malaysian Journal of Science, 38(2), 18–30. https://doi.org/10.22452/mjs.vol38no2.2
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References
Aplin, K.A., Brown, P.R., Jacob, J., Krebs, C.J. and Singleton, G.R. (2003). Field methods for rodent studies in Asia and the Indo Pacific, Australian Centre for International Agricultural Research Monograph. pp. 26. Australia: Australian Centre for International Agricultural.
Barnett, S.A. (1963). The Rat: A Study in Behaviour. Chapter 3, Feeding Behaviour. Chicago, USA: Aldine Publishing Company.
Barnett, S.A., and Spencer, M.M. (1951). Feeding, social behaviour and interspecific competition in wild rats. Behaviour 3: 229–242.
Bealin, C., and Lawton, C. (2018). Bait preference in Irish small rodents. Irish Naturalists’ Journal 36(1): 9-13.
Behnke, J.M., Bajer, A., Sinski, E. and Wakelin, D. (2001). Interactions involving intestinal nematodes of rodents: experimental and field studies. Parasitology 122: 39 – 49.
Brooks, J.E. and Bowerman, A.M. (1973). Preference of wild Norway rats for grain, seeds and legumes. Pest Control 41: 13-39
Bullard, R.W. (1985). Isolation and characterization of natural products that attract or repel wild vertebrates. In: Acree, T., and Soderlund, D. (Eds.). Semiochemistry: Flavors and Pheromones. Walter de Gruyter & Co., New York, pp. 65–93.
Castillo, E., Priotto, J., Ambrosio, A.M., Provensal, M.C., Pini, N., Morales, M.A., Steinmann, A., Polop, J.J. (2003). Commensal and wild rodents in an urban area of Argentina. International Biodeterioration & Biodegradation 52: 135-141.
Cavia, R., Cueto, G.R. and Suarez, O.V. (2009). Changes in rodent communities according to the landscape structure in an urban ecosystem. Landscape and Urban Planning 90: 11-19
Child, J.E., Mclafferty, S.L., Sadek, R., Miller, G.L., Khan, A.S., Dupree, E.R., Advani, R. and Glass, G.E. (1998). Epidemiology of rodent bites and prediction of rat infestation in New York City. American Journal of Epidemiology 83: 590-598
Clapperton, B.K. (2006). A review of the current knowledge of rodent behaviour in relation to control devices. Wellington, Science and Technical Pub., Department of Conservation.
Diete, R.L., Meek, P.D., Dixon, K.M., Dickman, C.R., and Leung, L.K.-P. (2016). Best bait for your buck: bait preference for camera trapping north Australian mammals. Australian Journal of Zoology 63: 376-382. DOI: http://dx.doi.org/10.1071/ZO15050
Field, A.J., Copsey, J.A., Tragett, C.E.E., and Gode, M. (2017). Improving capture rate for an invasive species in Mauritius: determining Asian musk shrew Suncus murinus bait preference. Conservation Evidence 14: 16-19.
Gardner Santana, L.C., Norris, D.E., Fornadel, C.M., Hinson, E.R., Klein, S.L. and Glass, G.E. (2009). Commensal ecology, urban landscape, and their influence on the genetic characteristics of city-dwelling Norway rats (Rattus norvegicus). Molecular Ecology 18(13): 2766-2778
Gómez, M.D., Priotto, J., Provensal, M.C., Steinmann, A., Castillo, E., and Polop, J.J. (2008). A population study of house mice (Mus musculus) inhabiting different habitats in an Argentine urban area. International Biodeterioration and Biodegradation 62:270–273.
Grainger, J.P., and Fairley, J.S. (1978). Studies on the biology of the pygmy shrew Sorex minutus in the west of Ireland. Journal of Zoology 186:109–141.
Hamilton, W.J. (1940). The biology of the smoky shrew (Sorex fumeus fumeus Miller). Zoologica 25:307–310.
Harper, G.A., and Bunbury, N. (2015). Invasive rats on tropical islands: their population biology and impacts on native species. Global Ecology and Conservation 3: 607–627.
Harrison, J.L., and Woodville, H.C. (1950). Notes on the feeding habits of house-rats in Rangoon, Burma. Annals of Applied Biology 37: 296–304.
