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Could the Blood Parasite Leucocytozoon Deter Mallard Range Expansion? Author(S): Dave Shutler, C

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Could the Blood Parasite Leucocytozoon Deter Mallard Range Expansion? Author(S): Dave Shutler, C Could the Blood Parasite Leucocytozoon Deter Mallard Range Expansion? Author(s): Dave Shutler, C. Davison Ankney, Darrell G. Dennis Source: The Journal of Wildlife Management, Vol. 60, No. 3 (Jul., 1996), pp. 569-580 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3802074 Accessed: 04/12/2008 06:50 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=acg. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. Allen Press is collaborating with JSTOR to digitize, preserve and extend access to The Journal of Wildlife Management. http://www.jstor.org COULDTHE BLOODPARASITE LEUCOCYTOZOON DETER MALLARD RANGEEXPANSION? DAVESHUTLER,1 Department of Zoology, Universityof WesternOntario, London, Ontario, N6A 5B7, Canada C. DAVISONANKNEY, Department of Zoology, Universityof WesternOntario, London, Ontario, N6A 5B7, Canada DARRELLG. DENNIS,Canadian Wildlife Service, 153 NewboldCourt, London, Ontario, N6E 1Z7, Canada Abstract: We investigated whether the blood parasite Leucocytozoon simondi could slow mallard (Anas platyrhynchos)population growth in the east that has been associatedwith Americanblack duck (A. rubripes; hereafter black duck) population decline. Susceptibility to parasites was compared among F1 ducklings produced from crossesbetween mallard and black ducks from areas of Leucocytozoonendemicity (Ontario), and between mallards from an area free of Leucocytozoon (Saskatchewan). We produced 6 "types" of ducklings: Ontario black duck x Ontario black duck (OB x OB), Ontario black duck x Ontario mallard (OB x OM), Ontario black duck x Saskatchewan mallard (OB x SM), OM x OM, OM x SM, and SM x SM. We predicted that because of probable coevolution of black ducks and Leucocytozoon, black duck ducklings would have resistance to the parasite. We also predicted that Ontario genes would confer some resistance to ducklings because these ducklings' parents had survived exposure to Leucocytozoon. In contrast, we predicted that mallard and Saskatchewan genes would not confer resistance, i.e., OB x OB ducklings would have greatest resistance to Leucocytozoon, SM x SM ducklings would have least, and remaining duckling types would have intermediate resistance. Of 169 ducklings exposed in 2 years in 3 geographically separate locales, none died, showed noticeable symptoms, or otherwise behaved abnormally. Nonetheless, weekly blood smears indicated that 91% of ducklings became infected, and many developed intense para- sitemias. However, infection intensities were not different among the 6 duckling types. In addition, hematocrits were not lowered by intense infections. These results suggest that the effects of Leucocytozoon on wild waterfowl populations have been overestimated, and that Leucocytozoon will not prevent further range expansionof mallards. J. WILDL.MANAGE. 60(3):569-580 Key words: American black ducks, Anas platyrhynchos, Anas rubripes, blood parasites,hybrids, Leuco- cytozoon simondi, mallard, Ontario, Saskatchewan. Mallards appear to be displacing American Leucocytozoon has the potential to prevent or black ducks in eastern North America through slow mallard encroachment into those parts of competition and/or introgressive hybridization the black duck range in which Leucocytozoon (Rogers and Patterson 1984, Brodsky and occurs. Weatherhead 1984, Ankney et al. 1987, Mer- Leucocytozoon is transmitted by black flies endino et al. 1993). Thus, continued mallard (Simuliidae) (Shewell 1955) and can infect 100% range expansion could lead to local or complete of mallards and black ducks in some populations extinction of black ducks. However, the blood in northeastern North America (O'Roke 1934, parasite Leucocytozoon simondi may be a bar- Nelson and Gashwiler 1941, Chernin 1952, rier to this outcome (Khan and Fallis 1968, Den- Trainer et al. 1962, Bennett et al. 1974, 1975, nis 1974). Leucocytozoon is endemic in much 1991). In contrast, Leucocytozoon is rarely re- of the historic range of black ducks, and the ported from central North America (Savage and parasite reportedly causes substantial duckling Isa 1955, Burgess 1957). Differences in Leuco- mortality (O'Roke 1934, Fallis et al. 1956, 