Ecological Archives M075-022-A5

Robert E. Ricklefs, Bethany L. Swanson, Sylvia M. Fallon, Alejandro Martínez-Abraín, Alexander Scheuerlein, Julia Gray, and Steven C. Latta. 2005. Community relationships of avian malaria parasites in southern Missouri. Ecological Monographs 75:543–559.

Appendix E. Evidence for species distinction in malaria parasites from host distribution.

Species limits are difficult to define in malaria parasites without evidence of linkage disequilibrium of independent genetic markers (Bensch et al. 2004) . Species names are based primarily on broad host taxon and morphological characters of the parasite gametophytes seen in blood smears (Valkiunas 2005) . Evidence for species distinction can also be inferred from distribution of closely related parasite lineages on different host taxa in the same region (Ricklefs et al. 2004) . Host distributions of the most closely related lineages represented in our sample from the Missouri Ozarks are shown in Table E1. The clearest differentiation of host distributions occurs in the comparison of lineages 14 and 25 (boldface indicates Plasmodium), which differ by 2 nucleotides (0.6%). Lineage 14 is widely distributed in North American and the Greater Antilles and occurs in a variety of nine-primaried oscine passerines (warblers, finches, and icterids) whereas lineage 25 has been recovered only from Tufted Titmice (BBI: Baeolophus bicolor) in the Ozarks and Michigan. Lineage 3 (Haemoproteus), recovered from seven warbler individuals of three species and a Carolina wren (TLU: Thryothorus ludovicianus), differs from lineage 33, recovered from two cardinals, a common grackle, and a western bluebird in four different locations, by four nucleotides. Host preferences of the other lineage pairs are not distinctive. Lineages 12 and 13 have both been recovered from Red-eyed Vireos (VOL: Vireo olivaceus); lineages 26 and 27 from vireos, albeit different species; lineages 30 and 31 from Piranga tanagers (POL and PRU); and lineages 16 and 21 from White-eyed Vireos (VGR: Vireo griseus). Without additional information, we treat these lineages as separate parasite species, although future genetic analyses might erase these distinctions.

Table E1. Host distributions of pairs of parasite lineages differing by four or fewer nucleotides out of about 350. The columns marked lineages A and B refer to the identification numbers of the lineages being compared (e.g., 12 and 13) and D refers to the number of nucleotide differences between the lineages.






Hosts of lineage A

Hosts of lineage B




OZ: 1 DDO, 4 VOL; PR: 4 VAL; CY: 1 MNI





OZ: 1 VGR; KZ: 1 VSO





OZ: 1 POL, 1 VOL





OZ: 2 VGR, 2 HVE, 1 IVI; CT: 2 DCA, 1 SNO

OZ: 1 VOL, 1 VGR; PR: 14 LPO, 12 CFA




OZ: 4 PAM, 2 DDO, 1 SAU, 1 TLU

OZ: 1 CCA; CT: 1 QQU; TY: 1 CCA; NM: 1 SME




OZ: 1 MVA, 1 DDI, 1 CCA, 1 SAU; BRC: 2 QQU, 1 CBR; CT: 1 DCA, 2 GTR, 1 MAT; KZ: 1 CUS, 1 DCO, 2 DMA; DR: 1 DPA; CY: 1 PLU

OZ: 2 BBI; KZ: 3 BBI

   Note: Locations (boldface type) are: BRC, Bird Rehabilitation Center, St. Louis, Missouri; CY, Cayman Islands; CT, Connecticut; DR, Dominican Republic; KZ, Kalamazoo, Michigan; NM, New Mexico; OZ, Ozarks; PR, Puerto Rico; TY, Tyson Research Station, St. Louis, Missouri.

Species acronyms: BBI = Baeolophus bicolor Tufted Titmouse, CBR = Corvus brachyrhynchos Common Crow, CCA = Cardinalis cardinalis Northern Cardinal, CFA = Coereba flaveola Bananaquit, CUS = Catharus ustulatus Swainson’s Thrush, DCA = Dendroica castanea Bay-breasted Warbler, DCO = Dendroica coronata Yellow-rumped Warbler, DDI = Dendroica discolor Prairie Warbler, DDO = Dendroica dominica Yellow-throated warbler, DMA = Dendroica magnolia Magnolia Warbler, DPA = Dendroica palmarum Palm Warbler, GTR = Geothlypis trichas Common Yellowthroat, HVE = Helmitheros vermivorus Worm-eating Warbler, IVI = Icteria virens Yellow-breasted Chat, LPO = Loxigilla portoricensis Puerto Rican Bullfinch, MAT = Molothrus ater Brown-headed Cowbird, MNI = Melopyrrha nigra Cuban Bullfinch, MVA = Mniotilta varia Black-and-White Warbler, PAM = Parula Americana Northern Parula, PLU = Piranga ludoviciana Western Tanager, POL = Piranga olivacea Scarlet Tanager, PRU = Piranga rubra Summer Tanager, QQU = Quicalus quiscula Common Grackle, SAU = Seirurus aurocapillus Ovenbird, SME = Sialia mexicanus Western Bluebird, SNO = Seiurus novaboracensis Northern Waterthrush, TLU = Thryothorus ludovicianus Carolina Wren, VAL = Vireo altiloquus Black-whiskered Vireo, VFL = Vireo flavifrons Yellow-throated Vireo, VGR = Vireo griseus White-eyed Vireo, VOL = Vireo olivaceus Red-eyed Vireo, VSO = Vireo solitarius Blue-headed Vireo.


Bensch, S., J. Pérez-Tris, J. Waldenström, and O. Hellgren. 2004. Linkage between nuclear and mitochondrial DNA sequences in avian malaria parasites: multiple cases of cryptic speciation? Evolution 58:1617–1621.

Ricklefs, R. E., S. M. Fallon, and E. Bermingham. 2004. Evolutionary relationships, cospeciation, and host switching in avian malaria parasites. Systematic Biology 52:111–119.

Valkiunas, G. 2005. Avian Malaria Parasites and Other Haemosporidia. CRC Press, Boca Raton, Florida.

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