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      A large 28S rDNA-based phylogeny confirms the limitations of established morphological characters for classification of proteocephalidean tapeworms (Platyhelminthes, Cestoda)

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          Abstract

          Abstract

          Proteocephalidean tapeworms form a diverse group of parasites currently known from 315 valid species. Most of the diversity of adult proteocephalideans can be found in freshwater fishes (predominantly catfishes), a large proportion infects reptiles, but only a few infect amphibians, and a single species has been found to parasitize possums. Although they have a cosmopolitan distribution, a large proportion of taxa are exclusively found in South America. We analyzed the largest proteocephalidean cestode molecular dataset to date comprising more than 100 species (30 new), including representatives from 54 genera (80%) and all subfamilies, thus significantly improving upon previous works to develop a molecular phylogeny for the group. The Old World origin of proteocephalideans is confirmed, with their more recent expansion in South America. The earliest diverging lineages are composed of Acanthotaeniinae and Gangesiinae but most of the presently recognized subfamilies (and genera) appear not to be monophyletic; a deep systematic reorganization of the order is thus needed and the present subfamilial system should be abandoned. The main characters on which the classical systematics of the group has been built, such as scolex morphology or relative position of genital organs in relation to the longitudinal musculature, are of limited value, as demonstrated by the very weak support for morphologically-defined subfamilies. However, new characters, such as the pattern of uterus development, relative ovary size, and egg structure have been identified, which may be useful in defining phylogenetically well-supported subgroups. A strongly supported lineage infecting various snakes from a wide geographical distribution was found. Although several improvements over previous works regarding phylogenetic resolution and taxon coverage were achieved in this study, the major polytomy in our tree, composed largely of siluriform parasites from the Neotropics, remained unresolved and possibly reflects a rapid radiation. The genus Spasskyellina Freze, 1965 is resurrected for three species of Monticellia bearing spinitriches on the margins of their suckers.

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          Accounting For Alignment Uncertainty in Phylogenomics

          Uncertainty in multiple sequence alignments has a large impact on phylogenetic analyses. Little has been done to evaluate the quality of individual positions in protein sequence alignments, which directly impact the accuracy of phylogenetic trees. Here we describe ZORRO, a probabilistic masking program that accounts for alignment uncertainty by assigning confidence scores to each alignment position. Using the BALIBASE database and in simulation studies, we demonstrate that masking by ZORRO significantly reduces the alignment uncertainty and improves the tree accuracy.
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            Majority-rule reduced consensus trees and their use in bootstrapping.

            Bootstrap analyses are usually summarized with majority-rule component consensus trees. This consensus method is based on replicated components and, like all component consensus methods, it is insensitive to other kinds of agreement between trees. Recently developed reduced consensus methods can be used to summarize much additional agreement on hypothesised phylogenetic relationships among multiple trees. The new methods are "strict" in the sense that they require agreement among all the trees being compared for any relationships to be represented in a consensus tree. Majority-rule reduced consensus methods are described and their use in bootstrap analyses is illustrated with a hypothetical and a real example. The new methods provide summaries of the bootstrap proportions of all n-taxon statements/partitions and facilitate the identification of hypotheses of relationships that are supported by high bootstrap proportions, in spite of a lack of support for particular components or clades. In practice majority-rule reduced consensus profiles may contain many trees. The size of the profile can be reduced by constraints on minimal bootstrap proportions and/or cardinality of the included trees. Majority-rule reduced consensus trees can also be selected a posteriori from the profile. Surrogates to the majority-rule reduced consensus methods using partition tables or tree pruning options provided by widely used phylogenetic inference software are also described. The methods are designed to produce more informative summaries of bootstrap analyses and thereby foster more informed assessment of the strengths and weaknesses of complex phylogenetic hypotheses.
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              Interrelationships and evolution of the tapeworms (Platyhelminthes: Cestoda).

              Interrelationships of the tapeworms (Platyhelminthes: Cestoda) were examined by use of small (SSU) and large (LSU) subunit ribosomal DNA sequences and morphological characters. Fifty new complete SSU sequences were added to 21 sequences previously determined, and 71 new LSU (D1-D3) sequences were determined for the complementary set of taxa representing each of the major lineages of cestodes as currently understood. New sequences were determined for three amphilinidean taxa, but were removed from both alignments due to their excessively high degree of divergence from other cestode sequences. A morphological character matrix coded for supraspecific taxa was constructed by the modification of matrices from recently published studies. Maximum-parsimony (MP) analyses were performed on the LSU, SSU, LSU+SSU, and morphological data partitions, and minimum-evolution (ME) analyses utilizing a general time reversible model of nucleotide substitution including estimates of among-site rate heterogeneity were performed on the molecular data partitions. Resulting topologies were rooted at the node separating the Gyrocotylidea from the Eucestoda. The LSU data were found to be more informative than the SSU data and were more consistent with inferences from morphology, although nodal support was generally weak for most basal nodes. One class of transitions was found to be saturated for comparisons between the most distantly related taxa (gyrocotylideans vs cyclophyllideans and tetrabothriideans). Differences in the topologies resulting from MP and ME analyses were not statistically significant. Nonstrobilate orders formed the basal lineages of trees resulting from analysis of LSU data and morphology. Difossate orders were basal to tetrafossate orders, the latter of which formed a strongly supported clade. A clade including the orders Cyclophyllidea, Nippotaeniidea, and Tetrabothriidea was supported by all data partitions and methods of analysis. Paraphyly of the orders Pseudophyllidea, Tetraphyllidea, and Trypanorhyncha was consistent among the molecular data partitions. Inferences are made regarding a monozoic (nonsegmented) origin of the Eucestoda as represented by the Caryophyllidea and for the evolution of the strobilate and acetabulate/tetrafossate conditions having evolved in a stepwise pattern. Copyright 2001 Academic Press.
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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2015
                27 April 2015
                : 500
                : 25-59
                Affiliations
                [1 ]Natural History Museum of Geneva, CP 6434, CH - 1211 Geneva 6, Switzerland
                [2 ]Natural History Museum, Life Sciences, Cromwell Road, London SW7 5BD, UK
                [3 ]Institute of Parasitology, Biology Centre of the Czech Academy of Sciences, Branišovská 31, 370 05 České Budějovice, Czech Republic
                [4 ]Department of Genetics and Evolution, University of Geneva, CH - 1205 Geneva, Switzerland
                Author notes
                Corresponding author: Jean Mariaux ( jean.mariaux@ 123456ville-ge.ch )

                Academic editor: B. Georgiev

                Article
                10.3897/zookeys.500.9360
                4432239
                25987870
                52ba1b02-1ed2-4622-b5be-4144f49682b6
                Alain de Chambrier, Andrea Waeschenbach, Makda Fisseha, Tomáš Scholz, Jean Mariaux

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 8 February 2015
                : 23 March 2015
                Categories
                Research Article

                Animal science & Zoology
                eucestoda,proteocephalidae,systematics,molecular phylogeny,host-parasite associations,spasskyellina

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