The effects of invasion on parasite dynamics and communities

Functional Ecology

Volumn 26, Issue 6, 2012, Pages 1288-1299

  Telfer, Sandra ^a   Bown, Kevin ^b  

^a UNIVERSITY OF ABERDEEN   (United Kingdom)

^b UNIVERSITY OF SALFORD   (United Kingdom)

Author keywords

Density dependent; Disease emergence; Frequency dependent; Multihost; Non indigenous species; Pathogen pollution; Spillback; Spillover; Transmission

Indexed keywords

BIOLOGICAL INVASION; COMMUNITY RESPONSE; DENSITY DEPENDENCE; DISEASE INCIDENCE; DISEASE VECTOR; ECOLOGICAL IMPACT; EMPIRICAL ANALYSIS; FREQUENCY DEPENDENCE; HOST; INVASIVE SPECIES; MORTALITY; NATIVE SPECIES; PARASITE TRANSMISSION; PATHOGEN; POPULATION DYNAMICS; RELATIVE ABUNDANCE;

EID: 84869864675     PISSN: 02698463     EISSN: 13652435     Source Type: Journal    
DOI: 10.1111/j.1365-2435.2012.02049.x     Document Type: Article

References (101)

1. Albon, S.D., Stien, A., Irvine, R.J., Langvaten, R., Ropstad, E. & Halvorsen, O. (2002) The role of parasites in the dynamic of a reindeer population. Proceedings of the Royal Society B, 269, 1625-1632.
2. Altizer, S., Dobson, A., Hosseini, P., Hudson, P. & Pascual, M. (2006) Seasonality and the dynamics of infectious diseases. Ecology Letters, 9, 467-484.
3. Begon, M., Bennett, M., Bowers, R.G., French, N.P., Hazel, S.M. & Turner, J. (2002) A clarification of transmission terms in host-microparasite models: numbers, densities and areas. Epidemiology and infection, 129, 147-153.
4. Beldomenico, P.M. & Begon, M. (2009) Disease spread, susceptibility and infection intensity: vicious circles? Trends in Ecology & Evolution, 25, 21-27.
5. Brooks, C.P. & Zhang, H. (2010) A null model of community disassembly effects on vector-borne disease risk. Journal of Theoretical Biology, 264, 866-873.
6. Brown, C.R., Padhi, A., Moore, A.T., Brown, M.B., Foster, J.E., Pfeffer, M., O'Brien, V.A. & Komar, N. (2009) Ecological divergence of two sympatric lineages of Buggy Creek virus, an arbovirus associated with birds. Ecology, 90, 3168-3179.
7. Byers, J.E. & Goldwasser, L. (2001) Exposing the mechanism and timing of impact of non-indigenous species on native species. Ecology, 82, 1330-1343.
8. Cattadori, I.M., Boag, B. & Hudson, P.J. (2008) Parasite co-infection and interaction as drivers of host heterogeneity. International Journal for Parasitology, 38, 371-380.
9. Chapman, J.W., Dumbauld, B.R., Itani, G. & Markham, J.C. (2012) An introduced Asian parasite threatens northeastern Pacific estuarine ecosystems. Biological Invasions, 14, 1221-1236.
10. Chen, H.W., Shao, K.T., Liu, C.W.J., Lin, W.H. & Liu, W.C. (2011) The reduction of food web robustness by parasitism: fact and artefact. International Journal for Parasitology, 41, 627-634.
11. Christiano, R.S.C. & Scherm, H. (2007) Quantitative aspects of the spread of Asian soybean rust in the southeastern United States, 2005 to 2006. Phytopathology, 97, 1428-1433.
12. Cleaveland, S., Hess, G.R., Dobson, A.P., Laurenson, M.K., McCallum, H.I., Roberts, M.G. & Woodroffe, R. (2002) The role of pathogens in biological conservation. The Ecology of Wildlife Diseases (eds P.J. Hudson, A. Rizzoli, B.T. Grenfell, J.A.P. Heesterbeek & A.P. Dobson), pp. 139-150. Oxford University Press, Oxford.
13. Cobb, R.C., Filipe, J.A.N., Meentemeyer, R.K., Gilligan, C.A. & Rizzo, D.M. (2012) Ecosystem transformation by emerging infectious disease: loss of large tanoak from California forests. Journal of Ecology, 100, 712-722.
