title: Hidden in plain sight: Cryptic and endemic malaria parasites in North American white-tailed deer (Odocoileus virginianus).
abstract: Malaria parasites of the genus Plasmodium are diverse in mammal hosts, infecting five mammalian orders in the Old World, but were long considered absent from the diverse deer family (Cervidae) and from New World mammals. There was a description of a Plasmodium parasite infecting a single splenectomized white-tailed deer (WTD; Odocoileus virginianus) in 1967 but none have been reported since, which has proven a challenge to our understanding of malaria parasite biogeography. Using both microscopy and polymerase chain reaction, we screened a large sample of native and captive ungulate species from across the United States for malaria parasites. We found a surprisingly high prevalence (up to 25%) and extremely low parasitemia of Plasmodium parasites in WTD throughout the eastern United States. We did not detect infections in the other ungulate species nor in western WTD. We also isolated the parasites from the mosquito Anopheles punctipennis. Morphologically, the parasites resemble the parasite described in 1967, Plasmodium odocoilei. Our analysis of the cytochrome b gene revealed two divergent Plasmodium clades in WTD representative of species that likely diverged 2.3 to 6 million years ago, concurrent with the arrival of the WTD ancestor into North America across Beringia. Multigene phylogenetic analysis placed these clades within the larger malaria parasite clade. We document Plasmodium parasites to be common in WTD, endemic to the New World, and as the only known malaria parasites from deer (Cervidae). These findings reshape our knowledge of the phylogeography of the malaria parasites and suggest that other mammal taxa may harbor infection by endemic and occult malaria parasites.
authors.affiliation: Center for Conservation and Evolutionary Genetics, Smithsonian Conservation Biology Institute, National Zoological Park, Washington, DC 20013-7012, USA.
authors.forename: Ellen S
authors.initials: ES
authors.lastname: Martinsen
authors.affiliation: Center for Conservation and Evolutionary Genetics, Smithsonian Conservation Biology Institute, National Zoological Park, Washington, DC 20013-7012, USA.
authors.forename: Nancy
authors.identifier: 0000-0002-6519-7671
authors.initials: N
authors.lastname: McInerney
authors.affiliation: Center for Conservation and Evolutionary Genetics, Smithsonian Conservation Biology Institute, National Zoological Park, Washington, DC 20013-7012, USA.
authors.forename: Heidi
authors.identifier: 0000-0002-0072-6237
authors.initials: H
authors.lastname: Brightman
authors.affiliation: Rock Creek National Park, 3545 Williamsburg Lane, NW, Washington, DC 20008, USA.
authors.forename: Ken
authors.initials: K
authors.lastname: Ferebee
authors.affiliation: Department of Pathology, Smithsonian Institution, National Zoological Park, Washington, DC 20560, USA.
authors.forename: Tim
authors.initials: T
authors.lastname: Walsh
authors.affiliation: Conservation Ecology Center, Smithsonian Conservation Biology Institute, National Zoological Park, 1500 Remount Road, Front Royal, VA 22630, USA.
authors.forename: William J
authors.identifier: 0000-0002-8102-0200
authors.initials: WJ
authors.lastname: McShea
authors.affiliation: Conservation Ecology Center, Smithsonian Conservation Biology Institute, National Zoological Park, 1500 Remount Road, Front Royal, VA 22630, USA.
authors.forename: Tavis D
authors.identifier: 0000-0001-6762-4115
authors.initials: TD
authors.lastname: Forrester
authors.affiliation: Department of Conservation Medicine, Smithsonian Conservation Biology Institute, National Zoological Park, Front Royal, VA 22630, USA.
authors.forename: Lisa
authors.initials: L
authors.lastname: Ware
authors.affiliation: Department of Conservation Medicine, Smithsonian Conservation Biology Institute, National Zoological Park, Front Royal, VA 22630, USA.
authors.forename: Priscilla H
authors.identifier: 0000-0002-2908-8726
authors.initials: PH
authors.lastname: Joyner
authors.affiliation: Sackler Institute for Comparative Genomics, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA.
authors.forename: Susan L
authors.initials: SL
authors.lastname: Perkins
authors.affiliation: Department of Biological Sciences, Behavioral and Molecular Ecology Research Group, University of Wisconsin-Milwaukee, 3209 North Maryland Avenue, Milwaukee, WI 53211, USA.
authors.forename: Emily K
authors.initials: EK
authors.lastname: Latch
authors.affiliation: Southeastern Cooperative Wildlife Disease Study, Department of Population Health, College of Veterinary Medicine, University of Georgia, Athens, GA 30602, USA.; Warnell School of Forestry and Natural Resources, University of Georgia, Athens, GA 30602, USA.
authors.forename: Michael J
authors.identifier: 0000-0003-2452-5015
authors.initials: MJ
authors.lastname: Yabsley
authors.affiliation: Department of Biology, Marsh Life Sciences Building, University of Vermont, Burlington, VT 05405, USA.
authors.forename: Joseph J
authors.initials: JJ
authors.lastname: Schall
authors.affiliation: Center for Conservation and Evolutionary Genetics, Smithsonian Conservation Biology Institute, National Zoological Park, Washington, DC 20013-7012, USA.
authors.forename: Robert C
authors.identifier: 0000-0002-2792-7055
authors.initials: RC
authors.lastname: Fleischer
mesh_terms: D000818:Animals; D000852:Anopheles / Q000469:parasitology; D003670:Deer / Q000469:parasitology*; D058507:Host Specificity; D007303:Insect Vectors / Q000469:parasitology; D008288:Malaria / Q000469:parasitology / Q000662:veterinary*; D010802:Phylogeny; D058974:Phylogeography; D010961:Plasmodium / Q000145:classification / Q000235:genetics / Q000302:isolation & purification; D014481:United States
keywords: Malaria parasites; Odocoileus; Plasmodium; haemosporidians