1974; cytozoon prevalence arise because, in the latter Barrow et al. 1968) that is higher among mallard area, black fly vectors have less swift water than black duck ducklings (Khan and Fallis breeding habitat and are often insufficiently nu- 1968). In this paper, we test whether differential merous to spread the parasite (Herman 1968). susceptibility of mallards and black ducks to Historically, mallards occupied western and central North America, whereas black ducks oc- the eastern of the continent Present address:Canadian Wildlife Service, 115 cupied part (Bellrose Perimeter Road, Saskatoon,Saskatchewan, S7N 0X4, 1976). This historical distribution pattern is con- Canada. sistent with current distributions of Leucocy- 569 570 LEUCOCYTOZOONAND DUCKS * Shutler et al. J. Wildl. Manage. 60(3):1996 tozoon (Khan and Fallis 1968). Because mal- his crew provided guidance in the ways of Leu- lards have not coevolved with Leucocytozoon cocytozoon; the staff of Canadian Wildlife Ser- but are closely related to black ducks genetically vice in London provided logistical assistance (J. (Ankney et al. 1986, Avise et al. 1990), they Robinson, N. North, and J. Sullivan); B. Clark should be more susceptible to the parasite (Ewald of Canadian Wildlife Service in Saskatchewan 1983, 1993; Van Riper et al. 1986, Black 1992, sent us ducklings; A. Toline provided logistical Bennett et al. 1993). assistance in Algonquin Park, and the Ontario If a Leucocytozoon-mediated barrier to mal- Ministry of Natural Resources provided logis- lards still exists, however, it is necessary to ex- tical assistance at Swan Lake (in particular, H. plain continuing mallard range expansion into Anderson, J. Rice, D. Strickland, and M. Wil- areas where the parasite is endemic (Johnsgard ton). Drs. A. M. Fallis and R. A. Khan provided and DiSilvestro 1976, Rusch et al. 1989, Mer- historical details of earlier studies, and com- endino and Ankney 1994). One possibility is that mented on our conclusions. Funding was pro- mallards have acquired genes for Leucocyto- vided by the Natural Sciences and Engineering zoon resistance, either from introgressive hy- Research Council of Canada, the Black Duck bridization with resistant black ducks, or via Joint Venture, the Ontario Federation of An- natural selection that has culled non-resistant glers and Hunters, and the University of West- mallards (collectively, the resistance hypothe- ern Ontario. sis). A second possibility is that mallards are able to survive in habitats within the boreal only METHODS forest where Leucocytozoon is not present (e.g., areas without fast running water [the suscepti- Initial Preparations bility hypothesis]). The null hypothesis is that a In April 1991, we advertised rewards to in- Leucocytozoon-mediated barrier does not and dividuals finding active wild mallard and black did not exist, and that no differences in suscep- duck nests in an area of Leucocytozoon endem- tibility occur between duck species. icity in Ontario bounded approximately by Par- Our objective was to determine whether ry Sound in the west (80?0'W), North Bay in the duckling mortality could limit mallard range north (46?30'N), Ottawa in the east (75?30'W), expansion into areas of Leucocytozoon endem- and Haliburton in the south (45?0'N). Eggs we icity. We tested whether greater parasite resis- collected from these nests were transported to tance occurred among ducklings with a greater Lake St. Clair, an area where vectors of Leu- proportion of genes from presumably coevolved cocytozoon are not found. In addition, we re- black ducks, or from parents recently exposed ceived 40 Saskatchewan mallard ducklings (Can. to Leucocytozoon; i.e., resistance of black ducks Wildl. Serv., Saskatchewan). Eggs were hatched > mallards, and resistance of Ontario birds > in incubators, and we gave each duckling in- resistance of Saskatchewan birds. Our protocol dividually numbered web tags. Ducklings were was to expose ducklings to black fly vectors of sexed by cloacal inspection (Schemnitz 1980) Leucocytozoon in seminatural conditions, and within a day after hatching, and those that sur- then observe ducklings for differences in mor- vived to 4 weeks were given individually num- tality or signs of illness. We first ensured that bered leg bands. Because we needed hybrids for sex, initial date of exposure, and site of exposure our experiments on Leucocytozoon,
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