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      Co-invaders: The effects of alien parasites on native hosts

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          Graphical abstract

          Highlights

          • Co-introduced parasites are transported to a new area with an alien host.

          • Co-invading parasites are those which are co-introduced and switch from alien to native hosts.

          • Co-invading parasites with indirect life cycles are reported as frequently as those with direct life cycles.

          • Co-invaders are usually more virulent in native hosts than in alien hosts.

          Abstract

          We define co-introduced parasites as those which have been transported with an alien host to a new locality, outside of their natural range, and co-invading parasites as those which have been co-introduced and then spread to new, native hosts. Of 98 published studies of co-introductions, over 50% of hosts were freshwater fishes and 49% of parasites were helminths. Although we would expect parasites with simple, direct life cycles to be much more likely to be introduced and establish in a new locality, a substantial proportion (36%) of co-introductions were of parasites with an indirect life cycle. Seventy-eight per cent of co-introduced parasites were found in native host species and can therefore be classed as co-invaders. Host switching was equally common among parasites with direct and indirect life cycles. The magnitude of the threat posed to native species by co-invaders will depend, among other things, on parasite virulence. In 16 cases where co-introduced parasites have switched to native hosts and information was available on relative virulence, 14 (85%) were more virulent in native hosts than in the co-introduced alien host. We argue that this does not necessarily support the naïve host theory that co-invading parasites will have greater pathogenic effects in native hosts with which they have no coevolutionary history, but may instead be a consequence of the greater likelihood for parasites with lower virulence in their natural host to be co-introduced.

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          Most cited references88

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          Introduced species and their missing parasites.

          Damage caused by introduced species results from the high population densities and large body sizes that they attain in their new location. Escape from the effects of natural enemies is a frequent explanation given for the success of introduced species. Because some parasites can reduce host density and decrease body size, an invader that leaves parasites behind and encounters few new parasites can experience a demographic release and become a pest. To test whether introduced species are less parasitized, we have compared the parasites of exotic species in their native and introduced ranges, using 26 host species of molluscs, crustaceans, fishes, birds, mammals, amphibians and reptiles. Here we report that the number of parasite species found in native populations is twice that found in exotic populations. In addition, introduced populations are less heavily parasitized (in terms of percentage infected) than are native populations. Reduced parasitization of introduced species has several causes, including reduced probability of the introduction of parasites with exotic species (or early extinction after host establishment), absence of other required hosts in the new location, and the host-specific limitations of native parasites adapting to new hosts.
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            How should pathogen transmission be modelled?

            Host-pathogen models are essential for designing strategies for managing disease threats to humans, wild animals and domestic animals. The behaviour of these models is greatly affected by the way in which transmission between infected and susceptible hosts is modelled. Since host-pathogen models were first developed at the beginning of the 20th century, the 'mass action' assumption has almost always been used for transmission. Recently, however, it has been suggested that mass action has often been modelled wrongly. Alternative models of transmission are beginning to appear, as are empirical tests of transmission dynamics.
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              Identifying Reservoirs of Infection: A Conceptual and Practical Challenge

              (2002)
              Many infectious agents, especially those that cause emerging diseases, infect more than one host species. Managing reservoirs of multihost pathogens often plays a crucial role in effective disease control. However, reservoirs remain variously and loosely defined. We propose that reservoirs can only be understood with reference to defined target populations. Therefore, we define a reservoir as one or more epidemiologically connected populations or environments in which the pathogen can be permanently maintained and from which infection is transmitted to the defined target population. Existence of a reservoir is confirmed when infection within the target population cannot be sustained after all transmission between target and nontarget populations has been eliminated. When disease can be controlled solely by interventions within target populations, little knowledge of potentially complex reservoir infection dynamics is necessary for effective control. We discuss the practical value of different approaches that may be used to identify reservoirs in the field.
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                Author and article information

                Contributors
                Journal
                Int J Parasitol Parasites Wildl
                Int J Parasitol Parasites Wildl
                International Journal for Parasitology: Parasites and Wildlife
                Elsevier
                2213-2244
                24 April 2014
                24 April 2014
                August 2014
                : 3
                : 2
                : 171-177
                Affiliations
                [a ]Freshwater Fish Group and Fish Health Unit, School of Veterinary and Life Sciences, Murdoch University, Murdoch 6150, Western Australia, Australia
                [b ]Fisheries Department, Faculty of Agriculture, Gonbad Kavous University, Iran
                Author notes
                [* ]Corresponding author. Tel.: +61 8 9360 7509; fax: +61 8 9360 7512. a.lymbery@ 123456murdoch.edu.au
                Article
                S2213-2244(14)00012-1
                10.1016/j.ijppaw.2014.04.002
                4145144
                25180161
                b8b97922-7480-453d-b2c9-39596fecee42
                © 2014 Published by Elsevier Ltd on behalf of Australian Society for Parasitology.

                This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/3.0/).

                History
                : 20 November 2013
                : 9 April 2014
                : 9 April 2014
                Categories
                Invited Review

                invasive species,co-introduction,co-invasion,host-switching,virulence

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