14. Colautti, R.I., Ricciardi, A., Grigorovich, I.A. & MacIsaac, H.J. (2004) Is invasion success explained by the enemy release hypothesis? Ecology Letters, 7, 721-733.
15. Condeso, T.E. & Meentemeyer, R.K. (2007) Effects of landscape heterogeneity on the emerging forest disease sudden oak death. Journal of Ecology, 95, 364-375.
16. Cronin, J.P., Welsh, M.E., Dekkers, M.G., Abercrombie, S.T. & Mitchell, C.E. (2010) Host physiological phenotype explains pathogen reservoir potential. Ecology Letters, 13, 1221-1232.
17. Daszak, P., Cunningham, A.A. & Hyatt, A.D. (2000) Emerging infectious diseases of wildlife - threats to biodiversity and human health. Science, 287, 443-449.
18. Dobson, A. (2004) Population dynamics of pathogens with multiple host species. American Naturalist, 164, S64-S78.
19. Duplantier, J.M., Duchemin, J.B., Chanteau, S. & Carniel, E. (2005) From the recent lessons of the Malagasy foci towards a global understanding of the factors involved in plague re-emergence. Veterinary Research, 36, 437-453.
20. Ezenwa, V.O. & Jolles, A.E. (2011) From host immunity to pathogen invasion: the effects of helminth coinfection on the dynamics of microparasites. Integrative and Comparative Biology, 51, 540-551.
21. Ezenwa, V.O., Etienne, R.S., Luikart, G., Beja-Pereira, A. & Jolles, A.E. (2010) Hidden consequences of living in a wormy world: nematode-induced immune suppression facilitates tuberculosis invasion in African buffalo. American Naturalist, 176, 613-624.
22. Fabiszewski, A.M., Umbanhowar, J. & Mitchell, C.E. (2010) Modeling landscape-scale pathogen spillover between domesticated and wild hosts: Asian soybean rust and kudzu. Ecological Applications, 20, 582-592.
23. Foley, P. & Foley, J. (2010) Modeling susceptible infective recovered dynamics and plague persistence in Californian rodent - flea communities. Oikos, 10, 59-67.
24. Gassó, N., Sol, D., Pino, J., Dana, E.D., Lloret, F., Sanz-elorza, M. & Sobrino, E. (2009) Exploring species attributes and site characteristics to assess plant invasions in Spain. Diversity and Distributions, 5, 0-58.
25. Gendron, A.D., Marcogliese, D.J. & Thomas, M. (2011) Invasive species are less parasitized than native competitors, but for how long? The case of the round goby in the Great Lakes-St. Lawrence Basin Biological Invasions, 14, 367-384.
26. Gilbert, L., Norman, R.A., Laurenson, M.K., Reid, H. & Hudson, (2001) Disease persistence and apparent competition in a three-host community: an empirical and analytical study of large-scale, wild populations. Journal of Animal Ecology, 70, 1053-1061.
27. Graham, A.L., Cattadori, I.M., Lloyd-smith, J.O., Ferrari, M.J. & Bjørnstad, O.N. (2007) Transmission consequences of coinfection: cytokines writ large? Trends in Parasitology, 23, 284-291.
28. Hamer, G.L., Chaves, L.F., Anderson, T.K., Kitron, U.D., Brawn, J.D., Ruiz, M.O., Loss, S.R., Walker, E.D. & Goldberg, T.L. (2011) Fine-scale variation in vector host use and force of infection drive localized patterns of West Nile Virus transmission. PLoS ONE, 6, e23767.
29. Harris, R.N., Brucker, R.M., Walke, J.B., Becker, M.H., Schwantes, C.R., Flaherty, D.C., Lam, B.A., Woodhams, D.C., Briggs, C.J., Vredenburg, V.T. & Minbiole, K.P.C. (2009) Skin microbes on frogs prevent morbidity and mortality caused by a lethal skin fungus. Microbial Ecology, 3, 818-824.
30. Haydon, D.T., Cleaveland, S., Taylor, L.H. & Laurenson, M.K. (2002) Identifying reservoirs of infection: a conceptual and practical challenge. Emerging Infectious Diseases, 8, 1468-1473.
31. Hayes, K.R. & Barry, S. (2008) Are there any consistent predictors of invasion success? Biological Invasions, 10, 483-506.