Hice, C.L., and Velazco, P.M. (2013). Relative effectiveness of several bait and trap types for assessing small mammal communities in Neotropical rainforest. Occasional Papers, Museum of Texas Tech University 316: 1–15.
Jackson, M., Hartley, S., and Linklater, W. (2016). Better food-based baits and lures for invasive rats Rattus spp. and the brushtail possum Trichosurus vulpecula: a bioassay on wild, free-ranging animals. Journal of Pest Science 89: 479–488. DOI: https://doi.org/10.1007/ s10340-015-0693-8.
Khan, J.A. (1974). Laboratory experiments on the food preferences of the black rat (Rattus rattus L.). Zoological Journal of Linnean Society 54: 167-172.
Kolody, N., Brosvic, G.M., Pak, D., and Loeffler, S. (1993). Taste preference behavior in Long-Evans rats and Egyptian spiny mice. Bulletin of the Psychonomic Society 31: 307–310.
Lam, Y.M., Lee, A.K., Tan, Y.P. and Mohan, E. (1982). A Case Warfarin Resistance in Rattus rattus diardii. Mardi Research Bulletin 10(3): 378-383.
Langton, S., Cowan, D. and Meyer, A. (2001). The occurrence of commensal rodents in dwellings as revealed by the 1996 English House Condition Survey. Journal of Applied Ecology 38: 699-709
Lim, B.L. (1970). Distribution, relative abundance, food habits and parasite patterns of Giant rats (Rattus) in West Malaysia. Journal of Mammalogy 51(4): 730-740.
Lim, B.L. (2015). The house rodents and house shrew in Malaysia and Southeast Asia. UTAR Agriculture Science Journal 1(2): 45-50
Marsh, R.E. (1994). Roof rats. In: Hygnstorm S.E., Timm, R.M. & Larson G.E. (eds.): Prevention and control of wildlife damage, pp. 125-135, California, USA.
Medway, L. & Lim, B.L. (1966). Fauna of Pulau Tioman: The specifics relations of Rattus tiomanicus. Bulletin of the National Museum Singapore 34: 33-38
Meehan, A.P. (1984). Rats and Mice: their biology and control. Sussex, UK: Rentokil.
Nelson, L. and Clark, F.W. (1973). Correction for sprung traps in catch/effort calculations of trapping results. Journal of Mammalogy 54(1): 295–298
Nolte, D.L. (1999). Impact of rodents on reforestation in the United States: problems and management. In: Zhi-bin, Z., Hinds, E., Singleton, G., and Zhang, Z. (Eds) Rodent biology and management. pp 84. In abstracts of papers presented at International Conference on Rodent Biology and Management, Beijing, China, 5–9 October 1998. ACIAR Technical Report 45.
Paramasvaran, S., Sani, R.A., Krishnasamy, M., Amal Nasir, M., Hassan, L., Mohd Zain, S.N., Jeffery, J., Santhana Raj, L., Latifa, I., Abdul Aziz, Vellayan, S. and Selvasan, S. (2012). An overview of nematodes infecting urban and wild rats (Muridae) in Malaysia. Malaysian Journal of Veterinary Research 3(1): 15–21
Parshad, V.R. (1999). Rodent control in India. Integrated Pest Management Reviews 4: 97-126
Partridge, L., and Maclean, R. (1981). Effects of nutrition and peripheral stimuli on preferences for familiar foods in the bank vole. Animal Behaviour 29: 217-220.
Paull, D.J., Claridge, A.W., and Barry, S.C. (2011). There’s no accounting for taste: bait attractants and infrared digital cameras for detecting small to medium ground-dwelling mammals. Wildlife Research 38: 188–195. DOI:10.1071/WR10203
Priyambodo, S. and Pelz, H.J. (2002). Studies on neophobic behaviour in Norway rats (Rattus norvegicus) from farms in Germany. In: Singleton, G.R., Hinds, L.A., Krebs, C.J. and Spratt, D.M. (eds.): Rats, mice and people: Rodent biology and management. Bogor, Indonesia, pp.564, Australian Centre for International Agricultural Research.
