Evolutionary relationships of avian Eimeria species among other apicomplexan protozoa: monophyly of the apicomplexa is supported

Abstract

Direct, reverse transcriptase-mediated, partial sequencing of the small-subunit (16S-like) ribosomal RNA (srRNA) of Eimeria tenella and E. acervulina was performed. Sequences were aligned by eye with six previously published, partial or complete srRNA sequences of apicomplexan protists (Plasmodium berghei, Theileria annulata, Cryptosporidium sp., Toxoplasma gondii, Sarcocystis muris, and S. gigantea). Six eukaryotic protists (a slime mold, a yeast, two dinoflagellates, and two ciliates) acted as an outgroup for a parsimony-based phylogenetic analysis (PAUP Ver. 3.0.). The 188 phylogenetically informative sites (i.e., those positions that neither were unvaried nor had only autapomorphic substitutions) supported a single tree topology 481 steps in length with a consistency index of 0.65 in which the monophyly of the Apicomplexa was supported. The two Eimeria species and S. muris, S. gigantea, and T. gondii formed a pair of monophyletic groups that were sister groups. The two Sarcocystis species were not hypothesized to be sister taxa. The genera Plasmodium and Cryptosporidium were hypothesized to form the sister group to these five coccidia and T. annulata. A priori data-editing techniques that deleted "variable" positions prior to analysis failed to recognize the monophyly of the Apicomplexa when the same parsimony-based tree-building algorithm was used. Inability of the outgroup taxa to root the well-supported ingroup tree (Apicomplexa) at a unique site when these taxa were used individually for this purpose reinforces the need for an appropriate, multiple-taxon outgroup in such analyses.