32. Hayes, E.B., Komar, N., Nasci, R.S., Montgomery, S.P., O'Leary, D.R. & Campbell, G.L. (2005) Epidemiology and transmission dynamics of West Nile virus disease. Emerging Infectious Diseases, 11, 1167-1173.
33. Hentonnen, H., Fuglei, E., Gower, C.N., Hauisalmi, V., Ims, R.A., Niemimaa, J. & Yoccoz, N.G. (2001) Echinococcus multilocularis on Svalbard: introduction of an intermediate host has enabled the local life-cycle. Parasitology, 123, 547-552.
34. Hess, G.R., Randolph, S.E., Arneberg, P., Chemini, C., Furlanello, C., Harwood, J., Roberts, M.G. & Swinton, J. (2002) Spatial aspects of disease dynamics. The Ecology of Wildlife Diseases (eds P.J. Hudson, A. Rizzoli, B.T. Grenfell, J.A.P. Heesterbeek & A.P. Dobson), pp. 102-118. Oxford University Press, Oxford.
35. Hingston, M., Goodman, S.M., Ganzhorn, J.U. & Sommer, S. (2005) Reconstruction of the colonization of southern Madagascar by introduced Rattus rattus. Journal of Biogeography, 32, 1549-1559.
36. Hoberg, E.P., Kutz, S.J., Nagy, J., Jenkins, E., Elkin, B., Branigan, M. & D., C. (2002) Protostrongylus stilesi (Nematoda: Protostrongylidae): ecological isolation and putative host-switching between Dall's sheep and muskoxen in a contact zone. Comparative Parasitology, 69, 1-9.
37. Holt, R.D., Dobson, A.P., Begon, M., Bowers, R.G. & Schauber, E.M. (2003) Parasite establishment in host communities. Ecology Letters, 6, 837-842.
38. Hudson, P.J., Dobson, A.P. & Newborn, D. (1998) Prevention of population cycles by parasite removal. Science, 282, 2256-2258.
39. Johnson, P.T.J. & Thieltges, D.W. (2010) Diversity, decoys and the dilution effect: how ecological communities affect disease risk. Journal of Experimental Biology, 213, 961-970.
40. Jolles, A.E., Ezenwa, V.O., Etienne, R.S., Turner, W.C. & Olff, H. (2008) Interactions between Macroparasites and microparasites drive infection patterns in free-ranging African buffalo. Ecology, 89, 2239-2250.
41. Jones, K.E., Patel, N.G., Levy, M.A., Storeygard, A., Balk, D., Gittleman, J.L. & Daszak, P. (2008) Global trends in emerging infectious diseases. Nature, 451, 990-994.
42. Keeling, M.J., Bjornstad, O. & Grenfell, B.T. (2004) Metapopulation dynamics of infectious disease. Ecology, Evolution and Genetics of Metapopulations (eds I. Hanski & O.E. Gaggioti), pp. 415-446. Elsevier, Amsterdam.
43. Keeling, M.J. & Gilligan, C.A. (2000) Metapopulation dynamics of bubonic plague. Nature, 407, 903-906.
44. Keesing, F., Holt, R.D. & Ostfeld, R.S. (2006) Effects of species diversity on disease risk. Ecology Letters, 9, 485-498.
45. Keesing, F., Brunner, J., Duerr, S., Killilea, M., LoGiudice, K., Schmidt, K., Vuong, H. & Ostfeld, R.S. (2009) Hosts as ecological traps for the vector of Lyme disease. Proceedings of the Royal Society B, 276, 3911-3919.
46. Keesing, F., Belden, L.K., Daszak, P., Dobson, A., Harvell, C.D., Holt, R.D., Hudson, P., Jolles, A., Jones, K.E., Mitchell, C.E., Myers, S.S., Bogich, T. & Ostfeld, R.S. (2010) Impacts of biodiversity on the emergence and transmission of infectious diseases. Nature, 468, 647-652.
47. Kelly, D.W., Paterson, R.A., Townsend, C.R., Poulin, R. & Tompkins, D.M. (2009a) Has the introduction of brown trout altered disease patterns in native New Zealand fish? Freshwater Biology, 54, 1805-1818.
48. Kelly, D.W., Paterson, R.A., Townsend, C.R., Poulin, R. & Tompkins, D.M. (2009b) Parasite spillback: a neglected concept in invasion ecology? Ecology, 90, 2047-2056.
49. Kilpatrick, A.M., Briggs, C.J. & Daszak, P. (2009) The ecology and impact of chytridiomycosis: an emerging disease of amphibians. Evolution, 10, 9-118.