Robards, G.E. and Saunders, G. (1998). Food preferences of house mice (Mus domesticus) and their implication for control strategies. Wildlife Research 25: 595-601.
Schlötelburg, A., Jakob, G., Bellingrath-Kimura, S., and Jacob, J. (2018). Natural bait additives improve trapping success of common voles, Microtus arvalis. Applied Animal Behaviour Science 208: 75-81. DOI: https://doi.org/10.1016/j.applanim.2018.08.013
Shahwar, D., Hussain, I., Anwar, M., Beg, M.A., Kawan, A., and Akrim, F. (2015). Development of cereal baits and comparative field efficacy of some additives as bait carrier for zinc phosphide and coumatetralyl against rodent pests of poultry farms in Rawalpindi-Islamabad, Pakistan. International Biodeterioration and Biodegradation 104: 460-471.
Singla, N., and Kanwar, D. (2014). Poultry egg components as cereal bait additives for enhancing rodenticide based control success and trap index of house rat, Rattus rattus. Asian Pacific Journal of Tropical Biomedicine 4(Suppl 1): S341-S347. DOI:10.12980/APJTB.4.2014C1260
Timm, R.M. (1994). Norway Rats. Prevention and Control of Wildlife Damage, 1994. Lincoln, USA: Institute of Agriculture and Natural Resources, University of Nebraska.
Tung, K.C., Hsiao, F.C., Wang, K.S., Yong, C.H. and Lai, C.H. (2013). Study of the endoparasitic fauna commensal rats and shrews caught in traditional wet markets in Taichung City, Taiwan. Journal of Microbiology, Immunology and Infection 46: 85-88
Vadell, M.V., Cavia, R., and Suárez, O.V. (2010). Abundance, age structure and reproductive patterns of Rattus norvegicus and Mus musculus in two areas of the city of Buenos Aires. International Journal of Pest Management 56(4): 327–336.
Walsh, J.F., Molyneux, D.H. and Birley, M.H. (1993). Deforestation effects on vector – borne diseases. Journal of Parasitology 106: 55–75.
Yabe, T. (1979). The relation of food habits to the ecological distributions of Norway rat (Rattus norvegicus) and the roof rat (Rattus rattus). Japanese Journal of Ecology 29: 235.
Barnett, S.A. (1963). The Rat: A Study in Behaviour. Chapter 3, Feeding Behaviour. Chicago, USA: Aldine Publishing Company.
Barnett, S.A., and Spencer, M.M. (1951). Feeding, social behaviour and interspecific competition in wild rats. Behaviour 3: 229–242.
Bealin, C., and Lawton, C. (2018). Bait preference in Irish small rodents. Irish Naturalists’ Journal 36(1): 9-13.
Behnke, J.M., Bajer, A., Sinski, E. and Wakelin, D. (2001). Interactions involving intestinal nematodes of rodents: experimental and field studies. Parasitology 122: 39 – 49.
Brooks, J.E. and Bowerman, A.M. (1973). Preference of wild Norway rats for grain, seeds and legumes. Pest Control 41: 13-39
Bullard, R.W. (1985). Isolation and characterization of natural products that attract or repel wild vertebrates. In: Acree, T., and Soderlund, D. (Eds.). Semiochemistry: Flavors and Pheromones. Walter de Gruyter & Co., New York, pp. 65–93.
Castillo, E., Priotto, J., Ambrosio, A.M., Provensal, M.C., Pini, N., Morales, M.A., Steinmann, A., Polop, J.J. (2003). Commensal and wild rodents in an urban area of Argentina. International Biodeterioration & Biodegradation 52: 135-141.
Cavia, R., Cueto, G.R. and Suarez, O.V. (2009). Changes in rodent communities according to the landscape structure in an urban ecosystem. Landscape and Urban Planning 90: 11-19
Child, J.E., Mclafferty, S.L., Sadek, R., Miller, G.L., Khan, A.S., Dupree, E.R., Advani, R. and Glass, G.E. (1998). Epidemiology of rodent bites and prediction of rat infestation in New York City. American Journal of Epidemiology 83: 590-598
Clapperton, B.K. (2006). A review of the current knowledge of rodent behaviour in relation to control devices. Wellington, Science and Technical Pub., Department of Conservation.
