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Phylogenetic systematics and the evolutionary history of some intestinal flatworm parasites (Trematoda : Digenea: Plagiorchioidea) of Anurans O'Grady, Richard Terence


Historical structuralism is presented as a research program in evolutionary biology. It uses patterns of common ancestry as initial hypotheses in explaining evolutionary history. Such patterns, represented by phylogenetic trees, or cladograms, are postulates of persistent ancestral traits. These traits are evidence of historical constraints on evolutionary change. Patterns and processes consistent with a cladogram are considered to be consistent with an initial hypothesis of historical constraint. As an application of historical structuralism, a phylogenetic analysis is presented for members of the digenean plagiorchioid genera Glypthelmins Stafford, 1905 and Haplometrana Lucker, 1931. The eight species studied are intestinal parasites of frogs and toads in North, Central, and South America. In a Wagner parsimony analysis of 21 morphological characters with both the PAUP and PHYSYS computer programs, a single phylogenetic tree with a consistency index of 84.8% can be inferred. This suggests strong historical constraint in the evolution of the characters examined. It is postulated that the eight species form a monophyletic group (clade), consisting of two less inclusive clades. Glypthelmins hyloreus and G. pennsylvaniensis comprise one of these clades; G. robustus, G. shastai, H. intestinalis, G. californiensis, G. quieta, and G. facioi comprise the other. G. robustus, found in Bufo marinus in Colombia, is both the southernmost and the most plesiomorphic member of its clade. Glypthelmins californiensis, G. quieta, and G. facioi form a clade, and parasitize frogs in the Rana pipiens complex in Mexico, eastern North America, and Central America, respectively. Glypthelmins shastai and H. intestinalis, the latter of which is the only member of its genus, form a western North American clade, and parasitize Bufo boreas and Rana pretiosa, respectively. The phylogenetic analysis includes a redescription of G. shastai, the synonymy of the genus Haplometrana with Glypthelmins, the redescription of H. intestinali s as G. intestinalis, an emended diagnosis of the genus Glypthelmins, and the first account of the life cycle of G. californiensis. Three aspects of phylogenetic analysis are examined in detail. These are the coding of multistate character trees, the use of parasite data to infer host relationships, and the properties of the Consistency Index and the F-Ratio. It is proposed that the Consistency Index be calculated without non-homoplasious autapomorphic characters. For the present study, this modification gives a value of 76.9%. Using the phylogenetic tree as a general reference system of patterns of common ancestry, it is inferred from developmental studies that (1) there is no conflict between the phylogenetic relationships indicated by only larval or only adult characters, and that (2) the evolution of some of the characters involved certain types of heterochrony. Paedomorphic heterochrony is inferred to have occurred in the evolution of the uterus in G. shastai, H. intestinalis, G. californiensis, G. quieta, and G. facioi. Peramorphic heterochrony is inferred to have occurred in the evolution of the penetration glands in G. facioi, and of the hindbody in H. intestinalis. The relatively longer hindbody of H. intestinalis was experimentally induced to show paedomorphic development by raising specimens of H. intestinalis in Bufo boreas, which is the host of G. shastai, its sister-species. By one year after infection, the relative length of the hindbody is shorter, and is equal to that of the primitive state found in G. shastai. The phylogenetic relationships among the anuran hosts are re-analyzed. There is 80% congruence between them and the postulated phylogenetic tree for their parasites, suggesting strong historical association between the parasite and host groups. This inference of coevolution is further supported by the concordance of the present geographical distributions of the parasites and their hosts with the historical geology of the areas in which they occur. This implies an historical association between the areas and the organisms.

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