50. van Kleunen, M., Weber, E. & Fischer, M. (2010) A meta-analysis of trait differences between invasive and non-invasive plant species. Ecology Letters, 13, 235-245.
51. Kohler, S.L. & Hoiland, W.K. (2001) Population regulation in an aquatic insect: the role of disease. Ecology, 82, 2294-2305.
52. Kolar, C.S. & Lodge, D.M. (2001) Progress in invasion biology: predicting invaders. Trends in Ecology & Evolution, 16, 199-204.
53. Komar, N., Langevin, S., Hinten, S., Nemeth, N., Edwards, E., Hettler, D., Davis, B., Bowen, R. & Bunning, M. (2003) Experimental infection of north American birds with the New York 1999 strain of West Nile virus. Emerging Infectious Diseases, 9, 311-322.
54. Kopp, K. & Jokela, J. (2007) Resistant invaders can convey benefits to native species. Oikos, 116, 295-301.
55. Lafferty, K.D. & Holt, R.D. (2003) How should environmental stress affect the population dynamics of disease? Ecology Letters, 6, 654-664.
56. Lafferty, K.D., Arim, M., Briggs, J., Leo, G.D., Dobson, P., Dunne, J.A., Johnson, P.T.J., Armand, M., Pablo, A., Pascual, M., Poulin, R. & Thieltges, D.W. (2008) Parasites in food webs: the ultimate missing links. Ecology, 53, 3-546.
57. Lee, K.A. & Klasing, K.C. (2004) A role for immunology in invasion biology. Trends in Ecology & Evolution, 19, 523-529.
58. Lello, J., Boag, B., Fenton, A., Stevenson, I.R. & Hudson, P.J. (2004) Competition and mutualism among the gut helminths of a mammalian host. Nature, 428, 840-844.
59. Lello, J., Norman, R.A., Boag, B., Hudson, P.J. & Fenton, A. (2008) Pathogen interactions, population cycles, and phase shifts. American Naturalist, 171, 176-182.
60. Lloyd-Smith, J.O., Schreiber, S.J., Kopp, P.E. & Getz, W.M. (2005) Superspreading and the effect of individual variation on disease emergence. Nature, 438, 355-359.
61. Mack, R.N., Simberloff, D., Lonsdale, W.M., Evans, H., Clout, M. & Bazzaz, F.A. (2000) Biotic invasions: causes, epidemiology, global consequences, and control. Ecological Applications, 10, 689-710.
62. Malmstrom, C.M., McCullough, A.J., Johnson, H.A., Newton, L.A. & Borer, E.T. (2005) Invasive annual grasses indirectly increase virus incidence in California native perennial bunchgrasses. Oecologia, 145, 153-164.
63. Martin, L.B., Hasselquist, D. & Wikelski, M. (2006) Investment in immune defense is linked to pace of life in house sparrows. Oecologia, 56, 5-575.
64. Martin, L.B., Weil, Z.M. & Nelson, R.J. (2007) Immune defense and reproductive pace of life in Peromyscus mice. Ecology, 251, 6-2528.
65. McCallum, H.I., Barlow, N. & Hone, J. (2001) How should pathogen transmission be modelled? Trends in Ecology & Evolution, 16, 295-300.
66. McGeoch, M.A., Butchart, S.H.M., Spear, D., Marais, E., Kleynhans, E.J., Symes, A., Chanson, J. & Hoffmann, M. (2010) Global indicators of biological invasion: species numbers, biodiversity impact and policy responses. Diversity and Distributions, 16, 95-108.
67. Michel, A.L., Bengis, R.G., Keet, D.F., Hofmeyr, M., Klerk, L.M., Cross, P.C., Jolles, A.E., Cooper, D., Whyte, I.J., Buss, P. & Godfroid, J. (2006) Wildlife tuberculosis in South African conservation areas: implications and challenges. Veterinary Microbiology, 112, 91-100.
68. Mitchell, C.E. & Power, A.G. (2003) Release of invasive plants from fungal and viral pathogens. Nature, 421, 625-627.
69. Moore, S.M., Manore, C.A., Bokil, V.A., Borer, E.T. & Hosseini, P.R. (2011) Spatiotemporal model of barley and cereal yellow dwarf virus transmission dynamics with seasonality and plant competition. Bulletin of Mathematical Biology, 73, 2707-2730.