Diete, R.L., Meek, P.D., Dixon, K.M., Dickman, C.R., and Leung, L.K.-P. (2016). Best bait for your buck: bait preference for camera trapping north Australian mammals. Australian Journal of Zoology 63: 376-382. DOI: http://dx.doi.org/10.1071/ZO15050
Field, A.J., Copsey, J.A., Tragett, C.E.E., and Gode, M. (2017). Improving capture rate for an invasive species in Mauritius: determining Asian musk shrew Suncus murinus bait preference. Conservation Evidence 14: 16-19.
Gardner Santana, L.C., Norris, D.E., Fornadel, C.M., Hinson, E.R., Klein, S.L. and Glass, G.E. (2009). Commensal ecology, urban landscape, and their influence on the genetic characteristics of city-dwelling Norway rats (Rattus norvegicus). Molecular Ecology 18(13): 2766-2778
Gómez, M.D., Priotto, J., Provensal, M.C., Steinmann, A., Castillo, E., and Polop, J.J. (2008). A population study of house mice (Mus musculus) inhabiting different habitats in an Argentine urban area. International Biodeterioration and Biodegradation 62:270–273.
Grainger, J.P., and Fairley, J.S. (1978). Studies on the biology of the pygmy shrew Sorex minutus in the west of Ireland. Journal of Zoology 186:109–141.
Hamilton, W.J. (1940). The biology of the smoky shrew (Sorex fumeus fumeus Miller). Zoologica 25:307–310.
Harper, G.A., and Bunbury, N. (2015). Invasive rats on tropical islands: their population biology and impacts on native species. Global Ecology and Conservation 3: 607–627.
Harrison, J.L., and Woodville, H.C. (1950). Notes on the feeding habits of house-rats in Rangoon, Burma. Annals of Applied Biology 37: 296–304.
Hice, C.L., and Velazco, P.M. (2013). Relative effectiveness of several bait and trap types for assessing small mammal communities in Neotropical rainforest. Occasional Papers, Museum of Texas Tech University 316: 1–15.
Jackson, M., Hartley, S., and Linklater, W. (2016). Better food-based baits and lures for invasive rats Rattus spp. and the brushtail possum Trichosurus vulpecula: a bioassay on wild, free-ranging animals. Journal of Pest Science 89: 479–488. DOI: https://doi.org/10.1007/ s10340-015-0693-8.
Khan, J.A. (1974). Laboratory experiments on the food preferences of the black rat (Rattus rattus L.). Zoological Journal of Linnean Society 54: 167-172.
Kolody, N., Brosvic, G.M., Pak, D., and Loeffler, S. (1993). Taste preference behavior in Long-Evans rats and Egyptian spiny mice. Bulletin of the Psychonomic Society 31: 307–310.
Lam, Y.M., Lee, A.K., Tan, Y.P. and Mohan, E. (1982). A Case Warfarin Resistance in Rattus rattus diardii. Mardi Research Bulletin 10(3): 378-383.
Langton, S., Cowan, D. and Meyer, A. (2001). The occurrence of commensal rodents in dwellings as revealed by the 1996 English House Condition Survey. Journal of Applied Ecology 38: 699-709
Lim, B.L. (1970). Distribution, relative abundance, food habits and parasite patterns of Giant rats (Rattus) in West Malaysia. Journal of Mammalogy 51(4): 730-740.
Lim, B.L. (2015). The house rodents and house shrew in Malaysia and Southeast Asia. UTAR Agriculture Science Journal 1(2): 45-50
Marsh, R.E. (1994). Roof rats. In: Hygnstorm S.E., Timm, R.M. & Larson G.E. (eds.): Prevention and control of wildlife damage, pp. 125-135, California, USA.
Medway, L. & Lim, B.L. (1966). Fauna of Pulau Tioman: The specifics relations of Rattus tiomanicus. Bulletin of the National Museum Singapore 34: 33-38
Meehan, A.P. (1984). Rats and Mice: their biology and control. Sussex, UK: Rentokil.