70. Mosmann, T.R. & Sad, S. (1996) The expanding universe of T-cell subsets: Th1, Th2 and more. Immunology Today, 17, 138-146.
71. O'Brien, V.A., Moore, A.T., Young, G.R., Komar, N., Reisen, W.K. & Brown, C.R. (2011) An enzootic vector-borne virus is amplified at epizootic levels by an invasive avian host. Proceedings of the Royal Society B, 278, 239-246.
72. Ostfeld, R.S., Glass, G.E. & Keesing, F. (2005) Spatial epidemiology: an emerging (or re-emerging) discipline. Trends in ecology & evolution, 20, 328-336.
73. Ostfeld, R.S. & Keesing, F. (2000) The function of biodiversity in the ecology of vector-borne zoonotic diseases. Canadian Journal of Zoology, 78, 2061-2078.
74. Pasternak, Z., Diamant, A. & Abelson, A. (2007) Co-invasion of a Red Sea fish and its ectoparasitic monogenean, Polylabris cf. mamaevi into the Mediterranean: observations on oncomiracidium behavior and infection levels in both seas. Parasitology Research, 100, 721-727.
75. Pedersen, A.B. & Fenton, A. (2006) Emphasizing the ecology in parasite community ecology. Trends in Ecology & Evolution, 22, 133-139.
76. Poulin, R., Paterson, R.A., Townsend, C.R., Tompkins, D.M. & Kelly, D.W. (2011) Biological invasions and the dynamics of endemic diseases in freshwater ecosystems. Freshwater Biology, 56, 676-688.
77. Rachowicz, L.J. & Briggs, C.J. (2007) Quantifying the disease transmission function: effects of density on Batrachochytrium dendrobatidis transmission in the mountain yellow-legged frog Rana muscosa. Journal of Animal Ecology, 76, 711-721.
78. Randolph, S.E. & Rogers, D.J. (2010) The arrival, establishment and spread of exotic diseases: patterns and predictions. Nature Reviews Microbiology, 8, 361-371.
79. Randolph, S.E., Chemini, C., Furlanello, C., Genchi, C., Hails, R.S., Hudson, P.J., Jones, L.D., Medley, G., Norman, R.A., Rizzoli, S.P., Smith, G. & Woolhouse, M.E.J. (2002) The ecology of tick-borne infections in wildlife reservoirs. The Ecology of Wildlife Diseases (eds P.J. Hudson, A. Rizzoli, B.T. Grenfell, J.A.P. Heesterbeek & A.P. Dobson), pp., 11, 9-138.
80. Roche, B. & Guégan, J.-François. (2011) Ecosystem dynamics, biological diversity and emerging infectious diseases. Comptes Rendus Biologies, 334, 385-392.
81. Rudolf, V.H.W. & Antonovics, J. (2005) Species coexistence and pathogens with frequency dependent transmission. American Naturalist, 166, 112-118.
82. Rushton, S.P., Lurz, P.W.W., Gurnell, J., Nettleton, P., Bruemmer, C., Shirley, M.D.F. & Sainsbury, A.W. (2006) Disease threats posed by alien species: the role of a poxvirus in the decline of the native red squirrel in Britain. Epidemiology and Infection, 134, 521-533.
83. Sakai, A.K., Allendorf, F.W., Holt, J.S., Lodge, D.M., Molofsky, J., With, K.A., Baughman, S., Cabin, R.J., Cohen, J.E., Ellstrand, N.C., McCauley, D.E., O'Neil, P., Parker, I.M., Thompson, J.N. & Weller, S.G. (2001) The population biology of invasive species. Annual Review of Ecology and Systematics, 32, 305-332.
84. Smith, M.J., Telfer, S., Kallio, E.R., Burthe, S., Cook, A.R., Lambin, X. & Begon, M. (2009) Host - pathogen time series data in wildlife support a transmission function between density and frequency dependence. Proceedings of the National Academy of Sciences of the USA, 106, 7905-7909.
85. Stuart, P., Mirimin, L., Cross, T.F., Sleeman, D.P., Buckley, N.J., Telfer, S., Birtles, R.J., Kotlik, P. & Searle, J.B. (2007) The origin of Irish bank voles Clethrionomys glareolus assessed by mitochondrial DNA analysis. Irish Naturalists' Journal, 28, 440-446.