Nelson, L. and Clark, F.W. (1973). Correction for sprung traps in catch/effort calculations of trapping results. Journal of Mammalogy 54(1): 295–298
Nolte, D.L. (1999). Impact of rodents on reforestation in the United States: problems and management. In: Zhi-bin, Z., Hinds, E., Singleton, G., and Zhang, Z. (Eds) Rodent biology and management. pp 84. In abstracts of papers presented at International Conference on Rodent Biology and Management, Beijing, China, 5–9 October 1998. ACIAR Technical Report 45.
Paramasvaran, S., Sani, R.A., Krishnasamy, M., Amal Nasir, M., Hassan, L., Mohd Zain, S.N., Jeffery, J., Santhana Raj, L., Latifa, I., Abdul Aziz, Vellayan, S. and Selvasan, S. (2012). An overview of nematodes infecting urban and wild rats (Muridae) in Malaysia. Malaysian Journal of Veterinary Research 3(1): 15–21
Parshad, V.R. (1999). Rodent control in India. Integrated Pest Management Reviews 4: 97-126
Partridge, L., and Maclean, R. (1981). Effects of nutrition and peripheral stimuli on preferences for familiar foods in the bank vole. Animal Behaviour 29: 217-220.
Paull, D.J., Claridge, A.W., and Barry, S.C. (2011). There’s no accounting for taste: bait attractants and infrared digital cameras for detecting small to medium ground-dwelling mammals. Wildlife Research 38: 188–195. DOI:10.1071/WR10203
Priyambodo, S. and Pelz, H.J. (2002). Studies on neophobic behaviour in Norway rats (Rattus norvegicus) from farms in Germany. In: Singleton, G.R., Hinds, L.A., Krebs, C.J. and Spratt, D.M. (eds.): Rats, mice and people: Rodent biology and management. Bogor, Indonesia, pp.564, Australian Centre for International Agricultural Research.
Robards, G.E. and Saunders, G. (1998). Food preferences of house mice (Mus domesticus) and their implication for control strategies. Wildlife Research 25: 595-601.
Schlötelburg, A., Jakob, G., Bellingrath-Kimura, S., and Jacob, J. (2018). Natural bait additives improve trapping success of common voles, Microtus arvalis. Applied Animal Behaviour Science 208: 75-81. DOI: https://doi.org/10.1016/j.applanim.2018.08.013
Shahwar, D., Hussain, I., Anwar, M., Beg, M.A., Kawan, A., and Akrim, F. (2015). Development of cereal baits and comparative field efficacy of some additives as bait carrier for zinc phosphide and coumatetralyl against rodent pests of poultry farms in Rawalpindi-Islamabad, Pakistan. International Biodeterioration and Biodegradation 104: 460-471.
Singla, N., and Kanwar, D. (2014). Poultry egg components as cereal bait additives for enhancing rodenticide based control success and trap index of house rat, Rattus rattus. Asian Pacific Journal of Tropical Biomedicine 4(Suppl 1): S341-S347. DOI:10.12980/APJTB.4.2014C1260
Timm, R.M. (1994). Norway Rats. Prevention and Control of Wildlife Damage, 1994. Lincoln, USA: Institute of Agriculture and Natural Resources, University of Nebraska.
Tung, K.C., Hsiao, F.C., Wang, K.S., Yong, C.H. and Lai, C.H. (2013). Study of the endoparasitic fauna commensal rats and shrews caught in traditional wet markets in Taichung City, Taiwan. Journal of Microbiology, Immunology and Infection 46: 85-88
Vadell, M.V., Cavia, R., and Suárez, O.V. (2010). Abundance, age structure and reproductive patterns of Rattus norvegicus and Mus musculus in two areas of the city of Buenos Aires. International Journal of Pest Management 56(4): 327–336.
Walsh, J.F., Molyneux, D.H. and Birley, M.H. (1993). Deforestation effects on vector – borne diseases. Journal of Parasitology 106: 55–75.
Yabe, T. (1979). The relation of food habits to the ecological distributions of Norway rat (Rattus norvegicus) and the roof rat (Rattus rattus). Japanese Journal of Ecology 29: 235.