86. Swei, A., Ostfeld, R.S., Lane, R.S. & Briggs, C.J. (2011) Impact of the experimental removal of lizards on Lyme disease risk. Proceedings of the Royal Society B, 278, 2970-2978. ST - Impact of the experimental removal.
87. Swinton, J., Woolhouse, M.E.J., Begon, M.E., Dobson, A.P., Ferroglio, E., Grenfell, B.T., Guberti, V., Hails, R.S., Heesterbeek, J.A.P., Lavazza, A., Roberts, M.G., White, P.J. & Wilson, K. (2002) Microparasite transmission and persistence. The Ecology of Wildlife Diseases (eds P.J. Hudson, A. Rizzoli, B.T. Grenfell, J.A.P. Heesterbeek & A.P. Dobson), pp. 83-101. Oxford University Press, Oxford.
88. Tatem, A.J., Hay, S.I. & Rogers, D.J. (2006) Global traffic and disease vector dispersal. Proceedings of the National Academy of Sciences of the USA, 103, 6242-6247.
89. Telfer, S., Bown, K.J., Sekules, R., Begon, M., Hayden, T. & Birtles, R. (2005) Disruption of a host-parasite system following the introduction of an exotic host species. Parasitology, 130, 661-668.
90. Telfer, S., Birtles, R., Bennett, M., Lambin, X., Paterson, S. & Begon, M. (2008) Parasite interactions in natural populations: insights from longitudinal data. Parasitology, 135, 767-787.
91. Telfer, S., Lambin, X., Birtles, R.J., Beldomenico, P., Burthe, S., Paterson, S. & Begon, M. (2010) Species interactions in a parasite community drive infection risk in a wildlife population. Science, 243, 243-246.
92. Thieltges, D.W., Reise, K., Prinz, K. & Jensen, K.T. (2008) Invaders interfere with native parasite-host interactions. Biological Invasions, 11, 1421-1429.
93. Thrall, P.H., Antonovics, J. & Hall, D.W. (1993) Host and pathogen coexistence in sexually transmitted and vector-borne diseases characterized by frequency-dependent disease transmission. American Naturalist, 142, 543-552.
94. Thuiller, W., Richardson, D.M., Rouget, M., Proches, S. & Wilson, J.R.U. (2006) Interactions between environment, species traits, and human uses describe patterns of plant invasions. Ecology, 87, 1755-1769.
95. Tollenaere, C., Rahalison, L., Ranjalahy, M., Duplantier, J., Rahelinirina, S. & Telfer, S. (2010) Susceptibility to Yersinia pestis experimental infection in wild Rattus rattus, reservoir of Plague in Madagascar. EcoHealth, 7, 242-247.
96. Tollenaere, C., Duplantier, J.-M., Rahalison, L., Ranjalahy, M. & Brouat, C. (2011) AFLP genome scan in the black rat (Rattus rattus) from Madagascar: detecting genetic markers undergoing plague-mediated selection. Molecular Ecology, 20, 1026-1038.
97. Tompkins, D.M., White, A.R. & Boots, M. (2003) Ecological replacement of native red squirrels by invasive greys driven by disease. Ecology Letters, 6, 189-196.
98. Tompkins, D.M., Dunn, A.M., Smith, M.J. & Telfer, S. (2011) Wildlife diseases: from individuals to ecosystems. Journal of Animal Ecology, 80, 19-38.
99. Torchin, M.E., Byers, J.E. & Huspeni, T.C. (2005) Differential parasitism of native and introduced snails: replacement of a parasite fauna. Biological Invasions, 7, 885-894.
100. Torchin, M.E., Lafferty, K.D., Dobson, A.P., McKenzie, V.J. & Kuris, A.M. (2003) Introduced species and their missing parasites. Nature, 421, 628-630.
101. Wilson, K., Bjørnstad, O.N., Dobson, A.P., Merler, S., Poglayen, G., Read, A.F. & Skorping, A. (2002) Heterogeneities in macroparasite infections: patterns and processes. The Ecology of Wildlife Diseases (eds P.J. Hudson, A. Rizzoli, B.T. Grenfell, J.A.P. Heesterbeek & A.P. Dobson), pp. 6-44. Oxford University Press, Oxford.